RUBIACEAE

qian cao ke
Chen Tao ()1, Zhu Hua ()2, Chen Jiarui ( Chen Chia-jui)3;
Charlotte M. Taylor4, Friedrich Ehrendorfer5, Henrik Lantz6, A. Michele Funston4, Christian Puff5
Trees, shrubs, annual or perennial herbs, subshrubs, vines, or lianas, infrequently monocaulous or creeping and rooting at nodes,
terrestrial or infrequently epiphytic, with bisexual flowers, infrequently dioecious, or rarely polygamo-dioecious (Diplospora,
Galium, Guettarda, perhaps Brachytome) or monoecious (Galium), evergreen or sometimes deciduous (Hymenodictyon), sometimes
armed with straight to curved spines (formed by modified stems or peduncles), infrequently with elongated principal stems bearing
lateral short shoots (i.e., brachyblasts; Benkara, Catunaregam, Ceriscoides, Himalrandia, Leptodermis, Serissa), infrequently with
lateral branches or short shoots spinescent (i.e., prolonged, sharp, and leafless at apex), infrequently with reduced internodes that give
an appearance of verticillate leaf arrangement (Brachytome, Damnacanthus, Duperrea, Rothmannia, Rubovietnamia), infrequently
with buds resinous (Gardenia) or mucilaginous (Scyphiphora), infrequently with tissues fetid when bruised, [rarely with swollen
hollow stems or leaf bases housing ants (Neonauclea)]; branchlets terete to angled or quadrate, in latter two cases often becoming
terete with age, or rarely flattened (Wendlandia) or winged (Hedyotis, Rubia), buds conical or rounded with stipules valvate or imbricate, or infrequently flattened with stipules erect and pressed together (Cinchona, Haldina, Nauclea, Neonauclea). Raphides present
or absent. Leaves opposite, verticillate, or apparently verticillate (i.e., closely set due to reduced internodes), decussate or occasionally distichous, petiolate to sessile, infrequently somewhat to strongly anisophyllous, rarely punctate- or striate-glandular (Galium);
margins flat to occasionally undulate or crisped, entire or rarely lobed (Hymenodictyon, Morinda) to denticulate or serrate (Hymenodictyon, Leptomischus, Ophiorrhiza, Wendlandia); secondary veins pinnate or rarely triplinerved or palmate (Hedyotis, Rubia), free
(i.e., eucamptodromous) or uniting near margins (i.e., brochidodromous) in weak to well-developed or rarely substraight submarginal
vein, sometimes with foveolate (i.e., pitted or cryptlike) and/or tufted (i.e., pubescent) domatia (i.e., structures that house mites) in
abaxial axils, these rarely also present in axils of tertiary veins (Morinda), with presence of domatia often variable within a species;
tertiary and/or quaternary venation rarely arranged in regular squares (Guettarda), regular rectangles (i.e., clathrate; Urophyllum), or
lineolate (i.e., closely parallel within each areole; Timonius); petiole rarely articulate at base (Ixora); stipules persistent with leaves,
deciduous before leaves, or quickly caducous, interpetiolar and infrequently fused to adjacent petioles or leaf bases, sometimes
united around stem into a sheath, rarely completely united into a conical cap (i.e., calyptrate; Gardenia), with interpetiolar portion
variously triangular in general shape to truncate, with apex entire or bilobed, multifid, lacerate, setose, or laterally appendaged, with
apex, lobes, setae, and/or appendages sometimes glandular (Chassalia, Hedyotis, Hymenodictyon, Knoxia, Mitchella, Mycetia,
Neanotis, Ophiorrhiza, Pentas, Pseudopyxis, Psychotria, Trailliaedoxa), internally (i.e., adaxially) with small to well-developed colleters (i.e., glandular trichomes), these infrequently persistent after stipules fall (Psychotria), or stipules rarely expanded into 1 to
several leaflike segments and then apparently absent due to leaflike form that gives an appearance of verticillate leaves (Argostemma,
Asperula, Galium, Microphysa, Phuopsis, Rubia). Inflorescences terminal, axillary (i.e., borne at both axils at a node), or pseudoaxillary (i.e., borne consistently in 1 axil per node; lateral), sometimes apparently leaf-opposed due to marked anisophylly, or rarely
superaxillary (Damnacanthus, Diplospora) or cauline (Mycetia), variously cymose to thyrsiform, corymbiform, paniculiform,
racemiform, spiciform, fasciculate, or capitate and few to many flowered or occasionally reduced to a solitary flower, pedunculate
(peduncle here used for basalmost axis supporting inflorescence or solitary flower) to sessile, when sessile often with 3 principal axes
(i.e., tripartite), bracteate or bracts sometimes reduced or apparently absent, with bracts (here usually including bracts borne on pedicels or next to flowers, i.e., bracteoles) generally triangular to linear or sometimes leaflike (i.e., similar to normal or somewhat
reduced leaves) and rarely enlarged, petaloid, and resembling calycophylls (Dunnia, Neohymenopogon), infrequently fused and involucral, occasionally fused in pairs (i.e., forming a calyculus, or calyculate), infrequently thickened and spatulate to clavate or conical
(genera of Naucleeae), or infrequently stipuliform, rarely glandular (Damnacanthus, Mycetia), multifid to fimbriate (Damnacanthus,
Kelloggia, Spermacoce) or spinescent (Phuopsis). Flowers sessile to pedicellate (pedicel here used for ultimate axis immediately
supporting a single flower, except when this is a peduncle), bisexual and monomorphic, distylous, or rarely tristylous (Chassalia,
Pentas), unisexual with 2 forms generally similar except for corolla size and hypanthium development, or rarely cleistogamous
(Ophiorrhiza), actinomorphic or rarely zygomorphic (Argostemma), sometimes with ovaries of individual flowers partially to fully
fused (Mitchella, Morinda, Mouretia, Nauclea), variously diurnal or nocturnal, usually sweetly fragrant, protandrous [or rarely protogynous], occasionally with secondary pollen presentation (e.g., Ixora, Pavetta, Phuopsis, Scyphiphora, genera of Naucleeae). Calyx
gamosepalous and fused to inferior ovary in hypanthium or ovary portion, this generally ellipsoid, turbinate, obconic, cylindrical, or
occasionally subglobose to hemispherical, glabrous and smooth, pubescent, or rarely tuberculate (Galium) or with unusual flattened
(Dentella) or hooked, sometimes glandular trichomes (Galium, Kelloggia), or rarely longitudinally ridged to winged (Gardenia,
1 Fairy Lake Botanical Garden, 160 Xianhu Road, Liantang, Shenzhen, Guangdong 518004, Peoples Republic of China.
2 Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences, Menglun Town, Mengla County, Xishuangbanna, Yunnan 666303, Peoples Republic of

China.

3 Institute of Botany, Chinese Academy of Sciences, 20 Nanxincun, Xiangshan, Beijing 100093, Peoples Republic of China.
4 Missouri Botanical Garden, P.O. Box 299, Saint Louis, Missouri 63166-0299, U.S.A.
5 Department of Systematic and Evolutionary Botany, Faculty of Life Sciences, University of Vienna, Rennweg 14, A-1030 Vienna, Austria.
6 Museum of Evolution, Botany Section (Fytoteket), Evolutionary Biology Center, Uppsala University, Norbyvgen 16, SE-752 36 Uppsala, Sweden.

57

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RUBIACEAE

Spiradiclis), above this developed into a free limb portion, this limb variously tubular to cupular or infrequently reduced to obsolete
(Asperula, Coffea, Galium, Leptunis, Microphysa, Ophiorrhiza, Phuopsis, Rubia), truncate to 4- or 5(9)-denticulate (i.e., lobes
reduced to tiny projections along a generally truncate margin) or shallowly to deeply 4- or 5(9)-lobed, open in bud or rarely with
lobes markedly imbricate (Emmenopterys, Keenania), [or rarely spathaceous (i.e., fused into a conical cap that splits irregularly)],
inside variously near base and/or at sinuses between lobes with few to numerous small colleters, rarely densely or markedly veined
(Clarkella, Myrioneuron, Pseudopyxis), rarely on margins with well-developed, sessile to stalked glands (Mycetia), lobes generally
triangular to linear, occasionally obtuse to lanceolate or oblanceolate, or rarely prolonged into a slender shaft bearing a thickened
apical portion (Neonauclea), occasionally slightly to markedly unequal on an individual flower with all lobes of different lengths or
infrequently in unequal pairs of similar lengths (Diodia, Mitracarpus, Spermacoce), infrequently with 1(to 5, Mussaenda) lobe on
some (or all, Mussaenda) flowers of an inflorescence enlarged into a calycophyll (i.e., a membranous to papery, petaloid, veined,
white to colored blade borne on a generally well-developed stipe; Emmenopterys, Morinda, Mussaenda, Schizomussaenda). Corolla
large and often showy to reduced, gamopetalous, white, yellow, orange, red, blue, purple, and/or pale green, when nocturnal often
white at anthesis becoming yellow with age, variously funnelform, salverform (i.e., hypocrateriform), tubular, campanulate, or occasionally rotate to infrequently urceolate (i.e., swollen in basal part of tube; Canthium, Lasianthus) or inflated (i.e., markedly swollen
in middle or upper part of tube; Keenania, Leptomischus), infrequently curved in tube and/or gibbous (i.e., asymmetrically swollen at
very base of tube; Chassalia, Guettarda, Mycetia, Ophiorrhiza), infrequently differing in shape between long-styled and short-styled
forms (Antirhea), infrequently markedly fleshy to leathery (Caelospermum, Damnacanthus, Fosbergia, Rothmannia, Timonius,
Urophyllum), inside glabrous to variously pubescent with pubescence frequently confined to throat, outside infrequently ridged to
winged (Cinchona, Ophiorrhiza), rarely fenestrate in tube (i.e., with longitudinal slits; Damnacanthus, Paederia), lobes (3 or)4 or
5(11), shorter than or occasionally longer than tube, acute or less often obtuse to rounded at apex, generally spreading to somewhat
reflexed at anthesis, infrequently with margins crisped to irregular, densely ciliate, and/or appendaged (Cinchona, Luculia, Rondeletia, Saprosma, Serissa), in bud imbricate (and usually quincuncial), valvate, valvate-induplicate, valvate-reduplicate, or convolute (i.e., contorted) to left or rarely to right (Coptosapelta, Rothmannia), usually with aestivation consistent within a genus, occasionally with wings, ridges, and/or rounded to hornlike thickenings or protuberances on back or at apex (Lerchea, Ophiorrhiza),
rarely cucullate (Lerchea). Stamens adnate to corolla, free or rarely fused to stigma (Acranthera), alternate to corolla lobes and isomerous (i.e., equal in number to lobes) or rarely more numerous (Gardenia) [to rarely fewer], inserted variously in corolla throat,
tube, or infrequently at base and sometimes appearing free (Galium), included to exserted, with point of insertion and position of
anthers usually differing between long-styled and short-styled forms of distylous flowers, with staminodes of pistillate flowers generally similar to stamens but smaller; filaments well developed to reduced or obsolete, free or rarely coherent (Argostemma) or fused
(Acranthera, Argostemma), variously glabrous to pubescent, occasionally markedly flattened (Hymenodictyon, Kelloggia); anthers
free or rarely coherent or fused (Argostemma), 4-thecal or rarely 2-thecal (Hymenodictyon), 2-celled, in outline generally narrowly
oblong, linear (i.e., narrowly fusiform), narrowly lanceolate, or narrowly elliptic, at base occasionally bifid (i.e., sagittate; Caelospermum, Cephalanthus, Duperrea, Hamelia, Neohymenopogon, Scyphiphora), dorsifixed (i.e., medifixed to dorsifixed near base) to
occasionally basifixed, infrequently pubescent (Hyptianthera, Lerchea), with dehiscence introrse by longitudinal slits or rarely by
apical pores (Argostemma), with connective infrequently prolonged into an apical and/or sometimes basal appendage (Acranthera,
Argostemma, Hyptianthera, Morinda, Rubovietnamia, Wendlandia), appendages rarely fused into a cone (Acranthera); pollen variously 3- or 4(or 5)-colpate and generally subglobose or occasionally 3- or 4-porate, 525-colpate and disk-shaped or ellipsoidal, in
tetrahedral tetrads and 3- or 4-porate, cylindrical with 2 pores, or inaperturate. Ovary inferior [or rarely secondarily superior], sometimes fused between flowers (Mitchella, Morinda, Mouretia), (1 or)2(10)-celled (i.e., locular), with ovules 1 or 2 to numerous in
each cell (i.e., locule) on basal, axile (i.e., inserted on septum), apical, or infrequently parietal (Ceriscoides, Gardenia) placentas, in
staminate flowers usually with ovary reduced and pistillode composed of structures similar to but smaller than style and stigma, in
distylous flowers usually with ovary similarly developed but style and stigma differing in size and position in flower and sometimes
stigmas also differing in shape between long-styled and short-styled forms; style 1, terminal on ovary, developed or rarely reduced
(Galium, Microphysa), variously glabrous to pubescent, surrounded at base by well-developed fleshy disk [or this rarely reduced],
this disk variously annular, conical, 2-parted, or shallowly lobed, glabrous or rarely pubescent (Clarkella, Mouretia, Timonius);
stigmas free or rarely fused to anther connectives (Acranthera), 1- or 2(10)-lobed (i.e., these lobes often equivalent to stigmas 1 or
2 to 10 of some authors), with whole stigma or lobes variously capitate, linear, spatulate, clavate, lobed, or infrequently mitriform
(i.e., shaped like a drinking glass) to cylindrical with recessed attachment (Canthium, Mitragyna, Pyrostria), glabrous or infrequently
pubescent (Clarkella, Hyptianthera), variously included to exserted, with receptive surfaces introrse, apical, or internal. Infructescences generally similar to inflorescences but occasionally with part or all changing color, orientation, shape, and/or other characteristics as the fruit mature. Fruit simple or rarely multiple (i.e., a syncarp; Mitchella, Morinda, Nauclea), variously capsular with dehiscence loculicidal, septicidal, circumscissile (i.e., around equator or middle; Mitracarpus), or through an apical beak (i.e., prolonged
disk portion, sometimes to give appearance of partially superior ovary; Hedyotis, Neanotis, Neohymenopogon, Ophiorrhiza, Pentas,
Spiradiclis) or operculum (i.e., circular lid; Argostemma, Leptodermis, Leptomischus, Mouretia, Pseudopyxis); or fleshy, small to
quite large, and baccate (i.e., with numerous seeds enclosed by fleshy to juicy pulp or endocarp and usually indehiscent); or drupaceous (i.e., with 1 to several seeds enclosed in pyrenes), fleshy to occasionally dry, and indehiscent or infrequently dehiscent releasing
pyrenes (Paederia, Serissa); or schizocarpous (i.e., dry and separating into segments) with mericarps (i.e., segments, cocci, nutlets)
indehiscent (e.g., Asperula, Cephalanthus, Richardia); calyx limb persistent or deciduous usually leaving a circular scar, sometimes
with persistent carpophore or septum (Adina); pyrenes (i.e., seeds enclosed in and dispersing with endocarp layer) when present and

RUBIACEAE

59

ovules all developed 110-locular and 110-seeded (i.e., fruit containing several pyrenes and each pyrene with 1 seed in 1 locule, or
pyrenes solitary in each fruit and comprising entire ovary), ellipsoid to subglobose, plano-convex (i.e., hemispherical), concavoconvex, lenticular, or angled (i.e., narrow with 2 large inner faces and a small outer face), with outer wall hard to cartilaginous (Caelospermum) or infrequently papery (Coffea, Pavetta), smooth to ridged or sulcate on dorsal (i.e., abaxial) surface, without (Psychotria) or usually with evident preformed germination slits, pores, and/or opercula generally on ventral (i.e., adaxial) surface, rarely
winged (Paederia) or pubescent (Caelospermum, Paederia); seeds 1 to numerous, small (0.11.9 mm), medium-sized (25 mm) to
large (5.120 mm), variously ellipsoid, lenticular, flattened, oblanceoloid, angled, or plano-convex, smooth to variously winged,
foveolate, tuberculate, papillose, and/or striate; endosperm (i.e., albumen) fleshy, oily, corneous (i.e., horny) and entire or infrequently ruminate (Psychotria), or rarely absent (Antirhea); embryo variously shaped. x = 617, most commonly 11, less frequently 9
or 12.
About 660 genera and 11,150 species: cosmopolitan family, with most genera and species in humid tropical regions; 97 genera (three endemic,
ten introduced) and 701 species (352 endemic, 23 introduced, six of unconfirmed occurrence) in China.
This is one of the largest families of flowering plants and is represented nearly worldwide though it is most abundant in the tropical regions of
both hemispheres. Important economic members of the family are coffee (Coffea), quinine (Cinchona), madder (Rubia), ipecac (Carapichea Aublet),
and various horticultural plants, notably Gardenia, Ixora, Mussaenda, and Pentas. Several genera include pantropical weeds, notably Mitracarpus,
Richardia, and Spermacoce, of neotropical origin, and Oldenlandia (Hedyotis) of African origin. A number of species are used for various medicinal
purposes.
Genera, tribes, and subfamilies of Rubiaceae characteristically either have or lack raphides; this character is generally considered informative
taxonomically in this family. Raphides are tiny, needle-shaped crystals that are produced in groups inside some cells of leaf, stem, flower, and fruit
tissues. They apparently function to protect the plant from herbivores. Raphides are often visible with a hand lens on dried specimens, particularly in
soft tissues such as petals, the calyx hypanthium, and the young leaves at the apex of the stem, but may be difficult to see especially in succulent
plants.
Leaf domatia are common in Rubiaceae and are sometimes taxonomically informative at least for separation of species. These domatia provide
protection for tiny mites that live on the leaves and that help protect the plant from herbivores and fungi by eating insect eggs, small insects, and
spores. These domatia occasionally take the form of foveolae or crypts, but more often are formed by small groups or tufts of pubescence, which is
sometimes different in form from pubescence found on other parts of the plant.
At least half of the species of Rubiaceae have distylous flowers: an individual species has two flower forms, the long-styled form (or pin) with
the stigmas borne above the anthers and the short-styled form (or thrum) with the anthers borne above the stigmas. An individual plant bears only one
flower form, and the flowers are incompatible with other flowers of similar form whether borne on the same or another plant. Heterostyly is unusual in
the flowering plants and is found in several other plant families that mostly have three floral forms instead of two with the third form (the
homostylous) with the stigmas and anthers borne at the same level. Tristyly has only been documented very rarely for Rubiaceae, although it was
noted by FRPS for Chassalia curviflora in China.
A number of species of many genera of Rubiaceae have calyx lobes that are unequal in size on an individual flower, and in some species of
several genera one or more of the calyx lobes are markedly different, much enlarged and also often petaloid. These structures are found on flowers,
where they are typically brightly colored and apparently function to attract pollinators, and/or on the fruit, where they are dry and brown and
apparently function in dispersal of the fruit or seeds. These structures have variously been called petaloid calyx lobes, a morphologically descriptive
term; semaphylls, a term that refers to the assumed function in pollination; and calycophylls, the term used here that refers to the leaflike shape of
the calyx lobe. These structures are sometimes mistaken for bracts; a few species of Rubiaceae do also have petaloid bracts (e.g., Dunnia), but those
structures are inserted on the inflorescence axes or immediately below the ovary of the flower.
The woody Rubiaceae are better represented (i.e., more diverse) in moist temperate regions of China than in comparable habitats in North
America, as detailed by Latham and Ricklefs (in Ricklefs & Schluter, Spec. Diversity Ecol. Communities, 294314. 1993). They concluded that this
pattern, which is also found in a number of other dicotyledonous families, is due largely to historical factors. The principal factors they identified are
the extinction of woody Rubiaceae in these other regions, while the plants were able to persist in E Asia (e.g., Emmenopterys), and the long-term
direct connection of the moist temperate and moist tropical zones in E Asia vs. the separation of these zones by deserts and large water bodies in other
regions. They agreed with Wolfe (in Nelson & Rosen, Vicariance Biogeogr. 413427. 1981) that the connections of plant ranges between E North
America and E Asia were broken by the end of the Eocene, and thus that species of the several genera that are found in both regions (e.g.,
Cephalanthus) have had separate evolutionary histories for some time. Several SE Asian Rubiaceae genera, all found in China, appear to represent the
most basal living lineages of the family, notably Acranthera, Coptosapelta, and Luculia (Bremer et al., Syst. Biol. 48: 413435. 1999; Bremer &
Manen, Pl. Syst. Evol. 225: 4372. 2000; Rydin et al., Pl. Syst. Evol. 278: 101123. 2009).
A number of species of Rubiaceae are cultivated as ornamentals; most of these are mentioned and some are keyed and described here.
Additional information on many of the cultivated Rubiaceae was presented by Puff and Chamchumroon (Thai Forest Bull., Bot. 31: 7594.
2003).
Complete Rubiaceae floras for E Asia are very few. The Chinese Rubiaceae were treated comprehensively in 1999 by FRPS; the treatment here
is based primarily on that work. The Fl. URSS was published in English in 2000, but this is an unedited, unmodified translation of the 1958 work. The
Fl. Japan has been published in several editions but includes relatively few genera and is rather idiosyncratic. The genus overview within the
Rubiaceae of Thailand: A Pictorial Guide to Indigenous and Cultivated Genera by Puff et al. (43240. 2005) is a useful recent reference.
Some morphological features that are characteristic of the Rubiaceae have been described, measured, and named very differently by different
authors, and the differing terminology has generated some confusion. In particular, the terminology and measurements used for the calyx and
gynoecium have differed widely. The ovary of Rubiaceae flowers is typically inferior, and the calyx is gamosepalous with its free portion, or limb,
arising from the top of the ovary portion; this limb varies from reduced (i.e., not developed) to developed, and truncate to deeply lobed. Most
Rubiaceae authors have distinguished the inferior ovary portion of the flower from the free calyx limb; the ovary portion has sometimes been called

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RUBIACEAE

simply an ovary, but sometimes considered to include the fused calyx and corolla tissue covering it and called a hypanthium. However, several
authors, including in FRPS, have treated as one structure the inferior ovary together with the unlobed lower portion of the calyx limb, when this is
present, as the calyx tube and the lobed portion of the calyx as a separate structure, the calyx lobes. The calyx tube is often clearly delimited and
measurable, but the inclusion of the ovary in this means that this measurement can sometimes include ovaries that have been fertilized and are
developing into fruit, thus are larger than and not comparable to the ovaries at anthesis even though the corolla may be still attached to the flowers.
Here the ovary or hypanthium portion of the flower is measured separately from the limb; the term hypanthium is used to link these measurements
to those of FRPS. The limb is measured separately here because the length of the tubular portion of this often includes taxonomically useful
information, and while the calyx tube measurement is lost when the fruit are developed, the length of the tubular portion of the free calyx limb is
usually still evident on the fruit thus facilitating identification. Another portion of the gynoecium that has been regarded differently by different
authors is the distinction between the stigma and style: the structures regarded as 2-lobed stigmas by some authors are regarded as a style with 2
separate stigmas by as many other authors. Morphological study of the androecium and gynoecium here during preparation of a flora treatment has
not been detailed and no focused effort has been made here to systematize these morphological interpretations.
Inflorescence terminology is complicated in most flowering plant families. The terms bract and bracteole have been carefully and regularly
applied by some Rubiaceae authors, with the term bracteole used only for the bracts that immediately subtend the flowers or in some cases that are
found anywhere along the pedicels, and the term bract used for the remaining structures. However, other authors have not distinguished these
structures, or have not distinguished them regularly or consistently. The term floral bract here refers to the structures found on the pedicels or that
immediately subtend sessile flowers. Bracts and leaves are generally considered homologous structures, and in Rubiaceae these often vary
continuously along the inflorescences. In particular, Rubiaceae commonly have somewhat reduced leaves subtending the inflorescences, and bracts
that are somewhat enlarged and occasionally leaflike. These structures have been variously called reduced leaves or foliose bracts or foliaceous
bracts (here called leaflike bracts). In particular, many species with branched inflorescences sometimes have rather large, leaflike bracts subtending
the basalmost pair of secondary axes; these have been various interpreted as enlarged bracts at the top of the peduncle, or vegetative leaves that are
borne at the base of a sessile, tripartite inflorescence.
Significant measurements found on plants from outside China are noted in the discussions following the species descriptions and/or are included
within brackets in the descriptions in accordance with Flora of China style. Authors in FRPS described the seeds and embryos of some Rubiaceae but
not others; their descriptions are reported. In many cases no information is available about these features for Chinese Rubiaceae. Authors in FRPS also
described the pollen of a few genera; these reports are not included here because a number of very detailed, extensively documented pollen
descriptions are available elsewhere, which show that much variation is found in pollen morphology at the species level in Rubiaceae and as yet
Rubiaceae genera are not well characterized in terms of pollen. Thus, the very limited information available only at the genus level here does not seem
well integrated into this flora treatment. The counts presented here for leaf veins apply to the secondary veins, as done by the majority of Rubiaceae
authors. Many species of Rubiaceae also frequently have rather well-developed though shorter intersecondary veins, which often vary widely in
number and degree of development between populations or plants; these apparently were occasionally combined with the secondary veins in the
measurements given by FRPS, but such measurements are subject to significant misinterpretation by authors working in other regions and the varied
usage in that work adds yet more confusion.
Where data were missing from the descriptions in the first draft for the Flora of China, the terms unknown, not known, not noted, and
not seen were retained (with not known consisted to unknown). Where these terms were used in the habitats and elevational ranges they were
deleted in accordance with the style of previous Flora of China volumes. Where missing data were denoted with question marks, e.g., shape??, or
with constructions such as __ __ , these were deleted because it was not apparent which of the above categories of missing data applied.
The genus Khasiaclunea Ridsdale (Blumea 24: 347. 1979) is known with certainty from India and Myanmar and provisionally from N Vietnam;
this may also be present in China, but no confidently determined specimens have yet been seen. The single species, K. oligocephala (Haviland)
Ridsdale (Blumea 24: 347. 1979; Adina oligocephala Haviland, J. Linn. Soc., Bot. 33: 46. 1897), was reported by Ridsdale (loc. cit.) as probably
found in China; however, this genus is not treated here.
Lo Hsienshui, Ko Wancheung, Chen Weichiu, Hsue Hsianghao & Wu Hen. 1999. Rubiaceae (1). In: Chen Weichiu, ed., Fl. Reipubl. Popularis
Sin. 71(1): ixvii, 1432; Lo Hsienshui, Ko Wancheung, Chen Weichiu & Ruan Yunzhen. 1999. Rubiaceae: Rubioideae. In: Chen Weichiu, ed., Fl.
Reipubl. Popularis Sin. 71(2): ixvi, 1377; Wan Wenhao. 2000. Theligonaceae. In: Chen Chia-jui, ed., Fl. Reipubl. Popularis Sin. 52(2): 147151.

Key 1
1a. Leaves apparently whorled and estipulate, 4 to numerous at each node; herbs, twiners, and low shrubs; calyx limb
reduced to a thin rim or absent .................................................................................................................................................... Key 2
1b. Leaves paired to whorled with stipules developed between each pair; calyx limb well developed to reduced or
apparently absent.
2a. Acaulescent to caulescent herbs, soft subshrubs, or herbaceous vines or clambering plants, if tall subshrubs then
stems annual ............................................................................................................................................................................. Key 3
2b. Low to tall woody shrubs, trees, or lianas with well-developed secondary growth and perennial stems .............................. Key 4
Key 2
1a. Flowers 5-merous and fruit fleshy, dispersing as 1 drupaceous structure, red to orange or black ...................................... 79. Rubia
1b. Flowers 4-merous or 5-merous, fruit dry though sometimes inflated, dispersing as 1 structure or splitting into
2 structures.
2a. Fruit inflated, dispersing as 1 structure; flowers 4-merous ..................................................................................... 50. Microphysa
2b. Fruit not inflated, schizocarpous and dispersing as 2 partially to completely separated mericarps; flowers
4- or 5-merous.
3a. Corollas rotate to campanulate or broadly funnelform ............................................................................................... 28. Galium
3b. Corollas tubular to salverform or funnelform.

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61

4a. Corollas 5-merous ................................................................................................................................................. 70. Phuopsis

4b. Corollas 4-merous (in our species).
5a. Leaves narrowly elliptic to lanceolate, with developed blades; mericarps ellipsoid and generally straight ....... 7. Asperula
5b. Leaves linear, with blades hardly developed; mericarps obovoid and markedly curved .................................. 46. Leptunis
Key 3
1a. Flowers fused in pairs by their ovaries; fruit fused in pairs ............................................................................................ 51. Mitchella
1b. Flowers and fruit free.
2a. Plants monoecious, with unisexual anemophilous flowers, staminate with 3 or 5 corolla lobes, 630 stamens,
and no ovary, pistillate with 3 corolla lobes, reduced staminodes, and a 1-celled ovary; calyx limb reduced
to absent .................................................................................................................................................................... 91. Theligonum
2b. Plants with bisexual insect-pollinated flowers, with 46 corolla lobes, with 46 stamens, and 25-celled
ovary both developed; calyx limb well developed to reduced or absent.
3a. Calyx limb reduced to a thin truncate rim or absent.
4a. Flowers 5-merous and fruit fleshy, orange to red or black ......................................................................................... 79. Rubia
4b. Flowers 4- or 5-merous and fruit dry, schizocarpous or capsular, green to brown.
5a. Ovules numerous in each cell; fruit capsular, mitriform and laterally markedly flattened (this shape
often evident in young developing fruit), with numerous small seeds ........................................................ 65. Ophiorrhiza
5b. Ovules 1 per cell; fruit indehiscent or schizocarpous with 2 mericarps, ellipsoid to subglobose,
sometimes flattened, with 2 seeds.
6a. Fruit inflated, dispersing as a single structure ............................................................................................ 50. Microphysa
6b. Fruit not inflated, schizocarpous, dispersing as 2 partially to completely separated mericarps.
7a. Corolla salverform to tubular or funnelform ................................................................................................. 7. Asperula
7b. Corolla rotate to campanulate or broadly funnelform ................................................................................... 28. Galium
3b. Calyx limb developed, with lobes sometimes reduced but still evident.
8a. Ovary and fruit densely covered by well-developed, unusual, flattened or curved and glandular trichomes
or projections.
9a. Ovary and fruit covered with flattened trichomes, these sometimes with apical thickenings; flowers
solitary, terminal or pseudoaxillary, sessile or subsessile ................................................................................... 21. Dentella
9b. Ovary and fruit covered with glandular curved trichomes; flowers several and pedicellate in terminal
and pseudoaxillary, irregularly branched cymes .............................................................................................. 41. Kelloggia
8b. Ovary and fruit smooth and glabrous to variously pubescent but trichomes neither regularly glandular nor
markedly flattened.
10a. Plants extensively twining, climbing, or clambering, with stems slender and herbaceous or suffrutescent
and flexuous.
11a. Plants climbing by adventitious roots (P. serpens); stipules deciduous after distalmost few nodes;
fruit fleshy, drupaceous, red ....................................................................................................................... 74. Psychotria
11b. Plants not climbing by adventitious roots, these sometimes scattered along stem but not principal
supports; stipules persistent or deciduous; fruit dry, green or becoming brown or black.
12a. Plants not fetid when bruised; fruit capsular with several to numerous small angled seeds;
corollas with 4 or 5 valvate lobes .............................................................................................................. 35. Hedyotis
12b. Plants usually with fetid odor when bruised; fruit schizocarpous, dry, inflated, with 2 flattened
winged pyrenes; corollas with 46 valvate-induplicate lobes ................................................................. 66. Paederia
10b. Plants weak to erect, creeping on ground, self-supporting or weakly shortly clambering.
13a. Cultivated plants; corolla white to pink or red, 2025 mm; fruit usually not developing .............................. 68. Pentas
13b. Wild plants; corolla white to red or purple, 155 mm; fruit usually developing.
14a. Stipules setose, laciniate, multifid, fimbriate, and/or dentate to markedly erose, if lobed then lobes
4 or more and entire or erose.
15a. Calyx and corolla lobes each 6; fruit schizocarpous with 3 mericarps ............................................... 76. Richardia
15b. Calyx and corolla lobes 35; fruit indehiscent, schizocarpous with 2 mericarps, or capsular.
16a. Flowers solitary and sessile, or several and sessile in sessile heads or glomerules.
17a. Fruit a circumscissile capsule; calyx lobes usually with hyaline margins; seeds 2 with
cruciform attachment scar ......................................................................................................... 52. Mitracarpus
17b. Fruit indehiscent, schizocarpous, or capsular and splitting longitudinally or apically;
calyx lobes with margins not hyaline; seeds 2 to numerous with simple rounded to
linear attachment scar.
18a. Seeds 1 per cell, 2 per fruit, fruit indehiscent, schizocarpous splitting into mericarps,
or capsular splitting longitudinally and deeply.
19a. Fruit indehiscent or splitting into 2 indehiscent mericarps ......................................................... 22. Diodia

62

RUBIACEAE

19b. Fruit capsular, splitting septicidally into 2 segments with 1 or both segments then
splitting loculicidally ........................................................................................................... 86. Spermacoce
18b. Seeds 2 or more per cell, 4 or more per fruit, fruit indehiscent, schizocarpous splitting
into mericarps, or capsular splitting primarily through apical beak or apical half.
20a. Plants without fetid odor when bruised, usually drying green or brownish green; seeds
angled to lenticular .................................................................................................................... 35. Hedyotis
20b. Plants usually with fetid odor when bruised, usually drying black to gray; seeds peltate
to plano-convex ......................................................................................................................... 60. Neanotis
16b. Flowers solitary to numerous, sessile to pedicellate or pedunculate in fascicles or cymes,
if flowers sessile then borne severally in cymes or in pedunculate heads or glomerules.
21a. Fruit schizocarpous, flattened, with 2 suborbicular indehiscent mericarps ....................................... 42. Knoxia
21b. Fruit capsular, subglobose to ovoid or mitriform, opening to release several to numerous
seeds.
22a. Fruit mitriform and laterally markedly flattened (this shape often evident in young
developing fruit) ..................................................................................................................... 65. Ophiorrhiza
22b. Fruit subglobose to ovoid, angled to ridged but strongly flattened laterally.
23a. Fruit opening through apical lid that develops from disk .................................................. 73. Pseudopyxis
23b. Fruit dehiscent through linear openings across top and often extending down sides.
24a. Flowers 5-merous ............................................................................................................... 88. Spiradiclis
24b. Flowers 4- or 5-merous.
25a. Plants without fetid odor when bruised, usually drying green or brownish green;
seeds angled to lenticular ................................................................................................... 35. Hedyotis
25b. Plants usually with fetid odor when bruised, usually drying black to gray; seeds
peltate to plano-convex ...................................................................................................... 60. Neanotis
14b. Stipules entire, 2-lobed, or 3-lobed, lobes entire.
26a. Fruit a fleshy drupe with 24 pyrenes; principal stems creeping, with short erect reproductive
stems.
27a. Inflorescences several flowered, capitate, and pedunculate; leaves cordiform to ovate .................. 30. Geophila
27b. Flowers solitary, sessile or subsessile; leaves elliptic ........................................................................... 64. Nertera
26b. Fruit dry, or fleshy and baccate with several to numerous seeds; habit erect to creeping with
or without short erect reproductive stems.
28a. Stipules densely covered with numerous, closely set, parallel or palmate fibers or veins ......... 58. Myrioneuron
28b. Stipules smooth, without evident veins or fibers or 1- or 2-costate.
29a. Fruit dry and indehiscent or schizocarpous.
30a. Fruit indehiscent, obconical, several seeded; small herbs with well-developed tubers ............. 17. Clarkella
30b. Fruit schizocarpous with 2 flattened, suborbicular mericarps; erect to rather tall
herbs without tubers ........................................................................................................................ 42. Knoxia
29b. Fruit dry and capsular, or fleshy and indehiscent.
31a. Fruit mitriform and laterally markedly flattened (this shape often evident in young
developing fruit) ..................................................................................................................... 65. Ophiorrhiza
31b. Fruit subglobose to ovoid, laterally not or only weakly flattened.
32a. Fruit an operculate capsule (dehiscence mode unknown in Keenania).
33a. Flowers 1 to several in cymes or fascicles, all of them generally well separated by
each other on inflorescence axes and/or pedicels.
34a. Corolla rotate to campanulate; ovary and fruit 2-celled ................................................ 6. Argostemma
34b. Corolla tubular-funnelform to salverform; ovary and fruit 4- or 5-celled ................... 73. Pseudopyxis
33b. Flowers several to numerous in heads or congested cymes, many or all of them
closely grouped together.
35a. Ovules and seeds borne on stipitate placentas near base of septum .......................... 45. Leptomischus
35b. Ovules and seeds borne on peltate placentas near middle of septum.
36a. Plants apparently without raphides; disk apparently glabrous ...................................... 40. Keenania
36b. Raphides present; disk puberulent ................................................................................. 55. Mouretia
32b. Fruit a capsule dehiscent through linear openings, or fleshy and indehiscent.
37a. Ovary and fruit 5-celled; plants often with fetid odor when bruised .......................... 87. Spermadictyon
37b. Ovary and fruit 1- or 2-celled; plants without or sometimes with fetid odor
when bruised.
38a. Fruit baccate, fleshy to dry, indehiscent (unknown in Keenania).
39a. Flowers solitary on paired axillary short shoots (i.e., brachyblasts); corollas
ca. 55 mm ...................................................................................................................... 1. Acranthera
39b. Flowers 2 to numerous on axillary or terminal peduncles; corollas 310 mm.

RUBIACEAE

63

40a. Corolla lobes convolute in bud; plants nearly to completely acaulescent ..... 32. Guihaiothamnus
40b. Corolla lobes valvate in bud; plants with stems (so far as known).
41a. Inflorescences cymose, branched; raphides present and evident ............................. 47. Lerchea
41b. Inflorescences capitate or congested-cymose, not or sparingly branched;
plants perhaps with raphides but these not readily evident .................................... 40. Keenania
38b. Fruit capsular, septicidal or loculicidal (unknown in Keenania).
42a. Flowers with 4 calyx and corolla lobes and stamens or some flowers occasionally
with 5.
43a. Plants without fetid odor when bruised, usually drying green or brownish green;
seeds angled to lenticular ............................................................................................. 35. Hedyotis
43b. Plants usually with fetid odor when bruised, usually drying black to gray; seeds
peltate to plano-convex ................................................................................................ 60. Neanotis
42b. Flowers with 5 calyx and corolla lobes and stamens.
44a. Inflorescences capitate or congested-cymose; ovary and fruit smooth ..................... 40. Keenania
44b. Inflorescences branched, cymose; ovary and fruit ridged to winged ...................... 88. Spiradiclis
Key 4
1a. Leaves with tertiary or quaternary venation finely, regularly lineolate (this visible best on abaxial surface);
inflorescences axillary; fruit a fleshy drupe.
2a. Fruit with 1 pyrene, this with 25 cells (and seeds); corollas 511 mm, with 4 or 5 lobes ........................................... 5. Antirhea
2b. Fruit with 50 or more pyrenes, each with 1 cell; corollas 1015 mm, with 6 lobes ................................................... 92. Timonius
1b. Leaves with tertiary and higher order venation not visible or irregularly to somewhat regularly areolate;
inflorescences axillary, terminal, or in other positions; fruit dry to fleshy, drupaceous, baccate, capsular,
schizocarpous, or other form.
3a. Plants mangroves, growing along seashores in areas regularly inundated by saltwater, with succulent tissues,
usually with stilt roots, usually with young growth covered with mucilage or resin ............................................ 83. Scyphiphora
3b. Plants of terrestrial or freshwater riverine habitats, with succulent to very thin tissues, without stilt roots,
with young growth covered with resin or not.
4a. Flowers fused together by their ovaries, at least shortly at their bases; fruit multiple (i.e., formed from
more than 1 flower).
5a. Stipules twisted or imbricate in a hemispherical to conical bud; fruit drupaceous, each with 2 or 4 pyrenes,
each pyrene with 1 seed .......................................................................................................................................... 54. Morinda
5b. Stipules held erect and pressed together in flattened bud; fruit baccate, each fruit with numerous seeds ............ 59. Nauclea
4b. Flowers free, sometimes tightly packed on a common supporting structure but not actually fused; fruit
free (i.e., each formed from only 1 flower).
6a. Flowers in 1 to several symmetrical, globose heads, heads sessile to pedunculate, terminal or axillary,
flowers sessile to pedicellate.
7a. Lianas with recurved spines ................................................................................................................................. 94. Uncaria
7b. Shrubs or trees, unarmed.
8a. Some or all leaves truncate to cordate at base.
9a. Inflorescences all axillary; stipules ovate to elliptic or spatulate, unlobed, held erect and pressed
together in a flattened bud ............................................................................................................................. 33. Haldina
9b. Inflorescences terminal and sometimes also in uppermost leaf axils; stipules ovate in outline and
deeply bilobed, imbricate in a conical bud ................................................................................................ 85. Sinoadina
8b. Leaves obtuse to acute at base.
10a. Fruit indehiscent; large trees, frequently cultivated for lumber ....................................................... 62. Neolamarckia
10b. Fruit schizocarpous splitting into 2 mericarps or capsular with septicidal then loculicidal
dehiscence; shrubs and trees, perhaps occasionally cultivated.
11a. Fruit schizocarpous, splitting into 2 indehiscent mericarps; leaves opposite or whorled;
shrubs .............................................................................................................................................. 13. Cephalanthus
11b. Fruit capsular, valves opening to release seeds; leaves opposite; shrubs and trees.
12a. Stipules held erect and pressed together in a flattened bud.
13a. Inflorescence heads sessile or subsessile with peduncles up to 0.5 cm; stigmas cylindrical
with style attached in a recessed portion (i.e., upside-down mitriform) ...................................... 53. Mitragyna
13b. Inflorescence heads pedunculate, peduncles 18 cm; stigmas globose to ovoid ...................... 63. Neonauclea
12b. Stipules imbricate or valvate in a conical to hemispherical bud.
14a. Seeds subglobose to trigonous or ellipsoid, not winged ................................................................ 49. Metadina
14b. Seeds flattened to fusiform or angled, usually winged.
15a. Calyx lobes ca. 0.5 mm; fruit 57 mm ....................................................................................... 85. Sinoadina

64

RUBIACEAE

15b. Calyx lobes 12 mm; fruit 1.55 mm.

16a. Stipules deeply bilobed for more than 1/2 their length ................................................................... 2. Adina
16b. Stipules triangular and entire to shortly bilobed for 1/4 or less their length ...................... 69. Pertusadina
6b. Flowers variously arranged in heads, cymes, panicles, or other types of groups, these groups
sometimes umbelliform but heads then not symmetrically globose, terminal, axillary, pseudoaxillary,
or in other positions.
17a. Fruit capsular, opening through an apical operculum and later sometimes splitting into 5 valves,
with each seed individually enclosed in a fibrous, netlike aril .................................................................. 44. Leptodermis
17b. Fruit indehiscent, baccate, drupaceous, or capsular but then not operculate and with fewer than
5 valves, seeds without aril or with aril continuous, fleshy.
18a. Inflorescences not borne at stem apex but regularly axillary (i.e., borne in both axils at a node),
superaxillary, and/or apparently axillary due to regularly produced axillary or lateral short
shoots (i.e., brachyblasts).
19a. Stems and abaxial surface of leaf densely sericeous, hirsute, or strigose with remarkably long
silky trichomes .................................................................................................................................. 97. Xanthophytum
19b. Stems and abaxial surface of leaf variously glabrous to pubescent with trichomes of various
types, but not densely and consistently long and silky.
20a. Inflorescence borne on regular paired axillary short shoots (i.e., brachyblasts), each short
shoot with several stipulate nodes; fruit fleshy and drupaceous or baccate, or dry and
indehiscent or schizocarpous.
21a. Stipules with 18 stiffly spiniform bristles ............................................................................................ 84. Serissa
21b. Stipules entire or with several to numerous bristles or projections but these not spiny.
22a. Bracts multifid with segments glandular.
23a. Corolla lobes valvate in bud; fruit a fleshy drupe .............................................................. 20. Damnacanthus
23b. Corolla lobes convolute in bud; fruit a dry schizocarp ........................................................ 93. Trailliaedoxa
22b. Bracts absent or entire and not glandular.
24a. Stipules bilobed .................................................................................................................. 72. Prismatomeris
24b. Stipules triangular.
25a. Flowers with 4 calyx lobes, corolla lobes, and stamens.
26a. Petioles and inflorescence axes articulate at their bases; stigmas fusiform ................................ 39. Ixora
26b. Petioles and inflorescence axes not articulate; stipules lobed.
27a. Plants without fetid odor; corolla salverform, with lobes convolute in bud ....................... 67. Pavetta
27b. Plants with or without fetid odor when bruised; corolla campanulate, tubular, or
funnelform, with lobes valvate or valvate-induplicate in bud ......................................... 81. Saprosma
25b. Flowers with 5 calyx lobes, corolla lobes, and stamens.
28a. Calyx lobes 58 mm ...................................................................................................... 12. Catunaregam
28b. Calyx lobes 0.54 mm.
29a. Ovules and seeds 14 in each cell; plants without spines; flowers sessile and
solitary ......................................................................................................................... 36. Himalrandia
29b. Ovules and seeds more than 4 in each cell; plants with or without spines;
flowers sessile to pedicellate, solitary to several in fascicles.
30a. Plants with bisexual flowers; ovary with ovules and seeds attached to septum
(i.e., axile) ........................................................................................................................... 8. Benkara
30b. Plants dioecious; ovary with ovules and seeds attached to outer wall
(i.e., parietal); nodes of stems mostly reduced .......................................................... 14. Ceriscoides
20b. Inflorescences borne on peduncles directly from main stems, these with no more than
1 reduced basal node.
31a. Inflorescences racemiform or spiciform, with well-developed primary axes directly
bearing numerous pedicels ....................................................................................................... 37. Hymenodictyon
31b. Inflorescences capitate to cymose, without well-developed primary axes or with axes
of several orders developed, with higher order axes or all axes bearing few to numerous
pedicels.
32a. Woody twiners or climbers; fruit a woody capsule, subglobose, with winged seeds ............. 19. Coptosapelta
32b. Erect shrubs and trees; fruit fleshy and indehiscent or capsular, papery to cartilaginous,
flattened, with unwinged seeds.
33a. Fruit capsular, papery to cartilaginous, mitriform and laterally markedly flattened
(this shape often evident in young developing fruit) ............................................................. 65. Ophiorrhiza
33b. Fruit indehiscent and fleshy.

RUBIACEAE

65

34a. Corolla lobes imbricate in bud; fruit a drupe with a single, 49-celled subglobose
pyrene, this 23 cm in diam.; plants of coastal thickets ......................................................... 31. Guettarda
34b. Corolla lobes convolute or valvate in bud or imbricate in some species of Lasianthus;
fruit baccate with numerous seeds or drupaceous with 1-celled pyrenes 2 to numerous;
plants of interior habitats.
35a. Cultivated plants; corolla lobes convolute in bud; fruit drupaceous with 2 plano-convex
pyrenes with papery walls; bracts fused in calyculate pairs ..................................................... 18. Coffea
35b. Native plants; corolla lobes convolute, valvate, or imbricate in bud; fruit baccate or
drupaceous with 2 to numerous plano-convex to angled, ellipsoid-oblong, ellipsoid,
or flattened, hard-walled pyrenes; bracts free or fused in calyculate pairs.
36a. Fruit a berry with several to numerous seeds.
37a. Seeds numerous; corolla lobes valvate in bud ........................................................... 95. Urophyllum
37b. Seeds 38; corolla lobes convolute in bud.
38a. Ovules and seeds 13 per cell; stigmas papillose; calyx lobes minute or
up to 0.5 mm ............................................................................................................ 23. Diplospora
38b. Ovules and seeds 36 per cell; stigmas pubescent; calyx lobes 12 mm ............ 38. Hyptianthera
36b. Fruit a drupe, with 2 to numerous pyrenes, each containing 1 seed.
39a. Flower buds acute to rounded at apex; pyrenes flattened to triangular; plants
often with fetid odor when bruised .............................................................................. 43. Lasianthus
39b. Flower buds sharply acute to acuminate at apex; pyrenes plano-convex to
ellipsoid-oblong or ellipsoid; plants without fetid odor.
40a. Plants without or sometimes with spines; stipules pilose to sericeous inside
(i.e., adaxially) ........................................................................................................... 11. Canthium
40b. Plants without spines; stipules glabrous inside ............................................................ 75. Psydrax
18b. Inflorescences terminal, pseudoaxillary (i.e., regularly borne in only 1 axil at a node), or apparently
pseudoaxillary or lateral due to anisophylly or an undeveloped internode.
41a. Stipules with 18 sharp spines or spiny bristles ........................................................................................... 84. Serissa
41b. Stipules entire to lobed or multifid, sometimes aristate or caudate but not spiny.
42a. Stipules with several well-developed, fibrous, closely set veins, these parallel to palmate .......... 58. Myrioneuron
42b. Stipules smooth, 13-costate but not densely veined.
43a. Calyx limb with developed truncate tube; flowers several to numerous, all pedicellate,
borne in umbelliform cymules .................................................................................................. 10. Caelospermum
43b. Calyx limb reduced to developed and at least shortly lobed or denticulate; flowers
variously arranged but not uniformly pedicellate in umbelliform cymules.
44a. Corolla lobes convolute in bud and fruit a berry with fleshy to leathery outer wall.
45a. Fruit and usually ovary portion of calyx 58-ridged or -winged, ridges continuous with
midrib of each calyx lobe ............................................................................................................ 29. Gardenia
45b. Fruit and ovary portion of calyx smooth to tuberculate.
46a. Some or all inflorescences regularly borne in pseudoaxillary, leaf-opposed, and/or
lateral position with only 1 inflorescence per node (i.e., not paired, borne on only
one side of stem), this arrangement due to anisophylly or a reduced internode
producing apparently alternate leaves or a 3-leaved node.
47a. Inflorescences all apparently leaf-opposed or borne at leafless nodes (due to
extreme anisophylly), stems with paired or apparently alternate leaves ...................................... 3. Aidia
47b. Inflorescences mostly borne on apparently 3-leaved nodes, stems with some
3-leaved nodes.
48a. Flowers 12 or more per inflorescence ............................................................................... 25. Duperrea
48b. Flowers 110 per inflorescence.
49a. Some or all leaf apices with short filamentous arista or projection ..................... 80. Rubovietnamia
49b. Leaf apices without filamentous projections.
50a. Plants dioecious, with unisexual flowers; fruit 515 mm in diam.; corollas
(so far as known) 58 mm ........................................................................................ 9. Brachytome
50b. Plants with bisexual flowers; fruit ca. 35 mm in diam.; corollas 6875 mm ....... 78. Rothmannia
46b. Inflorescence borne at stem apices and sometimes in adjacent uppermost leaf axils,
sometimes displaced to pseudoaxillary by subsequent overtopping stem growth,
produced at stems generally with number and arrangement of leaves similar at
all nodes.
51a. Low shrubs, up to 0.5 m tall, with many internodes reduced and relatively
small leaves.

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RUBIACEAE

52a. Flowers solitary ........................................................................................................... 36. Himalrandia

52b. Flowers in cymes of 612 .......................................................................................... 93. Trailliaedoxa
51b. Shrubs, low (ca. 0.3 m tall) to tall, or trees.
53a. Corolla densely sericeous (i.e., with long, silky, appressed trichomes) on outside .... 71. Porterandia
53b. Corolla glabrous to puberulent, spreading pubescent, or strigillose (i.e., with
short appressed trichomes) on outside.
54a. Inflorescences with 17 flowers; fruit 411 cm ........................................................... 27. Fosbergia
54b. Inflorescences with 5 to numerous flowers; fruit 2 cm or shorter.
55a. Petioles and inflorescence axes articulate; ovules and seeds 1 per cell, 2 per
ovary or fruit; stigmas unlobed .......................................................................................... 39. Ixora
55b. Petioles and inflorescence axes not articulate; ovules and seeds 1 to numerous
per cell, 2 to numerous per ovary or fruit; stigmas lobed.
56a. Bark of older stems becoming coppery red and circumferentially wrinkled
to peeling ............................................................................................................ 90. Tarennoidea
56b. Bark of older stems smooth, persistent, green to gray or brown, sometimes
becoming longitudinally striate or fissured with age.
57a. Ovules 1 per cell and 2 per ovary; fruit drupaceous with 2 plano-convex
pyrenes ..................................................................................................................... 67. Pavetta
57b. Ovules 2 or more per cell and 4 or more per ovary; fruit baccate with seeds
2 or more, ellipsoid, obovoid, or compressed globose.
58a. Seeds 26 per fruit, 45 mm in diam., compressed globose, fruit yellowish
white; plants without spines ........................................................................... 4. Alleizettella
58b. Seeds 4 to numerous per fruit, 35 mm in diam., compressed globose to
ellipsoid or obovoid, fruit purplish black, black, or sometimes white;
plants with or without spines.
59a. Plants without or usually with spines; calyx limb persistent or deciduous
on fruit ............................................................................................................... 8. Benkara
59b. Plants without spines; calyx limb deciduous on fruit ..................................... 89. Tarenna
44b. Corolla lobes valvate, valvate-induplicate, imbricate, or convolute in bud and fruit a drupe,
berry, capsule, or schizocarp, if corolla lobes convolute then fruit not a berry.
60a. Stigmas, cells of ovary, and cells of fruit 4 or 5.
61a. Corollas tubular with relatively short lobes, these 10% or less of total corolla length;
plants cultivated ornamentals .................................................................................................... 34. Hamelia
61b. Corollas funnelform, tubular-funnelform, or salverform with lobes developed, 15% or
more of total corolla length; plants native or cultivated.
62a. Fruit dry, schizocarpous or capsular, leathery; plants often with fetid odor when
bruised .......................................................................................................................... 87. Spermadictyon
62b. Fruit fleshy and drupaceous or baccate, or capsular and thinly textured; plants
without fetid odor.
63a. Plants not particularly succulent, sometimes with spines, usually with some
internodes not expanded; calyx without stalked glands .......................................... 20. Damnacanthus
63b. Plants sometimes succulent, without spines, with all internodes expanded; calyx
sometimes with stalked glands ............................................................................................ 57. Mycetia
60b. Stigmas 1 or 2 and cells of ovary and fruit 2, or sometimes any of these 3 on some
but not all flowers in an inflorescence.
64a. Cultivated ornamental plants; herbs and low shrubs to small trees; corollas red, orange,
or yellow.
65a. Stigmas unlobed; corolla lobes convolute in bud; petioles and inflorescence axes
articulate ....................................................................................................................................... 39. Ixora
65b. Stigmas 2- or 3-lobed; corolla lobes valvate, valvate-induplicate, or imbricate in bud;
petioles and inflorescence axes not articulate.
66a. Plants often with petaloid calycophylls; corolla lobes valvate-induplicate in bud ....... 56. Mussaenda
66b. Plants without petaloid calycophylls; corolla lobes valvate or imbricate in bud.
67a. Herbs and low shrubs; corolla with white pubescence in throat, lobes obtuse
to acute, with margins flat .................................................................................................. 68. Pentas
67b. Shrubs; corolla with yellow pubescence in throat, lobes broadly obtuse to
rounded, with margins flat to crisped .......................................................................... 77. Rondeletia
64b. Native plants; low shrubs to large trees; corollas white, pink, purple, yellow, or red.

RUBIACEAE

67

68a. Fruit indehiscent, drupaceous, or baccate, fleshy or leathery; corolla lobes convolute,
imbricate, or valvate in bud; ovules and seeds 1 to numerous per cell, 2 to numerous
per fruit.
69a. Stigmas unlobed; petioles and inflorescence axes articulate at base ....................................... 39. Ixora
69b. Stigmas 2- or 3-lobed; petioles and inflorescence axes not articulate.
70a. Fruit baccate, with numerous seeds in each cell, these enclosed in soft tissues.
71a. Corolla lobes valvate-reduplicate; calyx sometimes with calycophylls,
without stalked glands; seeds flattened; plants not succulent ................................. 56. Mussaenda
71b. Corolla lobes valvate-induplicate; calyx without calycophylls, sometimes
with stalked glands; seeds angled; plants often succulent ........................................... 57. Mycetia
70b. Fruit indehiscent or drupaceous, with 1 seed in each cell, this enclosed in a pyrene.
72a. Stipules caducous or rather quickly deciduous, often exposing a ring of
persistent trichomes that usually dry red-brown (use magnification) ...................... 74. Psychotria
72b. Stipules persistent to caducous, not enclosing persistent trichomes or these
reduced and/or drying white.
73a. Corolla lobes convolute in bud.
74a. Fruit drupaceous, with pyrenes flattened to concavo-convex ................................ 67. Pavetta
74b. Fruit schizocarpous with segments or valves oblanceoloid ......................... 93. Trailliaedoxa
73b. Corolla lobes valvate in bud.
75a. Corolla tubular-funnelform with base usually gibbous and tube usually
curved .................................................................................................................. 15. Chassalia
75b. Corolla tubular-funnelform or salverform, generally straight at base and
in tube.
76a. Stipules persistent and becoming hardened and yellowed with age, often
fragmenting when older .................................................................................. 74. Psychotria
76b. Stipules caducous, or persistent but then not becoming hardened,
yellowed, or fragmented.
77a. Plants often with fetid odor when bruised; corolla lobes mostly 4, or
on a few flowers of some plants 5 or 6 ........................................................ 81. Saprosma
77b. Plants without fetid odor; corolla lobes regularly 5, on a few flowers
of some plants 4.
78a. Bracts glandular-multifid .................................................................. 20. Damnacanthus
78b. Bracts entire, not glandular ............................................................... 72. Prismatomeris
68b. Fruit dry and capsular (i.e., dehiscent), indehiscent, or schizocarpous (i.e., splitting
into sections); corolla lobes valvate, valvate-induplicate, or imbricate in bud; ovules
and seeds several to numerous in each cell and fruit, or 1 per cell and 2 per fruit in
Trailliaedoxa.
79a. Inflorescences racemiform or spiciform, with 1 or a few well-developed main
axes bearing pedicels ............................................................................................... 37. Hymenodictyon
79b. Inflorescences subcapitate to cymose, with axes of various orders developed
to different degrees and flowers usually borne from axes of various orders.
80a. Fruit mitriform and laterally markedly flattened (this shape often evident in
young developing fruit) ............................................................................................. 65. Ophiorrhiza
80b. Fruit ellipsoid to subglobose.
81a. Fruit capsular, septicidal, 850 mm with at least some fruit 10 mm or
longer; seeds flattened and often winged; corollas 570 mm.
82a. Stipules persistent; shrubs, often epiphytic ............................................... 61. Neohymenopogon
82b. Stipules caducous; terrestrial shrubs or trees.
83a. Corolla lobes valvate in bud, acute; cultivated trees, without petaloid
calycophylls ......................................................................................................... 16. Cinchona
83b. Corolla lobes imbricate in bud, obtuse to rounded; native trees perhaps
sometimes cultivated, sometimes with petaloid calycophylls.
84a. Calyx lobes 22.5 mm; corolla lobes ca. 7 mm; plants sometimes
with calycophylls ..................................................................................... 26. Emmenopterys
84b. Calyx lobes 820 mm; corolla lobes 1118 mm; plants without
calycophylls ......................................................................................................... 48. Luculia
81b. Fruit indehiscent, schizocarpous, or capsular, septicidal or loculicidal,
18 mm; seeds flattened and sometimes winged to angled and unwinged;
corollas 220 mm.

68

RUBIACEAE

85a. Corolla lobes imbricate in bud; fruit capsular, loculicidal, subglobose,

woody .................................................................................................................. 96. Wendlandia
85b. Corolla lobes valvate, valvate-induplicate, or convolute in bud; fruit
indehiscent, schizocarpous, or capsular and septicidal or loculicidal,
subglobose to ellipsoid, papery to woody, if loculicidal then corolla
lobes valvate.
86a. Corolla lobes convolute in bud; low shrubs; ovules and seeds 1 per cell .... 93. Trailliaedoxa
86b. Corolla lobes valvate or valvate-induplicate in bud; low to large shrubs;
ovules and seeds several to numerous in each cell.
87a. Corolla lobes valvate-induplicate in bud; plants often with petaloid
calycophylls ......................................................................................... 82. Schizomussaenda
87b. Corolla lobes [simple-]valvate in bud; plants with or without
petaloid calycophylls.
88a. Inflorescences usually with 1 or several petaloid bracts and/or
calycophylls; corolla yellow; capsules septicidal; stipules
interpetiolar, persistent ..................................................................................... 24. Dunnia
88b. Inflorescences without petaloid bracts or calycophylls; corolla white,
pink, blue, purple, or yellow; capsules septicidal or loculicidal or fruit
indehiscent; stipules interpetiolar or fused to petiole bases, persistent
or caducous.
89a. Fruit dehiscent through apical part or splitting into 2 indehiscent
valves ........................................................................................................... 35. Hedyotis
89b. Fruit deeply dehiscent into 2 or 4 separate valves ................................... 88. Spiradiclis

1. ACRANTHERA Arnott ex Meisner, Pl. Vasc. Gen. 1: 162; 2: 115. 1838, nom. cons.
jian yao hua shu
Chen Tao (); Charlotte M. Taylor, Christian Puff
Herbs or subshrubs, unbranched or little branched, unarmed. Raphides absent. Leaves opposite and often clustered at ends of
stems, without domatia; stipules persistent or deciduous, interpetiolar, triangular to oblong. Inflorescences terminal or sometimes
displaced to pseudoaxillary, borne on principal stems or sometimes short shoots at stem apex or in 1 or both leaf axils, 1-flowered [or
cymose to fasciculate and several flowered], bracteate [or bracts reduced]. Flowers pedicellate or subsessile, bisexual, apparently
monomorphic. Calyx with ovary portion usually relatively prolonged; limb lobed essentially to base; lobes 4 or 5, often with 1 welldeveloped colleter in each sinus. Corolla pale green or white to blue or purple, funnelform to campanulate, glabrous inside; lobes 4 or
5, valvate or reduplicate-valvate in bud. Stamens 5, inserted near base of corolla tube, included or exserted; filaments short to well
developed, glabrous, free or fused in basal portion; anthers linear, at apex with sharply acute or spurred appendage, these connate into
a tube surrounding stigma and united with it at tops of both structures. Ovary 1-celled, ovules many per cell on 2 T-shaped parietal
placentas; stigma 1, clavate, relatively large, 10-ridged, sometimes fused at top to anther appendages. Fruit baccate or perhaps
occasionally tardily capsular with irregular dehiscence, fleshy, ovoid to cylindrical or turbinate, color not reported, with calyx limb
persistent; seeds numerous, reddish brown or nearly black, small, compressed or lenticular; endosperm fleshy; embryo small,
straight.
About 40 species: China, India, Indonesia, Malaysia, Sri Lanka, Thailand, Vietnam, with most species apparently in Borneo; one species
(endemic) in China.
This genus was monographed by Bremekamp (J. Arnold Arbor. 28: 261308. 1947), who named a number of subgenera and series. C. Y. Wus
new species did not fit into Bremekamps classification; consequently, he described a new monotypic subgenus for it, Acranthera subg. Sinacranthera
C. Y. Wu, distinguished from A. subg. Acranthera and A. subg. Androtropis Bremekamp by its funnelform corollas, inflorescences borne on opposite
brachyblasts bearing two rudimentary leaves, and red to purple corolla color (vs. basal part of corolla cylindrical, inflorescences borne at stem apices,
and blue corollas in A. subg. Acranthera and inflorescences borne at stem apices and pale green corollas in A. subg. Androtropis) and from the other
subgenera by its well-developed, subglobose disk (vs. inconspicuous and presumably flattened). Acranthera is considered based on recent molecular
and some morphological data to be a rather isolated basal lineage in Rubiaceae: its closest relative is probably Coptosapelta, and these two genera are
probably most closely related to Luculia (Alejandro et al., Amer. J. Bot. 92: 544557. 2005; Rydin et al., Pl. Syst. Evol. 278: 101123. 2009).
Puff et al. (Ann. Missouri Bot. Gard. 82: 357382. 1995) concluded that the flowers have an anther-style and stigma complex that is unique in
Rubiaceae and suggested that at least some Acranthera species may be buzz-pollinated. Bremekamp described the ovaries as 2-celled with laminar
axile placentas, but Puff et al. showed that the ovaries are actually 1-celled, with parietally inserted placentas that are T-shaped and meet closely
enough in the middle to appear superficially to be borne on a median septum, though there is none.

RUBIACEAE

1. Acranthera sinensis C. Y. Wu, Acta Phytotax. Sin. 6: 295.

1957.
zhong hua jian yao hua
Herbs or subshrubs, 40(100) cm tall, with tissues often
turning black when dry; branches generally quadrangular,
densely hirsute to strigose and strigillose becoming glabrescent
with age. Petiole 17 cm, strigose to strigillose; leaf blade thinly
papery to membranous, elliptic or obovate, 822 4.59 cm,
adaxially sparsely to moderately hirsute to hispid, abaxially
strigillose to strigose or hispid with pubescence denser along
principal veins, base acute or cuneate, margins ciliolate, apex
acuminate or acute; secondary veins 911 pairs; stipules persistent, broadly ovate to triangular, 13 mm, glabrescent, cuspidate or 2- or 3-dentate. Inflorescences terminal, pseudoaxillary, and/or axillary on short shoots, 1-flowered, short shoot
reduced to developed (appearing as base of articulate pedun-

69

cle), to 2 mm; peduncle (i.e., directly subtending flower) 34

cm; bracts lanceolate, 23 mm, acute. Calyx strigillose to strigose or pilose; ovary portion cylindrical to obconical, 34 cm;
lobes linear-lanceolate, 2.74 cm. Corolla purple outside and
pink inside, funnelform, outside hirtellous to tomentulose;
tube ca. 45 mm; lobes ovate, ca. 10 mm, obtuse then abruptly
acuminate with tip ca. 1.5 mm. Filaments free; anthers 68 mm,
with connective spurred at apex. Berry compressed cylindrical, 4045 ca. 5 mm, bisulcate; seeds with testa reticulate. Fl.
AprJun, fr. JunOct.
Forests on mountain slopes; 10001500 m. SE Yunnan.
Wu in the protologue explicitly designated two syntypes, H. T.
Tsai 55229 collected on 19 May 1934 as the typus florifer and H. T.
Tsai 60568 collected on 29 Jun 1934 as the typus fructifer; his species
was published in 1957, so this name is validly published in spite of the
lack of a single holotype.

2. ADINA Salisbury, Parad. Lond. t. 115 [116]. 1808.

shui tuan hua shu
Chen Tao (); Charlotte M. Taylor
Shrubs or small trees, unarmed, often deciduous; buds conical. Raphides absent. Leaves opposite, distichous or decussate, usually with domatia; stipules persistent or deciduous, interpetiolar, bifid, nodes with a ring of persistent trichomes exposed after stipules
fall. Inflorescences terminal and/or in axils of uppermost leaves, capitate with 1[to several] globose heads, pedunculate with peduncles usually articulate in basal half and bearing stipuliform bracts at middle, many flowered, bracteate; bracteoles linear to linearspatulate. Flowers sessile, bisexual, monomorphic. Calyx limb 5-lobed. Corolla white, salverform to slenderly funnelform, glabrous
inside; lobes 5, valvate in bud. Stamens 5, inserted in upper part of corolla tube, partially to fully exserted; filaments short, glabrous;
anthers basifixed. Ovary 2-celled, ovules many in each cell, pendulous on axile placentas attached to upper 1/3 of septum; stigma
globose to obovoid, exserted. Fruiting heads globose, at maturity shattering easily. Fruit capsular, obconic, septicidally then loculicidally dehiscent into 4 valves separating from base to apex from each other and from septum, cartilaginous to papery, with septum
persistent or tardily deciduous, with calyx limb persistent on septum; seeds numerous, small, fusiform to oblanceoloid, flattened to
rounded, with embryo portion obovoid, ellipsoid, oblong, or trigonous, sometimes with short wing at one or both ends.
Four species: China, Japan, Korea, Thailand, Vietnam; three species in China.

1a. Deciduous low shrubs; leaves distichous, subsessile to shortly petiolate, petioles to 2 mm, leaf blade
2.54 0.81.2 cm; flowering heads borne separately on axillary or sometimes apically paired peduncles ................ 3. A. rubella
1b. Evergreen shrubs or small trees; leaves decussate, petiolate, petioles 212 mm, leaf blade 418 1.56 cm;
flowering heads borne separately on axillary or sometimes apically paired peduncles and/or infrequently or
often in cymes of 3 or 5.
2a. Leaf blade 412 1.53 cm, abaxially glabrous or sparsely puberulent on costa, petioles 26 mm; stipules
persistent to caducous, 38 mm, puberulent to glabrous, lobes lanceolate to ovate; flowering heads
infrequently in cymes of 3 ...................................................................................................................................... 1. A. pilulifera
2b. Leaf blade 618 2.36 cm, abaxially sparsely to densely puberulent on veins to throughout, petioles
312 mm; stipules caducous, 45 mm, glabrous, lobes narrowly triangular; flowering heads often in
cymes of 3 or 5 .................................................................................................................................................... 2. A. pubicostata
1. Adina pilulifera (Lamarck) Franchet ex Drake, J. Bot.
(Morot) 9: 207. 1895.
shui tuan hua
Cephalanthus pilulifer Lamarck, Encycl. 1: 679. 1785;
Adina globiflora Salisbury; A. globiflora var. tonkinensis Pitard;
A. pilulifera var. tonkinensis (Pitard) Merrill ex H. L. Li.
Shrubs or small trees, evergreen, 15(10) m tall; branches
angled to terete, puberulent to glabrescent, often lenticellate,
with bark usually gray. Leaves decussate; petiole 26 mm, gla-

brous or puberulent; blade drying papery to stiffly papery, narrowly elliptic, elliptic-lanceolate, obovate-oblong, oblanceolate,
or obovate-oblanceolate, 412 1.53 cm, adaxially glabrous,
abaxially glabrous or sparsely puberulent on costa, base acute to
cuneate or obtuse, apex acute to acuminate with tip usually ultimately blunt; secondary veins 612 pairs, usually with foveolate and/or pilosulous domatia; stipules persistent to caducous,
in outline ovate, 38 mm, puberulent to glabrous, deeply bifid, lobes lanceolate to ovate, acute to acuminate, erect to
spreading. Inflorescences puberulent to glabrous; peduncles 25
cm, with bracts ca. 2 mm; flowering heads borne separately on

RUBIACEAE

70

axillary or sometimes apically paired peduncles and/or infrequently in cymes of 3, 46 mm in diam. across calyces, 812
mm in diam. across corollas; bracteoles linear to linear-clavate,
1.31.8 mm. Calyx puberulent to glabrescent; ovary portion
obconic, 0.51 mm, surrounded at base by a strigose ring; limb
deeply lobed, lobes linear-oblong or spatulate, 11.8 mm, with
apex thickened. Corolla white, narrowly funnelform, outside
puberulent or hirtellous to glabrous; tube 23.5 mm; lobes
ovate-oblong, 0.51 mm, obtuse. Stigma globose to obovoid,
ca. 0.2 mm, exserted for 35 mm. Fruiting heads 711 mm in
diam. Capsules obcuneate, 25 mm, puberulent to glabrescent;
seeds ca. 2 mm. Fl. JunSep, fr. JulDec.
Thickets or sparse forests at streamsides, at roadsides, or in valleys; 200400 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Jiangsu, Jiangxi, Yunnan, Zhejiang [Japan, Vietnam].
H. L. Li (J. Arnold Arbor. 25: 317. 1944) recognized Adina
pilulifera var. tonkinensis for Chinese plants from Guangxi, Hainan, and
Yunnan but did not there explain their separation from the typical variety; no subsequent authors, including H. H. Hsue and H. Wu (in FRPS
71(1): 274275. 1999), have recognized infraspecific taxa within this
widespread species.

2. Adina pubicostata Merrill, J. Arnold Arbor. 21: 385. 1940.

mao mai shui tuan hua
Shrubs or small trees, evergreen, 18 m tall; branches
angled to terete, puberulent to glabrescent, often lenticellate,
with bark usually grayish to reddish brown. Leaves decussate;
petiole 312 mm, glabrous to densely puberulent; blade drying
papery to stiffly papery, narrowly elliptic, elliptic-oblong, lanceolate, or oblanceolate, 618 2.36 cm, adaxially glabrous,
abaxially sparsely to densely puberulent on veins to throughout,
base acute to cuneate or obtuse, apex acute to acuminate with
tip usually ultimately blunt; secondary veins 612 pairs, usually
with foveolate and/or pilosulous domatia; stipules caducous, in
outline ovate, 45 mm, glabrous, deeply bifid, lobes narrowly
triangular, acute, erect to spreading. Inflorescences densely puberulent; peduncles 25 cm, with bracts ca. 2 mm; flowering
heads borne separately on axillary or sometimes apically paired
peduncles and/or often in cymes of 3 or 5, 6.57 mm in diam.
across calyces, 1011 mm in diam. across corollas; bracteoles
linear to linear-clavate, 12 mm. Calyx puberulent to glabrescent; ovary portion obconic, 0.51 mm, surrounded at base by a
pilosulous ring; limb deeply lobed; lobes narrowly spatulate, ca.
1 mm, with apex cucullate to thickened. Corolla white, narrowly funnelform, outside puberulent to glabrous; tube 23
mm; lobes ovate, 11.5 mm, subacute. Stigma globose to obovoid, ca. 0.2 mm, exserted for 35 mm. Fruiting heads 89 mm
in diam. Capsules obcuneate, 22.5 mm, pilosulous at apex to

glabrescent in lower portions, with persistent calyx lobes (on

persistent septum) 1.22 mm; seeds 1.22 mm. Fl. Jun, fr. Jul
Nov or rarely into Jan.
Forests; 4001200 m. Guangxi, Hunan [Vietnam].
This species has previously been included in Adina pilulifera
(Ridsdale, Blumea 24: 357358. 1979; H. H. Hsue & H. Wu, FRPS
71(1): 275. 1999); however, with more material now available it can be
clearly distinguished from that species.

3. Adina rubella Hance, J. Bot. 6: 114. 1868.

xi ye shui tuan hua
Low shrubs, deciduous, 13(6) m tall; branches angled
to terete, puberulent to tomentulose or glabrescent, sometimes
lenticellate, with bark usually reddish brown. Leaves distichous,
subsessile to shortly petiolate; petiole to 2 mm; blade drying
papery, lanceolate, ovate-lanceolate, lanceolate-elliptic, or
ovate-oblong, 2.54 0.81.2 cm, adaxially puberulent to glabrous, abaxially glabrescent to puberulent or sparsely pilosulous
at least along veins, base obtuse to rounded, apex acute or
acuminate; secondary veins 57 pairs, with pilosulous domatia;
stipules persistent to deciduous, ligulate to ovate in outline, 25
mm, puberulent to glabrous, deeply bilobed, lobes narrowly triangular, acute to acuminate, erect to reflexed. Inflorescences
densely puberulent or strigillose to glabrescent; peduncles 15
cm, with bracts 24 mm; flowering heads borne separately on
axillary or sometimes apically paired peduncles, 47 mm in
diam. across calyces, 1012 mm in diam. across corollas; bracteoles linear or linear-clavate, 12 mm. Calyx puberulent to glabrescent; ovary portion obconic, 0.81 mm; limb deeply
lobed, lobes spatulate or spatulate-clavate, 12 mm, at apex
thickened, rounded, papillose. Corolla white or flushed with
pink or purple on lobes, slenderly funnelform, outside glabrescent, puberulent, or hirtellous with trichomes often purple-red;
tube 23 mm; lobes deltoid to ligulate, 0.51 mm, acute to
obtuse. Stigma obovoid, 0.20.3 mm, exserted for 34 mm.
Fruiting heads 812 mm in diam. Capsules oblong-obcuneate,
34 mm, puberulent, hirtellous, or glabrescent; seeds ca. 1.5
mm. Fl. and fr. MayDec.
Wet sites at streamsides, riversides, and sand beaches; below 100
600 m. Fujian, Guangdong, Guangxi, Hunan, Jiangsu, Jiangxi, Shaanxi,
Zhejiang [Korea].
This is a widespread and frequently collected species. The stipules
are sometimes completely split, with the lobes falling separately, and
can be confused with triangular entire stipules if care is not taken. How
(Sunyatsenia 6: 249. 1946) noted that this species is effective for stabilization of slopes.

3. AIDIA Loureiro, Fl. Cochinch. 1: 143. 1790.

qian shu shu
Chen Tao (); Charlotte M. Taylor
Shrubs or trees [or rarely vines], unarmed. Raphides absent. Leaves opposite, isophyllous or at nodes with inflorescences often
markedly anisophyllous with smaller leaf reduced to a scale and/or deciduous to give an appearance of alternate leaf arrangement,
usually with domatia; stipules persistent or usually caducous, interpetiolar or shortly united around stem, triangular. Inflorescences
pseudoaxillary, usually produced at nodes with markedly anisophyllous leaves thus appearing leaf-opposed, fasciculate to usually

RUBIACEAE

71

cymose, few to many flowered, sessile to pedunculate, bracteate. Flowers sessile or pedicellate, bisexual, monomorphic. Calyx limb
cupular or campanulate, 4- or 5-lobed or -denticulate. Corolla white, yellow, or green sometimes flushed with pink or red, salverform, pilosulous to villosulous in throat; lobes 4 or 5, convolute in bud, usually strongly reflexed at anthesis. Stamens 4 or 5,
inserted in corolla throat, exserted; filaments very short; anthers dorsifixed, exserted. Ovary 2(or 3)-celled, ovules several to many in
each cell on axile placentas; stigma clavate, fusiform, or 2-lobed with lobes sometimes coherent or free. Fruit red to orange, baccate,
fleshy, globose and smooth or ridged, with calyx limb deciduous; seeds several to numerous, medium-sized, angled to compressed,
embedded in pulp.
About 50 species: tropical Africa, S and SE Asia, Oceania; eight species (one endemic) in China.
The Asian and Malesian portion of this genus was reviewed in detail by Ridsdale (Blumea 41: 135179. 1996), who recognized five sections; all
of our species fall in his Aidia sect. Aidia. An additional species, A. shweliensis, was included in Aidia by W. C. Chen (in FRPS 71(1): 350. 1999) but
is here treated as Fosbergia shweliensis. The morphology of Aidia was studied in some detail by Tirvengadum and Sastre (Bull. Mus. Natl. Hist. Nat.,
B, Adansonia 8: 257296. 1986), who also provided an overview of the taxonomy. Ridsdale (Reinwardtia 12: 289. 2008) treated A. canthioides in the
genus Benkara, but it seems anomalous in that genus and is here retained in Aidia. The species circumscriptions of Ridsdale differ from those of W. C.
Chen (loc. cit.: 348356); neither author is completely followed here. Following Ridsdale, here plants with 4-merous flowers are distinguished from
those with 5-merous flowers, which results in the separation of A. racemosa and A. cochinchinensis, both 5-merous representatives of primarily
Malesian and Vietnamese species, from the 4-merous, primarily Chinese species A. henryi. Following W. C. Chen, A. henryi and A. merrillii are here
considered conspecific. Following Ridsdale, A. densiflora (Wallich) Masamune is not treated here as a species found in China, and the names Merrill
(Lingnan Sci. J. 14: 61. 1935) considered synonyms of that species (Randia densiflora (Wallich) Bentham, R. oppositifolia Koorders, and Webera
oppositifolia Roxburgh) are here considered misapplied names that do not correspond to any species in the Chinese flora.

1a. Young branches, leaf blade abaxially, and inflorescences densely hirtellous, pilosulous, or tomentulose, with
trichomes spreading ..................................................................................................................................................... 5. A. pycnantha
1b. Young branches, leaf blade abaxially, and inflorescences glabrous, puberulent, or strigillose, with trichomes
mostly appressed or leaves sometimes pilose below on principal veins in A. salicifolia.
2a. Inflorescences fasciculate, or subsessile and congested-cymose with axes short and often monochasial,
with bracts nearly as long as internodes of axes, and with pedicels much longer than axes (i.e., at first
glance appearing fasciculate or fungoid).
3a. Pedicels 517 mm; calyx limb 35 mm; corolla with tube 89 mm, lobes 47 mm and shorter
than tube ....................................................................................................................................................... 1. A. canthioides
3b. Pedicels 14 mm; calyx limb 12.5 mm (unknown in A. salicifolia); corolla with tube ca. 3 mm,
lobes 45 mm and longer than tube (corolla unknown in A. salicifolia).
4a. Leaf blade narrowly lanceolate to narrowly elliptic, 8.523 0.53 cm, with secondary veins
912 pairs; Guangxi ................................................................................................................................. 7. A. salicifolia
4b. Leaf blade elliptic-lanceolate or oblong-lanceolate, 6.217.5 2.76.5 cm, with secondary
veins 710 pairs; Yunnan ...................................................................................................................... 8. A. yunnanensis
2b. Inflorescences cymose, subsessile to pedunculate, branched to several orders with axes sometimes
dichasial or monochasial, with internodes of axes exceeding bracts, with pedicels absent or shorter
than peduncle plus branched portion of inflorescence.
5a. Calyx limb 55.5 mm (including lobes); corolla lobes 910 mm ................................................................ 4. A. oxyodonta
5b. Calyx limb 12.5 mm; corolla lobes 58 mm.
6a. Calyx lobes 4; corolla lobes 4; stipules 610 mm; widespread in China ...................................................... 3. A. henryi
6b. Calyx lobes 5; corolla lobes 5; stipules 35 mm; Hainan (A. cochinchinensis also in Yunnan).
7a. Inflorescence axes dichasial .................................................................................................... 2. A. cochinchinensis
7b. Inflorescence axes with branching dichasial at basal nodes but markedly monochasial at
distal nodes ......................................................................................................................................... 6. A. racemosa
1. Aidia canthioides (Champion ex Bentham) Masamune,
Trans. Nat. Hist. Soc. Formosa 28: 118. 1938.
xiang nan
Randia canthioides Champion ex Bentham, Hookers J.
Bot. Kew Gard. Misc. 4: 194. 1852; Aidia canthioides var.
lanceolata Masamune; Benkara canthioides (Champion ex
Bentham) Ridsdale; Fagerlindia canthioides (Champion ex
Bentham) Ridsdale.
Shrubs or trees, 112 m tall; branches flattened to subterete, glabrous. Petiole 518 mm, glabrous; leaf blade drying papery or thinly leathery, oblong-elliptic, oblong-lanceolate, or
lanceolate, 4.518.5 28 cm, both surfaces glabrous, base

cuneate to obtuse or shortly rounded, sometimes inequilateral,

apex acute to acuminate; secondary veins 37 pairs, in abaxial
axils usually with foveolate and/or pilosulous domatia; stipules
deciduous or sometimes persistent on distalmost nodes, broadly
triangular, 38 mm, glabrous, apex acute or acuminate. Inflorescences fasciculate or shortly congested-monochasial and
subsessile, 23 35 cm, several flowered, glabrescent; bracts
ovate, often fused in pairs, 0.51 mm, acute to obtuse; pedicels
517 mm. Calyx densely to sparsely strigillose; ovary portion
obconic, 11.5 mm; limb with basal tubular portion 35 mm;
lobes 5, triangular to deltoid, 0.52 mm, acute. Corolla white or
yellowish white, glabrous outside; tube 89 mm; lobes 5, narrowly spatulate-oblong, 47 mm, acute. Berry 58 mm in diam.,

RUBIACEAE

72

sparsely strigillose or glabrous; seeds 6 or 7, flattened, angled,

23 mm. Fl. AprJun, fr. MayFeb.
Thickets or forests on hills, on mountain slopes, or at streamsides
in valleys; below 1001500 m. Fujian, Guangdong, Guangxi, Hainan,
Taiwan, Yunnan [Japan, Vietnam].
This species was excluded from Aidia and treated as Fagerlindia
canthioides by Ridsdale (Blumea 41: 176. 1996), then as Benkara canthioides by Ridsdale (Reinwardtia 12: 289. 2008). This species is here
included in Aidia because of its apparent lack of lateral short shoots or
spines, inflorescences not terminal on developed stems, and apparently
bisexual flowers.

Thickets or forests at streamsides, on hills, or on mountain slopes;

below 1002400 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan,
Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang
[Japan, Thailand, Vietnam].
As noted above, this species is here circumscribed to generally
comprise the plants treated by W. C. Chen (in FRPS 71(1): 354. 1999)
as Aidia cochinchinensis. The reports of A. cochinchinensis (as Randia
cochinchinensis) from Zhejiang by Qiu and Zhong (Fl. Zhejiang 6: 103.
1986) and from Fujian (Fl. Fujian. 5: 165. 1993) are here treated as
reports of A. henryi.

2. Aidia cochinchinensis Loureiro, Fl. Cochinch. 1: 143. 1790.

4. Aidia oxyodonta (Drake) T. Yamazaki, J. Jap. Bot. 45: 339.

1970.

qian shu

jian e qian shu

Randia cochinchinensis (Loureiro) Merrill.

Shrubs or trees, 215 m tall; branches somewhat flattened
to terete, glabrous. Petiole 510 mm, glabrous; leaf blade
drying leathery or papery, elliptic to lanceolate, 915 35 cm,
both surfaces glabrous, base acute to obtuse, apex acute to acuminate; secondary veins 57 pairs, in abaxial axils usually with
pilosulous and/or foveolate domatia; stipules deciduous after
distalmost 2 or 3 nodes, lanceolate to narrowly triangular, 35
mm, glabrous, apex acuminate. Inflorescences cymose, 26 cm,
with axes usually regularly dichasial, glabrous to strigillose;
peduncle ca. 0.5 cm; bracts lanceolate, 12 mm; pedicels 12
mm. Calyx glabrous; ovary portion obconic, ca. 2 mm; limb ca.
2 mm, shallowly toothed; lobes 5. Corolla white, glabrous outside; tube 34 mm; lobes 5, narrowly spatulate-oblong, 56
mm, obtuse. Berry 46 mm in diam. Fl. Apr.
Open mountain slopes; 5001300 m. Hainan, Yunnan [Vietnam].

3. Aidia henryi (E. Pritzel) T. Yamazaki, J. Jap. Bot. 45: 338.

1970.
heng shi xiang nan
Randia henryi E. Pritzel, Bot. Jahrb. Syst. 29: 581. 1901;
Aidia merrillii (Chun) Tirvengadum; R. acutidens Hemsley &
E. H. Wilson; R. caudatifolia Merrill (1923), not Pitard (1923);
R. merrillii Chun.
Shrubs or trees, 215 m tall; branches somewhat flattened
to subterete, glabrous. Petiole 518 mm, glabrous; leaf blade
drying leathery or papery, elliptic-oblong, oblong-lanceolate, or
narrowly elliptic, 921.5 1.58 cm, both surfaces glabrous,
base cuneate to obtuse, apex acute to acuminate; secondary
veins 510 pairs, in abaxial axils usually with foveolate and/or
pilosulous domatia; stipules caducous or sometimes persisting
on distalmost nodes, lanceolate to narrowly triangular, 610
mm, glabrous, apex long acuminate. Inflorescences cymose, 1
7 110 cm, puberulent, strigillose, or glabrous, with axes
dichasial or sometimes congested; peduncle 210 mm; bracts
lanceolate to triangular or ovate, 0.52 mm, obtuse to acute;
pedicels 0.57 mm. Calyx glabrous to strigillose; ovary portion
obconic to cylindrical or narrowly ellipsoid, 11.5 mm; limb
with tubular portion 12.5 mm; lobes 4, triangular to narrowly
triangular or lanceolate, 0.51.5 mm. Corolla yellow, white, or
sometimes red, glabrous outside; tube 34 mm; lobes 4, narrowly spatulate-oblong, 58 mm, acute to obtuse or rounded.
Berry 56 mm in diam.; seeds ca. 2 mm. Fl. MarJun, fr. May
Feb.

Randia oxyodonta Drake, J. Bot. (Morot) 9: 218. 1895.

Shrubs or trees, 212 m tall; branches somewhat flattened
to subterete, puberulent to glabrous. Petiole 813 mm, glabrous;
leaf blade drying leathery and often pale yellow adaxially and
reddish brown abaxially, elliptic-oblong, lanceolate, or elliptic,
819 2.37.5 cm, glabrous on both surfaces, base obtuse to
acute, apex acuminate or acute; secondary veins 710 pairs, in
abaxial axils often with foveolate and/or pilosulous domatia;
stipules generally persistent, ovate to narrowly triangular, 515
mm, glabrous, apex long acuminate to aristate. Inflorescences
cymose with axes dichasial or often becoming monochasial
distally, 45 cm, glabrous; peduncle 0.81 cm; bracts ovate to
subulate, 23 mm, acute to acuminate; pedicels 25 mm. Calyx
glabrous; ovary portion obconic, ca. 1.5 mm; limb with tubular
portion campanulate, 55.5 mm; lobes 5, subulate or linearlanceolate, 24.5 mm, ciliate to glabrous. Corolla yellowish
white, outside glabrous; tube 45 mm; lobes oblong-spatulate,
910 35 mm, acuminate. Berry 713 mm in diam., glabrous;
seeds flattened, ca. 2.5 mm. Fl. AprNov, fr. MayOct.
Thickets or forests on hills or mountains; 1001000 m. SW
Guangdong, SE Guangxi, Hainan [Vietnam].
The Vietnam checklist (Checkl. Pl. Spec. Vietnam 3: 85. 2005)
recognized Aidia oxyodonta var. microdonta (Pitard) P. H. H (Ill. Fl.
Vietnam 3: 189. 1993), which is based on Randia oxyodonta var.
microdonta Pitard; presumably if this variety is recognized the Chinese
plants fall under A. oxyodonta var. oxyodonta, but so far Chinese
literature has not mentioned these varieties.

5. Aidia pycnantha (Drake) Tirvengadum, Nordic J. Bot. 3:

455. 1983.
duo mao qian cao shu
Randia pycnantha Drake, J. Bot. (Morot) 9: 218. 1895;
Aidia acuminatissima (Merrill) Masamune; R. acuminatissima
Merrill.
Shrubs or trees, 212 m tall; branches terete to flattened,
densely tomentulose, pilosulous, or hirtellous with pubescence
drying ferruginous, sometimes becoming glabrescent with age.
Petiole 515 mm, densely hirtellous or pilosulous; leaf blade
drying thinly leathery or papery, often reddish brown, ellipticoblong, oblong-lanceolate, or oblong-oblanceolate, 827.5 2
10 cm, adaxially glabrous and slightly shiny, abaxially hirtellous or pilosulous with pubescence usually denser on principal

RUBIACEAE

veins, base cuneate to obtuse and sometimes slightly inequilateral, apex acuminate to caudate-acuminate with tip to 2.5 cm
and sometimes falcate curved; secondary veins 1014 pairs, in
abaxial axils sometimes with weakly developed pilosulous
domatia; stipules deciduous or sometimes persisting on apical 2
or 3 nodes, interpetiolar, lanceolate to narrowly triangular, 812
mm, densely strigillose to hirtellous or tomentulose, apex acute
to acuminate. Inflorescences cymose with axes markedly dichasial, many flowered, 46 512 cm, branched to several
orders, densely hirtellous to pilosulous or tomentulose; peduncle 0.51.5 cm; bracts linear-lanceolate, 24 mm, acute; pedicels 14 mm. Calyx densely hirtellous to strigillose; ovary portion 11.5 mm; limb with tubular portion 23 mm; lobes 5,
narrowly triangular, 12 mm, acute to acuminate. Corolla white
or pale yellow, outside glabrous; tube ca. 4 mm, densely villous
in throat; lobes 5, oblong-oblanceolate or spatulate, 69 2
2.5 mm, obtuse to rounded. Berry 68 mm in diam., sparsely
strigillose to hirtellous or subglabrous; seeds ca. 2 mm. Fl.
MarSep, fr. AprDec.
Thickets or forests at streamsides, in fields or valleys, or on hills
or mountain slopes; near sea level to 1000 m. Fujian, Guangdong,
Guangxi, Hainan, Yunnan [Vietnam].

6. Aidia racemosa (Cavanilles) Tirvengadum, Nordic J. Bot. 3:

455. 1983.
zong zhuang qian cao shu

73

liu ye xiang nan

Randia salicifolia H. L. Li, J. Arnold Arbor. 24: 456. 1943.
Shrubs, ca. 1 m tall; branches slender, terete, glabrous.
Petiole 28 mm, glabrous; leaf blade drying papery, dark olivegreen adaxially, paler abaxially, narrowly lanceolate to narrowly elliptic, 823 0.53 cm, both surfaces glabrous or
sometimes pilose abaxially along principal veins, base cuneate
or acute, apex long acuminate; secondary veins 912 pairs, in
abaxial axils with foveolate domatia; stipules caducous, lanceolate to triangular, 310 mm, glabrous, apex long acute to acicular. Inflorescences not seen. Infructescences congested-cymose,
subsessile, 0.51.5 cm; bracts not described; pedicels in fruit ca.
4 mm. Berry 68 mm in diam. Fr. Nov.
Forests on mountains; 6001000 m. Guangxi.
Ridsdale (Blumea 41: 135179. 1996) noted that this species is
only reliably known from the type collection, though the additional,
more broad-leaved collections Steward et al. 544 and Steward et al. 806
(no herbarium given by him) from Kweichow might be conspecific.

8. Aidia yunnanensis (Hutchinson) T. Yamazaki, J. Jap. Bot.

45: 339. 1970.
dian qian shu
Randia yunnanensis Hutchinson in Sargent, Pl. Wilson. 3:
400. 1916.

Forests; elevation in China not noted on specimens [ca. 200 m to

probably higher]. Hainan [Indonesia, Malaysia, New Guinea, Philippines, Thailand; Australia, Pacific islands].

Shrubs or trees, 27 m tall; branches somewhat flattened

to subterete, glabrous. Leaves subsessile to petiolate; petiole to
6 mm, strigillose to glabrous; blade drying papery or thinly
leathery and sometimes reddish brown, elliptic-lanceolate or
oblong-lanceolate, 6.217.5 2.76.5 cm, adaxially glabrous,
abaxially glabrous or sometimes strigose or strigillose along
principal veins, base cuneate to shortly rounded, sometimes
slightly inequilateral, apex acuminate to caudate-acuminate;
secondary veins 710 pairs, in abaxial axils sometimes with
small foveolate and/or pilosulous domatia; stipules caducous,
interpetiolar or shortly united around stem, lanceolate to narrowly triangular, 47 mm, strigillose to glabrous, smooth to
keeled, apex acute to acuminate. Inflorescences fasciculate, several flowered, strigillose to hirtellous or strigose sometimes
becoming glabrescent with age; bracts triangular, 0.51 mm,
acute; pedicels 24 mm. Calyx strigillose; ovary portion obconic to ellipsoid, ca. 1 mm; limb 12.5 mm including lobes,
shallowly lobed; lobes 4, triangular to broadly triangular, 0.5
0.7 mm, acute. Corolla white, outside glabrous; tube ca. 3 mm,
villous in throat; lobes 4, narrowly oblong-spatulate, 45 ca. 2
mm, obtuse to rounded. Berry red, 58 mm in diam., strigillose
or glabrous. Fl. MarMay, fr. MayJan.

7. Aidia salicifolia (H. L. Li) T. Yamazaki, J. Jap. Bot. 45: 339.

1970.

Thickets or forests on hills or mountains; 5001700 m. S Yunnan

[Thailand (Maxwell 97-144, MO!)].

Stylocoryna racemosa Cavanilles, Icon. 4: 46. 1798;

Randia racemosa (Cavanilles) Fernndez-Villar (1880), not
Roxburgh (1824); R. suishaensis Hayata.
Trees to 25 m tall; branches somewhat flattened becoming
subterete, glabrous. Petiole 56 mm, glabrous; leaf blade drying
thinly leathery, lanceolate to elliptic-oblong, 712 24 cm,
glabrous on both surfaces, base acute to cuneate, apex acute;
secondary veins 4 or 5 pairs, in abaxial axils usually with
foveolate and/or pilosulous domatia; stipules caducous, shortly
united around stem, narrowly triangular, 35 mm, glabrous,
apex acuminate. Inflorescences cymose, ca. 3 46 cm, glabrescent, with axes becoming monochasial distally; peduncle
ca. 0.5 cm; bracts triangular, 11.5 mm, acute; pedicels 13
mm. Calyx glabrous; ovary portion ellipsoid, ca. 1 mm; limb
ca. 1.5 mm, shortly dentate to denticulate; teeth 5. Corolla white,
outside glabrous; tube ca. 4 mm; lobes 5, narrowly spatulateoblong, 55.5 mm, adaxially strigillose, obtuse to rounded.
Berry 48 mm in diam., glabrous.

4. ALLEIZETTELLA Pitard in Lecomte, Fl. Indo-Chine 3: 278. 1923.

bai xiang nan shu
Chen Tao (); Charlotte M. Taylor
Shrubs, sometimes scandent, perhaps sometimes dioecious, unarmed. Raphides absent. Leaves opposite, sometimes anisophyllous, usually with domatia; stipules caducous or persistent, interpetiolar or shortly united around stem, generally triangular. In-

RUBIACEAE

74

florescences terminal on principal stems and/or pseudoaxillary on reduced lateral branches, shortly cymose, several flowered, sessile
to pedunculate, bracteate. Flowers sessile to pedicellate, bisexual and monomorphic or perhaps sometimes unisexual. Calyx limb 5lobed. Corolla white, salverform to funnelform, inside pubescent in throat and upper part of tube; lobes 5, convolute in bud. Stamens
5, inserted at upper part of corolla or at throat, included and positioned near base of corolla tube; filaments short; anthers dorsifixed.
Ovary 2-celled, ovules 2 or 3 in each cell, on axile placentas; stigma 2-lobed with lobes linear, exserted. Fruit yellowish white, baccate, subglobose to oblate, fleshy, smooth, with calyx limb persistent or tardily deciduous; seeds few to several, medium-sized, ellipsoid to ovoid, embedded in pulp.
Two species: China, Vietnam; one species in China.
Most authors have described Alleizettella as having bisexual flowers, but Robbrecht and Puff (Bot. Jahrb. Syst. 108: 122123, table 7. 1986)
suggested it is sometimes dioecious and thus with unisexual flowers.

1. Alleizettella leucocarpa (Champion ex Bentham) Tirvengadum, Nordic J. Bot. 3: 455. 1983.

bai guo xiang nan
Randia leucocarpa Champion ex Bentham, Hookers J.
Bot. Kew Gard. Misc. 4: 194. 1852; Aidia leucocarpa
(Champion ex Bentham) T. Yamazaki.
Shrubs, perhaps sometimes scandent, 13 m tall; branches
compressed to subterete or subangled, strigose to strigillose or
pilosulous often becoming glabrescent. Leaves equal to somewhat unequal; petiole 412 mm, strigose to strigillose; blade
drying papery or thinly leathery, abaxially sometimes paler
and/or brownish black speckled, oblong-obovate, elliptic-oblong, narrowly elliptic, or lanceolate, 4.517 1.56 cm, adaxially glabrous or sometimes strigillose to hirtellous along principal veins, abaxially glabrous to usually puberulent or hirtellous
on blade and strigose to hirtellous along principal veins, base
acute to cuneate, apex subacuminate to acuminate; secondary

veins 47 pairs, in abaxial axils usually with pilosulous and

usually also foveolate domatia; stipules persistent, shortly united
around stem, broadly triangular, 47 mm, strigose to strigillose,
long acuminate. Inflorescences 12 cm, densely strigose, strigillose, or hirtellous; bracts triangular to lanceolate, 0.52 mm,
acute to acuminate; pedicels 23.5 mm. Calyx sparsely to
densely strigose to strigillose; ovary portion obconic, 11.5
mm; limb campanulate, basal tubular portion 1.52 mm; lobes
triangular to linear, 0.51 mm, acute. Corolla funnelform; tube
34 mm, outside strigose in upper part, inside villous at throat;
lobes subovate to ligulate, 1.52 mm, outside puberulent to glabrous, obtuse to acute. Infructescences usually displaced to
pseudoaxillary by subsequent stem growth. Fruit subglobose to
somewhat oblate, 813 mm in diam., strigillose, pilosulous, or
glabrous; seeds 24, compressed globose, 45 mm in diam. Fl.
AprJun, fr. JunFeb.
Forests or thickets at streamsides in valleys or on mountain slopes;
2001000 m. Fujian, Guangdong, Guangxi, Hainan [Vietnam].

5. ANTIRHEA Commerson ex Jussieu, Gen. Pl. 204. 1789.

mao cha shu
Chen Tao (); Charlotte M. Taylor
Guettardella Champion ex Bentham.
Trees or shrubs, dioecious, unarmed. Raphides absent. Leaves opposite [or sometimes whorled], often with domatia; quaternary
venation lineolate [or sometimes regularly areolate]; stipules caducous or persistent, interpetiolar, generally triangular to oblong.
Inflorescences axillary, cymose with axes dichotomous or often markedly scorpioid, few flowered with flowers often fewer on
pistillate plants, pedunculate, bracteate or bracts reduced. Flowers sessile, unisexual. Calyx sericeous outside; limb truncate or 4[or
5]-lobed; lobes often unequal. Corolla white or yellow, salverform in staminate flowers, funnelform in pistillate flowers, with tube
often prolonged and slender, inside glabrous or pubescent in throat; lobes 4[or 5], obtuse, imbricate in bud. Stamens 4[or 5], inserted
in corolla throat, partially exserted; filaments short or reduced; anthers dorsifixed. Ovary 28-celled, ovule 1 in each cell, apical and
pendulous, with funicle thickened; stigma capitate or 2- or 3-lobed, included. Fruit dark purple, drupaceous, thinly fleshy, ellipsoid to
subglobose and smooth, with calyx limb and subtending bracts persistent; pyrene 1, 28-celled with 1 seed in each cell, ellipsoid,
woody or bony; seeds cylindrical, medium-sized, without endosperm; cotyledons compressed and minute; radicle clavate and
ascending.
Thirty-six species: tropical Asia, Australia, Madagascar, Mascarene Islands, Pacific islands; one species (endemic) in China.
Antirhea has traditionally included both paleotropical and neotropical species, but the bisexual and polygamous neotropical plants are now
included in other genera; even so, some authors still incorrectly give a pantropical range for this genus.
Chaw and Darwin (Tulane Stud. Zool. & Bot. 28: 50, 59, 69. 1992) recognized three subgenera of Antirhea. Antirhea chinensis is the type of A.
subg. Guettardella (Champion ex Bentham) Chaw, the largest subgenus.

1. Antirhea chinensis (Champion ex Bentham) Bentham & J.

D. Hooker ex F. B. Forbes & Hemsley, J. Linn. Soc., Bot. 23:
384. 1888.
mao cha

Guettardella chinensis Champion ex Bentham, Hookers J.

Bot. Kew Gard. Misc. 4: 197. 1852.
Shrubs, erect, 12(4) m tall; branches subterete to somewhat flattened, moderately to densely strigose, strigillose, or to-

RUBIACEAE

mentulose, often with trichomes of 2 lengths, occasionally becoming glabrescent, often markedly sylleptic from nodes below
leaves with markedly elongated lowermost internode. Petiole
410 mm, densely strigose to tomentulose; leaf blade drying
papery, elliptic-oblong, oblong-lanceolate, oblanceolate, elliptic, or narrowly elliptic, 37(9) 12.5(3) cm, adaxially glabrous or sparsely strigillose, abaxially densely sericeous to
strigillose, base cuneate to acute, margin flat to thinly revolute,
apex acuminate; secondary veins 46 pairs, usually with welldeveloped pilosulous domatia; stipules usually persisting with
leaves, triangular to narrowly triangular, 2.56 mm, abaxially
strigose or strigillose to glabrescent, adaxially densely sericeous, acuminate. Inflorescences cymose, dichotomous or with

75

higher order axes often scorpioid, staminate several flowered,

pistillate few to several flowered, sericeous to strigillose; peduncles 13 cm; branched portion 13 13 cm; bracts linear
to subulate, 1.54 mm. Calyx densely sericeous; ovary portion
ellipsoid to obconic, 0.51 mm; limb deeply lobed, tubular part
0.30.5 mm; lobes linear or lanceolate, 0.51.5 mm, often
unequal. Corolla yellow, outside, densely sericeous; staminate
tube 69 mm, pistillate 36 mm, inside glabrous or sparsely
sericeous in lower portion; lobes ovate, 12 mm, obtuse. Drupe
ellipsoid-oblong or ellipsoid, 57 34 mm, sparsely strigose;
pyrene 25-celled, angled to ridged. Fl. AprJun, fr. OctNov.
Forests, thickets; 1001700 m. Fujian, Guangdong, Hainan.

6. ARGOSTEMMA Wallich in Roxburgh, Fl. Ind. 2: 324. 1824.

xue hua shu
Chen Tao (); Charlotte M. Taylor
Herbs, unarmed, often reduced and/or monocaulous, often rhizomatous or tuberous, usually notably fleshy; stems flattened to
subterete. Raphides present. Leaves opposite, apparently verticillate due to closely set stem nodes and/or leaflike stipules, or slightly
to markedly anisophyllous and apparently alternate, decussate or distichous, without domatia; stipules persistent or deciduous,
interpetiolar, entire or bilobed, sometimes leaflike and thus apparently absent. Inflorescences terminal or sometimes pseudoaxillary,
fasciculate, cymose-corymbiform, or umbelliform, several flowered, few flowered, or reduced to 1 flower, sessile to pedunculate,
bracteate with bracts sometimes fused into an involucre or sometimes with bracts very reduced. Flowers pedicellate or sessile,
bisexual or monomorphic, sometimes somewhat zygomorphic, usually nodding. Calyx limb deeply 4- or 5-lobed. Corolla white,
broadly rotate (i.e., Solanum-like) to campanulate (i.e., bell-shaped), glabrous inside; lobes 4 or 5, valvate in bud. Stamens 4 or 5,
inserted near base of corolla tube, partially to fully exserted; filaments reduced or developed and free, coherent, or fused in middle
portions into a tube; anthers free or usually coherent into a tube, with longitudinal slits or terminal pores, sometimes with connective
prolonged at apex. Ovary 2-celled, ovules numerous in each cell on axile placentas attached near top of septum; stigma capitate or 2lobed, included or exserted. Fruiting pedicels usually becoming thickened and erect. Fruit capsular, subglobose to obovoid, dehiscent
through apical operculum formed from disk portion, leathery or membranous, often becoming black, with calyx limb persistent, with
endocarp and septum becoming notably fibrous; seeds numerous, small, flattened to angled, with testa reticulate; endosperm fleshy.
At least 106 species: Bhutan, Cambodia, China, India, Indonesia, Japan, Laos, Malaysia, Myanmar, Nepal, New Guinea, Philippines, Thailand,
Vietnam; two species endemic to W Africa; six species (four endemic) in China.
The morphology and anatomy of the androecium of Argostemma were detailed by Puff et al. (Ann. Missouri Bot. Gard. 82: 358366. 1995),
who noted an absence of nectaries and probable buzz-pollination of flowers with both linear and poricidal anther dehiscence. Bremer (Ann. Missouri
Bot. Gard. 76: 749. 1989) reported that the inner layers of the fruit become fibrous and trap seeds inside, with some of them sprouting while still in
the capsule in an unusual form of vivipary. Argostemma was studied in Thailand by Sridith (Thai Forest Bull., Bot. 27: 86137. 1999) and Sridith and
Puff (Thai Forest Bull., Bot. 28: 123137. 2000). Argostemma species apparently vary widely in size of individual plants and vegetative organs
probably in relation to environmental factors.

1a. Leaves subsessile, most of them apparently in whorls of 4 at tops of stems, slightly to markedly unequal;
anthers free, opening by apical pores ...................................................................................................................... 5. A. verticillatum
1b. Leaves equal or unequal in pairs or sometimes apparently alternate due to marked reduction of 1 leaf of a
pair, at least larger leaf of a pair clearly petiolate; anthers coherent in a cone, opening by longitudinal slits.
2a. Calyx and fruit pilosulous, strigillose, villosulous, villous, or hirsute.
3a. Calyx and fruit pilosulous or strigillose with generally straight hairs; larger leaf of a pair 14 cm ................ 1. A. discolor
3b. Calyx and fruit densely villous, villosulous, or hirsute with usually crisped hairs; larger leaf of a
pair 2.510 cm ............................................................................................................................................ 4. A. solaniflorum
2b. Calyx and fruit glabrous or puberulent.
4a. Corolla lobes ovate, ca. 5 mm ............................................................................................................................ 3. A. saxatile
4b. Corolla lobes lanceolate, 8.511 mm.
5a. Leaf blade brownish yellow abaxially when dry, with secondary veins not visible; larger leaf
of a pair 14 0.61.5 cm .................................................................................................................... 2. A. hainanicum
5b. Leaf blade pale abaxially when dry, with secondary veins visible; larger leaf of a pair
26 1.52.5 cm ................................................................................................................................... 6. A. yunnanense

RUBIACEAE

76

1. Argostemma discolor Merrill, Philipp. J. Sci. 23: 265. 1923.

yi se xue hua
Herbs, 715 cm tall, little branched; stems prostrate and
rooting or with apices ascending, densely strigose to villosulous. Leaves opposite, markedly anisophyllous, in larger leaf of
a pair petiole 38 mm, villosulous; blade drying papery and
pale abaxially, elliptic, elliptic-oblong, or broadly ovate, 1040
720 mm, adaxially sparsely strigose to hispid along midrib
and near margins, abaxially glabrous on lamina and densely
strigose to hispid along veins, base cuneate to obtuse, margin
serrate-ciliate, apex acute to obtuse; secondary veins 46 pairs;
smaller leaf of a pair subsessile, suborbicular or broadly ovate,
510 mm; stipules persistent, ovate, suborbicular, or obovate,
26 mm, glabrescent, ciliate. Inflorescences 2- or 3-flowered,
strigose to hispid; peduncles 520 mm; bracts triangular to
linear, 13 mm; pedicels 1025 mm. Calyx pilosulous or strigillose; hypanthium portion obconic, 11.5 mm; lobes 5, triangular to lanceolate, 1.52 mm. Corolla white, rotate, glabrous;
tube ca. 2 mm; lobes 5, lanceolate, 67 mm, ciliate. Stamens 5;
anthers coherent, dehiscent by longitudinal slits, with connective prolonged. Capsules subglobose or obovoid, ca. 3 mm in
diam. Fl. MarMay, fr. SepOct.
Dense forests; 5001500 m. Hainan.

2. Argostemma hainanicum H. S. Lo, Bull. Bot. Res., Harbin

6(4): 46. 1986.
hai nan xue hua
Herbs, 1020 cm tall; stems grayish black when dry, suberect or creeping, densely strigose-villous becoming glabrescent. Leaves opposite, anisophyllous; blade drying black adaxially, dark brown abaxially, oblong-lanceolate or oblong-ovate,
1040 612(15) mm, both surfaces strigose on midrib and
sometimes near margins, base obtuse, margin entire or erose to
ciliate, apex acute or shortly acuminate; secondary veins not
visible; smaller leaf of a pair subovate, 28(10) mm. Inflorescence 1- or 2-flowered, villous to glabrescent; peduncles 12
cm; bracts lanceolate, 1.52 mm. Calyx puberulent or subglabrous; lobes 5. Corolla white, rotate, glabrous; lobes 5, lanceolate, ca. 1 cm. Stamens 5; anthers coherent, dehiscent by longitudinal slits, with connective prolonged. Capsules slightly cordate, ?maybe didymous, ca. 5 78 mm, glabrous. Fl. May.
Valleys or watersides in dense forests. Hainan (Ledong).

persistent, subovate, ca. 1 mm, glabrescent. Inflorescences umbelliform or racemiform, 24-flowered, glabrous; peduncles 1
or 2, 13 cm; bracts ovate, 24 mm; pedicels 13 cm. Calyx
glabrous; hypanthium portion obconic, 22.5 mm; lobes 5,
broadly triangular, 1.52 mm. Corolla white, apparently rotate,
glabrous or subglabrous; tube 1.52 mm; lobes 5, ovate, ca. 5
mm. Stamens 5; anthers coherent, dehiscent by longitudinal
slits, with prolonged connective. Capsules not seen. Fl. Mar.
Wet sites in dense forests; ca. 600 m. SW Guangxi.

4. Argostemma solaniflorum Elmer, Leafl. Philipp. Bot. 1: 2.

1906.
shui guan cao
Argostemma iriomotense Masamune; A. taiwanense S. S.
Ying.
Herbs, to 30 cm tall, sometimes branched; stems erect,
puberulent or villosulous to glabrescent. Leaves opposite,
slightly to markedly anisophyllous, in larger leaf of a pair petiole 0.53 cm, villosulous to glabrescent; blade drying membranous, ovate, lanceolate, or elliptic-oblong, 25100 1230
mm, adaxially glabrous or sparsely hirsute or pilose, abaxially
pilosulous or villosulous along principal veins and glabrous on
lamina, base cuneate to obtuse, margin entire and glabrous,
apex acute to weakly acuminate; secondary veins 7 or 8 pairs;
stipules persistent, oblong-elliptic or ovate, 37 mm, glabrescent, obtuse. Inflorescences cymose, 16-flowered, strigillose
or villosulous to glabrescent; peduncles 12.5 cm; bracts triangular and 13 mm or resembling reduced leaves and stipules;
pedicels 520 mm. Calyx villous, villosulous, or hirsute; hypanthium portion subglobose, ca. 1.5 mm; lobes 5, triangular, ca.
1.5 mm. Corolla white, rotate, glabrous; tube ca. 1.5 mm; lobes
5, narrowly lanceolate, ca. 7 mm, marginally densely ciliate.
Stamens 5; anthers coherent, 5.57 mm, dehiscent by linear
slits. Capsules cupuliform, 23 mm, leathery, densely hirsute or
villosulous, with persistent calyx limb to 3 mm. Fl. MarApr, fr.
Jul.
Shady and moist streamsides; below 100500 m. Taiwan [Japan
(Ryukyu Islands), Philippines].
This species appears to vary widely in size of the plants, leaf size,
and pubescence; it has sometimes been characterized (e.g., FRPS 71(1):
180182. 1999) based only on larger plants, as described in the protologue. The two synonymous names listed here were not included in
FRPS but were cited in Fl. Taiwan (ed. 2, 4: 247. 1998).

3. Argostemma saxatile Chun & F. C. How ex W. C. Ko, Fl.

Hainan. 3: 578. 1974.

5. Argostemma verticillatum Wallich in Roxburgh, Fl. Ind. 2:

325. 1824.

yan xue hua

xiao xue hua

Herbs, 515 cm tall, unbranched or sometimes few

branched; stems prostrate and rooting near base, ascending
at apex, villous. Leaves opposite, markedly anisophyllous,
in larger of a pair petiole 35 mm, pilose; blade drying thinly
papery and pale abaxially, oblong-elliptic or ovate, 1555 8
18 mm, adaxially sparsely strigose at least along midrib and
margin, abaxially strigose at least along veins, base cuneate to
obtuse, apex acute or rarely acuminate; secondary veins 57
pairs; smaller leaf of a pair subsessile, ovate, ca. 5 mm; stipules

Dwarf herbs, 27 cm tall, unbranched, borne from subglobose tubers; stems erect, puberulent to glabrous. Leaves clustered at stem apex and usually apparently 4-verticillate, anisophyllous, subsessile; blade drying membranous to thinly papery,
ovate, elliptic, elliptic-oblong, or obovate, 1070 725 mm,
both surfaces glabrous [or sometimes sparsely villosulous to
villous], base obtuse to acute, apex acute to obtuse or shortly
acuminate; secondary veins 47 pairs; stipules deciduous, elliptic to orbicular, 23 mm, obtuse to rounded. Inflorescences

RUBIACEAE

cymose to umbelliform, 2- to several flowered, glabrous; peduncles 13, 0.72 cm; bracts ligulate to ovate, 12 mm; pedicels 510 mm. Calyx glabrous; hypanthium portion broadly
campanulate to obconic, ca. 1 mm; lobes 5, subtriangular, ca. 1
mm. Corolla white, rotate, glabrous; tube ca. 1 mm; lobes 5, oblong-lanceolate to triangular, 35 mm. Stamens 5; anthers free,
23 mm, poricidal. Capsules obovoid, ca. 2 mm, with persistent
calyx limb to 2 mm. Fl. Jun.
Streamsides in forests; ca. 1500 m. S Yunnan [Bhutan, NE India,
Myanmar, Nepal, Vietnam].
This species was reported from Thailand by Sridith (Thai Forest
Bull., Bot. 27: 118. 1999) and as having filaments fused at their middles
into a tube, but according to the Chinese specimens studied and descriptions of this species in other regions (e.g., Fl. Bhutan 2(2): 756. 1999)
the filaments are free. The Thai distribution is provisionally excluded
here pending further study of the delimitation of this species.

6. Argostemma yunnanense F. C. How ex H. S. Lo, Bull. Bot.

Res., Harbin 6(4): 45. 1986.

77

dian xue hua

Herbs, 620 cm tall; stems procumbent and rooting with
apices ascending, densely crisped pubescent. Leaves opposite,
anisophyllous, in larger leaf of pair petiole to 1 cm; blade
drying pale abaxially, oblong-obovate, 2060 1525 mm,
adaxially sparsely pilose to subglabrous, abaxially pubescent
with pubescence denser on midrib and secondary veins, base
obtuse, apex acute; secondary veins 710 pairs; smaller leaf of
a pair subsessile, ovate to suborbicular, 46(10) mm; stipules
ovate, 57 mm. Inflorescences umbelliform, 2- or 3-flowered;
peduncles 22.5 cm; bracts lanceolate, 35 mm; pedicels 11.5
cm. Calyx glabrous; hypanthium ca. 2 mm; lobes 5, broadly
triangular, 11.2 mm. Corolla white, rotate; tube ca. 2.5 mm;
lobes 5, lanceolate, 8.511 mm, margins glabrous. Stamens 5;
anthers coherent, 44.5 mm, dehiscent by longitudinal slits,
with prolonged connective ca. 3 mm. Capsules not seen. Fl.
May.
Dense forests; ca. 900 m. Yunnan.

7. ASPERULA Linnaeus, Sp. Pl. 1: 103. 1753, nom. cons.

che ye cao shu
Chen Tao (); Friedrich Ehrendorfer
Asperula sect. Chlorostemma Lange; Chlorostemma (Lange) Fourreau.
Subshrubs, perennial, or annual herbs. Raphides present. Leaves opposite, usually with leaflike stipules in whorls of 414, sessile to shortly petiolate, without domatia; leaflike stipules rarely reduced. Inflorescences thyrsoid, paniculiform to capitate, with terminal and often also axillary pedunculate to sessile cymes, bracteate with bracts often fused and sometimes involucral. Flowers
pedicellate to sessile, with prophylls, bisexual, monomorphic. Calyx limb reduced, practically absent. Corolla blue, pink, purple, or
yellow to greenish or white, salverform, funnelform, campanulate, or sometimes rotate, glabrous inside; lobes 4 or 5, valvate in bud.
Stamens 4 or 5, inserted in corolla tube, exserted (or sometimes included); filaments developed to short; anthers dorsifixed. Ovary
inferior (hypanthium), 2-celled, ovules 1 in each cell, erect and axile; stigma globose to clavate, often 2-lobed, included or exserted.
Fruit schizocarpous, generally didymous, dry; mericarps 2, indehiscent, with 1 seed, subglobose, ellipsoid-oblong, or reniform,
smooth to tuberculate, glabrous to pubescent (but never with uncinate hairs); seeds small, with membranous testa; endosperm
corneous; embryo curved; cotyledons leaflike; radicle terete, hypogynous.
About 200 species: widespread throughout N Africa, C and SW Asia, and Europe, extending into Australia and New Zealand, greatest species
diversity in the dry regions of SW Asia and the E Mediterranean; two species (one introduced) in China.
The circumscription and relationships of Asperula were discussed most recently by Ehrendorfer et al. (Fl. Iranica 176: 105161. 2005). Short
references to the position of the genus within the Rubieae-Rubiinae are found in the introduction to the genus Galium of the present volume and its
Chinese species are keyed out there.
Originally, the Linnaean genera Asperula and Galium were separated from each other on the basis of their salverform to campanulate vs. rotate
corollas only. Sixty years of critical morphological and later DNA-analytical studies (see Natali et al., Opera Bot. Belg. 7: 193203. 1996; Soza &
Olmstead, Taxon 59: 755771. 2010) have shown that this differentiation often does not reflect true phylogenetic relationships. In some obvious cases
(e.g., A. odorata Linnaeus to G. odoratum (Linnaeus) Scopoli in G. sect. Hylaea (Grisebach) Ehrendorfer or G. purpureum Linnaeus to A. purpurea
(Linnaeus) Ehrendorfer in A. sect. Thliphthisa Grisebach), the problem could be solved by a simple nomenclatorial transfer, but in several other cases
the problems persist. Even after an effort to redefine the two genera with the help of the presence of prophylls (bracteoles) at the pedicels in Asperula
vs. their absence in Galium (Ehrendorfer et al., Fl. Europaea 4: 338. 1976) the two genera are still phylogenetically interdigitated and heterogeneous.
Thus, one is still left with a partly provisional classification of Asperula as proposed by Ehrendorfer et al. (loc. cit. 2005). Here, we follow FRPS
(71(2): 213. 1999) and do not combine the genus Leptunis with Asperula (as in loc. cit. 2005).
As in Galium, the sectional classification of Asperula by Ehrendorfer et al. (loc. cit. 2005) does not fully agree with that of Pobedimova et al.
(Fl. URSS 23: 205285. 1958), which was followed by FRPS. In particular, Ehrendorfer et al. (loc. cit. 2005: 131142, 157158) placed A. oppositifolia in A. sect. Oppositifoliae Schishkin ex Schnbeck-Temesy and A. orientalis in A. sect. Asperula (A. sect. Sherardianae Candolle). The two species are keyed out below but are also included in the key to all taxa of Chinese Rubieae found in the present volume under Galium.

1a. Perennials, with stems originating from a woody rootstock; leaves at all nodes opposite, 36(15)
0.51.5(3) mm, glabrous, stipules reduced; flowers pink to purple ..................................................................... 1. A. oppositifolia

RUBIACEAE

78

1b. Herbaceous annuals, with stems from fibrous roots; leaves and leaflike stipules at upper nodes in whorls of
48, 1.22.5 0.20.5 cm, glabrous to sparsely hispidulous on lamina and densely antrorsely aculeolate on
veins and margins; flowers pale to clear purplish blue ................................................................................................ 2. A. orientalis
1. Asperula oppositifolia Regel & Schmalhausen in Regel,
Descr. Pl. Nov. Rar. 42. 1882.
dui ye che ye cao
Subshrubs, perennial, originating from a woody rootstock.
Stems often woody at base, up to 40 cm tall, erect, weakly 4angled, glabrous to puberulent. Leaves opposite, subsessile;
blade drying stiffly papery, linear to linear-lanceolate, 36(15)
0.51.5(3) mm, glabrous throughout or hairy, base and
apex acute; secondary veins not evident; stipules 24 per leaf
pair, reduced and never more than 0.3 mm. Inflorescences terminal and axillary, dichasial, branched to 14 orders, glabrous
to puberulent; peduncles 110(30) mm; bracts linear to narrowly elliptic (i.e., leaflike), 14 mm; pedicels 03 mm. Corolla
pink to purple, funnelform, glabrous to sparsely pilose outside;
tube ca. 2 mm; lobes 4, ovate-oblong, ca. 1.5 mm. Ovary subglobose to narrowly ellipsoid, 0.51 mm, glabrous to densely
pilosulous. Mericarps ovoid, 1.52.5 mm, glabrous to densely
pilosulous. Fl. JunJul, fr. JulAug.
Gravel on mountain slopes; ca. 3700 m. Xizang (Zanda) [Afghanistan, Pakistan, Tajikistan].
Asperula oppositifolia is a polymorphic species with several subspecies and belongs to the very variable A. sect. Oppositifoliae (Ehrendorfer et al., Fl. Iranica 176: 131142. 2005). As we have not seen material from China, the data presented here are mostly taken from FRPS
(71(2): 214. 1999) and do not allow an exact determination. Outside of
China, populations of A. oppositifolia are found at elevations down to
1350 m. Their leaves are narrow and short relative to the internodes and
sometimes deciduous. Thus, the plants appear to consist only of photosynthetic stems with small terminal groups of flowers.

2. Asperula orientalis Boissier & Hohenacker in Boissier,

Diagn. Pl. Orient., ser. 1, 3: 30. 1843.

lan hua che ye cao

Asperula arvensis Linnaeus subsp. orientalis (Boissier &
Hohenacker) Thibaud; A. azurea Jaubert & Spach.
Herbs, annual, from fibrous roots. Stems few or solitary, to
30(40) cm tall, 4-angled, erect, often regularly divaricately
branched, glabrous to scaberulous or hispidulous. Leaves and
leaflike stipules at middle stem regions in whorls of 48, subsessile; blade drying papery, lanceolate, linear-lanceolate, or
spatulate, (7)1225(30) (1.5)25(10) mm, glabrous to
sparsely hispidulous on lamina and densely scaberulous to
antrorsely aculeolate on veins and margins, base acute, apex
obtuse to rounded; secondary veins not evident. Inflorescences
terminal, capitate to subcapitate; peduncles 1.54 cm; involucral bracts leaflike, 112 mm, white ciliate at margins. Flowers
sessile. Corolla pale to clear purplish blue, salverform, outside
densely papillose; tube 810 mm, dilated in throat around anthers; lobes 4, elliptic to ovate, 23 mm, obtuse. Ovary ovoid,
ca. 1 mm, glabrescent. Mericarps 11.8 mm, glabrous. Fl. Jun
Jul, fr. AugSep.
Cultivated ornamental in Anhui, Jiangsu (Nanjing), and Shaanxi
(Xian) [native to SW Asia (Georgia, Iraq, Lebanon, Syria, Turkey)].
Asperula orientalis is a butterfly-pollinated SW Asiatic member
of A. sect. Asperula. Pobedimova et al. (Fl. URSS 23: 283. 1958) erroneously gave the name A. azurea priority over A. orientalis. The other
closely related taxa of A. sect. Asperula are the smaller-flowered and
widespread A. arvensis Linnaeus (the conserved type of the genus) and
A. setosa Jaubert & Spach. The latter has very small flowers, is obviously autogamous, and grows at higher elevations (12003200 m)
from SW Asia to the Tian Shan and Pamir-Alai. It could be found in the
mountains of W China.

8. BENKARA Adanson, Fam. Pl. 2: 85, 525. 1763.

le qian shu
Chen Tao (); Charlotte M. Taylor
Fagerlindia Tirvengadum; Griffithia Wight & Arnott.
Shrubs or small trees, erect to clambering or perhaps scandent, usually armed with paired, ascending, straight to slightly curved,
axillary thorns, often with short shoots. Raphides absent. Leaves opposite or sometimes congested to apparently fascicled on lateral
short shoots, usually with domatia; stipules caducous, interpetiolar or shortly united around stem, generally triangular, acute to
cuspidate. Inflorescences terminal on lateral branches and/or short shoots, 1-flowered or 2- to several flowered and fasciculate to
cymose, sessile to pedunculate, bracteate or bracts reduced. Flowers pedicellate, bisexual, monomorphic. Calyx limb 5-lobed, usually
with well-developed basal tubular portion. Corolla white to yellow, salverform, usually pubescent in throat; lobes 5, convolute in
bud, usually spreading to reflexed. Stamens 5, inserted in corolla throat, partially to fully exserted; filaments short; anthers dorsifixed. Ovary 2-celled, ovules several to many in each cell on axile placentas; stigma clavate to fusiform or 2-lobed with lobes coherent, included to partially exserted. Fruiting pedicels sometimes elongating. Fruit purplish black, baccate, fleshy to leathery or infrequently woody, globose to ellipsoid, smooth, with calyx limb tardily deciduous; seeds several to numerous, medium-sized, angled or
ellipsoid, embedded in pulp; testa weakly areolate.
About 19 species: China, India, Indonesia, Japan, Malaysia, Myanmar, Philippines, Thailand, Vietnam; seven species (four endemic) in China.
Ridsdale (Reinwardtia 12: 289300. 2008) recently reviewed Oxyceros and Fagerlindia and concluded by re-circumscribing Oxyceros Loureiro
(Fl. Cochinch. 1: 150. 1790) and synonymizing Fagerlindia with Benkara. In this new circumscription, Oxyceros comprises species distinguished by
markedly recurved thorns (or spines), notably O. horridus Loureiro, and is not represented (as far as now known) in the Chinese flora. Ridsdale (loc.

RUBIACEAE

79

cit.) transferred all the Chinese species previously included in Oxyceros to Benkara, with straight or only slightly curved spines or thorns. One
Chinese species included by Ridsdale in Fagerlindia and later Benkara is here instead treated as Aidia canthioides.
Benkara griffithii (J. D. Hooker) Ridsdale (Reinwardtia 12: 298. 2008; Randia griffithii J. D. Hooker, Fl. Brit. India 3: 112. 1880; Oxyceros
griffithii (J. D. Hooker) W. C. Chen) is not treated here. See the discussion under B. forrestii below.

1a. Flowers subsessile to very shortly pedicellate, on pedicels 0.51.5 mm ....................................................................... 7. B. sinensis
1b. Flowers pedicellate or pedunculate, on pedicels or peduncles 115 mm with at least some of them more than
2 mm.
2a. Branchlets and calyx glabrous; corolla white, with tube 1422 mm and lobes 512 mm.
3a. Secondary leaf veins 46 pairs; calyx lobes 0.51.2 mm; plants drying with characteristic yellowed
color ................................................................................................................................................................. 5. B. rectispina
3b. Secondary leaf veins 2 or 3 pairs; calyx lobes 1.52 mm; plants drying greenish to grayish ........................ 6. B. scandens
2b. Branchlets and calyx glabrous to puberulent, pilosulous, strigillose, or hirtellous; corolla white or yellow,
with tube 37 mm and lobes 58 mm.
4a. Calyx limb with tubular portion 3.54 mm; corolla white ........................................................................ 1. B. depauperata
4b. Calyx limb with tubular portion 13 mm; corolla white to yellow.
5a. Flowers 1 or 2 in fascicles, these sometimes congested and appearing as a cyme; leaves obtuse
to rounded and sometimes shortly mucronate at apex ................................................................................ 2. B. evenosa
5b. Flowers 3 to several in cymes branched to 2 or more orders; leaves acute to acuminate at apex
with tips sharp to slightly blunt.
6a. Corolla with tube 56 mm and lobes ca. 5 mm; leaves 1.55 cm wide, usually at least some
leaves wider than 3 cm ......................................................................................................................... 3. B. forrestii
6b. Corolla with tube ca. 7 mm and lobes ca. 8 mm; leaves 1.53 cm wide ...................................... 4. B. hainanensis
1. Benkara depauperata (Drake) Ridsdale, Reinwardtia 12:
298. 2008.

Randia evenosa Hutchinson in Sargent, Pl. Wilson. 3: 400.

1916; Oxyceros evenosus (Hutchinson) T. Yamazaki.

duo ci le qian

Shrubs, ca. 3 m tall; branches compressed to terete, puberulent to glabrous, with thorns 214 mm. Petiole 35 mm,
puberulent to glabrous; leaf blade drying thinly leathery to
stiffly papery and dark brown, obovate or elliptic, 26 1.53
cm, both surfaces glabrous, base cuneate to obtuse, apex obtuse
to rounded and sometimes shortly mucronate; secondary veins
35 pairs, in abaxial axils with foveolate and/or pilosulous domatia; stipules triangular to ovate, 2.53 mm, glabrous, acute.
Inflorescences 1- or 2-flowered, fasciculate, sometimes clustered on shortened internodes and appearing cymose, puberulent to glabrous; peduncles 510 mm, sometimes articulate near
base; bracts triangular, ca. 1 mm, ciliate, acute. Calyx glabrous;
ovary portion ellipsoid to obconic, 1.52 mm; limb with tube
23 mm; lobes triangular to linear, 0.51 mm. Corolla yellow,
glabrous outside; tube 35 mm; lobes oblong-elliptic, ca. 6 3
3.5 mm, obtuse. Fruiting pedicels 1017 mm. Berry globose, 5
7 mm in diam., glabrous. Fr. SepOct.

Randia depauperata Drake, J. Bot. (Morot) 9: 217. 1895;

Canthium spinosissimum Merrill; Fagerlindia depauperata
(Drake) Tirvengadum.
Shrubs, 13 m tall; branches compressed to terete, puberulent to hirtellous or strigillose, with thorns 415 mm. Petiole 2
6 mm, puberulent to hirtellous or strigillose; leaf blade drying
thinly papery, ovate, ovate-orbicular, lanceolate, or ovate-lanceolate, 18.2 0.83 cm, adaxially glabrous, abaxially glabrous or strigillose to hirtellous at least on principal veins, base
rounded to obtuse or broadly cuneate, apex acute to caudateacuminate; secondary veins 24 pairs, in abaxial axils usually
with pilosulous domatia; stipules triangular to narrowly triangular, 34 mm, strigillose to puberulent, acuminate to aristate. Inflorescences 1-flowered or cymose and 2- or 3-flowered, puberulent or strigillose to glabrescent; peduncles 310 mm, articulate when flowers solitary; bracts lanceolate, 23 mm; pedicels
610 mm. Calyx limb hirtellous to strigillose; ovary portion
obconic, 11.5 mm; limb with tube 3.54 mm; lobes triangular
to linear, 12 mm, apex acute. Corolla white, glabrous outside;
tube 34.5 mm; lobes elliptic-oblong to ligulate, 55.5 mm,
ciliolate, acute. Fruiting pedicels 515 mm. Berry globose, 56
mm in diam., pilosulous or strigillose to glabrescent; seeds
ellipsoid to angled, ca. 3 mm. Fl. Apr, fr. MayJan.
Forests or thickets on hills; below 100300 m. Fujian, Guangxi,
Hainan [Vietnam].

2. Benkara evenosa (Hutchinson) Ridsdale, Reinwardtia 12:

298. 2008.
wu mai le qian

Forests on mountains; 13001600 m. Yunnan.

3. Benkara forrestii (J. Anthony) Ridsdale, Reinwardtia 12:

299. 2008.
dian le qian
Randia forrestii J. Anthony, Notes Roy. Bot. Gard. Edinburgh 18: 204. 1934.
Shrubs or trees, 25 m tall; branches hirtellous or pilosulous to glabrescent, compressed to angled or subterete, with
thorns 310 mm. Petiole 2.510 mm, hirtellous or pilosulous to
glabrous; leaf blade drying papery to stiffly papery, brownish
green, oblong-ovate, elliptic-lanceolate, or ovate, 3.57 1.55
cm, adaxially glabrous and shiny, abaxially glabrous or sparsely
strigillose on principal veins, base cuneate to obtuse or sub-

80

RUBIACEAE

rounded, apex acute to acuminate; secondary veins 46 pairs, in

abaxial axils with foveolate and/or pilosulous domatia; stipules
lanceolate to triangular, 25 mm, strigillose to glabrous, acuminate. Inflorescences cymose, 2.53 34 cm, 5- to several
flowered, branched to several orders, pilosulous or hirtellous to
glabrous; peduncle 410 mm; bracts and bracteoles lanceolate
or triangular, 24 mm, acute; pedicels 13 mm. Calyx puberulent or strigillose to glabrous; ovary portion obconic, ca. 1 mm;
limb 23 mm, partially lobed; lobes triangular, 11.5 mm,
acute. Corolla white to greenish white or perhaps yellow, glabrous outside; tube 56 mm; lobes spatulate, ca. 5 mm, obtuse.
Fruiting pedicels ca. 6 mm. Berry globose, 58 mm in diam.,
glabrous. Fl. AprJun, fr. MayDec.
Forests or thickets at streamsides, on hills, or on mountain
slopes; 10002400 m. Yunnan.
Ridsdale (loc. cit.: 298299) treated Benkara griffithii and B.
forrestii as two different species, without commentary or a key, and
synonymized Randia hainanensis under B. forrestii. W. C. Chen (in
FRPS 71(1): 346. 1999) treated these names as synonyms of Oxyceros
griffithii. However, B. forrestii and R. hainanensis were recognized as
separate species by Tirvengadum (in herb.), and these appear morphologically distinct as outlined in the key to species above and thus are
separated here. These are provisionally treated as endemic pending
further study of this genus and specimens from surrounding countries.

4. Benkara hainanensis (Merrill) C. M. Taylor, comb. nov.

hai nan le qian
Basionym: Randia hainanensis Merrill, Lingnan Sci. J. 11:
58. 1932.
Shrubs, 0.52.5 m tall; branches slender, compressed to
subterete, glabrous, with thorns 47 mm. Petiole 2.513 mm,
pilosulous to glabrous; leaf blade drying papery, brownish
green to dark brown, oblong-ovate, elliptic-lanceolate, or lanceolate, 38 1.53 cm, both surfaces glabrous, base cuneate
or obtuse to sometimes subrounded, apex acute to acuminate;
secondary veins 36 pairs, in abaxial axils with foveolate and/or
pilosulous domatia; stipules lanceolate to triangular, 24 mm,
glabrous, acuminate. Inflorescences cymose, 2.53 34 cm,
3- to several flowered, branched to several orders, glabrous; peduncle 47 mm; bracts and bracteoles lanceolate to triangular,
26 mm, acute; pedicels 23 mm. Calyx glabrous; ovary portion obconic, ca. 1 mm; limb 13 mm, partially lobed; lobes triangular, 11.5 mm, acute. Corolla white or yellow, glabrous
outside; tube ca. 7 mm; lobes spatulate, ca. 8 mm, obtuse and
often mucronulate. Fruiting pedicels to 9 mm. Berry globose,
58 mm in diam., glabrous. Fl. AprJun, fr. MayDec.
Forests or thickets at streamsides, on hills, or on mountain
slopes; 200600 m. Hainan.
W. C. Chen (in FRPS 71(1): 346. 1999) treated this species as a
synonym of Benkara griffithii (as Oxyceros griffithii), and Ridsdale
(Reinwardtia 12: 299. 2008) treated it as a synonym of B. forrestii; see
comments above under B. forrestii regarding its separation here.

Randia rectispina Merrill, Lingnan Sci. J. 14: 60. 1935;

Oxyceros rectispinus (Merrill) T. Yamazaki.
Shrubs, 24 m tall, sometimes scandent; branches weakly
compressed to subterete, glabrous, rather stout, often yellowish
white, with thorns 614 mm. Petiole 36 mm, glabrous; leaf
blade drying leathery and usually straw- to brownish yellow,
ovate, elliptic-ovate, or elliptic, 1.56.5 13 cm, both surfaces
glabrous and rather shiny, base cuneate, obtuse, or subrounded,
apex obtuse or slightly rounded or rarely acute; secondary veins
46 pairs, in abaxial axils usually with foveolate and/or pilosulous domatia; stipules lanceolate to triangular, ca. 2 mm, acute
to caudate or aristate. Inflorescences 1- or less often 2-flowered,
glabrescent; peduncles 28 mm; bracteoles absent or 2, broadly
ovate-triangular, 1.52 mm, acute, after falling leaving a persistent pilose ring; pedicels 58 mm. Calyx limb glabrous; ovary
portion subglobose, ca. 2 mm; limb campanulate, ca. 4.5 3
mm, shallowly lobed; lobes triangular, 0.51.2 mm, cuspidate.
Corolla white, outside glabrous; tube 1822 mm; lobes ellipticovate, 56 mm, obtuse. Fruiting pedicels 512 mm. Berry globose, 610 mm in diam., glabrous; seeds 34 mm. Fl. AprJul,
fr. SepJan.
Forests or thickets at seasides or on hills; near sea level to 300
m. Hainan.

6. Benkara scandens (Thunberg) Ridsdale, Reinwardtia 12:

300. 2008.
nong zi mo li
Gardenia scandens Thunberg, Gardenia, 17. 1780; Fagerlindia scandens (Thunberg) Tirvengadum; Plectronia levinei
Merrill; Randia accedens Hance; R. scandens (Thunberg)
Lamarck.
Shrubs, 13 m tall; branches terete to flattened, glabrous,
with thorns 612 mm. Petiole 25 mm, glabrous; leaf blade
drying papery or thinly leathery, ovate, lanceolate, broadly
elliptic, or elliptic-oblong, 0.65.5 0.42.5 cm, both surfaces
glabrous, base cuneate, margins sometimes thinly revolute,
apex obtuse or acute; secondary veins 2 or 3 pairs, in abaxial
axils usually with foveolate and/or pilosulous domatia; stipules
triangular to ovate, 1.52 mm, glabrous, acuminate to cuspidate. Inflorescences 1-flowered or cymose and 2- or 3-flowered, glabrescent; peduncles 210 mm, articulate when flowers
solitary; bracts lanceolate, 1.52.5 mm; pedicels 310 mm.
Calyx glabrous; ovary portion obconic, 1.21.5 mm; limb with
tube somewhat campanulate, 3.54 mm; lobes narrowly triangular, 1.52 mm, acute to acuminate. Corolla white, outside puberulent to usually glabrous; tube 1420 mm; lobes lanceolate,
612 mm, acute. Fruiting pedicels 312 mm. Berry globose, 5
7 mm in diam., glabrous; seeds ellipsoid to angled, 23 mm. Fl.
MarMay, fr. MayDec.
Thickets in fields or on hills at low elevations. Guangdong,
Guangxi, Hainan, Yunnan [Vietnam].

5. Benkara rectispina (Merrill) Ridsdale, Reinwardtia 12: 299.

2008.

7. Benkara sinensis (Loureiro) Ridsdale, Reinwardtia 12: 300.

2008.

zhi ci le qian

le qian

RUBIACEAE

Oxyceros sinensis Loureiro, Fl. Cochinch. 1: 151. 1790;

Aidia sinensis (Loureiro) Masamune; Fagerlindia sinensis
(Loureiro) Tirvengadum; Randia sinensis (Loureiro) Schultes.
Shrubs or small trees, sometimes scandent, many
branched, 17 m tall; branches compressed to terete, rather
stout, yellowish brown hirtellous or -pilosulous, with thorns 4
15 mm. Petiole 515 mm, yellowish hirtellous to -pilosulous or
glabrescent; leaf blade drying papery to thickly papery, ovateelliptic, elliptic-oblong, or ovate, 221 1.59.5 cm, adaxially
glabrous, abaxially glabrescent to strigillose, hirtellous, or pilosulous at least on principal veins, base cuneate or slightly
rounded, apex acute or shortly acuminate; secondary veins 58
pairs, in abaxial axils with pilosulous domatia; stipules triangular to narrowly triangular, 35 mm, strigillose to pilosulous,
hirtellous, or glabrescent, long acuminate. Inflorescences congested-cymose, often umbelliform, several to many flowered,
2.54 34.5 cm, densely pilosulous to strigose or strigillose;

81

peduncle 35 mm; bracts lanceolate to ovate, 1.52.5 mm,

acute to acuminate; pedicels 0.51.5 mm. Calyx densely strigose to strigillose; ovary portion ellipsoid to cylindrical, 11.2
mm; limb 34 mm, shallowly to deeply lobed; lobes narrowly
triangular or ovate-triangular, 14 mm, acute. Corolla white or
yellow, outside puberulent to glabrous; tube (12)1524 14
mm; lobes elliptic-oblong to oblanceolate, 59 44.5 mm,
acute. Fruiting pedicels to 5 mm. Berry globose, 812 mm in
diam., pilosulous to strigillose or glabrous; seeds ca. 5 mm. Fl.
MarDec, fr. MayFeb.
Forests, forest margins, or thickets on hills, on mountains, or in
fields; near sea level to 1200 m. Fujian, Guangdong, Guangxi, Hainan,
Taiwan, Yunnan [Japan, ?Thailand (Larsen et al. 43560, MO!), Vietnam].
The shorter measurements here are taken from W. C. Chen (in
FRPS 71(1): 345346. 1999); the specimens studied all have corolla
tubes 1524 mm.

9. BRACHYTOME J. D. Hooker, Hookers Icon. Pl. 11: 70. 1871.

duan e chi mu shu
Chen Tao (); Charlotte M. Taylor
Shrubs or small trees, dioecious or perhaps polygamo-dioecious, unarmed, often with some internodes reduced. Raphides
absent. Leaves opposite or often appearing ternate due to reduced internodes and marked anisophylly grouping 2 leaves at 1 node
plus a single leaf at another node, apparently without domatia; stipules persistent or deciduous after terminal 24 nodes, interpetiolar
or shortly united around stem, triangular. Inflorescences pseudoaxillary, usually borne opposite a single leaf of an anisophyllous pair
and appearing leaf-opposed or sometimes just above an undeveloped internode at an apparent 3-leaved node, cymose, few to several
flowered, subsessile to pedunculate, bracteate. Flowers subsessile to pedicellate, unisexual or perhaps sometimes bisexual. Calyx
limb cupular, 5-lobed or -dentate. Corolla white to cream or pale yellow, funnelform to tubular or subrotate, glabrous inside; lobes 5,
convolute in bud. Stamens 5, inserted in corolla throat, exserted, with staminodes included; filaments short; anthers dorsifixed. Ovary
2-celled, ovules many in each cell on peltate axile placentas; stigma 2-lobed, grooved striate, partially exserted. Fruit red to orange,
baccate, fleshy, globose to ellipsoid, smooth, with calyx limb persistent, often with fruit base and/or pedicels elongating into stipes;
seeds numerous, small, cuneate to flattened, with testa reticulate; endosperm fleshy; embryo small, subterete.
About five species: Bangladesh, Cambodia, China, India, Malaysia, Myanmar, Vietnam; three species in China.
The morphology and branching of this genus were studied by Tirvengadum and Sastre (Bull. Mus. Natl. Hist. Nat., B, Adansonia 8: 257296.
1986).
Brachytome was described as polygamo-dioecious in FRPS (71(1): 360. 1999), but this has not been reported by other sources; the genus was
described as unisexual by Puff et al. (Rubiaceae of Thailand, 68. 2005).

1a. Branches densely strigose, strigillose, hirtellous, or hispidulous ................................................................................. 2. B. hirtellata

1b. Branches glabrous or sparsely to moderately strigose to strigillose when young, becoming glabrescent with age.
2a. Infructescences 38 35 cm, with axes and pedicels mostly well developed; stipules 38 mm; berry
subglobose, ca. 6 5 mm .................................................................................................................................. 1. B. hainanensis
2b. Inflorescences and infructescences 35 35 cm, with axes and pedicels short to well developed;
stipules 615 mm; berry ellipsoid, 1020 815 mm ........................................................................................... 3. B. wallichii
1. Brachytome hainanensis C. Y. Wu ex W. C. Chen, Guihaia
7: 298. 1987.
hai nan duan e chi mu
Shrubs, 2.53 m tall; branches flattened becoming subterete, glabrous. Petiole 0.41.5 cm, glabrous; leaf blade drying
papery and usually brown, elliptic-oblong, elliptic, or ellipticlanceolate, 720.5 2.57 cm, adaxially glabrous, abaxially
glabrous or puberulent, base cuneate to obtuse, apex acute to
acuminate; secondary veins 812 pairs; stipules triangular to

broadly triangular, 38 mm, glabrous, acuminate or cuspidate.

Inflorescences not seen. Flowers not seen. Infructescences corymbiform-cymose, 38 35 cm, several to many fruited; peduncle 1.52 cm; bracts triangular to ovate, 0.51 mm, mostly
situated at nodes (i.e., branching points); pedicels 39 mm.
Berry red, ellipsoid to subglobose except with narrow cylindrical base, rounded portion 56 ca. 5 mm plus basal narrow
portion ca. 1 mm; seeds yellow, ca. 1.5 mm wide. Fr. Mar.
Forests. Hainan [Vietnam].

RUBIACEAE

82

2. Brachytome hirtellata Hu, Bull. Fan Mem. Inst. Biol. 10:

164. 1940.

2a. Brachytome hirtellata var. glabrescens W. C. Chen, Acta

Phytotax. Sin. 22: 147. 1984.

dian duan e chi mu

shu mao duan e chi mu

Shrubs, ca. 3 m tall; branches somewhat flattened becoming angled and sometimes shallowly channeled, sometimes
flexuous, densely hirtellous or hispidulous to strigillose or strigose. Petiole 212 mm, densely hirtellous to strigillose or strigose; leaf blade drying membranous or thinly papery, green to
yellowish green adaxially, brown or purplish brown abaxially,
oblanceolate or oblong-lanceolate to elliptic, 721.5 26 cm,
adaxially glabrous or scaberulous to sparsely hirtellous, hispidulous, or strigillose, abaxially glabrescent or densely to sparsely
hirtellous or strigillose, base cuneate or acute, apex acute to
acuminate; secondary veins 1220 pairs; stipules ovate to triangular, 48 mm, densely strigillose or strigose to glabrous, acute
to acuminate. Inflorescences congested-cymose, 11.5 11.5
cm, usually 510-flowered, densely hirtellous to strigose or
strigillose, sessile or subsessile; bracts ovate to triangular or
linear, 0.58 mm, situated at nodes, along axes, and usually in
pairs and 0.52.5 mm immediately below flowers. Flowers
sessile to shortly pedicellate, with pedicels to 1 mm. Calyx
strigillose to pilosulous; ovary portion obconic, 0.51.5 mm;
limb 22.5 mm, lobed for ca. 1/2 its length; lobes triangular to
linear-lanceolate, acute. Corolla white or yellowish white, funnelform to tubular, outside glabrous; tube 46 mm; lobes ovate,
12 mm. Berry globose, ca. 8 mm in diam., densely hirtellous
or hispidulous to strigillose, with stipe or pedicel elongating
above paired bracts, up to 12 mm; seeds reddish yellow, ca. 1.5
11.5 mm. Fl. MarJun, fr. MayMar.

Leaf blade adaxially smooth and glabrous or subglabrous,

abaxially subglabrous or sparsely hirtellous or strigillose with
pubescence denser along veins. Fl. MayJun, fr. JunMar.

Forests or thickets at streamsides in valleys; 4002200 m. Xizang,

Yunnan [Vietnam].
These varieties were recognized by W. C. Chen in FRPS (71(1):
362. 1999); some recently collected specimens seem to be intermediate
but this species is poorly documented in general, so these are treated
here at least for reference.

1a. Leaf blade adaxially smooth and

glabrous or subglabrous, abaxially
subglabrous or sparsely hirtellous or
strigillose with pubescence denser
along veins .......................................... 2a. var. glabrescens
1b. Leaf blade adaxially scaberulous and
glabrous to sparsely hirtellous,
strigillose, or hispidulous, abaxially
densely grayish brown hirtellous
to strigillose ............................................. 2b. var. hirtellata

Forests at streamsides in valleys; 4002200 m. Xizang (Mdog),

Yunnan [Vietnam].

2b. Brachytome hirtellata var. hirtellata

() dian duan e chi mu (yuan bian zhong)
Leaf blade adaxially scaberulous and glabrous to sparsely
hirtellous, strigillose, or hispidulous, abaxially densely grayish
brown hirtellous to strigillose. Fl. MarJun, fr. MayDec.
Forests or thickets at streamsides in valleys; 7001600 m. Yunnan.

3. Brachytome wallichii J. D. Hooker, Hookers Icon. Pl. 11:

70. 1871.
duan e chi mu
Shrubs or small trees, 1.53 m tall; branches flattened to
subterete, glabrous or sparsely strigillose or strigose becoming
glabrescent with age. Petiole 0.31 cm, glabrous; leaf blade
drying papery, green adaxially, often flushed with reddish
brown abaxially, elliptic-oblong, lanceolate, or oblanceolate, 9
14 2.54.5 cm, glabrous on both surfaces, base cuneate to
acute, apex acuminate; secondary veins 810 pairs; stipules triangular to narrowly triangular, 615 mm, glabrous to puberulent, acute to acuminate and sometimes falcate. Inflorescences
34 35 cm, several flowered, glabrous; peduncle ca. 0.5 cm;
bracts triangular and often fused in pairs, 12 mm, usually situated at nodes and perhaps immediately below flowers; pedicels 510 mm. Calyx glabrous; ovary portion obconic, 11.5
mm; limb 11.5 mm, denticulate to lobed for up to 1/3 of its
length; lobes broadly triangular. Corolla funnelform, outside
glabrous; tube ca. 6 mm; lobes triangular, ca. 1 mm, obtuse.
Berry red, ellipsoid, 1020 815 mm, with stipe or pedicel
elongating, up to 20 mm; seeds 11.5 12 mm. Fr. SepOct.
Forests in valleys; 12002000 m. SW Yunnan [Bangladesh, Cambodia, India, Myanmar, Vietnam].
This species was reported by W. C. Chen (in FRPS 71(1): 361.
1999) from Bhutan, but Springate et al. (Fl. Bhutan 2(2): 784. 1999)
treated this as reported in error.

10. CAELOSPERMUM Blume, Bijdr. 994. 18261827.

xue guo mu shu
Chen Tao (); Charlotte M. Taylor
Lianas or sometimes shrubs or small trees, unarmed. Raphides present. Leaves opposite or rarely ternate, decussate on
ascending branches but distichous on horizontal branches, with domatia; stipules persistent at least on apical 2 or 3 nodes and usually
becoming hardened with age, at least shortly united around stem [or rarely interpetiolar], triangular and acute to truncate, sometimes
shortly 2-toothed. Inflorescences terminal and sometimes in axils of uppermost leaves, corymbiform to paniculiform with flowers
grouped in small heads or umbelliform cymes, many flowered, pedunculate, bracteate or bracts reduced. Flowers pedicellate to
sessile, free [sometimes fused by their ovaries], bisexual, apparently monomorphic. Calyx with ovary portion usually hemispherical

RUBIACEAE

83

or campanulate; limb tubular, truncate or 46-denticulate. Corolla white becoming yellowed when old, salverform or funnelform,
fleshy, variously pubescent inside; lobes 4 or 5(or 6), valvate in bud. Stamens 4 or 5(or 6), inserted in corolla tube, exserted; filaments developed; anthers dorsifixed near middle, 2-parted at base. Ovary 2-celled with ovules 2 in each cell or 4-celled via secondarily developing septa and ovules 1 in each cell, ovules inserted on middle of septum, anatropous or pendulous; stigmas 2, linear to
spatulate, exserted or included. Fruit simple [sometimes multiple due to fusion of flowers], dark red to dark purple, drupaceous,
fleshy, subglobose to ellipsoid, smooth, with calyx limb persistent; pyrenes 2, 3, or usually 4, 1-celled with 1 seed in each cell, ellipsoid to plano-convex, cartilaginous, densely puberulent; seeds medium-sized; endosperm oily; embryo straight.
Seven to ten species: from tropical Asia to Australia; one species in China.
This genus is similar and has long been considered related to Morinda and perhaps will be synonymized eventually, but it has provisionally been
maintained by recent workers (Razafimandimbison et al., Molec. Phylogen. Evol. 48: 207223. 2008). This genus was studied by Johansson (Blumea
3: 265297. 1988), who synonymized the two species recognized by Y. Z. Ruan (in FRPS 71(2): 166167. 1999) in China together with several other
SE Asian names into a wide-ranging, morphologically variable species with the variation apparently continuous.

1. Caelospermum truncatum (Wallich) Baillon ex K. Schumann in Engler & Prantl, Nat. Pflanzenfam. 4(4): 136. 1891.
xue guo mu
Webera truncata Wallich in Roxburgh, Fl. Ind. 2: 538.
1824; Caelospermum kanehirae Merrill; C. morindiforme Pierre
ex Pitard; C. scandens Blume.
Lianas or weak shrubs, to 15 m high; branches weakly
flattened to subterete or quadrangular, glabrous or glabrescent.
Leaves opposite; petiole 1025 mm, glabrous to puberulent;
blade drying leathery or thickly papery and brownish yellow to
pale black, elliptic, elliptic-oblong, lanceolate-oblong, ovate, or
obovate, 715 310 cm, adaxially glabrous, abaxially paler
and glabrous to puberulent, base cuneate to rounded, apex
acute, obtuse, or rounded; secondary veins 47 pairs, with pilosulous domatia; stipules broadly triangular, 14.5 mm, truncate or 2-denticulate. Inflorescence a cymose panicle of 39

corymbs or umbels, 617 cm, mealy puberulent; peduncles 2.5

4 cm; umbels 320-flowered; bracts reduced. Flowers free,
subsessile to pedicellate, pedicels to 6(11) mm. Calyx mealy
puberulent to glabrescent; ovary portion subglobose to cupuliform, 11.5 mm; limb 12 mm, truncate or 4- or 5-denticulate.
Corolla salverform, outside glabrous to sparsely papillose; tube
(4)56(9) mm, pubescent in upper part and throat; lobes
linear to oblong-linear, (4.5)56(12) mm. Drupes subglobose,
812(27) mm in diam. Fl. AprMay, fr. JulSep.
Thickets or sparse forests on hills or mountains; sea level to 1900
m. Guangxi, Hainan [Cambodia, Indonesia (Borneo, Java, Sumatra),
Malaysia, Thailand, Vietnam].
Measurements in parentheses here come from Johanssons
(Blumea 3: 265297. 1988) description of this species throughout its
range. He noted that the most widely used name for this species is
Caelospermum scandens, here included as an aid to relating the Chinese
species to the SE Asian flora.

11. CANTHIUM Lamarck, Encycl. 1: 602. 1785.

yu gu mu shu
Chen Tao (); Charlotte M. Taylor, Henrik Lantz
Meyna Roxburgh ex Link.
Shrubs or small trees, sometimes dioecious, sometimes with short shoots, unarmed or sometimes with paired straight supraaxillary thorns. Raphides absent. Leaves opposite, with or without domatia; stipules persistent or caducous, interpetiolar, shortly
united around stem, or fused to petiole bases, triangular, internally (i.e., adaxially) sometimes sericeous to pilose. Inflorescences axillary, cymose to fasciculate, few to several flowered, sessile to pedunculate, bracteate with bracts sometimes fused in calyculate pairs.
Flowers subsessile to pedicellate, bisexual and monomorphic (Canthium s.s.) or sometimes unisexual. Calyx with ovary portion often
subglobose to hemispherical; limb very short, truncate or 4- or 5-lobed. Corolla green to white or pale yellow, tubular, urceolate, or
funnelform, with tube often constricted at top, inside variously pubescent but usually with ring of introrse hairs in tube; lobes 4 or 5,
often long acuminate or aristate at apex, in bud valvate and often with apices held erect and pressed together forming apiculate projection, at anthesis notably reflexed. Stamens 4 or 5, inserted at corolla throat, partially to fully exserted; filaments short or reduced;
anthers dorsifixed near base, elliptic to ovate, at anthesis reflexed. Ovary 25-celled, ovules 1 in each cell, pendulous from apical
placenta; stigma included or exserted, capitate to cupular, entire to variously lobed, usually with style attachment recessed. Fruit
brown, yellow, orange, or red, drupaceous, subglobose, ellipsoid, or often dicoccous when fully developed or reniform with only 1
seed, fleshy, with calyx limb caducous or infrequently persistent; pyrenes 25, 1-celled with 1 seed in each cell, ellipsoid to reniform,
bony or crustaceous; seeds medium-sized to large, ellipsoid, cylindrical, or plano-convex; testa membranous; endosperm fleshy;
radicle ascending.
About 30 species: tropical and subtropical Africa and Asia; four species (one endemic) in China.
The circumscription and separation of genera of Vanguerieae and especially Canthium, Meyna, Psydrax, Pyrostria Commerson ex Jussieu,
Vangueria Jussieu, and their closely related genera are problematic, controversial, and far from a full resolution. Lantz and Bremer (Bot. J. Linn. Soc.
146: 257283. 2004) provided revised circumscriptions of several genera; in particular, they synonymized Meyna with Canthium and moved some
species from Canthium to Psydrax. The Asian lineages have not yet been studied deeply, and the Chinese species may yet be reclassified.

RUBIACEAE

84

The name Canthium cavaleriei H. Lveill (Repert. Spec. Nov. Regni Veg. 10: 434. 1912) is based on a specimen from Guizhou; this name was
not treated in FRPS. As the specimen has not been seen and its description is scanty, its identity is unknown. It has been suggested to be related to
Lasianthus, but according to Lauener and Ferguson (Notes Roy. Bot. Gard. Edinburgh 32: 103. 1972) that is incorrect.

1a. Plants unarmed or usually armed with thorns; flowers borne in fascicles on peduncles 12 mm ............................. 3. C. horridum
1b. Plants without thorns; flowers borne in fascicles on peduncles 48 mm, or in cymes with peduncles 825 mm
and pedicels 28 mm.
2a. Leaves 919 48.5 cm; inflorescences corymbiform to cymose, 2.53 ca. 2 cm in flower, to 7 9 cm in
fruit; fruit 1020 915 mm ....................................................................................................................................... 4. C. simile
2b. Leaves 39 1.55 cm; inflorescences fasciculate to subumbelliform, 13.5 cm in flower and fruit; fruit
810 mm in diam.
3a. Flowers unisexual on dioecious plants; secondary leaf veins 3 or 4 pairs; stipules persistent; corolla
tube ca. 1.5 mm and lobes 26 mm; ovary locules and pyrenes 2 .......................................................... 1. C. gynochthodes
3b. Flower bisexual; secondary leaf veins 5 or 6 pairs; stipules caducous; corolla tube ca. 3 mm and
lobes ca. 3 mm; ovary locules and pyrenes 3 or 4 ....................................................................................... 2. C. hainanense
1. Canthium gynochthodes Baillon, Adansonia 12: 199. 1878.
po lai mu
Canthium cumingii Vidal; Plectronia cumingii (Vidal)
Merrill; P. gynochthodes (Baillon) Merrill; P. moluccana
Merrill; P. umbellata K. Schumann.
Small trees, height not noted, often rather succulent;
branches flattened becoming subterete, glabrous. Petiole 15
mm, glabrous; leaf blade drying thinly leathery, elliptic to elliptic-oblong or obovate, 38 1.55 cm, glabrous on both
surfaces, base acute to obtuse, apex obtuse to shortly acute;
secondary veins 3 or 4 pairs, in abaxial axils with foveolate and
sometimes also pilosulous domatia; stipules persistent, shortly
united around stem, triangular to ovate, 24.5 mm, glabrous,
aristate to cuspidate. Inflorescences glabrous, 13.5 cm, several
flowered; peduncles 0.82.5 cm, terminating in a pair of bracteoles, these triangular, ca. 1 mm, shortly fused; pedicels 47
mm. Flowers unisexual on dioecious plants. Calyx glabrous;
ovary portion of pistillate flowers ca. 1.2 mm; limb ca. 0.5 mm,
truncate to undulate. Corolla white to pale green, outside glabrous; tube ca. 1.5 mm; lobes 4, 22.5(6) mm, acute. Ovary 2celled; stigma ca. 0.3 mm. Drupes orangish yellow, oblate or
obovoid to dicoccous, ca. 8 10 mm, smooth, glabrous, with
calyx limb persistent; pyrenes 2. Fl. Aug.
Forests on mountain ridges. Taiwan [Philippines].
Only very young flower buds have been seen so far from China;
the descriptions here of the mature flowers and the fruit are based in
large part on the description by Liu and Yang (Fl. Taiwan, ed. 2, 4: 247
248. 1998) and on Philippine specimens. The synonymy here is taken
from Merrill (Enum. Philipp. Pl. 3: 536537. 1923). The accepted
species epithet was incorrectly spelled as gynochodes by Liu and
Yang (loc. cit.: 247).
Canthium dicoccum var. obovatifolium should probably be compared with this species. The variety is poorly known and is here included with the species in which it was described, now called Psydrax
dicocca; but its inclusion in that species is questionable, and its description suggests it is very similar to C. gynochthodes.

2. Canthium hainanense (Merrill) Lantz, comb. nov.

qiong mei
Basionym: Meyna hainanensis Merrill, Lingnan Sci. J. 14:
57. 1935.

Small trees, 36 m tall; branches weakly flattened to subterete, glabrous or puberulent, at nodes with stipule bases together with bases of petioles usually markedly thickened with
thickened portion sometimes tardily splitting interpetiolarly,
usually with stout lateral short shoots 0.51 cm, these densely
covered with scalelike old stipule bases. Leaves borne at nodes
near apex of developed stems or clustered on short shoots;
petiole 58 mm, glabrous, sometimes weakly articulate near
base; blade drying papery or thinly papery, ovate, oblong-ovate,
or elliptic-oblong, 39 1.55 cm, adaxially glabrous or
rarely strigillose, abaxially strigillose or strigose, base obtuse
to rounded, apex acute or acuminate; secondary veins 5 or 6
pairs, in abaxial axils with pilosulous domatia; stipules shortly
united around stem with basal portion persistent becoming
thickened and upper portion caducous, triangular to lanceolate,
25 mm, often keeled, acuminate to aristate. Inflorescences
fasciculate to subumbelliform and subsessile, 11.5 cm, few to
several flowered, puberulent or hirtellous to glabrescent; peduncles 48 mm; bracts reduced. Calyx puberulent; ovary portion
obconic, ca. 0.5 mm; limb with lobes (4 or)5, triangular, 0.50.8
mm, acute. Corolla color not noted, funnelform, outside glabrous; tube ca. 3 mm, inside pubescent in upper half; lobes (4
or)5, lanceolate to spatulate-triangular, ca. 3 mm, acute to acuminate. Ovary 3- or 4-celled; stigma ca. 0.8 mm. Drupes with
color not noted, subglobose, 810 mm in diam., puberulent to
glabrescent, with calyx limb persistent; pyrenes 3 or 4. Fl. Apr
Aug, fr. Jun.
Broad-leaved forests; 200300 m. Hainan.
The illustration of this species presented by W. C. Ko (in FRPS
71(2): 8, t. 3. 1999) shows four calyx lobes and four corolla lobes;
however, these structures both were described as five in the protologue,
and the specimens seen have five calyx lobes.

3. Canthium horridum Blume, Catalogus, 45. 1823.

zhu du mu
Canthium hebecladum Candolle; C. pauciflorum Blanco;
Plectronia horrida (Blume) Bentham & J. D. Hooker.
Shrubs, 23 m tall; branches flattened to subterete, often
rather slender, strigillose to strigose or pilosulous, sometimes
with lateral short shoots to 0.5 cm; thorns slender to stout, 330
mm, straight, or sometimes absent. Leaves paired along developed stems or sometimes clustered on lateral short shoots;

RUBIACEAE

petiole 23 mm, strigillose to pilosulous; blade drying papery,

ovate to lanceolate or elliptic, 26 13.5 cm, adaxially glabrous or strigillose to hispidulous along principal veins or
throughout, abaxially puberulent to strigillose or hispidulous,
base rounded or obtuse, apex obtuse, acute, or weakly acuminate; secondary veins 2 or 3 pairs, in abaxial axils with pilosulous and/or foveolate domatia; stipules deciduous after distalmost several nodes, triangular to ovate, 23 mm, densely
strigillose to strigose, acute. Inflorescences fasciculate, 0.51
cm, few flowered, strigillose to glabrescent; peduncles 13
mm, each with a pair of triangular bracteoles ca. 1 mm and
fused in pairs. Calyx puberulent to glabrous; ovary portion obconic, ca. 0.8 mm; limb 0.50.8 mm, truncate to undulate. Corolla white, funnelform, outside glabrous; tube suburceolate,
ca. 2 mm; lobes 5, triangular-oblong, ca. 3 mm, acuminate.
Ovary 2-locular; stigma ca. 0.5 mm. Drupes yellow, ovoid to
subglobose, obovoid, or somewhat dicoccous, laterally somewhat flattened, 1525 1020 mm, smooth, glabrous, with calyx limb persistent; pyrenes 2, weakly tuberculate. Fl. AprJun,
fr. JulNov.
Thickets at low elevations; near sea level to 500 m. Guangdong,
Guangxi, Hainan, Yunnan [India, Malaysia, Thailand, Vietnam].

85

branches glabrous, somewhat compressed becoming terete.

Leaves borne along developed stems; petiole 58 mm, glabrous; blade drying thinly to thickly papery, ovate to ovate-oblong or elliptic-oblong, 919 48.5 cm, adaxially shiny and
glabrous or sometimes pustulose, abaxially glabrous to pilosulous or pilose, base obtuse to rounded, apex shortly acuminate;
secondary veins 68 pairs, in abaxial axils with small foveolate
domatia; stipules deciduous, shortly united around stem, triangular, 28 mm, sparsely pilosulous to glabrous, keeled, cuspidate to acuminate. Inflorescences corymbiform to cymose, 2.5
3 ca. 2 cm, several flowered, puberulent to hirtellous; peduncle 1014 mm; bracts reduced; pedicels 23 mm. Calyx puberulent to glabrous; ovary portion obconic to hemispherical, 11.5
mm; limb ca. 0.5 mm, 5-lobed; lobes broadly ovate-triangular,
acute. Corolla white, urceolate-tubular, outside glabrous; tube
1.52 mm, inside villous in throat; lobes 5, triangular to spatulate, 1.53 mm, acute. Ovary 2-locular; stigma ca. 1.2 mm. Infructescences expanded, to 7 9 cm; fruiting pedicels to 12
mm. Drupes with color not noted, obovoid, laterally compressed, often somewhat dicoccous, 1020 915 mm, glabrescent, with calyx limb deciduous; pyrenes 2, plano-convex.
Fl. JanMar, fr. JunJul.

The name Canthium parvifolium Roxburgh has sometimes been

applied to the Chinese plants that are here called C. horridum. W. C. Ko
(in FRPS 71(2): 10. 1999) considered it a misapplied name, listing it as
Canthium parvifolium auct. non Roxb.

Broad-leaved forests at low to middle elevations; 2001500 m.

Guangdong, Guangxi, Hainan, Yunnan [Vietnam].

4. Canthium simile Merrill & Chun, Sunyatsenia 2: 19. 1934.

The measurements of the flower parts given by Merrill and Chun

in the protologue do not agree with the measurements inferred from
their figure 2; their figure seems to have been reproduced at twice the
size they planned.

da ye yu gu mu
Erect shrubs to small trees, 410(18) m tall, unarmed;

This species is newly reported here from Vietnam, based on

Thorel 1294 (GH).

12. CATUNAREGAM Wolf, Gen. Pl. 75. 1776.

shan shi liu shu
Chen Tao (); Charlotte M. Taylor
Xeromphis Rafinesque.
Shrubs or small trees, often with short shoots, often armed with spines or spinescent short shoots. Raphides absent. Leaves
opposite or often clustered and apparently fasciculate on short shoots, subsessile to petiolate, usually with domatia; stipules deciduous or caducous, interpetiolar, triangular. Inflorescences terminal on short shorts or these sometimes reduced giving an appearance of
axillary position, 1-flowered, or cymose to fasciculate and 26-flowered, sessile to pedunculate, bracteate or bracts reduced. Flowers
subsessile to pedicellate, bisexual, monomorphic. Calyx limb deeply 5-lobed, lobes spatulate to obovoid. Corolla white to cream or
pale green, campanulate to subrotate, usually densely sericeous outside, variously pubescent inside; lobes 5[10], convolute in bud.
Stamens 5, inserted in corolla throat, partially to fully exserted; filaments short; anthers dorsifixed. Ovary 2-celled, ovules many in
each cell on axile placentas attached at center of septum; stigma ellipsoid or 2-lobed, exserted. Fruit generally yellowish brown,
baccate, leathery to thickly fleshy or infrequently hard, globose, ellipsoid, or ovoid-globose, with calyx limb persistent; seeds numerous, medium-sized, ellipsoid, angled, or reniform, embedded in fleshy or mucilaginous pulp.
About five to perhaps ten species: widespread in Africa, also in Bhutan, China, Cambodia, India, Indonesia, Kashmir, Laos, Madagascar, Malaysia, Myanmar, Nepal, Pakistan, Sri Lanka, Thailand, Vietnam; one species in China.
The calyx limb enlarges markedly during the development of the fruit. At least some species of Catunaregam in Africa have nocturnal sweetly
fragrant flowers (Bridson & Verdcourt, Fl. Trop. E. Africa, Rub. (Pt. 2), 496500. 1988), and Catunaregam in China may also. Many species of
related genera also have secondary pollen presentation, and these features may be present in Catunaregam also.

1. Catunaregam spinosa (Thunberg) Tirvengadum, Bull. Mus.

Natl. Hist. Nat., Sr. 3, Bot. 35: 13. 1978.
shan shi liu

Gardenia spinosa Thunberg, Gardenia, 16. 1780; G. dumetorum Retzius; Randia dumetorum (Retzius) Lamarck; Randia
spinosa (Thunberg) Poiret; Xeromphis retzii Rafinesque; X.
spinosa (Thunberg) Keay.

RUBIACEAE

86

Shrubs or small trees, 110 m tall; branches rather stout,

hirsute, pilose, pilosulous, or puberulent to glabrescent, armed
with axillary stout paired thorns 15 cm. Petiole 28 mm,
pilose, pilosulous, or hirtellous to glabrous; leaf blade drying
papery or subleathery, obovate or oblong-obovate or rarely
ovate to spatulate, 1.811 15.7 cm, both surfaces glabrous to
strigillose, strigose, hirtellous, or sparsely hirsute at least along
principal veins, base cuneate and sometimes decurrent, margins
entire or often shortly ciliate, apex acute; secondary veins 47
pairs, often with pilosulous domatia in abaxial axils; stipules
caducous, ovate to broadly triangular, 34.5 mm, acute to
aristate. Inflorescences terminal on lateral short shoots together
with tufted leaves, 13-flowered; pedicels 25 mm, brown villous or -hirtellous. Calyx brown villous, -hirtellous, or -strigose;
ovary portion ovoid to ellipsoid, 3.57 45.5 mm; limb
slightly dilated, deeply lobed; lobes broadly elliptic to oblan-

ceolate or obovoid, 58 36 mm, acute to rounded. Corolla

white, becoming pale yellow with age, campanulate; tube 56
mm, sparsely villous in throat; lobes ovate or ovate-oblong, 6
11 (5.5)89 mm, spreading, rounded to subtruncate. Anthers
ca. 3 mm, fully exserted. Style 46 mm; stigma fusiform, with 2
coherent lobes, ca. 2 mm. Berry globose, 24 cm in diam., glabrous or sparsely pilose or strigose; seeds 45 mm. Fl. Mar
Jun, fr. MayJan.
Thickets or forests at streamsides, on hills or mountain slopes, or
in valleys or fields; near sea level to 1600 m. Fujian, Guangdong,
Guangxi, Hainan, Taiwan, Yunnan [Cambodia, India, Indonesia, Kashmir, Laos, Malaysia, Myanmar, Nepal, Pakistan, Sri Lanka, Thailand,
Vietnam; Africa, Madagascar].
W. C. Chen (in FRPS 71(1): 338340. 1999) noted that this
species is sometimes climbing, but this has not been noted by any other
authors nor seen on specimens.

13. CEPHALANTHUS Linnaeus, Sp. Pl. 1: 95. 1753.

feng xiang shu shu
Chen Tao (); Charlotte M. Taylor
Shrubs or trees, unarmed; buds conical. Raphides absent. Leaves verticillate or opposite, distichous, usually with domatia;
stipules persistent [or sometimes caducous], interpetiolar, triangular, at apex sometimes with a black gland. Inflorescences terminal
and sometimes in axils of uppermost leaves, capitate with several globose heads, many flowered, pedunculate, bracteate; bracteoles
clavate to clavate-spatulate. Flowers sessile, bisexual, monomorphic. Calyx limb 4(or 5)-lobed. Corolla white to cream, salverform to
funnelform, variously pubescent inside; lobes 4, imbricate (and quincuncial) in bud. Stamens 4, inserted in corolla throat, partially
exserted; filaments short; anthers dorsifixed, bifid at base. Ovary 2-celled, ovules 1 in each cell, apical and pendulous, anatropous;
stigma clavate to capitate, exserted. Fruiting heads globose. Fruit schizocarpous, obconic to turbinate, dry, with calyx limb persistent;
mericarps 2, indehiscent, with 1 seed, obconic, stiffly papery; seeds medium-sized, oblong-ellipsoid, with white spongy aril.
Six species: three in the Americas, two in Asia, one in Africa; one species in China.

1. Cephalanthus tetrandrus (Roxburgh) Ridsdale & Bakhuizen f., Blumea 23: 182. 1976 [tetrandra].
feng xiang shu
Nauclea tetrandra Roxburgh, Fl. Ind. 2: 125. 1824;
Cephalanthus glabrifolius Hayata; C. naucleoides Candolle; C.
ratoensis Hayata.
Deciduous shrubs or small trees, 15 m tall; branches subangled to terete, densely pilosulous or tomentulose to usually
glabrous. Leaves opposite or in whorls of 35, with number
often irregular on a stem; petiole 510 mm, densely tomentulose or pilosulous to usually glabrous; blade drying stiffly papery to subleathery, ovate to ovate-lanceolate, elliptic, or elliptic-oblong, 6.515 36 cm, adaxially glabrous to sparsely
puberulent and often rugulose, abaxially densely pilosulous to
usually glabrous, base obtuse, rounded, or subcordate, apex
acute to acuminate; secondary veins 812 pairs, often with pilosulous domatia in axils or along costa; stipules persistent or
often deciduous leaving a persistent truncate basal portion,
broadly ovate, 35 mm, adaxially enclosing a ring of persistent
white trichomes 0.51 mm, abaxially glabrous to occasionally
densely pilosulous, apex cuspidate, often with a black terminal
gland. Inflorescence densely tomentulose or pilosulous to usu-

ally glabrous; peduncles 2.56 cm; flowering heads 110, 812

mm in diam. across calyces, 2025 mm in diam. across corollas; bracteoles linear-clavate, ca. 2 mm, strigose. Calyx with
ovary portion obconic, ca. 1 mm, glabrous, surrounded at base
by ring of pilose trichomes; limb with tubular portion 12 mm,
glabrous to sparsely puberulent; lobes spatulate-oblong, 12
mm, densely puberulent, obtuse to rounded, in sinuses usually
with stipitate or sessile black gland. Corolla outside glabrous;
tube 712 mm, pubescent inside; lobes spatulate-oblong, 12
mm, obtuse or rounded, in sinuses often with stipitate black
gland. Stigma ellipsoid or clavate, ca. 0.5 mm, exserted by 46
mm. Fruiting head 1020 mm in diam. Mericarps 46 mm, glabrous; seeds brown, 35 mm. Fl. JunSep, fr. JulSep.
Shady sites at roadsides or streamsides; sea level to 700 m. Fujian,
Guangdong, Guangxi, Hainan, Hunan, Jiangxi, Taiwan, Yunnan, Zhejiang [Bangladesh, India, Laos, Myanmar, Thailand, Vietnam].
This species was long considered conspecific with Cephalanthus
occidentalis Linnaeus of North America; consequently, that name has
been frequently but erroneous used for the Asian plants. Ridsdale
(Blumea 23: 177188. 1976) concluded that only one species of Cephalanthus is found in Asia and it is distinct from the North American
species. Ridsdale (loc. cit.: 180181) reported that this species is tolerant of wet soils and sometimes cultivated for ground stabilization.

RUBIACEAE

87

14. CERISCOIDES (Bentham & J. D. Hooker) Tirvengadum, Bull. Mus. Natl. Hist.
Nat., Sr. 3, Bot. 35: 13. 1978.
mu gua lan shu
Chen Tao (); Charlotte M. Taylor
Gardenia sect. Ceriscoides Bentham & J. D. Hooker, Gen. Pl. 2: 90. 1873.
Shrubs or small trees, dioecious [or polygamo-dioecious], often with short shoots, unarmed or with spines or spinescent short
shoots. Raphides absent. Leaves opposite but usually crowded on short shoots and apparently fasciculate, sometimes with domatia;
stipules caducous, interpetiolar, triangular. Inflorescences terminal on lateral branches or short shoots [sometimes cauliflorous],
sessile to pedunculate, bracteate, staminate 2-flowered or 2- or 3-flowered and cymose, pistillate 1-flowered. Flowers sessile to pedicellate, unisexual [sometimes bisexual and monomorphic]. Calyx with ovary portion hemispherical to ellipsoid in pistillate flowers,
markedly turbinate with base narrowed in staminate flowers; limb subtruncate to 5(7)-lobed. Corolla white to pale green, tubularcampanulate or funnelform, glabrous inside; lobes 5(7), convolute in bud. Stamens 5(7), inserted in corolla throat, included,
staminodes markedly reduced; filaments short; anthers dorsifixed. Ovary 1-celled, ovules numerous in each cell on 24[6] parietal
placentas; stigmas 2[6]-lobed, included. Fruit brown, baccate, fleshy, globose to ellipsoid, smooth, with calyx limb persistent; seeds
numerous, large, ellipsoid to lenticular, embedded in fleshy pulp.
About 11 species: China, India, Indonesia, Laos, Malaysia, Myanmar, Philippines, Sri Lanka, Thailand, Vietnam; one species (endemic) in
China.
Ceriscoides was recently studied by Azmi (Harvard Pap. Bot. 7(2): 443464. 2003). H. S. Lo (in FRPS 71(1): 337. 1999) described the
placentas as 24 and the stigmas of our species as 2, but Azmi (loc. cit.: 445) described both of these as 36.

1. Ceriscoides howii H. S. Lo, Bull. Bot. Res., Harbin 18: 281.

1998.
mu gua lan
Shrubs, 34 m tall; branches angled, pale red or pale yellow, often stout, armed with spines 2.54 mm. Petiole 25 mm,
glabrous; leaf blade drying olive-green and abaxially paler, narrowly elliptic or oblong-lanceolate, 511 1.54 cm, both surfaces glabrous, base obtuse to acute or rarely subrounded, apex
acuminate to slightly obtuse; secondary veins 5 or 6 pairs, with
pilosulous domatia in abaxial axils; stipules triangular, ca. 2

mm. Inflorescences glabrescent; bracteoles squamiform, erose.

Flowers subsessile. Calyx glabrous; limb ca. 4.5 mm, subtruncate or 5-denticulate. Corolla pale yellow, tubular-campanulate
to tubular-funnelform, glabrous outside; tube at base ca. 3 mm
in diam., in throat 67 mm in diam., glabrous inside, in staminate flower 1011.5 mm, in pistillate flower 1313.5 mm; lobes
5, suborbicular, 3.54 mm, obtuse. Style stout, 56 mm. Berry
brown, ovoid to subglobose, 3.54.5 cm, glabrous, smooth;
seeds dark brown, oblong or ellipsoid, 810 mm, rather shiny.
Fl. Oct, fr. Apr of following year.
Forests in valleys; 400500 m. Hainan.

15. CHASSALIA Commerson ex Poiret in Lamarck, Encycl. Suppl. 2: 450. 1812.

wan guan hua shu
Chen Tao (); Charlotte M. Taylor
Shrubs, subshrubs, or small trees, unarmed. Raphides present. Leaves opposite or in whorls of 3, sometimes with domatia; stipules persistent or sometimes with top portion deciduous, interpetiolar or united around stem, triangular to bifid, sometimes with basal
portion becoming hardened, sometimes glandular at apex or on short appendages. Inflorescence terminal, thyrsiform to cymose,
many flowered, sessile to pedunculate, bracteate. Flowers sessile or pedicellate, bisexual, usually distylous. Calyx limb 5-lobed [to
truncate or denticulate]. Corolla white to pink or pale yellow, tubular to funnelform, with tube often curved, variously glabrous or
pubescent inside; lobes 5, valvate in bud. Stamens 5, inserted in corolla tube, included or exserted; filaments short or absent; anthers
dorsifixed. Ovary 2-celled, ovules 1 in each cell, basal; stigmas 2, linear, included or exserted. Infructescence frequently becoming
purple with axes swollen and jointed. Fruit purple to black, drupaceous, fleshy, ellipsoid, subglobose, or ovoid, with calyx limb
persistent; pyrenes 2, plano-convex, 1-celled, each with 1 seed, smooth on dorsal surface, on ventral surface with a large concave
excavation; seeds medium-sized, compressed orbicular; endosperm fleshy; radicle cylindrical, basiscopic.
About 40 species: widespread in tropical Africa, Asia, and Madagascar, and the Mascarene Islands; one species in China.

1. Chassalia curviflora (Wallich) Thwaites, Enum. Pl. Zeyl.

150. 1859.
wan guan hua
Subshrubs, erect, 12 m tall; branches weakly flattened to
subterete, glabrous or rarely sparsely puberulent. Leaves oppo-

site; petiole 14 cm, glabrous; blade drying membranous to

thinly papery and often yellowish green, oblong-elliptic, elliptic, oblanceolate, or narrowly lanceolate, 627 1.57.5 cm,
glabrous, base cuneate to attenuate, apex acuminate to long acuminate; secondary veins 817 pairs, without domatia; stipules
persistent, united shortly around stem, with interpetiolar portion

RUBIACEAE

88

broadly ovate or broadly triangular, 1.54.5 mm, acute or obtuse, entire or usually shortly bifid, with 1 or 2 bristles 0.31
mm, often gland-tipped. Inflorescence cymose, pyramidal to
rounded, several to many flowered, puberulent; peduncle 15
cm; branched portion 37 cm; axes weakly flattened; bracts lanceolate to triangular or usually multifid, 0.53 mm. Flowers
subsessile, trimorphic: with anthers exserted and stigmas included, with anthers included and stigmas exserted, or with anthers and stigmas both exserted. Calyx with hypanthium portion
ellipsoid to obovoid, 11.5 mm, glabrous; limb 5-lobed, 0.51
mm; lobes 0.30.5 mm, acute. Corolla white with pink, red, or
orange on lobes, outside glabrous to sparsely puberulent and
longitudinally ridged to winged along tube then midribs of
lobes; tube shallowly to markedly curved, straight or bent at
base, 1015 mm, pubescent inside; lobes (4 or)5, ovate-triangular, 22.5 mm, at apex thickened. Infructescence axes becoming swollen and red. Fruit purple, oblate to globose or
weakly didymous, 57 69 mm. Fl. AprJun, fr. AprJan.
Wet places, forest understories, at low elevations; 1002000 m.
Guangdong, Guangxi, Hainan, Xizang, Yunnan [Bangladesh, Bhutan,
Borneo, Cambodia, E and NE India (including Andaman Islands), Indonesia, Malaysia, Philippines, Singapore, Sri Lanka, Thailand, Vietnam].
At anthesis the corolla tube curves upward so that the base of the
tube is suberect, the middle portion of the tube arcs toward the horizontal, and the mouth of the corolla opens outward to the side (Puff et
al., Rubiaceae of Thailand, 101. 2005). Other authors have found this
species distylous with the flowers 5-merous or infrequently 4-merous
on aberrant individual flowers, as in many Rubiaceae; H. S. Lo (in
FRPS 71(2): 61. 1999) described it as tristylous with often 4 corolla
lobes.

1a. Leaves oblong-elliptic, elliptic, or

oblanceolate, 627 2.57.5 cm, with
secondary veins distinct on adaxial
surface .................................................... 1a. var. curviflora

1b. Leaves narrowly lanceolate,

1327 1.54.5 cm, with secondary
veins distinct to indistinct on adaxial
surface ..................................................... 1b. var. longifolia
1a. Chassalia curviflora var. curviflora
() wan guan hua (yuan bian zhong)
Psychotria curviflora Wallich in Roxburgh, Fl. Ind. 2: 167.
1824.
Leaves oblong-elliptic, elliptic, or oblanceolate, 627
2.57.5 cm; secondary veins distinct on adaxial surface. Fl.
AprJun, fr. AprJan.
Wet places, forest understories; 1002000 m. Guangdong, Guangxi, Hainan, Xizang, Yunnan [Bangladesh, Bhutan, Borneo, Cambodia, E
and NE India (including Andaman Islands), Indonesia, Malaysia, Philippines, Singapore, Sri Lanka, Thailand, Vietnam].

1b. Chassalia curviflora var. longifolia J. D. Hooker, Fl. Brit.

India 3: 177. 1880.
jian ye wan guan hua
Psychotria longifolia Dalzell, Hookers J. Bot. Kew Gard.
Misc. 2: 133. 1850, not Hoffmannsegg ex Roemer & Schultes
(1819), nor Sprengel (1824); Chassalia longifolia K. M. Wong.
Leaves narrowly lanceolate, 1327 1.54.5 cm; secondary veins distinct or often indistinct on adaxial surface. Fl. May,
fr. Jul.
Wet places, forest understories; 1002000 m. Guangdong, Guangxi, Hainan, Xizang, Yunnan [Bangladesh, Bhutan, Borneo, Cambodia, E
and NE India (including Andaman Islands), Indonesia, Malaysia, Philippines, Singapore, Sri Lanka, Thailand, Vietnam].
Plants that match this variety are uncommonly collected and are
provisionally separated here.

16. CINCHONA Linnaeus, Sp. Pl. 1: 172. 1753.

jin ji na shu
Chen Tao (); Charlotte M. Taylor
Kinkina Adanson; Quinquina Boehmer.
Shrubs or usually trees, unarmed; buds flattened with stipules erect and pressed together; bark usually notably bitter. Raphides
absent. Leaves opposite, decussate, usually with well-developed domatia; stipules caducous, interpetiolar or shortly united around
stem, ligulate to obovate, entire. Inflorescences terminal and often also in axils of uppermost leaves, cymose to paniculiform, many
flowered, pedunculate, bracteate. Flowers pedicellate, bisexual, fragrant, usually distylous. Calyx limb 5-lobed. Corolla yellow, pink,
purple, red, or occasionally white, salverform or funnelform, inside glabrous or pubescent in throat, with tube often weakly 5-ridged
outside; lobes 5, valvate in bud, with margins densely ciliate to villous. Stamens 5, inserted in corolla tube, included to partially
exserted; filaments short to developed, glabrous; anthers dorsifixed. Ovary 2-celled, ovules many in each cell on axile placentas;
stigma 2-lobed, lobes capitate to linear. Fruit capsular, ovoid to cylindrical or ellipsoid, septicidally dehiscent into 2 valves from base
or sometimes from apex with valves then loculicidal through septum, stiffly papery to woody, often lenticellate, with calyx limb
persistent; seeds numerous, medium-sized, ellipsoid to fusiform and somewhat flattened with membranous marginal wing and
elliptic central seed portion; endosperm fleshy; cotyledons ovate.
Twenty-three species: Central America (Costa Rica, Panama) and South America (Bolivia, Colombia, Ecuador, Peru, Venezuela) and cultivated
as species and hybrids in tropical regions worldwide; two species (both introduced) in China.
Several species of Cinchona are the natural source of quinine, which has long been used worldwide as a treatment for malaria. Quinine is found
along with several other alkaloids in high concentrations in some species of Cinchona, particularly the bark; these alkaloids give the plants their bitter
taste. Cinchona is native to South America, where its species are not all well differentiated, are morphologically variable, and hybridize freely
especially in cultivation, where numerous artificial hybrids have been created. Cinchona was recently monographed by Andersson (Mem. New York
Bot. Gard. 80: 175. 1998), followed here, who clarified the identities of the commonly cultivated species.

RUBIACEAE

89

Cinchona officinalis (Linnaeus, Sp. Pl. 1: 172. 1753; zheng ji na shu) is native to South America (Ecuador) and perhaps occasionally
cultivated in tropical regions worldwide. No confirmed documentation of this species has been seen from China. The name has long been incorrectly
used in cultivation for plants treated here as Cinchona calisaya (Andersson, loc. cit.: 5557). It is included for reference in the key to species.

1a. Leaf blade usually relatively broad, ovate, ovate-elliptic, or elliptic-oblong, 5.517 cm wide, abaxially
moderately to densely hirtellous at least when young and with pilosulous domatia but without crypt
domatia in axils of veins .............................................................................................................................................. 2. C. pubescens
1b. Leaf blade usually of average or relatively narrow width, lanceolate, oblong-lanceolate, elliptic,
obovate-lanceolate, or elliptic-oblong, 211 cm wide, abaxially glabrous or puberulent to hirtellous
and with or without crypt domatia.
2a. Leaf blade without crypt domatia .......................................................................................... C. officinalis (see comment above)
2b. Leaf blade with crypt domatia.
3a. Leaves with domatia best developed in proximal part of blade; calyx lobes comprising more
than 1/2 length of calyx limb; capsules stiffly papery to woody ...................................................................... 1. C. calisaya
3b. Leaves with domatia best developed in distal part of blade; calyx lobes comprising
less than or up to ca. 1/2 length of calyx limb; capsules stiffly papery .......................... C. officinalis (see comment above)
1. Cinchona calisaya Weddell, Ann. Sci. Nat., Bot., sr. 3, 10:
6. 1848.

matches the cultivated hybrid C. lucumifolia Pavon ex Lindley C.

pubescens (Andersson, loc. cit.: 60).

jin ji na shu

2. Cinchona pubescens Vahl, Skr. Naturhist.-Selsk. 1: 19. 1790.

Cinchona calisaya var. ledgeriana Howard; C. ledgeriana

(Howard) Bernelot Moens ex Trimen; Quinquina calisaya
(Weddell) Kuntze; Q. ledgeriana (Howard) Kuntze.

ji na shu

Trees or shrubs, to 15(25) m tall; bark grayish brown,

thin, with many shallow fissures; branches flattened to subquadrangular, puberulent to hirtellous or glabrescent. Petiole 3
20(30) mm, glabrous or hirtellous or puberulent; leaf blade
drying papery or thinly leathery, oblong-lanceolate, elliptic-oblong, or lanceolate, 716(21.5) 2.56(11) cm, both surfaces
glabrous or sparsely puberulent to hirtellous abaxially, base
acute to cuneate, apex obtuse to rounded or rarely acute;
secondary veins 711 pairs, usually with crypt domatia, these
best developed in proximal part of blade; stipules 1020 mm,
glabrous to puberulent or hirtellous, obtuse to rounded. Inflorescences 523 518 cm, densely hirtellous to puberulent;
bracts triangular, 0.53 mm; pedicels 18 mm. Calyx densely
sericeous; ovary portion ellipsoid, 1.52 mm; limb 12 mm,
sparsely puberulent, partially lobed; lobes ovate-triangular, 0.5
1 mm. Corolla white, pale yellow, or pale pink, glabrous to
puberulent outside; tube cylindrical, 59 mm, glabrous inside;
lobes lanceolate, 34(6) mm, acute. Capsules 830 38 mm,
stiffly papery to woody, puberulent or pilosulous to glabrescent;
seeds 310 1.63.7 mm (including wing). Fl. JunFeb.
Cultivated. Hainan, Taiwan, S Yunnan [native to South America
(Bolivia, Peru); cultivated in tropical regions worldwide].
Plants cultivated under the trade name Cinchona ledgeriana were
considered to belong to C. calisaya by Andersson (Mem. New York.
Bot. Gard. 80: 5557. 1998). The description of the plants treated as C.
ledgeriana by W. C. Chen (in FRPS 71(1): 224225. 1999) agrees with
Anderssons classification of C. lancifolia Mutis and also generally

Cinchona succirubra Pavon ex Klotzsch.

Trees and shrubs, to 12 m tall; bark grayish brown,
sometimes striped with white and/or longitudinally fissured;
branches somewhat flattened to subterete or angled, densely
pilosulous or hirtellous to puberulent or glabrescent. Petiole
1.54 cm, glabrous or puberulent to hirtellous; leaf blade drying
papery, ovate, ovate-elliptic, or elliptic-oblong, 1024.5 5.5
17 cm, adaxially glabrous to pilosulous or puberulent, abaxially
hirtellous to pilosulous or puberulent and often reddened at
least when young, base obtuse to rounded or truncate, apex obtuse to rounded; secondary veins 611 pairs, with pilosulous
domatia; stipules 1025 mm, sparsely pilosulous, obtuse to
rounded. Inflorescences 623 623 cm, pilosulous or hirtellous to puberulent; bracts triangular, 0.53 mm; pedicels 13
mm. Calyx with ovary portion ellipsoid, 23 mm, densely pilosulous; limb 1.53 mm, sparsely to densely pilosulous or hirtellous, shallowly lobed; lobes triangular, 0.51 mm. Corolla
white or pink, outside glabrescent to pilosulous or puberulent;
tube cylindrical, 8.514 mm; lobes ovate-lanceolate, 46 mm,
acute. Capsules 1018(41) 57 mm, pilosulous to glabrescent; seeds 712 23 mm (including wing). Fl. and fr. Jun
Feb.
Cultivated. Guangxi (Nanning), Hainan, Taiwan, S Yunnan [native
to Central America (Costa Rica) and South America (Bolivia, Colombia, Ecuador, Peru, Venezuela); often cultivated in tropical regions
worldwide].
This is the most frequently cultivated species and hybrid parent of
Cinchona. It is variable morphologically in cultivation and also in its
native range.

17. CLARKELLA J. D. Hooker, Fl. Brit. India 3: 46. 1880.

yan shang zhu shu
Chen Tao (); Charlotte M. Taylor
Small herbs from tubers, apparently perennial, unarmed, usually with a cluster of prophylls at stem base. Raphides present.
Leaves opposite, isophyllous or sometimes markedly anisophyllous at basalmost node, without domatia; stipules persistent, inter-

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petiolar and fused to petioles, triangular, perhaps sometimes glandular on margins. Inflorescences terminal, cymose, umbelliform, or
subfasciculate, few to several flowered, pedunculate or sessile and tripartite, bracteate. Flowers pedicellate, bisexual with biology
unknown. Calyx limb well developed, markedly reticulate veined, 5(7)-lobed for ca. 1/2 length. Corolla white, slenderly salverform
or funnelform-salverform, glabrous inside; lobes 5, valvate in bud. Stamens 5, inserted at base of corolla tube, included; filaments
short. Ovary 2-celled, ovules numerous in each cell on axile placentas inserted not far below middle of septum; disk pilosulous; stigma 2-lobed, pubescent. Fruit indehiscent, obconical, dry, with funnelform calyx limb persistent; seeds numerous, small, subellipsoid,
black papillose or -granular.
One species: China, N India, N Myanmar, Thailand.
This is a poorly known species or genus that apparently prefers wet limestone substrates and is similar to Pseudopyxis.

1. Clarkella nana (Edgeworth) J. D. Hooker, Fl. Brit. India 3:

46. 1880.
yan shang zhu
Ophiorrhiza nana Edgeworth, Trans. Linn. Soc. London
20: 60. 1846; Clarkella siamensis Craib; O. pellucida H.
Lveill.
Low herbs, to 10 cm tall; tubers ellipsoid-oblong, 11.5
cm, scaly; stems unbranched, subterete, glabrous to densely villosulous. Leaves opposite or sometimes only 1 leaf developing
at basalmost node; petiole 0.55 cm, glabrous to puberulent;
blade drying membranous or thinly papery, ovate to broadly
ovate, 16 14 cm, both surfaces glabrous to puberulent or
villosulous, base obtuse to cordulate and sometimes oblique,

apex acute to rounded; secondary veins 48 pairs; stipules triangular to narrowly triangular, 0.20.5 mm. Inflorescence 0.83
cm, 325-flowered, villosulous to puberulent; bracts ovate
to narrowly oblong, 0.512 mm; pedicels 13.5 mm. Calyx
densely puberulent to villosulous; hypanthium portion 12 mm;
lobes triangular to ovate, 13 mm, markedly unequal on an
individual flower. Corolla outside villosulous to puberulent;
tube 1014 mm; lobes elliptic to suborbicular, ca. 3 mm, ciliolate. Fruit 78 mm, villosulous, with calyx limb becoming 2
4.5 mm with increase in size principally in growth of basal,
unlobed portion. Fl. and fr. Aug.
On wet rocks, usually limestone; ca. 1400 m. Guangdong (Liannan), N Guangxi (Nandan), C Guizhou, N Yunnan [N India, N Myanmar, Thailand].

18. COFFEA Linnaeus, Sp. Pl. 1: 172. 1753.

ka fei shu
Chen Tao (); Charlotte M. Taylor
Cafe Adanson.
Shrubs or small trees, unarmed, often resinous on young growth; lateral branches usually spreading horizontally. Raphides
absent. Leaves opposite or rarely in whorls of 3, distichous at least on lateral branches, often with foveolate and/or pilosulous domatia; stipules persistent, shortly united around stem, generally triangular, sometimes aristate. Inflorescences axillary, in each axil with 1
to several capitate to fasciculate, 1- to several-flowered cymes, these sessile to shortly pedunculate, bracteate; bracts often fused in
cupulate pairs (i.e., forming a calyculus). Flowers sessile or shortly pedicellate, bisexual, monomorphic. Calyx limb obsolete or
occasionally truncate or 46-toothed. Corolla white or pink, salverform or funnelform, inside glabrous or villous in throat; lobes 49,
convolute in bud. Stamens 48, inserted in corolla throat, exserted; filaments absent or short; anthers dorsifixed near base. Ovary 2celled, ovules 1 in each cell, attached at middle of septum; stigma 2-lobed, exserted. Fruit red, yellow, orange, blue, or black, drupaceous, globose to ellipsoid, fleshy or infrequently dry, with calyx limb when developed persistent; pyrenes 2, each 1-celled, with 1
seed, plano-convex, leathery or papery, on ventral (i.e., adaxial) face with longitudinal groove; seeds medium-sized to large,
longitudinally grooved on ventral face; radicle terete, basiscopic.
About 103 species: native to tropical Africa, Madagascar, and the Mascarene Islands, several species and hybrids cultivated in moist tropical
regions worldwide; five species (all introduced) in China.
Several species of Coffea are widely cultivated as a source of the drink coffee, a leading world commodity. Species limits and identifications are
often difficult for wild plants, due to the complexity of the genus, its evolutionary behavior, and its numerous reduced morphological features; and the
taxonomy of cultivated plants is additionally complicated by extensive, sometimes poorly documented hybridization for crop improvement during
several centuries. The genus is native to Africa, Madagascar, and the Mascarenes; plants found outside this region are cultivated. Cultivated plants of
Coffea generally persist after active cultivation is abandoned but do not generally establish growing permanent populations or spread. Cultivated
Coffea is surveyed usefully by Purseglove (Trop. Crops: Dicot. 451492. 1968). Coffea arabica is the most valuable species, producing highest
quality coffee; this is a tetraploid species (Stoffelen et al., Opera Bot. Belg. 7: 237248. 1996). Coffea canephora is generally the most productive
species, producing a lower quality coffee; this is a diploid species (Purseglove, loc. cit.: 482488). Coffea liberica also produces a lower quality coffee
than C. arabica, is also diploid (Purseglove, loc. cit.: 488491), and is less often cultivated. W. C. Ko (in FRPS 71(2): 2225. 1999) additionally
treated two species, C. congensis and C. stenophylla, that have been hybridized with commercial coffee (Purseglove, loc. cit.: 458) and may persist
from old plantations, but these are also two names that have been widely confused in cultivation with C. arabica and C. canephora (Davis et al., Bot.
J. Linn. Soc. 152: 483, 497. 2006). Some frequently used synonymous names are included here for reference.
Coffea is similar to Psilanthus J. D. Hooker, and some species have been variously treated in each genus depending on the current circum-

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91

scriptions. Davis et al. (Monogr. Syst. Bot. Missouri Bot. Gard. 104: 398420. 2005) addressed this problem and concluded by separating the genera;
in their circumscription Psilanthus is not known from China. Traditional Coffea descriptions often retain characters of Psilanthus, including that of W.
C. Ko (loc. cit.: 2025). Coffea benghalensis B. Heyne ex Schultes and C. jenkinsii J. D. Hooker were included in the Fl. Xizang. (4: 445447. 1985).
Davis et al. (loc. cit. 2006: 501) treated the first of these species as P. benghalensis (B. Heyne ex Schultes) J.-F. Leroy; Purseglove (loc. cit.: 458) listed
it as a native species of SE Asia and Sumatra sometimes cultivated for coffee in India. Davis et al. (loc. cit. 2006: 504) treated C. jenkinsii as a species
of Nostolachma T. Durand: N. jenkinsii (J. D. Hooker) Deb & Lahiri.
The fruit of Coffea are sometimes described informally as berries because of their size and fleshy texture, but morphologically they are similar
to other drupes of Rubiaceae; thus, this technical terminology is used here. The fruit of Coffea are also commercially sometimes called cherries.

1a. Blades of outer, unshaded leaves larger, usually 1540 622 cm, with apex acuminate to obtuse; fruit red.
2a. Leaf blade without domatia or with glabrous foveolate domatia in abaxial vein axils; fruit ovoid-globose
with length generally equal to width, 1012 1012 mm ................................................................................. 2. C. canephora
2b. Leaf blade with glabrous or pilosulous foveolate domatia in abaxial vein axils; fruit ellipsoid, longer than
wide, 1921 1517 mm ......................................................................................................................................... 4. C. liberica
1b. Blades of outer, unshaded leaves smaller, mostly shorter than or up to 15 7 cm, with apex acute to at least
shortly acuminate; mature fruit red to black or bluish black.
3a. Inflorescences with cymes 1 or 2 in each axil, each cyme 24-flowered; mature fruit black or bluish
black; leaf blade narrowly elliptic-oblong to narrowly oblanceolate, 410 1.52.5 cm; corolla
with 69 lobes ..................................................................................................................................................... 5. C. stenophylla
3b. Inflorescences with cymes 14 per axil, each cyme 25-flowered; mature fruit red; leaf blade elliptic,
elliptic-oblong, ovate-lanceolate, ovate, or lanceolate-elliptic, 38.5 cm wide; corolla with 46 lobes.
4a. Stipules aristate at apex; leaves of unshaded outer branches with lateral veins 713 pairs, with
glabrous foveolate domatia in abaxial vein axils; fruit smooth or usually drying with a shallow
sulcus or indentation along septum .................................................................................................................... 1. C. arabica
4b. Stipules obtuse to acute at apex; leaves of unshaded outer branches with lateral veins 69 pairs,
with pilosulous domatia in abaxial vein axils; fruit drying smooth ............................................................... 3. C. congensis
1. Coffea arabica Linnaeus, Sp. Pl. 1: 172. 1753.
xiao li ka fei
Small trees or large shrubs, 58 m tall; branches flattened
to subterete, glabrous. Petiole 815 mm, glabrous; leaf blade
drying thinly leathery, elliptic, elliptic-oblong, or occasionally
ovate-lanceolate, (2)614(22) 3.55(8.5) cm, glabrous on
both surfaces, base cuneate to obtuse or rarely rounded, margins
occasionally crisped-undulate, apex acuminate with tip usually
1015 mm; secondary veins 710(13) pairs, without domatia
or with glabrous foveolate domatia; stipules broadly triangular,
38(12) mm, aristate at least on youngest branches. Inflorescences with 1 to several cymes in each axil, each cyme subcapitate to fasciculate, (1 or)25-flowered, sessile to pedunculate with peduncles to 4 mm; bracts cupuliform, 12 mm; pedicels to 2 mm. Calyx glabrous; ovary portion ellipsoid, 13 mm;
limb truncate to undulate or denticulate, 0.21 mm. Corolla
white, funnelform, outside glabrous; tube 515 mm; lobes (4
or)5(or 6), spatulate-elliptic, 920 mm, obtuse. Drupe red, ellipsoid to subglobose, 1116 914 mm, when dry smooth or
sometimes weakly didymous, glabrous. Fl. MarJul, fr. Oct
Jan.
Cultivated in moist, usually cool tropical regions; 200700 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, Yunnan
[native to E Africa (Ethiopia, N Kenya, Sudan); cultivated worldwide].
This species is tetraploid and is the source of Arabica Coffee. It
has been widely hybridized with several other Coffea species to produce
commercial coffee plants; flower size varies markedly among many of
these.

2. Coffea canephora Pierre ex Froehner, Notizbl. Knigl. Bot.

Gart. Berlin 1: 237. 1897.

zhong li ka fei
Coffea robusta L. Linden.
Small trees or shrubs, 48 m tall; branches flattened becoming subterete, glabrous. Petiole 1020 mm, glabrous; leaf
blade drying thickly papery, elliptic, elliptic-oblong, or occasionally ovate-oblong, (12)1530(40) (4.5)612(22) cm,
glabrous on both surfaces, base cuneate to obtuse, margins flat
or occasionally crisped-undulate, apex acuminate with tip 10
18 mm; secondary veins (8)1012(17) pairs, without domatia
or with glabrous foveolate domatia; stipules triangular, 618
mm, obtuse to acute, aristate. Inflorescences with cymes 1
3(7) in each axil, each cyme subcapitate to fasciculate, 3
6-flowered, subsessile to pedunculate with peduncles to 7 mm;
bracts cupular, 13 mm; pedicels to 2 mm. Calyx glabrous;
ovary portion ellipsoid, 12 mm; limb reduced or denticulate,
0.10.5 mm. Corolla white to pink, funnelform, outside glabrous; tube 516 mm; lobes (4 or)57(or 8), spatulate to narrowly elliptic, 819 mm, obtuse to rounded. Drupe red, subglobose, 1012 1012 mm, smooth when dry, glabrous. Fl. Apr
Jun, fr. OctDec.
Cultivated in moist, often warm tropical regions. Fujian, Guangdong, Hainan, Yunnan [widespread in tropical Africa; commonly cultivated worldwide].
This species is diploid and is cultivated as Robusta Coffee. It
has been widely hybridized with several other species to produce
commercial coffee plants.

3. Coffea congensis Froehner, Notizbl. Knigl. Bot. Gart. Berlin 1: 235. 1897.
gang guo ka fei

RUBIACEAE

92

Shrubs, 26 m tall; branches flattened, glabrous. Petiole 5

10 mm, glabrous; leaf blade drying thinly leathery, elliptic-oblong to ovate or lanceolate-elliptic, 815 37 cm, glabrous on
both surfaces, base cuneate or obtuse to rounded, apex acute to
shortly acuminate with tip 510 mm; lateral veins 69 pairs,
usually with pilosulous domatia; stipules triangular to broadly
triangular, 25 mm, obtuse to acute but not aristate. Inflorescences with cymes 14 in each axil, each cyme subcapitate to
fasciculate, 24-flowered, sessile to pedunculate with peduncles
to 4 mm; bracts cupuliform, 13 mm; pedicels to 3 mm. Calyx
glabrous; ovary portion cylindrical-ellipsoid, 11.5 mm; limb
reduced or undulate, 0.10.5 mm. Corolla white, funnelform,
outside glabrous; tube 710 mm; lobes 5 or 6, spatulate to narrowly elliptic, 710 mm, obtuse to rounded. Drupe red, ellipsoid to ovoid-oblong, 1012 810 mm, smooth when dry, glabrous. Fr. Dec.
Cultivated in moist forest regions. Hainan [native to Africa (Congo River basin: Democratic Republic of Congo, Republic of Congo),
growing as rheophyte; widely planted in tropical regions, though perhaps not so often in recent decades].
This species is presumably diploid and has been occasionally
hybridized with other species to produce commercial coffee (Purseglove, Trop. Crops: Dicot. 458. 1968; Bridson, Fl. Trop. E. Africa, Rub.
(Pt. 2), 703. 1988).

4. Coffea liberica W. Bull ex Hiern, Trans. Linn. Soc. London,

Bot. 1: 171. 1876.
da li ka fei
Coffea dewevrei De Wildeman & T. Durand.
Small trees or large shrubs, 615 m tall; branches flattened
to weakly angled, often rather stout, glabrous. Petiole 820 mm,
often rather stout, glabrous; leaf blade drying thinly leathery to
stiffly papery, elliptic to obovate or obovate-elliptic, 1438
5.512(20.5) cm, glabrous on both surfaces, base cuneate to
obtuse, margins flat, apex obtuse to shortly acuminate with tip
410 mm; secondary veins 710(13) pairs, without domatia or
with glabrous to pilosulous foveolate domatia; stipules broadly
triangular, 24.5 mm, obtuse to acute but not aristate. Inflorescences with cymes 13 per axil, each cyme fasciculate to
subcapitate, 210-flowered, subsessile; bracts cupuliform, 13
mm; pedicels to 1 mm. Calyx glabrous; ovary portion ellipsoid-

cylindrical, 1.53.5 mm; limb reduced or glandular-denticulate,

to 0.2 mm. Corolla white, funnelform, outside glabrous; tube 4
13 mm; lobes 68, spatulate to lanceolate or narrowly elliptic,
816 mm, obtuse to rounded. Drupe red, ellipsoid, 1921 15
17 mm, smooth when dry, glabrous. Fl. JanMay, fr. presumably AugNov.
Cultivated in moist, warm to cool regions. Fujian, Guangdong,
Hainan, Yunnan [widespread in tropical Africa; widely but not intensively cultivated in tropical regions worldwide].
This species is diploid and is cultivated as Liberica Coffee. It
has been hybridized with several other species to produce commercial
coffee plants.

5. Coffea stenophylla G. Don, Gen. Hist. 3: 581. 1834.

xia ye ka fei
Shrubs to small trees, 36 m tall; branches flattened to
subterete, glabrous. Petiole 35 mm, glabrous; leaf blade drying
thinly leathery, narrowly oblanceolate to narrowly elliptic-oblong, 410 1.52.5 cm, glabrous on both surfaces, base acute
to cuneate, margins generally flat, apex acuminate with tip 10
20 mm; secondary veins 710 pairs, without domatia or with
glabrous foveolate domatia; stipules broadly triangular, 23.5
mm, obtuse to acute and sometimes shortly mucronate. Inflorescences with cymes 1 or 2 per axil, each branched to subcapitate or fasciculate, 24-flowered, subsessile to pedunculate
with peduncles to 2.5 mm; bracts cupuliform, 12 mm; pedicels
to 6 mm. Flower buds resinous. Calyx glabrous; ovary ellipsoid, 11.5 mm; limb reduced, truncate. Corolla white or pale
pink, funnelform, outside glabrous to puberulent; tube 68 mm;
lobes 68(or 9), spatulate, 1215 mm, obtuse. Drupe black or
bluish black, subglobose to ovoid, 1213 810 mm. Fl. Mar
Apr, fr. Dec.
Cultivated in moist, probably warm forest regions. Hainan
(Chengmai) [native to W Africa; formerly occasionally cultivated in Old
World tropics, apparently primarily in coffee research stations].
This species is diploid and is cultivated as Highland Coffee of
Sierra Leone. It has been occasionally hybridized with other species to
produce commercial coffee plants and reported as cultivated by older
authors (Purseglove, Trop. Crops: Dicot. 459. 1968; Bridson, Fl. Trop.
E. Africa, Rub. (Pt. 2), 703. 1988, with documentation).

19. COPTOSAPELTA Korthals, Ned. Kruidk. Arch. 2(2): 112. 1851.

liu su zi shu
Chen Tao (); Charlotte M. Taylor
Lianas or scandent shrubs, unarmed. Raphides absent. Leaves opposite, sometimes with domatia; stipules caducous, interpetiolar, triangular. Inflorescences axillary [to terminal], 1-flowered [or cymose and several to many flowered], pedunculate, bracteate. Flowers sessile, pedunculate, or pedicellate, bisexual, monomorphic. Calyx limb 5-lobed. Corolla pale yellow or white becoming
yellowed with age, salverform, variously glabrous or pubescent inside; lobes 5, convolute to right in bud. Stamens 5, inserted in
corolla throat, exserted; filaments short; anthers basifixed. Ovary 2-celled, ovules numerous in each cell on axile placentas; stigma
fusiform, exserted. Fruit capsular, subglobose, loculicidally dehiscent with valves sometimes secondarily separating along septum,
woody, with calyx limb persistent; seeds several, medium-sized to large, flattened, suborbicular, bordered by a regularly erose wing;
endosperm fleshy; embryo straight; cotyledons short; radicle terete, basiscopic.
Sixteen species: China, Indonesia, Japan, Laos, Malaysia, Myanmar, New Guinea, Philippines, Thailand, Vietnam; one species in China.

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93

The 35-porate pollen of Coptosapelta was studied by Verellen et al. (J. Pl. Res. 117: 5768. 2003), who also noted that the genus has secondary
pollen presentation and accumulates aluminum in the leaves. Based on molecular and some morphological characters, Coptosapelta is now considered
rather isolated and one of the most basal lineages in Rubiaceae (Bremer et al., Syst. Biol. 48: 413435. 1999; Bremer & Manen, Pl. Syst. Evol. 225:
4372. 2000), with its closest relative probably being Acranthera and these two genera most closely related to Luculia (Rydin et al., Pl. Syst. Evol.
278: 101123. 2009).

1. Coptosapelta diffusa (Champion ex Bentham) Steenis,

Amer. J. Bot. 56: 806. 1969.
liu su zi
Thysanospermum diffusum Champion ex Bentham,
Hookers J. Bot. Kew Gard. Misc. 4: 168. 1852.
Lianas or scandent shrubs, 25 m or taller, often much
branched; branches densely hirtellous or strigillose to strigose
often becoming glabrescent with age. Petiole 25 mm, hirsute
to strigose or rarely glabrous; leaf blade drying papery to leathery and often yellowish green, ovate or ovate-oblong to lanceolate, 29.5 0.83.5 cm, adaxially rather shiny and glabrous
except strigose to hirsute along costa, abaxially glabrous except
strigose to hirsute on principal veins, base rounded to obtuse,
margins sometimes sparsely ciliate and/or thinly revolute, apex
acute or acuminate; secondary veins 4 or 5 pairs; stipules lanceolate to narrowly triangular, 27 mm, acute. Inflorescences

with peduncles slender, 320 mm, glabrous or usually strigose

to hirtellous; bracts 1 or 2 on upper part of peduncle, 0.51 mm.
Calyx glabrous to strigillose; ovary portion subglobose, 12
mm; limb deeply lobed; lobes ovate-triangular, 0.81 mm. Corolla outside glabrous to strigillose or sericeous, inside hirtellous in upper part of tube and usually onto basal half of lobes;
tube cylindrical to somewhat funnelform, 815 mm; lobes elliptic-oblong to spatulate, 46 mm, obtuse to rounded. Anthers
3.54 mm. Stigma 2.53 mm. Capsule brownish yellow, compressed globose or compressed ellipsoid, weakly didymous, 4
6 58 mm; seeds 1.52 mm in diam. Fl. MayJul, fr. Jun
Dec.
Thickets or forests on mountains or hills; 1001500 m. Anhui,
Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan (Ryukyu Islands)].
This is a commonly collected species in China.

20. DAMNACANTHUS C. F. Gaertner, Suppl. Carp. 18. 1805.

hu ci shu
Chen Tao (); Charlotte M. Taylor
Tetraplasia Rehder.
Shrubs, sometimes with paired infrastipular or superaxillary spines; branches sometimes with complex sympodial growth with
reduced internodes and prophylls (Damnacanthus indicus); roots at least sometimes moniliform (i.e., nodose or node-like constricted). Raphides present. Leaves opposite, apparently without domatia; stipules persistent and becoming hardened or sometimes
falling by fragmentation, interpetiolar or shortly united around stem, generally triangular, acute or shortly bifid to multifid. Inflorescences pseudoaxillary, superaxillary, apparently terminal, and/or paired on short shoots giving an appearance of being axillary, 1flowered or usually cymose to fasciculate and 24-flowered, subsessile to apparently shortly pedunculate (i.e., borne on a leafless
short shoot), bracteate with bracts usually small and glandular-fimbriate. Flowers subsessile to pedicellate and often nodding, bisexual, monomorphic or distylous. Calyx limb cupular or campanulate, 4-lobed (or 5-lobed, D. henryi). Corolla white to yellow or pale
purple, tubular-funnelform, often leathery, inside densely pubescent in throat to throughout; tube rarely fenestrate (D. henryi); lobes 4
(or 5, D. henryi), valvate in bud. Stamens 4, inserted in upper part of corolla tube, included or exserted; filaments short; anthers
dorsifixed. Ovary 4-celled (or 2-celled, D. henryi), ovules 1 in each cell and attached near top of septum, campylotropous; stigmas 4
(or 2, D. henryi), linear, included or exserted. Fruit red, drupaceous, globose to ellipsoid or oblate, fleshy, with calyx limb persistent;
pyrenes 4(or 2, D. henryi), each with 1 seed, plano-convex, subglobose, ellipsoid, or obtusely trigonous; seeds medium-sized, subglobose to plano-convex; endosperm corneous; embryo small; radicle hypogeous.
About 13 species: China, N India, Japan, Korea, Laos, Myanmar, Vietnam; 11 species (six endemic) in China.
The morphology of Damnacanthus was reviewed in detail by Robbrecht et al. (Blumea 35: 307345. 1991), who described its complex,
sympodial growth pattern and variations found in different species. They considered the stems of Damnacanthus to be composed of sympodial units,
with varying degrees of development of the individual parts in different species. The most characteristic and apparently complicated growth is found
in D. indicus, in which each sympodial unit comprises a basal node bearing a pair of prophylls, similar to bud scales or reduced leaves; then, a
developed internode; then, a node bearing a pair of normally developed foliage leaves, decussate in orientation to the prophylls; then, a node (without
an intervening internode) bearing a pair of thorns, decussate in orientation to the leaves. Growth of the stem continues from one of the axillary buds of
the foliage leaves, which gives the thorns the appearance of being stipular or superaxillary in position. The alternating prophylls and foliage leaves
produce the characteristic heterophyllous growth of this genus. Species of Damnacanthus vary in the characteristic number of nodes in each
sympodial unit and in the development (or not) of the thorns. Robbrecht et al. (loc. cit.) interpreted the characteristic spines of the genus as reduced
shoot systems produced from the axillary buds subtending two undeveloped (and thus missing) leaves. They considered the characteristic 2- or 4flowered inflorescences of Damnacanthus to be formed of one or two sympodial growth units, produced from one or both axils of a node bearing
foliage leaves, with each of these units comprising three congested nodes, lacking separating internodes, with the basalmost nodes bearing bractlike
scales and the terminal node producing a flower in each axil then stopping growth, thus comprising a 2-flowered cymule. They also noted that,
although previous authors have described the ovules of Damnacanthus as amphitropous or pendulous, in fact the ovules are unique in the Rubiaceae
in being campylotropous.

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Damnacanthus is represented by at least three species in Japan (Fl. Japan 3a: 224225. 1993), several of them apparently common and
hybridizing, and its taxonomy has been rather intensively studied there and in Taiwan (e.g., Koidzumi, Acta Phytotax. Geobot. 3: 155160. 1934),
where it apparently has some medicinal use. Damnacanthus was revised for China by H. S. Lo (Acta Phytotax. Sin. 17(3): 104109. 1979), then
treated comprehensively by Y. Z. Ruan in FRPS (71(2): 167176. 1999) in an essentially monographic work. Koidzumi (loc. cit.) recognized three
sections within Damnacanthus and treated Tetraplasia as a separate genus, based largely on root characters, but these were not mentioned again until
Y. Z. Ruan (loc. cit.: 169) recognized two sections in Chinese Damnacanthus, one of them under an unpublished name. The Chinese plants with
spines were included in D. sect. Damnacanthus; the unarmed plants were separated by Koidzumi in Tetraplasia and were included by Ruan in his
second, unnamed section.
Naiki and Nagamasu (J. Pl. Res. 116: 105113. 2003; Amer. J. Bot. 91: 664671. 2004) surveyed the breeding biology of several Japanese and
Chinese species of Damnacanthus and found variation in breeding system among species, discovered a correlation between ploidy with breeding
system but not leaf size, and reported distyly and dimorphic pollen in this genus.

1a. Branches glabrous, hispidulous, hirtellous, or puberulent when young, with spines in axils of stipules or leaves,
with at least shortly developed spines at apices (these may appear to be stipule bristles if not observed carefully).
2a. Spines 16 mm, persistent or deciduous when new leaves come out; leaf blade with midrib flat to impressed
or thinly prominulous adaxially.
3a. Leaf blade lanceolate or oblong-lanceolate, 415 cm, with or usually without microphylls (i.e., prophylls;
reduced leaves), midrib flat to impressed adaxially; spines 12 mm; young branches and petioles
puberulent or hispidulous; corollas 1518 mm; stipules caducous ................................................................ 2. D. giganteus
3b. Leaf blade ovate, lanceolate, oblong-ovate, or oblong-lanceolate, 38 cm, with or without
microphylls, midrib thinly prominulous adaxially; spines 26 mm; young branches and petioles
sparsely hispidulous, puberulent, hirtellous, or glabrescent; corollas 1015 mm; stipules
persistent at least on distalmost nodes ..................................................................................................... 8. D. macrophyllus
2b. Spines 325 mm, persistent; leaf blade with midrib thinly prominulous adaxially.
4a. Leaf blade lanceolate or oblong-lanceolate, 37.5 cm, with 510 pairs of secondary veins ............................... 11. D. tsaii
4b. Leaf blade cordiform, ovate, elliptic, broadly elliptic, broadly ovate, or elliptic-ovate, 0.54 cm, with
24 pairs of secondary veins.
5a. Leaf blade 0.53 cm, with secondary veins 24 pairs; spines 320 mm, 1/2 or more as long as
leaf blade ....................................................................................................................................................... 6. D. indicus
5b. Leaf blade 34 cm, with secondary veins 35 pairs; spines 310 mm, less than 1/2 as long as
leaf blade ......................................................................................................................................................... 9. D. major
1b. Branches glabrous, without spines.
6a. Low to dwarf shrubs, 0.51 m tall; corolla 810.5 mm; Taiwan .................................................................... 1. D. angustifolius
6b. Low to tall shrubs, 0.45 m tall; corolla 816 mm; mainland and Hainan.
7a. Leaf blade lanceolate-linear, apex tapered to acute or acuminate tip.
8a. Leaf blade when dry straw-yellow adaxially, olive-green abaxially, and thickly leathery, with
secondary veins flat and indistinct adaxially, 69 pairs; calyx teeth sharply triangular ................... 3. D. guangxiensis
8b. Leaf blade when dry gray, gray-green, or brownish green adaxially, gray-green, brownish green,
or straw-yellow abaxially, and papery, with secondary veins flat and indistinct to prominulous
adaxially, 916 pairs; calyx teeth broadly triangular ................................................................................. 7. D. labordei
7b. Leaf blade lanceolate, narrowly elliptic, linear, elliptic, elliptic-ovate, elliptic-oblong, or
oblong-lanceolate, apex acute to cuspidate or long but rather abruptly acuminate.
9a. Ovary 2-celled, stigmas 2; corolla lobes variably 4 or 5 .............................................................................. 5. D. henryi
9b. Ovary 4-celled, stigmas 4; corolla lobes regularly 4.
10a. Leaves isomorphic (i.e., prophylls not present), with blade elliptic-ovate, elliptic-oblong,
or oblong-lanceolate; Hainan ...................................................................................................... 4. D. hainanensis
10b. Leaves dimorphic (i.e., regularly with prophylls), with blade linear at lower part of stem
to elliptic, elliptic-oblong, or oblong-lanceolate at upper part of stem; mainland .................... 10. D. officinarum
1. Damnacanthus angustifolius Hayata, J. Coll. Sci. Imp.
Univ. Tokyo 25(19): 113. 1908.
tai wan hu ci
Damnacanthus angustifolius var. altimontanus J. C. Liao;
D. angustifolius f. stenophyllus (Koidzumi) T. Yamazaki; D.
angustifolius var. stenophyllus (Koidzumi) Masamune; D.
stenophyllus (Koidzumi) Masamune; Tetraplasia angustifolia
(Hayata) Koidzumi; T. stenophylla Koidzumi.
Dwarf to low shrubs, 0.51 m tall. Branches glabrous,

without spines, when young subquadrate or usually with 8

alternately thick and thin longitudinal ridge lines, becoming 4angled and yellow. Petiole of developed leaves 1.55 mm, glabrous; leaf blade drying stiffly papery, narrowly lanceolate, narrowly elliptic, linear-lanceolate, elliptic, or lanceolate-elliptic,
514 0.53 cm, glabrous throughout or sometimes sparsely
puberulent to hispidulous adaxially, base acute to cuneate, margins flat and entire or irregularly serrulate, apex acute to acuminate; midrib thinly prominulous adaxially; secondary veins 59
pairs; stipules caducous, interpetiolar, triangular to spatulate,
0.51 mm, glabrous, acute to glandular-fimbriate. Inflores-

RUBIACEAE

cences glabrous. Pedicels 25 mm. Calyx glabrous; hypanthium

portion turbinate, 1.21.5 mm; limb ca. 1 mm, lobed for ca. 1/2;
lobes triangular. Corolla white, glabrous outside; tube 68 mm;
lobes triangular, 22.5 mm. Drupes oblate, ca. 4 6 mm. Fl.
JanApr, Oct, fr. Jan, Jun, OctNov.
Primary forests; 10002500 m. Taiwan.
Damnacanthus angustifolius var. altimontanus is recognized as
distinct in the Kew Rubiaceae checklist (Govaerts et al., World Checkl.
Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on 15 Sep
2010) but was formally synonymized in the Fl. Taiwan (ed. 2, 4: 251.
1998), which is followed here.

95

out spines. Petiole ca. 6 mm, glabrous; leaf blade drying thickly
leathery, straw-yellow adaxially, olive-green abaxially, lanceolate-linear, 1322 12 cm, adaxially glabrous, abaxially
sparsely pilosulous, base cuneate, margins entire and flat, apex
acute then shortly tapered-acuminate; midrib prominent adaxially; secondary veins 69 pairs; stipules caducous or deciduous
through fragmentation, interpetiolar, triangular, acute to glandular-fimbriate. Pedicels ca. 2 mm. Calyx glabrous; hypanthium
portion cupuliform, ca. 1 mm; limb ca. 1 mm, lobed for ca. 1/2;
teeth 4 or 5, narrowly triangular, acuminate. Corolla white, outside glabrous; tube ca. 8 mm, pubescent inside; lobes ovate to
triangular, ca. 4 mm. Fruit unknown. Fl. winterspring.

This species was reported to be distylous by Naiki and Nagamasu

(Amer. J. Bot. 91: 664671. 2004). This species was reported from
Guangdong by Merrill and Chun (Sunyatsenia 1(1): 80. 1930) but said
to be restricted to Taiwan by Y. Z. Ruan in FRPS (71(2): 173174.
1999); the Guangdong plants were apparently included by Ruan (loc.
cit.: 174175), followed here, as Damnacanthus labordei.

4. Damnacanthus hainanensis (H. S. Lo) H. S. Lo ex Y. Z.

Ruan, Fl. Reipubl. Popularis Sin. 71(2): 176. 1999.

2. Damnacanthus giganteus (Makino) Nakai, Trees Shrubs

Japan, 412. 1922.

Damnacanthus henryi (H. Lveill) H. S. Lo subsp. hainanensis H. S. Lo, Acta Phytotax. Sin. 17(3): 108. 1979.

duan ci hu ci

Shrubs, 14 m tall. Branches brown, angulate or ridged,

glabrous, without spines. Petiole of developed leaves 28
mm, glabrous; leaf blade drying black, elliptic-ovate, oblong,
or oblong-lanceolate, 611 24.5 cm, glabrous, base rounded
or cuneate, margins thinly revolute, apex cuspidate; midrib
prominulous adaxially; secondary veins 57 pairs; stipules
caducous, interpetiolar, triangular. Pedicels ca. 2 mm. Calyx
limb subtruncate or lobed; teeth triangular. Corolla white,
outside glabrous; tube ca. 14 mm; lobes 4, ovate-lanceolate.
Drupes ca. 8 mm in diam., glabrous. Fl. May, fr. Nov.

Damnacanthus indicus C. F. Gaertner var. giganteus

Makino, Bot. Mag. (Tokyo) 18: 33. 1904; D. macrophyllus
Siebold ex Miquel f. giganteus (Makino) T. Yamazaki; D.
macrophyllus var. giganteus (Makino) Koidzumi; D. subspinosus Handel-Mazzetti.
Shrubs or rarely small trees, 0.52(7.5) m tall. Roots
moniliform. Branches usually 4-angulate or terete to flattened,
dark green and sparsely hispidulous or puberulent to glabrous
when young, becoming grayish yellow and glabrous, with few
spines 12 mm and deciduous or sometimes persistent. Petiole
of developed leaves 25 mm, sparsely puberulent to glabrous;
leaf blade drying leathery, lanceolate or oblong-lanceolate, 4
14(15) 23(5) cm, adaxially glabrous, abaxially glabrous or
rarely densely puberulent along veins, base obtuse to rounded,
margins flat to usually revolute and/or sometimes crisped, apex
acuminate or acute; midrib flat to impressed adaxially; secondary veins 57 pairs; stipules caducous, interpetiolar, 12 mm,
puberulent to glabrous, acute to bifid, thickened. Inflorescences
strigillose to glabrescent. Pedicels 12 mm. Calyx strigillose to
puberulent; hypanthium portion turbinate, 1.21.5 mm; limb 1
1.5 mm, lobed for 1/41/2; teeth broadly triangular. Corolla
white, glabrous outside; tube 1316 mm; lobes ovate-triangular,
ca. 2 mm. Drupes ca. 4 58 mm, with pedicels sometimes
elongated to 3 mm. Fl. MarMay, fr. OctJan.
Sparse or dense forests or thickets; 5001100 m. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Yunnan, Zhejiang
[Japan].
Naiki and Nagamasu (Amer. J. Bot. 91: 664671. 2004) reported
the breeding biology of this species as monomorphic with pin type
flowers (i.e., with stigmas exserted and anthers included).

3. Damnacanthus guangxiensis Y. Z. Ruan, Fl. Reipubl. Popularis Sin. 71(2): 320. 1999.
guang xi hu ci
Shrubs, height not noted. Branches terete, glabrous, with-

Forests on mountains; ca. 1200 m. Guangxi (Lingyun).

hai nan hu ci

Forests, forest margins; 8001800 m. Hainan.

5. Damnacanthus henryi (H. Lveill) H. S. Lo, Acta Phytotax. Sin. 17(3): 108. 1979.
yun gui hu ci
Canthium henryi H. Lveill, Repert. Spec. Nov. Regni
Veg. 13: 178. 1914; Prismatomeris brevipes Hutchinson; P.
henryi (H. Lveill) Rehder.
Shrubs or small trees, 1.55 m tall. Branches 4-angled to
flattened, pale brown, smooth, glabrous. Petiole of developed
leaves 25 mm, glabrous; leaf blade drying papery or leathery,
lanceolate, narrowly elliptic, elliptic, or elliptic-oblong, 513
14 cm, glabrous, base acute or cuneate and often decurrent,
margins entire and flat to usually thinly revolute, apex acute to
long acuminate; midrib thinly prominulous adaxially; secondary veins 57 pairs; stipules caducous, interpetiolar, triangular
to narrowly triangular, 11.5 mm, glabrous, acute. Inflorescences glabrous. Pedicels 23.5 mm. Calyx glabrous; hypanthium portion turbinate, 11.5 mm; limb ca. 0.8 mm, lobed for
2/33/4; lobes 4 or 5, narrowly triangular, sometimes separated
by subtruncate sinuses. Corolla white or pale purple, glabrous
outside; tube 912 mm, fenestrate at base; lobes 4 or 5, ovatelanceolate, 34 mm. Drupes 58 mm in diam.; pyrenes 2, subglobose. Fl. Oct, fr. DecFeb.
Dense forests on mountains; 12002500 m. Guangxi, Guizhou,
Yunnan.

96

RUBIACEAE

This species is here provisionally included in Damnacanthus

based on its inflorescence morphology. It was reported to be distylous
by Naiki and Nagamasu (Amer. J. Bot. 91: 664671. 2004).

6. Damnacanthus indicus C. F. Gaertner, Suppl. Carp. 18.

1805.
hu ci
Damnacanthus esquirolii H. Lveill; D. formosanus
(Nakai) Koidzumi; D. indicus var. formosanus Nakai; D. indicus var. lancifolius Makino; D. lancifolius (Makino) Koidzumi.
Shrubs, 0.31.5 m tall. Roots fleshy, moniliform. Branches
densely hispidulous to hirtellous, sometimes becoming glabrescent, terete or sometimes 4-angled, with numerous persistent
spines 320 mm. Petiole of developed leaves 0.53 mm, strigillose, hispidulous, or glabrescent; leaf blade drying stiffly papery to leathery and discolorous, ovate, cordiform, ellipticovate, elliptic, or broadly elliptic, 0.52(3) 0.51(1.5) cm,
adaxially glabrous, abaxially glabrous or sparsely hirtellous to
strigillose along veins, base obtuse to rounded, truncate, or cordulate, sometimes oblique, margins entire and flat, apex acute;
midrib thinly prominulous adaxially; secondary veins 2 or 3(or
4) pairs; stipules quickly fragmenting or caducous, interpetiolar,
narrowly to broadly triangular, 0.31 mm, strigillose to glabrescent, acute to glandular-fimbriate. Inflorescences strigillose to
hispidulous. Pedicels 0.58 mm. Calyx strigillose to glabrous;
hypanthium portion turbinate, 11.5 mm; limb 0.82 mm,
lobed for 1/44/5; lobes broadly triangular to narrowly triangular. Corolla white, glabrous outside; tube 79 mm; lobes
elliptic to lanceolate-elliptic, 2.55 mm. Drupes 46 mm in
diam. Fl. MarJun, fr. MarJan.
Sparse or dense forests on hills or mountains, rocky thickets; 100
1500 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan,
Jiangsu, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [N and
NE India, Japan, Korea].
Naiki and Nagamasu (Amer. J. Bot. 91: 664671. 2004) reported
that populations they studied are either monomorphic, with only pintype flowers, or distylous. Koidzumi (Acta Phytotax. Geobot. 3: 158.
1934) reported Damnacanthus lancifolius from Yunnan and treated D.
esquirolii as a synonym of that name; these names were apparently
overlooked by Y. Z. Ruan (in FRPS 71(2): 169. 1999).

7. Damnacanthus labordei (H. Lveill) H. S. Lo, Acta Phytotax. Sin. 17(3): 107. 1979.
liu ye hu ci
Canthium labordei H. Lveill, Repert. Spec. Nov. Regni
Veg. 13: 178. 1914; Lasianthus labordei (H. Lveill) Rehder;
Prismatomeris labordei (H. Lveill) Merrill ex Rehder; P.
linearis Hutchinson.
Small shrubs, 0.42 m tall. Roots fleshy, moniliform.
Branches 4-angled to flattened and usually with a well-developed longitudinal ridge descending from stipule midrib on each
side, yellow to pale brown, glabrous, without spines. Petiole of
developed leaves 26 mm, glabrous; leaf blade drying papery,
gray to brownish green adaxially, gray-green, brownish green,
or straw-yellow abaxially, lanceolate to lanceolate-linear, 521
0.62.5 cm, glabrous or sometimes pubescent along veins
adaxially, base cuneate or acute, margins entire or irregularly

serrulate and flat to thinly revolute, apex tapered and acuminate

with tip often flexuous; midrib thinly prominulous adaxially;
secondary veins 916 pairs; stipules caducous, interpetiolar, triangular, 0.51 mm, acute to glandular-fimbriate. Inflorescences
glabrous. Pedicels 23 mm. Calyx glabrous; hypanthium portion turbinate, ca. 1 mm; limb 0.51 mm, lobed for 1/41/2;
lobes broadly triangular. Corolla white to yellow, outside glabrous; tube 59 mm; lobes ovate, ca. 3 mm. Drupes ca. 8 mm in
diam. Fl. FebMar, OctDec, fr. SepDec.
Sparse or dense forests or thickets; 8001800 m. N Guangdong, N
Guangxi, Guizhou, Hunan, Sichuan, Yunnan [Vietnam].
This species was reported to be distylous by Naiki and Nagamasu
(Amer. J. Bot. 91: 664671. 2004).

8. Damnacanthus macrophyllus Siebold ex Miquel, Ann.

Mus. Bot. Lugduno-Batavi 3: 110. 1867 [macrophylla].
zhe wan hu ci
Damnacanthus indicus C. F. Gaertner f. macrophyllus (Siebold ex Miquel) Makino; D. indicus var. macrophyllus (Siebold
ex Miquel) Makino; D. indicus var. parvispinus Koidzumi; D.
major Siebold & Zuccarini var. macrophyllus (Siebold ex
Miquel) Maximowicz; D. major var. parvispinus (Koidzumi)
Koidzumi; D. major var. submitis Maximowicz ex Regel; D.
moniliformis Koidzumi; D. minutispinus Koidzumi; D. shanii
K. Yao & M. B. Deng.
Shrubs, 12 m tall. Roots fleshy, moniliform. Branches
puberulent to hispidulous, with 8 alternately thick and thin
striae, with few deciduous or persistent spines 26 mm. Petiole
of developed leaves 12 mm, glabrous or sparsely puberulent to
puberulent or hirtellous; leaf blade drying stiffly papery, ovate
to oblong-ovate, lanceolate, or oblong-lanceolate, 36(8) 1
2.5(3) cm, adaxially glabrous, abaxially glabrous or puberulent along veins, base cuneate to rounded, margins flat to thinly
revolute, apex shortly acuminate or acute; midrib thinly prominent adaxially; secondary veins 3 or 4(7) pairs; stipules persistent, interpetiolar, triangular, 0.31 mm, puberulent to glabrescent, acute to glandular-multifid or -fimbriate. Inflorescences strigillose to puberulent. Pedicels 12 mm. Calyx strigillose to glabrescent; hypanthium portion obconic, 1.21.5 mm;
limb 11.5 mm, lobed for 1/31/2; lobes triangular to ovate.
Corolla white, outside glabrous; tube 8(13) mm; lobes ovatetriangular, ca. 2 mm. Drupes ca. 5 mm in diam., with pedicels
sometimes elongated to 5 mm. Fl. AprJun, fr. OctDec.
Streamsides in sparse or dense forests on mountains; 8001000 m.
Anhui, Fujian, Guangdong, Guizhou, Yunnan, Zhejiang [Japan].
Naiki and Nagamasu (Amer. J. Bot. 91: 664671. 2004) reported
the breeding biology of this species as monomorphic with pin type
flowers.
Damnacanthus subspinosus var. salicifolius (M. B. Deng & K.
Yao, Bull. Bot. Res., Harbin 10(4): 2. 1990) belongs here but was not
validly published because two gatherings were designated as types
(Vienna Code, Art. 37.2).

9. Damnacanthus major Siebold & Zuccarini, Abh. Math.Phys. Cl. Knigl. Bayer. Akad. Wiss. 4(3): 177. 1846.
da luan ye hu ci

RUBIACEAE

Damnacanthus indicus C. F. Gaertner f. major (Siebold &

Zuccarini) Makino; D. indicus subsp. major (Siebold & Zuccarini) T. Yamazaki; D. indicus var. major (Siebold & Zuccarini) Makino ex Nakai.
Shrubs, 12 m tall. Roots fleshy, white or pale purple,
moniliform. Branches densely hispidulous to hirtellous when
young, sometimes becoming glabrescent, terete or sometimes
4-angled, with numerous spines 310 mm, persistent. Petiole of
developed leaves 14 mm, strigillose to hirtellous; leaf blade
drying papery, broadly ovate, ovate, or elliptic-ovate, 34
1.52 cm, adaxially glabrous, abaxially glabrous or sometimes
sparsely hispidulous along veins, base obtuse or rounded, margins flat and entire, apex acute; midrib thinly prominent adaxially; secondary veins 35 pairs; stipules quickly fragmenting or
caducous, interpetiolar, broadly triangular, 0.51 mm, strigillose
to glabrescent, obtuse to acute. Inflorescences strigillose. Pedicels ca. 1 mm. Calyx strigillose or puberulent; hypanthium portion obconic, 12 mm; limb ca. 2 mm, deeply lobed; lobes narrowly to broadly triangular. Corolla white, outside glabrous;
tube ca. 11 mm; lobes ovate-triangular, ca. 4 mm. Drupes 510
mm in diam. Fl. Apr, fr. winter.
Sparse forests and thickets on mountains; 600700 m. Guangdong, Zhejiang [Japan, Korea].
These plants were treated as a subspecies of Damnacanthus indicus in the Fl. Japan (3a: 224225. 1993) and also considered there to be
restricted to Japan and Korea.

10. Damnacanthus officinarum C. C. Huang in H. S. Lo, Acta

Phytotax. Sin. 17(3): 108. 1979.
si chuan hu ci
Shrubs, 12.5 m tall. Roots fleshy, moniliform. Branches
slightly flattened when young becoming terete, glabrous, without spines. Petiole of developed leaves ca. 5 mm, glabrous; leaf
blade drying leathery, brown or straw-yellow adaxially, strawyellow or olive-green abaxially, elliptic, elliptic-oblong, or oblong-lanceolate on upper part of stem, to linear on lower part of

97

stem, 513(16) 24(6) cm, glabrous, base cuneate to acute,

margins entire and flat to thinly revolute, apex acute to acuminate; midrib thinly prominent adaxially; secondary veins 68
pairs; stipules caducous, interpetiolar, triangular, ca. 1 mm, glabrous, acute. Inflorescences glabrous. Pedicels ca. 2 mm. Calyx
glabrous; hypanthium portion cupuliform, ca. 1.5 mm; limb
0.51 mm, undulate to lobed for ca. 1/2; lobes broadly triangular. Corolla in bud pale green, glabrous outside, 1012 mm.
Drupes 67 mm in diam. Fl. winterspring, fr. OctDec.
Thickets or forests on hills; 700900 m. Hubei, Hunan, Sichuan.
This species was reported to be distylous by Naiki and Nagamasu
(Amer. J. Bot. 91: 664671. 2004).

11. Damnacanthus tsaii Hu, Bull. Fan Mem. Inst. Biol. 6: 178.
1935.
xi nan hu ci
Shrubs, 13 m tall. Branches densely hirtellous, with
spines 425 mm, persistent. Petiole of developed leaves 12
mm, glabrous or pilosulous; leaf blade drying papery, lanceolate
or oblong-lanceolate, 37.5 0.92.4 cm, adaxially glabrous,
abaxially glabrous or hirtellous along veins when young, base
cuneate or rounded, margins entire to irregularly serrulate and
thinly revolute, apex acuminate; midrib thinly prominulous
adaxially; secondary veins 58(10) pairs; stipules quickly
deciduous, interpetiolar, broadly triangular, usually aciculate.
Pedicels ca. 2 mm. Calyx glabrous; hypanthium portion obconic, ca. 1 mm, lobed for ca. 1/2; lobes 4(or 5), triangular to
subulate-triangular. Corolla white, ca. 12 mm, outside glabrous;
lobes ovate-triangular. Drupes ca. 5 mm in diam. Fl. Apr, fr.
winterspring.
Forests, forest margins, roadsides, rocky mountains; 10002500
m. Sichuan, Yunnan.
Naiki and Nagamasu (Amer. J. Bot. 91: 664671. 2004) reported
that all flowers of this species seen resemble the long-styled form of
distylous species.

21. DENTELLA J. R. Forster & G. Forster, Char. Gen. Pl. 13. 1775.
xiao ya cao shu
Chen Tao (); Charlotte M. Taylor
Herbs, annual or perennial, prostrate, unarmed. Raphides present. Leaves opposite, usually relatively small, without domatia;
stipules persistent, interpetiolar and frequently fused to petioles, triangular, entire to erose, often scarious. Inflorescences terminal or
displaced to pseudoaxillary, 1-flowered, ebracteate, subsessile or pedunculate. Flowers bisexual, remarkably reduced. Calyx with
ovary portion sometimes papillose-villous with distinctive flattened trichomes; limb tubular, 5-lobed or -toothed. Corolla white to
pink, funnelform, inside frequently pubescent in throat; lobes 5, valvate in bud. Stamens 5, inserted at middle or perhaps base of
corolla tube, included; filaments short; anthers basifixed or perhaps dorsifixed, included. Ovary 2-celled, ovules numerous in each
cell inserted on axile subglobose placentas; stigmas 2, filiform, included. Fruit indehiscent, subglobose to ellipsoid, dry, papery,
sometimes papillose-villous with distinctive flattened trichomes, with calyx limb persistent; seeds numerous, small, angled; testa
granulate; endosperm fleshy; embryo minute.
About ten species: S Asia to Oceania, with one species apparently adventive in North America; one species in China.
The seeds and capsules of this species were described in detail by Terrell and Robinson (J. Bot. Res. Inst. Texas 1(1): 373384. 2007).

1. Dentella repens (Linnaeus) J. R. Forster & G. Forster, Char.

Gen. Pl. 13. 1775.
xiao ya cao

Oldenlandia repens Linnaeus, Syst. Nat., ed. 12, 2: 126;

Mant. Pl. 1: 40. 1767; Dentella matsudae Hayata; D. repens
var. grandis Pierre ex Pitard.

RUBIACEAE

98

Dwarf herbs, creeping, much branched, rather fleshy;

branches flattened to angled, glabrous or puberulent, often
rooting at nodes. Leaves subsessile to shortly petiolate; petiole
to 1[4] mm; blade drying stiffly papery, oblong-lanceolate,
oblanceolate, or spatulate, 47[10] 12[4] mm, glabrous or
sparsely hirtellous at least along margins and principal veins,
base cuneate to acute, margins flat, apex obtuse to acute;
secondary veins not visible; stipules triangular, ca. 1 mm, entire
to erose. Flowers solitary at forks of branchlets or rarely along
stems, subsessile or with pedicel to 1[2] mm. Calyx with
ovary portion subglobose, ca. 1 11.5 mm, densely papillosevillous with flattened trichomes; limb 11.2 mm, deeply lobed;
lobes narrowly triangular. Corolla white or pale yellow, 38[
12] mm, glabrous outside, sparsely villous inside throat; lobes
triangular-ovate, 12 mm, acute. Fruit compressed globose, 35
35 mm, densely villous with multicellular, transparent, flat-

tened, bulbous-tipped trichomes; seeds 0.30.5 mm. Fl. winter,

fr. summer.
Wet sites in fields; sea level or higher. Guangdong, Hainan, Taiwan, Yunnan (Xishuangbanna) [India, Indonesia, Malaysia, Myanmar,
Nepal, Philippines, Sri Lanka, Thailand, Vietnam; Australia, Oceania;
adventive in North America (E United States, Mexico)].
The Fl. Taiwan (ed. 2, 4: 252. 1998) described markedly larger
leaves, petioles, pedicels, and corollas than seen on specimens studied
for this work or found in other descriptions; these measurements are
included above provisionally in brackets. Two varieties were recognized
by Xing and Wu (Fl. Nansha Islands, 200201. 1996): Dentella repens
var. repens, with flowers ca. 3 mm, and D. repens var. grandis, with
flowers 68 mm; their description of the second variety corresponds to
the larger flower measurements given in Fl. Taiwan. These varieties
were subsequently synonymized by H. S. Lo (in FRPS 71(1): 21. 1999)
and are provisionally synonymized here.

22. DIODIA Linnaeus, Sp. Pl. 1: 104. 1753.

shuang jiao cao shu
Chen Tao (); Charlotte M. Taylor
Herbs [or sometimes small shrubs], annual or perennial, unarmed. Raphides present. Leaves opposite or sometimes apparently
verticillate due to clustered leaves on unexpanded axillary stems, sessile to shortly petiolate, without domatia; stipules persistent,
interpetiolar and fused to petioles, sheath truncate to rounded and often membranous, setose. Inflorescences axillary and sometimes
also terminal, glomerulate or capitate, few to several flowered, sessile, bracteate or bracts reduced. Flowers sessile, bisexual, monomorphic. Calyx limb deeply 2- or 4-lobed; lobes sometimes unequal in pairs. Corolla white, pink, or pale purple, funnelform or salverform, inside variously glabrous or pubescent; lobes 4, valvate in bud. Stamens 4, inserted in corolla throat, exserted; filaments
developed; anthers dorsifixed. Ovary 2-celled, ovules 1 in each cell, axile and attached at middle of septum; stigmas 2, capitate to
linear, exserted. Fruit schizocarpous or indehiscent, ellipsoid to obovoid, papery to cartilaginous, bony, or corky, with calyx limb
persistent; mericarps (i.e., cocci) 2, indehiscent, with 1 seed, ellipsoid to plano-convex; seeds ellipsoid-oblong, medium-sized, often
sulcate to convex on adaxial (i.e., ventral) surface; endosperm corneous; embryo straight; cotyledons broad; radicle hypogynous.
About 50 species: warm temperate and tropical America and Africa, with several American species naturalized in the Old World tropics; two
species (both introduced) in China.
Measurements and characters in brackets below are found in plants of other regions and are included here to aid future identifications of these
species, which may be introduced more than once in China and have more variation here than currently documented.

1a. Plants erect; fruit papery to bony, 33.5 mm, usually separating and dispersing as 2 indehiscent mericarps ................... 1. D. teres
1b. Plants prostrate to weakly ascending; fruit corky, 69 mm, usually not separating into mericarps .......................... 2. D. virginiana
1. Diodia teres Walter, Fl. Carol. 87. 1788.
shan dong feng hua cao
Borreria shandongensis F. Z. Li & X. D. Chen; Diodella
teres (Walter) Small; Spermacoce shandongensis (F. Z. Li & X.
D. Chen) Govaerts.
Herbs, annual, erect, to 30[50] cm tall; stems somewhat
flattened to slightly [or sharply] 4-angled, pilosulous [and/or
pilose to glabrescent]. Leaves sessile; blade drying papery,
linear-lanceolate, [8]2040 [1]35[7] mm, both surfaces
hispidulous, base obtuse [to rounded or subcordate], margin
weakly [to strongly] revolute, apex acute to acuminate; secondary veins indistinct; stipule sheaths truncate, 12.5 mm, pilosulous to glabrescent, with 59 setae 17 mm. Flowers 1[3] per
axil or 1[6] per node; bracts reduced. Calyx pilosulous to glabrescent; ovary portion obovoid, 0.50.8 mm; limb deeply
lobed; lobes lanceolate, ca. 1 mm, pilosulous to glabrescent.
Corolla pink [to pale purple or blue], funnelform, hispidulous to

glabrescent outside; tube [1.5]4[7] mm, glabrescent inside;

lobes elliptic-oblong, 0.52 mm. Fruit obovoid, [1.5]33.5
mm, papery to bony, hispid to hispidulous, separating into 2
mericarps each with a Y-shaped sulcus on adaxial surface; seeds
ca. 2.5 mm, yellowish brown when dry, longitudinally 1grooved. Fl. and fr. AugSep.
Disturbed, often degraded open ground. S Fujian (Jinmen), Shandong (Qingdao) [native to Antilles and North and South America; adventive in N Africa, Japan, Korea, and Madagascar].
Diodia teres has not been widely reported previously from Asia,
but it is known from Japan (Honshu, 18 Oct 1995, S. Tsuagaru & G.
Murata 22898, MO!) at a similar latitude to its occurrence in China and
probably should be expected elsewhere.

2. Diodia virginiana Linnaeus, Sp. Pl. 1: 104. 1753.

shuang jiao cao
Herbs, perennial, prostrate to weakly ascending, to 60 cm
tall; stems 4-angled, retrorse pilose or -pilosulous along angles.

RUBIACEAE

Leaves petiolate; petiole ca. 3 mm; blade drying papery, elliptic-lanceolate to oblanceolate, 2032 48 mm, both surfaces
glabrescent, base cuneate and sometimes oblique, apex acute;
secondary veins 4 or 5 pairs; stipule sheaths truncate to
rounded, 23 mm, glabrous, with 35 setae 14 mm. Flowers 1
or sometimes 2 or 3 per axil, 2 or sometimes 46 per node;
bracts reduced. Calyx villous to pilose; ovary portion obconic,
ca. 1 mm; lobes narrowly triangular-lanceolate, [2]57[10]
mm, often unequal on an individual flower. Corolla white, glabrous inside and outside; tube 5.56 mm; lobes triangular, 4
5[6] mm, sometimes pubescent adaxially. Fruit corky, ellipsoid, 69 46 mm, distinctly 8-ridged, pilose or villous to glabrescent, usually not separating into mericarps; seeds 56 23
mm, reticulate. Fl. and fr. AugSep.
Bamboo forest sides. Naturalized in Taiwan [native to C and E

99

North America; perhaps naturalized in Mexico and Central America;

adventive in Japan].
This species is rather widely but infrequently and locally naturalized in tropical and subtropical areas, especially near seaports (Taylor,
Monogr. Syst. Bot. Missouri Bot. Gard. 85(3): 22062284. 2001), and
probably should be expected elsewhere in Asia. The plants are usually
found in microsites with moving water, and apparently the fruit are frequently water-dispersed.
Hsieh and Chaw (Bot. Bull. Acad. Sin. 28: 4445. 1987) gave the
number of calyx lobes as 2; this is the first report of this condition for
Diodia virginiana, and their illustration of the flowers suggest that there
may at least sometimes be 4 lobes. They also gave some unusual measurements for the corolla of their plant, with the tube said to be 15 mm
and the limb to 18 mm wide; both of these are exceptionally large measurements that are unknown in other plants of this species.

23. DIPLOSPORA Candolle, Prodr. 4: 477. 1830.

gou gu chai shu
Chen Tao (); Charlotte M. Taylor
Shrubs or small trees, sometimes polygamo-dioecious, unarmed; branches sometimes superaxillary. Raphides absent. Leaves
opposite, sometimes with domatia; stipules persistent or deciduous, shortly united around stem, triangular. Inflorescences axillary or
sometimes supra-axillary, cymose to fasciculate or glomerulate, few to many flowered, sessile to pedunculate, bracteate. Flowers
subsessile to pedicellate, bisexual and monomorphic or unisexual. Calyx limb truncate to 4(or 5)-lobed or -denticulate. Corolla white,
pale green, or pale yellow, salverform, inside pubescent in throat and sometimes on lower part of lobes; lobes 4(or 5), convolute in
bud. Stamens 4(or 5), inserted in corolla throat, exserted; filaments short; anthers dorsifixed. Ovary 2-celled, ovules 13(6) in each
cell on axile placentas; stigma 2-lobed, shortly exserted. Fruiting pedicels often elongating notably. Fruit yellow, orange, or red,
baccate, subglobose or ellipsoidal-globose, fleshy or leathery, with calyx limb usually persistent; seeds several, medium-sized,
angled, subglobose, or somewhat flattened, embedded in pulp derived from placenta; hilum linear or comma-shaped; radicle
hypogynous.
About 20 species: tropical and subtropical Asia; three species (one endemic) in China.
This genus was reviewed by Robbrecht and Puff (Bot. Jahrb. Syst. 108: 114119. 1986), then in more detail by Ali and Robbrecht (Blumea 35:
279305. 1991). The description of the sexuality of the plants and flowers here follows Ali and Robbrecht (loc. cit.) and Puff et al. (Rubiaceae of
Thailand, 84. 2005).

1a. Leaves drying leathery or thickly papery, glabrous and rather shiny on both surfaces, abaxially with secondary
veins plane or thinly prominulous, higher order venation hardly or not visible, without domatia; petioles
glabrous ............................................................................................................................................................................... 1. D. dubia
1b. Leaves drying papery or thinly leathery, glabrescent, strigillose, puberulent, tomentulose, or hirtellous abaxially
or on both surfaces, abaxially with secondary veins prominent, higher order venation well marked and usually
thinly raised, usually with domatia; petioles glabrescent, strigillose, or tomentulose.
2a. Leaves abaxially pubescent along principal veins but glabrescent or very sparsely pubescent on blade;
stipules usually deciduous after distalmost 1 or 2 nodes ....................................................................................... 2. D. fruticosa
2b. Leaves densely tomentulose abaxially; stipules usually persistent ..................................................................... 3. D. mollissima
1. Diplospora dubia (Lindley) Masamune, Trans. Nat. Hist.
Soc. Formosa 29: 269. 1939.
gou gu chai
Canthium dubium Lindley, Bot. Reg. 12: t. 1026. 1826;
Diplospora buisanensis Hayata; D. tanakae Hayata; D. viridiflora Candolle; Tricalysia dubia (Lindley) Ohwi; T. lutea Handel-Mazzetti; T. viridiflora (Candolle) Masamune; T. viridiflora
var. buisanensis (Hayata) Yamamoto; T. viridiflora var. tanakae
(Hayata) Yamamoto.
Shrubs or trees, 112 m; branches terete to somewhat flattened, glabrous. Petiole 415 mm, glabrous; leaf blade drying

leathery or rarely thickly papery, and yellowish green to brown,

ovate-oblong, elliptic-oblong, elliptic, or lanceolate, 419.5
1.58 cm, both surfaces glabrous and rather shiny, base acute,
cuneate, or obtuse, sometimes slightly oblique, margins often
thinly revolute, apex acute to acuminate with tip often ultimately obtuse; secondary veins 511 pairs, without domatia;
stipules persistent on distalmost 1 or 2(4) nodes, ovate to
triangular, 58 mm, glabrous, densely white sericeous inside,
acute, keeled in upper part with keel often extending into an
arista 14 mm. Inflorescences subcapitate to congested-fasciculate, 610 mm, puberulent to strigillose, tomentulose, or puberulent, sessile or subsessile; bracts triangular, ca. 1 mm, frequently fused in pairs, acute; pedicels 14 mm. Calyx strigillose

RUBIACEAE

100

to glabrescent; ovary portion obconic to obovoid, 0.51 mm;

limb ca. 1 mm, 4-denticulate, sometimes also irregularly splitting. Corolla white or yellow, glabrous outside or sometimes
strigose on tube; tube 23 mm; lobes elliptic-oblong to narrowly elliptic, 34 mm, obtuse. Fruiting pedicels to 8 mm.
Berry red, subglobose, 49 mm in diam., sparsely strigillose to
glabrous; seeds dark red, subovoid, 36 34 mm. Fl. Apr
Aug, fr. MayFeb of following year.
Thickets or forests on hillsides, ravines, fields; near sea level to
1500 m. Anhui, Fujian, Guangdong, Guangxi, Hainan, Hunan, Jiangsu,
Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Vietnam].
This is one of the most commonly collected Rubiaceae species in
China.

2. Diplospora fruticosa Hemsley, J. Linn. Soc., Bot. 23: 383.

1888.
mao gou gu chai
Discospermum fruticosum (Hemsley) Kuntze; Tricalysia
fruticosa (Hemsley) K. Schumann ex E. Pritzel.
Shrubs or trees, 18(15) m tall; branches compressed to
terete, densely puberulent to pilosulous or strigillose sometimes
becoming glabrescent. Petiole 313 mm, strigillose or pilosulous to glabrescent; leaf blade drying papery or thinly leathery,
elliptic-oblong, oblong-lanceolate, obovate, oblanceolate, or
narrowly elliptic, 5.522 2.58 cm, adaxially glabrous or
along principal veins sometimes strigillose to puberulent,
abaxially hirtellous to strigillose at least on principal veins, base
acute, cuneate, or rarely rounded, sometimes slightly oblique,
apex acuminate with tip sometimes curved; secondary veins 7
13 pairs, in abaxial axils usually with well-developed pilosulous
domatia; stipules usually deciduous after distalmost 1 or 2
nodes, lanceolate to ovate, 48 mm, strigillose or puberulent to
glabrescent, acute, keeled in upper part or with ridges in form
of upside-down Y, keel extended into arista 0.52 mm. Inflorescences shortly cymose, 11.5 cm, subsessile, densely
puberulent to strigillose; bracts ovate to triangular, 0.51 mm,
mostly united in pairs; pedicels 0.53 mm. Calyx strigillose or
puberulent to glabrous; ovary portion obconic to ellipsoid, ca. 1

mm; limb 0.51 mm, lobed shallowly or for up to 1/2 its length;
lobes broadly triangular. Corolla white or seldom yellow, glabrous outside; tube 23.5 mm; lobes oblong-elliptic, ca. 3 mm,
obtuse to rounded. Fruiting pedicels to 10 mm. Fruit red, subglobose, 57 mm in diam., strigillose to glabrescent; seeds ca. 3
mm. Fl. MarMay, fr. JunFeb of following year.
Thickets or forests in ravines; 2002000 m. Guangdong, Guangxi,
Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Xizang, Yunnan [Vietnam].

3. Diplospora mollissima Hutchinson in Sargent, Pl. Wilson. 3:

401. 1916.
yun nan gou gu chai
Tricalysia mollissima (Hutchinson) Hu.
Shrubs or trees, 28 m tall; branches compressed to terete
or angled, densely tomentulose or hirtellous. Petiole 410 mm,
densely tomentulose to glabrescent; leaf blade drying papery or
thinly papery, elliptic-oblong, lanceolate, elliptic, or oblonglanceolate, 524 27.5 cm, adaxially glabrous except midrib
and sometimes principal veins tomentulose, abaxially tomentulose or hirtellous on principal veins and hirtellous to glabrous
on lamina, base cuneate, obtuse, or rounded, sometimes slightly
oblique, apex acute or acuminate with tip often ultimately obtuse; secondary veins 710 pairs, in abaxial axils with pilosulous domatia; stipules persistent on distalmost 13 nodes, triangular to ovate, 810 mm, densely tomentulose to glabrescent,
acute or acuminate, keeled in upper part with keel extending
into arista 36 mm. Inflorescences congested-cymose to glomerulate, 11.5 cm in diam., densely strigillose to tomentulose,
subsessile; bracts triangular, 0.51 mm, often united in pairs;
pedicels 0.10.5 mm. Calyx strigillose to tomentulose; ovary
portion turbinate to obconic, ca. 0.5 mm; limb 11.3 mm, undulate to shallowly lobed; lobes broadly triangular. Corolla
white, glabrous outside; tube 23 mm; lobes oblong-elliptic, 3
3.5 1.52 mm, obtuse. Fruiting pedicels to 8 mm. Fruit red,
subglobose, 57 mm in diam., strigillose to glabrescent; seeds
34 mm. Fl. MayJun, fr. JunDec.
Forests on mountains or at streamsides; 7001900 m. Yunnan.

24. DUNNIA Tutcher, J. Linn. Soc., Bot. 37: 69. 1905.

xiu qiu qian shu
Chen Tao (); Charlotte M. Taylor
Shrubs, unarmed; branches sometimes rather stout. Raphides present. Leaves opposite, without domatia; stipules persistent,
interpetiolar, generally triangular, acute to bifid. Inflorescences terminal, corymbose-cymose, several to many flowered, pedunculate,
with peduncles usually elongated and flexuous, bracteate, with some bracteoles on most inflorescences fused to base of hypanthium,
expanded, and petaloid thus appearing to be a calycophyll. Flowers subsessile to shortly pedicellate, bisexual, distylous. Calyx limb
4- or 5-denticulate, sometimes 1 or a few flowers with 1 petaloid calycophyll. Corolla yellow, salverform or funnelform, villosulous
inside; lobes 4 or 5, valvate in bud. Stamens 4 or 5, inserted in upper part of corolla tube, included or partially exserted; filaments
short; anthers apparently dorsifixed. Ovary 2-celled, ovules numerous, position of placentas unknown; stigma 2-lobed, included.
Fruit capsular, subglobose to obovoid, apically prolonged into a short beak, stiffly cartilaginous or leathery, septicidally dehiscent
into 2 valves with each valve sometimes later splitting into 2 parts, with calyx limb, petaloid bracts, and calycophylls persistent;
seeds small, flattened, with marginal wing membranous, irregular or lacerate; endosperm abundant; embryo minute.
Two species: China, India; one species (endemic) in China.
The stipitate petaloid structures on the inflorescence are similar to the petaloid calycophylls of other Rubiaceae genera; however, these appear to

RUBIACEAE

101

comprise two different morphological structures. Most of these structures appear to be inserted at the base of the ovary and thus can be considered
bracts, but some appear to be enlarged calyx lobes inserted above the ovary.
Dunnia was revised by Ridsdale (Blumea 24: 367368. 1979).

1. Dunnia sinensis Tutcher, J. Linn. Soc., Bot. 37: 70. 1905.

xiu qiu qian cao
Shrubs, 0.32.5 m tall; branches rather stout, subterete to
angled, puberulent to pilosulous sometimes becoming glabrescent with age. Petiole 0.72.5 cm, pilosulous or puberulent; leaf
blade drying papery to leathery, narrowly lanceolate, narrowly
elliptic, or oblanceolate, 723 16 cm, both surfaces puberulent to strigillose with pubescence denser on principal veins,
sometimes becoming glabrescent with age, abaxially epidermis
often with apparently enlarged cells, base acute to cuneate,
margins often thinly revolute, apex acuminate, acute, or rarely
obtuse; secondary veins 1117 pairs, extending to very close to
margins, with intersecondary veins often very well developed;
stipules ovate or triangular, 68 810 mm, strigillose to puberulent, acute or often bifid for up to 1/3. Inflorescences 812
35 cm, moderately to densely strigillose or pilosulous; peduncle 26 cm; branched portion 24 35 cm; bracts trian-

gular, 11.5 mm, petaloid bracts similar to calycophylls; pedicels to 3 mm. Calyx sparsely to densely puberulent or strigillose; ovary portion obconic, 11.5 mm; limb deeply lobed;
lobes deltoid, 0.30.5 mm; petaloid calycophyll white to cream,
elliptic-oblong to lanceolate, 25.5 12.3 cm, puberulent to
glabrescent, obtuse to rounded, with 3 longitudinal veins and
well-developed reticulate venation, with stipe 0.81.5 cm. Corolla outside sparsely to densely pilosulous or puberulent; tube
810 mm; lobes triangular-ovate, ca. 2 mm, acute to obtuse.
Capsule 35 mm in diam. including beak ca. 1 mm, smooth,
pilosulous to glabrescent; seeds 0.61 mm in diam. Fl. and fr.
AprNov.
Thickets or forests in ravines; 200900 m. Guangdong.
In conservation status, this species has been considered seriously
threatened due primarily to habitat destruction (Ge et al., Conservation
Genet. 3: 351362. 2002; Chiang et al., Molec. Biol. Evol. 19: 1367
1375. 2002).

25. DUPERREA Pierre ex Pitard in Lecomte, Fl. Indo-Chine 3: 334. 1924.

chang zhu shan dan shu
Chen Tao (); Charlotte M. Taylor
Shrubs or small trees, unarmed. Raphides absent. Leaves opposite or sometimes markedly anisophyllous and apparently ternate
due to reduced internodes grouping 2 leaves at 1 node and an apparently single leaf produced on a single very short axillary branch,
without domatia or these few, pilosulous, and rudimentary; stipules persistent, interpetiolar or sometimes shortly united around stem,
triangular, acuminate to aristate. Inflorescence terminal on principal branches or on reduced axillary branches, quickly becoming
overtopped by growth of 1 or 2 axillary branches thus appearing axillary or sometimes with subtending leaf caducous and then
appearing pseudoaxillary, corymbiform, many flowered, pedunculate, bracteate. Flowers pedicellate, bisexual, monomorphic. Calyx
limb deeply 5(or 6)-lobed. Corolla white, salverform with tube slender, glabrous inside; lobes 5(or 6), convolute in bud. Stamens 5(or
6), inserted in corolla throat, partially exserted; filaments reduced; anthers dorsifixed, bifid at base, with connective prolonged
apically into a short conical appendage. Ovary 2-celled, ovules 1 in each cell on peltate axile placentas; stigma subglobose to ellipsoid, long exserted. Fruit black, drupaceous but sometimes appearing baccate, compressed subglobose to somewhat didymous,
fleshy, with calyx limb deciduous; pyrenes 2, 1-celled, with 1 seed, plano-convex to oblate, papery or cartilaginous; seeds mediumsized, oblate or concave on ventral face; endosperm corneous; embryo small; cotyledons ovate; radicle basiscopic.
Two species: Cambodia, China, India, Laos, Myanmar, Thailand, Vietnam; one species in China.
Puff et al. (Rubiaceae of Thailand, 88. 2005) observed that Duperrea pavettifolia is quite variable morphologically and suggested that this genus
probably comprises only one variable species.

1. Duperrea pavettifolia (Kurz) Pitard in Lecomte, Fl. IndoChine 3: 334. 1924 [pavettaefolia].
chang zhu shan dan
Mussaenda pavettifolia Kurz, Forest Fl. Burma 2: 57.
1877 [pavettaefolia]; Ixora pavettifolia (Kurz) Craib.
Erect shrubs to small trees, 1.56 m tall; branches slightly
compressed, strigillose to strigose. Petiole 38 mm, strigillose;
leaf blade drying membranous to papery, oblanceolate to
obovate, elliptic, or elliptic-oblong, 725 38.5 cm, adaxially
glabrous to puberulent, abaxially puberulent on blade and strigillose along veins, base cuneate to obtuse or rounded, apex
acuminate; secondary veins 712 pairs; stipules ovate, 610

mm, strigillose to strigose, acuminate to shortly aristate. Inflorescence densely strigillose, hirsute, or strigose; peduncle 12.5
cm; branched portion 25 2.56 cm; bracts linear, 1.55 mm;
pedicels 35 mm. Calyx densely hirsute to strigillose; ovary
portion cylindrical to obconic, ca. 1 mm; limb with basal tubular portion 12 mm; lobes linear, 45 mm, acuminate. Corolla
densely strigillose outside; tube 1620 1.52 mm; lobes
oblong-elliptic to elliptic or suborbicular, 45 mm, obtuse to
rounded. Fruit 710 1012 mm; pyrenes 56 56 mm,
smooth to shallowly ridged. Fl. AprJun, fr. SepDec.
Broad-leaved forests at low to middle elevations; 1001100 m.
Guangxi, Hainan, Yunnan [Cambodia, Laos, Myanmar, Thailand, Vietnam].

RUBIACEAE

102

26. EMMENOPTERYS Oliver, Hookers Icon. Pl. 19: t. 1823. 1889.

xiang guo shu shu
Chen Tao (); Charlotte M. Taylor
Trees, unarmed. Raphides absent. Leaves opposite, with domatia; stipules caducous, interpetiolar, triangular, acute. Inflorescences terminal, corymbose to paniculiform, many flowered, pedunculate, bracteate with bracts reduced. Flowers pedicellate, bisexual, monomorphic, fragrant. Calyx limb 5-lobed nearly to base, with lobes imbricate in bud, some flowers of most inflorescences
with 1 lobe expanded into a white petaloid calycophyll. Corolla white or yellow, funnelform with tube notably narrowed near base,
glabrous inside; lobes 5, imbricate in bud. Stamens 5, inserted below corolla throat, included; filaments developed, tomentose; anthers dorsifixed, included. Ovary 2-celled, ovules numerous in each cell on peltate axile placentas; stigma capitate or shallowly 2lobed, included. Fruit capsular, oblong-cylindrical to fusiform with disk portion shortly prolonged into a rounded beak, septicidally
dehiscent from apex in apical half, woody, with calyx limb deciduous except with calycophyll sometimes persistent; seeds numerous,
medium-sized, fusiform, flattened, winged; testa spongy, reticulate-areolate; albumen rich; embryo minute; cotyledons cylindric.
One species: China.
Emmenopterys is currently known only from moist temperate E Asia, with its range wholly inside China, but fossils of the genus are known
from W North America; Latham and Ricklefs (in Ricklefs & Schluter, Spec. Diversity Ecol. Communities, 294314. 1993) discussed the
biogeography and putative history of this distribution.
Emmenopterys henryi is considered a rare species of conservation concern in China. Its habitat, population status, and distribution were studied
by Chen et al. (Acta Bot. Yunnan. 29: 461466. 2007) in one site, where the population appears to be declining; additional studies are cited there, and
its classification as a Grade II Chinese National Protected Wild Plant was reviewed by Liu (Gansu Sci. Technol. 19(10): 151152. 2003).

1. Emmenopterys henryi Oliver, Hookers Icon. Pl. 19: t.

1823. 1889.
xiang guo shu
Mussaenda cavaleriei H. Lveill; M. mairei H. Lveill.
Trees, deciduous, to 30 m tall, to 1 m d.b.h., with grayish
brown bark; branches angled to terete, often lenticellate, rather
stout, glabrous. Petiole 2080 mm, glabrous to hirtellous or
strigillose; leaf blade drying papery or leathery and paler below,
broadly elliptic, broadly ovate, or ovate-oblong, 630 3.5
14.5 cm, adaxially glabrous or sparsely strigose, abaxially glabrous to strigillose or hirtellous throughout or only on principal
veins, base acute to cuneate or obtuse, apex acute, abruptly acuminate, or rarely obtuse; secondary veins 59 pairs, in abaxial
axils with pilosulous domatia; stipules triangular-ovate, 610

mm, acute. Inflorescences hirtellous to glabrous; peduncle 2.5

5.5 cm; branched portion 518 620 cm; bracts caducous,
narrowly triangular, 18 mm, acute; pedicels 1.55 mm. Calyx
puberulent to usually glabrous; ovary portion ellipsoid, 34
mm; lobes suborbicular, 22.2 mm, ciliate, rounded; calycophyll white, pink, or pale yellow, blade drying papery or
leathery, spatulate to broadly elliptic or ovate, 1.58 16 cm,
parallel veined, on stipe 13 cm. Corolla outside densely
tomentulose; tube 1323 mm, narrow at base then quickly
dilated near middle; lobes suborbicular, 57 mm, densely ciliolate, rounded. Capsules 35 11.5 cm, smooth or longitudinally weakly ribbed; seeds 68 1.52 mm, broadly winged.
Fl. JunAug, fr. AugNov.
Forests in valleys; 4001600 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shanxi, Sichuan, Yunnan, Zhejiang.

27. FOSBERGIA Tirvengadum & Sastre, Biogeographica (Paris) 73(2): 88. 1997.
da guo qian shu
Chen Tao (); Charlotte M. Taylor
Trees or shrubs, unarmed; bark gray or reddish brown and scaly. Raphides absent. Leaves opposite but sometimes crowded at
stem apices, often with domatia; stipules generally persistent, interpetiolar or shortly united around stem, triangular. Inflorescences
terminal or displaced to pseudoaxillary, 27-flowered and cymose or reduced to 1 flower, pedunculate, bracteate. Flowers subsessile
to pedicellate, apparently bisexual and monomorphic. Calyx limb shallowly 5-lobed. Corolla white, salverform, fleshy to leathery,
inside variously pubescent; lobes 5, convolute in bud. Stamens 5, inserted in corolla throat, included; filaments short; anthers perhaps
dorsifixed. Ovary 2-celled, ovules numerous in each cell on axile placentas; stigma fusiform to clavate, shallowly bilobed, partially
exserted. Fruit baccate, thickly fleshy, globose to ellipsoid, smooth or infrequently ridged or tuberculate, color at maturity unknown,
with calyx limb tardily deciduous; seeds numerous, medium-sized to large, broadly angled, ovoid, or compressed, embedded in pulp.
At least five species: China, Myanmar, Thailand, Vietnam; three species (one endemic) in China.
Tirvengadum and Sastre (loc. cit.: 8794) implied without directly stating so that the flowers are hermaphroditic, and Puff et al. (Rubiaceae of
Thailand, 62. 2005) reported this condition tentatively. The pollen is reportedly simple, 3-porate; the ovary wall contains crystal sands present in
small clusters in mesocarp; and the testa cells have tube-like trabecular thickenings, [with] inner wall irregularly thickened.

RUBIACEAE

103

1a. Stems strigillose at least when young; leaves elliptic to oblong-elliptic, lanceolate-oblong, or oblanceolate,
9.515 24.5 cm; stipules keeled or ornamented throughout their length, with a straight median keel or
sometimes a ridge and/or line of pubescence in shape of an upside-down Y; calyx margins uniform, similar
in texture to rest of limb; fruit ca. 11 9 cm ............................................................................................................. 2. F. shweliensis
1b. Stems glabrous; leaves oblanceolate to obovate or broadly obovate, 1024 3.512 cm; stipules smooth or
keeled only in upper half, glabrous in lower half; calyx margins uniform to hyaline; fruit 46 45 cm.
2a. Leaves obovate to oblanceolate, with domatia in abaxial vein axils; flowers 35 in cymes, with developing
fruit solitary; calyx limb with margins hyaline and lobes aristate ........................................................................... 1. F. petelotii
2b. Leaves broadly obovate, without domatia; flowers solitary; calyx limb with margins uniform and lobes
acute but tips not prolonged ................................................................................................................................. 3. F. thailandica
1. Fosbergia petelotii Merrill ex Tirvengadum & Sastre, Biogeographica (Paris) 73(2): 89. 1997.
zhong yue da guo qian
Trees, 610 m tall; bark reddish brown; branches compressed to terete, glabrous. Petiole 510 mm, strigose; leaf
blade drying papery, oblanceolate to obovate, 1024 3.57
cm, adaxially glabrous, abaxially strigose at least on veins, base
cuneate to acute, apex acuminate to long acuminate; secondary
veins 811 pairs, in abaxial axils with pilosulous domatia; stipules triangular, 47 mm, glabrous, abaxially keeled in upper
half, acute and aristate, arista 23 mm. Inflorescences 315flowered, glabrous; peduncle 0.51 cm; bracts triangular to suborbicular, 13 mm; pedicels 815 mm. Calyx glabrous; ovary
portion obovoid to ellipsoid, ca. 4 mm; limb 1015 mm, lobed
for ca. 1/2; lobes triangular to narrowly triangular, marginally
hyaline, acute and aristate, arista 12 mm. Corolla white, outside apparently glabrous; tube 614 mm, inside glabrescent or
sparsely hairy at base; lobes narrowly triangular, 1317 mm.
Stigmas ca. 3 mm. Berries solitary, globose to ovoid-globose,
46 45 cm, sparsely to rather densely tuberculate or perhaps
smooth at least when young; seeds orbicular, laterally compressed, 68 56 2.53 mm, bony. Fl. MarApr.
Evergreen forests on mountain slopes or in valleys; 14001900 m
[as low as 11001200 m in Vietnam]. Yunnan (Jinping, Lchun, Maguan) [Vietnam].
Zhang et al. (Acta Phytotax. Sin. 45: 9093. 2007) noted that the
Chinese plants differ from the Vietnamese plants in having leaf blades
glabrous adaxially and hairy along the veins abaxially and corolla tubes
11.4 cm (vs. glabrous on both surfaces and corolla tube up to 6 mm in
the Vietnamese plants).

2. Fosbergia shweliensis (J. Anthony) Tirvengadum & Sastre,

Biogeographica (Paris) 73(2): 88. 1997.
rui li qian shu
Randia shweliensis J. Anthony, Notes Roy. Bot. Gard.
Edinburgh 18: 205. 1934; Aidia shweliensis (J. Anthony) W. C.
Chen.
Trees, 820 m tall; branches angled to terete, strigillose
becoming glabrescent. Petiole 510 mm, sparsely strigillose to
glabrous; leaf blade drying papery, elliptic to oblong-elliptic,
lanceolate-oblong, or oblanceolate, 9.515 24.5 cm, adaxially glabrous, abaxially sparsely strigillose or strigose along
principal veins, base cuneate to acute and sometimes slightly
inequilateral, apex shortly acuminate to acuminate; secondary

veins 811 pairs, usually with pilosulous domatia in abaxial

axils; stipules triangular to ovate, 510 mm, keeled or with
ridges in upside-down-Y pattern, strigillose, slenderly acute,
sometimes with 1 or 2 aristae 13 mm. Inflorescences 210flowered, strigillose to strigose or glabrescent; peduncle 11.5
cm; bracts linear-lanceolate, 24 mm, aristate; pedicels 618
mm. Calyx glabrous; ovary portion obovoid to ellipsoid, 34
mm; limb 813 mm, inside villosulous to tomentulose, lobed
for ca. 1/3; lobes triangular to linear-lanceolate, acute and terminating in arista 12 mm. Corolla creamy white to pale green
or pale yellow, outside glabrous; tube 2025 mm, inside tomentose; lobes lanceolate, 1526 mm, acuminate to subaristate,
marginally hyaline. Fruiting peduncle to 3.5 m. Berry subglobose to ellipsoid, ca. 11 9 cm, smooth; seeds ovoid to oblongangular, ca. 12 7 mm. Fl. MayJun, fr. May.
Open thickets or sparse moist primary broad-leaved forests in
valleys; 11002200 m. W Yunnan.
Li et al. (Acta Phytotax. Sin. 44: 707711. 2006) illustrated this
species in their figure 1 and reported that the flower buds are formed
nearly a year before the flowers open and that the fruit may take two
years to mature. They also detailed errors in the localities of Chinese
specimens given by Tirgengadum and Sastre in their protologue and
excluded one of the specimens treated by them, C. W. Wang 78281 (A),
which they included instead in Fosbergia thailandica. Additionally, they
reported that the Gaoligong Shan population of this species does not
appear to be reproducing itself, although they stop short of considering
this a species of conservation concern.

3. Fosbergia thailandica Tirvengadum & Sastre, Biogeographica (Paris) 73(2): 89. 1997.
tai guo da guo qian
Trees; bark thin, exfoliating in small pieces, grayish or
brownish on older parts when dry; branches glabrous. Petiole to
2 cm, glabrous; leaf blade broadly obovate, 1423 412 cm,
both surfaces glabrous except abaxially strigose on principal
veins, base acute to attenuate, apex acuminate to long acuminate; secondary veins 811 pairs, without domatia; stipules
ovate, 510 mm, glabrous, smooth at base and keeled in upper
part, obtuse to acute and aristate, arista ca. 2 mm. Flowers solitary. Calyx glabrous; ovary portion obovoid, ca. 1 cm; limb 35
mm, deeply lobed; lobes narrowly triangular. Corolla not seen.
Fruiting peduncle ca. 2.5 cm. Immature berry globose, 45 cm
in diam., glabrous, smooth.
Valleys, mixed woods, lower montane evergreen forests; 1500
1900 m. Yunnan (Jinghong) [N Thailand (Nan)].

104

RUBIACEAE

28. GALIUM Linnaeus, Sp. Pl. 1: 105. 1753.

la la teng shu
Chen Tao (); Friedrich Ehrendorfer
Subshrubs to perennial or annual herbs. Stems often weak and clambering, often notably prickly or sticky (i.e., retrorsely
aculeolate, velcro-like). Raphides present. Leaves opposite, mostly with leaflike stipules in whorls of 4, 6, or more, usually sessile
or occasionally petiolate, without domatia, abaxial epidermis sometimes punctate- to striate-glandular, mostly with 1 main nerve,
occasionally triplinerved or palmately veined; stipules interpetiolar and usually leaflike, sometimes reduced. Inflorescences mostly
terminal and axillary (sometimes only axillary), thyrsoid to paniculiform or subcapitate, cymes several to many flowered or infrequently reduced to 1 flower, pedunculate to sessile, bracteate or bracts reduced especially on higher order axes [or bracts sometimes leaflike and involucral], bracteoles at pedicels lacking. Flowers mostly bisexual and monomorphic, hermaphroditic, sometimes
unisexual, andromonoecious, occasionally polygamo-dioecious or dioecious, pedicellate to sessile, usually quite small. Calyx with
limb nearly always reduced to absent; hypanthium portion fused with ovary. Corolla white, yellow, yellow-green, green, more rarely
pink, red, dark red, or purple, rotate to occasionally campanulate or broadly funnelform; tube sometimes so reduced as to give
appearance of free petals, glabrous inside; lobes (3 or)4(or occasionally 5), valvate in bud. Stamens (3 or)4(or occasionally 5),
inserted on corolla tube near base, exserted; filaments developed to reduced; anthers dorsifixed. Inferior ovary 2-celled,
didymous, ovoid, ellipsoid, or globose, smooth, papillose, tuberculate, or with hooked or rarely straight trichomes, 1 erect and axile
ovule in each cell; stigmas 2-lobed, exserted. Fruit on pedicels sometimes elongating during development, green, gray, or infrequently white (to red, orange, or black), mostly dry to leathery schizocarps, infrequently spongy, rarely fleshy and berrylike, ellipsoid to subglobose; schizocarps separating into 2 indehiscent mericarps, each with 1 seed, subglobose, ellipsoid-oblong, or reniform,
smooth and glabrous to tuberculate and/or covered with trichomes often hooked and clinging; seeds small, grooved ventrally (i.e.,
adaxially); testa membranous; endosperm corneous; embryo curved; cotyledons leaflike; radicle terete, inferior.
More than 600 species: worldwide, mostly in meridional to temperate but also in alpine and arctic regions or in subtropical and tropical zones at
higher elevations; 63 species (23 endemic, four of unconfirmed occurrence) in China.
Galium is by far the largest and most widespread genus within the tribe Rubieae (subfamily Rubioideae). According to the most recent contributions (Natali et al., Opera Bot. Belg. 7: 193203. 1996; Ehrendorfer et al., Fl. Iranica 176: 1287. 2005; Bremer & Eriksson, Int. J. Pl. Sci. 170:
766793. 2009; Soza & Olmstead, Taxon 59: 755771. 2010), this tribe is closest to Theligoneae, Putorieae, and Paederieae, and includes the
following genera treated (or mentioned) in the present flora: Asperula, Cruciata Miller, Galium, Kelloggia, Leptunis, Microphysa, Phuopsis, Rubia,
and Sherardia Linnaeus.
So far, the genera Cruciata and Sherardia have not been found in China yet but may be expected there because of their partly weedy character
and widely adventive occurrence. They are included in the key below for future reference but not among the full generic presentations. Sherardia
arvensis Linnaeus is widely distributed in warm temperate and high-elevation tropical regions and can be separated from Asperula, Phuopsis,
Leptunis, or Galium by its terminal capitate inflorescences enclosed by leaflike bracts, its clearly developed calyx with 6 acute lobes, and its pink or
violet corollas with well-developed funnelform tubes and 4 lobes.
Among the few Cruciata species, the W Eurasiatic C. pedemontana (Bellardi) Ehrendorfer appears occasionally as an adventive in warm
temperate regions. It is common, e.g., in SE North America, and could be found in China too. Cruciata can be separated from Galium by its flowering
stems with vegetative apices and the inflorescences consisting only of lateral axillary cymes on middle and lower stem nodes. These cymes are equal
to or shorter than the subtending leaves when fully developed. In contrast, the inflorescences are mostly terminal and axillary and longer than the
leaves in Galium.
The characters relevant for the taxonomy of Galium and other Rubieae deserve some comments. Life and growth forms are important,
particularly with respect to the differentiation into half-shrubs, herbaceous perennials, and annuals. Stem and leaf posture, consistency, shape, and
indumentum (e.g., pubescent or retrorsely aculeolate with recurved microhairs) are often quite diverse and may vary within species or even
populations. The true leaves are always opposite and 2, but interpetiolar stipules may vary from inconspicuous and divided or simple to enlarged and
leaflike, forming whorls of 4 or up to 6 and more. During seedling and shoot development all these taxa pass through the 2- and 4-whorl stage, but
some taxa remain at this stage, while others continue to develop more numerous whorl elements toward the middle of their stems. This is a most
informative differential character within Rubieae. Other relevant features relate to leaf shape, venation, texture, and particularly indumentum. Here,
the presence of longer or shorter microhairs (use a lens) on surfaces and particularly margins as well as their forward or backward direction is of taxonomic importance.
Other morphological characters decisive for Rubieae taxonomy concern the inflorescences (e.g., the position and structure of the cymes). Flower
shape is essential for the traditional separation of the genera Asperula (with salverform, funnelform, or cup-shaped corollas) and Galium (with rotate
corollas). It is now clear that there are transitions between these character states and that even closely related taxa may differ in this respect. So far, it
has been possible to provisionally maintain Asperula and Galium by the transfer of obviously misplaced taxa and by using the presence or absence of
bracts and bracteoles as a differential character for the two genera (see Ehrendorfer et al., loc. cit. 2005).
The indumentum of ovaries and fruit as well as fruit consistency also vary strongly within Rubieae. Informative are, for example, fleshy
berries (as in Relbunium Bentham & J. D. Hooker, Rubia, and certain Galium taxa) vs. dry schizocarps or the presence vs. absence of hairiness and
whether the trichomes are hooked (i.e., the fruit disperse as stick-tights on animals) vs. straight. However, the distinction between all these structures
is arbitrary, and there are even transitions between trichomes and tuberculate protuberances of various shapes as well as between hairy and glabrous.

RUBIACEAE

105

All this is well illustrated by Yang and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: f. 1. 1998). Furthermore, ovary and fruit indumentum and surface
structures may change during development and sometimes vary genetically within species or even within populations, as in several Galium species. In
general, authors in other regions have documented infraspecific variation from glabrous to densely hairy or tuberculate fruit but traditionally have only
separated plants with hooked trichomes into different species. However, intrepid Chinese authors have easily combined these latter morphotypes, e.g.,
in G. dahuricum sensu W. C. Chen (in FRPS 71(2): 255. 1999), whereas Fl. Japan (3a: 238239. 1993) distinguished G. manshuricum on the basis of
this character. Only careful studies and field observations can clarify such cases, as in G. spurium, where the infraspecific variation of fruit, either
smooth, tuberculate, or covered with hooked hairs, has been proven.
Further differential characters for the taxonomy of Rubieae come from the fields of palynology (e.g., number of colpi), karyology (e.g.,
deviations from the normal chromosome base number x = 11 in Asperula sect. Cynanchicae (Candolle) Boissier with x = 10 or in Galium sect.
Aparinoides (Jordan) Grenier with x = 12; common occurrence of polyploidy), and reproductive biology. Most of the perennial Rubieae taxa have
conspicuous hermaphroditic or andromonoecious flowers and inflorescences and are insect-pollinated and self-incompatible outbreeders (e.g.,
Phuopsis or G. boreale and G. verum). Nevertheless, for several annuals with small and inconspicuous flower aggregates selfing and autogamy have
been documented (e.g., G. aparine, G. spurium, and Sherardia arvensis). Furthermore, polygamodioecy and dioecy occur in some groups (e.g., G.
elegans). Up to now, only few and insufficient data from all these fields are available for Asian Rubieae species and have not been mentioned in
FRPS. Nevertheless, such data are significant and will have to be addressed in more detailed future systematic Rubieae studies from this region.
The -taxonomy of Rubieae in E Asia is still in a problematic state. A general survey of the collections at the herbaria KUN, MO, PE, W, and
WU has revealed the existence of many very polymorphic, complex, and insufficiently understood species groups. Therefore, the present treatment
has to be regarded as provisional.
A particularly critical case concerns several Galium species described by H. Lveill from 19041917 (see Lauener & Ferguson, Notes Roy.
Bot. Gard. Edinburgh 32: 103115. 1973). These descriptions are most fragmentary and the relevant types are not yet studied sufficiently (but see
Mill, Edinburgh J. Bot. 53: 193213. 1996). Relevant taxa in alphabetical order are G. blinii (see under that name), G. bodinieri (see under G. blinii and
G. rebae), G. cavaleriei (see under G. asperifolium), G. comarii (see under G. dahuricum), G. esquirolii (see under G. asperifolium), G. hongnoense (see
under G. spurium), G. mairei (see under G. elegans), G. martini (see under G. bungei), G. quinatum (see under G. blinii), G. remotiflorum (see under G.
bungei), and G. venosum (see under G. bungei).
The treatment of Galium for the Flora of Taiwan by Yang and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: 101117. 1998; Fl. Taiwan, ed. 2, 4:
254259. 1998) is not satisfactory in several aspects: keys and descriptions are rather idealized and lack carefully observed ranges of morphological
variation for the taxa; species are circumscribed more narrowly and based on different characters than used by other authors in the region (e.g.,
presence vs. absence of leaf indumentum is considered variable within species by most other authors); the treatment is not well reconciled with
continental Galium taxonomy (e.g., there are no references to the Russian floras, and names synonymized by others are used without explanation); and
at least two names based on types from Taiwan are missing.
With respect to a more natural and general taxonomic classification of the Rubieae and Galium, a number of recent morphological,
karyological, palynological, and particularly DNA-analytical studies (e.g., Natali et al., loc. cit.; Robbrecht & Manen, Syst. & Geogr. Pl. 76: 85146.
2006; Bremer & Eriksson, loc. cit.; Soza & Olmstead, loc. cit.) are available. They show that Theligonum should be placed into a separate tribe
(Theligoneae), that the tribe Rubieae is monophyletic, and that Kelloggia (as subtribe Kelloggiinae, still with normal Rubiaceae stipules, calyx teeth,
and 3-colpate pollen, but already with hooked trichomes on the dry mericarps) occupies a basal position in Rubieae. The Central American genus
Didymaea J. D. Hooker (still with normal stipules but with the calyx already lacking, 5-coplate pollen, and seeds separating from the fleshy pericarp)
represents a link to the genus Rubia in the true Rubiinae. Their stipules are nearly always leaflike, the pollen is polycolpate, and the seeds never
separate from the pericarp. Rubia, a well-circumscribed and certainly monophyletic genus, is always perennial, has 5-lobed corollas, and berrylike
fruit.
The remaining Rubiinae are also monophyletic as a whole, but their traditional genera Asperula, Bataprine Nieuwland, Callipeltis Steven,
Crucianella Linnaeus, Cruciata, Galium, Leptunis, Mericarpaea Boissier, Microphysa, Phuopsis, Relbunium, Sherardia, Valantia Linnaeus, and
Warburgina Eig are all essentially interdigitated. They are difficult to separate and can hardly be brought into concordance with available phylogenetic
data. These advanced Rubiinae tend to develop more and more apomorphic character profiles, i.e., change from perennial to annual, increase in
numbers of leaflike stipules from 4 to numerous, loss of bracts and prophylls in the inflorescences, reduction from 5-lobed to (3 or)4-lobed corollas,
specialization of mericarps, etc. As shown by the most comprehensive phylogram available so far (Natali et al., loc. cit.: f. 2; Soza & Olmstead, loc.
cit.: f. 1, 2) and new findings (unpubl.), these more apomorphic Rubiinae form a polytomy or a grade with seven parallel clades. The most basal clade
(1) consists of the monotypic Galium sect. Cymogalia Pobedimova only. The following Sherardia clade (2) includes Crucianella, Phuopsis,
Sherardia, and several sections of Asperula together with Leptunis. Separate clades are formed by G. sect. Depauperata Pobedimova (3), A. sect.
Glabella Grisebach, including G. sect. Aparinoides (4), and A. sect. Asperula (5). The Cruciata clade (6) consists not only of the genera Cruciata and
Valantia but also of all sections of Galium (including the traditional genera Bataprine, Microphysa, and Relbunium) that form whorls of 2 leaves and
normally not more than 2 even-sized leaflike stipules. Finally, the G. sect. Galium clade (7) comprises this and various other sections of Galium, which
regularly develop whorls of leaves and leaflike stipules with 5 to more elements.
From the above data and the fact that a number of major groups of Rubiinae have not been DNA-analyzed yet, it is obvious that it is still difficult
and partly impossible to harmonize DNA-supported clades with the traditional genera and sections. Thus, extensive changes are expected for generic
and sectional circumscriptions within Rubiinae in the future. Therefore, we refrain from taxonomic changes for the present flora, list taxa in alphabetical order, and only supplement phylogenetic comments. Thus, the present treatment in principle follows FRPS (71(2): 216286. 1999), mainly
based on Pobedimova et al. (Fl. URSS 23: 287381. 1958), but also considers Ehrendorfer et al. (loc. cit. 2005). In order to make comparison with
available phylogenetic data and present infrageneric classification easier, relevant information is inserted as a Taxonomic Conspectus before the
individual species descriptions. It was not until this volume was ready for the press that the need for the nomen novum, Galium glabriusculum, was
discovered; therefore, this species alone is outside of the alphabetical order.

RUBIACEAE

106

Here the key to species of Galium is extensively revised from that of FRPS. It includes all of the Chinese Galium species with full ranges of
differential character variation. Furthermore, it keys out all other Rubieae genera which are easily confused with Galium and are documented or can
be expected in China. Details on the genera Asperula, Leptunis, Microphysa, Phuopsis and Rubia can be found where they are listed in alphabetical
order, references to Cruciata and Sherardia appear in the comments above.
Several species are keyed out more than once in the present key because they are circumscribed by combinations of characters rather than by
unique features. Furthermore, many Galium species are markedly variable because of genetic differentiation (e.g., G. bungei, G. elegans) but also
because of phenetic plasticity due to different environmental conditions. References to the number of leaves and leaflike stipules in whorls as well as
leaf measurements refer to middle stem regions. Measurements of organs with hairy surfaces (e.g., leaves, fruit, and mericarps) here apply to the solid
surface of the structure and do not include the trichomes. The terms leaf whorl, ovary, and uncinate trichome follow common usage in Galium.
Infraspecific taxa are adopted from FRPS in order to facilitate future and more detailed work on this group and comparison with other floras. They are
not included in the following main key but are subordinated under the relevant species in alphabetical order and keyed out there.

Taxonomic conspectus of the Rubieae (excluding Kelloggia and Rubia)

In FRPS (71(2): 216286. 1999) the taxa of Galium were placed in the following sections (designated here by capital letters): G. sect.
Depauperata (A), G. sect. Aparine (B), G. sect. Pseudaparine Lange (C), G. sect. Cymogalia (D), G. sect. Trachygalium (E), G. sect. Leptogalium
Lange (F), G. sect. Platygalium (G), G. sect. Galium (H), G. sect. Leiogalium (I), G. sect. Trichocarpa (Pobedimova) Pobedimova (J), G. sect. Asperuloides Pobedimova (K), and G. sect. Brachyantha (Boissier) Pobedimova (L). Some species of uncertain position were classified as dubious (M).
Species accepted in the present treatment but lacking in FRPS are designated as (Z). For comparison, a current (e.g., Jelenevsky et al., Novosti Sist.
Vyssh. Rast. 35: 174187. 2003; Ehrendorfer et al., Fl. Iranica 176: 1287. 2005), DNA-analytically supported (e.g., Natali et al., Opera Bot. Belg. 7:
193203. 1996; Soza & Olmstead, Taxon 59: 755771. 2010; Ehrendorfer, unpubl.) but still provisional taxonomic conspectus is presented below. It
lists all species here accepted under their clades and sections. For the clades 16 one should compare the comments above, for the sections compare
Ehrendorfer et al. (loc. cit.). The placement of Galium species into sections (or lack of placement) by FRPS is indicated by showing the relevant letters
used above in parentheses after the species names.
Clade 1
Galium sect. Cymogalia Pobedimova s.s.
39. Galium paradoxum (D)
Clade 2
Phuopsis stylosa
Sherardia arvensis
Leptunis trichodes
Asperula oppositifolia
Clade 3
Galium sect. Depauperata Pobedimova
16. Galium exile (A; incl. G. songaricum
sensu FRPS)
Clade 4
Galium sect. Aparinoides (Jordan) Grenier
27. Galium karakulense (E)
25. Galium innocuum (as G. trifidum: E;
incl. G. palustre)
Clade 5
Asperula sect. Asperula
Asperula orientalis
Clade 6
Galium sect. Platygalium W. D. J. Koch s.l.
9. Galium bungei (E; incl. G. martini:
M)
45. Galium salwinense (E)
12. Galium crassifolium (A)
31. Galium linearifolium (E)
21. Galium hirtiflorum (Z)
20. Galium glandulosum (A)
18. Galium forrestii (D)
44. Galium rupifragum (Z)
35. Galium morii (D)
54. Galium tarokoense (A)
34. Galium minutissimum (M)

36. Galium nankotaizanum (M; incl. G.

maborasense)
47. Galium serpylloides (A)
29. Galium kinuta (G)
24. Galium hupehense (M)
30. Galium kunmingense (Z)
17. Galium formosense (as G. kwanzanense: M)
15. Galium elegans (D)
63. Galium yunnanense (M)
26. Galium kamtschaticum (D)
10. Galium chekiangense (as G. nakaii:
G)
Microphysa elongata
40. Galium platygalium (K)
32. Galium maximoviczii (K)
7. Galium boreale s.l. (G)
60. Galium turkestanicum (M)
Clade 7
[Galium s.s.]
Galium sect. Hylaea (Grisebach) Ehrendorfer
s.l.
37. Galium odoratum (J)
4. Galium asperuloides (A)
22. Galium hoffmeisteri (as subsp. of G.
asperuloides: A)
14. Galium echinocarpum (A)
53. Galium takasagomontanum (M)
59. Galium triflorum (A)
58. Galium trifloriforme (Z)
Galium sect. Trachygalium K. Schumann s.l.
48. Galium sichuanense (Z)
13. Galium dahuricum (as G. davuricum: F; incl. G. comarii, G. niewerthii, G. pseudoasprellum)
56. Galium tokyoense (as var. of G.
davuricum: F)

41. Galium prattii (M)

52. Galium taiwanense (M)
3. Galium asperifolium (I)
6. Galium blinii (as syn. of G. asperifolium
var.: I; incl. G. quinatum: M)
51. Galium sungpanense (A)
42. Galium pusillosetosum (A)
1. Galium acutum (M)
43. Galium rebae (Z)
33. Galium megacyttarion (Z)
5. Galium baldensiforme (A)
49. Galium glabriusculum (A)
28. Galium karataviense (as G. rivale s.l.:
K)
61. Galium uliginosum (F)
Galium sect. Leiogalium Ledebour
38. Galium paniculatum (M; incl. G. xinjiangense: J)
Galium sect. Orientigalium Ehrendorfer
8. Galium bullatum (I)
Galium sect. Galium
23. Galium humifusum (L)
62. Galium verum (H)
11. Galium consanguineum (as G. majmechense: H)
46. Galium saurense (M)
Galium sect. Aparine W. D. J. Koch s.s.
50. Galium spurium (as G. aparine var.
tenerum: B)
2. Galium aparine (B)
Galium sect. Kolgyda Dumortier s.s.
57. Galium tricornutum (as G. tricorne: B)
Galium sect. Microgalium Grisebach
19. Galium ghilanicum (Z)
55. Galium tenuissimum (C)

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107

Key to species of Galium and to related genera of the Rubieae

1a. Interpetiolar stipules inconspicuous, multifid or fimbriate, not leaflike and not forming whorls with true
leaves; corolla funnelform, (4 or)5-lobed; ovary and dry mericarps with hooked trichomes ........................ Kelloggia (see p. 183)
1b. Interpetiolar stipules mostly leaflike and in whorls with true leaves, rarely reduced.
2a. Corolla lobes regularly 5; fruit fleshy, mericarps berrylike, 2(or 1, by non-development), often
dispersed together ................................................................................................................................................. Rubia (see p. 305)
2b. Corolla lobes usually 4 (rarely 3); fruit dry or leathery, mericarps mostly 2, nearly always separating
for dispersal.
3a. Leaves in middle stem region opposite, with stipules reduced or leaflike and in whorls of
4 but then always clearly smaller than true leaves.
4a. Corolla pink, funnelform, with well-developed tube longer than lobes; fruit smooth ........ Asperula oppositifolia (see p. 78)
4b. Corolla white, rotate, with tube shorter than lobes; fruit with uncinate trichomes.
5a. Perennial herbs; leaves 530 523 mm, obtuse to truncate at base, on petioles
1.510 mm; flowers 311 in cymes; corolla with 4 lobes ......................................................................... 39. G. paradoxum
5b. Annual herbs; leaves 212 14 mm, acute to cuneate at base, subsessile or on short petioles;
flowers solitary at each node; corolla mostly with 3 lobes .................................................................................. 16. G. exile
3b. Leaves in middle stem region opposite and with very similar leaflike stipules in whorls of 416.
6a. Leaf apex rounded, obtuse, or blunt, never acute or with a hyaline mucro; leaves in whorls
of 46, linear to broadly oblong, 1-nerved, dried blackening; ripe mericarps globose, didymous
and only with a short zone of contact, glabrous; corolla cup-shaped to slightly campanulate,
3- or 4-merous.
7a. Inflorescences with many-flowered cymes; corolla 4-lobed, 2.54 mm in diam.; leaves mostly
1520 58 mm ....................................................................................................................................... 27. G. karakulense
7b. Inflorescences with 13(or 4)-flowered cymes; corolla mostly 3-lobed, 11.8 mm in diam.; leaves
mostly 38 12 mm ................................................................................................................................... 25. G. innocuum
6b. Leaf apex mostly acute, often with a hyaline mucro; leaves in whorls of 416, sometimes broader
and with 35 palmate nerves; ripe mericarps ovoid to subglobose, with a longer zone of contact
and with diverse surface structures; corolla diverse, but often rotate and always 4-merous.
8a. Leaves and leaflike stipules in middle stem region never in whorls of more than 4 (if
rarely in whorls of up to 6 then leaves with 35 palmate principal veins), from linear
to broadly ovate.
9a. Stem apex vegetative, with few- to several-flowered lateral cymes only in leaf axils
and shorter than or equal to subtending leaves, nodding in fruit .................................. Cruciata (see comments above)
9b. Stem apex usually floriferous, with terminal and axillary cymes, often longer than
subtending leaves and mostly not nodding in fruit.
10a. Condensed plants of rocks or high elevations; stems usually less than 10 cm tall,
glabrous or with spreading (but never retrorse) hairs; leaves mostly ovate,
(1)38(20) (0.8)24(10) mm, with 13 main veins; corolla rotate,
often only 1.52 mm in diam.; mericarps with spreading (rarely appressed)
hooked or straight trichomes.
11a. Mericarps with straight hairs, 22.5 mm in diam.; stems mostly pilose or hirtellous.
12a. Fruiting pedicels straight; Xizang ............................................................................................. 47. G. serpylloides
12b. Fruiting pedicels nodding; Taiwan ....................................................................................... 36. G. nankotaizanum
11b. Mericarps with weakly to strongly curved and uncinate trichomes; stems partly glabrous.
13a. Corolla ca. 3 mm in diam.; stems glabrous .................................................................. 53. G. takasagomontanum
13b. Corolla 1.22 mm in diam.; stems hairy or glabrous.
14a. Stems hairy.
15a. Leaves ovate to broadly lanceolate, acute, up to 3.5 mm wide; Yunnan ............................ 44. G. rupifragum
15b. Leaves broadly elliptic to obovate, obtuse and mucronate, up to 10 mm wide;
Taiwan .................................................................................................................................. 17. G. formosense
14b. Stems glabrous; Taiwan.
16a. Leaves very small, only 0.81 mm wide, with 1 main vein only; fruit hairs
spreading .......................................................................................................................... 34. G. minutissimum
16b. Leaves wider, with 1 or 3 main veins; fruit hairs appressed.
17a. Leaves with 3 main veins; corolla only ca. 1.2 mm in diam. .................................................... 35. G. morii
17b. Leaves with 1 main vein; corolla ca. 2 mm in diam. ........................................................ 54. G. tarokoense
10b. Taller plants, usually of lower elevations with larger leaves (if plants condensed then stem
hairs retrorsely curved or fruit hairs appressed but not hooked).

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RUBIACEAE

18a. Leaves with 1 principal vein or 2 lateral veins only weakly visible and not extending past
middle of blade.
19a. Open corollas funnelform, 2.53 mm, tube somewhat shorter than lobes; fruit
with pericarp smooth to granular, becoming slightly inflated, enclosing
both mericarps at dispersal ................................................................................ Microphysa elongata (see p. 216)
19b. Open corollas rotate, fused basal part much shorter than lobes; mericarps
clearly separated.
20a. Stems strigose-hirsute, with hairs retrorse (but not retrorsely aculeolate);
leaves ovate or elliptic to linear-lanceolate, broadest in middle, at lower side
usually with glandlike spots; flowers unisexual, usually yellowish, greenish,
or reddish; fruit normally with uncinate trichomes.
21a. Plants usually less than 15 cm tall, strongly branched from base; leaves often less
than 8 mm, mostly glabrescent or glabrous, subleathery; inflorescences with
few-flowered, bracteate cymes .......................................................................................... 20. G. glandulosum
21b. Plants usually more than 15 cm tall, little branched; leaves usually longer than 8 mm,
hairy on both sides; inflorescences paniculate to corymbiform, little bracteate.
22a. Leaves linear-elliptic to narrowly lanceolate, mostly 817 12.5 mm, dried rather
papery; inflorescence paniculate ....................................................................................... 21. G. hirtiflorum
22b. Leaves ovate-elliptic, mostly 812 35 mm, dried rather subleathery; inflorescence
corymbiform ........................................................................................................................... 18. G. forrestii
20b. Stems glabrous or with indumentum, but not with retrorse hairs; leaves often broadest above
middle and thinner, without glandlike spots; flowers usually bisexual.
23a. Leaves linear, often longer than 20 mm, in addition to 1 principal, with 2 weaker lateral
veins; corolla 45 mm in diam.
24a. Leaves linear-spatulate, 14 mm wide; inflorescences loose, broadly paniculiform;
ovaries and fruit glabrous and smooth ........................................................................... 31. G. linearifolium
24b. Leaves linear-lanceolate, 39 mm wide; inflorescences dense, elongate-paniculate;
ovaries and fruit with sparse hooked trichomes or glabrous ....................................... 60. G. turkestanicum
23b. Leaves not linear, mostly shorter than 20 mm; corolla smaller.
25a. Leaves ovate, length/breadth index 2 or less, in addition to 1 principal, with 2 weaker
lateral veins; corolla larger; fruit with spreading hooked or straight hairs; Taiwan.
26a. Mericarps with straight trichomes; corolla 22.5 mm in diam.; stems pilose or
glabrescent .............................................................................................................. 36. G. nankotaizanum
26b. Mericarps with hooked trichomes; corolla ca. 3 mm in diam.; stems
glabrous ............................................................................................................ 53. G. takasagomontanum
25b. Leaves ovate-oblong to lanceolate, length/breadth index 2 or more, with
only 1 principal vein; corolla 1.52 mm in diam.; ovaries and fruit glabrous
or with various indumentum.
27a. Leaves dried subleathery; fruit with appressed, curved (but not uncinate)
hairs; Shanxi ................................................................................................................. 12. G. crassifolium
27b. Leaves dried papery; fruit with various indumentum.
28a. Plants ascending, weak, sparsely hairy or glabrous; inflorescence few flowered,
peduncles and pedicels very thin and elongated, latter mostly 48 mm; fruit
with spreading uncinate trichomes ............................................................................. 45. G. salwinense
28b. Plant erect, more robust, indumentum diverse; inflorescences many flowered,
peduncles and pedicels thicker and shorter, latter mostly 24 mm; fruit
tuberculate, with appressed or spreading hooked trichomes, or more rarely
smooth .................................................................................................................................. 9. G. bungei
18b. Leaves with 35 palmate principal veins, lateral veins well marked and extending for more
than half of blade length.
29a. Corolla funnelform or cup-shaped, 25 mm in diam., with fused lower part as long as
lobes; ovaries and fruit glabrous.
30a. Corolla cup-shaped or campanulate, 22.7 mm in diam.; cauline leaves broadly
lanceolate, always in whorls of 4 .......................................................................................... 30. G. kunmingense
30b. Corolla funnelform or campanulate, 2.55 mm in diam.; middle stem leaves ovate to
elliptic, in whorls of 46.
31a. Open corollas 3.55 mm in diam.; cauline leaves usually in whorls of 4,
1228 mm ............................................................................................................................ 40. G. platygalium

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109

31b. Open corollas 2.53.5 mm in diam.; cauline leaves in whorls of 46(8),

2353 mm ......................................................................................................................... 32. G. maximoviczii
29b. Corolla rotate, (1)25 mm in diam., with fused base much shorter than lobes.
32a. Leaves linear to linear-lanceolate, 2740 39 mm, in addition to 1 principal, with
2 weaker lateral veins; corolla 45 mm in diam.; ovaries and fruit with sparse
uncinate trichomes or glabrous ........................................................................................... 60. G. turkestanicum
32b. Leaves lanceolate to ovate, shorter, 3 principal veins mostly readily visible.
33a. Fruit glabrous, smooth to granular-papillose, or with appressed and hooked or with
spreading and straight (but never with spreading and hooked) trichomes.
34a. Open corollas 34 mm in diam.; stems (except nodes) mostly glabrous and smooth.
35a. Leaves ovate-lanceolate to ovate, papillose, length/breadth index mostly 2.5 or
less; cymes rather few flowered; ovaries and fruit with appressed, apically
somewhat bent trichomes .......................................................................................... 10. G. chekiangense
35b. Leaves mostly rather narrowly lanceolate, smooth or hairy, length/breadth
index mostly 3 or more; cymes many flowered; ovaries and fruit glabrous or
with various indumentum ...................................................................................................... 7. G. boreale
34b. Open corollas (1)22.5 mm in diam.; stems glabrous or hairy.
36a. Cauline leaves broadly to narrowly lanceolate, length/breadth index often
3.5 or more.
37a. Stems hairy throughout; leaves lanceolate; fruit with straight hairs or rarely
glabrous ........................................................................................................................ 24. G. hupehense
37b. Stems (except nodes) glabrous; leaves ovate-lanceolate (sometimes also broader),
apex subacute to acuminate, striate-punctate glandular below; fruit glabrous
and smooth .......................................................................................................................... 29. G. kinuta
36b. Cauline leaves narrowly to broadly ovate-lanceolate, length/breadth index usually
3 or less.
38a. Leaves 630 320 mm; fruiting pedicels straight; fruit glabrous or scaberulous;
mainland ........................................................................................................................... 15. G. elegans
38b. Leaves 410 25 mm; fruiting pedicels nodding; fruit with straight
trichomes; Taiwan ................................................................................................ 36. G. nankotaizanum
33b. Fruit with spreading and uncinate trichomes.
39a. Open corollas 3 mm or more in diam.; stems (except nodes) often glabrous and smooth.
40a. Leaves lanceolate to ovate-lanceolate or elliptic, length/breadth index 3.5 or more,
apex acute to acuminate ......................................................................................................... 7. G. boreale
40b. Leaves ovate, length/breadth index less than 3.
41a. Leaf apex obtuse to rounded, usually mucronate; mainland ............................... 26. G. kamtschaticum
41b. Leaf apex acuminate; Taiwan ....................................................................... 53. G. takasagomontanum
39b. Open corollas 2.5 mm or less in diam.; stems often hairy.
42a. Leaves ovate-lanceolate to narrowly elliptic, with acute apex, length/breadth
index normally more than 2.5 ....................................................................................... 63. G. yunnanense
42b. Leaves ovate to broadly elliptic, with obtuse to rounded apex, entire or shortly
mucronate, length/breadth index normally less than 2.5.
43a. Leaves up to 20 mm wide; plants slender to usually rather robust; mainland ................ 15. G. elegans
43b. Leaves up to 10 mm wide; slender low plants; Taiwan ............................................ 17. G. formosense
8b. Leaves and leaflike stipules in middle stem region regularly in whorls of more than 4, i.e., in
whorls of 516, with only 1 principal vein, linear to broadly lanceolate or elliptic, but never
ovate or with a length/breadth index of less than 2.5.
44a. Inflorescences capitate and enclosed by leaflike bracts; corolla funnelform or salverform,
415 mm, with 4 or 5 lobes.
45a. Plants perennial, 2070 cm tall; calyx limb obsolete; corolla 5-lobed, 1214 mm ...... Phuopsis stylosa (see p. 291)
45b. Plants annual; corolla 4-lobed, shorter.
46a. Calyx teeth well developed; corolla pink, tube 45 mm ....................... Sherardia arvensis (see comments above)
46b. Calyx lacking; corolla bluish, tube 713(15) mm .................................................. Asperula orientalis (see p. 78)
44b. Inflorescences branched, not enclosed by bracts; corolla rotate, campanulate,
or funnelform, 0.513 mm, mostly with 4 (rarely 3) lobes.
47a. Medium stem leaves marginally (and often on upper side) with microhairs directed forward
(use 20 lens), thus antrorsely ciliolate or aculeolate; stems mostly not retrorsely aculeolate.
48a. Ovaries and fruit densely covered with uncinate trichomes; plants perennial with smooth stems.

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49a. Corolla funnelform, with tube as long as lobes ......................................................................... 37. G. odoratum
49b. Corolla rotate, with tube markedly shorter than lobes.
50a. Robust, procumbent to ascending plants often longer than 50 cm; inflorescences terminal
and lateral with cymes in upper 24 nodes ................................................................................ 59. G. triflorum
50b. Slender erect plants, less than 50 cm tall; inflorescences predominantly terminal.
51a. Leaves 625 27 mm; open corollas ca. 2 mm in diam.; Taiwan (cf. also
53. G. takasagomontanum) .............................................................................................. 14. G. echinocarpum
51b. Leaves often larger; open corollas 1.53.5 mm in diam.; mainland.
52a. Leaves mostly in whorls of 7 or 8, (ob)lanceolate, length/breadth index mostly
3.54.5, subpetiolate; fruit with uncinate trichomes 0.60.8 mm; expected
in Xizang .......................................................................................................................... 4. G. asperuloides
52b. Leaves mostly in whorls of 6, narrowly obovate to broadly oblanceolate,
length/breadth index mostly 2.53.5, clearly petiolate; fruit with uncinate
trichomes 0.81.2 mm or longer; widespread ................................................................. 22. G. hoffmeisteri
48b. Ovaries and fruit glabrous or hairy, but never with uncinate trichomes; plants perennial
or annual.
53a. Plants annual, slender; flowers on pedicels and peduncles often longer than 5 mm and
in lax, diffuse inflorescences.
54a. Leaves filiform, 2030 mm, ascending; corolla funnelform, pink to red; ovary
and mericarps with dense, short and curved hairs ............................................. Leptunis trichodes (see p. 213)
54b. Leaves linear to oblanceolate, 420 mm, spreading to reflexed; corolla rotate,
whitish, yellowish, or greenish; ovary and fruit glabrous or slightly tuberculate.
55a. Inflorescences broadly ovate, diffuse and intricate, with fruiting pedicels elongated
to 20 mm ............................................................................................................................. 55. G. tenuissimum
55b. Inflorescences rather narrowly thyrsoid, not diffuse and intricate, with fruiting
pedicels only up to 4 mm ..................................................................................................... 19. G. ghilanicum
53b. Plants perennial, slender to robust; flowers on pedicels and peduncles 0.55 mm, in lax
to congested inflorescences.
56a. Corolla funnelform, lobed for 1/22/3, white.
57a. Inflorescences lax, ebracteate; stems erect, smooth; main stem leaves 1565
312 mm ............................................................................................................................. 38. G. paniculatum
57b. Inflorescences congested, strongly bracteate; stems procumbent, mostly hairy;
main stem leaves 523 12(5) mm ................................................................................. 23. G. humifusum
56b. Corolla rotate, lobed for 3/4 or more, often yellowish; plants erect to ascending.
58a. Leaves in middle stem region in whorls of not more than 6; plants of (sub)alpine region,
not taller than 30 cm ................................................................................................................. 46. G. saurense
58b. Leaves in middle stem region in whorls of more than 6 and up to 12.
59a. Open corollas 3.55 mm in diam., white; fruit somewhat spongy or fleshy, 33.5 mm,
with a dry pericarp separating from rest of fruit ..................................................................... 8. G. bullatum
59b. Open corollas ca. 3 mm in diam., yellow to whitish; fruit with dry mericarps,
1.52 mm, with pericarp dark and firmly attached to rest of fruit.
60a. Leaves mostly 2.55 mm wide, glabrous abaxially; fruit ca. 1.5 mm ................... 11. G. consanguineum
60b. Leaves 12.5 mm wide, glabrous to densely pubescent abaxially; fruit 1.52 mm ............. 62. G. verum
47b. Medium stem leaves marginally (not on leaf surface) with microhairs directed backward
(use 20 lens), thus retrorsely aculeolate or completely glabrous and smooth; stems often
retrorsely aculeolate.
61a. Annuals, often in disturbed, weedy habitats; stems and leaf margins retrorsely aculeolate;
fully developed mericarps subspherical, 26 mm; open corollas 12 mm in diam.
62a. Fruit becoming pendulous on arching peduncles and pedicels, verrucose to spinulose;
leaves glabrescent above ............................................................................................................ 57. G. tricornutum
62b. Fruit on divaricate straight peduncles and pedicels (only latter sometimes bent just
beneath fruit), mostly with uncinate trichomes; leaves hairy above.
63a. Open corollas 1.52 mm in diam.; individual mature mericarps 2.55 mm in diam., with
trichomes arising from tuberculate bases ....................................................................................... 2. G. aparine
63b. Open corollas 11.5 mm in diam.; individual mature mericarps 13 mm in diam., with
trichomes straight from base ......................................................................................................... 50. G. spurium
61b. Perennials, in natural habitats; fully developed mericarps ellipsoid, 1.53 mm; open corollas
14 mm in diam.

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64a. Middle stem leaves narrowly obovate to oblanceolate, mostly 1828 510 mm; stems
slightly retrorsely aculeolate; inflorescences of medium size, with terminal and lateral,
few- to several-flowered, rather loose cymes with small bracts, in fruit stiffly divaricate;
corolla rotate, 1.52 mm in diam.; fruit with hooked trichomes .............................................. 58. G. trifloriforme
64b. Middle stem leaves mostly smaller; inflorescences different, usually more bracteate; fruit
smooth, verrucose, or with hooked trichomes.
65a. Corolla funnelform to subcampanulate, with tube as long as or slightly shorter than
lobes, whitish; ovaries and fruit glabrous, smooth or verrucose; leaves papery to
subleathery and glossy; stems rough, retrorsely aculeolate, procumbent to clambering.
66a. Corolla funnelform; plants robust, 0.61.2 m, often forming mats; main stem leaves
850 28 mm .................................................................................................................. 28. G. karataviense
66b. Corolla subcampanulate; plants slender and erect, 1060 cm tall; main stem leaves
316 13 mm ..................................................................................................................... 61. G. uliginosum
65b. Corolla rotate, fused basal part much shorter than lobes; ovaries and fruit with hooked
trichomes, tuberculate, or glabrous; stems glabrous, rough, or hairy.
67a. Middle stem leaves larger, (5)1035(50) (1)2.510 mm, mostly hairy, at least
margins retrorsely aculeolate; cymes lateral and terminal, many flowered; ovaries
and fruit glabrous or with diverse indumentum; plants from lower elevations,
relatively robust, erect or clambering, stems up to 0.7 m tall.
68a. Stems branched from base; cymes leafy, with bracts to last branches; stems and
leaves (nodes and margins excepted) glabrous and smooth; mericarps with
spreading uncinate trichomes .......................................................................................... 48. G. sichuanense
68b. Stems normally branched from middle; cymes usually less leafy; stems and leaves
mostly with more indumentum; fruit glabrous, papillose, tuberculate, or with
uncinate trichomes.
69a. Peduncles and pedicels slender, filiform and often flexuose, with inconspicuous
bracts; pedicels up to 5 mm, in fruit elongated to 10 mm or more; flowers never
reddish; plants usually clambering; ovary and fruit surfaces diverse.
70a. Fruit often with uncinate trichomes; mainland .......................................................... 13. G. dahuricum
70b. Fruit glabrous; Taiwan ................................................................................................ 52. G. taiwanense
69b. Peduncles and pedicels rather stiff and often divaricate and bracteate; pedicels
shorter than 5 mm and hardly elongated in fruit; ovary and fruit surfaces diverse,
but often glabrous.
71a. Corolla red to purple (very rarely maroon or white), 1.52.5 mm in diam. ......................... 6. G. blinii
71b. Corolla whitish, yellowish, or greenish.
72a. Corolla small, 1.52 mm in diam., yellow to greenish white, lobes aristate;
inflorescences divaricate and regularly bracteate with bracts similar to but
smaller than leaves, giving a diffuse miniature aspect; ovary and fruit surface
variable; plants often clambering ........................................................................... 3. G. asperifolium
72b. Corolla larger, mostly more than 2 mm in diam.; inflorescences ebracteate or
with bracts reduced and irregularly distributed; ovary and fruit surface
smooth to tuberculate; plants erect, hardly clambering.
73a. Leaves lanceolate, gradually narrowed into acute apex .............................................. 41. G. prattii
73b. Leaves subspatulate to obovate, apex rounded and abruptly narrowed into
a mucro ................................................................................................................... 56. G. tokyoense
67b. Middle stem leaves uniformly small, 212(15) 0.33.5 mm, glabrous and smooth
to hairy; cymes lateral and terminal, few flowered; ovaries and fruit with uncinate
(very rarely straight) trichomes or glabrous; plants from high elevations, usually
reduced and weak, caespitose to procumbent, stems only up to 0.3 m.
74a. Ovaries and fruit with uncinate (very rarely straight) trichomes.
75a. Leaves and stems densely hispid and often retrorsely aculeolate; stems with
4 conspicuous whitish angles .................................................................................. 42. G. pusillosetosum
75b. Leaves completely glabrous and smooth or only slightly hairy and/or retrorsely
aculeolate; stems with 4 inconspicuous angles.
76a. Leaves dried blackening, papery and thin, oblanceolate to narrowly obovate,
with flat margins, hardly longer than 7 mm ............................................................ 5. G. baldensiforme
76b. Leaves dried greenish-brownish, with revolute margins, often longer
than 7 mm.
77a. Plants nearly always smooth; leaves subleathery; ovary in flower
ca. 1 mm ............................................................................................................ 49. G. glabriusculum

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77b. Plants retrorsely aculeolate at least on margins and lower side of papery leaves;
ovary in flower 0.50.8 mm ................................................................................. 51. G. sungpanense
74b. Ovaries and fruit glabrous, smooth, papillose, or verrucose; Himalaya.
78a. Plants weak to procumbent but not mat-forming; cells of adaxial leaf surface
relatively large, readily visible individually with 20 lens; corolla mostly
whitish ...................................................................................................................... 33. G. megacyttarion
78b. Plants procumbent and often mat-forming; cells of adaxial leaf surface small,
not or hardly visible individually with 20 lens.
79a. Stems densely hairy and/or retrorsely aculeolate, with 4 conspicuous whitish
angles .................................................................................................................... 42. G. pusillosetosum
79b. Stems glabrous or only slightly retrorsely aculeolate, with inconspicuous angles.
80a. Leaves on main stems 28.5 mm; inflorescence cymes 1- to few flowered,
fascicled; corolla white, pale green, or pale yellow, with upper surface of
lobes papillose ................................................................................................................. 1. G. acutum
80b. Leaves on main stems 510.5 mm; inflorescence cymes 16-flowered; corolla
nearly always red or purple, with upper surface of lobes glabrous and smooth
except puberulent on margins and central vein ............................................................ 43. G. rebae
1. Galium acutum Edgeworth, Trans. Linn. Soc. London 20:
61. 1846.
jian ban la la teng
Herbs, perennial, procumbent, much branched, matforming. Stems up to 30 cm, 4(or 6)-angled, glabrous, smooth
or sometimes with scattered (very rarely more dense) short and
straight hairs. Leaves in whorls of up to 6, sessile; blade drying
papery and blackish, linear-oblanceolate to narrowly ellipticoblanceolate, 28.5 0.31.5 mm, glabrous and smooth, occasionally with straight hairs, base cuneate, margins flat to thinly
revolute, very rarely antrorsely aculeolate, apex acute, contracted and mucronate; vein 1. Inflorescences with terminal and
axillary cymes, 1- to few flowered; peduncles (1.5)38(10)
mm; pedicels (0.1)0.52(3) mm, glabrous, smooth. Ovary
ellipsoid-obovoid, ca. 0.5 mm, didymous, glabrous. Corolla
white, pale greenish, or yellowish, rotate, 1.23.5 mm in diam.,
glabrous to puberulent, lobed for 2/3 or more; lobes 4, lanceolate-spatulate, inside (i.e., adaxially) papillose, shortly acuminate. Mericarps ellipsoid, ca. 1 0.40.6 mm, glabrous, smooth
or granular-verruculose, often on elongating pedicels. Fl. and fr.
JulOct.
Mountain rocks and slopes; 20004100 m. ?Sichuan, Xizang,
?Yunnan [India, Nepal, Pakistan].
Galium acutum is a (sub)alpine Himalayan member of the G.
asperifolium group (see under that species). This group of taxa can be
divided into subgroups: (1) from lower elevations and (2) from higher
elevations. The latter subgroup is represented in the W Himalaya of
Pakistan by G. acutum alone (Nazimuddin & Ehrendorfer, Pl. Syst.
Evol. 155: 7175. 1987). Mill (Edinburgh J. Bot. 53: 193213. 1996; Fl.
Bhutan 2(2): 825834. 1999) has analyzed both subgroups in detail with
emphasis on their E Himalayan members. Among subgroup (2) he
recognized three species: G. acutum and the newly described G. rebae
and G. megacyttarion. The only material from China incorporated in
Mills study are specimens of G. rebae from Xizang deposited at E and
BM. Additional species from subgroup (2) from the C and E Himalaya
and adjacent China treated here are G. baldensiforme, G. pusillosetosum,
G. glabriusculum, and G. sungpanense. They appear well separable from
G. acutum.
Galium acutum and G. rebae are very closely related taxa. Mill
(loc. cit. 1996: 199) presented a differential table which has been incor-

porated into the present descriptions. Nevertheless, from the material

available now, it appears that only flower color (white or greenish white
in G. acutum and reddish crimson in G. rebae) is really decisive for their
separation. Furthermore, at lower elevations, G. acutum appears linked
to G. asperifolium var. sikkimense. Cufodontis (Oesterr. Bot. Z. 89: 239.
1940) has described such transitional forms with longer and retrorsely
aculeolate stems and hairy corolla lobes from the Indian Himalaya as G.
acutum var. trichanthum Cufodontis.
Mill (loc. cit. 1996: 194198) considered Galium acutum to be
restricted to the NW Himalaya except for one provisionally identified
specimen from Sikkim. In contrast, specimens studied by us from the
herbaria PE, KUN, and WU clearly show that G. acutum extends much
further to the east, reaching Yunnan and Sichuan.
Galium himalayense was regarded as a synonym of G. acutum by
Cufodontis (loc. cit.: 239243). Mill (loc. cit. 1996: 195; loc. cit. 1999:
831832) agreed but maintained the taxon as G. acutum var. himalayense and described its sympatric occurrence with G. acutum var. acutum
throughout the NW Himalaya. As Mill did not consider the possible
occurrence of flower dimorphism in G. acutum, it remains uncertain
whether the flower and stigma size differences listed are possibly correlated with male and female plants or simply correspond to hermaphroditic variants within the morphological amplitude of the species. In
order to stimulate such studies and to clarify the distribution of the two
taxa in China, a key and descriptions (according to Mill, loc. cit. 1996)
follow:

1a. Open corollas 2.33.5 mm in diam.;

stigmas united to near middle, in total
length subequal to stamens ......................... 1a. var. acutum
1b. Open corollas 1.22.3 mm in diam.;
stigmas united only shortly at base, in
total length shorter than stamens ....... 1b. var. himalayense
1a. Galium acutum var. acutum
() jian ban la la teng (yuan bian zhong)
Galium asperifolium Wallich var. setosum Cufodontis.
Leaves of main stems with blades mucronate at apex,
mucro 0.20.45 mm. Corolla 2.33.5 mm in diam.; lobes 1.94
as long as wide, mucronate with mucro 0.150.3 mm. Stigmas united to near middle, in total length subequal to stamens.
Fl. and fr. JulOct.

RUBIACEAE

Mountain rocks and slopes; 20004100 m. ?Sichuan, Xizang,

?Yunnan [India, Nepal, Pakistan].

1b. Galium acutum var. himalayense (Klotzsch & Garcke) R.

R. Mill, Edinburgh J. Bot. 53: 195. 1996.
xi ma la ya jian ban la la teng
Galium himalayense Klotzsch & Garcke, Bot. Ergebn.
Reise Waldemar, 88. 1862.
Leaves of main stems with blade submucronate at apex,
mucro 0.150.2 mm. Corolla 1.22.3 mm in diam.; lobes 1.4
2.75 as long as wide, acute or submucronate with mucro to
0.1 mm. Stigmas united only shortly at base, in total length
shorter than stamens. Fl. and fr. JulOct.
Mountain rocks and slopes; 20004100 m. ?Sichuan, Xizang,
?Yunnan [India, Nepal].

2. Galium aparine Linnaeus, Sp. Pl. 1: 108. 1753, s.s.

yuan la la teng
Herbs, annual, procumbent or clambering. Stems 3090
cm high, 4-angled, 14 mm in diam., branched from base,
retrorsely aculeate along angles, glabrescent to pilose at nodes.
Leaves at middle stem region in whorls of 610, subsessile;
blade drying papery, narrowly oblanceolate to narrowly oblongoblanceolate, 1060 310 mm, usually somewhat pilosulous
or hispidulous adaxially, retrorsely aculeolate along midrib
abaxially, base acute, margins flat to thinly revolute, retrorsely
aculeolate, apex acute and shortly mucronate; vein 1. Inflorescences terminal and axillary, cymes 2- to several flowered; axes
glabrous to aculeolate; bracts leaflike or none, 15 mm;
peduncles 15 cm; pedicels 130 mm, finally elongating and
sometimes curved directly under fruit. Ovary subglobose, 0.3
0.5 mm, with uncinate trichomes. Flowers hermaphroditic. Corolla yellowish green or white, rotate, 1.52 mm in diam.; lobes
4, triangular to ovate, acute. Mericarps subglobose to kidneyshaped, 2.55 mm, with a dense cover of uncinate trichomes
0.41.2 mm from swollen base. Fl. MarJul, fr. AprNov.
Forest margins, riversides, meadows, open fields, farmlands; near
sea level to 2500 m. Evidently rare in China and possibly only introduced [originally in W Eurasia and the Mediterranean, but today nearly
worldwide as an adventive].
The Galium aparine group (G. sect. Aparine, formally part of G.
sect. Kolgyda s.l.) forms an annual, extremely polymorphic, and predominantly autogamous polyploid complex, also called G. aparine s.l.
or G. aparine agg. One has to consider as possible perennial ancestors
the morphologically very close E Asiatic taxa (e.g., G. sungpanense: see
there) and other annuals, such as the Aegean endemic G. monachinii
Boissier & Heldreich (2x, 2n = 22) and the Eurasian and African G.
spurium (2x and 4x, 2n = 20, 40). By allopolyploidy they apparently
have contributed in the Mediterranean and W Eurasia to G. aparine s.s.
(4x, 6x, and 8x with euploid and slightly oscillating aneuploid
chromosome numbers), which today has become a nearly worldwide
weed (Ehrendorfer et al., Fl. Iranica 176: 239. 2005).
Many authorities, including Cufodontis (Oesterr. Bot. Z. 89: 245
247. 1940) and W. C. Chen (in FRPS 71(2): 234237. 1999), have
treated all these plants under Galium aparine s.l. and recognized four
varieties: var. aparine, var. echinospermum, var. leiospermum, and var.
tenerum. Whereas the first refers to G. aparine s.s. described above, the

113

latter three should be assigned to G. spurium (see there). Here, we

follow the narrow circumscription of G. aparine s.s. and the specific
separation of G. spurium outlined above, in spite of occasional difficulties in separating the two taxa on the basis of flower and mericarp
size. A relevant survey of extensive Chinese material at the herbaria PE
and KUN has clearly shown the common and wide occurrence of G.
spurium in comparison with the rare and only occasional documentation
of G. aparine s.s. Only future karyosystematic studies on the group in E
Asia will clarify their distribution and ecological position. With respect
to the common confusion of members of the G. aparine group with
other annual and perennial taxa of Galium see G. spurium.

3. Galium asperifolium Wallich in Roxburgh, Fl. Ind. 1: 381.

1820.
xie ye l
?Galium cavaleriei H. Lveill; ?G. esquirolii H. Lveill.
Herbs, perennial, weak to climbing or trailing, usually
much branched. Stems 2070 cm, 4-angled to 4-winged, villosulous to hirtellous and/or sparsely aculeolate to smooth.
Leaves on main stems in whorls of up to 6(8), sessile or with
very short (ca. 1 mm) petiole; blade drying papery to leathery,
adaxially dark green and shiny, abaxially paler, oblanceolateoblong, oblanceolate, or obovate, (5)1020(25) (1)1.5
4(6) mm, adaxially scaberulous, hirtellous to glabrous, abaxially densely villosulous, hirsute, pilose to glabrous, base acute
to cuneate, margins retrorsely aculeolate and hairy, flat to
thinly revolute, apex obtuse, rounded, truncate, or emarginate
and shortly mucronate; vein 1. Inflorescences paniculate, up
to 18 cm, expanding through growing season, with terminal and
axillary, several- to many-flowered cymes; peduncles glabrous
to rarely villosulous, regularly spreading to divaricate, with a
dichasial branching pattern, at most nodes with leaflike bracts
(14 mm); pedicels 0.22.5 mm. Ovary obovoid, 0.20.3 mm,
mostly glabrous or smooth, but sometimes also verrucose, hirtellous, or with undeveloped uncinate trichomes. Corolla
greenish white or yellow, rotate, 1.52 mm in diam., glabrous,
lobed for 2/3 or more; lobes 4, triangular-ovate, filamentousaristate (rarely only acute). Mericarps ellipsoid, 12 mm, glabrous and smooth or rarely granular-tuberculate, hirtellous, or
with appressed to spreading hooked trichomes, on pedicels
often slightly elongating to 4 mm. Fl. and fr. (May)Jun
Sep(Oct).
Mountain slopes, farmland sides, riversides and beaches, grasslands, forests, thickets, ditch sides, open fields, meadows; 4003500 m.
Guangxi, Guizhou, Hubei, Hunan, Sichuan, Xizang, Yunnan [Afghanistan, Bangladesh, Bhutan, India, Nepal, Pakistan, Sri Lanka, Thailand].
Galium asperifolium is an exceedingly variable and widespread
species and was the first described from a larger assembly of taxa, here
called G. asperifolium group and provisionally placed into G. sect. Trachygalium (but certainly not into G. sect. Leptogalium as in Yamazaki,
Fl. Japan 3a: 238239. 1993, or into G. sect. Leiogalium as in W. C.
Chen, FRPS 71(2): 271. 1999). According to Cufodontis (Oesterr. Bot.
Z. 211251. 1940), Nazimuddin and Ehrendorfer (Pl. Syst. Evol. 155:
7175. 1987), Mill (Edinburgh J. Bot. 53: 193213. 1996; Fl. Bhutan
2(2): 825834. 1999), Ehrendorfer et al. (Fl. Iranica 176: 194. 2005),
and the present treatment, the center of diversity of the G. asperifolium
group lies in the E Himalaya and SW China. Within this area, two subgroups of taxa can be recognized, one with larger plants, longer than 10
mm middle stem leaves, and many-flowered cymes from lower eleva-

114

RUBIACEAE

tions (1), the other with more condensed growth, shorter middle stem
leaves, and 1- to few-flowered cymes from higher elevations (2). Both
subgroups include taxa with whitish, yellowish to greenish, and others
with reddish, purplish, or brownish flower color. The (sub)alpine subgroup (2) is briefly surveyed under G. acutum. Subgroup (1), discussed
here, is represented by taxa with whitish flowers: G. subfalcatum Nazimuddin & Ehrendorfer and G. campylotrichum Nazimuddin & Ehrendorfer in the W Himalaya of Pakistan and the widespread G. asperifolium with its var. sikkimense (= G. sikkimense) in the C and E Himalaya (including Bhutan). In addition, subgroup (1) includes taxa with a
more easterly distribution, extending from China into the remaining
parts of E Asia (including Japan): G. dahuricum (including G. comarii,
G. manshuricum, G. niewerthii, and G. pseudoasprellum), G. prattii, G.
taiwanense, and G. tokyoense. Subgroup (1) taxa with reddish flowers
are G. blinii in SW China and the newly described Bhutan endemic G.
craticulatum R. R. Mill (see also Mill, loc. cit. 1996; loc. cit. 1999).
Because of its great variability with respect to habit and the indumentum of stems, leaves, and fruit, Galium asperifolium is often not
easily separable from its closest relatives, and transitional forms occur.
Its best differential characters are the many-flowered, divaricate, distally dichasial branching, and strongly bracteate cymes and the small
yellowish-greenish flowers with aristate corolla lobes. Galium blinii
mainly deviates by larger, reddish flowers and non-aristate corolla
lobes. The filiform and flexuose peduncles and pedicels separate G.
dahuricum, and the less bracteate inflorescences and larger flowers
separate G. prattii, G. taiwanense, and G. tokyoense. To the taxa of the
(sub)alpine subgroup (2, e.g., G. acutum) G. asperifolium is linked
particularly through its var. sikkimense.
In addition to its natural complexity, the taxonomy of the Galium
asperifolium group is rendered difficult by a number of badly described
and insufficiently documented species created by H. Lveill. On the
basis of the studies by Cufodontis (loc. cit.), Lauener and Ferguson
(Notes Roy. Bot. Gard. Edinburgh 32: 103115. 1973), Mill (loc. cit.
1996), and our own judgment, we suggest to dispose of them in the
following way: Galium blinii is maintained as a separate species (possibly with G. bodinieri and G. quinatum as synonyms), whereas G.
cavaleriei and G. esquirolii (and G. cuneatum H. Lveill, though a
nomen nudum) are provisionally assigned as synonyms to G. asperifolium s.l. (including var. sikkimense); G. comarii and G. niewerthii are
treated as synonyms of G. dahuricum.
The following schematic key to the varieties of Galium asperifolium corresponds to W. C. Chen (loc. cit.: 271274) who mainly followed Cufodontis (loc. cit.: 239240). Only G. asperifolium var. setosum has been eliminated because it clearly is a synonym of G. acutum.
Individuals with uncinate trichomes on ovaries and fruit, but otherwise
identical to typical G. asperifolium, are reported here for the first time.
They still lack a varietal name and are provisionally accommodated
under var. asperifolium. In contrast to Mill (loc. cit. 1996; loc. cit.
1999), G. sikkimense is here again reduced to varietal status under G.
asperifolium, following Cufodontis (loc. cit.: 241). Forms of this species
with glabrescent stems dominate to the east of its wide distribution
area, but intraspecific and local variation of stem indumentum is so extensive and continuous as to make this character useless as a basis for
specific separation. Mill (loc. cit. 1996: 201212) assumed G. asperifolium var. asperifolium to be most common in C and W Nepal and to
be replaced by var. sikkimense toward the east. This statement is in conflict with the data on distribution in China from Chen (loc. cit.) and our
own observations presented below. In view of the taxonomical complexity of the G. asperifolium group and the common misinterpretation
of its members, further careful studies are obligatory.

1a. Plants stout, often clambering; stems

usually villosulous to hirtellous and densely

retrorsely aculeolate; leaf blade relatively

large, often oblanceolate-oblong, hairy
and marginally retrorsely aculeolate;
inflorescence large, many flowered;
corolla lobes aristate.
2a. Ovary and fruit glabrous, smooth,
granular-tuberculate, or with
uncinate trichomes .................... 3a. var. asperifolium
2b. Ovary and fruit hirtellous .......... 3b. var. lasiocarpum
1b. Plants often smaller; stems with
reduced indumentum; leaf blade
smaller, often more lanceolate, more
weakly retrorsely aculeolate to smooth
along margins; inflorescence with fewer
flowers; corolla lobes apiculate to acute.
3a. Mericarps smooth ......................... 3c. var. sikkimense
3b. Mericarps granular-tuberculate
................................................ 3d. var. verrucifructum
3a. Galium asperifolium var. asperifolium
() xie ye l (yuan bian zhong)
Galium pseudohirtiflorum H. Li.
Plants stout, often clambering. Stems 2070 cm, much
branched, villosulous or hirtellous, retrorsely aculeolate. Leaf
blade often larger, oblanceolate-oblong, mostly hirsute and
with margins retrorsely aculeolate. Inflorescences large, with
many-flowered cymes. Corolla lobes filamentous-aristate.
Ovary and fruit glabrous and smooth, verrucose, or rarely with
uncinate trichomes. Fl. and fr. JunSep.
Mountain slopes; 12003000 m. Guizhou, Sichuan, Xizang, Yunnan [Afghanistan, Bangladesh, India, Nepal, Pakistan, Sri Lanka, Thailand].

3b. Galium asperifolium var. lasiocarpum W. C. Chen, Acta

Phytotax. Sin. 28: 303. 1990.
mao guo xie ye l
Ovary and fruit hirtellous. Fl. and fr. MayOct.
Mountain slopes, farmland sides, riversides, forests; 14003200
m. Guangxi, Guizhou, Yunnan.
Mill (loc. cit. 1996) commented on the extensive indument variation on all organs of Galium asperifolium throughout most of its range
but did not mention any occurrence of straight hairs on fruit in this
species group, nor has anyone else besides W. C. Chen. Therefore, this
variety is only provisionally included here. Its densely hirsute ovary and
fruit suggests that it may belong to another species, possibly G. pusillosetosum.

3c. Galium asperifolium var. sikkimense (Gandoger) Cufodontis, Oesterr. Bot. Z. 89: 241. 1940.
xiao ye l
Galium sikkimense Gandoger, Bull. Soc. Bot. France 66:
307. 1920.
Herbs, perennial, weak to clambering or trailing. Stems
2060 cm, usually much branched, sparsely hairy and retrorsely
aculeolate to smooth. Leaves often smaller and narrower, less

RUBIACEAE

hairy and retrorsely aculeate to glabrous and smooth. Inflorescences large to medium sized with many- to several-flowered cymes. Corolla lobes apiculate to acute. Ovary and fruit
glabrous and smooth. Fl. JunSep, fr. JulOct.
Mountain slopes, river beaches, ditch sides, open fields, grasslands, meadows, thickets, forests; 4003200 m. Guangxi, Guizhou, Hubei, Hunan, Sichuan, Xizang, Yunnan [Bhutan, India, Nepal].

3d. Galium asperifolium var. verrucifructum Cufodontis,

Oesterr. Bot. Z. 89: 241. 1940.
dian xiao ye l
Similar to Galium asperifolium var. sikkimense but with
granular-tuberculate fruit. Fl. and fr. AugOct.
Mountain slopes, grasslands, thickets; 23003500 m. Sichuan,
Xizang, NW Yunnan.
The type material of this variety from Yunnan (particularly Handel-Mazzetti 9697, WU) consists of transitional forms toward Galium
acutum.

4. Galium asperuloides Edgeworth, Trans. Linn. Soc. London

20: 61. 1846.
che ye l
Herbs, perennial, emerging from filiform reddish rhizomes. Stems weak but generally erect, 1045 cm tall, 4-angled, glabrous and smooth, except hispidulous at nodes. Middle
stem leaves and leaflike stipules in whorls of (6 or)7 or 8, subsessile to very shortly petiolate; blade drying papery or membranous, remaining green, elliptic to narrowly oblong-oblanceolate or lanceolate, (10)2050(60) 313 mm, length/breadth
index mostly 3.54.5, glabrescent, base acute or cuneate, midrib
smooth or rarely retrorsely aculeolate, margins and upper leaf
side with antrorse microhairs, apex obtuse or rounded and
abruptly apiculate; vein 1. Inflorescences terminal and sometimes in axils of upper leaves with few- to several-flowered
cymes; axes glabrous, smooth; bracts none or few, 12 mm;
pedicels 0.55 mm. Ovary ovoid, 0.50.8 mm, with uncinate
trichomes. Corolla white or light greenish, rotate, 2.53.8 mm
in diam., lobed for 3/4 or more; lobes 4, ovate, acute. Mericarps
ellipsoid, 1.82.5 mm, with dense uncinate trichomes 0.60.8
mm, on fruiting pedicels elongating to 10 mm. Fl. AprAug, fr.
MaySep.
Forests on mountain slopes, thickets, ditch sides, along rivers,
meadows; 15002800 m. Expected in Xizang [Afghanistan, India,
Kashmir, Pakistan].
Galium asperuloides was previously circumscribed more broadly
to include as subspecies plants treated here as G. hoffmeisteri. The specific status of the latter is well justified (Ehrendorfer et al., Fl. Iranica 176:
193194. 2005; see comments and differential characters under that
species). When the two taxa are classified as one species, the typical
plants have to be called G. asperuloides subsp. asperuloides. Vegetative
plants are very similar to G. odoratum. Together with G. echinocarpum
from Taiwan and others they constitute G. sect. Hylaea.

5. Galium baldensiforme Handel-Mazzetti, Symb. Sin. 7:

1029. 1936.
yu long la la teng

115

Herbs, perennial, tender, caespitose. Stems ascending, 3

12(25) cm tall, 4-angled, glabrous and smooth or rarely
slightly retrorsely aculeolate. Leaves in whorls of up to 5 or 6,
sessile; blade drying papery and blackish, oblanceolate to narrowly obovate, 27 13 mm, mostly glabrous and smooth,
but occasionally with straight hairs abaxially or marginally
slightly retrorsely aculeolate, base cuneate, margin flat, apex
acute and often shortly mucronate; vein 1. Inflorescences with
cymes terminal and in axils of upper leaves, 1- or usually 2- or
3-flowered; peduncles up to 18 mm and pedicels 25 mm, glabrous and smooth, elongating during fruit development. Ovary
subglobose, ca. 0.7 mm, densely covered by undeveloped trichomes. Corolla pale green, rotate, ca. 2 mm in diam., glabrous;
lobes 4, triangular, acute. Mericarps ellipsoid, ca. 2 mm, with
dense brownish yellow uncinate trichomes ca. 0.7 mm. Fl. Aug,
fr. AugOct.
Rocky slopes, meadows, frost heave sites, river floodplains in
mountains; 28004300 m. Qinghai, Sichuan, Xizang, Yunnan (Lijiang).
Galium baldensiforme belongs to the alpine subgroup (2) of the G.
asperifolium complex within G. sect. Trachygalium s.l. (see under G.
acutum and G. asperifolium). It has been widely misidentified with related taxa, e.g., with G. glabriusculum from which it differs in its more
oblanceolate, thinner leaves, blackening when dried, and a slight tendency toward more indumentum.
Two collections from Xizang (H. Li 1978-07-22 and Y. T. Chang
& Lang, Nie-La-Mu, 1966-06-25, both from PE) differ from typical
Galium baldensiforme by purple flowers (reminiscent of G. rebae),
straight whitish hairs on its (still young) ovaries, and scattered straight
hairs on the upper and lower leaf sides (mainly midvein, but glabrous
along leaf margins). After closer inspection and field studies, these populations may very well deserve species rank.
A specimen from Sichuan (Dege Co., anonymous collector 7029,
PE) with stronger indumentum, subleathery, broadly lanceolate leaves
with retrorsely aculeolate margins, and aristate corolla lobes apparently
links Galium baldensiforme with G. asperifolium var. sikkimense.

6. Galium blinii H. Lveill, Bull. Acad. Int. Gogr. Bot. 25:

48. 1915.
wu ye la la teng
?Galium bodinieri H. Lveill; ?G. quinatum H. Lveill
& Vaniot.
Herbs, perennial, weak to climbing, trailing, or matted.
Stems usually much branched, 2070 cm, 4-angled, retrorsely
aculeolate to glabrescent. Middle stem leaves in whorls of 68,
subsessile; blade drying papery or leathery, often blackening,
linear-oblong to broadly (ob)lanceolate, (5)1022(30)
(1)24.5(5.5) mm, adaxially and particularly abaxially
rough, base acute to cuneate, margin flat to thinly revolute,
densely retrorsely aculeolate, gradually narrowed into acute
apex; vein 1. Inflorescences with terminal and axillary, severalflowered cymes 25 cm; axes glabrous, often slightly divaricate, with small bracts on lower branches only; pedicels
(0.2)13(5) mm. Ovary obovoid, 0.20.3 mm, glabrous or
with undeveloped trichomes. Corolla red to purple or violet
(only very rarely white), rotate, 1.52.5(3) mm in diam., glabrous; lobes 4, triangular-ovate, acute. Mericarps ovoid, 12
mm, glabrous, smooth to verrucose, or sometimes with appressed or spreading uncinate trichomes. Fl. JunSep, fr. Jul
Oct.

116

RUBIACEAE

Mountain slopes, river beaches, ditch sides, open fields, grasslands, meadows, thickets, forests; 8003000 m. Guizhou, Hubei, Shaanxi, Sichuan, Xizang, Yunnan.

with appressed, ascending, or spreading, straight or curved,

but hardly truly uncinate trichomes 0.30.5 mm. Fl. May
Aug(Sep), fr. (May)JunOct.

Galium blinii is a critical taxon that belongs to the lower elevation

subgroup (1) of the G. asperifolium group (see there). It was considered
a synonym of G. asperifolium var. sikkimense by Cufodontis (Oesterr.
Bot. Z. 89: 241. 1940), Lauener and Ferguson (Notes Roy. Bot. Gard.
Edinburgh 32: 106. 1973), and W. C. Chen (in FRPS 71(2): 273. 1999).
Only Mill (Edinburgh J. Bot. 53: 204. 1996) commented on its reddish
purple flowers and other differential characters, regarded it as a distinct
species, and proposed to use its old but so far neglected name. Up to
now, specimens of this rather widespread taxon were named as G.
asperifolium var. sikkimense, G. pseudoasprellum var. densiflorum, etc.
Analyses of a considerable number of relevant specimens from PE,
KUN, and WU support Mills interpretation and led to the above, more
elaborate description. It shows that G. blinii is quite variable with respect to leaf shape and ovary/fruit indumentum but relatively well characterized not only by its reddish purple flowers but also by height, leaf
size, retrorsely aculeolate stems and leaf margins, and the usually medium-sized and only small-bracteate cymes, relatively short and firm postfloral peduncles and pedicels, and larger flowers with acute (but not
aristate) lobes. This allows separation from its closest relatives, G. prattii
and G. asperifolium (where transitional forms occur), but also from G.
dahuricum and G. tokyoense.

Open forests and thickets, mountain slopes, grasslands, meadows,

open fields, ditch sides, river valleys and beaches, swamps, farmland
sides, wastelands; 2004600 m. Gansu, Hebei, Heilongjiang, Henan,
Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong,
Shanxi, Sichuan, Xinjiang, Xizang, Yunnan [Afghanistan, India, Japan,
Kashmir, Korea, Mongolia, Pakistan, Russia; SW Asia (Armenia, Iran),
Europe, North America].

Further studies will have to show to what extent the following,

also reddish purplish flowering taxa from SW China can be separated
from Galium blinii: G. craticulatum was described as an endemic from
the high mountains of Bhutan (Mill, loc. cit.: 202) and said to differ
from G. blinii in its present circumscription by completely glabrous
stems with conspicuous, vein-marked wings, less retrorsely aculeolate
leaves, larger flowers, and longer filaments; G. bodinieri, also with reddish flowers, according to Mill (loc. cit.: 204205) is reminiscent of G.
craticulatum (particularly in its broadly winged stems) and may
represent a link between G. blinii and the alpine, more condensed and
shorter leaved G. rebae; the purple-flowered G. quinatum (not mentioned by Mill) was very poorly described and is listed above as a possible synonym of G. blinii, but types have neither been seen by Lauener
and Ferguson (loc. cit.: 107) nor by us.

7. Galium boreale Linnaeus, Sp. Pl. 1: 108. 1753, s.l.

bei fang la la teng
Herbs, perennial, erect, 2065 cm tall. Stems 4-angled,
usuually glabrous, rarely shortly hairy, hispidulous at nodes,
angles thickened. Leaves in whorls of 4, sessile or subsessile;
blade drying papery or thinly leathery, linear-lanceolate or lanceolate to ovate, (10)1540(80) (1)315 mm, glabrous or
sparsely puberulent to hispidulous and/or pilose, abaxially never
with striate to punctate glandular idioblasts, base cuneate to subrounded, margins usually revolute and antrorsely scaberulous to
hispidulous, apex acute or usually narrowly tapered then obtuse
to rounded at very tip; principal veins palmate, 3. Inflorescences terminal, elongate or broadly paniculiform, 215 cm,
with several- to many-flowered cymes in axils of uppermost
leaves and terminal; peduncles glabrous or puberulent at nodes,
smooth or scaberulous; bracts ligulate, lanceolate, or elliptic, 1
4 mm; pedicels 0.52 mm elongating in fruit to 3.5 mm. Ovary
subglobose, 0.81 mm, glabrous or sparsely to densely strigillose to pilosulous. Corolla white or pale yellow, rotate, 34 mm
in diam., glabrescent, lobed for 3/4 or more; lobes 4, ovate-lanceolate, acute. Mericarps subglobose, 12 mm, pericarp firmly
attached but sometimes inflated, glabrous or densely hairy

The name Galium boreale, as used here in a wide sense, corresponds to a widespread and polymorphic, still insufficiently studied
N Hemisphere polyploid complex (Ehrendorfer et al., Fl. Iranica 176:
179181. 2005) within G. sect. Platygalium s.l. In China, another
species of this section with much smaller flowers, G. kinuta, can be
separated from this G. boreale aggregate only with difficulties, because
the two are linked by intermediate (and possibly hybrid) populations
(see under G. kinuta).
Within the Galium boreale aggregate and the flora of China, W. C.
Chen (in FRPS 71(2): 260263, 285. 1999) recognized only G. boreale
Linnaeus s.l. with numerous infraspecific taxa and G. turkestanicum,
whereas 11 species in three series were listed for the flora of the former
Soviet Union by Pobedimova et al. (Fl. URSS 23: 345354. 1958).
From these only G. turkestanicum is fully accepted here (G. ussuriense
and G. rubioides are cited as synonyms under G. boreale var. lanceolatum and G. boreale var. rubioides). Furthermore, and according to
Pobedimova et al. (loc. cit.), G. amblyophyllum Schrenk, G. amurense
Pobedimova, and G. septentrionale Roemer & Schultes can be expected
to occur in China. With the exception of the briefly mentioned G. septentrionale, they were not considered by W. C. Chen in FRPS and are
only mentioned here. As a competent treatment of the G. boreale aggregate is not yet possible, we follow the schematic taxonomic differentiation proposed by Cufodontis (Oesterr. Bot. Z. 89: 225228. 1940) and
accepted by W. C. Chen (loc. cit.). This scheme defines numerous
varieties according to leaf shape and the density, type, and distribution
of indumentum on leaves, ovaries, and fruit. These varieties form a
morphologically continuous series, linking the extremes: G. boreale
var. rubioides with large ovate leaves and a broadly paniculate inflorescence and G. boreale var. intermedium with much smaller lanceolate
leaves and an elongated narrow inflorescence. The following key and
short descriptions are presented here for reference, to facilitate comparison, and to stimulate future studies.

1a. Ovary and fruit glabrous.

2a. Leaf blade pilose or scabrous
abaxially at least along veins.
3a. Leaf blade linear-lanceolate
or narrowly lanceolate ...... 7h. var. lancilimbum
3b. Leaf blade ovate-lanceolate
or ovate ................................... 7k. var. rubioides
2b. Leaf blade glabrous abaxially.
4a. Leaf blade linear-lanceolate
or narrowly lanceolate .... 7d. var. hyssopifolium
4b. Leaf blade broadly
lanceolate or
ovate-lanceolate ................. 7g. var. lanceolatum
1b. Ovary and fruit hairy.
5a. Ovary and fruit sparsely hirtellous
or scabrous.
6a. Leaf blade linear-lanceolate
or narrowly lanceolate ...... 7e. var. intermedium
6b. Leaf blade broadly lanceolate
or ovate-lanceolate ....... 7j. var. pseudorubioides

RUBIACEAE

5b. Ovary and fruit densely hirsute or tomentose.

7a. Leaf blade sparsely pubescent
or scabrous at least along veins
abaxially.
8a. Leaf blade linear-lanceolate
or narrowly lanceolate ..... 7c. var. ciliatum
8b. Leaf blade broadly
lanceolate or
ovate-lanceolate .... 7f. var. kamtschaticum
7b. Leaf blade glabrous abaxially.
9a. Leaf blade less than
4 mm wide ......................... 7b. var. boreale
9b. Leaf blade 415 mm wide.
10a. Leaf blade 46
mm wide ......... 7a. var. angustifolium
10b. Leaf blade wider
than 6 mm .............. 7i. var. latifolium
7a. Galium boreale var. angustifolium (Freyn) Cufodontis,
Oesterr. Bot. Z. 89: 226. 1940.
xia ye zhen cao
Galium rubioides Linnaeus var. angustifolium Freyn,
Oesterr. Bot. Z. 45: 341. 1895.
Leaf blade narrowly lanceolate, 46 mm wide, abaxially
glabrous. Ovary and mericarps densely hirsute or tomentose. Fl.
JunAug, fr. JulSep.
Mountain slopes, river valleys, swamps, grasslands, meadows;
5003900 m. Hebei, Heilongjiang, Nei Mongol, Sichuan, Xinjiang,
Xizang [Japan, Kashmir, Russia; NE Europe].

7b. Galium boreale var. boreale

() bei fang la la teng (yuan bian zhong)
Galium boreale var. vulgare Turczaninow.
Leaf blade narrowly lanceolate or linear-lanceolate, 13.9
mm wide, glabrous. Ovary and mericarps densely hirsute or
tomentose with slightly curved, white trichomes. Fl. MayAug,
fr. JunOct.
Forests, thickets, or tussocks on mountain slopes, ditch sides,
grasslands, meadows; 7003900 m. Gansu, Hebei, Heilongjiang, Jilin,
Liaoning, Nei Mongol, Qinghai, Shandong, Shanxi, Sichuan, Xinjiang,
Xizang [India, Korea, Pakistan, Russia; Europe, North America].

7c. Galium boreale var. ciliatum Nakai, J. Jap. Bot. 15: 340.
1939.
ying mao la la teng
Leaf blade linear-lanceolate or narrowly lanceolate, 16
mm wide, abaxially sparsely hairy or scabrous at least along
veins. Ovary and mericarps densely hirsute or tomentose. Fl.
JunSep, fr. JulOct.
Mountain slopes, river beaches, ditch sides, open fields, meadows;
2004600 m. Gansu, Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol,
Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan
[Japan, Russia; Europe (Finland, Romania), North America].

7d. Galium boreale var. hyssopifolium (Hoffmann) Candolle,

Prodr. 4: 600. 1830.
fei suo pu zhen cao

117

Galium hyssopifolium Hoffmann, Deutschl. Fl., Dritter

Jahrgang, 71. 1800; G. boreale f. hyssopifolium (Hoffmann) B.
Boivin; G. boreale subsp. hyssopifolium (Hoffmann) Schbler
& G. Martens; G. rubioides var. hyssopifolium (Hoffmann) Persoon.
Leaf blade linear-lanceolate or narrowly lanceolate, 16
mm wide, glabrous abaxially. Ovary and mericarps glabrous.
Fl. and fr. JunAug.
Mountain slopes, grasslands; 18002300 m. Sichuan, Xinjiang
[Europe].
This variety probably has a wider geographic range.

7e. Galium boreale var. intermedium Candolle, Prodr. 4: 601.

1830.
xin zhen cao
Leaf blade linear-lanceolate or narrowly lanceolate, 16
mm wide, sparsely pubescent to glabrescent. Ovary and mericarps sparsely hirtellous or scabrous. Fl. JunJul, fr. JulOct.
Mountain slopes, wastelands, forests, grasslands, meadows; 1500
1800 m. Gansu, Heilongjiang, Xinjiang [Russia; Europe].

7f. Galium boreale var. kamtschaticum (Maximowicz) Nakai,

Bot. Mag. (Tokyo) 23: 103. 1909.
kan cha jia la la teng
Galium boreale f. kamtschaticum Maximowicz, Mm.
Acad. Imp. Sci. St.-Ptersbourg Divers Savans 9 [Prim. Fl.
Amur.]: 141. 1859; G. boreale var. koreanum Nakai.
Leaf blade broadly lanceolate or ovate-lanceolate, 610
mm wide, abaxially sparsely hairy or scabrous at least along
veins. Ovary and mericarps densely hirsute or tomentose. Fl.
and fr. JunSep.
Mountain slopes, farmland sides, riversides, river valleys, grasslands, meadows; 8002400 m. Heilongjiang, Henan, Jilin, Liaoning,
Nei Mongol, Shaanxi, Shanxi, Sichuan, Xinjiang [Kashmir, Korea,
Mongolia, Russia; C Asia (Turkestan), NE Europe].
Nakai (Bot. Mag. (Tokyo) 23: 103. 1909) commented on Japanese
plants and presented a key to Galium, including Galium boreale var.
kamtschaticum Maximowicz with no further information. Some authors have considered this a validly published combination (e.g., in Govaerts et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on 16 Nov 2007), whereas others have not (e.g., W. C.
Chen, loc. cit.: 263) and have instead cited J. Coll. Sci. Imp. Univ.
Tokyo 31: 498. 1911 (sometimes called Fl. Koreana 2) as the place of
publication. Nakais new combination was indeed validly published in
1909 because, before 1953, merely citing an authors name could constitute indication of a basionym (Vienna Code, Art. 33.2). Pobedimova
et al. (loc. cit.: 419) erroneously attributed this varietal name to Maximowicz, 1859; it was published as a forma there.

7g. Galium boreale var. lanceolatum Nakai, J. Jap. Bot. 15:

341. 1939.
guang guo zhen cao
Galium boreale var. leiocarpum Nakai; G. ussuriense Pobedimova.
Leaf blade broadly lanceolate or ovate-lanceolate, 612
mm wide, glabrous abaxially. Ovary and mericarps glabrous.
Fl. and fr. MaySep.

RUBIACEAE

118

Mountain slopes, open fields, grasslands; 9001900 m. Heilongjiang, Jilin, Xinjiang [Korea, Russia; C Asia (Turkestan)].
Nakais two varieties were published simultaneously; the choice
of epithet was apparently made by Cufodontis (Oesterr. Bot. Z. 89: 227.
1940).

7h. Galium boreale var. lancilimbum W. C. Chen, Acta

Phytotax. Sin. 28: 302. 1990.
pi zhen ye zhen cao
Leaf blade linear-lanceolate or narrowly lanceolate, 16
mm wide, abaxially pilosulous or scabrous at least along veins.
Ovary and mericarps glabrous. Fl. and fr. summer and autumn.
Mountain slopes, grasslands, meadows, ditch sides, wastelands;
18003000 m. Gansu, Heilongjiang, Sichuan, Xinjiang.

7i. Galium boreale var. latifolium Turczaninow, Bull. Soc.

Imp. Naturalistes Moscou 18: 315. 1845.
kuan ye la la teng
Leaf blade ovate-lanceolate, 615 mm wide, glabrous
abaxially. Ovary and mericarps densely hirsute or tomentose.
Fl. and fr. JunSep.
Mountain slopes, grasslands, meadows, farmland sides, river
beaches; 7002700 m. Gansu, Heilongjiang, Jilin, Liaoning, Nei
Mongol, Ningxia, Shanxi, Xinjiang [Kashmir, Korea, Russia; C Asia
(Turkestan)].

7j. Galium boreale var. pseudorubioides Schur, Enum. Pl.

Transsilv. 280. 1866.
jia qian zhen cao
Galium boreale subsp. pseudorubioides (Schur) So.
Leaf blade broadly lanceolate or ovate-lanceolate, 512
mm wide, sparsely pubescent to glabrescent abaxially. Ovary
and mericarps sparsely hirtellous or scabrous. Fl. and fr. Jun
Jul.
Mountain slopes, meadows; ca. 1400 m. Heilongjiang, Jilin,
Xinjiang [Russia; C Asia (Turkestan), Europe].

7k. Galium boreale var. rubioides (Linnaeus) elakovsky,

Prodr. Fl. Bhmen 2: 281. 1872.
qian zhen cao
Galium rubioides Linnaeus, Sp. Pl. 1: 105. 1753.
Leaf blade ovate-lanceolate or ovate, 46 mm wide, abaxially pilosulous or scabrous at least along veins. Ovary and
mericarps glabrous. Fl. and fr. JunSep.
Mountain slopes, grasslands; 11001400 m. Hebei, Heilongjiang,
Henan, Jilin, Liaoning, Xinjiang [Russia; Europe].
This broad- and large-leaved taxon is quite distinct from Galium
boreale s.s. in Europe and is usually treated there as a separate species.
Contrary to the above distribution data given by W. C. Chen (loc. cit.:
261) and according to Pobedimova et al. (loc. cit.: 420) it does not
extend into Asia.

8. Galium bullatum Lipsky, Trudy Imp. S.-Peterburgsk. Bot.

Sada 13: 300. 1894.

pao guo la la teng

Subshrubs, perennial, erect or ascending, sometimes caespitose, 540 cm tall. Rootstock stout, woody. Stems 4-angled,
very shortly pilose at base, glabrous and smooth above. Leaves
in whorls of 58, drying blackish, linear to linear-oblanceolate,
1227 12 mm, glabrous or sparsely ciliolate toward acute
apex; vein 1. Inflorescences terminal on main and short lateral
branches, cymose to corymbiform, lax, few to several flowered;
axes glabrous and smooth; bracts reduced or none; pedicels 14
mm. Ovary ovoid, ca. 1.5 mm, glabrous. Corolla white, cupshaped to subrotate, 3.55 mm in diam., glabrous; lobes 4, lanceolate-oblong, slightly mucronulate. Fruit usually from 1 mericarp only, subglobose, 33.5 mm in diam., glabrous, smooth,
white, with pericarp inflated, spongy to fleshy. Fl. JunJul, fr.
JulAug.
Grasslands, meadows; ca. 500 m. ?Xinjiang [SW Asia (Armenia,
?Iran, Nakhichevan)].
Galium bullatum is a member of G. sect. Orientigalium centered
in SW Asia and characterized by slightly cup-shaped corollas, never
retrorsely aculeolate stems, etc. The above diagnosis is taken from the
original description and a collection by Szovits in W (in Persia
borealis). We have not seen a specimen from China. The description by
W. C. Chen (in FRPS 71(2): 274. 1999), evidently based on plants from
Xinjiang, deviates from the authentic material in W by describing the
stems as retrorsely hispidulous along angles and the corolla as rotate.
Species of G. sect. Orientigalium usually are rather locally distributed
(Ehrendorfer et al., Fl. Iranica 176: 205231. 2005), and the distance
between Nakhichevan and Xinjiang is enormous. All this makes it quite
unlikely that G. bullatum (or even other related members of G. sect.
Orientigalium) really occurs in China. A definite decision has to wait
until voucher specimens become available for comparison.

9. Galium bungei Steudel, Nomencl. Bot., ed. 2, 1: 657. 1840.

si ye l
Herbs, perennial, 550 cm tall, erect from tender reddish
rootstock or filiform rhizome. Stems often caespitose, 4-angled,
unbranched or little branched, smooth, glabrous and smooth or
pilosulous to pilose, rarely retrorsely aculeolate, at nodes hispidulous. Leaves in whorls of 4, subsessile; blade drying papery, ovate-oblong, ovate-lanceolate, lanceolate-oblong, elliptic-oblong, or narrowly oblanceolate, (6)820(34) (2)3
7(10) mm, length/breadth index usually 35, glabrous and
sometimes antrorsely aculeolate on midrib and near margins, to
pilosulous or pilose throughout, lower side sometimes glandular-punctate or striate, base cuneate, apex acute or slightly obtuse; 1 principal vein, 2 lateral veins usually inconspicuous. Inflorescences terminal and/or axillary, cymose to paniculate,
congested to lax, cymes few to several flowered, 15 cm; peduncles glabrous, smooth; bracts none or few, spatulate to narrowly elliptic, 15 mm; pedicels (1)24(7) mm. Ovary subglobose to ellipsoid, laterally somewhat flattened, 0.40.8 mm,
glabrous to strigillose, smooth to tuberculate. Corolla yellowish
green or white, rotate, 1.52.5 mm in diam., glabrescent; lobes
4, ovate or oblong, acute to acuminate. Mericarps ellipsoid, 12
mm, tuberculate, aculeolate or with appressed and curved to
spreading and uncinate trichomes ca. 0.3 mm, rarely glabrous
and smooth. Fl. AprSep, fr. MayJan.

RUBIACEAE

Forests, thickets, or meadows on mountains, hills, open fields,

farmlands, ditch sides, riversides and beaches, streamsides; near sea
level to 3600 m. Anhui, Fujian, Gansu, Guangdong, Guangxi, Guizhou,
Hebei, Heilongjiang, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Liaoning,
Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi, Sichuan, Taiwan,
Yunnan, Zhejiang [Japan, Korea].
Galium bungei designates a group of tender perennial herbs from
G. sect. Platygalium, widespread and common at low to middle elevations throughout China. Galium bungei is also used medicinally there.
The group is very variable with respect to habit and inflorescence, as
well as stem, leaf, and fruit indumentum (the latter from tuberculate to
spreading hooked trichomes). The small and inconspicuous flowers suggest autogamous reproduction. All this has caused the recognition of
several species. In view of the gradual nature of this variation and the
partly simple genetic basis of the underlying differences, we give them
less taxonomic weight, follow Cufodontis (Oesterr. Bot. Z. 89: 219
223. 1940) and the Kew Rubiaceae checklist (Govaerts et al., World
Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on
15 Sep 2010), and include them all as synonyms under G. bungei s.l.
Only the closely related G. salwinense, endemic in Sichuan and Yunnan,
is maintained on the basis of its elongated and slender pedicels and the
constant hooked fruit trichomes.
Forms of Galium bungei with broader leaves in Sichuan (e.g., var.
punduanoides) develop, in addition to the principal vein, stronger side
veins somewhat approaching the larger G. yunnanense, typically with 3veined leaves, which occurs in the same area.
To bring some schematic order into the extreme variation of a
broadly circumscribed Galium bungei s.l., Cufodontis (loc. cit.: 221
222) created six varieties, without giving much weight to differences in
fruit surface. These varieties were taken up by W. C. Chen in FRPS
(71(2): 247250. 1999) and are also presented here. In contrast to this
approach, Yamazaki (J. Jap. Bot. 61: 51. 1991; Fl. Japan 3a: 236237.
1993) recognized several of these varieties as species for the Flora of
Japan: G. pogonanthum (corresponding to G. bungei var. setuliflorum),
separated by having appressed upcurved short hairs on its fruit rather
than spreading hooked trichomes as G. bungei s.s.; G. gracilens (corresponding to G. bungei var. bungei), characterized by short appressed
punctate fruit hairs and slender inflorescences; and G. trachyspermum
(corresponding to G. bungei var. trachyspermum), with short appressed
hooked fruit hairs and more condensed inflorescences. For each of these
segregate species Yamazaki (loc. cit. 1991; loc. cit. 1993) also created
several new additional varieties not considered here. For their treatment
of Taiwanese members of G. bungei s.l. Yang and Li (Bull. Natl. Mus.
Nat. Sci., Taichung 11: 105106. 1998; Fl. Taiwan, ed. 2, 4: 255256.
1998) accepted two species: G. gracilens with tuberculate fruit and G.
fukuyamae with appressed uncinate fruit hairs. Below, we present the
schematic infraspecific classification of W. C. Chen in FRPS. Descriptions are sketchy due to the limited material available. Nevertheless, this
may help as a reference and basis for urgently needed future studies on
this polymorphic and phylogenetically important group.

1a. Stems pubescent.

2a. Pubescence with trichomes shorter
than diam. of stems .......................... 9c. var. hispidum
2b. Pubescence with trichomes longer
than diam. of stems ................. 9d. var. punduanoides
1b. Stems glabrous, hairy only at nodes.
3a. Corolla lobes sparsely pubescent
at least in bud .............................. 9e. var. setuliflorum
3b. Corolla lobes glabrous.
4a. Leaf blade broadly elliptic,
obovate, or broadly lanceolate;

119

inflorescences crowded,
congested to subcapitate
........................................ 9f. var. trachyspermum
4b. Leaf blade narrowly
lanceolate, linear-lanceolate,
or ovate-lanceolate;
inflorescences lax.
5a. All leaves narrowly
lanceolate or
linear-lanceolate,
to 3 cm ..................... 9a. var. angustifolium
5b. Lower stem leaves
ovate-lanceolate,
upper stem leaves
narrower, often less
than 2 cm ............................ 9b. var. bungei
9a. Galium bungei var. angustifolium (Loesener) Cufodontis,
Oesterr. Bot. Z. 89: 221. 1940.
xia ye si ye l
Galium gracile f. angustifolium Loesener, Beih. Bot. Centralbl., Abt. 2, 37: 182. 1920.
Leaf blade narrowly lanceolate or linear-lanceolate, to 3
cm. Fl. JunJul, fr. AugOct.
Anhui, Fujian, Gansu, Hebei, Henan, Jiangsu, Jiangxi, Shaanxi,
Shandong, Shanxi, Zhejiang.

9b. Galium bungei var. bungei

() si ye l (yuan bian zhong)
Galium fukuyamae Masamune; G. gracile Bunge (1833),
not Wallroth (1822); G. gracile var. miltorrhizum (Hance)
Loesener; G. gracilens (A. Gray) Makino; G. lutchuense Nakai;
G. miltorrhizum Hance; G. remotiflorum H. Lveill & Vaniot;
G. trachyspermum A. Gray var. gracilens A. Gray.
Stems glabrous. Leaf blade ovate-lanceolate at lower part,
attenuate at upper part. Inflorescence paniculate and lax. Corolla glabrous. Fl. AprSep, fr. MayJan.
Forests or meadows on hills or mountains, open fields, farmlands,
ditch sides; below 1002600 m. Anhui, Fujian, Gansu, Guangdong,
Guangxi, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jiangsu,
Jiangxi, Liaoning, Nei Mongol, Ningxia, Shaanxi, Shandong, Shanxi,
Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Korea].

9c. Galium bungei var. hispidum (Matsuda) Cufodontis,

Oesterr. Bot. Z. 89: 222. 1940.
ying mao si ye l
Galium gracile f. hispidum Matsuda, Bot. Mag. (Tokyo)
26: 130. 1912; ?G. martini H. Lveill & Vaniot; G. trachyspermum var. hispidum (Matsuda) Kitagawa.
Stems soft pubescent, trichomes shorter than diam. of
stem. Fl. AprJun, fr. MaySep.
Forests or meadows on mountain slopes, river beaches, open
fields; 1003400 m. Anhui, Fujian, Gansu, Henan, Hubei, Jiangsu,
Shaanxi, Shanxi, Sichuan, Yunnan, Zhejiang.

RUBIACEAE

120

In the protologue Cufodontis correctly cited the basionym of his

variety but incorrectly gave his varietal name as var. hispidum (Kitag.)
Cufodontis when the basionym author was actually Matsuda.
Galium martini was accepted as a dubious species by W. C. Chen
in FRPS (loc. cit.: 282). The protologue (Bull. Soc. Bot. France 55: 58.
1908) is quite incomplete but probably refers to a strongly hairy plant of
G. bungei s.l. from Guizhou. Therefore, the name is provisionally placed
here as a possible synonym of var. hispidum or var. punduanoides until
a detailed study of the type (E) will clarify the matter.

9d. Galium bungei var. punduanoides Cufodontis, Oesterr.

Bot. Z. 89: 222. 1940.
mao si ye l
?Galium martini H. Lveill & Vaniot.
Stem soft to stiffly pubescent, trichomes longer than diam.
of stems. Inflorescence often more congested and terminal. Fl.
JunJul, fr. JulAug.
Forests, thickets, or meadows on mountains, open fields, riversides; 9003600 m. Gansu, Jiangsu, Sichuan, Yunnan.

9e. Galium bungei var. setuliflorum (A. Gray) Cufodontis,

Oesterr. Bot. Z. 89: 221. 1940.
mao guan si ye l
Galium trachyspermum var. setuliflorum A. Gray, Mem.
Amer. Acad. Arts, n.s., 6: 393. 1859; G. pogonanthum Franchet
& Savatier; G. pogonanthum var. setuliflorum (A. Gray) H.
Hara; G. setuliflorum (A. Gray) Makino.
Corolla lobes sparsely pubescent at least in bud.
Jiangsu, Shanxi [Japan, Korea].

9f. Galium bungei var. trachyspermum (A. Gray) Cufodontis,

Oesterr. Bot. Z. 89: 221. 1940.
kuo ye si ye l
Galium trachyspermum A. Gray in Perry, Narr. Exped.
China Japan 2: 313. 1856; G. venosum H. Lveill.
Leaf blade broadly elliptic, obovate, or broadly lanceolate.
Inflorescences congested to subcapitate. Fl. AprMay, fr. Apr
Jul.
Forests or meadows on hills, open fields, streamsides; near sea
level to 800 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hebei,
Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Shandong, Sichuan, Zhejiang
[Japan, Korea].

10. Galium chekiangense Ehrendorfer, Novon 20: 270. 2010.

zhe jiang la la teng
Herbs, perennial, with thin rhizomes. Stems 24 from a
common base, erect, usually unbranched, 2030(40) cm tall,
with only 4 or 5 internodes from base to first inflorescence
node, with 4 prominent, rounded, and whitish angles, glabrous
and smooth except for short and stiff hairs at nodes. Leaves
and leaflike stipules in whorls of 4, subsessile; blade drying
somewhat leathery and light brownish green, broadly elliptic
to ovate, (15)2330(50) (8)1115(25) mm, length/
breadth index 2.5 or less, gradually narrowed into base,
broadest near middle, gradually narrowed into apex, mostly

shortly apiculate but without a hyaline point, glabrous except

for antrorse microhairs (0.10.2 mm) along 3 main veins,
mostly on lower but also on upper side and along slightly
revolute margins, papillose on upper side under a strong lens
(20), lower side clearly marked by numerous darker and linear
idioblasts. Inflorescences loosely thyrsoid and elongate, from
uppermost 2 or 3 nodes, clearly longer than subtending leaves,
cymes lateral and terminal, somewhat divaricate, 14 cm, with
few and inconspicuous linear bracts and rather few flowered;
peduncles 12 cm and pedicels 0.11 cm. Ovary obovoid, 0.8
1 mm. Corolla greenish white, rotate, ca. 3 mm in diam.,
glabrous; lobes 4, acuminate. Ripe mericarps ovoid, 1.52(3)
mm, densely covered by stiff and appressed to slightly divergent microhairs, 0.150.25 mm and with an acute and bent
apex. Fl. Jul, fr. Aug.
Lower montane forests; ca. 1400 m. Fujian, Zhejiang.
Specimens of Galium chekiangense from the province of Fujian
were included and described in FRPS (71(2): 265. 1999) under the
name of G. nakaii Kud ex H. Hara (J. Jap. Bot. 9: 517. 1933). These
Fujian vouchers were not available, but two fruiting specimens from the
adjacent province of Zhejiang (formerly Chekiang: Xi ming shan) in the
herbarium PE could be studied. They were determined as G. kamtschaticum and closely correspond to the description of G. nakaii in
FRPS. These PE specimens deviate clearly from authentic Japanese
specimens of G. nakaii as well as from G. kamtschaticum and G.
oreganum Britton. This has justified the description of G. chekiangense
as a new and endemic Chinese species and makes G. nakaii an endemic
of Japan.
Galium chekiangense clearly belongs to G. sect. Platygalium s.l.
(Ehrendorfer et al., Fl. Iranica 176: 175. 2005) and apparently is a
member of the G. kamtschaticum species group, which includes the
amphi-Beringian G. kamtschaticum (in China limited to elevations of
15002300 m in the NE provinces of Heilongjiang and Jilin), the
Japanese G. nakaii from Hokkaido and N Honshu, and the W North
American G. oreganum. These three latter species differ from G.
chekiangense by their leaf blades drying dark brownish (not light
brownish green), thin, membranous and smooth, neither papillose
above nor with glandular-striate idioblasts below, and by their ripe
mericarps with much longer uncinate trichomes (0.81 mm, not 0.15
0.25 mm). In addition, G. nakaii has inflorescence cymes mostly
shorter (not clearly longer) than the subtending leaves.
For further and more detailed studies of the Galium kamtschaticum group, one should refer to the wide circumscription of G. kamtschaticum (with three varieties) and the confused, partly contradictory
description of the fruit indumentum of G. nakaii in Yamazaki (Fl. Japan
3a: 234235. 1999).

11. Galium consanguineum Boissier, Diagn. Pl. Orient., ser. 1,

6: 69. 1846.
juan bian la la teng
Galium consanguineum subsp. majmechense (Bordzilowski) A. D. Mikheev; G. majmechense Bordzilowski; G. verum
Linnaeus var. consanguineum (Boissier) Boissier.
Herbs, perennial, often caespitose from a stout and woody
rootstock with rhizomes. Stems erect, to 1 m tall, 4-angled,
glabrous to puberulent at least at nodes, smooth. Leaves in
whorls of 612, sessile; blade drying papery, linear-oblanceolate to linear, 2028 13 mm, glabrous, more rarely hairy,

RUBIACEAE

base acute to straight, margin shortly to strongly revolute and

usually antrorsely aculeolate, apex acute and mucronate; vein 1.
Inflorescences narrowly paniculate with main stems and short
lateral and terminal, few- to several- or many-flowered, rather
congested cymes; peduncles glabrous, smooth; pedicels 0.53
mm, subtended by leaflike bracts. Ovary subglobose to obovoid, 0.50.8 mm, smooth, glabrous or hispidulous with
straight hairs. Corolla yellow, rotate, ca. 3 mm in diam., glabrous, lobed for 3/4 or more; lobes 4(or 5), lanceolate-oblong,
acute to acuminate. Mericarps ellipsoid to obovoid, ca. 1.5 1
mm, glabrous or hispidulous with straight trichomes. Fl. Jul
Aug, fr. AugSep.
Thickets; [1300]1700[2800] m. Xinjiang [SW Asia (Armenia,
Azerbaijan, Iran, Iraq, Lebanon, E Turkey)].
Galium consanguineum was treated as G. majmechense (a
younger synonym) by W. C. Chen (FRPS 71(2): 269. 1999). It belongs to the polymorphic G. verum group or complex (see additional
comments under that species) and apparently links it (as a hybrid
taxon?) to more broadly leaved and glabrous members of G. sect.
Orientigalium (Ehrendorfer et al., Fl. Iranica 176: 205207. 2005).
Galium consanguineum can be separated from G. verum by its broader
(more than 2.5 mm), glabrous leaves, but intermediates occur.

12. Galium crassifolium W. C. Chen, Acta Phytotax. Sin. 28:

299. 1990.
hou ye la la teng
Herbs, perhaps perennial, ascending, ca. 10 cm tall. Stems
4-angled, caespitose, glabrous and smooth or sparsely puberulent. Leaves in whorls of 4, sessile or subsessile; blade drying
leathery, elliptic or ovate, 38 24 mm, scabrous with microhairs, base cuneate or subobtuse, margins antrorsely ciliolate,
apex obtuse and mucronate; vein 1. Inflorescences terminal
and/or axillary, cymose, few flowered, up to 1 cm; axes glabrous and smooth, somewhat bracteate; pedicels ca. 1 mm.
Flowers unknown. Mericarps ovoid, 0.51 mm, with appressed,
slightly curved microhairs, ca. 0.3 mm. Fl. AugSep, fr. Oct.
Valleys, open habitat; ca. 800 m. Shanxi (Zhenba).
We have seen no authentic material of Galium crassifolium, but
the original description is accompanied by a good drawing. In the
protologue similarities with the Taiwanese G. tarokoense are suggested.
Both species are distantly related, but apparently G. crassifolium falls
within the morphological amplitude of the polymorphic G. bungei. Its
distinctness with respect to reduced plants from dry habitats should be
checked in the future.

13. Galium dahuricum Turczaninow ex Ledebour, Fl. Ross. 2:

409. 1844.

121

gins, base acute to cuneate, apex acute to obtuse and mucronate; vein 1. Inflorescences paniculate, with axillary and terminal, several- to many-flowered, usually very lax and up to 7 cm
long cymes; axes filiform and often flexuous, sparsely aculeolate to glabrous; bracts few, lanceolate; pedicels slender, in flower 25 mm. Ovary obovoid, ca. 0.8 mm, with dense spreading
or appressed undeveloped trichomes, or glabrous. Corolla white
or pale green, rotate, of quite different sizes, (1)1.53(4) mm
in diam., glabrous; lobes 4, triangular, obtuse to acute or minutely apiculate. Mericarps ellipsoid, ca. 2 mm, with appressed
or spreading and uncinate trichomes (0.30.5 mm), tuberculate
to completely glabrous and smooth, on pedicels elongating to
10 mm or more. Fl. JunSep, fr. JulNov.
Humid forests, thickets, ditch sides, grasslands, meadows; 200
3400 m. Fujian, Gansu, Guizhou, Hebei, Heilongjiang, Henan, Hubei,
Hunan, Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai,
Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan, Zhejiang [Japan,
Korea, Russia].
Within Galium sect. Trachygalium and the extremely polymorphic
species group of G. asperifolium (see there) Cufodontis (Oesterr. Bot. Z.
89: 239243. 1940), Yamazaki (Fl. Japan 3a: 206240. 1993), and W. C.
Chen (in FRPS 71(2): 255258. 1999) differentiated the closely related
taxa G. dahuricum (in FRPS as G. davuricum, the spelling used in
the protologue by Ledebour), G. tokyoense, G. pseudoasprellum, and G.
manshuricum mainly according to the lack (in the two former) and the
presence (in the two latter) of appressed or spreading uncinate trichomes on the mericarps. As this character apparently often varies
within populations of these taxa, the present treatment relies on the
much more stable feature of slender, filiform, and flexuose vs. more
stiff and divaricate peduncles and pedicels. The former state characterizes G. dahuricum (including G. manshuricum and G. pseudoasprellum), the latter G tokyoense. This species assembly has its distribution
center in E Asia but extends with G. asprellum Michaux s.s. into E
North America.
Galium pseudoasprellum was accepted as a species by Cufodontis
(loc. cit.: 237238), W. C. Chen (loc. cit.: 254255), and Yamazaki (loc.
cit.: 238), and the latter two also maintained G. niewerthii. In our
opinion and because of their similar inflorescences, both taxa should be
regarded as synonyms of G. dahuricum, the former under var. lasiocarpum, the latter under var. dahuricum. Even if we have not seen authentic
specimens of G. niewerthii, all of its characters listed fall within the
limits of G. dahuricum; thus, we regard it as a glabrous-fruited form of
that variable species. So far, G. comarii has been a badly understood
taxon (see Cufodontis, loc. cit.: 241; Lauener, Notes Roy. Bot. Gard.
Edinburgh 32: 107. 1972; Mill, Edinburgh J. Bot. 53: 193213. 1996).
Because of its slender inflorescence, extremely long pedicels, and
rugose fruit mentioned in the protologue, it can now be safely assigned
as another synonym of G. dahuricum var. dahuricum.

da ye zhu yang yang

The following infraspecific taxa of Galium dahuricum (and G.

pseudoasprellum), accepted by Cufodontis (loc. cit.: 237238, 243244)
and W. C. Chen (loc. cit.), are keyed and listed here for comparison.

Herbs, perennial, from a slender reddish rootstock. Stems

erect to ascending, weak to procumbent and often climbing,
sometimes up to 2.5 m, 4-angled, sparsely to densely retrorsely
aculeolate along angles and at nodes, rarely glabrescent.
Leaves on main stems in whorls of 5 or 6, subsessile; blade
drying papery, of very variable shape, from obovate and elliptic-oblong to narrowly oblanceolate, (11)1540(55) (2)3
10(14), sparsely to densely retrorsely aculeolate adaxially
along midrib, abaxially, and along flat to thinly reflexed mar-

1a. Ovary and mericarps glabrous or

tuberculate .......................................... 13a. var. dahuricum
1b. Ovary and mericarps with spreading
or appressed uncinate trichomes.
2a. Inflorescences with loosely
branched cymes, with filiform
and flexuose pedicels of up
to 5 mm, in fruit up to
10 mm ...................................... 13b. var. lasiocarpum

RUBIACEAE

122

2b. Inflorescences more congested,

pedicels shorter than 5 mm ....... 13c. var. densiflorum

14. Galium echinocarpum Hayata, J. Coll. Sci. Imp. Univ.

Tokyo 30(1): 147. 1911.

13a. Galium dahuricum var. dahuricum

ci guo zhu yang yang

() da ye zhu yang yang (yuan bian zhong)

Herbs, perennial, emerging from reddish, filiform rhizomes. Stems ascending to erect, 1040 cm tall, 4-angled, glabrous and smooth. Leaves in whorls of (4 or)5 or 6, subsessile;
blade drying papery, oblanceolate, obovate, narrowly elliptic, or
narrowly oblanceolate, 625 27 mm, glabrous or sometimes
sparsely hispid to strigillose, base acute, margins flat, smooth or
antrorsely aculeolate, apex acute to obtuse or rounded and
abruptly mucronate; vein 1. Inflorescences terminal and in axils
of upper leaves, with lax, few- to several-flowered cymes; axes
glabrous, smooth; bracts none or leaflike, 26 mm; pedicels
0.52 mm. Ovary subglobose, 0.50.7 mm, densely strigillose
with undeveloped trichomes. Corolla white, rotate, ca. 2 mm in
diam., glabrous, lobed for 3/4 or more; lobes 4, triangular, apex
obtuse. Mericarps subglobose to ellipsoid, ca. 2 mm, with dense
uncinate trichomes ca. 1 mm, on pedicels elongating to 10 mm.
Fl. May, fr. MayDec.

Galium asprellum Michaux var. dahuricum (Turczaninow

ex Ledebour) Maximowicz; G. comarii H. Lveill & Vaniot; G.
dahuricum var. leiocarpum Nakai; G. niewerthii Franchet &
Savatier.
Stems and nodes often glabrescent. Leaf blade with margins retrorsely aculeolate. Inflorescences lax; peduncles and
pedicels strongly elongated. Ovary and mericarps glabrous or
tuberculate. Fl. JunJul, fr. AugOct.
Forests, grasslands; 7001000 m. Fujian, Guizhou, Hebei, Heilongjiang, Hubei, Hunan, Jilin, Liaoning, Nei Mongol, Sichuan, Xinjiang, Yunnan [Korea, Russia].
This variety was listed as Galium niewerthii for the Flora of Japan (Yamazaki, loc. cit.: 238). Forms with many-flowered inflorescences, somewhat shorter pedicels, and glabrous ovaries and fruit
have been seen from Yunnan; they apparently tend toward G. asperifolium and/or G. prattii. Galium taiwanense (see there) is very close to
G. dahuricum var. dahuricum and evidently replaces it on Taiwan.

Montane forest regions, grassy fields, along drainage ditches;

9003500 m. Taiwan.

13b. Galium dahuricum var. lasiocarpum (Makino) Nakai, J.

Coll. Sci. Imp. Univ. Tokyo 31: 498. 1911.

Galium echinocarpum is very similar to G. hoffmeisteri and

replaces it on Taiwan. Galium takasagomontanum may belong here as a
synonym (see there).

dong bei zhu yang yang

15. Galium elegans Wallich in Roxburgh, Fl. Ind. 1: 382. 1820.

Galium asprellum var. lasiocarpum Makino, Bot. Mag.

(Tokyo) 17: 76. 1903; G. dahuricum var. manshuricum (Kitagawa) H. Hara; G. manshuricum Kitagawa; G. pseudoasprellum
Makino.

xiao hong shen

Stem nodes and leaf blade retrorsely aculeolate. Inflorescences lax; peduncles and pedicels elongated. Ovary and mericarps with dense appressed or spreading uncinate trichomes. Fl.
JunJul, fr. AugOct.
Forests, meadows, ditch sides; 3001100 m. Gansu, Hebei, Heilongjiang, Henan, Jiangsu, Jilin, Liaoning, Qinghai, Shaanxi, Shanxi,
Sichuan, Yunnan [Japan, Korea].

13c. Galium dahuricum var. densiflorum (Cufodontis) Ehrendorfer, Novon 20: 277. 2010.
mi hua la la teng
Galium pseudoasprellum var. densiflorum Cufodontis,
Oesterr. Bot. Z. 89: 237. 1940.
Stems often lower and leaf blade smaller. Inflorescences
shorter and more congested; pedicels rarely longer than 5 mm;
bracts larger, similar to leaves. Ovary and mericarps with dense
spreading uncinate trichomes. Fl. JulAug, fr. AugNov.
Forests, thickets, meadows on mountains; 7003400 m. Gansu,
Guizhou, Hebei, Henan, Jiangxi, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan.
The above geographic indications for this variety are uncertain
because of confusion with Galium tokyoense, etc. At least in part, G.
dahuricum var. densiflorum may refer to transitional (?hybrid) forms of
G. dahuricum var. lasiocarpum with G. asperifolium, G. blinii, G. prattii,
and/or G. sungpanense.

Herbs, perennial, climbing or procumbent to usually erect,

0.11 m tall, from a slender rootstock with purplish rhizomes.
Stems somewhat stout, 4-angled, smooth, sparsely to densely
hirsute, villous, or villosulous and often densely puberulent at
nodes, angles thickened. Leaves in whorls of 4, subsessile or
petiole to 1.5 mm; blade drying papery to leathery, green to
gray, or dark brown, ovate to broadly elliptic, 630 320 mm,
length/breadth index mostly 2 or less, sparsely to densely hirtellous, villosulous, or hispidulous to scaberulous at least on
principal veins, abaxially often glandular-punctate and/or striate, base rounded to acute, margins antrorsely ciliate to ciliolate and flat to thinly revolute, apex rounded to obtuse; principal veins palmate, 3(or 5). Inflorescences thyrsoid to paniculiform, with several- to many-flowered, 210 cm long cymes in
uppermost leaf axils and terminal; peduncles glabrescent to
sparsely scaberulous, hirtellous, puberulent, or villosulous;
bracts narrowly spatulate to narrowly elliptic, 13 mm; pedicels 0.52.5 mm. Flowers dioecious, polygamo-dioecious, or
sometimes ?hermaphroditic. Ovary obovoid, in staminate flowers ca. 0.5 mm and glabrous to scaberulous or sparsely strigillose, in pistillate and bisexual flowers 0.81 mm and usually
moderately to densely strigillose, particularly on their lateral
side. Corolla white or pale yellow, rotate, 22.5 mm in diam.,
glabrous; lobes 4, ovate-triangular, acute to rounded. Mericarps
ellipsoid, 11.5 mm, with sparse to dense and spreading
uncinate trichomes 0.50.8 mm, rarely scaberulous or glabrous. Fl. AprAug(Oct), fr. MayDec.
Forests, thickets, meadows on mountain slopes, streamsides, open
fields, on rocks; 2003500 m. Anhui, Fujian, Gansu, Guangxi, Guizhou,

RUBIACEAE

Hunan, Qinghai, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bangladesh, Bhutan, India, Kashmir, Myanmar, Nepal, Pakistan, Thailand].
Galium elegans is a widely ranging, polymorphic species that may
not be completely distinct from several other related taxa. It is here circumscribed more narrowly than by Cufodontis (Oesterr. Bot. Z. 89:
228232. 1940) and W. C. Chen (in FRPS 71(2): 242245. 1999),
which reduces its variation a bit. These aspects are discussed below.

123

thickly textured, not black

when dry ................................. 15a. var. elegans
3b. Stems pilose or glabrescent at
least on upper parts; leaf blade
often large and thinly textured,
blackish brown when dry
....................................... 15b. var. glabriusculum

Plants with narrower leaves are separated here as Galium yunnanense. This species comprises two of the varieties included by Cufodontis in G. elegans, i.e., G. elegans var. angustifolium and G. elegans
var. nemorosum. Separation of the two taxa is not always easy, as transitional specimens occasionally occur.

15a. Galium elegans var. elegans

Plants of Galium elegans with shortened pedicels and more congested cymes (e.g., from Sichuan, Shimian Xian) may approach the
Himalayan G. confertum Royle ex J. D. Hooker.

Stems sparsely or densely hirsute. Leaf blade drying

thickly textured, not black, ovate to ovate-lanceolate or broadly
elliptic, length/breadth index usually less than 2, apex obtuse or
slightly acuminate. Mericarps with spreading uncinate trichomes. Fl. AprAug, fr. MayDec.

Galium nephrostigmaticum was described as a species by Diels,

an opinion still maintained by some authors. Here, it is treated as a
variety of G. elegans, following W. C. Chen (Acta Phytotax. Sin. 28:
301. 1990) who referred to its glabrous to scaberulous ovaries and fruit,
as noted in Dielss protologue. In contrast to this, Cufodontis (loc. cit.)
synonymized G. nephrostigmaticum with G. elegans var. elegans. In an
extensive discussion he demonstrated that G. elegans is dioecious and
that G. nephrostigmaticum was based on a male plant with staminate
flowers and reduced glabrous to smooth ovaries and sessile stigmas. In
contrast, pistillate flowers of G. elegans have hairy ovaries and fruit,
well-developed styles, and reduced stamens. Thus, according to Cufodontis, G. nephrostigmaticum does not merit taxonomic recognition.
Ehrendorfer et al. (Fl. Iranica 176: 177. 2005) did not contradict the
conclusions of Cufodontis but noted that some plants of G. elegans are
monoecious or have bisexual flowers. This shows that the reproductive
biology of this species apparently is more complex than thought before.
Provisionally, G. nephrostigmaticum is treated here as a variety, pending
more detailed studies of this critical group.
The still uncertain relationships between Galium elegans on the
Chinese mainland and G. formosense on Taiwan are discussed under the
latter taxon. In the present treatment their separation is maintained provisionally.
The traditional varieties of Galium elegans are separated schematically by the density, type, and distribution of the indumentum on
the vegetative organs and have doubtful taxonomic value. W. C. Chen in
FRPS (loc. cit.) also used leaf size and apex shape as characters to differentiate these varieties. For reference and to facilitate comparison with
other works, we present a key to these infraspecific taxa below.

1a. Ovary (and fruit?) glabrous to

scaberulous ............................ 15c. var. nephrostigmaticum
1b. Ovary and fruit with uncinate
trichomes.
2a. Leaf blades in middle stem
region ovate-lanceolate,
length/breadth index 22.5,
apex acute or obtuse and
shortly acuminate, with a dense
and fine indumentum ................... 15d. var. velutinum
2b. Leaf blade ovate to ovate-lanceolate
or broadly elliptic, length/breadth
index usually less than 2, apex
usually obtuse or slightly acuminate.
3a. Stems sparsely or densely
hirsute; leaf blade small and

() xiao hong shen (yuan bian zhong)

Galium petiolatum Geddes.

Forests, thickets, meadows on mountain slopes, streamsides, open

fields, on rocks; 6003500 m. Anhui, Gansu, Guizhou, Hunan, Qinghai,
Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bangladesh, Bhutan, India, Kashmir, Myanmar, Nepal, Pakistan, Thailand].

15b. Galium elegans var. glabriusculum Requien ex Candolle, Prodr. 4: 600. 1830.
guang xi la la teng
Galium elegans f. glabriusculum (Requien ex Candolle)
H. Hara ex H. Ohba.
Stems pilose or glabrescent at least on upper parts. Leaf
blade drying blackish brown, often thinly textured, ovate,
ovate-lanceolate, or elliptic, 1033 518 mm, length/breadth
index usually 2 or less, apex mostly obtuse. Mericarps with
spreading uncinate trichomes. Fl. JulAug, fr. JulOct.
Forests or meadows on mountains and at streamsides; 11002900
m. Guangxi, Guizhou, Sichuan, Xizang, Yunnan [India, Nepal].
We have not seen authentic material of this taxon.

15c. Galium elegans var. nephrostigmaticum (Diels) W. C.

Chen, Acta Phytotax. Sin. 28: 301. 1990.
shen zhu la la teng
Galium nephrostigmaticum Diels, Notes Roy. Bot. Gard.
Edinburgh 5: 279. 1912.
Similar to Galium elegans var. elegans, but ovary (and
fruit?) glabrous to scaberulous. Fl. AprOct, fr. AugDec.
Forests, meadows; 2003000 m. Gansu, Guizhou, Sichuan,
Yunnan.
This taxon apparently refers to male plants only (see above).

15d. Galium elegans var. velutinum Cufodontis, Oesterr. Bot.

Z. 89: 230. 1940.
mao la la teng
Galium mairei H. Lveill.
Plants densely and finely pubescent, trichomes slender,
spreading. Leaf blade lanceolate or ovate-lanceolate, 715 ca.

RUBIACEAE

124

3 mm, length/breadth index 22.5, apex acute or obtuse and

shortly acuminate. Mericarps with spreading uncinate trichomes. Fl. and fr. Jul.
Meadows or on rocks on mountain slopes; 21002300 m. Sichuan, Yunnan.

16. Galium exile J. D. Hooker, Fl. Brit. India 3: 207. 1881.

dan hua la la teng
Galium handelii Cufodontis (1940), non Nblek (1923).
Herbs, annual, slender, procumbent to weak, 420 cm
tall. Roots slender, reddish when dry. Stems slender, 4-angled,
somewhat branched, sparsely retrorsely aculeolate to glabrous.
Middle stem leaves opposite with clearly smaller, leaflike stipules in whorls of 4; blades drying papery, obovate or oblanceolate to linear-elliptic, (2)3.510(12) 13.5(5) mm, adaxially with sparse appressed hairs or glabrous, margins mostly
antrorsely ciliolate, otherwise glabrous, base acute, cuneate, or
shortly petiolate, apex obtuse to acute but not mucronate; principal vein 1, with inconspicuous pinnate-reticulate lateral veins.
Flowers mostly solitary; pedicels 13 mm, glabrous. Ovary
subglobose, ca. 1 mm, densely covered with undeveloped trichomes. Corolla white, rotate, 11.5 mm in diam.; lobes 3(or
4), ovate, obtuse. Mericarps ovoid to elongated, 22.5 mm, with
dense, white to yellowish brown, uncinate trichomes 0.20.5
mm, on pedicels elongating to 10 mm and curved near apex. Fl.
JunJul, fr. AugSep.
Rock crevices on mountain slopes, sand and gravel drifts on
grassy plains; 12004800 m. Gansu, Nei Mongol, Ningxia, Qinghai,
Shaanxi, Shanxi, Sichuan, Xinjiang, Xizang, Yunnan [India, Nepal].
In his description of Galium handelii Cufodontis (Oesterr. Bot. Z.
89: 234235. 1940) referred to the close G. songaricum Schrenk (in
Fischer & C. A. Meyer, Enum. Pl. Nov. 1: 57. 1841) but overlooked the
older homonym by Nblek and the Himalayan G. exile. This latter annual has a much wider distribution than thought before and is quite
variable in China with respect to leaf shape and hairiness, length of
pedicels, shape of mericarps, etc. In view of its remarkably small flowers and high fruit set, it very likely is autogamous.
Within the morphologically and DNA-analytically very isolated
Galium sect. Depauperata (Ehrendorfer et al., Fl. Iranica 176: 231232.
2005) G. exile is morphologically very close to the W North American
G. bifolium S. Watson and particularly to G. songaricum, described from
the C Asiatic mountain system of Alatau. This latter species, treated in
FRPS as G. soongoricum, is assumed to differ by its 1- or 2(or 3)flowered cymes, the 4-lobed flowers, and the strongly elongating
fruiting pedicels. In FRPS (71(2): 224227. 1999) both taxa are
accepted and indicated for very much the same area in China. Nevertheless, a first analysis of all Chinese specimens in PE, KUN, MO, and
WU has not revealed reliable differential characters. Even the inflorescence character and the corolla lobe number varies in some specimens.
All these findings do not exclude the possibility that further and more
detailed studies will allow to separate the populations from the mountains of C and N Asia as typical G. songaricum. But for the Chinese
flora and the present treatment we recognize only one species, G. exile.
In case that the two taxa cannot be separated as species in the future G.
songaricum will have to replace G. exile for priority reasons.
As noted already by Cufodontis (loc. cit.), both Galium exile and
G. songaricum have sometimes been misidentified as G. pauciflorum, a
synonym of G. spurium from the G. aparine group. In contrast to G.

exile, the latter always has more than 4 leaf whorl elements, retrorsely
aculeolate leaf margins, and more than 1-flowered cymes.

17. Galium formosense Ohwi, Repert. Spec. Nov. Regni Veg.

36: 55. 1934.
guan shan zhu yang yang
Galium kwanzanense Ohwi.
Herbs, perennial, procumbent to erect, (5)820(30) cm
tall. Stems 4-angled, sparsely to rather densely pilose, angles
thickened. Leaves in whorls of 4, sessile; blade drying submembranous, blackish green, broadly elliptic to obovate, 420 3
10 mm, length/breadth index 2 or less, both surfaces sparsely to
densely pilose at least along veins, base cuneate to obtuse, apex
obtuse to rounded and mucronate; principal veins 3, palmate.
Inflorescences with terminal and axillary, few- to many-flowered, 13 cm long cymes; peduncles sparsely pilose to glabrous
and smooth; bracts spatulate to ovate, 1.53 mm; pedicels 14
mm. Flowers ?hermaphroditic. Ovary obovoid, ca. 0.5 mm,
densely pubescent with uncinate trichomes. Corolla yellowish
white, rotate, 12 mm in diam., glabrous; lobes 4, ovate, 0.4
0.8 mm, acute. Mericarps ovoid, ca. 1 mm, with dense white to
yellowish uncinate trichomes 0.40.5 mm. Fl. JunSep, fr. Jun
Nov.
Mountains, along trails and roads, fields, open ditches; 600
3000 m. Taiwan (Gaoxiong).
Ohwi described the relatively tall (2030 cm) Galium formosense from lower elevations and the condensed G. kwanzanense (510
cm) from an exposed higher peak of Taiwan. The technical differences
indicated mainly relate to flower diameter (1 mm in the former, 2 mm in
the latter). In their study of Taiwanese Galium Yang and Li (Bull. Natl.
Mus. Nat. Sci., Taichung 11: 101117. 1998; Fl. Taiwan, ed. 2, 4: 254
259. 1998) formally synonymized the taxa and demonstrate a considerable ecological amplitude of G. formosense s.l. Furthermore, the
specific separation of G. formosense from two other Taiwan mountain
endemics, with glabrous stems, G. morii and G. tarokoense, needs better
documentation.
In FRPS (71(2): 243. 1999), W. C. Chen treated Galium formosense as a synonym of G. elegans. He referred to Cufodontis (Oesterr.
Bot. Z. 89: 228. 1940) who supported the occurrence of G. elegans in
Taiwan based on Hayatas report of G. rotundifolium Linnaeus (in J.
Coll. Sci. Tokyo 30(1): 148. 1911) and to J. M. Chao (in Fl. Taiwan 4:
261. 1978) who considered G. elegans to be the same as G. formosense.
In their study of Taiwanese Galium Yang and Li (loc. cit. 1998; loc. cit.
1999) did not mention G. elegans nor compare G. formosense to it. This
rather suggests that they were unaware of Cufodontiss work than that
they concluded the two species to be distinct. Similarly, Cufodontis (loc.
cit.: 211251), studying only mainland material, did not mention G.
formosense, already described in 1934. The Taiwanese specimens at
MO (studied by C. M. Taylor) appear to represent a distinct species but
fall within G. elegans as more broadly circumscribed by Cufodontis
(loc. cit.: 228232). Thus, G. formosense is here provisionally separated
and regarded as replacing G. elegans on Taiwan. In future studies, it will
be of particular importance to clarify whether the dioecy or polygamodioecy found in G. elegans (see there) also occurs in G. formosense.

18. Galium forrestii Diels, Notes Roy. Bot. Gard. Edinburgh 5:

279. 1912.
li jiang la la teng
Herbs, perennial, 1525 cm tall. Stems little branched, 4-

RUBIACEAE

angled, retrorsely strigose hairy. Leaves in whorls of 4, subsessile or shortly petiolate; blade rather subleathery, pale abaxially, ovate-elliptic, 812 35 mm, strigose to hirsute, abaxially yellowish brown glandular-punctate or striate, base cuneate, apex acute or apiculate; principal vein 1, 2 lateral veins
weak. Inflorescences terminal, corymbiform, with terminal and
axillary several-flowered and somewhat bracteate cymes; pedicels ca. 1.5 mm. Ovary obovoid, hispidulous. Flowers ca. 2.8
mm in diam., probably sexually differentiated (dioecious or
polygamo-dioecious?). Corolla yellowish green, dark brown
when dry, rotate, lobed for 3/4 or more; lobes 4, subovate, apiculate at apex. Fruit unknown, but probably with uncinate trichomes. Fl. JunAug.
Meadows on mountain slopes; 30003200 m. Sichuan (Yajiang),
Yunnan (Lijiang).
Because of its uncertain fruit indumentum, Galium forrestii is in
need of further studies. We have not seen authentic material but agree
with Cufodontis (Oesterr. Bot. Z. 89: 232. 1940) that it is obviously
close (or even identical?) with G. glandulosum and/or G. hirtiflorum.
Their characteristic stem indumentum and other similarities link these
taxa as members of the G. hirtiflorum group within G. sect. Platygalium
s.l. (see under G. hirtiflorum).

19. Galium ghilanicum Stapf, Denkschr. Kaiserl. Akad. Wiss.,

Wien. Math.-Naturwiss. Kl. 50: 53. 1886.
ji lan la la teng
Galium parisiense Linnaeus var. brachypodum Boissier;
G. transcaucasicum Stapf.
Herbs, annual, ascending, branching from base. Stems
(4)830(40) cm tall, tender, 4-angled, with retrorsely aculeolate angles and numerous, rather short internodes. Leaves at
middle stem region in whorls of (5 or)68(or 9), sessile; blade
drying papery, linear-elliptic to narrowly oblanceolate, mostly
glabrous but margins and sometimes abaxial vein sparsely to
densely antrorsely aculeolate, base acute, apex acute-acuminate.
Inflorescences narrowly thyrsoid, with axillary and terminal
cymes mostly 26-flowered; peduncles as long or 24 as long
as subtending leaves, slightly divaricate, with 1 or 2 bracts,
smooth; pedicels thin, 0.54 mm, reflexed and hardly elongated
in fruit. Flowers hermaphroditic. Ovary obovoid to ellipsoid, ca.
0.8 mm, glabrous. Corolla white or greenish white, slightly cupshaped, 0.81.2 mm in diam.; lobes ovate, acute to shortly apiculate. Mericarps subglobose to kidney-shaped, 0.81.5 mm,
colliculate, glabrous.
Open habitats; ca. 700 m. Xinjiang (Yining) [Afghanistan, Nepal,
N Pakistan, Tajikistan; SW Asia].
Galium ghilanicum belongs to the annual G. sect. Microgalium
and is a taxon widespread in SW Asia. It is here recorded for the first
time for China. In FRPS (71(2): 237. 1999) it was misidentified as G.
aparine var. leiospermum (= G. aparine f. leiocarpum, G. spurium),
from which it is clearly separable by its antrorsely (and not retrorsely)
aculeolate leaf margins. The other Chinese representative of G. sect.
Microgalium, G. tenuissimum, differs from G. ghilanicum mainly by its
strongly elongated peduncles and pedicels.

20. Galium glandulosum Handel-Mazzetti, Symb. Sin. 7:

1028. 1936.

125

xian ye la la teng
Herbs, perennial, caespitose, procumbent or erect, 515
cm tall. Stems numerous from reddish rhizomes and roots,
usually strongly branched, 4-angled, densely retrorsely strigose hairy, hispidulous at nodes. Leaves in whorls of 4, sessile or subsessile; blade drying subleathery, quite variable in
shape, ovate to oblong or lanceolate, (2.5)410(14) (0.6)
13(4.5) mm, sometimes with scattered hairs adaxially, on margins and on midrib abaxially, or mostly glabrescent, adaxially
slightly shiny and papillose, abaxially matte and usually minutely glandular-punctate or -striate, base cuneate, margins revolute, apex acute or subobtuse; principal vein 1, lateral veins 2,
weak. Inflorescences terminal and in axils of upper leaves, with
few- to several-flowered and up to 2 cm long cymes; peduncles
hairy, bracteate, divaricate in fruit; pedicels 12(5) mm.
Flowers usually sexually differentiated (dioecious or polygamodioecious?). Ovary obovoid, ca. 0.8 mm, with curved trichomes. Corolla yellowish, greenish, or brownish-reddish,
rotate, 1.82.7 mm in diam.; lobes 4, ovate, obtuse or slightly
acute. Mericarps reniform, 1.52 mm, mostly with uncinate
trichomes of ca. 0.3 mm (very rarely also glabrous?), on
straight or curved, up to 4 mm long pedicels. Fl. JunAug, fr.
AugSep.
Mountain slopes, river beaches, open shrublands and forests,
grasslands, on rocks; 23003900 m. Sichuan, Xizang, Yunnan.
Galium glandulosum is mainly characterized by condensed habit,
short and predominantly retrorse stem hairs, subleathery leaves, often
with glandlike idioblasts on abaxial leaf sides, strongly bracteate
cymes, sexual differentiation of flowers, and hooked trichomes on fruit.
Glabrous-fruited specimens included by W. C. Chen in FRPS (71(2):
228. 1999) may belong to other species. Galium glandulosum and G.
forrestii are members of the G. hirtiflorum group (see there) within G.
sect. Platygalium s.l.

21. Galium hirtiflorum Requien ex Candolle, Prodr. 4: 600.

1830.
mao hua la la teng
Herbs, perennial, weak, procumbent or suberect, 1060 cm
tall. Rhizome and roots red, filiform. Stems numerous, 4angled, with retrorse and/or spreading hairs to glabrescent.
Leaves and leaflike stipules in whorls of 4; blade drying papery
or membranous, linear-elliptic to narrowly lanceolate, (3)8
17(25) (0.5)12.5(3.5) mm, both surfaces hirsute or only
midrib and margins with straight or slightly curved hairs, base
gradually and shortly attenuate, apex obtuse to subacute or
shortly acuminate; vein 1. Inflorescences terminal and axillary,
with several- to many-flowered cymes on peduncles longer than
leaves; pedicels 16 mm. Flowers evidently sexually differentiated (dioecious or polygamo-dioecious?). Ovary obovoid, ca.
0.5 mm, hairy. Corolla light greenish, marked with reddish, rotate, 22.5 mm in diam.; lobes 4, ovate, cuspidate, sparsely hirsute toward apex. Mericarps ellipsoid, 11.5 mm, with dense
uncinate trichomes 0.50.7 mm. Fl. JulAug, fr. SepOct.
Among shrubs and stones; 17003000 m or higher. Expected in
Xizang [Bhutan, India, Nepal].
Galium hirtiflorum was not included in the Chinese flora by W. C.
Chen but is likely to occur in Xizang. According to Ehrendorfer et al.

126

RUBIACEAE

(Fl. Iranica 176: 179. 2005) the group of G. hirtiflorum s.l. includes a
series of vicarious Himalayan taxa: G. subtrinervium Ehrendorfer &
Schnbeck-Temesy in Pakistan (Swat) and Kashmir, G. hirtiflorum s.s.
extending eastward to Bhutan and possibly adjacent China, and finally
G. glandulosum and G. forrestii reaching Yunnan and Sichuan. Their
common group characters are short and retrorse stem hairs, subleathery
leaves with glandlike idioblasts abaxially, strongly bracteate cymes, sexual differentiation of flowers, and fruit with uncinate trichomes.

Together with G. asperuloides, all these taxa belong to G. sect. Hylaea

as shown by Ehrendorfer et al. (loc. cit.: 181185). Galium triflorum is
rare in China and closely related to G. trifloriforme (see under these
species). The latter may be a hybridogenous taxon linking G. sect.
Hylaea and G. sect. Trachygalium. But to synonymize G. trifloriforme
with G. hoffmeisteri (e.g., W. C. Chen in FRPS 71(2): 230. 1999;
Govaerts et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/
rubiaceae/; accessed on 15 Sep 2010) is certainly not correct.

22. Galium hoffmeisteri (Klotzsch) Ehrendorfer & Schnbeck-Temesy ex R. R. Mill, Edinburgh J. Bot. 53: 95. 1996.

Slender plants of Galium hoffmeisteri are distantly reminiscent of

G. kikumugura Ohwi (= G. brachypodum Maximowicz (1874), not
Jordan (1846)) from Japan (see also under G. sichuanense). Yamazaki
(Fl. Japan 3a: 236. 1993) described G. kikumugura as having leaf whorls
of 4. In reality, it always develops whorls of up to 5 or 6 in the middle
stem region. These leaves are quite similar to those of G. hoffmeisteri
and have margins slightly antrorsely aculeolate or smooth. Nevertheless, G. kikumugura strongly deviates from G. hoffmeisteri and other
members of G. sect. Hylaea by its cymes nearly exclusively lateral in
leaf axils, usually consisting only of a filiform peduncle, a single bract,
and 2 small flowers (ca. 1.5 mm in diam.) or sometimes a single flower.
Furthermore, its kidney-shaped (not ellipsoid) mericarps (1.82 0.91
mm) differ by having scattered short and hooked hairs only ca. 0.1 mm.
These latter characters are reminiscent of G. sect. Trachygalium and
particularly of G. bungei (in G. sect. Platygalium s.l., where leaf whorls
have only 4 elements). Thus, G. kikumugura is an isolated and aberrant
species of the genus, possibly better placed into a separate monotypic
section.

liu ye l
Asperula hoffmeisteri Klotzsch, Bot. Ergebn. Reise Waldemar, 87. 1862; Galium asperuloides Edgeworth subsp. hoffmeisteri (Klotzsch) H. Hara; G. asperuloides var. hoffmeisteri
(Klotzsch) Handel-Mazzetti; G. asperulopsis H. J. P. Winkler;
G. japonicum Makino (1895), not (Maximowicz) Makino &
Nakai (1908); G. triflorum Michaux var. hoffmeisteri (Klotzsch)
J. D. Hooker.
Herbs, perennial, from filiform reddish rhizomes. Stems
generally erect, (10)1530(40) cm tall, 4-angled, glabrous
and smooth, sometimes hispidulous at nodes. Middle stem
leaves and leaflike stipules in whorls of up to 6 (in weak
plants rarely only up to 4), with petioles up to 3 mm; blade
drying papery or membranous, narrowly elliptic-oblong to
broadly oblanceolate, (10)1530(40) (4)510(12) mm,
length/breadth index mostly 2.53.5, glabrescent, smooth or
rarely retrorsely aculeolate on abaxial midrib, base acute to
obtuse, margins antrorsely aculeolate, apex obtuse to rounded
and abruptly apiculate; vein 1. Inflorescences terminal and
sometimes in axils of upper leaves, with few- to severalflowered cymes; peduncles glabrous, smooth; bracts none or
few, 12 mm; pedicels 0.33 mm. Ovary obovoid to subglobose, 0.50.8 mm, strigillose with undeveloped trichomes.
Corolla white or light green, rotate, 2.53 mm in diam., glabrescent, lobed for 3/4 or more; lobes 4, ovate, acute. Mericarps ellipsoid, 1.22 mm, with dense uncinate trichomes 0.8
1.2 mm, on pedicels elongating and up to 10 mm. Fl. Apr
Aug, fr. MaySep.
Forests on mountain slopes, thickets, along rivers, ditch sides,
meadows; 4004000 m. Anhui, Gansu, Guizhou, Hebei, Heilongjiang,
Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi, Shanxi, Sichuan, Xizang, Yunnan, Zhejiang [Afghanistan, Bhutan, India, ?Japan, Kashmir,
Korea, Myanmar, Nepal, Pakistan].
Galium hoffmeisteri, together with G. bungei, G. spurium, and G.
verum, is one of the most commonly collected species of Galium in
China. Previous authors usually have included it as a subspecies under
G. asperuloides. Only recently, the two taxa were discussed and reestablished on the species level by Ehrendorfer et al. (Fl. Iranica 176:
183185. 2005) and by Mill (loc. cit.). All specimens seen so far from
China belong to G. hoffmeisteri. Nevertheless, as G. asperuloides s.s.
ranges in the W Himalaya from Afghanistan to N Pakistan and N India,
it can also be expected in SW China (in particular, Xizang). Accordingly, it is included in the present treatment for reference.
In Japan Galium hoffmeisteri is replaced by G. nipponicum
Makino (G. trifloriforme var. nipponicum (Makino) Nakai), but some of
the Japanese specimens greatly approach G. hoffmeisteri. Another very
similar vicariant is G. echinocarpum from Taiwan. Further relatives are
the Eurasian disjunct G. odoratum and the circumboreal G. triflorum.

23. Galium humifusum M. Bieberstein, Fl. Taur.-Caucas. 1:

104. 1808.
man sheng la la teng
Asperula humifusa (M. Bieberstein) Besser.
Herbs, perennial, sometimes slightly woody at base, clambering to procumbent, from a thick rootstock with slender,
trailing reddish rhizomes. Stems up to 1 m tall, 4-angled to
subterete, often caespitose, glabrescent to white pilosulous, hirtellous, and/or pilose often with mixed trichome types, smooth
or sparsely scaberulous. Leaves in whorls of 610, sessile, frequently reflexed; blade drying papery, from linear and narrowly
oblong-oblanceolate to oblong-elliptic or ligulate, (5)1028(
32) (1)1.53(6) mm, adaxially glabrous and scaberulous,
abaxially glabrous to densely white pilosulous or -pilose, base
straight to cuneate, margin antrorsely aculeolate and usually
markedly revolute, apex obtuse to acute and mucronate with tip
to 2 mm; vein 1. Inflorescences with numerous terminal and
axillary, congested to fasciculate, leaflike and many-flowered
cymes; peduncles glabrous to hirtellous and/or pilosulous, with
reduced leaves and leaflike bracts, 1.53 mm; pedicels 14
mm. Ovary ellipsoid, 0.81 mm, glabrous to hispidulous with
straight trichomes. Corolla yellowish white to white, funnelform, 1.52.5 2.53 mm, glabrous to sometimes hairy on outside; lobes 4, ca. 1/2 as long as tube, triangular-ovate, acute to
apiculate. Mericarps ellipsoid to reniform, 11.5 1.52 mm,
glabrous and smooth, granulate or hispidulous, becoming separated in middle as fruit expand but remaining attached at top
and bottom. Fl. and fr. MayOct.
Riversides and beaches, forests, grasslands, farmland sides, wastelands, meadows, mountain slopes; 4002200 m. Xinjiang [Afghanistan,
Kazakhstan, Mongolia, Pakistan, Russia, Turkistan; SW Asia (Armenia,
Azerbaijan, Georgia, Iran, Iraq), E Europe (Balkan Peninsula, Ukraine)].

RUBIACEAE

This species has often been included in Asperula (e.g., Pobedimova et al., Fl. URSS 23: 276. 1958) because of its funnelform, relatively large, white corollas, but its affinities are clearly with members of
Galium, particularly G. verum, though the flowers are distinct. Rarely
the two species form a hybrid, which has been called G. himmelbauerianum (Ronniger) So, and both should be placed into G. sect. Galium.
Galium humifusum is a widespread diploid species, very variable
due to modificational plasticity and genetic diversity (Ehrendorfer et
al., Fl. Iranica 176: 197. 2005), but at present it does not appear possible
to recognize infraspecific taxa.

24. Galium hupehense Pampanini, Nuovo Giorn. Bot. Ital. 17:

719. 1910.
hu bei la la teng
Galium boreale Linnaeus var. molle Hemsley; G. hemsleyanum Beauverd; G. hupehense var. molle (Hemsley) Cufodontis.
Herbs, perennial, erect, with slender rhizome. Stems 4angled, pilosulous. Leaves in whorls of 4, subsessile; blade lanceolate, 3050 612 mm, length/breadth index above 4, adaxially hispidulous to scaberulous, abaxially pilose at least on
principal veins, apex acuminate to subacute; principal veins 3,
palmate. Inflorescence terminal, paniculiform, 1520 48 cm,
with many-flowered cymes; peduncles and pedicels hairy to
glabrescent. Ovary ovoid, densely hairy. Corolla yellowish
white, rotate, ca. 2 mm in diam., glabrous or pilose; lobes 4,
ovate, hairy outside, acute. Mericarps with straight trichomes
(and/or ?glabrous). Fl. Jul, fr. AugSep.
Mountains; ca. 2000 m. Hubei (Yichang), Jiangsu (Kunshan).
Galium hupehense, possibly endemic to EC China, is evidently
related to the also small-flowered G. kinuta (Cufodontis, Oesterr. Bot. Z.
89: 223224. 1940) but has hairy stems, whereas G. chekiangense and
G. boreale differ i.a. by their larger flowers (34 mm in diam.). As we
have seen no authentic material of G. hupehense, the above description
is based on available literature sources only. Its original description
gives no information on ovary and fruit indumentum, whereas straight
(and ?multicellular) hairs are indicated for the certainly synonymous G.
hemsleyanum. Possibly by mistake, W. C. Chen (in FRPS 71(2): 281
282. 1999) reported plants with glabrous (var. hupehense) and with
densely hairy ovaries (var. molle). Both are recorded from Yichang,
only the latter from more condensed plants in Kunshan. These uncertainties and the status of G. hupehense with its varieties need to be clarified in the future.

25. Galium innocuum Miquel, Fl. Ned. Ind. 2: 341. 1857.

xiao zhu yang yang
Galium modestum Diels; G. trifidum Linnaeus var. modestum (Diels) Cufodontis.
Herbs, perennial, weak to procumbent, from slender rhizomes. Stems (7)1040(60) cm, 4-angled, caespitose, glabrous and smooth to sparsely retrorsely aculeolate on angles.
Leaves in whorls of 4(6), subsessile; blade drying papery,
blackish or green, linear-lanceolate to oblanceolate, 38(10)
12 mm, glabrous and smooth to sparsely retrorsely aculeolate
on margins and midrib, base acute to attenuate, apex rounded or
obtuse; vein 1. Inflorescences terminal and axillary, cymes 1

127

3.5 cm, with 13(or 4) flowers; peduncles glabrous and smooth;

bracts oblanceolate to narrowly elliptic; fruiting pedicels (3)5
8(10) mm, straight and divaricate. Ovary didymous, glabrous, smooth. Corolla white, cup-shaped to slightly campanulate, 11.8 mm in diam.; lobed to 1/2 or slightly more; lobes
3(or 4), ovate and rounded at tip. Fruit markedly didymous,
mericarps (sub)globose, 22.8 mm, glabrous, smooth to slightly
tuberculate. Fl. and fr. MarAug.
Swampy or wet localities at lower to upper montane elevations.
Fujian, Sichuan, Taiwan, Yunnan, and ?elsewhere [India, Indochina,
Indonesia (Java, Sumatra), New Guinea].
In the available floras of China and Taiwan, W. C. Chen (in FRPS
71(2): 252253. 1999) and Yang and Li (Bull. Natl. Mus. Nat. Sci.,
Taichung 11: 101117. 1998; Fl. Taiwan, ed. 2, 4: 254259. 1998) have
completely ignored Galium innocuum, classifying most of the relevant
specimens under G. trifidum. In the Kew Rubiaceae checklist (Govaerts
et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/;
accessed on 15 Sep 2010) G. trifidum var. modestum appears as a
synonym under G. innocuum, with a range from India to China and
through Taiwan to SE Asia and New Guinea. Originally, G. innocuum
was described from Java. In the critical revision of G. sect. Aparinoides
by Puff (Canad. J. Bot. 54: 19111925. 1976), not considered by the
above authors, G. innocuum is accepted as a valid species and regarded
as a southern member of the G. trifidum group. The above diagnosis and
distribution data correspond to Puffs revision. He differentiated the two
species mainly by their fruiting pedicels: relatively short, straight, and
divaricate in G. innocuum but slender, elongated, and conspicuously
arcuate in G. trifidum s.s. According to Puff (loc. cit.: 19221923) only
G. innocuum but none of the subspecies of G. trifidum occur in China.
This is in strong conflict with W. C. Chen (loc. cit.: 253) who described
the distribution of G. trifidum in China by listing the provinces Anhui,
Fujian, Guangdong, Guangxi, Guizhou, Hebei, Heilongjiang, Hunan,
Jiangsu, Jiangxi, Jilin, Liaoning, Nei Mongol, Shanxi, Sichuan, Taiwan,
Xizang, Yunnan, and Zhejiang. As we have seen only limited Chinese
material of G. sect. Aparinoides, the question remains whether G.
innocuum extends from S to N China or is replaced further north by
populations of the G. trifidum group not mentioned by Puff. Furthermore, one has to consider that species from other sections of Galium
have often been misidentified as members of G. sect. Aparinoides, e.g.,
G. bungei. In view of these uncertainties, we accept only G. innocuum
but not G. trifidum for the present Chinese flora.
Another problematic taxon in Galium sect. Aparinoides for the
Flora of China is G. palustre Linnaeus. In spite of the critical comments
by Cufodontis (Oesterr. Bot. Z. 89: 232. 1940), this species has been
included in FRPS by W. C. Chen (loc. cit.: 250). According to Puff (loc.
cit.: 19231924) this species belongs to the G. palustre group of taxa,
with leaves often in whorls of more than 4, many-flowered cymes
(more than 3 or 4 flowers), and smooth pedicels. Its natural distribution
is verified from temperate and boreal Europe to W Siberia, whereas
occurrences in E North America (and elsewhere) are obviously adventive. Considering the common confusion of G. sect. Aparinoides taxa (in
China particularly G. innocuum in the south and G. karakulense in the
north) and the lack of authentic specimens seen by us, we exclude G.
palustre from China in the present text.

26. Galium kamtschaticum Steller ex Schultes & J. H. Schultes, Mant. 3: 186. 1827.
san mai zhu yang yang
Herbs, perennial, erect, 525 cm tall, emerging from filiform rhizomes. Stems mostly unbranched, 4-angled, glabrous to
sparsely hispidulous. Leaves in whorls of 4, sessile or subses-

128

RUBIACEAE

sile; blade drying blackish brown and papery, broadly elliptic,

ovate, or suborbicular, 1025 617 mm, glabrous to hispidulous at least on veins, otherwise adaxially smooth, abaxially not
glandular-striate, base cuneate to obtuse, margins antrorsely
ciliolate or hispidulous, apex rounded and usually mucronate; principal veins 3, palmate. Inflorescences small, thyrsoid,
cymes terminal and in axils of uppermost leaves, 26 cm, few
to several flowered, lax; peduncles glabrous, smooth; bracts
leaflike or ligulate to narrowly elliptic, 0.54 mm; pedicels 15
mm. Ovary subglobose, ca. 1 mm, densely pubescent with
spreading uncinate trichomes. Corolla white or greenish yellow,
rotate, 2.53(4) mm in diam., glabrous, lobed for 3/4 or more;
lobes 4, elliptic-lanceolate or ovate-triangular, acute. Mericarps
ovoid, 1.52 mm, with dense uncinate trichomes 0.81 mm, on
pedicels usually elongating to 15 mm. Fl. and fr. JulSep.
Forests on mountains, tussocks at ditch sides; 15002300 m [as
low as ca. 100 m in N Japan]. Heilongjiang, Jilin [Japan, Korea, NE
Russia; NW North America].
Galium kamtschaticum is an amphi-Beringian member of G. sect.
Platygalium and forms a related species group with G. chekiangense in
SE China, G. nakaii in Japan, and G. oreganum in W North America. In
the Flora of Japan (Yamazaki, Fl. Japan 3a: 206240. 1993) three varieties are recognized, one of them endemic to Japan, the second extending to Sakhalin and the Kuriles, and only var. kamtschaticum more
widespread and extending to the NE provinces of China. Specimens
from SE China, Zhejiang, that were determined as G. kamtschaticum
and those from Fujian that were listed by W. C. Chen (in FRPS 71(2):
265. 1999) as G. nakaii differ from typical G. kamtschaticum by their
leaves drying somewhat leathery, greenish-brownish, papillose, glandular-striate abaxially, and by their fruit with short hairs 0.10.2 mm, with
a bent but not hooked tip. In the present flora they are treated as a new
species, G. chekiangense (see the comments under that species).

27. Galium karakulense Pobedimova in Schischkin, Fl. URSS

23: 712. 1958.
cu zhao la la teng
Herbs, perennial, weak to procumbent, 4070 cm tall.
Rootstock slender, with prolonged rhizomes. Stems 4-angled,
flaccid, much branched, retrorsely aculeolate, hispidulous or
glabrescent at nodes. Leaves in whorls of 4(or 5), subpetiolate;
blade drying papery, elliptic to oblanceolate, (12)1520(30)
(2)58(12) mm, densely antrorsely aculeolate adaxially and
along margins, retrorsely aculeolate on abaxial midrib, base
attenuate to cuneate, margins thinly revolute, apex rounded to
bluntly pointed; vein 1. Inflorescences paniculate, terminal and
lateral cymes with several to many flowers (usually more than
4); peduncles scabrous and divaricate; bracts ovate to elliptic,
37 13 mm; pedicels 24.5 mm, rough, elongated in fruit.
Ovary didymous, glabrous. Corolla white, cup-shaped, 3
4(4.5) mm in diam., 4-lobed to ca. 1/2. Mericarps (sub)globose, 1.52.5(3.5) mm, glabrous, slightly verrucose. Fl. Jul, fr.
AugSep.
Swamps and riversides at low to middle elevations. Xinjiang
(Chabuchaer) [Afghanistan, Kazakhstan, Kyrgyzstan, Russia, Tajikistan,
Uzbekistan].
Galium karakulense represents G. sect. Aparinoides in C Asia and
is considered as a link between the G. trifidum group in the N
Hemisphere and the predominantly Mediterranean-European G. palustre

group (Puff, Canad. J. Bot. 54: 19231924. 1976). It is also treated in

Pobedimova et al. (loc. cit.) and Ehrendorfer et al. (Fl. Iranica 176: 174.
2005) and may be responsible for some of the erroneous indications of
G. palustre for N China, as discussed under G. innocuum. The latter is
the second verified species of the section in the more southerly part of
the Chinese flora. In comparison with G. karakulense, G. innocuum is a
much smaller and more slender plant with few-flowered cymes and
smooth pedicels.

28. Galium karataviense (Pavlov) Pobedimova, Novosti Sist.

Vyssh. Rast. 7: 278. 1971.
ka la tao la la teng
Asperula karataviensis Pavlov, Vestn. Akad. Nauk Kazakhsk. S.S.R. 4: 95. 1951; A. aparine M. Bieberstein, s.l.; A.
rivalis Sibthorp & Smith, s.l.; Galium rivale (Sibthorp &
Smith) Grisebach, s.l.
Herbs, perennial, procumbent and often matted or lodged,
from slender, reddish brown rhizomes. Stems 0.61.2 m, much
branched, densely retrorsely aculeolate on 4 angles. Leaves in
whorls of 610, sessile or subsessile; blade drying papery or
somewhat leathery and glossy, narrowly (ob)lanceolate or
narrowly elliptic, (6)1525(50) (2)2.54(8) mm, glabrescent, both surfaces sparsely to densely aculeolate on midrib,
base acute to cuneate, margins flat to narrowly revolute, densely
retrorsely aculeolate, apex acute and shortly mucronate; vein 1.
Inflorescences terminal and axillary, to 12 9 cm, with severalto many-flowered cymes; peduncles elongating as inflorescences develop, becoming much longer than subtending leaves;
axes glabrescent, sparsely to densely retrorsely aculeolate;
bracts elliptic or oblong-lanceolate, 1.53 0.51 mm; pedicels
13 mm. Ovary ellipsoid to obovoid, 0.50.8 mm, glabrous.
Corolla bluish to violet (rarely white?), shortly funnelform, 1.5
2.5 mm, tube 11.5 as long as lobes; lobes 4, triangular-spatulate. Mericarps subglobose to ellipsoid, 1.52 1.72 mm,
glabrous, smooth or often tuberculate. Fl. and fr. JunSep.
Humid forests, riversides, beaches, wet grasslands; 7003300 m.
Gansu, Hebei, Heilongjiang, Nei Mongol, Ningxia, Qinghai, Shanxi,
Sichuan, Xinjiang [C Asia].
Galium karataviense was treated in FRPS (71(2): 280. 1999) as G.
rivale. In a wider sense the latter name and its synonyms apply to a
polymorphic polyploid complex (2x, 4x, 6x), ranging from NE, E, and
SE Europe to SW and C Asia. Because of its funnelform corollas this
group formerly was treated as part of the genus Asperula, either as A.
aparine or as A. rivalis (e.g., in Pobedimova et al., Fl. URSS 23: 275.
1958). More recently, morphological analyses (see Ehrendorfer et al.,
Fl. Iranica 176: 188. 2005) and DNA-data have clearly shown that it is
closely related to G. uliginosum in G. sect. Trachygalium. Similarities
with the annual G. sect. Euaparine are homoplasies.
In former treatments (e.g., Fl. Europaea 4: 20. 1976) Galium
rivale was circumscribed in a wide sense. On the basis of differences in
floral (relative length of corolla tube and lobes, color), fruit (mericarp
epidermis with rounded or acute cells), and other characters, several still
insufficiently understood microspecies have been suggested (Pobedimova et al., loc. cit.: 327, under Asperula; Ehrendorfer & Schnbeck,
Pl. Syst. Evol. 174: 200202. 1991, under G. anguineum; Ehrendorfer et
al., loc. cit., under G. pseudorivale Tzvelev). Accordingly, among the
vicarious microspecies of G. rivale s.l., G. anguineum Ehrendorfer &
Schnbeck-Temesy from Iraq and Iran, with white corollas and divaricate fruiting axes, is replaced toward the east in C Asia by G. karatav-

RUBIACEAE

iense, with bluish to violet corollas and more convergent fruiting axes.
Further studies will have to demonstrate whether species status is really
justified for all these taxa and how they correspond to the different
cytotypes encountered in this polyploid complex.

29. Galium kinuta Nakai & H. Hara, J. Jap. Bot. 9: 518. 1933.
xian mai la la teng
Galium boreale Linnaeus var. japonicum Maximowicz; G.
japonicum (Maximowicz) Makino & Nakai (1908), not Makino
(1895).
Herbs, perennial, erect, 2060 cm tall. Stems with 4 thickened angles, glabrous and smooth, hispidulous only at nodes.
Leaves in whorls of 4, subsessile or petiole to 2 mm; blade
drying mostly somewhat leathery, remaining green, oblanceolate to ovate-lanceolate, sometimes even narrowly elliptic or
ovate, 2080 420 mm, length/breadth index (2)35(6),
strigillose or hispidulous at least along veins to glabrescent,
sparsely to densely punctate- to striate-glandular abaxially, base
acute to rounded, margins flat to thinly revolute, antrorsely
ciliolate to hispidulous, apex subacute to acute, but hardly concave and long acuminate; principal veins 3, palmate. Inflorescences paniculiform, to 25 15 cm, cymes in uppermost leaf
axils and terminal, many flowered, lax and often somewhat
divaricate; peduncles smooth and glabrous or hispidulous at
nodes; bracts oblanceolate to narrowly elliptic, 1.53 mm; pedicels 1.53 mm. Ovary subglobose to obovoid, ca. 0.8 mm,
smooth, glabrous. Corolla white to purplish, rotate, 22.5 mm
in diam., glabrous; lobes 4, ovate, acuminate. Mericarps subglobose to obovoid, ca. 2.5 mm, glabrous and smooth. Fl.
MayJul, fr. AugSep.
Mesic, generally rich forests on mountain slopes, rocks at watersides, open grasslands, meadows; 5002100 m. Gansu, Hebei, Henan,
Hubei, Liaoning, Shaanxi, Shanxi, Sichuan, Xinjiang [Japan, Korea].
Galium kinuta, described from Japan, was first reported for China
by Cufodontis (Oesterr. Bot. Z. 89: 223224. 1940). Nevertheless, the
more numerous samples now available suggest certain differences: in
Chinese specimens the leaves are more leathery (not paperlike) when
dry, abaxially more markedly punctate-striate-glandular (not inconspicuously so), and apically acute (but hardly concave and long attenuate).
It is still uncertain whether these differences merit taxonomic separation
of the Japanese and Chinese populations. Another critical aspect is the
occurrence of specimens intermediate between Chinese G. kinuta and
local G. boreale s.l., as observed, e.g., from Henan, Shaanxi, and Shanxi. Local studies will have to show whether this is due to hybridization
and whether it is linked to the extreme variability of G. kinuta in leaf
shape, ranging from narrowly lanceolate to ovate.
Galium kinuta may be related to the still badly understood G.
hupehense (see there). Similarities also exist with G. platygalium, which
differs by funnelform corollas. Galium hoffmeisteri (= G. japonicum
Makino (1895)) and G. kinuta (= G. japonicum (Maximowicz) Makino
& Nakai (1908)) have been widely confused because of similar habit
and name confusion.

30. Galium kunmingense Ehrendorfer, Novon 20: 270. 2010.

kun ming la la teng
Herbs, perennial, erect, (12)1520(25) cm tall. Stems
with 4 prominent and rounded angles, with scattered antrorsely
(or sometimes also retrorsely) curved short hairs, more dense at

129

nodes; internodes 1.54 cm at middle stem regions, longer or

somewhat shorter than leaves. Leaves in whorls of 4; blade
drying leathery, broadly lanceolate, (10)1525(35) (4.5)6
7.5(9) mm, length/breadth index (2)2.53.5(4), glabrous,
adaxially papillose, abaxially without glandular idioblasts, base
cuneate, margins revolute, antrorsely aculeolate, apex acute but
not acuminate; 3 prominent principal veins extending into apex
region. Inflorescences pyramidal, with cymes from middle to
upper stem nodes and terminal, several to many flowered; axes
glabrous, somewhat divaricate; peduncles mostly 1.54 cm;
pedicels (0.5)15 mm; bracts lanceolate, small and inconspicuous. Flowers hermaphroditic. Ovary obovoid, ca. 0.5 mm, glabrous, smooth. Corolla white to greenish, cup-shaped or campanulate, (2)2.32.5(2.7) mm in diam., fused at base for
length of 4 free lobes, 0.81 mm, lanceolate, acute but not apiculate. Mericarps ovoid, 1.53 mm, glabrous and smooth. Fl.
JunAug, fr. JulSep.
Open grasslands and rocky slopes; 19002500 m. C Yunnan.
The new Galium kunmingense clearly belongs to G. sect. Platygalium. Among species in the section with corollas basally fused to ca.
1/2 the length Microphysa elongata deviates by leaves with only one
main vein, somewhat inflated fruit mericarp, and funnelform corollas.
Galium platygalium and G. maximoviczii have similar corollas but
deviate by their broader and 35-veined leaves. Closer relationships can
be assumed for some E Asiatic Galium species with rotate corollas: G.
kinuta has glabrous fruit but narrower, adaxially punctate-striate glandular leaves, slender, more floriferous inflorescences, and larger flowers. Apparent relatives with rotate and smaller flowers are G. hupehense
with spreading straight fruit hairs, G. chekiangense with appressed
curved fruit hairs, and particularly G. yunnanense, with spreading uncinate fruit hairs. This latter is sympatric with G. kunmingense but differs not only by its rotate (not campanulate/cup-shaped) and smaller corollas and uncinate fruit hairs, but also by its more hairy and abaxially
punctate-striate glandular leaves. It is remarkable that G. kunmingense, a
quite conspicuous species that evidently was not too rare in the surroundings of the capital of Yunnan, has remained unnoticed up to now.

31. Galium linearifolium Turczaninow, Bull. Soc. Imp. Naturalistes Moscou 7: 152. 1837.
xian ye la la teng
Herbs, perennial, erect, sometimes slightly woody at base.
Stems up to 65 cm tall, 4-angled, hirtellous or puberulent to
glabrescent or smooth and glabrous. Leaves in whorls of 4, sessile or subsessile; blade drying leathery, linear-spatulate, often
slightly falcate, 1060 14 mm, adaxially glabrous, weakly
shiny, antrorsely aculeolate along midrib and/or near margins,
abaxially glabrous or sparsely hirtellous along midrib, base
cuneate or obtuse, margin antrorsely aculeolate or pubescent,
revolute, apex obtuse to acute; vein 1. Inflorescences terminal, paniculiform, with few- to many-flowered, 1.55 cm long
cymes; peduncles hirtellous to glabrous, smooth; bracts narrowly elliptic, 13 mm or often lacking; pedicels 1.56 mm.
Ovary ellipsoid to obovoid, ca. 0.8 mm, glabrous, smooth. Corolla white, rotate, ca. 4 mm in diam.; lobes 4, lanceolate, acute.
Mericarps ellipsoid to subglobose, 2.53 mm, glabrous and
smooth. Fl. JunAug, fr. JulSep.
Grassy slopes, forests, thickets, mountain meadows; 4001800 m.
Hebei, Hubei, Liaoning [Korea].

130

RUBIACEAE

Galium linearifolium seems to be a rare species and is easily confused with G. boreale. The latter has leaves usually somewhat lanceolate
and with 3 main veins. Cufodontis (Oesterr. Bot. Z. 89: 219223. 1940)
mentioned a certain affinity of G. linearifolium with G. bungei. Although
G. linearifolium is said in FRPS to have leaves in whorls of 4, the
relevant figure (71(2): 251, t. 56, f. 3. 1999) shows them in whorls of 5,
evidently a mistake.

32. Galium maximoviczii (Komarov) Pobedimova, Novosti

Sist. Vyssh. Rast. 7: 277. 1971 [maximowiczii].
yi ye lun cao
Asperula maximoviczii Komarov, Trudy Glavn. Bot. Sada
39: 109. 1923, based on A. platygalium Maximowicz var. pratensis Maximowicz, Bull. Acad. Imp. Sci. Saint-Ptersbourg 19:
284. 1874.
Herbs, perennial, from thin creeping rhizomes. Stems
erect, 0.31 m tall, 4-angled, smooth; nodes hispidulous or glabrous. Leaves in whorls of 46(8), subsessile or with petiole
up to 6 mm; blade drying papery, lanceolate-oblong, lanceolateelliptic, or ovate to ovate-lanceolate, (23)3540(53) (7)9
10(18) mm, glabrous or sparsely to moderately hispidulous, at
least on principal veins, margins antrorsely ciliolate or -aculeolate, base acute to cuneate, apex tapered and shortly obtuse to
acute; veins 35, palmate. Inflorescences broadly paniculate, 4
20 215 cm, lax and many flowered, with cymes in axils of
uppermost leaves and terminal; peduncles glabrous; bracts
linear to narrowly elliptic, 15 mm; pedicels 24 mm. Ovary
obovoid and laterally flattened, ca. 0.8 mm, glabrous, smooth.
Corolla white, campanulate; tube as long as lobes, 2.53.5
mm in diam.; lobes 4, ovate-oblong, obtuse. Mericarps ellipsoid, 22.5 mm, glabrous, smooth to granular-papillose. Fl.
JunJul, fr. JulOct.
Forests, thickets, or grasslands on mountains, open fields, ditch
sides; 16003800 m. Anhui, Hebei, Heilongjiang, Henan, Jilin, Liaoning, Nei Mongol, Shaanxi, Shandong, Shanxi, Zhejiang [Korea,
Russia].
Galium maximoviczii differs from the quite close G. platygalium
(see there) by somewhat smaller campanulate flowers and larger leaves
in whorls of up to 6(8). Within G. sect. Platygalium the two species
form an isolated group limited to the temperate E Asiatic mainland,
characterized by its whorls of leaves and leaflike stipules often with
more than 4, up to 6, or rarely even 8 elements, with 35 palmate principal veins, and campanulate to funnelform corollas. In Pobedimova et
al. (Fl. URSS 23: 273. 1958) the two taxa were placed in the artificial
Asperula sect. Galioideae Pobedimova ser. Paniculatae Pobedimova.

33. Galium megacyttarion R. R. Mill, Edinburgh J. Bot. 53:

200. 1996.
da bao la la teng
Herbs, perennial, weak to procumbent. Stems 640 cm,
sharply 4-angled, glabrous, smooth, rough or sparsely retrorsely
aculeolate; internodes 6.533 mm; nodes hairy. Middle stem
leaves in whorls of up to 6, sessile; blade drying papery, often
blackening, linear-oblanceolate to narrowly elliptic, 212.5
0.42 mm, glabrous, smooth or sparsely retrorsely aculeolate on
midrib abaxially, adaxially with relatively large epidermal cells
(use 20 lens), base acute, margins flat to thinly revolute, apex
acute then contracted and mucronate; vein 1. Inflorescences
axillary, with 1- or occasionally 3-flowered cymes; pedicels

0.21.5 mm, glabrous, smooth. Ovary ellipsoid-obovoid, ca. 0.5

mm, glabrous, smooth. Corolla white or pale green (perhaps
sometimes drying pink), rotate, 1.52.7 mm in diam.; lobes 4,
lanceolate-spatulate, glabrous beneath, puberulent above, with
shortly acuminate apex, clearly longer than stamens. Mericarps
ellipsoid, 0.71.1 11.5 mm, glabrous, granular-verruculose,
with pedicels often elongating to 3.5 mm. Fl. and fr. JulSep.
Open places, forests; 18003100 m. ?Sichuan, Xizang [Bhutan,
India, Nepal].
Galium megacyttarion (type from Uttar Pradesh, Raizada 7326,
E) belongs to the high elevation Himalayan subgroup (2) of the G.
asperifolium group, which includes G. acutum (see additional comments
under these species). The protologue of G. megacyttarion describes the
flowers as having stamens shorter than the corolla; comparable information is not yet available for the majority of the Chinese Galium species.
We have seen no material cited in the original description nor plants
which undoubtedly belong here. Nevertheless, one very condensed
provenance (Duthie 7492, from Bhutan, the Black Mountain Expedition
1888, WU) exhibits the large leaf epidermal cells described for G.
megacyttarion; but it deviates by having antrorse microhairs on the
adaxial leaf side and glabrous leaf margins, finely rough stems, manyflowered cymes, and glabrous petals. It was determined by Cufodontis
(Oesterr. Bot. Z. 89: 241243. 1940) erroneously as G. asperifolium var.
sikkimense. Another plant with large epidermal cells has been seen from
Sichuan (W. C. Chen, 23 Jun 1988, PE), but this corresponds in all other
characters to G. pusillosetosum.

34. Galium minutissimum T. Shimizu, J. Fac. Textile Sci.

Technol. Shinsu Univ., A, 36(12): 58. 1963.
wei xiao la la teng
Herbs, perennial (not annual), ascending, caespitose, minute, 23 cm tall. Stems 4-angled, branched, glabrous or sometimes hispidulous at nodes. Leaves in whorls of 4; blade rhombic-oblong, 23 0.81 mm, glabrous or abaxially hispidulous
along midrib, base attenuate, apex obtuse; vein 1. Inflorescence
with terminal and partly axillary few-flowered cymes, glabrous
axes, and ca. 2 mm long pedicels. Flowers unknown. Mericarps reniform, sparsely hispid with apically weakly curved
trichomes.
Mountains; 18002400 m. Taiwan (Hualian).
Galium minutissimum was accepted as a species by W. C. Chen
(in FRPS 71(2): 283. 1999) but was not treated or mentioned by Yang
and Li in their publication on Galium in Taiwan (Bull. Natl. Mus. Nat.
Sci., Taichung 11: 101117. 1998) or in the subsequent second edition
of Fl. Taiwan (4: 254259. 1998). We have seen no authentic material,
and the specific status of G. minutissimum remains uncertain. The taxon
obviously belongs to G. sect. Platygalium and the closely related G.
morii group (see there) from the high mountains of Taiwan.

35. Galium morii Hayata, Icon. Pl. Formosan. 7: 32. 1918.

sen shi zhu yang yang
Galium sigeyosii Masamune.
Herbs, perennial, erect, 510 cm tall. Stems slender, 4angled, glabrous. Leaves in whorls of 4, sessile or subsessile;
blade drying papery, obovate, ovate, elliptic, or elliptic-oblong,
16 1.510 mm, glabrous or sparsely hairy abaxially, base
obtuse, margins smooth, apex obtuse or apiculate-acute; principal veins 3, palmate. Inflorescences terminal or sometimes axil-

RUBIACEAE

lary, with few-flowered cymes of 0.51.5 cm; peduncles and

bracts glabrous; pedicels 12 mm. Ovary densely strigillose
with undeveloped trichomes. Corolla ?white, rotate, ca. 1.2 mm
in diam., lobed for 3/4 or more; lobes 4, ovate. Mericarps subglobose, ca. 1 mm, with dense, appressed uncinate trichomes.
Mountains; 25003400 m. Taiwan (Jiayi).
Galium morii was described as a very small plant from Yu Shan
(Mt. Morrison) in Taiwan. We have seen no authentic material. The
present description combines information from the protologue, FRPS
(71(2): 241. 1999), and Yang and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: 106107. 1998; Fl. Taiwan, ed. 2, 4: 256. 1998). But there are
certain conflicts: whereas the leaves were characterized by FRPS as 1or indistinctly 3-veined, the protologue and Yang and Li said they were
3-veined.
Galium morii was the first species to be described from an obviously closely related assembly of G. sect. Platygalium taxa growing in
the high mountains of Taiwan, which is here called the G. morii group
and also includes G. formosense, G. minutissimum, G. nankotaizanum,
and G. tarokoense. On the mainland, the newly described G. rupifragum
from Yunnan obviously also belongs here. This G. morii group is characterized by low and condensed growth, small ovate to elliptic or
broadly lanceolate leaves, reduced inflorescences, and hairy fruit.
Affinities obviously exist with the aggregates of G. elegans and G.
serpylloides.
Characters used to differentiate the taxa of the Galium morii group
are stem indumentum, number of leaf veins (13), uncinate to straight
fruit hairs, etc., but many taxonomical problems remain. As an example:
G. morii may not be clearly separable from G. tarokoense by its smaller
and sparsely pubescent, 3-nerved leaves (vs. larger, completely glabrous, and rather 1-nerved leaves of G. tarokoense); the two taxa are
geographically separated according to Yang and Li (loc. cit. 1998: 107,
110). Furthermore, relationships between the alpine core group (as G.
morii) and other taxa with taller growth from lower elevations (as G.
formosense) evidently need more attention in the future. Transitional
states in the above characters make it necessary to place several species
in two or three different positions in our key.

36. Galium nankotaizanum Ohwi, Repert. Spec. Nov. Regni

Veg. 36: 56. 1934.
nan hu da shan zhu yang yang
Galium maborasense Masamune.
Herbs, perennial, erect, often reduced, 512(20) cm tall.
Stems 4-angled, sparsely to moderately pilose becoming glabrescent, angles thickened. Leaves in whorls of 4, sessile to
subsessile; blade drying membranous, ovate, elliptic, or broadly
lanceolate, 410 25 mm, length/breadth index 23, both surfaces sparsely hirsute at least along midrib, base cuneate to obtuse, apex acute to obtuse, sometimes apiculate; principal vein
1, 2 lateral veins weakly developed. Inflorescences terminal and
in upper leaf axils, with cymes 2- to several flowered, 216
mm, shorter to somewhat longer than subtending leaves; peduncles glabrescent, bracteate; pedicels 0.84 mm. Flowers hermaphroditic. Ovary obovoid, ca. 0.5 mm, densely appressed
hairy. Corolla white, rotate, 22.5 mm in diam., lobed for 1/2
2/3; lobes 4, ovate. Fruit on pedicels becoming deflexed to nodding; mericarps obloid to oblate, 0.81 mm, densely grayish
yellow hairy with straight or slightly curved trichomes ca. 0.8
mm. Fl. Jul, fr. AugSep.

131

Under shrubs and in rock crevices; 30003500 m. Taiwan

(Hualian, Nantou, Yilan).
Galium nankotaizanum is closely related to a group of species
described from the high mountains of Taiwan (see under G. morii).
Galium maborasense, treated as a separate but dubious species by W. C.
Chen (in FRPS 71(2): 284. 1999), was explicitly synonymized by Yang
and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: 107. 1998) with G.
nankotaizanum. This publication apparently was not available to Chen,
but here we follow these Taiwanese authors.
Galium nankotaizanum is mainly characterized by its straight or
slightly curved fruit trichomes. This links it to the high-alpine Himalayan group of G. serpylloides s.l. (see there). Otherwise, G. nankotaizanum
also appears similar to taxa with hooked fruit trichomes, e.g., G. tarokoense, with glabrous stems and only 1 principal leaf vein, from which it
apparently differs by larger flowers.

37. Galium odoratum (Linnaeus) Scopoli, Fl. Carniol., ed. 2,

1: 105. 1771.
che zhou cao
Asperula odorata Linnaeus, Sp. Pl. 1: 103. 1753.
Herbs, perennial, from slender and elongated rhizomes.
Stems erect, 1050 cm tall, 4-angled, glabrous and smooth
except hispidulous at nodes. Leaves in whorls of 610, sessile
or petiole to 1 mm; blade drying papery, oblanceolate, oblonglanceolate, or narrowly elliptic, (6)1550(65) (3)4.5
15(17) mm, length/breadth index ca. 4, glabrous except antrorsely aculeolate on margins and with antrorse microhairs on
upper side and sometimes on abaxial midrib, base acute to cuneate, margins flat, apex acute or usually obtuse then abruptly
mucronate; vein 1. Inflorescences terminal, with several- to
many-flowered cymes; axes glabrous, smooth; bracts none or
leaflike, 13 mm; pedicels 14 mm. Ovary ellipsoid to obovoid,
ca. 0.8 mm, densely hispidulous. Corolla white or bluish white,
broadly funnelform, 4.56.5 37 mm, glabrous, lobed for
ca. 1/2; lobes 4, triangular-spatulate, acute. Mericarps subglobose, 22.5 mm, with dense uncinate trichomes 11.2 mm. Fl.
and fr. JunSep.
Mountain forests; 15002800 m. Gansu, Heilongjiang, Jilin, Liaoning, Ningxia, Qinghai, Shandong, Shanxi, Sichuan, Xinjiang [Japan,
Korea, Russia; NW Africa, SW Asia, Europe; introduced in North
America].
Dried plants of Galium odoratum have a sweet coumarine odor,
which is still evident on herbarium specimens; its name refers to this.
On account of its funnelform corollas (with the tube as long as the
lobes), G. odoratum usually has been treated as a member of Asperula.
As shown by Ehrendorfer et al. (Fl. Iranica 176: 183. 2005) and verified
by DNA-analytical studies, it belongs to G. sect. Hylaea and is closely
related to G. asperuloides and G. hoffmeisteri. Without flowers it is difficult to separate, particularly from the former with more narrow leaves,
but generally G. odoratum is more robust. As a constant element of
temperate deciduous forests (often with Fagus), it has an extensive but
rather disjunct distribution area throughout Eurasia, with diploid cytotypes in E Asia, replaced by tetraploids in Europe.

38. Galium paniculatum (Bunge) Pobedimova, Novosti Sist.

Vyssh. Rast. 7: 277. 1971.
yuan zhui la la teng
Asperula paniculata Bunge in Ledebour, Fl. Altaic. 1: 140.
1829; Galium xinjiangense W. C. Chen.

RUBIACEAE

132

Herbs, perennial, often somewhat caespitose from elongated, much branched, and ca. 1 mm thick rhizomes. Stems
erect, to 60 cm tall, 4-angled, little branched, glabrous and
smooth, only sometimes puberulent at nodes. Leaves in whorls
of up to 6, subsessile; blade drying papery, discolorous (more
pale abaxially), lanceolate or narrowly lanceolate, (15)25
60(70) (3)510(12) mm, glabrous, smooth or mostly
somewhat antrorsely ciliolate on margins and midrib, base
acute to cuneate, apex acute to acuminate; vein 1. Inflorescences terminal, 816 816 cm, corymbiform to paniculate,
lax, with several- to many-flowered cymes; axes glabrous,
smooth with few lanceolate, 13 mm long bracts and 1.56 mm
long pedicels. Ovary obovoid, ca. 1 mm, glabrous. Corolla
white, drying often yellowish brown, campanulate to funnelform, ca. 4 mm in diam., glabrous, lobed for ca. 1/2; lobes 4,
triangular, acute. Fruit on elongating pedicels with ellipsoid, ca.
2 mm, glabrous and smooth mericarps. Fl. JunJul, fr. Aug
Sep.
Montane river valleys, open forests, grasslands, rocky slopes and
talus; 13001900 m. Xinjiang [Russia].
When W. C. Chen described Galium xinjiangense, he compared it
only with the completely different G. odoratum, not being aware of the
certainly conspecific G. paniculatum. Because of its corolla shape, this
characteristic and relatively isolated taxon was originally described as
Asperula. But there is no affinity to any group of Asperula as presently
circumscribed. Instead, there are similarities with G. ser. Nemoralia M.
Popova of G. sect. Leiogalium and with some members of G. sect.
Orientigalium. Therefore, the transfer of A. paniculata to Galium by
Pobedimova was fully justified.
Pobedimova et al. (Fl. URSS 23: 271. 1958) also discussed the
disjunct distribution of this relict species, which extends from its center
in the Altai to the middle Yenisei and to the Dzungarian Alatau in NW
China (Xinjiang).

39. Galium paradoxum Maximowicz, Bull. Acad. Imp. Sci.

Saint-Ptersbourg 19: 281. 1874.

4000 m. Anhui, Gansu, Guangxi, Guizhou, Hebei, Heilongjiang, Henan,

Hubei, Hunan, Jilin, Liaoning, Qinghai, Shanxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Japan, Korea, Nepal, Russia].
Galium paradoxum is a rather unusual species of the genus because of its broad, petiolate, and pinnately nerved opposite leaves,
forming whorls of 4 with 2 smaller leaflike stipules. Therefore, it can be
confused with other genera, in particular Kelloggia, which differs by its
calyx teeth and its never-leaflike stipules. Because of its isolated
position, G. paradoxum was made the type species of G. sect. Cymogalia and placed into a monotypic series, G. ser. Paradoxa (see
Pobedimova et al., Fl. URSS 23: 326. 1958). As shown by Ehrendorfer
et al. (Fl. Iranica 176: 232. 2005) and unpublished DNA analyses, it is
only distantly related to G. kamtschaticum, G. rotundifolium, and G. elegans in G. sect. Platygalium s.l. or to members of G. sect. Hylaea, as G.
hoffmeisteri.
Schnbeck-Temesy and Ehrendorfer (in Tan et al., Davis & Hedge
Festschrift, 111114. 1989) commented on the morphological variation
and biogeography of Galium paradoxum and recognized three subspecies, of which subsp. paradoxum and subsp. duthiei occur in China
as keyed out and described below. The third, G. paradoxum subsp.
franchetianum Ehrendorfer & Schnbeck-Temesy, is restricted to Japan
and can be recognized by its relatively larger leaves with relatively short
trichomes and often acute apices, and by its relatively large flowers. A
survey of many new collections now available from the herbaria PE,
KUN, MO, and WU shows that ranges of variation in all differential
features used overlap and that many intermediates occur. With respect to
the two Chinese taxa one can suspect not only an allopatric but also an
altitudinal differentiation pattern.

1a. Leaves usually lanceolate-ovate,

(10)1230(40) (5)715(23)
mm; corolla lobes obtuse to
subapiculate ................................... 39a. subsp. paradoxum
1b. Leaves broadly ovate to suborbicular,
(5)610(17) (3.5)47(10) mm;
corolla lobes acute to acuminate .......... 39b. subsp. duthiei
39a. Galium paradoxum subsp. paradoxum

lin zhu yang yang

() lin zhu yang yang (yuan ya zhong)

Herbs, perennial, ascending from filiform rhizomes. Stems

erect, slender, 425 cm tall, 4-angled and narrowly winged, glabrous and smooth, only nodes slightly shortly hairy. Middle
stem leaves opposite and with 2 leaflike but clearly smaller
stipules in whorls of 4, at lower nodes stipules linear, 1.53
mm; petiole 1.510 mm; leaf blade membranous, suborbicular,
broadly ovate to ovate-lanceolate, or elliptic-oblong, (5)6
30(40) (3.5)515(23) mm, adaxially with scattered, appressed, short hairs, abaxially glabrescent, base attenuate, obtuse to truncate, margins antrorsely hispidulous-ciliolate, apex
acute to rounded; single principal vein with 24 pairs of pinnate
lateral veins. Inflorescences terminal and in axils of upper
leaves with 311-flowered cymes; axes trichotomous and divaricate; bracts narrowly elliptic or ligulate, 0.83 mm; pedicels
13 mm. Ovary ovoid, ca. 0.5 mm, with undeveloped uncinate
hairs. Corolla white, rotate, 2.53 mm in diam., lobed for 1/2
2/3; lobes ovate, obtuse, subapiculate to acute or acuminate.
Mericarps ovoid, 12 mm, densely covered with uncinate
yellowish brown trichomes 0.81 mm, on pedicels thickening
and elongating up to 11 mm. Fl. MayAug, fr. JunSep.

Leaves ovate-lanceolate or sometimes ovate or elliptic-oblong, attenuate at base, (10)1230(40) (5)715(23) mm.
Corolla lobes obtuse to subapiculate. Fl. MayAug, fr. JunSep.

Forests, meadows, near water, on shady (sub)alpine rocks; 1200

Forests, meadows, near water; 12003000 m. Anhui, Gansu,

Guangxi, Guizhou, Hebei, Heilongjiang, Henan, Hubei, Hunan, Jilin,
Liaoning, Qinghai, Shanxi, Sichuan, Yunnan, Zhejiang [Bhutan, Korea,
Russia].

39b. Galium paradoxum subsp. duthiei Ehrendorfer & Schnbeck-Temesy in Tan et al., Davis & Hedge Festschrift, 113.
1989.
da shi lin zhu yang yang
Leaves broadly ovate to suborbicular, (5)610(17)
(3.5)47(10) mm, truncate at base. Corolla lobes acute to
acuminate. Fl. JulAug, fr. JulSep.
On shady (sub)alpine rocks; 27004000 m. Hubei, Sichuan, Xizang, Yunnan [Bhutan, India, Nepal].
In an otherwise quite typical specimen of this subspecies (e.g.,
Sichuan, Lutinghsien Mujiaogou: T. P. Wang 18-9-1938, PE) the stipules at the mid-stem region are quite comparable to true leaves, only

RUBIACEAE

slightly smaller. This results in a considerable similarity to small plants

of Galium hoffmeisteri from G. sect. Hylaea, which differ by their
leaves and leaflike stipules in whorls of up to 5 or 6.

40. Galium platygalium (Maximowicz) Pobedimova, Novosti

Sist. Vyssh. Rast. 7: 277. 1971.
luan ye lun cao
Asperula platygalium Maximowicz, Bull. Acad. Imp. Sci.
Saint-Ptersbourg 19: 284. 1874.
Herbs, perennial, from thin creeping rhizomes. Stems
erect, 2035 cm tall, with 4 thickened angles, smooth, at nodes
glabrous or hispidulous. Leaves in whorls of 4(6), subsessile,
with petiole up to 2 mm; blade drying papery to leathery,
elliptic or elliptic-oblong to ovate, (12)2025(28) (7)10
11(15) mm, glabrous or hirtellous to hispidulous along principal veins, abruptly narrowed into short petiole, margins
antrorsely ciliolate, apex obtuse to subacute; principal veins 3
5, palmate. Inflorescences umbel-like thyrsoid and many-flowered panicles with terminal and lateral cymes from uppermost
leaf axils; peduncles smooth and glabrous; bracts narrowly oblanceolate or elliptic to ligulate, 13 0.51.5 mm; pedicels
0.51.5 mm. Ovary ellipsoid, ca. 0.8 mm, glabrous. Corolla
white, funnelform, with tube as long as lobes, 45 mm in
diam.; lobes 4, elliptic-oblong. Mericarps ovoid, 1.72.8 mm,
glabrous and smooth. Fl. and fr. JulSep.
Open forests on mountain slopes; ca. 1700 m. Heilongjiang, Jilin,
Shanxi [Korea, Russia].
Galium platygalium is closely related to G. maximoviczii (see
there) but has larger flowers and whorls of opposite leaves and leaflike
stipules with 4 or more (rarely up to 6) elements. Because of their
slightly elongated, fused corolla base, the two species in the older literature were placed into the genus Asperula, where they had no other
close relatives. In spite of their increase in leaf whorl elements, the two
species have been placed by Ehrendorfer et al. (Fl. Iranica 176: 175.
2005) into G. sect. Platygalium s.l. Though this section normally is characterized by leaf whorls with only up to 4 elements, this placement may
be justified because the two species correspond otherwise quite well, as
in habit, inflorescences, and leaves (with 35 palmate principal veins).
Furthermore, Galium platygalium and G. maximoviczii exhibit
similarities with Microphysa elongata (see p. 216), also with a welldeveloped corolla tube but with leaves in whorls of only 4 and only 1
main vein. Thus, the monotypic genus Microphysa, maintained here,
is another obvious member of G. sect. Platygalium.

41. Galium prattii Cufodontis, Oesterr. Bot. Z. 89: 244. 1940.

kang ding la la teng
Herbs, perennial, erect to weak, up to 50 cm tall. Stems
obtusely 4-angled, usually branched, retrorsely aculeolate on
angles or glabrous. Leaves in whorls of 6, sessile; blade drying
papery, lanceolate, 1535 2.56 mm, glabrous, adaxially
smooth or scaberulous along midrib and near margins, abaxially
sparsely retrorsely aculeolate along midrib, base cuneate, margins sparsely to densely retrorsely aculeolate, flat to thinly
revolute, gradually narrowed into acute to acuminate apex; vein
1. Inflorescences terminal and axillary, cymes many flowered;
peduncles glabrous, smooth; bracts none or few, 12 mm; pedicels 14 mm. Ovary obovoid, 0.50.8 mm, glabrous. Corolla

133

white to greenish, rotate, 2.53 mm in diam., lobed for 2/3 or

more, glabrous; lobes 4, triangular-spatulate, acute to shortly
acuminate. Mericarps ovoid, ca. 1 mm, glabrous, smooth or minutely granulose. Fl. Jun, fr. Aug.
Valleys, open habitats of the montane zone; 31003700 m.
Sichuan (Kangding).
Galium prattii apparently is endemic in Sichuan. It is often so
close to forms of the reddish-purplish flowering G. blinii that only the
longer lanceolate leaves, the more flower-rich cymes, and the white to
greenish flower color of G. prattii help to separate them. Otherwise, G.
prattii seems to link G. asperifolium and G. tokyoense. From the first it
differs by less-bracteate inflorescences and less-acuminate corolla lobes,
from the second by its leaves, which are lanceolate and apically more
gradually narrowed (vs. subspatulate and apically rounded, mucronate).

42. Galium pusillosetosum H. Hara, J. Jap. Bot. 51: 134. 1976.

xi mao la la teng
Herbs, perennial, procumbent and ascending, caespitose,
(5)1020(40) cm tall. Stems with 4 prominent whitish angles,
variable from densely hispid with straight trichomes of ca. 0.8
mm to retrorsely (very rarely also antrorsely) aculeolate. Middle stem leaves in whorls of up to 5 or 6 (never only 4), subsessile; blade drying blackish, narrowly oblanceolate to narrowly lanceolate, (3)510(17) 0.82(4.2) mm, hispid,
retrorsely aculeolate on midrib and margins or glabrescent, base
cuneate, apex acute and shortly mucronate; vein 1, whitish. Inflorescences with axillary and/or terminal cymes, few to several
flowered; peduncles divaricate, with leaflike bracts, 13 mm;
pedicels 0.53 mm. Ovary ovoid, ca. 0.6 mm, densely hispidulous, glabrescent or glabrous. Corolla purple, yellowish green,
or white, rotate, 2.53 mm in diam., glabrous or scaberulous
inside; lobes 4, ovate, acute. Fruit subglobose, ca. 2 mm in
diam., with dense to sparse uncinate trichomes 0.50.7 mm or
glabrous and smooth. Fl. and fr. MayAug.
Mountain slopes, open ground and grasslands; 21003900 m.
Gansu, Nei Mongol, Ningxia, Qinghai, Shanxi, Sichuan, Xinjiang, Xizang [Bhutan, Nepal].
Galium pusillosetosum is an obvious alpine member of the G.
asperifolium group (G. sect. Trachygalium s.l.). Main differential characters are its relatively dense leaf and stem indumentum and its conspicuous whitish stem angles. These separate it from G. acutum, G.
baldensiforme, G. megacyttarion, G. rebae, G. glabriusculum, and G.
sungpanense. Nevertheless, its variability with respect to leaf epidermal
cell size, flower color, and ovary and fruit indumentum is remarkable.

43. Galium rebae R. R. Mill, Edinburgh J. Bot. 53: 195. 1996.

rui ba la la teng
?Galium bodinieri H. Lveill.
Herbs, perennial, procumbent and mat-forming. Stems
(5)1430(45) cm, 4-angled or -sulcate, branched, glabrous
and smooth, but sometimes with scattered straight hairs. Leaves
in whorls of up to 6, sessile; blade drying papery, blackish,
linear-lanceolate to narrowly oblanceolate, 2.510.5 0.41.4
mm, with inconspicuous epidermal cells, glabrous and smooth,
rarely with a few straight hairs on abaxial midvein, base
cuneate, margins entire and smooth, flat to thinly revolute, apex
acute, contracted and shortly mucronate; vein 1. Inflorescences

RUBIACEAE

134

with predominantly axillary cymes, 16-flowered; axes glabrous, smooth; pedicels 0.13.2 mm. Ovary ellipsoid-obovoid,
ca. 0.5 mm, glabrous, smooth. Corolla red, purple, or occasionally white, rotate, 1.73.6 mm in diam., glabrous to papillose; lobes 4, lanceolate-spatulate, adaxially glabrous except
puberulent on margins and central vein, apex acute to shortly
acuminate. Mericarps ellipsoid, ca. 1.5 0.7 mm, glabrous and
smooth or granular-verruculose, on pedicels often elongating to
5 mm. Fl. and fr. JunNov.
Damp banks under evergreen forests, alpine meadows, on rocks;
20004000 m. Sichuan, Xizang, Yunnan [Bhutan, India (Sikkim), Nepal].
Galium rebae belongs to the Himalayan and SW Chinese complex
of (sub)alpine taxa from the G. asperifolium group studied by Mill
(Edinburgh J. Bot. 53: 193213. 1996; Fl. Bhutan 2(2): 825834. 1999).
It is closely related to G. acutum and often only separable by its flower
color (see comments under that species). At the same time, it appears
linked to the likewise reddish flowering but larger G. blinii at lower
elevations. Reddish flowers also occur in G. pusillosetosum, which
differs by its dense stem, leaf, and fruit indumentum.

44. Galium rupifragum Ehrendorfer, Novon 20: 273. 2010.

ping bian la la teng
Herbs, perennial, densely caespitose, emerging from a
slender branching rootstock; all vegetative parts very fragile
when dried and with a loose indumentum of soft hairs, 0.50.8
mm, straight and spreading, on upper leaf surface slightly retrorse. Stems ascending or erect, 510 cm tall, 4-angled, hairy,
with 1218 internodes, increasing in length from 28(15) mm
upward, with some short vegetative lateral branches from middle region to inflorescence base. Leaves in whorls of 4; blade
thinly papery and remaining greenish when dried, ovate to
broadly lanceolate, 58 2.53.5 mm, loosely hairy on both
sides and marginally, base attenuate, margins flat or slightly
revolute, apex acute to apiculate; principal veins 3, palmate,
lateral weak. Inflorescences terminal, often with 3 cymes, each
with 35 flowers; bracts few and reduced; peduncles 45 mm
and pedicels 0.53 mm, glabrescent, somewhat elongated and
divaricate in fruit. Flowers hermaphroditic. Ovary ovoid, ca. 0.5
0.3 mm, with still undeveloped appressed hairs. Corolla
greenish white, rotate, ca. 1.5 mm in diam., with 4 triangular
and acute to slightly apiculate lobes. Mericarps 0.81 mm, with
spreading uncinate trichomes ca. 0.25 mm. Fl. and fr. JulSep.
Mountain regions, on rocks; ca. 1800 m. Yunnan (Pingbian).
The above description of Galium rupifragum is based on two
sheets collected by H. T. Tsai (H. T. Tsai 60986) on rocks at the type
locality and deposited in PE. Galium rupifragum belongs to G. sect.
Platygalium s.l. and exhibits affinities with the G. bungei group, in
particular with G. salwinense. Both share slender growth and uncinate
fruit hairs. But G. salwinense has fewer (only up to 10) and longer (up to
1020 mm) stem internodes, smaller leaves, only 1 (and not 3) main
leaf veins, and smaller flowers. Nevertheless, on Emei Shan (Sichuan)
typical G. salwinense occurs at lower elevations, whereas at higher
elevations reduced forms approach G. rupifragum. In addition, there are
also obvious similarities between G. rupifragum and representatives of
the G. morii group (see there), growing with five accepted species in the
high mountains of Taiwan, particularly with G. morii itself and with the
related G. formosense. Main differences from the latter are its lower
growth and smaller leaves, from the former its hirsute stems and the
more divaricate, often longer peduncles and pedicels.

45. Galium salwinense Handel-Mazzetti, Symb. Sin. 7: 1028.

1936.
nu jiang la la teng
Herbs, perennial, weak, procumbent to ascending, from
filiform rhizome, 850 cm tall. Stems 4-angled, glabrous or
with sparse spreading hairs. Middle stem leaves in whorls of 4,
sessile or subsessile; blade drying papery, dark green, narrowly
ovate or elliptic to oblong-lanceolate, (3)518(23) (1.5)3
7(11) mm, sparsely and at margins antrorsely hairy, base attenuate, apex slightly acute; vein 1. Inflorescences terminal and
axillary, slender and slightly divaricate, cymes 1.56 cm, rather
few flowered; axes glabrous, nearly ebracteate; pedicels (2)4
8(12) mm. Ovary obovoid, ca. 1 mm, appressed hairy. Corolla
yellowish, rotate, 11.3 mm in diam.; lobes 4, triangular,
obtuse. Mericarps ovoid, 11.5 mm, densely with short uncinate trichomes ca. 0.1 mm. Fl. and fr. JulOct.
Shady habitats, rocks in forests; 17002800 m. Sichuan, Yunnan
(Gongshan).
The regional Galium salwinense is apparently closely related to
the widespread G. bungei. Nevertheless, its habit, elongated pedicels,
and fruit surface, always with short and spreading hooked trichomes,
offer reliable differential characters.

46. Galium saurense Litvinov, Trudy Bot. Muz. Imp. Akad.

Nauk 7: 75. 1910.
xia xu la la teng
Galium densiflorum Ledebour var. saurense (Litvinov)
Tzvelev.
Herbs, perennial, caespitose, with stout rootstock and slender, woody rhizomes. Stems erect or ascending, 830 cm tall, 4angled and puberulent. Leaves in whorls of 4 in lower and of
6 in middle stem region, sessile; blade dark green adaxially,
pale green abaxially, linear or linear-oblong, 715 0.52.5
mm, smooth or usually sparsely to densely scaberulous adaxially, usually densely hairy abaxially, base acute to cuneate,
margins revolute, apex mucronate; vein 1. Inflorescences
narrowly paniculate with axillary and terminal, 315-flowered cymes; peduncles bracteose, densely pubescent (rarely
glabrescent), with 13 mm long pedicels. Ovary ellipsoid,
puberulent. Corolla yellow, rotate, 34 mm in diam., lobed for
3/4 or more; lobes 4, ovate-oblong, acute. Mericarps ellipsoid,
ca. 2 3 mm, densely puberulent (rarely glabrous or tuberculate). Fl. and fr. JulAug.
Alpine and subalpine habitats. Qinghai, Xinjiang [Kazakhstan,
Kyrgyzstan, Mongolia, Russia].
Galium saurense, a C Asiatic mountain taxon, belongs to G. sect.
Galium and the extremely polymorphic Eurasiatic G. verum group. It
refers to condensed alpine populations, in which the leaf whorls are
reduced to 46 elements. The Kew Rubiaceae checklist (Govaerts et
al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/;
accessed on 15 Sep 2010) treats it as a synonym of G. verum subsp.
verum, whereas Ehrendorfer et al. (Fl. Iranica 176: 199, 204. 2005) suggest to maintain its specific rank as long as the whole group has not
been studied more intensively. In Pobedimova et al. (Fl. URSS 23: 368
369. 1958) G. saurense is reported i.a. from the Tien Shan in the border
region of Kazakhstan and Xinjiang, but its description is in conflict with

RUBIACEAE

plate 21, figure 1, which rather corresponds to G. majmechense and G.

consanguineum. W. C. Chen (in FRPS 71(2): 285. 1999) included G.
saurense as a dubious species and suspected its occurrence in NW
China. We have seen no authentic specimens, but vouchers from Xinjiang and Qinghai (and possibly other provinces) fit its description quite
well. Nevertheless, their separation from G. verum s.s. is partly doubtful.

47. Galium serpylloides Royle ex J. D. Hooker, Fl. Brit. India

3: 207. 1881, s.l.
long zi la la teng
Herbs, perennial, ascending, 310 cm tall. Rootstock
woody, rhizomatous. Stems 4-angled, branched, shortly pubescent. Leaves in whorls of 4(or ?5), sessile; blade drying papery,
ovate or elliptic, 46 2.54 mm, glabrous or sparsely hirtellous, base rounded, margins thinly revolute, usually with antrorse microhairs, apex obtuse or slightly acute; principal vein 1
(or with 2 inconspicuous side veins). Inflorescences terminal
and in axils of upper leaves, with cymes 58 mm, few or usually 3-flowered; pedicels 12 mm. Flowers hermaphroditic to
subdioecious. Ovary appressed hairy. Corolla yellowish green,
rotate, ca. 2 mm in diam.; lobes 4, ovate-triangular, acute. Mericarps up to 2 mm, with straight lanate hairs. Fl. and fr. JunSep.
Mountain slopes; 36003800 m. Xizang (Lhnz) [India, Nepal].
Originally, Galium serpylloides was very broadly circumscribed
as a species widely distributed throughout the whole Himalayan range.
But a critical taxonomic analysis of the group by Schnbeck-Temesy
and Ehrendorfer (Pl. Syst. Evol. 155: 7787. 1987) revealed five vicarious and geographically well-separated species from Kashmir in the west
to Nepal in the east: G. gymnopetalum Ehrendorfer & SchnbeckTemesy, G. lakulense Ehrendorfer & Schnbeck-Temesy, G. serpylloides
s.s., G. saipalense Ehrendorfer & Schnbeck-Temesy, and G. nepalense
Ehrendorfer & Schnbeck-Temesy. The above cited locality from E
Xizang, Lhnz, published in FRPS (71(2): 228. 1999), is much further
to the east, i.e., SE of Lhasa and NE of Bhutan. We have neither seen
authentic material from this nor from other Xizang and Charme province collections in the herbarium BM (e.g., Ludlow, Sheriff & Taylor
6390). At the moment, it is not possible to decide whether they correspond to one of the above mentioned species of the G. serpylloides
group, or the related G. glandulosum group, or represent a still undescribed taxon. Therefore, we provisionally classify these SE Xizang
populations under G. serpylloides s.l.
Characteristic morphological features of the Galium serpylloides
group are the whorls of 4 leaves and leaflike stipules, the trend toward
palmate leaf veins and sexual differentiation of flowers, and the change
from hooked to straight fruit hairs. All this suggests relationships with
members of G. sect. Platygalium s.l., as the G. morii group (in particular
G. nankotaizanum) or the G. elegans group. These trends reappear
(probably as a homoplasy?) in the W North and South American G. sect.
Lophogalium K. Schumann, evidently derived from ancestors similar to
G. sect. Platygalium s.l.

48. Galium sichuanense Ehrendorfer, Novon 20: 275. 2010.

si chuan la la teng
Herbs, perennial, from filiform rhizomes ascending to
erect, ca. 30 cm tall. Stems single, strongly branched from base,
with 4 prominent and whitish angles, glabrous and smooth,
only slightly aculeolate at nodes. Middle stem leaves and leaflike stipules in whorls of 46; blade drying papery and
greenish-brownish, lanceolate, (12)1520(25) (3)46(7)
mm, glabrous but on adaxial side near margins with antrorse
appressed microhairs, on flat margins retrorsely (sometimes

135

also somewhat antrorsely) aculeolate, subsessile and gradually

narrowed into base, largest breadth near middle, apex cuspidate
with hyaline point; principal vein 1. Inflorescences broadly
ovate, many flowered, cymes terminal and lateral, 2.55 cm,
slender, leafy and bracteate to last branches, divaricate; axes
glabrous; peduncles 1.52 cm; pedicels 0.55 mm. Flowers
hermaphroditic. Ovary obovoid, 0.50.8 mm, with appressed
curved hairs. Corolla dried reddish brown, rotate, 1.52 mm in
diam.; lobes 4, triangular, cuspidate. Mericarps ovoid, 1.82.5
mm, with spreading uncinate trichomes 0.20.3 mm. Fl. Jul
Aug, fr. AugSep.
Mountain forests; 32004000 m. Sichuan (Daocheng).
Up to now, only two collections of Galium sichuanense are
known from the type locality. The new species is morphologically isolated and not closely related to any other taxon of the genus. Its character profile, particularly its perennial, hemicryptophytic growth form,
the partly retrorsely aculeolate leaf margins, and the fruit with uncinate
trichomes, designates it as a member of G. sect. Trachygalium, but there
are also features reminiscent of G. sect. Hylaea (cf. Ehrendorfer et al.,
Fl. Iranica 176: 181. 2005). Nevertheless, the combination of the
branching pattern, the membranous leaves, the predominant glabrescence, the leafy inflorescences, and the small flowers separate it clearly
from all other members of the two sections.
Another isolated and apparently relict species from E Asia has to
be compared with Galium sichuanense, the Japanese G. kikumugura
(see also under G. hoffmeisteri). The two share the leaves in whorls of
46, the small flowers, and the hooked fruit trichomes. In contrast, the
habit and the somewhat antrorsely rough or even aculeolate leaf margins of G. kikumugura are reminiscent of G. bungei (G. sect. Platygalium). Unique characters of G. kikumugura are the very few-flowered
cymes on long peduncles with a single bract and the elongate, curved
mericarps. Its taxonomic placement within Galium also is uncertain.

49. Galium glabriusculum Ehrendorfer, nom. nov.

wu geng la la teng
Replaced synonym: Galium smithii Cufodontis, Oesterr.
Bot. Z. 89: 236. 1940, not Galium smithii G. Don, Gen. Hist. 3:
660. 1834 [Sherardia erecta Smith, Fl. Graec. 2: 14. 1813].
Herbs, perennial, weakly procumbent to erect. Stems
(5)715(30) cm, 4-angled, glabrous and smooth, rarely
shortly hairy at nodes. Leaves in whorls of up to 5 or 6,
subsessile; blade greenish brown and subleathery when dried,
narrowly lanceolate to narrowly elliptic, 312 12.5 mm,
glabrous and smooth, very rarely with straight hairs adaxially or
slightly retrorsely aculeolate on margins, base cuneate, margins
thinly revolute, apex acute and shortly mucronate; vein 1. Inflorescences with terminal and lateral 13-flowered cymes on up
to 10 mm long peduncles (often elongating in fruit) and with
subsessile flowers on 0.53 mm long pedicels. Ovary subglobose, ca. 1 mm, densely covered with undeveloped trichomes.
Corolla white, rotate, ca. 1.5 mm in diam.; lobes 4, ovate, obtuse. Mericarps ellipsoid, ca. 3 mm, with dense spreading yellowish brown uncinate trichomes ca. 0.7 mm. Fl. and fr. Jul
Aug.
Meadows, alpine mountain slopes; 38004700 m. Gansu, Qinghai, Sichuan, Xinjiang.
Galium glabriusculum is a well-documented species of the alpine
subgroup (2) within the G. asperifolium complex (see under that species
and G. acutum). It is similar to G. sungpanense and G. baldensiforme

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136

and mainly differs from the former by its narrower and more leathery
leaves and from both by the nearly total lack of indumentum.

50. Galium spurium Linnaeus, Sp. Pl. 1: 106. 1753.

zhu yang yang
Galium agreste Wallroth; G. agreste var. echinospermum
Wallroth; G. agreste var. leiospermum Wallroth; G. aparine
Linnaeus var. echinospermum (Wallroth) T. Durand; G. aparine
f. leiocarpum Makino; G. aparine var. leiospermum (Wallroth)
T. Durand; G. aparine var. spurium (Linnaeus) W. D. J. Koch;
G. aparine var. tenerum (Grenier & Godron) H. G. Reichenbach; G. aparine var. vaillantii (Candolle) W. D. J. Koch; G.
hongnoense H. Lveill; G. oliganthum Nakai & Kitagawa; G.
pauciflorum Bunge (1833), not Willdenow ex Candolle (1830);
G. spurium var. echinospermum (Wallroth) Hayek; G. spurium
var. tenerum Grenier & Godron; G. vaillantii Candolle; G. wutaicum Hurusawa.
Herbs, annual, procumbent or climbing, 3050 cm tall.
Stems 4-angled, 0.52.5 mm in diam., branched from base,
retrorsely aculeate on angles, glabrescent to pilose at nodes.
Leaves at middle stem region in whorls of 68, subsessile;
blade drying papery, narrowly oblanceolate to narrowly oblongoblanceolate, 540 15(8) mm, usually pilosulous or hispidulous adaxially, retrorsely aculeolate along midrib abaxially
and along margins, base acute, margins flat to thinly revolute,
apex acute and shortly mucronate; vein 1. Inflorescences terminal and axillary, cymes 2- to several flowered; axes glabrous
to aculeolate; bracts leaflike or none, 15 mm; peduncles 14
cm; pedicels 0.515 mm, finally elongating and often curved
directly under fruit. Ovary subglobose, 0.30.5 mm, with uncinate trichomes or glabrous. Corolla yellowish green or white,
rotate, 11.5 mm in diam., lobed for 2/3 or more; lobes 4, triangular to ovate, acute. Mericarps subglobose to broadly kidneyshaped, 13 mm in diam., glabrous or often densely covered
with uncinate trichomes 0.11 mm from straight bases. Fl.
MarJul, fr. AprNov.
Open fields, riversides, farmlands, mountain slopes; near sea level
to 4600 m. Common and widespread throughout China except Hainan
and Nanhai Zhudao [Africa, Eurasia, and the Mediterranean; today
sporadically adventive worldwide].
This species is occasionally used medicinally.
Galium spurium consists of basal elements (2x and 4x, 2n = 20,
40) of a polymorphic polyploid complex, the G. aparine group or G.
aparine s.l. (see there). Following Ehrendorfer et al. (Fl. Iranica 176:
234. 2005), G. spurium is maintained here at the species level, with
particular reference to its differential characters in flower and fruit size,
and not included under the higher polyploid and aneuploid G. aparine
s.s., as in Cufodontis (Oesterr. Bot. Z. 89: 245247. 1940) and W. C.
Chen (in FRPS 71(2): 237. 1999). Galium spurium is very common and
widespread in China, in contrast to the rare and partly doubtful G.
aparine s.s. To our knowledge, no chromosome counts are yet available
from Chinese populations of G. spurium. Nevertheless, a report of 2n =
40 for this species from Novosibirsk (Krasnikov & Schaulo, Bot. urn.
75: 118120. 1990) suggests the occurrence of 4x G. spurium cytotypes
in Asia, corresponding to similar 4x-cytotypes reported from Africa.
From the varieties recognized by Cufodontis (loc. cit.) and
accepted by W. C. Chen (loc. cit.: 234237) Galium spurium var.
tenerum refers to reduced specimens, which can appear under extreme
conditions as modifications everywhere, and are taxonomically irrele-

vant. But as genetically fixed reduced alpine ecotypes they deserve a

name: G. spurium subsp. ibicinum (Boissier & Haussknecht) Ehrendorfer, described from high mountains in SW Asia (see Ehrendorfer
et al., loc. cit.: 236). Some condensed alpine Chinese specimens may
belong to this taxon.
In contrast, Galium spurium var. echinospermum vs. var. spurium
(= G. aparine var. leiospermum) refer to genetically fixed forms with
uncinate hairy vs. glabrous fruit, which in W Eurasia and the Mediterranean often occur together in the same population and can be separated
as taxonomic forms. In China we have seen only the echinospermum
type, whereas var. spurium apparently is missing there. The reference to
it by W. C. Chen (loc. cit.: 237) concerns the glabrous fruited G.
ghilanicum (see there). It is obvious that a more detailed analysis of the
G. aparine-G. spurium polyploid complex in E Asia is badly needed.
The enormous variability of Galium aparine and G. spurium has
caused its many synonyms and common misidentifications with other
annual and even perennial taxa of Galium. This applies in particular to
the annual G. tricornutum, which differs by strongly verrucose (but not
uncinate hairy) fruit, and to the perennial taxa with retrorsely aculeolate
stems and fruit with uncinate hairs, such as G. sungpanense (see there),
G. dahuricum, etc., which often have larger flowers.

51. Galium sungpanense Cufodontis, Oesterr. Bot. Z. 89: 238.

1940.
song pan la la teng
Herbs, perennial, slender, with tender rootstock. Stems
procumbent or ascending, up to 30 cm, 4-angled, retrorsely
aculeolate to somewhat hispidulous or smooth. Leaves in
whorls of up to 5 or 6, sessile or narrowed to very short petiole;
blade drying greenish brown and stiffly papery to subleathery,
oblanceolate or narrowly elliptic-oblanceolate, 3.512(15)
1.53.5 mm, glabrous to hispidulous, sparsely to densely retrorsely aculeolate along margins and sometimes also on abaxial side of midrib, margins thinly revolute, apex acute and
cuspidate; vein 1. Inflorescences with terminal and axillary
cymes, 13-flowered with lanceolate bracts; peduncles up to 10
mm, glabrous and smooth; pedicels 27 mm, straight and elongating in fruit. Ovary obovoid, 0.50.8 mm, densely covered by
undeveloped uncinate trichomes. Corolla pinkish or purplish,
rotate, 1.11.5 mm in diam., glabrous; lobes 4, triangular, obtuse. Fruit with obovoid mericarps, ca. 2.5 mm, densely covered with spreading yellowish brown uncinate trichomes 0.4
0.8 mm. Fl. and fr. JulSep.
Thickets or meadows, often in shady places; higher elevations
up to 3300 m. Hebei, Sichuan, Xinjiang.
The description of Galium sungpanense in FRPS (71(2): 233.
1999) includes some details that do not agree with the material seen and
may have been based in part on specimens of other taxa. This has been
corrected in the above description.
Galium sungpanense belongs to the throughout-perennial G. sect.
Trachygalium s.l. and the G. asperifolium group (see there). It appears to
link its montane (1) and alpine (2) subgroups and shares the few-flowered cymes with the latter. From G. baldensiforme and G. glabriusculum, both also with uncinate fruit hairs, it is separated by marginally
stronger retrorsely aculeolate and partly longer leaves.
Remarkable are the close affinities between Galium sungpanense
and members of the annual G. sect. Euaparine. The single decisive
difference is the tender perennial (and possibly short-lived?) rootstock
of G. sungpanense. Only its pinkish to purplish flowers and the nevercurved fruiting pedicels allow the separation of plants collected without

RUBIACEAE

subterranean organs from the common G. spurium. This suggests that G.

sect. Euaparine could have originated from G. sect. Trachygalium-like
ancestors.

52. Galium taiwanense Masamune, Trans. Nat. Hist. Soc. Formosa 29: 180. 1939.
tai wan zhu yang yang
Herbs, perennial, procumbent. Stems 4-angled, sparsely
scaberulous. Leaves on main stems in whorls of up to 6, sessile;
blade drying ?papery, oblanceolate or narrowly obovate-oblong,
(4)1020(31) (1)24 mm, glabrous and smooth adaxially,
glabrous or hairy and retrorsely aculeolate along midrib abaxially and along margins, base acute to cuneate, apex acute or
acuminate; vein 1. Inflorescences with terminal and axillary,
few- to several-flowered, usually regularly trichotomous cymes;
bracts linear, ca. 2 mm; pedicels slender, 35 mm. Ovary ovoid,
ca. 0.7 mm, glabrous. Corolla ?white, rotate, 22.5 mm in
diam., lobed for 2/3 or more; lobes 4, ovate, obtuse. Mericarps
ellipsoid, ca. 2 mm, glabrous. Fl. MayJul, fr. Jul.
Mountain slopes; 2002100 m. N Taiwan.
We have seen no material from this taxon. Judging from the information and the holotype photograph presented by Yang and Li (Bull.
Natl. Mus. Nat. Sci., Taichung 11: 109, pl. 4, 3+4. 1998), this appears to
be close to and possibly even identical with Galium dahuricum var.
dahuricum.

53. Galium takasagomontanum Masamune, Trans. Nat. Hist.

Soc. Formosa 26: 52. 1936.
shan di la la teng
Herbs, perennial (not annual), erect, ca. 30 cm tall. Stems
4-angled, slender, much branched, glabrous. Leaves in whorls
of 4 or 5, sessile; blade narrowly ovate or ovate, abaxially pilose along veins, apex acuminate; principal vein 1 or with 2
short and weakly developed lateral veins. Inflorescences terminal, with few-flowered trichotomous cymes; peduncles glabrous; pedicels 12 mm, glabrous. Ovary ovoid-subglobose, ca.
1 mm, with undeveloped trichomes. Corolla white, rotate, ca. 3
mm in diam.; lobes 4, acute. Mericarps subglobose, with dense
uncinate trichomes. Fl. and fr. Jul.
Conifer mountain forests; ca. 2800 m. Taiwan.
Galium takasagomontanum was not treated or mentioned by Yang
and Li (Bull. Natl. Mus. Nat. Sci., Taichung 11: 101117. 1998; Fl. Taiwan, ed. 2, 4: 254259. 1998), and we have not seen authentic material.
According to the protologue it may belong to the condensed Taiwan
mountain group of G. morii (see there), where it appears similar with its
weakly 3-nerved leaves to G. nankotaizanum but differs by glabrous
stems, somewhat larger flowers, and uncinate trichomes on mericarps.
Nevertheless, its quite tall growth, the partly 5-whorled leaves, the exclusively terminal inflorescences, and the flower size are more indicative of its being a synonym of the Taiwanese G. echinocarpum from G.
sect. Hylaea. A definite placement will only be possible after inspection
of the type material.

54. Galium tarokoense Hayata, Icon. Pl. Formosan. 7: 33.

1918.
tai lu ge zhu yang yang
Herbs, perennial, procumbent to erect, condensed, 58 cm

137

tall. Stems 4-angled, glabrous, smooth. Leaves in whorls of 4,

sessile or subsessile; blade drying papery, rhombic-elliptic, elliptic, or obovate, 36 23.5 mm, glabrous, base acute to cuneate, margins flat or thinly revolute, apex acute to somewhat
obtuse; vein 1. Inflorescences with terminal and axillary cymes,
1- or few flowered, 0.31.5 cm; peduncles glabrous, smooth,
ebracteate; pedicels 02 mm. Ovary obovoid, ca. 0.4 mm,
densely strigillose at sides. Corolla pale yellow, rotate, ca. 2
mm in diam.; lobes 4, triangular-oblong, acute. Mericarps ellipsoid, 11.5(2) mm, with dense, appressed uncinate trichomes
ca. 0.3 mm. Fl. and fr. summerwinter.
Shady sites on limestone substrates; 14002700 m. Taiwan.
Galium tarokoense belongs to a group of high mountain taxa from
Taiwan, including G. morii (see there). Its main differential characters
are the lack of indumentum on most parts, the only 1-nerved leaves, and
the short appressed fruit hairs.

55. Galium tenuissimum M. Bieberstein, Fl. Taur.-Caucas. 1:

104. 1808.
xian xi la la teng
Herbs, annual, erect. Stems (10)3050(60) cm tall, with
strong, intricate and divaricate branching from base, 4-angled,
usually retrorsely aculeolate, more rarely glabrescent to smooth.
Leaves at middle stem region in whorls of 68, subsessile or
sessile; blade drying papery, linear to oblanceolate, (4)8
15(20) 0.51(2) mm, mostly glabrous and sparsely to
densely antrorsely aculeolate near margins and on veins abaxially, base straight to acute, apex acute to acuminate-aristate;
vein 1. Inflorescences broadly paniculate, very lax and loosely
divaricate, cymes axillary and terminal, with 311(14) flowers; axes glabrous or sparsely hispidulous; bracts none or small,
leaflike; pedicels 315 mm, strongly elongating in fruit up to 20
mm. Ovary obovoid to ellipsoid, ca. 0.8 mm, glabrous. Corolla
whitish, pale yellow, or greenish, rotate to slightly cup-shaped,
1.52 mm in diam., glabrous; lobes 4, oblong-elliptic, acute to
aristate. Mericarps ellipsoid to obovoid, ca. 1 1.25 mm, glabrous, smooth or tuberculate. Fl. and fr. MayJul.
Open mountain slopes; 3002800 m. Xinjiang (Xinyuan) [Kashmir, Kyrgyzstan, Pakistan, Russia, Turkmenistan; SW Asia (Armenia,
Georgia, Iran, Iraq, Lebanon, Syria, Turkey), Europe (Balkan Peninsula,
Hungary)].
Galium tenuissimum is a member of the annual G. sect. Microgalium, differentiated by antrorsely aculeolate leaf margins. It is widespread through SE Europe, SW and C Asia, and reaches its eastern limit
in NW China.

56. Galium tokyoense Makino, Bot. Mag. (Tokyo) 17: 72.

1903.
dun ye la la teng
Galium asprellum Michaux var. tokyoense (Makino)
Nakai; G. dahuricum Turczaninow ex Ledebour var. tokyoense
(Makino) Cufodontis.
Herbs, perennial, erect or ascending but not clambering.
Stems 3070 cm tall, 4-angled, retrorsely aculeolate. Middle
stem leaves in whorls of 5 or 6, subsessile; blade drying papery,
subspatulate to obovate, (11)1735(40) (2.5)37(10)

138

RUBIACEAE

mm, mostly retrorsely aculeolate adaxially, along midrib abaxially and always along margins, base acute, apex rounded to
emarginate, abruptly cuspidate; vein 1. Inflorescences congested, cymes terminal and in axils of uppermost leaves, several
to many flowered, up to 4 cm; axes rough or glabrous and
smooth; bracts few and small, only on lower inflorescence
branches; pedicels 12 mm. Ovary obovoid, ca. 0.8 mm, glabrous. Corolla white, rotate, 1.33.5 mm in diam., glabrous;
lobes acute to obtuse. Mericarps obovoid, ca. 2 mm, glabrous,
smooth or tuberculate. Fl. and fr. JunJul.
Forests, grasslands, meadows, riversides, open fields; 200900 m.
Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Shandong [Japan,
Korea].
Cufodontis (Oesterr. Bot. Z. 89: 243244. 1940) and W. C. Chen
(in FRPS 71(2): 256. 1999) treated Galium tokyoense as a variety of G.
dahuricum (see discussion under that species), but in the recent Flora of
Japan (Yamazaki, Fl. Japan 3a: 239. 1993) it is again regarded as a
separate species. In view of its erect (not clambering) growth, the
abruptly acuminate leaf shape, the short and post-florally not elongated
pedicels, and the always glabrous fruit this rank appears quite justified.

57. Galium tricornutum Dandy, Watsonia 4: 47. 1957.

mai ren zhu
Herbs, annual, weakly ascending to procumbent or clambering. Stems 580 cm tall, 4-angled, often little branched,
glabrescent, densely retrorsely aculeolate on angles. Leaves in
whorls of 68, subsessile; blade drying papery, narrowly oblanceolate to narrowly elliptic, 1032 26 mm, glabrescent, upper side glabrous, lower side densely aculeolate along midrib,
base acute, margins densely retrorsely and antrorsely aculeolate, apex acute; vein 1. Inflorescences elongated thyrsoid,
cymes terminal and axillary on short lateral stems, mostly 35flowered; axes retrorsely aculeolate; bracts none or leaflike and
35 mm; pedicels 0.32 mm. Ovary ellipsoid to didymous, 0.3
0.5 mm, smooth to verrucose or spinulose. Corolla white, rotate, 11.5 mm in diam., lobed for 2/3 or more; lobes triangular.
Mericarps subglobose, ca. 3 46 mm, becoming verrucose to
tuberculate but never with uncinate trichomes, pendulous on
arching pedicels to 7 mm. Fl. AprJun, fr. MayMar.
Adventive weeds in meadows on mountain slopes, open fields,
river beaches, ditch sides; 4004000 m. Anhui, Gansu, Guizhou, Henan,
Hubei, Jiangsu, Jiangxi, Shaanxi, Shanghai, Shanxi, Sichuan, Xinjiang,
Xizang [India, Pakistan; N Africa, SW Asia, Europe, North America].
In general aspect and habit the weedy annual Galium tricornutum
from G. sect. Kolgyda strongly resembles G. spurium and G. aparine but
can be separated by its above glabrous leaves and its verrucose fruit on
arching pedicels. Galium tricornutum apparently is rare in China. The
above wide distribution data from FRPS evidently is due to misidentifications of the common G. spurium.
Galium tricornutum has long been treated under the illegitimate
superfluous name G. tricorne Stokes, published in 1787. Stokess intent
was to transfer Valantia aparine Linnaeus (= G. verrucosum Hudson,
1767) to Galium, where the epithet aparine was blocked by G.
aparine Linnaeus. The specimens on which Stokes based his name belonged partly to G. verrucosum and partly to G. tricornutum, two close
but very well-separated species. However, when Stokes published his
article, the previously and validly published name in Galium by Hudson

(1767) already existed and made his name superfluous. That remained
unnoticed and G. tricorne was generally used for our species. Only in
1975 did Dandy clarify this situation, designating G. tricornutum as the
new name for the long known but misnamed G. tricorne.

58. Galium trifloriforme Komarov, Trudy Imp. S.-Peterburgsk. Bot. Sada 18: 428. 1901.
ni san hua la la teng
Herbs, perennial, from elongated rhizomes. Stems procumbent to erect, (10)2540(65) cm tall, 4-angled, mostly
slightly retrorsely aculeolate, hirtellous at nodes. Leaves on
main stems in whorls of up to 6(8), subsessile; blade drying
papery, blackish or green, narrowly obovate to oblanceolate,
(12)1828(50) (3)510(15) mm, with sparse antrorse
microhairs adaxially, moderately retrorsely aculeolate abaxially
on vein and at leaf margins, base acute to cuneate, margins flat
to thinly revolute, apex acute, obtuse, or rounded and abruptly
mucronate; vein 1. Inflorescences with axillary and terminal
cymes on upper 2 or 3 nodes, mostly 28-flowered; axes glabrous, smooth; bracts none or few, narrowly elliptic to narrowly
lanceolate, 25 mm; pedicels ca. 1.5 mm. Ovary obovoid, ca.
0.5 mm, densely hispidulous with undeveloped trichomes. Corolla white or pale green, rotate, 1.52 mm in diam., glabrous;
lobes 4, triangular, acute. Mericarps ellipsoid, 1.52.5 mm, with
dense uncinate trichomes ca. 1 mm, fruiting pedicels divaricate
and elongating to 10 mm. Fl. and fr. JulSep.
Mountain forests, open fields; 22003400 m. Heilongjiang, Jilin,
Nei Mongol, Qinghai [Japan, Korea, NE Russia].
Galium trifloriforme is a variable and problematic taxon from NE
Asia. It was either accepted as a separate species (e.g., by Pobedimova
et al., Fl. URSS 23: 303. 1958; Yamazaki, Fl. Japan 3a: 239. 1993) or
was treated as a synonym of G. triflorum (see Cufodontis, Oesterr. Bot.
Z. 89: 236237. 1940) or of G. hoffmeisteri (e.g., W. C. Chen in FRPS
71(2): 230. 1999, as G. asperuloides subsp. hoffmeisteri; Govaerts et al.,
World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on 15 Sep 2010). It differs from the very close typical G.
triflorum by its more condensed inflorescence with cymes terminal and
on the upper 1 or 2(or 3) nodes, its always retrorsely aculeolate leaf
margins, and its mostly rough stems.
These differential characters make Galium trifloriforme a link between members of G. sect. Hylaea, with smooth stems and antrorsely
directed microhairs on leaf margins, and of the G. asperifolium group of
G. sect. Trachygalium, mostly with retrorsely aculeolate stems and retrorse microhairs on leaf margins (but often also with antrorse
microhairs on the adaxial leaf surface). From the available material, it
appears that G. trifloriforme applies to the central part of this practically
continuous morphological series. This series begins with G. odoratum,
G. hoffmeisteri, G. echinocarpum, G. nipponicum, and typical G.
triflorum on the side of G. sect. Hylaea, continues via G. triflorum var.
asprelliforme Fernald and G. trifloriforme, and ends on the other side
with G. asprellum, G. dahuricum, G. blinii, and other typical members of
the G. asperifolium group of G. sect. Trachygalium. It is remarkable that
this transitional series apparently corresponds to a polyploid complex
with marginal taxa including 2x-, G. triflorum 4x- and 6x-, and G.
trifloriforme 10x-cytotypes. Thus, phylogenetic reticulation may have
caused the still insufficiently resolved taxonomic confusion around G.
triflorum, G. trifloriforme, and their relatives (cf. Pobedimova et al., loc.
cit.: 287381; Cufodontis, loc. cit.; Yamazaki, loc. cit.; Ehrendorfer et
al., Fl. Iranica 176: 182. 2005).

RUBIACEAE

Among the relatively few relevant PE, KUN, and WU specimens

we have seen from China and the Himalaya only two typical Galium
triflorum specimens with antrorsely aculeolate leaf margins were found
(see there); otherwise, only plants corresponding to the above description of G. trifloriforme with leaf margins retrorsely aculeolate were
seen. This finding is in conflict with W. C. Chen (loc. cit.) who accepted
only G. triflorum for the Chinese flora, but certainly needs verification
from more extensive studies including Korea (from where G. trifloriforme was described) as well as adjacent NE Siberia and Japan, where
both taxa apparently occur.

59. Galium triflorum Michaux, Fl. Bor.-Amer. 1: 80. 1803.

san hua la la teng
Herbs, perennial, from slender rhizomes. Stems procumbent, ascending or erect, (15)2580(125) cm tall, 4-angled,
usually glabrous and smooth (rarely somewhat retrorsely
aculeolate), hirtellous at nodes. Leaves in whorls of up to 6(8),
subsessile; blade drying papery, sometimes blackening, narrowly obovate to broadly (ob)lanceolate, (15)2035(45)
(3)612(16) mm, glabrous but with appressed and antrorse
microhairs adaxially and antrorse aculei along margins, base
acute to cuneate, margin flat to thinly revolute, apex acute or
rounded and abruptly mucronate; vein 1. Inflorescences terminal and axillary, with few- to several-flowered cymes at upper
24 stem nodes; axes glabrous and smooth; bracts few, narrowly elliptic to narrowly lanceolate, 25 mm; pedicels ca. 1.5
mm. Ovary obovoid, ca. 0.5 mm, densely hispidulous with
undeveloped trichomes. Corolla white or greenish, rotate, 1.52
mm in diam., lobed for 3/4 or more, glabrous; lobes 4, triangular, acute. Mericarps ellipsoid, 1.52.5 mm, with dense uncinate
trichomes ca. 1 mm, on pedicels elongating up to 10 mm. Fl.
and fr. JulSep.
Mountain forests; 15002000 m. Guizhou, Sichuan [Japan, Korea,
Russia; Europe, North America].
Galium triflorum is obviously rare in China and does not occur in
the Himalaya, as already suspected by Pobedimova et al. (Fl. URSS 23:
300303. 1958). We have seen only two non-flowering specimens from
SW China (Guizhou: Northern Qian [Guizhou] Team 907, PE; Sichuan:
G. H. Yang 54472, PE), which apparently belong here. Indications for
more northern provinces by W. C. Chen (FRPS 71(2): 232. 1999) have
been listed here under G. trifloriforme (see there), a taxon not recognized as distinct by W. C. Chen (loc. cit.: 230). Both taxa urgently need
more detailed study. Galium triflorum is very similar and morphologically subcontinuous with G. trifloriforme but differs from it by its more
elongate inflorescences, mostly smooth stems, and particularly by its
antrorsely aculeolate leaf margins. This latter character clearly places it
into G. sect. Hylaea and into the close neighborhood of G. hoffmeisteri
on the mainland, G. echinocarpum on Taiwan, and G. nipponicum in
Japan.

60. Galium turkestanicum Pobedimova in Schischkin, Fl.

URSS 23: 717. 1958.
zhong ya la la teng
Herbs, perennial, erect, to 0.55 m tall. Rhizomes slender,
reddish. Stems 4-angled, glabrous and smooth except puberulent to hispidulous at nodes. Leaves in whorls of 4, sessile;
blade drying papery, linear or linear-lanceolate, 2740 39
mm, glabrous or antrorsely scabrous along midrib and margins,
base obtuse to rounded, apex narrowed then shortly obtuse;
principal vein 1, with 2 weaker lateral veins reticulating near
middle of blade. Inflorescences terminal, paniculate, many

139

flowered, 215 cm; peduncles glabrous or scabrous; bracts few,

oblanceolate, 12 mm; pedicels 14 mm. Ovary ellipsoid and
laterally flattened, ca. 1 mm. Corolla whitish, rotate, 45 mm in
diam.; lobes 4(or 5), elliptic to lanceolate, acute or mucronulate.
Mericarps ellipsoid, at least 2 mm, glabrous or with sparse
hooked trichomes ca. 0.3 mm. Fl. JulAug, fr. AugSep.
Meadows and dry slopes in the (sub)alpine zone. Expected in Xinjiang [Kazakhstan, Russia].
Galium turkestanicum is included as a dubious species for W
China by W. C. Chen (FRPS 71(2): 285. 1999). It belongs to the extremely polymorphic G. boreale group of G. sect. Platygalium s.l.
and represents a particularly vigorous taxon with very narrow leaves
and lateral leaf veins only weakly developed (Ehrendorfer et al., Fl.
Iranica 176: 180. 2005). Pobedimova (loc. cit.) gives its distribution
as including the Tien Shan. Therefore, it probably also occurs in NW
China, Xinjiang.

61. Galium uliginosum Linnaeus, Sp. Pl. 1: 106. 1753.

zhao zhu yang yang
Herbs, perennial, tender, from slender rhizomes. Stems
1060 cm tall, somewhat retrorsely aculeolate along 4 angles.
Leaves of middle stem region in whorls of 68, sessile; blade
drying papery and glossy, narrowly oblanceolate or narrowly
elliptic-lanceolate, 316 13 mm, glabrous, retrorsely
aculeolate on veins and margins, base acute, apex acute; vein 1.
Inflorescences terminal and in upper leaf axils, with 2- to several-flowered cymes of 25 cm; axes glabrous, smooth; bracts
none or inconspicuous, 24 mm; pedicels 15 mm. Ovary ellipsoid, ca. 0.5 mm, glabrous. Corolla white, subcampanulate, 23
mm in diam., lobed for ca. 2/3; lobes 4, triangular, acute. Mericarps obovoid, ca. 1 1.52 mm, glabrous, smooth or granular.
Fl. and fr. JunAug.
Wet grasslands; ca. 2600 m. ?Sichuan, Xinjiang, ?Yunnan [Mongolia, Russia; C and SW Asia, Europe].
Galium uliginosum is a typical member of G. sect. Trachygalium
and closely related to the G. rivale group, represented in China by G.
karataviense. It differs from it by a much more slender habit and much
shorter corolla tubes. All other Chinese species of G. sect. Trachygalium
s.l. have rotate corollas.
We have seen specimens of Galium uliginosum from Xinjiang
only. In view of the more northerly distribution of the species, the indications for Sichuan and Yunnan (in FRPS 71(2): 258. 1999) may refer
to another taxon and should be verified.

62. Galium verum Linnaeus, Sp. Pl. 1: 107. 1753.

peng zi cai
Herbs, perennial, with rootstock and rhizomes. Stems
erect, (5)1570(120) cm tall, 4-angled, densely puberulent,
villosulous, or hirtellous to rarely glabrous and smooth. Leaves
in middle stem region in whorls of more than 6 and up to 12,
sessile; blade drying papery to subleathery, often blackening,
adaxially rather shiny, abaxially paler, linear to linear-oblong,
1030(50) 12(2.5) mm, adaxially glabrous to densely
hairy, smooth to sparsely aculeolate, abaxially usually densely
puberulent to tomentose, rarely glabrescent or glabrous, base
acute to cuneate, margins usually strongly revolute and antrorsely aculeolate, apex acute and shortly mucronate with tip to
1.5 mm; vein 1. Inflorescences thyrsoid or paniculate, terminal
and axillary cymes few to many flowered, rather dense and

RUBIACEAE

140

bracteose; axes usually densely puberulent, hirtellous, rarely

glabrous and smooth; bracts leaflike, 1.53 mm; pedicels 13
mm. Flowers fragrant, hermaphroditic. Ovary ellipsoid to subglobose, 0.50.8 mm, glabrous to densely hairy with straight
trichomes. Corolla yellow to white, rotate, ca. 3 mm in diam.,
glabrous, lobed for 3/4 or more; lobes 4, lanceolate-oblong,
subobtuse, acute to apiculate. Mericarps ellipsoid and laterally
flattened, 1.52 mm, glabrous to densely hispidulous with
straight trichomes. Fl. AprAug, fr. MayOct.
Mountains, grasslands, meadows, river beaches, open fields, ditch
sides, streamsides, wet places, forests, thickets, valleys; near sea level to
4100 m. Anhui, Gansu, Hebei, Heilongjiang, Henan, Hubei, Jiangsu,
Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong,
Shanxi, Sichuan, Xinjiang, Xizang, Zhejiang [India, Japan, Kashmir,
Kazakhstan, Korea, Mongolia, Pakistan, Russia, Turkmenistan, Uzbekistan; SW Asia, Europe; adventive in North America and elsewhere].
Galium verum is used medicinally and ranks among the most
commonly collected species of Galium in China, along with G. bungei,
G. spurium, and G. hoffmeisteri. Together with closely related taxa (as G.
saurense and G. consanguineum in the Chinese flora), it forms an extremely variable polyploid complex with 2x- and 4x-populations, which
is still very badly understood. Together with G. humifusum, with which
it can form a hybrid, it is placed into G. sect. Galium.
According to the considerable variation of Galium verum with
respect to habit, indumentum of leaves, ovaries, and fruit, and flower
color, Cufodontis (Oesterr. Bot. Z. 89: 216219. 1940) and subsequently W. C. Chen (in FRPS 71(2): 266269. 1999) have rather
schematically recognized a number of varieties, which often coexist in
one and the same population. Even if they do not correspond to natural
entities, they are provisionally treated here for reference, in particular as
to the range and adaptation to different habitats in China.

1a. Ovary and fruit hairy.

2a. Leaves pubescent and scabrous
adaxially .................................... 62e. var. tomentosum
2b. Leaves glabrous and smooth
adaxially ................................. 62f. var. trachycarpum
1b. Ovary and fruit glabrous.
3a. Corolla yellow.
4a. Leaves pubescent and
scabrous adaxially ........ 62g. var. trachyphyllum
4b. Leaves glabrous and
smooth adaxially.
5a. Plants to over 1 m tall;
leaves to 57 cm .......... 62a. var. asiaticum
5b. Plants to 45 cm tall;
leaves usually 1.53 cm .... 62h. var. verum
3b. Corolla pale yellow or white.
6a. Corolla pale yellow .......... 62c. var. leiophyllum
6b. Corolla white.
7a. Leaves pubescent and
scabrous adaxially .......... 62b. var. lacteum
7b. Leaves glabrous and
smooth adaxially ......... 62d. var. nikkoense
62a. Galium verum var. asiaticum Nakai, J. Jap. Bot. 15: 344.
1939.
chang ye peng zi cai
Galium verum subsp. asiaticum (Nakai) T. Yamazaki.
Plants stout, 50120 cm tall. Leaf blade to 57 cm, gla-

brous and smooth adaxially. Corolla yellow. Ovary and mericarps glabrous. Fl. and fr. JunAug.
Mountain grasslands, open fields, river beaches; below 1001700
m. Anhui, Gansu, Hebei, Heilongjiang, ?Henan, Hubei, Jiangsu, Jilin,
Liaoning, Nei Mongol, Shandong, Shanxi, Sichuan, Zhejiang [Japan,
Korea, Russia].
According to Yamazaki (Fl. Japan 3a: 240. 1993) Galium verum
subsp. asiaticum includes all Japanese forms of G. verum and differs
from the typical subspecies by longer leaves and hirsute (not minutely
pubescent) stems.

62b. Galium verum var. lacteum Maximowicz, Bull. Acad.

Imp. Sci. Saint-Ptersbourg 19: 283. 1874.
bai hua peng zi cai
Galium verum f. lacteum (Maximowicz) Nakai.
Leaf blade pubescent and scabrous adaxially. Corolla
white. Ovary and mericarps glabrous.
Wet places on mountains and in open fields; 5001000 m. Gansu,
Hebei, Heilongjiang, Jilin, Liaoning, Ningxia, Shaanxi [Japan, Korea].
Yamazaki (loc. cit.: 240) treated this as a form within Galium
verum subsp. asiaticum and listed G. verum f. nikkoense as a synonym
of G. verum f. lacteum.

62c. Galium verum var. leiophyllum Wallroth, Sched. Crit.

56. 1822.
dan huang peng zi cai
Corolla pale yellow. Ovary and mericarps glabrous. Fl.
JunJul.
Mountain grasslands, open fields; ca. 600 m. Hebei, Liaoning,
Shandong [?Japan; Europe].
Although W. C. Chen (loc. cit.: 268) included Japan in the distribution of this variety, its name is not mentioned in Fl. Japan.

62d. Galium verum var. nikkoense Nakai, J. Jap. Bot. 15: 347.
1939.
ri guang peng zi cai
Galium verum f. nikkoense (Nakai) Ohwi.
Leaf blade glabrous and smooth adaxially. Corolla white.
Ovary and mericarps glabrous.
Shandong [Japan].
Yamazaki (loc. cit.: 240) treated this variety as a synonym of
Galium verum f. lacteum.

62e. Galium verum var. tomentosum C. A. Meyer, Verz. Pfl.

Casp. Meer. 54. 1831.
mao peng zi cai
Galium verum f. tomentosum (C. A. Meyer) Nakai.
Leaf blade pubescent and scabrous adaxially. Ovary and
mericarps pilose. Fl. and fr. JunSep.
Forests on mountain slopes, farmland sides, grasslands; 4003100
m. Gansu, Hebei, Heilongjiang, Jilin, Liaoning, Nei Mongol, Qinghai,
Shanxi, Sichuan, Xinjiang [Japan].
The type of the name of this variety comes from SW Asia (S
Azerbaijan: Talysh) and may not fully conform to the Chinese popula-

RUBIACEAE

tions listed here. Although Nakai (Bot. Mag. (Tokyo) 34: 5051. 1920;
J. Jap. Bot. 15: 348. 1939) and W. C. Chen (loc. cit.: 269) included Japan in the distribution of this taxon, its name is not mentioned in Fl.
Japan. In the older literature, specimens with hairy ovaries and fruit,
corresponding to this and the following Galium verum var. trachycarpum, often have been called G. ruthenicum Willdenow.
With respect to the synonymy one has to consider the following:
Nakai (loc. cit. 1939) under Galium verum var. tomentosum Nakai,
comb. nov., cited Galium verum var. luteum f. tomentosum Nakai,
where (loc. cit. 1920) he cited G. verum var. c. Ledebour Fl. Ross. II. p.
415, where Ledebour wrote c. caule fructibusque tomento denso vestitis. C. A. Meyer l.c. Hab. in provinciis caucasicis [in m. Talsch, alt.
1100 hexap. (C. A. Meyer), giving the reference to Meyer on p. 414 as
Ind. cauc. p. 54, i.e., Verz. Pfl. Casp. Meer. 54. 1831, where Meyer
wrote [var.] tomentosum. caule et mericarpiis tomento denso vestitis.
In cacumine montium Talsch prope pagum Drych, in rupestribus siccis
sterilissimis (alt. 1100 hexap.) It is evident, therefore, that Nakai in
1920 was not publishing the name of a new taxon but a status novus at
the rank of forma with an indirect reference (allowed before 1953;
Vienna Code, Art. 33.2) to the basionym, i.e., Meyers varietal name.

62f. Galium verum var. trachycarpum Candolle, Prodr. 4:

603. 1830.
mao guo peng zi cai
Galium verum var. lasiocarpum Ledebour.
Leaf blade glabrous and smooth adaxially. Ovary and mericarps pilose. Fl. and fr. JunSep.
Forests, thickets, or grasslands on mountain slopes, streamsides,
open fields, river beaches; 1003900 m. Gansu, Hebei, Heilongjiang,
Henan, Jilin, Liaoning, Nei Mongol, Qinghai, Shanxi, Sichuan, Xinjiang, Xizang, Zhejiang [Japan, Korea, Russia; Europe].
This variety (including its synonym) was included by Ehrendorfer
et al. (Fl. Iranica 176: 200. 2005) in Galium verum f. subpubescens
Sergievskaya, to which it corresponds. Yamazaki (loc. cit.: 240) recognized G. verum f. album Nakai for white-flowered plants of this variety.

62g. Galium verum var. trachyphyllum Wallroth, Sched. Crit.

56. 1822.
cu cao peng zi cai
Leaf blade pubescent and scabrous adaxially. Corolla yellow. Ovary and mericarps glabrous. Fl. MayAug, fr. AugSep.
Forests or grasslands on mountain slopes or in valleys, open
fields, river beaches; 3004100 m. Anhui, Gansu, Hebei, Heilongjiang,
Henan, Jiangsu, Jilin, Liaoning, Nei Mongol, Ningxia, Qinghai, Shaanxi, Shandong, Shanxi, Sichuan, Xinjiang [Korea; Europe].

62h. Galium verum var. verum

() peng zi cai (yuan bian zhong)
Galium luteum Lamarck; G. verum var. leiocarpum Ledebour; G. verum var. luteum (Lamarck) Nakai.
Plants to 45 cm tall. Leaf blade usually 1.53 cm, glabrous
and smooth adaxially. Corolla yellow. Ovary and mericarps gla-

141

brous. Fl. AprAug, fr. MayOct.

Mountains, river beaches, open fields, ditch sides, grasslands,
meadows, thickets, forests; near sea level to 4000 m. Gansu, Hebei,
Heilongjiang, Jilin, Liaoning, Nei Mongol, Qinghai, Shandong, Shanxi,
Sichuan, Xinjiang, Xizang [India, Japan, Korea, Pakistan; SW Asia,
Europe; adventive in North America and elsewhere].
The citation here of Galium luteum, a long-recognized synonym
of G. verum based on a type from France, follows W. C. Chen (loc. cit.:
266). Ehrendorfer et al. (loc. cit.: 200201) recognized two forms within G. verum subsp. verum, separated by density and distribution of pubescence: f. verum with glabrous ovaries and fruit vs. f. subpubescens
with pubescent ovaries and fruit. Thus, their f. subpubescens corresponds to var. tomentosum and var. trachycarpum of Cufodontis
(loc. cit.: 216219) and W. C. Chen (loc. cit.: 268269).

63. Galium yunnanense H. Hara & C. Y. Wu, J. Jap. Bot. 61:

74. 1986.
dian la la teng
Galium elegans Wallich var. angustifolium Cufodontis; G.
elegans var. nemorosum Cufodontis.
Herbs, perennial, procumbent to scrambling or matted, up
to 1 m, from slender rhizomes. Stems glabrescent and smooth
to sparsely or moderately pilose to villous or retrorsely hispid,
nodes more densely hairy, angles 4, usually thickened. Leaves
in whorls of 4, subsessile; blade drying membranous, green,
elliptic, ovate-lanceolate, or lanceolate, 550 315 mm,
length/breadth index normally above 2.5, adaxially hispidulous
to hirsute, abaxially glabrescent to densely pilose and usually
glandular-punctate, base cuneate to obtuse, margins sparsely to
densely pilose or antrorsely ciliate, apex acute to acuminate and
often mucronulate; principal veins 3, palmate. Inflorescences
terminal and in axils of uppermost leaves, paniculate, many
flowered, 212 cm, diffusely branched; peduncles pilose to
glabrescent; bracts inconspicuous, ligulate to ovate, 1.52.5
mm, often lacking upward; pedicels 2.57 mm. Flowers dioecious, polygamous, or ?hermaphroditic. Ovary obovoid, ca. 0.5
mm, densely appressed hairy. Corolla white, rotate, 11.5 mm
in diam.; lobes 4, ovate, subacute. Mericarps ovoid, 1.52 mm,
with dense, uncinate, stiff and spreading, basally white to apically brown trichomes ca. 0.8 mm. Fl. and fr. JulNov.
Forests, meadows on mountains, riversides, streamsides; 700
3300 m. Gansu, Guangxi, Guizhou, Hunan, Sichuan, Yunnan.
As noted in the protologue, Galium yunnanense is similar to G.
elegans and comprises plants that previously have been included in a
more broadly circumscribed G. elegans. In particular, the two varieties
of G. elegans described by Cufodontis in 1940 and cited as synonyms
above now key to G. yunnanense. In spite of its variability and occasional forms approaching G. bungei (see there), the specific separation
of G. yunnanense from G. elegans by Chen (Acta Phytotax. Sin. 28: 301.
1990) seems well justified. It reduces the morphological variation
within G. elegans and results in a much clearer circumscription of the
two taxa.

29. GARDENIA J. Ellis, Philos. Trans. 51: 935. 1761, nom. cons., not Colden (1756).
zhi zi shu
Chen Tao (); Charlotte M. Taylor
Shrubs or rarely trees, sometimes with short shoots (Gardenia angkorensis, G. sootepensis), unarmed or with short shoots

142

RUBIACEAE

sometimes spinescent, with buds and young stem apices often resinous. Raphides absent. Leaves opposite or rarely ternate, sometimes clustered at stem apices, often with domatia; stipules persistent or deciduous, united shortly around stem or united completely
into a conical cap (i.e., calyptrate), triangular or when united into a cap then splitting along one side. Inflorescences pseudoaxillary
and/or terminal, fascicled to cymose and several flowered or reduced to 1 flower, pedunculate to sessile, bracteate. Flowers sessile to
pedicellate, bisexual, monomorphic, often showy. Calyx with ovary portion well developed and often longitudinally ridged; limb 5
8-lobed or sometimes fused into a tube or conical cap that splits irregularly as corolla elongates (i.e., spathaceous), often well
developed. Corolla white to cream, salverform or funnelform, glabrous or variously pubescent inside; lobes 512, convolute in bud.
Stamens 512, inserted in corolla throat, included or partially exserted; filaments very short or reduced; anthers dorsifixed. Ovary 1celled, ovules numerous on 26 parietal placentas; stigma clavate or 2-lobed, included or exserted. Fruit generally yellow to orange,
red-orange, or brown with pulp usually orange, baccate, leathery or fleshy, ellipsoid to subglobose, smooth or with longitudinal
ridges, with calyx limb usually persistent or sometimes upper part tardily deciduous; seeds numerous, medium-sized, ellipsoid,
compressed, embedded in pulp; testa leathery or membranous; endosperm usually corneous; embryo small or medium-sized; cotyledons broad, leaflike.
About 60200 or 250 species: tropical and subtropical regions of Africa, Asia, Madagascar, and Pacific islands; five species (one endemic) in
China.
The persistent calyx lobes apparently enlarge markedly as the fruit develop in many species; this may be confusing when the enlarged fruiting
calyx lobes are compared with descriptions of flowering calyx lobes. The flowers are often nocturnal and are usually strongly sweetly fragrant with an
odor of, well, gardenia. Several species from Asia and the Pacific are occasionally cultivated, but Gardenia jasminoidesnative to our regionis
very popular worldwide for its flowers.

1a. Leaf blade puberulent or pilosulous to glabrous adaxially, densely tomentose abaxially; stipules and calyx
limb each fused into a conical cap then splitting along one side; fruit with spathaceous upper portion of
calyx limb deciduous ................................................................................................................................................. 4. G. sootepensis
1b. Leaf blade glabrous to puberulent or pilosulous; stipules united in basal portion with apical portions free
or fused into a conical cap, calyx regularly lobed; fruit with calyx lobes persistent.
2a. Leaf blade 1.54 12.5 cm, obovate or spatulate, obtuse to rounded at apex; calyx lobes 45 mm in
flower, 58 mm in fruit ...................................................................................................................................... 1. G. angkorensis
2b. Leaf blade 325 0.48 cm, acute, acuminate, or obtuse at apex; calyx lobes 430 mm in flower, to
40 mm in fruit.
3a. Leaves ternate or sometimes opposite on a few nodes, with blade narrowly lanceolate or
linear-lanceolate, 0.42.3 cm wide, without domatia; fruit ellipsoid-oblong to ellipsoid,
1.52.5 11.3 cm, smooth or with 58 weak to developed longitudinal ridges ...................................... 5. G. stenophylla
3b. Leaves opposite or sometimes ternate on a few nodes, with blade oblanceolate, obovate-oblong,
elliptic-oblong, lanceolate-oblong, obovate, or elliptic, 1.58 cm wide, without or usually with
domatia; fruit ovoid-ellipsoid, subglobose, or ellipsoid, 1.57 1.22 cm, smooth or with
59 weak to well-developed longitudinal ridges.
4a. Trees; calyx lobes 47 mm in flower; corolla tube ca. 15 mm, shorter than lobes; fruit with 5
weak to well-developed longitudinal ridges ......................................................................................... 2. G. hainanensis
4b. Shrubs; calyx lobes 1030 mm in flower; corolla tube 3050 mm, equal to or usually longer
than lobes; fruit with 59 well-developed longitudinal ridges ............................................................. 3. G. jasminoides
1. Gardenia angkorensis Pitard in Lecomte, Fl. Indo-Chine 3:
252. 1923.
chi ye zhi zi
Shrubs, 13 m tall, with short shoots; branches terete,
glabrous, becoming grayish white. Leaves opposite, usually
crowded at ends of short shoots; petiole 14 mm, puberulent to
glabrous; blade drying stiffly papery, obovate or spatulate, 1.5
4 12.5 cm, adaxially glabrous and shiny, abaxially sparsely
puberulent to glabrous, base cuneate to acute, apex obtuse to
rounded; secondary veins 68 pairs, in abaxial axils with pubescent foveolate domatia; stipules united shortly around stem,
broadly triangular to ligulate, 23 mm, glabrous, obtuse to
rounded. Flowers solitary, terminal on short shoots, sessile or
subsessile. Calyx puberulent to glabrescent; ovary portion obconical, weakly ridged, 78 mm; limb lobed nearly to base;
lobes 6, narrowly spatulate, 45 mm, obtuse. Corolla outside
glabrous; tube 1315 mm, somewhat funnelform; lobes 6, spatulate, ca. 15 mm, obtuse. Fruiting pedicels to 3 mm. Berry ellip-

soid to subglobose, 1518 1015 mm, smooth to weakly

ridged, with persistent calyx lobes 78 mm; seeds ca. 5 34
mm. Fr. AugDec.
Forests or thickets at streamsides in valleys or on mountain slopes.
Hainan [Cambodia].

2. Gardenia hainanensis Merrill, Lingnan Sci. J. 9: 43. 1930.

hai nan zhi zi
Trees, 312 m tall; branches with internodes congested to
developed, compressed to terete, densely puberulent to glabrous, resinous at apices with usually distalmost several internodes coated with dried resin, becoming straw-brown. Leaves
opposite; petiole 0.21 cm, puberulent to glabrous; blade drying
thinly leathery, bright to dull or yellowed green adaxially, similar but darker or paler abaxially, obovate-oblong, elliptic-oblong, or oblanceolate, 519.5 28 cm, adaxially shiny and
glabrous, abaxially glabrous to puberulent, base cuneate to
acute, apex acute or shortly acuminate with tip often ultimately

RUBIACEAE

obtuse; secondary veins 1015 pairs, in abaxial axils with pilosulous domatia; stipules calyptrate, conical, 410 mm, splitting for 1/23/4 their length, puberulent to usually glabrous.
Flowers solitary, terminal or pseudoaxillary; peduncle 0.40.8
cm, densely puberulent. Calyx puberulent, usually covered with
resin; ovary portion broadly obconical to ellipsoid, 5-ridged, 5
6 mm; limb with tubular portion 34 mm; lobes 5, oblonglanceolate to linear or spatulate, 47 mm, 11.6 mm wide at
base, strongly keeled, narrowed then ultimately rounded at
apex. Corolla outside apparently glabrous, often covered with
resin; tube ca. 15 mm, somewhat funnelform; lobes 5, obovateoblong to elliptic, 2330 810 mm, acute to obtuse and apiculate. Fruiting peduncles to 2 cm. Berry yellow, subglobose or
ovoid-ellipsoid to ellipsoid, 1.63.3 1.51.6 cm, with 5 weak
to developed longitudinal ridges, with persistent calyx limb. Fl.
Apr, fr. MayOct.
Forests at streamsides in valleys or on mountain slopes; below
1001200 m. Guangxi (Shangsi), Hainan.

This is one of the most commonly collected species of Rubiaceae

in China. It is quite variable morphologically especially in leaf size,
calyx lobe size, and corolla size. Several varieties have been recognized
for Chinese plants (e.g., Qiu & Zhong, Fl. Zhejiang 6: 105. 1986) but
are not clearly separated or widely accepted outside this region. The
varieties recognized by W. C. Chen (in FRPS 71(1): 332335. 1999) are
outlined below for reference.

1a. Corolla doubled; flowers sterile, plants

not setting fruit; cultivated plants ......... 3a. var. fortuneana
1b. Corolla simple; flowers fertile, plants
setting fruit; wild and occasionally
cultivated plants .................................. 3b. var. jasminoides
3a. Gardenia jasminoides var. fortuneana (Lindley) H. Hara,
Enum. Sperm. Jap. 2: 15. 1952 [fortuniana].
bai chan
Gardenia florida Linnaeus var. fortuneana Lindley, Edwardss Bot. Reg. 32: t. 43. 1846 [fortuniana].

3. Gardenia jasminoides J. Ellis, Philos. Trans. 51: 935. 1761.

zhi zi
Shrubs, 0.33 m tall; branches terete to flattened, with internodes developed to shortened, glabrescent or usually densely
puberulent to pilosulous, becoming gray to grayish white, with
buds resinous and distalmost internodes often covered with
resin. Leaves opposite or rarely ternate, subsessile to petiolate;
petiole to 0.5(1) cm, densely puberulent or shortly pilosulous
to glabrous; blade drying thinly leathery to stiffly papery,
oblong-lanceolate, obovate-oblong, obovate, oblanceolate,
or elliptic, 325 1.58 cm, adaxially shiny and glabrous or
sometimes puberulent on principal veins, abaxially puberulent
or pilosulous to glabrous, base cuneate to acute, apex acute to
acuminate or obtuse then abruptly long acuminate; secondary
veins 815 pairs, in abaxial axils often with pilosulous domatia;
stipules calyptrate, cylindrical, 413 mm, splitting for ca. 3/4
their length, densely puberulent to glabrous. Flower solitary,
terminal; peduncle 110 mm, puberulent or pilosulous to glabrous. Calyx puberulent or pilosulous to glabrous; ovary portion obconic or obovoid, 58 mm, with (5 or)6(8) weak to
developed longitudinal ridges; limb with basal tubular portion
35 mm; lobes (5 or)6(8), lanceolate or linear-lanceolate to
spatulate, 1030 14 mm, often strongly keeled, acute. Corolla white to pale yellow, simple or in cultivation sometimes
doubled, outside glabrous; tube 3050 46 mm, cylindrical,
in throat pilose; lobes (5 or)6(8) or numerous when doubled,
obovate or obovate-oblong, 1540 628 mm, obtuse to
rounded. Fruiting peduncles apparently not much elongating.
Berry yellow or orange-yellow, ovoid, subglobose, or ellipsoid,
1.57 1.22 cm, with 59 longitudinal ridges, with persistent
calyx lobes to 40 6 mm; seeds suborbicular, weakly angled,
ca. 3.5 3 mm. Fl. MarJul, fr. MayFeb.
Thickets and forests at streamsides, on mountain slopes or hills, or
in valleys or fields; near sea level to 1500 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hebei, Hubei, Hunan, Jiangsu, Jiangxi, Shandong, Sichuan, Taiwan, Yunnan, Zhejiang; cultivated in Gansu,
Hebei, Shanxi [Bhutan, Cambodia, India, Japan, N Korea, Laos, Nepal,
Pakistan, Thailand, Vietnam; cultivated in Africa, Asia, Australia, Europe, North and South America, and Pacific islands].

143

Flowers with doubled corolla, sterile. Fl. throughout year.

Cultivated in gardens and green belts as an ornamental in provinces of S China (including Nanhai Zhudao) [cultivated worldwide, outdoors in tropical regions and indoors in temperate regions].

3b. Gardenia jasminoides var. jasminoides

() zhi zi (yuan bian zhong)
Gardenia florida Linnaeus, nom. illeg. superfl.; G. florida
f. oblanceolata Nakai; G. florida var. ovalifolia Sims; G. florida
f. simpliciflora Makino; G. grandiflora Loureiro; G. jasminoides
f. grandiflora (Loureiro) Makino; G. jasminoides var. grandiflora (Loureiro) Nakai; G. jasminoides var. longisepala (Masamune) F. P. Metcalf; G. jasminoides f. maruba (Siebold ex
Blume) Nakai ex Ishii; G. jasminoides var. maruba (Siebold ex
Blume) Nakai; G. jasminoides f. oblanceolata (Nakai) Nakai; G.
jasminoides f. ovalifolia (Sims) H. Hara; G. jasminoides var.
ovalifolia (Sims) Nakai; G. jasminoides var. radicans (Thunberg) Makino; G. jasminoides f. simpliciflora (Makino) Makino; G. jasminoides f. variegata (Carrire) Nakai; G. jasminoides
var. variegata (Carrire) Makino; G. maruba Siebold ex Blume;
G. radicans Thunberg; G. radicans var. variegata Carrire; G.
schlechteri H. Lveill (1911), not Bonati & Petitmengin (1907).
Flowers with simple corolla, fertile. Fl. MarJul, fr. May
Feb.
Thickets and forests at streamsides, on mountain slopes or hills, or
in valleys or fields; near sea level to 1500 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangsu, Jiangxi,
Shandong, Sichuan, Taiwan, Yunnan, Zhejiang; cultivated in Gansu,
Hebei, Shanxi [Cambodia, India, Japan, N Korea, Laos, Nepal, Pakistan, Vietnam; cultivated in Europe, North America, Pacific islands].

4. Gardenia sootepensis Hutchinson, Bull. Misc. Inform. Kew

1911: 392. 1911.
da huang zhi zi
Trees, 710 m tall, often with gelatinous secretions;
branches with both developed and shortened internodes, somewhat compressed to angled or subterete, densely puberulent,

RUBIACEAE

144

pilosulous, or tomentulose, becoming glabrescent. Leaves opposite; petiole 0.61.2 cm, puberulent or tomentulose; blade
drying papery or thinly leathery, obovate, obovate-elliptic,
broadly elliptic, or elliptic-oblong, 729 316 cm, adaxially
puberulent or pilosulous to glabrous, abaxially densely tomentose, base rounded to obtuse or cuneate, apex shortly acuminate
with tip acute or obtuse; secondary veins 1220 pairs, in abaxial
axils often with densely pilosulous domatia; stipules calyptrate,
conical, 0.51 cm, sericeous outside, densely puberulent or
tomentulose inside, apical portion triangular and caducous, basal portion truncate to broadly rounded and usually persisting
with leaves and sometimes becoming hardened. Flowers pseudoaxillary usually near branch apices, solitary; peduncle 11.5
cm, puberulent. Calyx densely puberulent to pilosulous externally; ovary portion ellipsoid, smooth, 56 mm; limb spathaceous, 1315 mm, splitting along one side for 2/33/4 of its
length, inside sericeous, often viscid or mucilaginous. Corolla
yellow or white, salverform; tube 5070 35 mm, cylindrical,
outside sparsely puberulent, inside glabrous; lobes 5, broadly
obovate, 4050 2030 mm, glabrous on both surfaces, obtuse
to acute. Berry ellipsoid or ellipsoid-oblong, 2.55.5 1.53.5
cm, puberulent, smooth or with 5 or 6 longitudinal lines or very
weak ridges, leathery to hard; seeds suborbicular, flattened, 34
mm in diam., foveolate. Fl. AprAug, fr. JunApr.
Forests at streamsides, at village margins, or on mountain slopes;
7001600 m. Yunnan [Laos, Thailand].
W. C. Chen (in FRPS 71(1): 335. 1999) described the flowers as

terminal on branchlets, but these are pseudoaxillary on all specimens

seen as described here.

5. Gardenia stenophylla Merrill, Philipp. J. Sci. 19: 678. 1922.

xia ye zhi zi
Shrubs, 0.53 m tall; branches generally slender, angled to
subterete, puberulent to glabrescent. Leaves opposite or ternate
with arrangement often variable on a branch, subsessile to
shortly petiolate; petiole to 5 mm, puberulent to glabrous; blade
drying thinly leathery, narrowly lanceolate, narrowly elliptic,
elliptic-spatulate, or linear-lanceolate, 312 0.42.3 cm, adaxially glabrous, abaxially puberulent to glabrous, base acute and
often decurrent, margin thickened and often thinly revolute,
apex acute to obtuse; secondary veins 913 pairs, in abaxial
axils without domatia; stipules calyptrate, conical, 710 mm,
splitting along one side for 3/4 or more of length, glabrous.
Flowers solitary, pseudoaxillary or terminal; peduncles 56
mm, glabrous. Calyx glabrous; ovary portion ellipsoid to cylindrical, 56 mm, longitudinally 58-ridged; limb with basal tubular portion 46 mm; lobes 58, linear to narrowly lanceolate,
715 mm, keeled, acute. Corolla outside glabrous; tube cylindrical, 3565 34 mm; lobes 58, oblong-obovate, 2535
1015 mm, obtuse. Fruiting pedicels to 2 cm. Berry yellow or
orange-red, ellipsoid-oblong to ellipsoid, 1.52.5 11.3 cm,
with 58 weak to developed longitudinal ridges, with persistent
calyx lobes to 2 cm; seeds 23 mm. Fl. AprAug, fr. MayJan.
Forests or thickets at streamsides in valleys, fields at riversides;
below 100800 m. Anhui, Guangdong, Guangxi, Hainan, Zhejiang
[Vietnam].

30. GEOPHILA D. Don, Prodr. Fl. Nepal. 136. 1825, nom. cons., not Bergeret (1803).
ai di cao shu
Chen Tao (); Charlotte M. Taylor
Geocardia Standley.
Herbs, perennial, unarmed; stems creeping and rooting at nodes. Raphides present. Leaves opposite, usually broadly ovate to
cordate and long petiolate, without domatia; stipules persistent, interpetiolar, generally triangular, entire [or 2-lobed]. Inflorescences
terminal or pseudoaxillary, cymose to capitate, few to several flowered, pedunculate, bracteate. Flowers sessile or subsessile,
bisexual, apparently monomorphic. Calyx limb 47-lobed. Corolla white, funnelform, inside pubescent in throat; lobes 47, valvate
in bud. Stamens 47, inserted in corolla tube, included or partly exserted; filaments developed; anthers dorsifixed. Ovary 2-celled,
ovules 1 in each cell, erect, basal; stigma 2-lobed, included or exserted. Fruit orange or red [or sometimes blue or black], drupaceous,
globose to ellipsoid, fleshy, with calyx limb persistent; pyrenes 2, 1-celled, each with 1 seed, hard, plano-convex and sometimes
longitudinally twisted, dorsally (i.e., abaxially) smooth to ridged, ventrally (i.e., adaxially) with a median ridge and 2 submarginal
longitudinal grooves; seeds medium-sized; testa membranous; endosperm corneous.
About 30 species: widespread in tropical Africa, Asia, Central, North, and South America, and Madagascar; one species in China.

1. Geophila repens (Linnaeus) I. M. Johnston, Sargentia 8:

281. 1949.
ai di cao
Rondeletia repens Linnaeus, Syst. Nat., ed. 10, 2: 928.
1759; Geophila herbacea (Jacquin) K. Schumann; G. reniformis
D. Don; Psychotria herbacea Jacquin .
Herbs, perennial, creeping; stems quadrate to subterete,
puberulent or hirtellous to glabrescent, vegetative creeping, 40
cm or longer, reproductive ascending, to 15 cm tall. Petiole 15

cm, densely puberulent to pilose or hirtellous often in lines; leaf

blade drying membranous to papery and sometimes yellowish
green, ovate to suborbicular, 13 13 cm, glabrescent on both
surfaces, base cordulate to cordate, margins often undulate,
apex obtuse to rounded; secondary veins 2 or 3 pairs, usually
grouped in basal half of blade; stipules broadly ovate, 12 mm.
Inflorescence with flowers 13 in umbelliform cymules, puberulent to hirtellous or glabrescent; peduncle 14 cm; bracts
linear to narrowly triangular, 1.53 mm. Calyx glabrous to puberulent; ovary portion ellipsoid, 1.52 mm; limb deeply lobed;
lobes 4, linear-lanceolate, 23 mm, ciliate. Corolla pilosulous to

RUBIACEAE

glabrescent outside; tube ca. 8 mm; lobes 4, ovate or ovate-lanceolate, ca. 3 mm, acute. Drupes red, subglobose to ovoid, 46
mm in diam., pilosulous to glabrescent; pyrenes dorsally weakly
rugulose. Fl. JulSep, fr. SepDec.

145

Forest and trail margins, ravines, damp places; 100600 m.

Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [widespread
in tropical Africa, Antilles, Asia, Central America, Madagascar, North
America (Mexico), and South America].

31. GUETTARDA Linnaeus, Sp. Pl. 2: 991. 1753.

hai an tong shu
Chen Tao (); Charlotte M. Taylor
Shrubs or trees, sometimes polygamo-dioecious, unarmed. Raphides absent. Leaves opposite or rarely ternate, decussate, usually with domatia; tertiary and/or quaternary venation often arranged in regular, often small rectangles or squares; stipules caducous,
interpetiolar, generally triangular. Inflorescences axillary, cymose with axes often markedly dichotomous or scorpioid, several to
many flowered, pedunculate with peduncle usually elongated [to sessile], bracts reduced [to well developed]. Flowers sessile to
shortly pedicellate, unisexual or bisexual and monomorphic or at least sometimes distylous, sweetly fragrant. Calyx limb tubular or
subcampanulate, truncate or irregularly denticulate. Corolla white or pink, salverform with tube infrequently curved, glabrous to
variously pubescent inside; lobes 49, obtuse, imbricate (and quincuncial) in bud. Stamens 49, inserted in corolla tube, included;
filaments short or reduced; anthers dorsifixed. Ovary 49-celled, ovules 1 in each cell, pendulous from apical placentas, anatropous,
with funicle thickened; stigma capitate, included. Fruit red, purple, black, or rarely green, drupaceous, thinly fleshy, oblate (i.e.,
depressed globose), ellipsoid, or subglobose, with calyx limb persistent; pyrene 1, 49-celled with 1 seed in each cell, oblate, ellipsoid, or subglobose, often 49-angled or -grooved, with preformed germination pore at apex of each cell; seeds medium-sized, ellipsoid, straight or curved; testa membranous; endosperm absent or scanty; embryo cylindrical or compressed; cotyledons small; radicle
ascending.
About 6080 species: tropical forests, most in tropical America and Pacific region, one species widespread on coasts of Indian Ocean and E
Pacific Ocean; one species in China.

1. Guettarda speciosa Linnaeus, Sp. Pl. 2: 991. 1753.

hai an tong
Small trees, 38 m tall; bark becoming black, smooth or
often lenticellate; branchlets rather stout, densely strigillose to
velutinous-tomentulose usually becoming glabrescent. Petiole
stout, 1.55 cm, densely strigillose or velutinous; leaf blade
drying thinly to stiffly papery, broadly obovate or broadly elliptic, 1120 818 cm, adaxially glabrescent, usually shiny, and
often rugulose, abaxially densely tomentulose or strigillose to
glabrescent, base obtuse, rounded, subcordate, or shortly cordate, apex obtuse or rounded sometimes with a short tip 35
mm; secondary veins 711 pairs, weakly but regularly looping
to connect; stipules ovate or lanceolate, 611 mm, moderately
to densely strigillose to strigose sometimes becoming glabrescent, obtuse to rounded. Inflorescences produced in with or
often below leaves, subcapitate to congested-cymose, densely
velutinous-tomentulose; peduncles 312 cm; branched portion
1.53 23.5 cm, axes scorpioid; bracts ovate, 57 mm, cadu-

cous. Flowers sessile. Calyx densely velutinous-tomentulose to

strigillose; ovary portion obconic to cupulate, 22.5 mm; limb
tubular, 23.5 mm, truncate. Corolla white, outside densely
velutinous-tomentulose to strigillose; tube 2.53 cm, inside
glabrous except sericeous in throat; lobes 7 or 8, obovate, 810
mm, obtuse to rounded. Drupes apparently green at maturity,
oblate, 23 cm in diam., sparsely strigillose or tomentulose to
eventually glabrescent; pyrene included in fibrous mesocarp. Fl.
AprJul.
Thickets on sandy and limestone coasts; sea level to near sea
level. Guangdong, Hainan, Taiwan [Borneo, India, Indonesia, Japan,
Malaysia, Philippines, Singapore, Sri Lanka, Thailand; coastal E Africa,
Australia, Madagascar, Pacific islands].
The flowers are nocturnal and open for only one night (Puff et al.,
Rubiaceae of Thailand, 130. 2005). The fruit of this species float and are
dispersed by water, and apparently they are green when mature. This
species is known to be distylous in Polynesia and E Africa (Bridson &
Verdcourt, Fl. Trop. E. Africa, Rub. (Pt. 2), 416. 1988).

32. GUIHAIOTHAMNUS H. S. Lo, Bull. Bot. Res., Harbin 18: 279. 1998.
gui hai mu shu
Chen Tao (); Charlotte M. Taylor
Subshrubs, unarmed; roots woody and stout. Raphides presumably absent. Leaves opposite, often congested or apparently fascicled at stem apices, without domatia; stipules caducous, apparently interpetiolar, triangular. Inflorescences terminal or pseudoaxillary, congested-cymose or subcapitate, several flowered, sessile to shortly pedunculate, bracteate with bracts linear. Flowers shortly
pedicellate or subsessile, presumably bisexual, with biology unknown. Calyx limb deeply 5-lobed. Corolla pink, salverform with
tube slender and abruptly expanded in throat, inside pubescent; lobes 5, convolute in bud. Stamens 5, inserted in corolla throat,
partially exserted; filaments short to reduced. Ovary 2-celled, ovules numerous in each cell on fleshy, peltate, axile placentas; stigma

RUBIACEAE

146

2-lobed with lobes capitate, exserted. Fruit purplish red, baccate, fleshy, subglobose, with calyx lobes tardily deciduous; seeds numerous, small, angled, reticulate.
One species: China.
The relationships of this genus and some of its morphological details are unknown. The presence or absence of raphides has not been explicitly
noted; absence is here inferred based on H. S. Los (in FRPS 71(1): ix. 1999) classification of this genus in Rondeletieae, but the one specimen seen
has scattered raphidelike structures deep within some intact tissues.

1. Guihaiothamnus acaulis H. S. Lo, Bull. Bot. Res., Harbin

18: 280. 1998.
gui hai mu
Plants 37 cm tall; stems quadrangular, densely villosulous, becoming glabrescent. Petiole 520 mm, villosulous; leaf
blade drying papery and grayish brown, broadly elliptic-oblong
to obovate, 4.511 2.57.5 cm, villosulous on both surfaces
or glabrescent adaxially, base cuneate to truncate and often
shortly decurrent, apex rounded or broadly obtuse and shortly
cuspidate; secondary veins 812 pairs, slender; stipules 12

mm, obtuse and partially reflexed. Inflorescences densely villosulous; bracts 1.51.7 mm. Calyx pilosulous, villosulous, or
glabrous; hypanthium portion obovoid-ellipsoid, ca. 1.7 mm;
lobes linear-lanceolate to narrowly triangular, 1.72 mm, acute
to acuminate. Corolla pale red, glabrous outside; tube ca. 19
mm, ca. 1 mm in diam. at base, 33.5 mm in diam. at throat,
sparsely villous inside; lobes elliptic or ovate-elliptic, ca. 3 mm,
rounded. Fruit purplish red, 2.53 mm in diam.; seeds 0.20.3
mm. Fl. Apr, fr. Jul.
Shady rocks; 100600 m. Guangxi (Rongshui).

33. HALDINA Ridsdale, Blumea 24: 360. 1979.

xin ye mu shu
Chen Tao (); Charlotte M. Taylor
Large trees, unarmed; buds flattened with stipules erect and pressed to each other. Raphides absent. Leaves opposite, decussate,
with domatia; stipules caducous, interpetiolar, ovate to oblong, entire. Inflorescences axillary, capitate with 1 or 2(5) fasciculate
globose heads, many flowered, pedunculate, bracteate; peduncles articulate and bracteate near top; bracteoles spatulate to spatulateclavate. Flowers sessile, bisexual, monomorphic. Calyx limb 5-lobed. Corolla pale to dark yellow, salverform, glabrous inside; lobes
5, imbricate in bud. Stamens 5, inserted in upper part of corolla tube, partially exserted; filaments reduced; anthers basifixed. Ovary
2-celled, ovules numerous in each cell, pendulous on axile placentas attached to upper third of septum; stigma ovoid to subglobose,
exserted. Fruiting heads globose, with fruit rather tightly packed. Fruit capsular, obconic to ellipsoid-oblong, septicidally then loculicidally dehiscent into 2 valves separating from base to apex and from persistent septum, cartilaginous to stiffly papery, with calyx
limb persistent on persistent septum; seeds numerous, medium-sized, ovoid to fusiform, slightly flattened, shortly winged on ends.
One species: Cambodia, China, India, Nepal, Sri Lanka, Thailand, Vietnam.
Ridsdale (loc. cit.) and Puff et al. (Rubiaceae of Thailand, 52. 2005) described the aestivation of the corolla lobes as, respectively, valvate but
strongly imbricate at apex and valvate in bud (but strongly imbricate at the apex); they also both describe the fruit as separating into 4 valves, but
only 2 valves have been seen on the specimens studied.

1. Haldina cordifolia (Roxburgh) Ridsdale, Blumea 24: 361.

1979.
xin ye mu
Nauclea cordifolia Roxburgh, Pl. Coromandel 1: 40. 1795;
Adina cordifolia (Roxburgh) J. D. Hooker ex B. D. Jackson.
Trees, deciduous, 730(40) m tall; trunk often buttressed
and fluted at base, with outer bark reddish brown, inner bark
dark red to brown; branches stout, with well-developed petiole
scars. Petiole 212 cm, densely tomentulose to pilosulous; leaf
blade drying thinly leathery, brown adaxially, and pale to yellowish green abaxially, broadly ovate, (5)816(25) (5)8
16(20) cm, adaxially sparsely hirtellous to puberulent or glabrescent, abaxially densely puberulent, tomentulose, pilosulous,
or glabrescent and with epidermis bullate inside areoles of quaternary veins, base cordate, apex acute to acuminate; secondary
veins 610 pairs, usually divaricately branched half way along

their length, sometimes with pilosulous domatia; stipules spatulate, ovate, or oblong-oblanceolate, 1012 512 mm, usually strongly keeled, densely strigillose to pilosulous, broadly
rounded. Inflorescence densely tomentulose to pilosulous; peduncle 26(10) cm, at articulation with 2 bracts 12 mm,
broadly rounded; flowering heads 58 mm in diam. across
calyces, ca. 20 mm in diam. across corollas; bracteoles linear to
clavate, ca. 2 mm. Calyx densely strigillose; ovary portion 12
mm, densely sericeous, surrounded at base by a dense ring of
trichomes 12 mm; limb deeply lobed, lobes narrowly triangular to linear, 1.31.8 mm, at apex clavate. Corolla outside
densely strigillose; tube 56 mm; lobes elliptic-oblong, 12
mm, obtuse to rounded. Stigma ovoid, ca. 0.2 mm, exserted for
57 mm. Fruiting heads 1015 mm in diam. Capsules 45 mm,
densely strigillose or glabrescent near base; seeds 23.5 0.51
mm. Fl. spring and summer.
Tropical rain forests; 3001000 m. Yunnan [Cambodia, India,
Nepal, Sri Lanka, Thailand, Vietnam].

RUBIACEAE

147

34. HAMELIA Jacquin, Enum. Syst. Pl. 2, 16. 1760.

chang ge mu shu
Chen Tao (); Charlotte M. Taylor
Duhamelia Persoon; Tangaraca Adanson.
Shrubs, unarmed. Raphides absent. Leaves opposite or verticillate, usually with domatia; stipules persistent, interpetiolar, triangular. Inflorescences terminal, cymose with axes often helicoid and secund, many flowered, bracteate or bracts reduced. Flowers sessile to pedicellate, bisexual, monomorphic. Calyx limb 5-lobed. Corolla red to orange or yellow, tubular [to ventricose or funnelform], glabrous inside; lobes 5, imbricate in bud. Stamens 5, inserted at base of corolla tube, included or partially exserted; filaments
short; anthers dorsifixed, 2-lobed at base, with connective flattened and prolonged at apex. Ovary 5-celled, ovules numerous in each
cell on axile placentas; stigmas 15, linear to clavate, included or exserted. Fruit becoming red then purple-black, baccate, fleshy to
juicy, ellipsoid to ovoid or subglobose, with calyx limb persistent; seeds numerous, small, irregularly angled to lenticular; testa membranous, reticulate.
Sixteen species: neotropical, from S United States and Mexico to Argentina; one species (introduced) in China.
H. S. Lo (in FRPS 71(1): 388. 1999) described the flower arrangement with short pedicels or subsessile, but the structures interpreted there as
pedicels are considered by one of us (C. M. Taylor) and by some other authors to be inflorescence axes and the flowers thus are sessile. Lo described
the stipules as multifid or bristlelike, but this condition is not otherwise known from the genus and has not been seen on any Chinese specimens. He
gave the number of calyx lobes, corolla lobes, and stamens as 46, but this has not been seen; these structures are 5 in all Hamelia species known. The
corollas were described as villous in throat, but all known species of Hamelia have corollas that are glabrous inside; as well, the corolla was described
as campanulate in shape, but the cultivated species has rather narrowly tubular flowers. The anthers were described as basifixed and entire at base, but
other authors all found them dorsifixed and sagittate at base.

1. Hamelia patens Jacquin, Enum. Syst. Pl. 16. 1760.

chang ge mu
Shrubs, deciduous, 1.54 m tall; branches angled to subterete, often becoming red, hirtellous or pilosulous to glabrescent. Leaves 2, 3, or 4(or 5) per node; petiole 1.54.5 cm,
pilosulous or hirtellous to glabrous; blade drying papery to
membranous, elliptic to oblanceolate, 720 46 cm, both surfaces pilosulous or hirtellous to glabrescent, base cuneate to
acute, apex acute to weakly acuminate; secondary veins 79
pairs, in abaxial axils frequently with pilosulous domatia; stipules narrowly triangular to subulate, 26 mm, hirtellous or pilosulous to glabrescent. Inflorescences corymbiform, villosulous or pilosulous to glabrescent; peduncle 18 cm; branched
portion corymbiform, 1.57 1.59 cm; bracts reduced or tri-

angular, 0.20.5 mm. Flowers subsessile to pedicellate; pedicels

to 2 mm. Calyx hirtellous to glabrous; ovary portion ellipsoid,
ca. 3 mm; limb deeply lobed; lobes triangular, 0.81 mm. Corolla red to red-orange or yellow, narrowly tubular, outside puberulent to glabrous; tube smooth to shallowly 5-ribbed, 1623
mm; lobes ovate-triangular, 12 mm, acute. Berry ovoid, 67
mm in diam., puberulent to glabrescent. Fl. MayDec.
Cultivated in gardens in Fujian, Yunnan [native and weedy in
Mexico, United States (Florida), and the Caribbean region, Central
America, and tropical South America; cultivated as a perennial in tropical regions and an annual or indoors in temperate regions worldwide].
In S China and in cultivation in general this species does not set
fruit; the fruit description here is based on wild plants. The flowers of
the cultivated plants range from yellow to dark scarlet red, and in recent
years many new cultivars have been developed.

35. HEDYOTIS Linnaeus, Sp. Pl. 1: 101. 1753, nom. cons.

er cao shu
Chen Tao (); Charlotte M. Taylor
Diplophragma (Wight & Arnott) Meisner; Exallage Bremekamp; Gonotheca Blume ex Candolle (1830), not Rafinesque
(1818); Hedyotis sect. Diplophragma Wight & Arnott; Metabolos Blume; Oldenlandia Linnaeus; Thecagonum Babu.
Herbs, subshrubs, or shrubs, annual or perennial, procumbent to erect or climbing, unarmed. Raphides present. Leaves opposite
[or rarely whorled], sometimes clustered at ends of stems, without domatia; secondary venation rarely triplinerved or palmate; stipules persistent, interpetiolar, fused to petiole bases, or united around stem, triangular to truncate, entire or ciliate to laciniate, erose, 1to several lobed and/or -setose. Inflorescences terminal, pseudoaxillary, and/or axillary, few to many flowered and fasciculate, cymose, paniculate, capitate, or glomerulate or reduced to 1 flower, sessile or pedunculate, bracteate or bracts reduced. Flowers
pedicellate or sessile, bisexual and monomorphic or distylous [to unisexual on dioecious plants]. Calyx limb shallowly to deeply
(2)4-lobed (or 5-lobed, Hedyotis hainanensis). Corolla white, pink, purple, or blue, tubular, funnelform, salverform, rotate, or
urceolate, variously glabrous or pubescent inside; lobes (2)4(or 5, H. hainanensis), valvate in bud. Stamens 4(or 5, H. hainanensis),
inserted in corolla tube or throat, included or exserted; filaments developed to reduced; anthers dorsifixed often near base. Ovary 2celled, ovules few to numerous or rarely 1 in each cell on axile placentas; stigma 2-lobed with lobes linear to clavate or rarely
undivided, included or exserted. Fruit indehiscent, schizocarpous, or capsular, generally subglobose to ovoid or dicoccous, crusta-

148

RUBIACEAE

ceous to membranous or leathery, when schizocarpous splitting into 2 mericarps, when capsular splitting partially to entirely septicidally and/or loculicidally, subsequently sometimes splitting other way, apically flattened or with short to well-developed beak (i.e.,
disk area inside calyx limb), sometimes dehiscent primarily through beak, with calyx limb persistent; seeds few to numerous, small,
angular or plano-convex; testa smooth, reticulate, or otherwise variously ornamented; endosperm fleshy; radicle clavate or terete.
About 500 species: tropical and subtropical regions worldwide, most in Africa and Asia, a few in warm temperate regions; 67 species (38
endemic) in China.
This is a very problematic genus or group of genera. Neither the overall identity and limits of this lineagedistributed throughout the tropics
and warm temperate regions of the world, with numerous species with often reduced morphologynor the evolutionary patterns within it are at all
understood or delineated. Widely differing taxonomies and species-level characters have long been used in different regions and floras. It is generally
accepted now that Hedyotis is closely related to or at least in some cases perhaps includes Houstonia Linnaeus, Kadua Chamisso & Schlechtendal,
Kohautia Chamisso & Schlechtendal, Neanotis, Oldenlandia, and a number of smaller segregate genera including Exallage, Oldenlandiopsis Terrell
& W. H. Lewis, Pentodon Hochstetter, Stenaria Terrell, Stenotis Terrell, and Thecagonum. The situation is far from resolution or even general consensus. This genus is treated broadly here, as done also by many recent authors working our flora region (Fukuoka, S. E. Asia Stud. 8(3): 305336.
1970; W. C. Ko in FRPS 71(1): 2677. 1999; Wang & Zhao, J. Trop. Subtrop. Bot. 9(3): 219228. 2001; Dutta & Deb, Taxon. Rev. Hedyotis. 2004).
Recently, some authors have separated Oldenlandia; but, as outlined by Terrell and Robinson (Taxon 52: 775782. 2003), recent molecular studies
have concluded that the circumscriptions and relationships of these two groups are less well understood than had been thought, and these groups are
probably paraphyletic and/or polyphyletic with relation to each other as well as several other genera.
The taxonomy of Hedyotis is further complicated by nomenclatural issues, in particular the designation of the type species. Dutta and Deb (loc.
cit. 2004 a late publication of a 1991 manuscript), following majority opinion of the time, considered H. auricularia as the type of Hedyotis;
however, subsequently, H. fruticosa Linnaeus instead was successfully proposed as the conserved type of the genus (Nicolson, Taxon 41: 564. 1992;
see Vienna Code, App. III, p. 343). The typification of Hedyotis and corresponding generic names were reviewed in detail by Terrell and Robinson
(loc. cit.).
Terrell and Robinson (loc. cit.) also summarized the infrageneric classification and species groups of Hedyotis, including those accepted by W.
C. Ko (loc. cit.), but without noting a few differences between Kos classifications and theirs, nor the use of some incorrect sectional names by Ko
(e.g., H. sect. Euoldenlandia would have been called H. sect. Oldenlandia if it had been published, but it was not; Kos H. sect. Diplophragma included the species that is now the type of the genus, thus this should have been called H. sect. Hedyotis, while this particular section was
synonymized by Terrell & Robinson). The genus circumscription as well as the infrageneric classification of Hedyotis are very far from understood at
present (Groeninckx et al., Scripta Bot. Belg. 44: 33. 2008).
The information available about Hedyotis bodinieri is inadequate to include this species in the key. Because of the complexity of this genus or
group of genera and the large number of species in China, the descriptions here are more detailed than in some other Rubiaceae genus treatments here.
W. C. Ko (loc. cit.) described the fruit of most species of Hedyotis as dehiscent into 2 mericarps at maturity, with mericarps vertically dehiscent at
ventral part, but these fruit are considered capsules by other authors. In some cases, this description was not entirely accurate because the fruit are
actually truly schizocarpous (i.e., with indehiscent mericarps) or primarily loculicidal.

1a. Stems pilose, hirsute, tomentose, or villous and sometimes also hirtellous or villosulous, with pubescence
sparsely to densely distributed and with some trichomes 1 mm or more.
2a. Plants scandent, climbing, or clambering; inflorescences terminal and in uppermost leaf axils, sometimes
on very short lateral stems.
3a. Inflorescences cymose, corymbiform-rounded; flowers all pedicellate, pedicels 1.53 mm ...................... 42. H. obliquinervis
3b. Inflorescences capitate, subcapitate, glomerulate, subglobose, or hemispherical; flowers sessile or
subsessile.
4a. Heads solitary on peduncles or glomerulate along stem and/or along peduncles ........................................ 24. H. dianxiensis
4b. Heads several in paniculate groups on each or at least most peduncles ........................................................ 13. H. capitellata
2b. Plants prostrate on ground, erect, or climbing; inflorescences terminal and/or axillary along middle or
lower stem nodes.
5a. Stipules entire; inflorescences and flowers sessile; Hainan, Taiwan.
6a. Annual herbs to 30 cm tall; calyx lobes ca. 1.5 mm; Taiwan ............................................................................ 11. H. butensis
6b. Perennial subshrubs or shrubs to 1 m tall; calyx lobes 2.53 mm; Hainan ........................................... 64. H. wuzhishanensis
5b. Stipules erose or 25-lobed or -setose; inflorescences and flowers sessile to pedicellate and/or
pedunculate; mainland and Hainan.
7a. Low annual herbs, prostrate to weakly erect; leaves 610 mm wide; flowers few or solitary,
subsessile to pedicellate; Hainan ....................................................................................................................... 58. H. trinervia
7b. Annual or perennial herbs or subshrubs, generally erect and robust; leaves 545 mm wide; flowers
several to numerous, sessile to pedicellate in pedunculate heads or cymes.
8a. Flowers subsessile to pedicellate; glomerules or cymes subsessile to pedunculate; calyx lobes
11.2 mm ........................................................................................................................................................... 34. H. lineata
8b. Flowers sessile or subsessile; cymes or heads pedunculate; calyx lobes 0.31 mm ........................................ 62. H. vestita
1b. Stems glabrous to puberulent, strigillose, hispidulous, strigose, hirtellous, or pilosulous and sometimes
hirsute or pilose in small scattered patches or lines, with trichomes generally all less than 1 mm.

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9a. Inflorescences mostly or all axillary or pseudoaxillary; stem apices generally vegetative.
10a. Inflorescences capitate, cymose, paniculate, fasciculate, or glomerulate, subsessile or on developed peduncles.
11a. Bracts lanceolate, ovate, elliptic, or broadly triangular, 112 mm, partly to fully enclosing flowers; flowers
sessile to shortly pedicellate in heads or congested or somewhat lax cymes, or on pedicels to 1.5 mm;
stipules triangular, 413 mm.
12a. Inflorescences congested-cymose to shortly paniculate or subglomerulate, sessile to pedunculate,
peduncle to 2 cm; leaves 1525 36 cm ............................................................................................... 15. H. cathayana
12b. Inflorescences capitate or glomerulate to laxly cymose, subsessile to pedunculate, peduncle to 8 cm;
leaves 1217 0.54 cm.
13a. Inflorescences capitate or glomerulate, on well-developed peduncles 2.57 cm ................................... 9. H. bracteosa
13b. Inflorescences cymose, branched, on peduncles 0.58 cm.
14a. Stipules 1.55 mm; calyx lobes 0.81.2 mm; corolla tube 2.23.5 mm ........................................... 1. H. acutangula
14b. Stipules 813 mm; calyx lobes 23 mm; corolla tube 4.55 mm ...................................................... 20. H. communis
11b. Bracts reduced, absent, or narrow to triangular, 15 mm, not enclosing flowers; flowers separately
pedicellate and/or pedunculate, solitary or in lax cymes; pedicels 130 mm; stipules reduced or
triangular to truncate, 0.33 mm.
15a. Flowers several (i.e., ca. 5) to many in terminal and axillary lax cymes or panicles, separately
pedicellate with pedicels 115 mm; leaves 735 mm wide.
16a. Pedicels 415 mm; stipules triangular, ovate, or subtruncate, entire or glandular-serrulate or
glandular-erose .......................................................................................................................................... 55. H. tenuipes
16b. Pedicels 13 mm; stipules entire, 46-lobed, -setose, and/or ciliolate.
17a. Stipules 46-lobed or -setose .................................................................................................................. 3. H. assimilis
17b. Stipules entire or ciliolate ................................................................................................................ 35. H. loganioides
15b. Flowers 15 per axil, all axillary on separate peduncles or in pedunculate cymes, pedicels and
peduncles 130 mm; leaves 15 mm wide.
18a. Capsule with developed beak 11.5 mm, conical, longer than calyx lobes; stipules reduced,
sheath portion reduced to a line or up to 0.3 mm; corolla tube 23 mm and lobes 0.51 mm ............. 30. H. herbacea
18b. Capsule flat on top or with beak rounded to broadly angled, up to 0.5 mm, shorter than or
equal to calyx lobes; stipules with sheath tubular to triangular, 0.52 mm; corolla tube
0.82.5 mm and lobes 0.81.5 mm.
19a. Stems quadrate to flattened with angles thickened to winged and usually pubescent to
scaberulous; flowers 15 per peduncle, all pedicellate; stigmas and anthers included .................... 22. H. corymbosa
19b. Stems terete to flattened or 4-angled when young, glabrous to rather evenly scaberulous
or pubescent; flowers 1 or 2 in each stem axil, subsessile to pedicellate; stigmas and
anthers exserted.
20a. Stems glabrous; peduncles and/or pedicels to 3 mm in flower, to 8 mm in fruit .......................... 8. H. brachypoda
20b. Stems puberulent and/or scaberulous; peduncles 420 mm ................................................................ 25. H. diffusa
10b. Inflorescences sessile or subsessile, unbranched, i.e., glomerulate, capitate, congested-cymose,
subcapitate, shortly fasciculate, or 1-flowered.
21a. Stipules at flowering nodes with sheath 330 mm and these and/or leaf bases partially to fully
enclosing and mostly hiding inflorescences.
22a. Leaves 1530 510 cm; stipules narrowly triangular, 830 mm; calyx lobes 68 mm; corolla
tube 1416 mm; Hainan ........................................................................................................................... 23. H. cryptantha
22b. Leaves 2.512 14 cm; stipules broadly triangular to obovate, 310 mm; calyx lobes 1.56 mm;
corolla tube 1.812 mm; mainland and Hainan.
23a. Calyx lobes 1.53 mm; corolla tube 1.82 mm ............................................................................... 33. H. lianshanensis
23b. Calyx lobes 26 mm; corolla tube 612 mm.
24a. Leaves rounded to obtuse at base and sessile or subsessile; inflorescence terminal and
pseudoaxillary; calyx lobes widest above middle ............................................................................ 40. H. merguensis
24b. Leaves rounded to acute at base and petiolate, petioles 318 mm; inflorescences axillary;
calyx lobes widest at or below middle ............................................................................................ 47. H. platystipula
21b. Stipules with sheath 4 mm or shorter (measured from line between bases of petioles), neither these
nor leaf bases enclosing inflorescence.
25a. Leaves linear, narrowly elliptic, narrowly spatulate, linear-lanceolate, or narrowly elliptic-oblong,
0.813 mm wide, with margins mostly straight, with secondary veins not visible.
26a. Fruit compressed globose or subglobose; calyx lobes 11.5 mm; flowers solitary or 2 per node,
pseudoaxillary ....................................................................................................................................... 8. H. brachypoda
26b. Fruit ovoid to lanceoloid; calyx lobes 12.5 mm; flowers 1 to several per node, axillary,
pseudoaxillary, and/or sometimes terminal.

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RUBIACEAE

27a. Inflorescences terminal and pseudoaxillary on short axillary stems; leaves linear to narrowly
spatulate, 0.82 mm wide ...................................................................................................................... 46. H. pinifolia
27b. Inflorescences all axillary; leaves narrowly elliptic to linear, 1.513 mm wide.
28a. Calyx glabrous; flowers 13 per node; leaves 1.54 mm wide .................................................... 54. H. tenelliflora
28b. Calyx densely hispidulous; flowers several per node; leaves 313 mm wide .............................. 61. H. verticillata
25b. Leaves narrowly to broadly elliptic, lanceolate, ovate, or elliptic-oblong, 445 mm wide, with
margins curved, with secondary veins visible or not.
29a. Plants prostrate, regularly rooting at nodes; leaves 0.83.2 0.51.4 cm; flowers few, subsessile
to pedicellate ...................................................................................................................................... 19. H. chrysotricha
29b. Plants erect to procumbent, clambering, or prostrate, rooting only at or near base; leaves
1.212 0.34.5 cm; flowers few to numerous, sessile to pedicellate.
30a. Calyx lobes 1.54 mm.
31a. Fruit indehiscent; inflorescences all produced at nodes below apex; corolla 34.5 mm ......... 33. H. lianshanensis
31b. Fruit dehiscent septicidally then loculicidally; inflorescences terminal and axillary at
uppermost stem nodes; corolla 4.55.5 mm .................................................................................... 59. H. uncinella
30b. Calyx lobes 0.81.5 mm.
32a. Leaves 1.24.5 cm wide, with secondary veins hardly or not visible; inflorescences axillary
at upper nodes and often also terminal; corolla 34.2 mm .............................................................. 48. H. prostrata
32b. Leaves 0.33 cm wide, with secondary veins clearly evident; inflorescences at apex and/or
nodes below apex; corolla 12.5 mm.
33a. Leaves subsessile to petiolate, 0.43 cm wide; stipules densely setose; inflorescences at
nodes well below stem apex; corolla lobes 0.51 mm; widespread ........................................... 4. H. auricularia
33b. Leaves sessile, 0.31 cm wide; stipules entire; inflorescences terminal and at uppermost
nodes; corolla lobes ca. 0.2 mm; N Taiwan .................................................................................... 11. H. butensis
9b. Inflorescences terminal, or terminal and in uppermost leaf axils.
34a. Flowers 5-merous; calyx lobes 33.5 mm .................................................................................................... 28. H. hainanensis
34b. Flowers 4-merous; calyx lobes 0.135 mm.
35a. Leaves linear, linear-lanceolate, narrowly elliptic, narrowly lanceolate, narrowly spatulate, or
narrowly elliptic-oblong, consistently 0.85.5 mm wide.
36a. Flowers 212 in lax cymes; pedicels 320 mm .............................................................................................. 32. H. koana
36b. Flowers 3 to numerous in heads, fascicles, or congested to lax cymes; flowers sessile or on
pedicels to 3 mm.
37a. Inflorescences with axes regularly dichotomous, 413 cm, with numerous flowers ...................... 57. H. tetrangularis
37b. Inflorescences up to 3 cm, capitate to cymose, with several flowers.
38a. Flowers on pedicels 0.51.2 mm; corolla tube 1.52.2 mm .............................................................. 10. H. brevicalyx
38b. Flowers sessile or on pedicels to 1 mm; corolla tube 34.2 mm .......................................................... 46. H. pinifolia
35b. Leaves variously shaped, linear to elliptic, lanceolate, elliptic-oblong, or ovate, 3120 mm wide
with at least some leaves more than 5 mm wide.
39a. Inflorescences 1-flowered or capitate, subcapitate, fascicled, or congested-cymose, with flowers few
to numerous and sessile to subsessile or shortly pedicellate, in heads or congested to lax cymes.
40a. Flowers solitary or 25 and fasciculate to very shortly cymose, sessile to shortly pedicellate; leaves
0.87 0.53.5 cm.
41a. Calyx lobes ca. 4 mm, longer than or as long as corolla tube ............................................................... 43. H. ovata
41b. Calyx lobes 33.5 mm, much shorter than corolla tube ................................................................. 50. H. pulcherrima
40b. Flowers 4 to numerous, sessile to shortly pedicellate in heads, congested cymes, or umbelliform
cymes; leaves 112 0.64.5 cm.
42a. Inflorescences with peduncles, with 1 to several heads, with subtending bracts developed to
leaflike or reduced.
43a. Calyx lobes 36 mm.
44a. Leaves 45.5 0.62.6 cm ...................................................................................................... 65. H. xanthochroa
44b. Leaves (at least well-developed ones) 812 34.5 cm ..................................................... 66. H. yangchunensis
43b. Calyx lobes 0.34 mm.
45a. Corolla tube ca. 7 mm; inflorescence with 1 congested-cymose head (as far as known);
flowers sessile ................................................................................................................................... 63. H. wangii
45b. Corolla tube 13 mm; inflorescence (at least usually) with 215 heads; flowers sessile
or subsessile.
46a. Plants erect; corolla lobes shorter than tube, tube ca. 3 mm and lobes ca. 1.3 mm .......... 14. H. capituligera
46b. Plants clambering to climbing; corolla lobes longer than tube, tube 12 mm and
lobes 35 mm.

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151

47a. Stems and leaves glabrescent or hirtellous, pilosulous, hispidulous, or tomentulose;

calyx limb lobed for 1/33/4 ................................................................................................ 13. H. capitellata
47b. Stems and leaves densely tomentulose or villous to glabrous; calyx limb deeply lobed ... 24. H. dianxiensis
42b. Inflorescences sessile or subsessile, in 1 head subtended by leaves or well developed to
leaflike bracts.
48a. Calyx lobes ca. 35 mm; leaves 412 cm wide ............................................................................. 67. H. yazhouensis
48b. Calyx lobes 15.5 mm; leaves 0.46 cm wide.
49a. Leaves with petioles 1015 mm ................................................................................................ 5. H. baotingensis
49b. Leaves sessile or with petioles to 10 mm, at least some shorter than 10 mm.
50a. Calyx lobes 11.2 mm; corolla tube 23 mm ........................................................................... 48. H. prostrata
50b. Calyx lobes 25.5 mm; corolla tube 312 mm.
51a. Corolla tube 912 mm ......................................................................................................... 40. H. merguensis
51b. Corolla tube 36.5 mm.
52a. Stipules acute or aristate.
53a. Leaves 0.41.2 cm wide, with petioles to 3 mm .......................................................... 37. H. longipetala
53b. Leaves 1.44.5 cm wide, with petioles to 10 mm.
54a. Stems glabrous; leaves 712 cm; corolla tube 45 mm ............................................ 45. H. paridifolia
54b. Stems densely villous to hirtellous; leaves 2.56.2 cm; corolla tube
5.56.5 mm ......................................................................................................... 64. H. wuzhishanensis
52b. Stipules laciniate or with several lobes or bristles.
55a. Leaves puberulent to glabrous adaxially; flowers sessile or subsessile; corolla
lobes 1.52 mm; fruit 1.82 mm in diam. ....................................................................... 59. H. uncinella
55b. Leaves glabrous or glabrescent adaxially; flowers sessile or on pedicels to 2 mm;
corolla lobes 22.5 mm; fruit 22.5 mm in diam.
56a. Corolla tube ca. 5 mm; fruit ca. 2 mm in diam. ........................................................... 17. H. cheniana
56b. Corolla tube ca. 3.5 mm; fruit ca. 2.5 mm in diam. .............................................. 56. H. terminaliflora
39b. Inflorescences branched, cymose to paniculate, with flowers few to numerous, at least some
of them regularly pedicellate (or separated on well-developed inflorescence axes).
57a. Plants low annuals or perennials, vegetative part (and usually entire plant) mostly up to 15 cm
tall, sometimes to 40 cm tall, often weak or scapose, often quite succulent; flowers few to
numerous in small to diffuse cymes.
58a. Leaves usually clustered at top of stem (i.e., at base of inflorescence) due to reduced internodes
and appearing verticillate; stems densely pilosulous, puberulent, or villosulous and often
glandular on uppermost nodes ............................................................................................................. 44. H. ovatifolia
58b. Leaves generally distributed along stem in pairs, or entire stem with reduced internodes; stems
glabrous to variously pubescent, not glandular.
59a. Hypanthium and fruit 4-winged, wings usually extending downward along pedicel .......................... 49. H. pterita
59b. Hypanthium and fruit smooth or with low ridges, or hypanthium sometimes narrowly winged
in flower but becoming smooth as fruit develops.
60a. Calyx lobes ovate, relatively broad, sometimes with evident venation ................................. 31. H. hermanniana
60b. Calyx lobes triangular to lanceolate, generally narrow, not veined.
61a. Calyx limb lobed essentially to base, with tubular portion reduced or just a line; fruit
with beak mostly as long as or longer than calyx lobes; plants not or somewhat
succulent, weedy in various habitats ............................................................................................... 6. H. biflora
61b. Calyx limb with tube developed, lobed for 1/22/3, remaining developed and connected
to lobes in fruit; fruit with beak much shorter than calyx lobes; plants succulent, mostly
on seashores and rocks at low elevations ................................................................................. 53. H. strigulosa
57b. Plants small to large, apparently perennials, vegetative portion usually at least 20 cm tall, robust,
erect to clambering or climbing, usually not scapose, not or sometimes slightly succulent;
flowers several to numerous in congested to expanded cymes or panicles.
62a. Plants clambering, scandent, lianescent, or climbing.
63a. Leaves with secondary veins plane and generally not evident abaxially ........................................ 51. H. scandens
63b. Leaves with secondary veins plane to prominulous and evident abaxially.
64a. Flowers pedicellate in dichotomous cymules, with pedicels 13.5 mm; axes and pedicels of
inflorescence generally spreading at 6090 .............................................................................. 2. H. ampliflora
64b. Flowers sessile to subsessile or pedicellate in congested to subcapitate cymules, with
pedicels to 2 mm; axes and pedicels of inflorescence generally ascending, at angles less
than 60.

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152

65a. Stems densely villous and/or tomentulose; flowers in distinct, discrete heads; calyx lobes
generally erect ......................................................................................................................... 24. H. dianxiensis
65b. Stems glabrous to papillose, pilosulous, or velutinous-strigillose; inflorescence at least
partly cymose; calyx lobes spreading to usually reflexed ....................................................... 29. H. hedyotidea
62b. Plants erect.
66a. Flowers all pedicellate, pedicels 415 mm.
67a. Calyx lobes 0.10.3 mm; corolla 11.5 mm ........................................................................... 18. H. chereevensis
67b. Calyx lobes 11.5 mm; corolla 4.57 mm ..................................................................................... 55. H. tenuipes
66b. Flowers sessile to subsessile, mixed sessile and pedicellate, or all pedicellate, pedicels to
5 mm but most less than 4 mm.
68a. Inflorescence axes and pedicels spreading at 90 or more; leaves usually borne at only
2 or 3 stem nodes ................................................................................................................................. 26. H. effusa
68b. Inflorescence axes and pedicels ascending to slightly spreading, generally at angles less
than 60 (or frequently to 90 or more in H. vachellii in fruit); leaves borne at few to
numerous stem nodes.
69a. Stipules setose, lobed, or laciniate, with 2 to several narrow lobes, awns, or bristles (at
least on lower nodes).
70a. Corolla tube longer than corolla lobes; inflorescences with several subcapitate to
congested-cymose heads or cymules.
71a. Flowers sessile to subsessile ........................................................................................... 14. H. capituligera
71b. Flowers sessile and pedicellate, pedicels to 4 mm ............................................................ 38. H. matthewii
70b. Corolla lobes longer than corolla tube; inflorescences with few- to many-flowered
cymes; pedicels 15 mm.
72a. Stipules 46-lobed or -setose ................................................................................................. 3. H. assimilis
72b. Stipules glandular-serrulate to 3-lobed or -setose ..................................................................... 39. H. mellii
69b. Stipules entire or sometimes with a terminal awn and/or serrate to serrulate-denticulate.
73a. Leaves with secondary veins prominulous and evident on abaxial surface ................................ 39. H. mellii
73b. Leaves with secondary veins plane and generally not evident on abaxial surface.
74a. Stems sharply angled to winged, at least on lower internodes; leaves sessile, often
clasping at base ................................................................................................................... 1. H. acutangula
74b. Stems rounded to flattened or angled, angles generally not sharp; leaves petiolate to
subsessile, at base narrow to rounded but usually not clasping.
75a. Calyx lobes ca. 4 mm ............................................................................................................. 43. H. ovata
75b. Calyx lobes 0.32.5 mm.
76a. Plants scapose, with leaves clustered at base; leaves 59 cm wide ..................... 52. H. shenzhenensis
76b. Plants with leaves distributed along stem; leaves 0.43.5 cm wide.
77a. Corolla tube 1.52.5 mm; calyx lobes 0.51.2 mm; inflorescence axes
often becoming spiciform and/or scorpioid.
78a. Calyx lobes 0.81.2 mm; leaves with secondary veins 46 pairs .................. 21. H. consanguinea
78b. Calyx lobes ca. 0.5 mm; leaves with secondary veins ca. 3 pairs .................... 36. H. longiexserta
77b. Corolla tube 2.510 mm; calyx lobes 12.5 mm; inflorescence axes
generally dichasial.
79a. Corolla with tube 410 mm, lobes 1/2 as long as tube or shorter.
80a. Corolla white or pink, with tube markedly funnelform ................................. 16. H. caudatifolia
80b. Corolla pink to purple, with tube cylindrical to slightly funnelform.
81a. Corolla tube ca. 6 mm ......................................................................................... 27. H. exserta
81b. Corolla tube 710 mm ......................................................................... 41. H. minutopuberula
79b. Corolla with tube 2.54 mm, lobes shorter than tube but more than
1/2 as long as tube.
82a. Pedicels to 4 mm; corolla tube 3.54 mm and lobes 23 mm .......................... 38. H. matthewii
82b. Pedicels to 1.5 mm; corolla tube 2.53.5 mm and lobes 22.2 mm.
83a. Inflorescences narrowly cylindrical to pyramidal, axes ascending;
fruit 23 mm in diam. ................................................................................ 12. H. cantoniensis
83b. Inflorescences broadly pyramidal to rounded, axes ascending to spreading
at 90 or more; fruit ca. 1.8 mm in diam. ........................................................ 60. H. vachellii
1. Hedyotis acutangula Champion ex Bentham, Hookers J.
Bot. Kew Gard. Misc. 4: 171. 1852.
jin cao

Oldenlandia acutangula (Champion ex Bentham) Kuntze.

Herbs or subshrubs, apparently biennial or perennial, erect,
2560 cm tall; stems sometimes slightly constricted at nodes, 4-

RUBIACEAE

angled to 4-winged, wings to 1.5 mm wide. Leaves sessile or

subsessile; blade drying stiffly papery to leathery, ovate-lanceolate, lanceolate, elliptic, or lanceolate-elliptic, 212 0.52.5
cm, glabrous, base cuneate to rounded, margins thickened, often
inrolled, sometimes glandular-serrulate, apex acute or shortly
acuminate; secondary veins indistinct; stipules fused to petiole
bases, ovate or triangular, 1.55 mm, glabrous, margins often
becoming revolute, entire or glandular-serrulate, apex acute
to acuminate. Inflorescences terminal, in axils of uppermost
leaves, and occasionally also axillary or pseudoaxillary at lower
nodes, paniculate or corymbose, 320 cm, many flowered,
glabrous, pedunculate; peduncle 28 cm; axes dichasial, sharply
4-angled to thinly 4-winged; bracts triangular to lanceolate
and 0.35 mm, or leaflike and larger. Flowers sessile, distylous. Calyx glabrous; hypanthium portion turbinate to ellipsoid,
0.81 mm; limb lobed nearly to base; lobes ovate, triangular, or
lanceolate, 0.81.2 mm. Corolla white to pink, funnelform, outside glabrous; tube 2.23.5 mm, inside tomentose above middle; lobes ovate-lanceolate, 23 mm. Anthers included or exserted, ca. 1 mm. Stigma included or exserted, 0.51 mm. Fruit
capsular, obovoid to ellipsoid, 22.5 11.2 mm, glabrous,
leathery, apically flat or with beak to 0.3 mm, septicidal and
then loculicidal along middle of septum; seeds several, black,
suborbicular, angled. Fl. MayDec, fr. JunDec.
Hill slopes, open fields; near sea level to 600 m. Fujian, Guangdong, Hainan (Wanning) [N Thailand, Vietnam].
Deb and Dutta (Taxon 34: 296297. 1985) briefly discussed this
species and formally synonymized Hedyotis elegans Wallich ex Kurz
with it; however, they gave no characters or discussion of either species
and gave its range as Burma and Malaysia even though its type is
from Hong Kong, so their conclusions are provisionally not included
here.

2. Hedyotis ampliflora Hance, J. Bot. 17: 11. 1879.

guang hua er cao
Oldenlandia ampliflora (Hance) Kuntze.
Herbs or shrubs, perennial, lianescent, clambering, twining,
or scandent, to perhaps 1 m; stems flattened to subterete,
smooth to sulcate, densely hirtellous or pilosulous throughout
or in lines, to glabrescent. Leaves petiolate; petiole 26 mm,
densely hirtellous to puberulent; blade drying papery, lanceolate, broadly lanceolate, elliptic, or narrowly elliptic, 27 0.8
3 cm, adaxially hispidulous or strigillose along midrib to
throughout, abaxially hispidulous or hirtellous to glabrous, base
cuneate to obtuse, apex acute or shortly acuminate; secondary
veins 3 or 4 pairs; stipules fused to petiole bases, densely hirtellous to hispidulous, sheath portion 13 mm, truncate to triangular, apex lacerate into 35 narrowly triangular to linear
bristles or lobes 16 mm. Inflorescences terminal and occasionally also in axils of uppermost leaves, compound-cymose, corymbiform, 312 cm, several to many flowered, densely to
sparsely hirtellous to puberulent, pedunculate; peduncles 13.5
cm; axes dichotomous, spreading; bracts triangular, lanceolate,
or narrowly elliptic, 17 mm; pedicels 13.5 mm. Flowers pedicellate, apparently distylous. Calyx glabrous to densely hirtellous; hypanthium portion subglobose or turbinate, 0.81 mm;
limb lobed for 3/44/5; lobes lanceolate to spatulate, 1.21.8

153

mm, acute, usually spreading to reflexed. Corolla white or

greenish white, tubular to salverform, outside granular-puberulent; tube ca. 1 mm, barbate in throat; lobes lanceolate to spatulate, 22.5 mm, acute. Anthers partially or long exserted, 0.81
mm. Stigmas in short-styled form not seen, or exserted and ca.
0.3 mm. Fruit capsular, subglobose to ovoid, sometimes slightly
flattened, 2.53 22.5 mm, glabrous to densely hirtellous,
cartilaginous to stiffly papery, loculicidal then septicidal across
top then tardily along sides, beak conical, ca. 1 mm; seeds numerous, blackish brown, angled. Fl. MayNov, fr. OctNov.
Sparse forests or thickets on mountain slopes; 100400 m.
Hainan.
The Kew Rubiaceae checklist (Govaerts et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on May 2007)
reports this species also from Vietnam; it is not listed in the Vietnam
checklist.

3. Hedyotis assimilis Tutcher, Rep. Bot. Dept. Hong Kong

1914: 32. 1915.
qing yuan er cao
Oldenlandia assimilis (Tutcher) Chun.
Herbs or subshrubs, erect, branched, 3040 cm tall; stems
quadrangular, scabrous. Petiole ca. 3 mm; leaf blade drying papery, lanceolate, 59 1.52.5 cm, scabrous, base cuneate,
apex long acuminate; secondary veins ca. 4 pairs, adaxially
impressed; stipules broadly triangular, ca. 3 mm, usually 46lobed or perhaps setose. Inflorescence axillary and terminal, in
cymes arranged into panicles, usually shorter than leaves, 68flowered; bracts small, narrow; pedicels 13 mm. Flowers pedicellate, with biology unknown. Calyx glabrous; lobes ovate,
acute. Corolla white, outside glabrous, inside and on filaments
densely bearded; lobes lanceolate, longer than tube, obtuse.
Style exserted, bearded below middle. Fruit capsular, ellipsoid,
ca. 3 mm including persistent calyx limb, septicidal then loculicidal through middle of septum; seeds numerous, black,
angled. Fl. AprMay.
Broad-leaved forests, slopes in ravines. Guangdong (Conghua,
Qingyuan).

4. Hedyotis auricularia Linnaeus, Sp. Pl. 1: 101. 1753.

er cao
Herbs, perennial, tufted to weak or procumbent, sometimes (var. auricularia) to usually (var. mina) rooting at
lowermost nodes, to 1 m tall or long; stems flattened, 4-angled,
or terete becoming angled and/or sulcate, densely hirtellous,
hispidulous, strigillose, pilosulous, or puberulent, to sometimes
glabrescent (var. auricularia), or glabrous except villosulous to
pilosulous and sometimes also hirsute to pilose inside grooves
(var. mina). Leaves subsessile to petiolate; petiole to 7 mm,
densely strigillose to pilosulous; blade drying subleathery to
papery, ovate, lanceolate, elliptic, or elliptic-oblong, 2.29
0.63 cm (var. auricularia) or 1.23 0.41.4 (var. mina),
adaxially glabrous or puberulent along midrib or scabrous,
abaxially glabrous on lamina and densely puberulent to strigillose or hispidulous along principal veins and margins, base
acute to obtuse and sometimes shortly decurrent, apex acute or

154

RUBIACEAE

acuminate; secondary veins 46 (var. auricularia) or 24 (var.

mina) pairs; stipules fused to petiole bases, densely puberulent,
strigillose, or pilosulous, sheath 1.53.5 mm, truncate to triangular or elliptic, in var. auricularia with 39 linear or setiform lobes 0.55 mm, often glandular at apex, in var. mina
entire and acute to aristate or sometimes with 2 setose lobes to 2
mm. Inflorescences axillary, glomerulate to congested-cymose,
512 mm in diam., several to many flowered (var. auricularia)
or 27-flowered (var. mina), puberulent, sessile; bracts lanceolate to stipuliform, 0.32 mm; pedicels to 1 (var. auricularia) or
3 (var. mina) mm. Flowers sessile to pedicellate, distylous.
Calyx puberulent (var. auricularia) or glabrous (var. mina);
hypanthium portion obconic, ca. 1 mm; limb lobed essentially
to base; lobes lanceolate to triangular, 0.81.2 mm, usually
ciliolate. Corolla white, tubular or tubular-funnelform, outside
pilosulous at least on lobes (var. auricularia) or glabrous
throughout (var. mina); tube 11.5 mm, barbate in throat; lobes
0.51 mm. Anthers narrowly oblong, ca. 1 mm, exserted or
included. Stigma ca. 1 mm, exserted or included. Fruit indehiscent, globose to ovoid, 1.21.5 mm in diam., sparsely hirtellous,
hispidulous, or puberulent (var. auricularia), or glabrous (var.
mina), sometimes with peduncles and/or pedicels elongating, to
2 mm; seeds 412, black, foveolate. Fl. and fr. MarDec.
Forest margins, thickets, grasslands, open fields, streamsides; below 1001500 m. Guangdong, Guangxi, Guizhou, Hainan, Yunnan
[India, Japan (Ryukyu Islands), Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand, Vietnam; Australia].

1a. Young stems densely hirtellous,

hispidulous, strigillose, pilosulous, or
puberulent to glabrescent; leaf blade
lanceolate to elliptic, 2.29 0.63 cm;
secondary veins 46 pairs .................... 4a. var. auricularia
1b. Young stems longitudinally sulcate,
glabrous except villosulous to
pilosulous and sometimes also hirsute
to pilose inside grooves; leaf blade
ovate to lanceolate, elliptic, or
elliptic-oblong, 1.23 0.41.4 cm;
secondary veins 24 pairs .............................. 4b. var. mina
4a. Hedyotis auricularia var. auricularia
() er cao (yuan bian zhong)
Exallage auricularia (Linnaeus) Bremekamp; Oldenlandia auricularia (Linnaeus) K. Schumann.
Stems densely hirtellous, hispidulous, strigillose, pilosulous, or puberulent to glabrescent with pubescence mostly uniform. Leaf blade lanceolate to elliptic, 2.29 0.63 cm;
secondary veins 46 pairs; stipules with 39 linear to setiform
lobes to 5 mm. Inflorescences glomerulate to congested-cymose, several to many flowered; pedicels to 1 mm. Fl. and fr.
MarDec.
Forest margins, thickets, grasslands; below 1001500 m. Guangdong, Guangxi, Guizhou, Hainan, Yunnan [India, Japan (Ryukyu
Islands), Malaysia, Myanmar, Nepal, Philippines, Sri Lanka, Thailand,
Vietnam; Australia].
The Chinese range of this taxon was not mentioned by Dutta
and Deb (Taxon. Rev. Hedyotis. 2004) but was long known to other
authors.

4b. Hedyotis auricularia var. mina W. C. Ko, Fl. Hainan. 3:

578. 1974.
xi ye ya po chao
Stems at least when young longitudinally sulcate, villosulous to pilosulous and sometimes also hirsute to pilose inside
grooves. Leaf blade ovate to lanceolate, elliptic, or elliptic-oblong, 1.23 0.41.4 cm; secondary veins 24 pairs; stipules
entire and acute to aristate or sometimes with 2 setose lobes to 2
mm. Inflorescences subcapitate to congested-cymose, 27flowered in each node; pedicels to 3 mm. Fl. almost year-round.
Wet or humid open fields, streamsides, thickets; ca. 200 m.
Guangdong, Guangxi, Hainan.

5. Hedyotis baotingensis W. C. Ko, J. S. China Agric. Univ.

16(4): 43. 1995.
bao ting er cao
Little branched herbs, prostrate, 4060 cm tall; stems terete to flattened, scabrous. Leaves petiolate, often congested and
appearing verticillate at top of stem; petiole 1015 mm; blade
drying papery, elliptic, elliptic-oblong, ovate, or obovate, 413
36 cm, adaxially glabrous, abaxially sparsely hirtellous at
least along veins, base obtuse then decurrent, apex obtuse then
sometimes abruptly and shortly acute; secondary veins 512
pairs; stipules narrowly triangular, long acuminate, marginally
erose. Inflorescence and flowers unknown. Infructescence terminal, capitate, subglobose, 22.5 cm in diam., sessile. Fruit indehiscent, sessile or subsessile, globose or ellipsoid, 11.5 mm
in diam., pilosulous; persistent calyx lobes narrowly spatulateoblong, narrowly lanceolate, or linear, 55.5 mm, striate; seeds
3 or 4, black, angled. Fr. Oct.
Broad-leaved forests on shady slopes. Hainan (Baoting).

6. Hedyotis biflora (Linnaeus) Lamarck, Tabl. Encycl. 1: 272.

1792.
shuang hua er cao
Oldenlandia biflora Linnaeus, Sp. Pl. 1: 119. 1753;
Hedyotis paniculata (Linnaeus) Lamarck; H. racemosa
Lamarck; O. crassifolia Candolle; O. paniculata Linnaeus.
Herbs, annual or perennial, erect to procumbent, to
15(30) cm tall; stems 4-angled to subterete and/or sulcate, glabrous. Leaves subsessile to petiolate; petiole to 0.5 cm, glabrous; blade drying papery to subleathery, elliptic-oblong, elliptic-ovate, obovate, oblanceolate, or elliptic, 14 0.31 mm,
glabrous, base acute to obtuse then abruptly decurrent, apex
acute to rounded; secondary veins indistinct; stipules fused to
petioles, triangular, 12 mm, glabrous, usually costate, acute to
long acuminate. Inflorescences terminal and in axils of uppermost leaves, cymose to compound-cymose, 16 cm, glabrous,
212-flowered, pedunculate; peduncles 0.53 cm; bracts triangular to lanceolate, 0.53 mm; pedicels 0.510 mm. Flowers
subsessile to pedicellate, apparently distylous. Calyx glabrous;
hypanthium portion turbinate, 0.51 mm; limb lobed essentially
to base; lobes triangular, 0.31 mm. Corolla white, sometimes
flushed with lavender, tubular to somewhat urceolate, outside
glabrous or puberulent in lines; tube 1.22 mm, barbate in

RUBIACEAE

throat; lobes spatulate-oblong, 11.5 mm. Anthers included or

exserted, 0.30.5 mm. Stigma exserted or apparently included,
ca. 0.2 mm. Fruit capsular, subglobose, oblate, or hemispherical, 2.53 mm in diam., loculicidally dehiscent across top, beak
0.51 mm high, pedicels usually elongating; seeds numerous,
black, foveolate. Fl. and fr. JanSep.
Limestone mountains, coastal areas, weedy fields, wastelands; sea
level to 1200 m. Fujian, Guangdong, Guangxi, Hainan, Jiangsu, Taiwan,
Yunnan [India, Indonesia, Malaysia, Nepal, Vietnam; SE Asia to Pacific
islands].
The circumscription and name of this species is controversial or,
at best, complicated. These are small herbs of ruderal sites and thus
probably respond markedly to local conditions, in particular growing to
a larger size with larger leaves, inflorescences, and fruit in sites with
better conditions. Similar species that have been variously synonymized
but are separated here include Hedyotis strigulosa and H. pterita; see
further discussion under H. strigulosa. Alternatively Biju et al. (Rheedea 2(1): 1118. 1992) separated H. biflora from H. racemosa (syn.
Oldenlandia paniculata); their treatment is carefully done but regional,
and because their key does not correspond well with the Chinese plants,
their conclusions are not accepted here.
The name Oldenlandia crassifolia was placed in synonymy with
both Hedyotis biflora and H. coreana (H. strigulosa) by W. C. Ko (in
FRPS 71(1): 39, 77. 1999). The only other author who studied this
name in any detail, Merrill (Enum. Philipp. Fl. Pl. 3(4): 492512; 3(5):
513576. 1923), cited this as a synonym of H. biflora and is followed
here.

7. Hedyotis bodinieri H. Lveill, Repert. Spec. Nov. Regni

Veg. 11: 64. 1912.
da mao shan er cao
Oldenlandia bodinieri (H. Lveill) Chun.
Herbs or subshrubs; stems pilosulous. Leaf blade drying
membranous, margins not revolute when dry. Inflorescence
shortly cymose. Flowers pedicellate. Calyx lobes sublinear,
acute, as long as corolla tube. Corolla white; tube slightly
longer than lobes. Anthers included. Fl. May.
Mountaintops. Hong Kong (Damao Shan).
This very poorly known species is not included in the key to
species here. It may will turn out to be an older name for an otherwiseknown species.

8. Hedyotis brachypoda (Candolle) Sivarajan & Biju, Taxon

39: 672. 1990.
ni ding jing cao
Oldenlandia brachypoda Candolle, Prodr. 4: 424. 1830.
Slender herbs, annual, diffusely branched, to 50 cm tall;
stems terete to slightly flattened, glabrous. Leaves sessile or
subsessile; blade drying membranous, linear, narrowly elliptic,
or narrowly spatulate, 736 14 mm, adaxially glabrous
(sometimes appearing papillose due to collapsed, large epidermal cells) to scaberulous and usually shiny, abaxially glabrous
and matte, base acute, margins usually revolute at least when
dry, apex acute; secondary veins not evident; stipules fused to
petiole bases, glabrous, truncate to broadly triangular, 0.82
mm, with 13 linear to setiform lobes 0.21 mm. Inflorescences

155

pseudoaxillary, 1-flowered (2-flowered and fasciculate), glabrous, sessile or with peduncle to 3 mm, ebracteate. Flowers
subsessile to shortly pedunculate, homostylous. Calyx glabrous;
hypanthium portion globose, 11.2 mm; limb lobed essentially
to base; lobes triangular, 11.5 mm. Corolla white, rotate,
outside glabrous; tube 11.5 mm, glabrous at throat; lobes triangular, 11.5 mm. Anthers ca. 0.3 mm, exserted. Stigma ca. 0.8
mm, exserted. Fruit capsular, membranous to papery, compressed globose to subglobose or somewhat dicoccous, ca. 2.5
34 mm, loculicidally dehiscent through flattened top, peduncles to 8 mm; seeds ca. 20, dark brown, angled, deeply and
thickly foveolate. Fl. and fr. (Feb)MarNov.
Paddy fields, ridges of farmlands, humid open fields; below 100
1500 m. Anhui, Guangdong, Guangxi, Hainan, Yunnan [Bangladesh,
Bhutan, India, Indonesia, Japan, Malaysia (Malacca), Nepal, Philippines, Vietnam].
The taxonomy of this and related species is complicated, and different authors have drawn very different conclusions (e.g., Sivarajan &
Biju, loc. cit.: 665674; Dutta & Deb, Taxon. Rev. Hedyotis. 2004). In
particular, Hedyotis corymbosa, H. diffusa, and H. erecta Manilal &
Sivarajan are related and have been variously circumscribed. Here,
these species are circumscribed generally, though not completely, following Sivarajan and Biju (loc. cit.) and W. C. Ko (in FRPS 71(1): 72,
75. 1999).

9. Hedyotis bracteosa Hance, J. Bot. 23: 323. 1885.

da bao er cao
Oldenlandia bracteosa (Hance) Kuntze.
Herbs, erect, to 50 cm tall; stems glabrous, 4-angled with
angles rounded to acute or winged, wings to 1 mm wide.
Leaves sessile or subsessile; blade drying papery, elliptic-oblong or oblong-lanceolate, 1215 24 cm, glabrous, base
obtuse to cordulate then abruptly narrowed, apex acute or
shortly acuminate; secondary veins ca. 7 pairs, weakly marked
or not visible; stipules fused to petiole bases, broadly triangular
to ovate, 412 mm, glabrous, with 2 winged flanges 12.5 mm
wide extending from each petiole base to apex, entire or glandular-serrulate, acute to acuminate. Inflorescence axillary, capitate, glabrous, pedunculate; peduncles 2.57 cm; involucral
bracts 2 or 4, ovate, 1025 918 mm; head hemispherical,
1016 mm in diam.; floral bracts narrowly lanceolate, 14 mm;
pedicels 0.21 mm. Flowers subsessile to shortly pedicellate,
distylous. Calyx glabrous; hypanthium portion ellipsoid, ca. 1
mm; limb lobed nearly to base; lobes ovate or lanceolate, 2.23
1.22 mm, often pinnately veined, sometimes ciliolate, obtuse
to acute. Corolla white, tubular or tubular-funnelform, outside
glabrous; tube ca. 4 mm, pilosulous inside in upper half and
throat; lobes lanceolate to triangular, ca. 1.5 mm, acute. Anthers
exserted and ca. 0.8 mm in short-styled form, in long-styled
form not seen. Stigmas ca. 0.5 mm and positioned near middle
of corolla tube in short-styled form, ca. 1 mm and exserted in
long-styled form. Fruit capsular, subglobose to ellipsoid, ca. 2
mm, glabrous, septicidal then perhaps rather tardily loculicidal,
with calyx limb persistent; seeds numerous, black, angled,
rugose. Fl. AprJul, fr. Jul.
Sparse forests on mountain slopes or on humid lands in
valleys/ravines. Guangdong.

156

RUBIACEAE

W. C. Ko (in FRPS 71(1): 56. 1999) described the corolla as

having lobes as long as the corolla tube, but this has not been seen on
specimens studied nor reported by other authors.

10. Hedyotis brevicalyx Sivarajan, Biju & P. Mathew, Kew

Bull. 48: 393. 1993.
san xing hua er cao
Oldenlandia umbellata Linnaeus, Sp. Pl. 1: 119. 1753;
Hedyotis umbellata (Linnaeus) Lamarck (1792), not Walter
(1788).
Herbs, annual or perennial, diffusely branched, to 40 cm
tall; stems 4-angled perhaps becoming terete, ribbed, scaberulous. Leaves opposite but often crowded and appearing verticillate, sessile; blade drying papery, linear to narrowly lanceolate,
1020 13 mm, adaxially punctate and glabrescent, abaxially
scaberulous along midrib, base acute, margins flat to revolute,
apex acute; secondary veins indistinct; stipules fused to petioles, glabrescent, truncate to rounded, 11.5 mm, with 25
bristles 12 mm. Inflorescence terminal and sometimes in uppermost axils, congested-cymose to congested-umbelliform,
several flowered, scaberulous, pedunculate; peduncles 12 cm;
branched portion ca. 3 cm; bracts stipuliform, ca. 1 mm; pedicels 0.51.2 mm. Flowers distylous, shortly pedicellate. Calyx
apparently glabrous; hypanthium portion subglobose to ovoid,
0.31 mm; limb lobed essentially to base; lobes lanceolate to
triangular, 12 mm, ciliate. Corolla white, tubular, outside glabrous; tube 1.52.2 mm, glabrous inside; lobes lanceolate, 1.5
2 mm, inside sparsely pubescent. Anthers 0.51 mm, included
or exserted. Stigmas 0.81 mm, included or exserted. Fruit
capsular, compressed globose to weakly didymous, 22.5 2
2.5 mm, loculicidally dehiscent across top, glabrous to scaberulous, perhaps papery; seeds several, blackish brown.
Sandy lands at sea beaches; sea level to near sea level [100900 m
in India]. Hainan (Lingshui) [India, Indonesia (Java), N Myanmar, Pakistan, Sri Lanka, Vietnam].

out; tube 0.82.3 mm; lobes oblong-linear, ca. 0.2 mm. Anthers
exserted. Fruit indehiscent or dehiscent across top, subglobose,
somewhat compressed laterally, 1.52 mm in diam.; seeds numerous, angled. Fl. JulSep.
Taiwan (Yilan).
This species is poorly known, and the available descriptions are
limited. It is included in the key to species twice, for the described
hirsute stems and the potentially glabrous stems, a condition eventually
found in most pubescent species of Hedyotis (and other Rubiaceae) and
one that can be confirmed as more specimens become available.

12. Hedyotis cantoniensis F. C. How ex W. C. Ko, J. S. China

Agric. Univ. 16(4): 42. 1995.
guang zhou er cao
Herbs or subshrubs, erect, perennial, to 60 cm tall; stems
subterete, sometimes striate, glabrous. Leaves subsessile to petiolate; petiole to 8 mm, glabrous; blade drying thinly leathery,
ovate, oblong-elliptic, narrowly elliptic, or narrowly lanceolate,
313 0.83 cm, glabrous, base acute to cuneate, apex acute to
shortly acuminate; secondary veins indistinct; stipules fused to
petiole bases, triangular to broadly triangular, 25 mm, glabrous, marginally entire to densely glandular serrate, acute to
aristate with tip to 4 mm. Inflorescences terminal and in axils of
uppermost leaves, cymose, paniculate and pyramidal to narrowly cylindrical or racemiform, 115 cm, several to many
flowered, glabrous, sessile and tripartite or pedunculate; peduncles 14.5 cm; bracts narrowly elliptic to triangular, 0.54 mm;
pedicels to 1 mm. Flowers sessile to pedicellate, distylous. Calyx glabrous; hypanthium portion subglobose to obconic, 11.2
mm; limb 12 mm, deeply lobed; lobes triangular to narrowly
triangular. Corolla salverform to tubular-funnelform, outside
glabrous; tube 2.53.5 mm, sparsely barbate in throat; lobes
oblong-lanceolate, 22.2 mm. Anthers included or exserted, ca.
1.1 mm. Stigmas included or exserted, ca. 0.3 mm when exserted. Fruit capsular, ellipsoid to subglobose, 23 mm, septicidally then loculicidally dehiscent, glabrous, cartilaginous to
somewhat woody, apex flat or with low rounded beak; seeds
several, angled. Fl. AprAug, fr. JunNov.

There is some disagreement over the identity and correct name of

this species; whether Hedyotis brevicalyx actually occurs in China is
unknown as yet, and the description here is taken from Dutta and Deb
(Taxon. Rev. Hedyotis. 2004, as H. puberula (G. Don) R. Brown ex
Arnott). The plant illustrated for this species (as H. umbellata) in FRPS
(71(1): 70, t. 13, f. 16. 1999) is actually H. pinifolia, and its description
there seems to combine some characters of both H. brevicalyx sensu
Dutta & Deb and H. pinifolia.

This species as provisionally circumscribed here is similar to

Hedyotis matthewii and slightly differently (and more narrowly) than
done by W. C. Ko (in FRPS 71(1): 48. 1999).

11. Hedyotis butensis Masamune, Trans. Nat. Hist. Soc. Formosa 28: 119. 1938.

13. Hedyotis capitellata Wallich ex G. Don, Gen. Hist. 3: 527.

1834.

tai wan er cao

tou hua er cao

Herbs, erect, annual, to 20 cm tall; stems terete to 4angled, hirsute. Leaves sessile; blade drying papery, elliptic-oblong, oblanceolate, ovate-lanceolate, or lanceolate, 1.56 0.3
1 cm, adaxially puberulent to scaberulous, abaxially densely
villous to hirsute, base cuneate to attenuate, apex acute to acuminate; stipules broadly triangular, ca. 3 mm, hirsute, acuminate. Inflorescences terminal and/or axillary, capitate or glomerulate; bracts linear, to 1.5 mm. Flowers sessile. Calyx hirsute;
hypanthium portion ovoid, ca. 1.5 mm; lobes linear to deltoid,
ca. 1.5 mm. Corolla white, funnelform, pubescent inside and

Herbs or shrubs, lianescent, climbing, clambering, or scandent, to 1 m; stems terete to weakly 4-angled, often longitudinally ridged and/or sulcate, glabrous or hirtellous to pilosulous or hispidulous at least in longitudinal lines, or tomentose in
var. mollissima. Leaves subsessile to petiolate; petiole to 4 mm,
glabrous to puberulent or hispidulous, or tomentose (var. mollissima); blade drying membranous, ovate, lanceolate, elliptic,
or elliptic-lanceolate, 212 14 cm, glabrous or sparsely to
moderately puberulent to hispidulous or hirtellous, or tomentose (var. mollissima), base acute to obtuse then often decurrent,

Sparse forests; 2001000 m. Guangdong.

RUBIACEAE

apex acute to shortly or long acuminate; secondary veins 3 or 4

pairs; stipules fused to petiole bases, glabrous to densely
puberulent, hispidulous, or strigillose, or tomentose (var. mollissima), triangular, 1.55 mm, entire or serrulate at margins,
acute to acuminate with tip 0.51.5 mm, often spreading to
recurved. Inflorescences terminal and often also in axils of uppermost leaves, 425 cm, with 315 globose, capitate to subcapitate heads in groups of 3, groups then paniculate, pyramidal, densely puberulent to hirtellous or glabrous, or tomentose (var. mollissima), pedunculate; peduncle 15 cm; bracts
subtending axes triangular to oblanceolate, 110 mm, bracts
subtending flowers reduced to a tuft of pubescence; heads 36
mm in diam. (not including corollas), 815 mm in diam. (including corollas). Flowers sessile or subsessile, distylous. Calyx
glabrous to densely puberulent; hypanthium portion campanulate to obconic, 0.81 mm; limb lobed for 1/33/4, 12 mm;
lobes oblong to narrowly triangular. Corolla white to pale blue,
funnelform, outside glabrous, inside densely bearded in throat
and onto lobes; tube 12 mm; lobes ligulate, 35 mm. Anthers
included or exserted, 11.2(2) mm. Stigmas linear, 12 mm.
Fruit capsular, ellipsoid to globose, 23 mm in diam., glabrescent, septicidally then loculicidally dehiscent (but see discussion below); seeds numerous, angled. Fl. almost year-round, fr.
not seen from China.
Broad-leaved forests in valleys, sunny mountain slopes; ca. 1500
m. Yunnan [India, Indonesia, Malaysia, Myanmar, Thailand, Vietnam].
The fruit is described above following W. C. Ko (in FRPS 71(1):
65. 1999); however, Dutta and Deb (Taxon. Rev. Hedyotis. 2004) described this differently, as loculicidal across the top beak portion, then
sometimes tardily septicidal.
W. C. Ko (loc. cit.: 6566) reported the varieties below from
China; they are treated following that work here for reference. Dutta
and Deb (loc. cit.) reported only Hedyotis capitellata var. subpubescens
Kurz from China; this name was not mentioned by Ko but presumably
corresponds to one of Dutta and Debs varieties.

1a. Plants glabrous .................................... 13a. var. capitellata

1b. Plants pubescent.
2a. Stems, inflorescences, calyx, and
leaf blade sparsely pubescent ............ 13b. var. mollis
2b. Stems, inflorescences, calyx, and
leaf blade densely pale yellow
tomentose .................................... 13c. var. mollissima
13a. Hedyotis capitellata var. capitellata
() tou zhuang hua er cao (yuan bian zhong)
Oldenlandia capitellata (Wallich ex G. Don) Kuntze; O.
capitellata var. glabra Pitard; O. rubioides Miquel.
Plants glabrous. Fl. May, fr. not seen from China.
Broad-leaved forests in valleys; ca. 1500 m [as low as below 100
m in India]. Yunnan [India, Indonesia, Malaysia, Myanmar, Thailand].

13b. Hedyotis capitellata var. mollis (Pierre ex Pitard) T. N.

Ninh in T. B. Nguyen, Fl. Taynguyen. Enum. 150. 1984.
shu mao tou zhuang hua er cao
Oldenlandia capitellata var. mollis Pierre ex Pitard in
Lecomte, Fl. Indo-Chine 3: 137. 1922.

157

Stems, leaf blade, inflorescences, and calyx sparsely pubescent. Fl. AprJul, fr. not seen from China.
Broad-leaved forests at middle elevations. Yunnan (Hekou) [India,
Indonesia, Malaysia, Vietnam].
Ninhs varietal combination was not known to W. C. Ko (loc. cit.:
66), who intended to publish it himself but was precluded.

13c. Hedyotis capitellata var. mollissima (Pitard) W. C. Ko,

Fl. Reipubl. Popularis Sin. 71(1): 66. 1999.
rong mao tou zhuang hua er cao
Oldenlandia capitellata var. mollissima Pitard in Lecomte,
Fl. Indo-Chine 3: 138. 1922.
Stems, leaf blade, inflorescences, and calyx densely pale
yellow tomentose. Fl. almost year-round, fr. not seen from
China.
Sunny mountain slopes at middle to high elevations. Yunnan (Xishuangbanna) [Vietnam].

14. Hedyotis capituligera Hance, J. Bot. 17: 12. 1879.

bai jiang er cao
Oldenlandia capituligera (Hance) Kuntze.
Leggy herbs, erect, to 80 cm tall; stems terete to 4-angled,
longitudinally usually sulcate, sparsely villosulous or hirtellous
to densely so at least in longitudinal lines, becoming glabrescent. Leaves subsessile to shortly petiolate; blade drying membranous, oblong-lanceolate, 3.511.5 0.83.5 cm, adaxially
glabrous or sparsely to densely puberulent, hirtellous, or hispid,
abaxially glabrescent on lamina and hispid to pilosulous or
pilose along veins, base acute to rounded, margins ciliolate,
apex obtuse to acute; secondary veins 58 pairs; stipules fused
to petiole bases, triangular, 1.54 mm, moderately to densely
hirtellous or villosulous, with 24 bristles or linear to narrowly
triangular lobes 1.54 mm. Inflorescences terminal on main
stems, short lateral stems, and sometimes also in axils of uppermost leaves, several to many flowered, with 18 congestedcymose to subcapitate heads or glomerules, these usually in
compound cymes 218 cm, densely villosulous to strigillose or
glabrescent, pedunculate; peduncles 110 cm; heads 0.81.5 cm
in diam.; bracts narrowly triangular to narrowly elliptic, 1.27
mm. Flowers subsessile. Calyx glabrous; hypanthium portion
subglobose, ca. 0.8 mm; limb lobed nearly to base; lobes triangular, 0.30.6 mm. Corolla white, funnelform, outside glabrous;
tube ca. 3 mm, stiffly barbate in throat; lobes ca. 1.3 mm, thickened at apex. Fruit capsular, compressed globose, dehiscent;
seeds several. Fl. JulAug.
Open grasslands. Guangdong, Guizhou, Yunnan.

15. Hedyotis cathayana W. C. Ko, Fl. Hainan. 3: 579. 1974.

zhong hua er cao
Herbs or subshrubs, presumably perennial, erect, to
80(100) cm tall; stems glabrous, 4-angled, angles sharp to
winged, wings to 2 mm wide. Leaves sessile or subsessile; blade
drying papery to membranous, purple on midrib at least when
young, elliptic-oblong, lanceolate, or oblanceolate, 1525 36

RUBIACEAE

158

cm, glabrous, base obtuse to rounded or truncate, apex acute to

long acuminate; secondary veins 7 or 8 pairs, hardly visible;
stipules apparently fused shortly around stem, broadly triangular, 412 mm, glabrous, with 2 winged flanges 12 mm wide
extending from each petiole base up to apex, acute to acuminate, entire, sparsely glandular-serrate, or with 1 to numerous
bristles 0.12 mm, gland-tipped. Inflorescence axillary, congested-cymose to shortly paniculate or subglomerulate, 1.53.5
cm, glabrous, subsessile to pedunculate; peduncles to 2 cm;
bracts linear-lanceolate, lanceolate, or narrowly elliptic, 112
mm; pedicels to 1.5 mm. Flowers subsessile to pedicellate,
?apparently distylous. Calyx glabrous; hypanthium turbinate,
11.3 mm; limb deeply lobed; lobes narrowly triangular to
spatulate, 23 mm, sometimes slightly unequal. Corolla white
or pale green, funnelform to tubular-funnelform, outside glabrous; tube 69 mm, bearded in throat; lobes spatulate-oblong,
2.54 mm. Anthers exserted, 11.5 mm. Stigma ca. 1.5 mm,
included, positioned just above middle of corolla tube. Fruit
capsular, subglobose to ovoid, 23 1.52 mm, septicidal
then loculicidal, papery, glabrous; seeds 46, blackish brown,
angled, papillose. Fl. and fr. almost year-round, often simultaneously.
On wet soil in valleys and ravines; ca. 500 m. Hainan.
The specimen H. Y. Liang 54367 (NY, Web!) was annotated as
Hedyotis Liangii Merr. & Chun, but this name was apparently never
published.

16. Hedyotis caudatifolia Merrill & F. P. Metcalf, J. Arnold

Arbor. 23: 228. 1942.
jian ye er cao
Hedyotis hui Diels.
Shrubs or perhaps subshrubs, erect, perennial, to 90 cm
tall; stems terete, shallowly striate, glabrous. Leaves petiolate;
petiole 1015 mm; blade drying leathery, grayish white abaxially, lanceolate, 613 1.53 cm, base cuneate or decurrent,
apex caudate-acuminate; secondary veins ca. 4 pairs but indistinct; stipules broadly ovate, 23 mm, entire or glandular-serrulate, acute. Inflorescences compound-cymose to paniculate,
several to many flowered; axes generally dichotomous; bracts
lanceolate or linear-lanceolate. Flowers shortly pedicellate. Calyx with hypanthium portion turbinate, ca. 3 mm; limb deeply
lobed; lobes ovate-triangular. Corolla white or pink, tubular,
outside glabrous; tube 48 mm, barbate in throat; lobes lanceolate. Stigmas exserted or included. Fruit capsular, ellipsoid-oblong or ellipsoid, ca. 2.5 2 mm, smooth, glabrous, dehiscent;
seeds several, black, subtriangular. Fl. MayJun.
Dry soil in jungles, thickets, on rock cliffs, on clay soil in grasslands. Fujian, Guangdong, Guangxi, Hunan, Jiangxi, Zhejiang.
W. C. Ko (in FRPS 71(1): 48. 1999) described the petioles of this
species as 1015 mm and the hypanthium as ca. 3 mm, but the accompanying figure (p. 50, t. 7, f. 710) does not show these features.

17. Hedyotis cheniana R. J. Wang, Novon 18: 264. 2008.

huan yong er cao
Subshrubs, erect, to 0.5 m tall; stems terete to sulcate,
glabrous to densely puberulent, strigillose, or pilosulous. Leaves

petiolate; petiole 35 mm, glabrous to densely pilosulous; blade

drying papery, ovate, elliptic, ovate-oblong, or suborbicular,
1.75 1.62.5 cm, adaxially glabrescent, abaxially puberulent
to pilosulous, base broadly obtuse to rounded or truncate, apex
acute or shortly acuminate; secondary veins 3 or 4 pairs, poorly
to well developed abaxially; stipules persistent, interpetiolar,
triangular, 23.5 mm, glabrous or densely pilosulous, lacerate
or with 59 linear lobes or bristles. Inflorescences terminal and
in uppermost leaf axils, subcapitate, 12 cm in diam., 1030flowered, puberulent to pilosulous, subsessile to pedunculate;
peduncles to 3 mm; bracts narrowly triangular to linear, 13
mm; pedicels 0.52 mm. Flowers distylous, shortly pedicellate.
Calyx glabrous; hypanthium portion subglobose-ellipsoid, 0.8
1 mm; limb lobed nearly to base; lobes narrowly triangular,
3.54 mm, entire to ciliolate, acuminate. Corolla white, salverform, outside glabrous; tube ca. 5 mm, barbate in throat; lobes
spatulate-oblong to lanceolate, ca. 2 mm. Anthers exserted or
situated in corolla throat, 11.5 mm. Stigmas included or exserted, ca. 1 mm. Fruit capsular, subglobose, ca. 2 mm, glabrous, loculicidal then septicidal; seeds 610, angled, reticulate.
Fl. JanFeb, fr. JunNov.
Dense forests; 6001000 m. Hainan.
The protologue described Hedyotis cheniana as being glabrous,
but two specimens cited there are in fact densely pubescent; the protologue also described the secondary veins as not visible, but in fact they
are well developed and usually prominulous abaxially on these specimens.

18. Hedyotis chereevensis (Pierre ex Pitard) Fukuoka, S. E.

Asian Stud. 8(3): 332. 1970.
yue nan er cao
Oldenlandia chereevensis Pierre ex Pitard in Lecomte, Fl.
Indo-Chine 3: 143. 1922.
Herbs, erect, perhaps perennial, to 50 cm tall; stems flattened to 4-angled, often ridged, glabrous or puberulent to hispidulous at and shortly below nodes. Leaves subsessile to petiolate; petiole to 2 mm, puberulent to hispidulous; blade drying
membranous, ovate, lanceolate, or elliptic, 27 0.83 cm,
adaxially glabrous or puberulent on principal veins, abaxially
sparsely to densely puberulent or hispidulous at least on principal veins, base cuneate to rounded then decurrent, apex acute
to acuminate; secondary veins 57 pairs; stipules fused to petiole bases, ovate-lanceolate to triangular, 35 mm, densely puberulent, hirtellous, or hispidulous, acuminate to aristate, marginally laciniate or with several filamentous lobes to 1 mm,
often glandular at apex. Inflorescences terminal and sometimes
in axils of uppermost leaves, cymose to paniculate, 37 cm,
several to many flowered, glabrous, usually slender, pedunculate; peduncles 12 cm; bracts lanceolate, 0.81 mm; pedicels
610 mm. Flowers all pedicellate. Calyx glabrous; hypanthium
portion obconical, 0.50.8 mm; limb lobed essentially to base;
lobes triangular, 0.10.3 mm. Corolla white, tubular to rotate,
outside glabrous; tube 0.51 mm, inside apparently glabrous;
lobes oblong-lanceolate, ca. 0.5 mm. Anthers exserted, ca. 0.5
mm. Stigma apparently exserted, ca. 0.2 mm. Fruit capsular,
obconical, turbinate, or subglobose, often somewhat didymous,
1.52 11.5 mm, glabrous, loculicidally dehiscent across top,

RUBIACEAE

beak rounded, ca. 0.4 mm high; seeds ca. 10, dark brown,
angled. Fl. MayJul.
On damp humus soil in forests; [below 1001300 m in Thailand].
Hainan [Cambodia, Thailand, Vietnam].
The name Hedyotis chereevensis has been attributed to W. C. Ko
(Fl. Hainan. 3: 306, 579. 1974), but this appears to be a later, superfluous combination.

19. Hedyotis chrysotricha (Palibin) Merrill, Lingnan Sci. J. 7:

322. 1931.
jin mao er cao
Anotis chrysotricha Palibin, Bull. Herb. Boissier, sr. 2, 6:
20. 1906; Oldenlandia chrysotricha (Palibin) Chun.
Herbs, perennial or perhaps annual, procumbent, usually
regularly rooting at nodes, to 40 cm; stems weakly 4-angled,
flattened, or terete, sometimes 2-sulcate, moderately to densely
hirtellous to pilosulous at least along grooves, sometimes glabrescent, trichomes often drying yellowed. Leaves subsessile to
petiolate; petiole 13 mm, moderately to densely pilosulous to
hirtellous; blade drying thinly papery, lanceolate, elliptic, or
ovate, 0.83.2 0.51.4 cm, adaxially sparsely hirtellous, hispidulous, or scaberulous at least along margins, abaxially
sparsely to densely hirtellous or pilosulous with pubescence
denser along principal veins or sometimes glabrous (e.g., Tsang
20859, MO!), base cuneate or obtuse, apex obtuse to acute or
shortly acuminate; secondary veins 2 or 3 pairs; stipules fused
to petiole bases, triangular, 12 mm, moderately to densely puberulent to hirtellous, with 13 linear lobes or bristles 1.53.5
mm, these sometimes glandular and/or bifid at apex. Inflorescences axillary, congested-cymose to fasciculate, 310 mm in
diam., (1 or)25-flowered in each axil, moderately to densely
pilosulous, sessile or subsessile; bracts linear, lanceolate, or narrowly elliptic, 0.54 mm; pedicels to 4 mm. Flowers subsessile
to pedicellate, distylous. Calyx sparsely to densely hirtellous;
hypanthium portion subglobose to obconic, ca. 1 mm; limb
lobed essentially to base; lobes lanceolate to triangular, ca. 2
mm, usually ciliolate to ciliate. Corolla white or purple, funnelform, outside pilosulous to glabrous; tube 2.54 mm, pubescent
inside; lobes linear-oblong to triangular, 2.53 mm. Anthers exserted or included, 11.2 mm. Stigma exserted or included, 1
1.2 mm. Fruit indehiscent, subglobose to ovoid, 1.52 22.5
mm, with calyx lobes to 3 mm; seeds several. Fl. and fr. yearround.
Broad-leaved forests in valleys, thickets on mountain slopes; 100
900 m. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei,
Hunan, Jiangsu, Jiangxi, Taiwan, Yunnan, Zhejiang [Japan, ?Philippines].

20. Hedyotis communis W. C. Ko, Fl. Hainan. 3: 579. 1974.

da zhong er cao
Subshrubs or herbs, erect, to 1.5 m tall; stems glabrous,
flattened to 4-angled or terete. Leaves sessile to subsessile;
blade drying papery, narrowly elliptic-oblong or narrowly elliptic, on vegetative stems, 1017 1.52.6 cm, glabrous, base
cuneate to acute then usually long decurrent, apex acute to acuminate; secondary veins 68 pairs, abaxially indistinct; stipules

159

shortly united around stem or sometimes fused to petiole bases,

triangular to narrowly triangular, 813 mm, glabrous, long acuminate, marginally sometimes sparsely glandular-serrulate or
-setose. Inflorescence axillary, paniculate-cymose with small
cymes distributed along axes with leaflike bracts and bracteal stipules, or along axillary branches with reduced leaves
(depending on interpretation), 724 cm, many flowered, glabrous, pedunculate; peduncle 0.54 cm; leaflike bracts or reduced leaves paired, lanceolate to narrowly elliptic, 1040 mm;
cymes 815 mm, several flowered; floral bracts linear or linearlanceolate, 28 mm; pedicels to 1 mm. Flowers subsessile to
pedicellate. Calyx glabrous; hypanthium portion ellipsoid, 1
1.2 mm; limb lobed nearly to base; lobes narrowly triangular,
23 mm. Corolla white, tubular, outside glabrous; tube 4.55
mm, bearded in throat; lobes spatulate-oblong to triangular, 2
2.5 mm, acute. Anthers exserted, ca. 1.2 mm. Stigma included,
ca. 0.8 mm. Fruit capsular, obovoid or subglobose, 23 22.5
mm, glabrous, septicidally then loculicidally dehiscent, cartilaginous; seeds 24, brownish black, tuberculate. Fl. almost
year-round, fr. May, Jun, Oct.
On soil rich in humus in valleys or at streamsides; 9001000 m.
Hainan.
The inflorescences are borne laterally along well-developed axes
with numerous leaflike bracts or reduced leaves, or alternatively along
lateral branches with reduced leaves; various morphological descriptions of these have been used by different authors.

21. Hedyotis consanguinea Hance, Ann. Sci. Nat., Bot., sr. 4,

18: 221. 1862.
ni jin cao
Hedyotis lancea Thunberg ex Maximowicz; Oldenlandia
consanguinea (Hance) Kuntze; O. lancea (Thunberg ex Maximowicz) Kuntze.
Herbs or subshrubs, erect, perennial, to 40 cm tall; stems
terete, flattened, or weakly 4-angled, smooth to sulcate, glabrous. Leaves subsessile to petiolate, sometimes appearing verticillate due to axillary groups of reduced leaves on unexpanded
lateral stems; petiole to 6 mm, glabrous; blade drying stiffly
papery to leathery, lanceolate, ovate, elliptic, lanceolate-elliptic,
or elliptic-oblong, 1.512 0.44 cm, glabrous, base acute to
obtuse and usually decurrent, margins plane to thinly revolute,
apex acute to usually acuminate; secondary veins 24 pairs but
mostly indistinct; stipules fused around stem, ovate to broadly
triangular, 25.5 mm, glabrous, marginally glandular-serrulate,
acute to acuminate or shortly aristate. Inflorescences terminal
and in axils of uppermost leaves, compound-cymose, paniculate, 315 cm, many flowered, glabrous, pedunculate; peduncle
17 cm; axes strictly dichotomous then often spiciform (due to
development of only 1 lateral axis, leaving 1 or 2 flowers at
node), or sometimes scorpioid (due to development of alternating axes); bracts triangular to elliptic, 0.85 mm or those
subtending principal axes sometimes leaflike; pedicels to 1 mm.
Flowers sessile to shortly pedicellate, apparently distylous. Calyx glabrous; hypanthium portion obconic, 0.81 mm; limb
deeply lobed; lobes lanceolate to triangular, 0.81.2 mm. Corolla white, salverform, outside glabrous; tube 1.52.5 mm,
inside sparsely villosulous; lobes lanceolate, 1.52.5 mm. An-

160

RUBIACEAE

thers included or exserted, ca. 0.8 mm. Stigma apparently included or exserted, ca. 0.3 mm. Fruit capsular, ellipsoid, ca. 2
2 mm, glabrous, woody, flattened to rounded on top, septicidally then loculicidally dehiscent; seeds numerous, blackish
brown, angled. Fl. and fr. JunNov.

though not completely following Sivarajan and Biju (loc. cit.) and W. C.
Ko (in FRPS 71(1): 72, 75. 1999). However, W. C. Ko described the
hypanthium of H. corymbosa as 11.2 mm in diam., which corresponds
to H. diffusa in most recent classifications rather than H. corymbosa.
The varieties recognized by W. C. Ko are presented here for reference.

Grasslands, ditch sides; 4001000 m. Fujian, Guangdong, Hainan, Zhejiang.

1a. Stems quadrangular, glabrous to

pubescent and/or scaberulous along
angles .................................................. 22a. var. corymbosa
1b. Stems terete to flattened, farinose
puberulent throughout ........................ 22b. var. tereticaulis

In the description here a distinction is made between the inflorescence axes and the pedicels; measurements in other descriptions sometimes include inflorescence axes that bear only a developed flower
together with an undeveloped bud on an apparently indeterminate segment as pedicels and, thus, are correspondingly longer.

22. Hedyotis corymbosa (Linnaeus) Lamarck, Tabl. Encycl. 1:

272. 1792.
san fang hua er cao
Slender herbs, annual, to 40 cm tall; stems 4-angled to
flattened, 2-sulcate, with sides glabrous and angles thickened to
winged and puberulent and/or scaberulous (var. corymbosa) or
terete to flattened and sparsely to densely scaberulous to puberulent throughout (var. tereticaulis). Leaves subsessile; blade
drying membranous, linear, narrowly lanceolate, or narrowly
elliptic, 0.82 0.10.5 cm, adaxially sparsely scaberulous to
glabrescent, abaxially glabrescent, base cuneate to acute, margins usually at least shortly revolute at least when dry, apex
acute; secondary veins not visible; stipules fused to petiole
bases, tubular, 12 mm, puberulent to glabrous, rounded to
triangular, with 15(or 7) linear lobes or bristles 0.22.5 mm,
these sometimes bifid. Inflorescence axillary, 1-flowered or usually cymose and 25-flowered, glabrous, pedunculate; peduncles 1(or 2) per axil, filiform, 116 mm; bracts lacking or
stipuliform and 11.2 mm; pedicels slender, 212 mm. Flowers
homostylous, pedicellate. Calyx glabrous to puberulent; hypanthium portion subglobose to narrowly ellipsoid, 0.50.8 mm;
limb lobed essentially to base; lobes narrowly triangular, 0.5
1.2 mm, entire to ciliate. Corolla white or pink, funnelform to
rotate; tube 0.81 mm, inside pubescent or glabrous; lobes spatulate-oblong to narrowly triangular, 0.50.8 mm. Anthers included, ca. 0.6 mm. Stigma 0.30.5 mm, included. Fruit capsular, subglobose, ovoid, or oblate, 1.22 1.22.2 mm, somewhat dicoccous, membranous, glabrous to puberulent, loculicidally dehiscent through flat to broadly rounded apex, beak
when present to 0.5 mm, peduncles and pedicels usually elongating rapidly and markedly as fruit develop; seeds 20 or more,
dark brown, angled, smooth. Fl. and fr. almost year-round.
Paddy fields, ridges of farmlands, humid grasslands; sea level to
900 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, Zhejiang [Africa and tropical Asia; widely adventive in the Americas and the Pacific region].
These plants are low annual weeds found in highly disturbed habitats and show wide morphological variation between sites and usually
among plants in one site in plant size, leaf width, number of peduncles
per axil, and number of flowers per peduncle. The taxonomy of this and
related species is complicated, and different authors have drawn very
different conclusions (e.g., Sivarajan & Biju, Taxon 39: 665674. 1990;
Dutta & Deb, Taxon. Rev. Hedyotis. 2004). In particular, Hedyotis
brachypoda, H. diffusa, and H. erecta are related and have been variously circumscribed. Here, these species are circumscribed generally

22a. Hedyotis corymbosa var. corymbosa

() san fang hua er cao (yuan bian zhong)
Oldenlandia corymbosa Linnaeus, Sp. Pl. 1: 119. 1753;
Pharaceum incanum Loureiro.
Stems quadrangular, glabrous to pubescent and/or scaberulous along angles. Fl. and fr. almost year-round.
Paddy fields, ridges of farmlands, humid grasslands. Fujian,
Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Zhejiang [Africa,
America, tropical Asia].

22b. Hedyotis corymbosa var. tereticaulis W. C. Ko, Fl. Hainan. 3: 580. 1974.
yuan jing er cao
Hedyotis burmanniana Wight & Arnott (1834), not
Schultes (1827); H. pseudocorymbosa Bakhuizen f.; Oldenlandia pseudocorymbosa (Bakhuizen f.) Raizada.
Stems terete to flattened, farinose puberulent throughout.
Fl. and fr. almost year-round.
Ridges of farmlands, humid open fields. S and SW China.
This variety may correspond to Hedyotis diffusa as circumscribed
by Sivarajan and Biju (loc. cit.) and to H. pseudocorymbosa as circumscribed by some other authors (e.g., Bakhuizen f., Fl. Java 2: 286. 1965;
Dutta & Deb, loc. cit.).

23. Hedyotis cryptantha Dunn, Bull. Misc. Inform. Kew 1912:

367. 1912.
bi hua er cao
Oldenlandia cryptantha (Dunn) Chun.
Herbs, perennial, procumbent, to 50 cm tall; stems 4-angled, sulcate, glabrous to puberulent, sometimes with wings to
4 mm wide. Leaves sessile to petiolate, sometimes clustered
near top of stem; petiole to 15 mm; blade with midrib often
purple, drying papery, elliptic, oblanceolate, or obovate, 1530
510 cm, glabrous (but sometimes when dry appearing puberulent due to collapsed large epidermal cells), base acute to
attenuate and long decurrent, apex acute to long acuminate;
secondary veins 57 pairs, sometimes with well-developed intersecondary veins; stipules interpetiolar, narrowly triangular,
830 mm, glabrous, laciniate or with 5 to numerous setiform
segments or lobes 120 mm, these sometimes apparently glandular at apex. Inflorescences axillary at nodes well below stem
apex, capitate, subcapitate, or congested-cymose, 1.52.5 cm in

RUBIACEAE

diam., several to many flowered, glabrous, sessile; bracts linearlanceolate, 58 mm, acute to erose or shortly laciniate. Flowers
sessile or subsessile, biology unknown. Calyx glabrous; hypanthium portion subglobose, ca. 2 mm; limb lobed essentially to
base; lobes linear-lanceolate, 68 mm. Corolla white to pale
purple, salverform or tubular, outside apparently glabrous; tube
1416 mm, inside tomentulose from above middle to throat;
lobes narrowly ovate, ca. 3 mm. Anthers included, 11.2 mm.
Fruit indehiscent, obovoid, laterally somewhat flattened, ca. 3
2 mm, glabrous, with calyx lobes to 10 mm; seeds angled,
black, foveolate. Fl. SepNov, fr. OctJan.
Forests in humid shady valleys, rock crevices at streamsides;
3001000 m. Hainan (Lingshui).

24. Hedyotis dianxiensis W. C. Ko, J. S. China Agric. Univ.

16(4): 44. 1995.
dian xi er cao
Herb, climbing to scandent; stems flattened to subterete,
often angled and/or sulcate, densely villous and usually also
tomentose. Leaves petiolate; petiole 12.5 mm, densely villous
or tomentose; blade drying papery, lanceolate, lanceolate-elliptic, or narrowly ovate, 29 0.63 cm, adaxially moderately to
densely hispidulous with pubescence denser on principal veins,
abaxially densely pilosulous to hirtellous throughout, base cuneate to rounded, apex acute to long acuminate; secondary
veins 35 pairs; stipules interpetiolar, broadly rounded to truncate, 13.5 mm, densely villosulous or pilosulous, with 39
bristles or linear lobes 16 mm. Inflorescences terminal and in
axils of uppermost leaves, capitate, subcapitate, or glomerulate,
densely pilosulous or hirtellous, sessile or pedunculate; peduncle 212 mm; heads 110, solitary or borne separately along
peduncles or axes, each subglobose, 12 cm in diam., 1530flowered; bracts reduced. Flowers sessile or subsessile, distylous. Calyx with hypanthium portion turbinate to subglobose,
ca. 1 mm, densely hispid; limb deeply lobed; lobes spatulateoblong, triangular, or lanceolate, 1.22 mm, hispidulous to glabrescent. Corolla yellowish green, funnelform, outside glabrous
to pilosulous, inside densely bearded in throat and onto lobes;
tube ca. 2 mm; lobes spatulate-oblong to triangular, ca. 4 mm.
Anthers 0.81.2 mm, exserted or included. Stigma 0.50.8 mm,
exserted or included. Fruit capsular, subglobose to obovoid,
3.54 mm, pilosulous or hirtellous, cartilaginous, septicidally
then later loculicidally dehiscent, with conical beak ca. 1 mm;
seeds several, angled. Fl. FebMay, fr. Jun.
Humid thickets at streamsides; 600800 m. Yunnan (Xishuangbanna).
Although all specimens seen of this species are pubescent, which
agrees with the descriptions available, it is keyed out here also with
glabrescent stems in case glabrous individuals are eventually found, as
in most pubescent Rubiaceae species.

25. Hedyotis diffusa Willdenow, Sp. Pl. 1: 566. 1798.

bai hua she er cao
Hedyotis herbacea Loureiro (1790), not Linnaeus (1753);
Oldenlandia diffusa (Willdenow) Roxburgh; O. herbacea (Linnaeus) Roxburgh var. uniflora Bentham.
Slender herbs, annual, ascending to procumbent, to 50 cm

161

tall; stems slightly flattened to terete or young stems sometimes

4-angled, sparsely to densely puberulent, scaberulous, or glabrescent to glabrous, similarly glabrous or pubescent on angles
and to sides. Leaves sessile or subsessile; blade drying membranous, linear, narrowly elliptic, or narrowly oblanceolate, 14
0.10.4 cm, adaxially glabrous and smooth or often scaberulous near margins, abaxially glabrous to scaberulous, base
acute, margins usually revolute at least when dry, apex acute;
secondary veins not visible; stipules fused to petiole bases, triangular to truncate, 0.51.5 mm, glabrescent, acute to aristate or
with 13 bristles 0.21 mm. Inflorescences axillary, 1-flowered
or fasciculate and 2-flowered, glabrous, pedunculate; peduncles
or pedicels 420 mm; bracts none or stipuliform, to 1 mm.
Flowers pedicellate, apparently homostylous. Calyx glabrous;
hypanthium portion subglobose, 11.2 mm; limb lobed essentially to base; lobes narrowly triangular, 12 mm, ciliolate. Corolla white, tubular, outside glabrous; tube 1.52.5 mm, glabrous inside; lobes ovate-oblong, 1.22 mm. Anthers ca. 0.8
mm, exserted. Stigma ca. 1.2 mm, exserted. Fruit capsular, compressed globose to oblate, 23 23 mm, sometimes somewhat
dicoccous, membranous, glabrous, loculicidally dehiscent on
flat to beaked top, beak rounded, to 0.5 mm, with peduncles
elongating rapidly and markedly as fruit mature, to 20 mm;
seeds ca. 20, dark brown, angled, deeply thickly foveolate. Fl.
and fr. MayOct.
Paddy fields, ridges of farmlands, humid open fields; sea level to
900 m. Anhui, Fujian, Guangdong, Guangxi, Hainan, Taiwan, Yunnan,
Zhejiang [Bangladesh, Bhutan, Indonesia, Japan, Malaysia, Nepal,
Philippines, Sri Lanka, Thailand].
This species is very commonly collected in China. The taxonomy
of this and related species is complicated, and different authors have
drawn very different conclusions (e.g., Sivarajan & Biju, Taxon 39:
665674. 1990; Dutta & Deb, Taxon. Rev. Hedyotis. 2004). In particular, Hedyotis brachypoda, H. corymbosa, and H. erecta are related and
have been variously circumscribed. Here, these species are circumscribed generally, though not completely, following W. C. Ko (in FRPS
71(1): 72, 75. 1999) and Dutta and Deb.

26. Hedyotis effusa Hance, J. Bot. 17: 11. 1879.

ding hu er cao
Oldenlandia effusa (Hance) Kuntze.
Herbs or subshrubs, perennial, erect, to 1 m or taller; stems
rounded, flattened, or 4-angled, glabrous. Leaves subsessile to
petiolate; petiole to 5 mm, glabrous; blade drying papery or
leathery, ovate, ovate-lanceolate, or lanceolate-elliptic, 413
1.56.5 cm, glabrous, base cuneate to rounded, apex acute;
secondary veins indistinct; stipules united around stem, broadly
triangular or truncate, 13 mm, glabrous, entire, sometimes partially costate, acute to mucronate. Inflorescences terminal, compound-cymose to paniculate, 515 410 cm, glabrous, several
to many flowered, pedunculate; peduncles 27 cm; axes regularly dichotomous, spreading at 90 or more; bracts triangular
to linear, 0.55 mm; pedicels 25 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion ovoid, 0.71 mm; limb 0.5
0.8 mm, lobed for 1/22/3, lobes triangular. Corolla white, funnelform to tubular, outside glabrous; tube ca. 1.5 mm, apparently barbate in throat; lobes narrowly triangular, ca. 1.5 mm.
Anthers and stigmas not seen. Fruit capsular, subglobose, 1.5

RUBIACEAE

162

2.5 mm, glabrous, stiffly papery to cartilaginous, septicidally

then loculicidally dehiscent; seeds several, angled. Fl. JulSep,
fr. AugMar.
Forests or streamsides in valleys, sometimes on humid hill
slopes; 200400 m. Guangdong, Guangxi.

27. Hedyotis exserta Merrill, Lingnan Sci. J. 13: 72. 1934.

chang hua zhou er cao
Herbs or subshrubs, erect, perennial, to 70 cm; stems terete
to flattened, smooth or sulcate, glabrous. Leaves petiolate; petiole 25 mm, puberulent; blade drying papery, lanceolate, narrowly elliptic, or narrowly lanceolate, 5.511 0.83 cm, adaxially glabrous or papillose at least near margins, abaxially glabrous, base cuneate to acute, apex acute to long acuminate;
secondary veins 46 pairs but indistinct; stipules fused to petiole bases, triangular to broadly triangular, 38 mm, glabrous or
papillose-puberulent, smooth or costate, entire or serrulate, acute
to caudate-acuminate. Inflorescences terminal and in uppermost
leaf axils, congested-cymose, fastigiate to subglobose, 23 cm,
densely puberulent, several to many flowered, pedunculate; peduncle 5.59 cm; bracts triangular, oblong-ovate, or narrowly
elliptic, 0.86 mm; pedicels 12 mm. Flowers pedicellate. Calyx densely papillose or farinose puberulent; hypanthium portion turbinate, 11.5 mm; limb deeply lobed; lobes oblongovate to triangular, 11.8 mm. Corolla purple or pink, salverform, outside farinose puberulent; tube 710 mm, inside pilose;
lobes oblong-ovate, 1.22 mm. Anthers positioned in throat.
Stigmas included, positioned near middle of corolla tube, ca. 2
mm. Fruit capsular, subglobose to ellipsoid and somewhat laterally flattened, 22.5 mm in diam., puberulent, flat on top, dehiscent; seeds several, blackish brown, angled. Fl. MaySep.
Slopes or thickets in valleys. Hainan [Vietnam].

28. Hedyotis hainanensis (Chun) W. C. Ko, Fl. Hainan. 3: 580.

1974.
hai nan er cao
Oldenlandia hainanensis Chun, Sunyatsenia 1: 311. 1934,
based on Hedyotis oligantha Merrill, Philipp. J. Sci. 23: 266.
1923, not Merrill (1921).
Shrubs, erect, to 30 cm tall; stems terete, densely puberulent to pilosulous. Leaves subsessile to petiolate; petiole to 2
mm, densely puberulent to pilosulous; blade drying papery,
subelliptic, broadly elliptic, or ovate, 12.5 0.81.5 cm, adaxially glabrous, abaxially densely puberulent at least along midrib, base obtuse to cuneate, margins sometimes thinly revolute
when dry, apex obtuse, acute, or shortly acuminate; secondary
veins ca. 3 pairs but indistinct; stipules fused to petiole bases,
triangular or ovate, ca. 0.5 mm, puberulent to pilosulous, with 3
short narrow lobes. Inflorescences terminal and also in axils of
uppermost leaves, capitate or 2-flowered, sessile. Flowers sessile. Calyx sparsely puberulent; hypanthium portion ovoid, ca. 2
mm; limb lobed nearly to base; lobes 5, narrowly lanceolate, 3
3.5 mm. Corolla blue or white, funnelform, outside glabrous;
tube ca. 8 mm, bearded in throat; lobes 5, oblong-lanceolate, ca.
4 mm. Anthers 5, included, ca. 1.8 mm. Stigmas not seen. Fruit
not seen. Fl. MayJun.
Dense forests. Hainan (Dingan).

The protologue did not describe the fruit; these were described by
W. C. Ko (in FRPS 71(1): 62. 1999, modified to format here) as Fruit
capsular, globose, laterally flattened, ca. 2 mm in diam., sparsely hispidulous, septicidally dehiscent then loculicidally dehiscent, with persistent calyx lobes with the same fruiting period as the flowers. However, this description exactly matches the fruit of a group of plants
that have been included in Hedyotis hainanensis but do not match the
type and are provisionally separated here.

29. Hedyotis hedyotidea (Candolle) Merrill, Lingnan Sci. J.

13: 48. 1934.
niu bai teng
Spermacoce hedyotidea Candolle, Prodr. 4: 555. 1830;
Hedyotis esquirolii H. Lveill; H. macrostemon Hooker &
Arnott; H. nantoensis Hayata; H. recurva Bentham; Oldenlandia esquirolii (H. Lveill) Chun; O. macrostemon (Hooker
& Arnott) Kuntze.
Shrubs or subshrubs, lianescent, clambering, scandent, or
climbing, to 5 m; stems flattened to subterete then often later 4angled, densely papillose to farinose puberulent or velutinousstrigillose. Leaves petiolate; petiole 110 mm, densely papillose, puberulent, or strigillose; blade drying membranous to papery, elliptic-oblong, lanceolate, elliptic, or ovate, 410 1.54
cm, adaxially glabrous or densely papillose to puberulent on
principal veins, abaxially glabrous or densely papillose to puberulent or occasionally velutinous-strigillose, base cuneate to
rounded, apex acute or shortly acuminate; secondary veins 35
pairs; stipules fused to petioles, truncate to broadly rounded,
1.83 mm, glabrous to densely papillose, puberulent, or velutinous-strigillose, entire to densely ciliolate and/or with 37 narrowly triangular to linear lobes or bristles 17 mm, at least
sometimes gland-tipped. Inflorescences terminal and in axils of
uppermost leaves, subcapitate, congested-cymose, compoundcymose, or paniculate, 116 cm, several flowered, with 110
congested to laxly cymose groups of flowers, densely granularpuberulent to tomentulose-velutinous, pedunculate; peduncles
0.52.5 cm; flower groups subglobose to hemispherical, 11.5
cm in diam. (not including corollas), 1.83 cm in diam. (including corollas); bracts triangular, narrowly elliptic, or narrowly lanceolate, 0.22 mm; pedicels 0.82 mm. Flowers pedicellate, distylous. Calyx densely puberulent; hypanthium portion turbinate, 11.2 mm; limb 22.5 mm, lobed essentially to
base; lobes linear-lanceolate to spatulate, usually reflexed at anthesis. Corolla white to pale yellow, tubular to funnelform, outside glabrous, inside densely bearded in throat and onto lobes;
tube 23 mm; lobes narrowly lanceolate to narrowly spatulate,
44.5 mm. Anthers exserted or long exserted, 1.22 mm. Stigmas exserted or long exserted, 0.81 mm in long-styled form,
ca. 2 mm in short-styled form. Fruit capsular, subglobose, obovoid, or oblong-ellipsoid, 2.53 mm, densely puberulent to glabrescent, cartilaginous to very stiffly papery, loculicidally then
septicidally dehiscent through apical beak, beak 0.51 mm;
seeds several, angled. Fl. and fr. AprDec.
Thickets in ravines, hill slopes; 2001000 m. Fujian, Guangdong,
Guangxi, Guizhou, ?Hainan, Taiwan, Yunnan [Cambodia, Thailand,
Vietnam].
The flowers appear to be sessile in heads, but closer study shows
they are shortly pedicellate. W. C. Ko (in FRPS 71(1): 67. 1999) de-

RUBIACEAE

scribed the leaves as scabrous above, but this has not been seen nor reported by other authors. Ko also described the anthers of the long-styled
flowers as included within the corolla, but they are all distinctly exserted on the specimens studied; this character has not been specifically
described by other authors.
W. C. Ko (loc. cit.) said that this species was treated as Hedyotis
fruticosa by Kuntze (Obs. 2: 8. 1781), but in fact Kuntzes name was
not applied to the same species described by Linnaeus. The name H.
nantoensis was overlooked by the Fl. Taiwan (ed. 2, 4: 265273. 1998)
but seems to be validly published and clear as to its identity. Fukuoka
(S. E. Asia Stud. 8(3): 326. 1970) reported this species from Hainan
based on plants here treated as H. obliquinervis, which he treated as a
variety of H. hedyotidea.

30. Hedyotis herbacea Linnaeus, Sp. Pl. 1: 102. 1753.

dan cao
Hedyotis heynii (G. Don) Beddome; Oldenlandia herbacea
(Linnaeus) Roxburgh; O. heynii G. Don.
Herbs, annual or biennial, generally erect, to 60 cm; stems
weakly to sharply 4-angled, glabrous to scaberulous at least on
angles. Leaves sessile or subsessile; blade linear or linear-lanceolate, 12.5 0.10.3 mm, glabrous to scaberulous, base
acute to obtuse, margins weakly to strongly revolute, apex
acute; secondary veins not visible; stipules reduced or fused to
petiole bases, glabrous to scaberulous, truncate to broadly triangular, to 0.3 mm, entire or with 15 triangular to linear lobes
or bristles 0.22 mm. Inflorescences axillary, 1-flowered or
several flowered and fasciculate to cymose, glabrous, apparently ebracteate, pedunculate; peduncles 13 per axil, 130
mm; pedicels 130 mm. Flowers homostylous or heterostylous,
subsessile to pedicellate. Calyx glabrous to puberulent; hypanthium portion subglobose to ovoid, 0.81 mm; limb lobed essentially to base; lobes narrowly triangular to linear, 0.51.5
mm. Corolla white to reddish or pale purple, funnelform, outside glabrous; tube 23 mm, glabrous at throat; lobes spatulateoblong, 0.51 mm. Anthers exserted or included, 0.20.4 mm.
Stigma ca. 0.8 mm, included or exserted. Fruit capsular, ovoid
to subglobose, 22.5 22.5 mm, loculicidally dehiscent
through apical beak 11.5 mm; seeds several, dark brown,
foveolate. Fl. and fr. Jan, MarApr.
On humid rocks. Fujian, Guangdong, Guangxi, Hainan, Jiangxi
[widespread in tropical Africa and Asia].
Very few specimens have been seen of this species from China (or
anywhere else east of Sri Lanka); the description here, therefore, is
based primarily on plants from India. W. C. Ko (FRPS 71(1): 73. 1999)
described the hypanthium as 1.82 mm, the calyx lobes as 2.52.8 mm
and fimbriate-serrulate, the corolla lobes as ca. 2 mm, and the anthers as
ca. 1.5 mm, but these features do not agree with specimens of Hedyotis
herbacea; they do apply to the rather similar species H. brachypoda and
H. diffusa.

163

adaxially sparsely or moderately hispidulous to glabrescent,

abaxially glabrous, base acute to obtuse, margins scaberulous,
apex obtuse to acute; secondary veins 2 or 3 pairs but mostly
indistinct; stipules fused to petiole bases, broadly triangular,
0.51.5 mm, hirtellous or puberulent to glabrescent, rounded to
truncate, sometimes apiculate or with 2 or 3 bristles to 1 mm.
Inflorescences terminal and in axils of uppermost leaves, fasciculate to loosely dichasial, few flowered, glabrous; peduncles
and/or pedicels 215 cm; bracts reduced or triangular to setiform, to 1 mm. Flowers pedicellate or pedunculate, floral biology unknown. Calyx glabrous; hypanthium obconic, ca. 1 mm;
limb lobed nearly to base; lobes ovate, ca. 1.2 mm, sometimes
with evident venation, ciliolate. Corolla white sometimes tinged
with pink, salverform, outside glabrous; tube ca. 1.5 mm, inside
glabrous; lobes ovate, ca. 0.5 mm. Anthers and stigmas not
seen. Fruit capsular, oblate-cupuliform, 2.53 3.54 mm, glabrous, smooth, loculicidally dehiscent across top then sometimes septicidal, with beak to 1 mm, with calyx lobes becoming
elliptic to ovate, to 2 mm; seeds numerous, subglobose.
Mountains; ca. 1600 m. Yunnan [India, Sri Lanka].
The description here is based on Chinese material. Dutta and Deb
(Taxon. Rev. Hedyotis. 2004) described the corolla of Hedyotis hermanniana in India as having tubes 34 mm and lobes 1.52 mm,
although their figure shows a corolla with a tube slightly over 1 mm and
the lobes ca. 2 mm.

32. Hedyotis koana R. J. Wang, Acta Phytotax. Sin. 45: 696.

2007.
yun zhang er cao
Herbs, erect, to 40 cm tall; stems 4-angled to terete, glabrous. Leaves sessile; blade drying subleathery, linear, 1.44
0.10.15 cm, glabrous, base cuneate, margins revolute, apex
acute; secondary veins not visible; stipules fused to petiole
bases, triangular, 11.8 mm, glabrescent, with 13 bristles 0.5
1 mm. Inflorescences terminal and sometimes in uppermost leaf
axils, cymose, 212-flowered, glabrous, pedunculate; axes ascending; bracts narrowly lanceolate, 15 mm, acute to aristate;
pedicels 320 mm. Flowers pedicellate, biology unknown. Calyx glabrous; hypanthium ovoid to subglobose, 11.5 mm; limb
deeply lobed; lobes triangular to lanceolate, 11.7 mm, acute to
acuminate. Corolla white, tubular, glabrous outside; tube 1.5
2.5 mm, glabrous inside; lobes elliptic-oblong to lanceolate,
0.91.2 mm. Fruit capsular, subglobose to ovoid, 1.22 23
mm, glabrous, stiffly papery, loculicidally dehiscent from top;
seeds several, angled. Fl. and fr. JunSep.
Weedy open areas; sea level to 200 m. Fujian, Guangdong,
Guangxi, Hainan, Hunan, Jiangxi.

33. Hedyotis lianshanensis W. C. Ko, J. S. China Agric. Univ.

16(4): 41. 1995.

31. Hedyotis hermanniana R. M. Dutta, J. Econ. Taxon. Bot.

23: 734. 1999.

lian shan er cao

he er man er cao

Subshrubs, erect, to 80 cm tall; stems hollow, 4-angled to

terete, smooth or sulcate, glabrous or usually sparsely hispidulous to puberulent just below stipules when young. Leaves subsessile to petiolate; petiole 35 mm, glabrous; blade drying papery, narrowly elliptic-oblong, narrowly elliptic, lanceolate, or
ovate, 612 1.54 cm, glabrous or rarely scaberulous adax-

Herbs, annual or perennial, erect to procumbent, to

15(20) cm tall; stems subterete to angled or sulcate, glabrous.
Leaves subsessile to petiolate; petiole to 2 mm; blade drying
papery, ovate, lanceolate, or elliptic, 12.5(5) 0.51.5 cm,

RUBIACEAE

164

ially along midrib, base cuneate to acute and usually decurrent,

apex acute to long acuminate; secondary veins 4 or 5 pairs;
stipules fused to petiole bases, obovate, elliptic, or triangular, 3
6 mm, glabrous or puberulent when young, lacerate or with 7 to
numerous narrow lobes or bristles 0.57 mm, these often glandular at apex. Inflorescence axillary, glomerulate, subglobose,
515 mm in diam., glabrous, sessile; bracts narrowly triangular
to linear, 12 mm. Flowers sessile to subsessile, biology unknown. Calyx glabrous to puberulent; hypanthium portion cupulate to subglobose, 11.2 mm; limb lobed essentially to base;
lobes lanceolate, narrowly triangular, narrowly ligulate, or linear, 1.53 mm, ciliolate. Corolla funnelform, outside glabrous;
tube 1.82 mm, glabrous inside; lobes lanceolate to triangular,
12.5 mm. Anthers partially exserted. Stigmas exserted. Fruit
indehiscent, ellipsoid to subglobose, 1.22 mm, glabrescent,
with pedicels to 2 mm; seeds numerous, angled. Fl. FebJul, fr.
JulDec.
Sparse forests in valleys; 200700 m. Guangdong (Liangshan).
The protologue described the flowers as shortly pedicellate, but on
the specimens studied they are sessile at anthesis with the pedicels or
stipes elongating as the fruit develop. W. C. Ko (in FRPS 71(1): 41.
1999) described the fruit as sometimes dehiscent across the top, but the
protologue described them as indehiscent, which agrees with the specimens studied.

34. Hedyotis lineata Roxburgh, Fl. Ind. 1: 369. 1820.

dong ya er cao
Exallage ulmifolia (Wallich) Bremekamp; Hedyotis ulmifolia Wallich; Oldenlandia lineata (Roxburgh) Kuntze.
Herbs or subshrubs, annual or perennial, clambering or
perhaps erect, to 35 cm tall; stems 4-angled, flattened, or subterete, sometimes sulcate, densely villous, hirsute, or pilose
with trichomes drying yellowed. Leaves sessile or petiolate;
petiole to 3 mm, hirsute or villous; blade drying papery, ovatelanceolate, elliptic-lanceolate, or elliptic, (0.5)1.53.5(4)
1.24.5 cm, adaxially sparsely to moderately strigillose to
strigose on blade and densely pilosulous on principal veins,
abaxially glabrescent on blade and moderately to densely strigose to pilose on principal veins, base obtuse to rounded, apex
acute to shortly acuminate; secondary veins 3 or 4 pairs; stipules interpetiolar or fused to petiole bases, broadly triangular to
subtruncate, 24 mm, densely villous to pilose, with 3 or 5
linear lobes or bristles 27 mm. Inflorescences axillary, congested-cymose to fasciculate or subglomerulate, subglobose, 5
15 mm in diam., several flowered, subsessile to pedunculate;
peduncles several in each axil, 0.54 mm; bracts linear, 12
mm; pedicels to 2 mm. Flowers subsessile to pedicellate, biology unknown. Calyx hirtellous; hypanthium portion subcupulate to obconic, ca. 0.8 mm; limb deeply (but not completely)
lobed; lobes triangular, 11.2 mm. Corolla white, salverform to
rotate, outside glabrous except lobes pilose at tip; tube 11.5
mm, pubescent in throat; lobes triangular, 11.5 mm. Anthers
0.50.8 mm, exserted. Stigma 0.30.5 mm. Fruit indehiscent,
subglobose to turbinate, 11.2 1.52 mm, hirtellous; seeds
3640, 0.81 mm. Fl. Jun, perhaps Nov, fr. Nov.
Broad-leaved forests at high elevations; [1001100 m in India].
Yunnan (Menghai) [Bangladesh, India, Myanmar, Nepal].

The names Hedyotis ulmifolia and H. lineata appear to have been

published simultaneously; the basis for choosing the name H. lineata is
not explained in the references consulted but seems to be consistent usage (e.g., Fl. Bhutan 2(2): 761. 1999; Dutta & Deb, Taxon. Rev. Hedyotis. 2004).
We have seen no corollas, anthers, or stigmas from China; the
description of these parts here is from Dutta and Deb (loc. cit.). W. C.
Ko (in FRPS 71(1): 34. 1999) described the corolla tube as 3 as long
as the lobes, which does not agree with the information given by any
other authors for this species. Also the elevational range of at high
elevations given for this species by Ko contrasts markedly with information on the specimens seen and the low to middle elevations noted
by other authors.

35. Hedyotis loganioides Bentham, Fl. Hongk. 149. 1861.

yue gang er cao
Oldenlandia loganioides (Bentham) Kuntze.
Herbs, erect, perennial, to 30 cm tall; stems terete, weakly
angled, or weakly flattened, glabrous. Leaves subsessile to
petiolate; petiole 0.54 mm, glabrous to puberulent; blade
drying papery, elliptic-oblong, lanceolate, or elliptic, 1.56 1
3.5 cm, glabrous, base acute to cuneate, apex acute to shortly
acuminate; secondary veins 35 pairs but hardly visible; stipules apparently interpetiolar, ovate-triangular to lanceolate, 2
2.5 mm, puberulent to glabrous, entire to shortly ciliolate, obtuse to rounded. Inflorescences axillary, cymose, 0.82 cm,
sometimes trichotomous and compact, 1020-flowered, glabrous; peduncles 210(20) mm; bracts triangular to narrowly
triangular, 0.53.5 mm; pedicels 12 mm. Flowers pedicellate,
floral biology unknown. Calyx glabrous; hypanthium portion
subglobose to flattened-ovoid, ca. 1 mm; limb lobed nearly to
base; lobes linear to narrowly triangular, 1.52.2 mm. Corolla
outside glabrous. Fruit capsular, ellipsoid to subglobose, ca. 2
22.3 mm, glabrous, cartilaginous to stiffly papery, smooth,
apparently septicidal then loculicidal, beak broadly triangular to
conical, up to 0.5 mm; seeds few. Fr. Dec.
Guangdong.

36. Hedyotis longiexserta Merrill & F. P. Metcalf, J. Arnold

Arbor. 23: 229. 1942.
shang si er cao
Herbs or subshrubs, erect, to 50 cm tall; stems terete, glabrous, smooth. Leaves petiolate; petiole 35 mm, glabrous;
blade lanceolate or oblong-lanceolate, 57 11.5 cm, base cuneate or obtuse, apex acuminate or acute; secondary veins ca. 3
pairs but very indistinct; stipules fused to petiole bases, broadly
ovate, ca. 3 mm, abruptly narrowed apically to acute, marginally weakly glandular-serrulate. Inflorescences terminal, compound-cymose, 712 cm, several to many flowered; secondary
axes few, racemiform, 34 cm, higher order axes shorter, becoming scorpioid; bracts linear-lanceolate or linear, 0.81.2 cm.
Flowers pedicellate and/or sessile. Calyx glabrous; hypanthium
portion ovoid, 11.2 mm; lobes triangular-ovate, ca. 0.5 mm.
Corolla white, outside glabrous; tube ca. 1.5 mm, inside densely
bearded; lobes broadly ovate, rounded. Fruit capsular, oblongellipsoid, 22.5 ca. 1.5 mm, septicidally then loculicidally
dehiscent; seeds several, black. Fl. JunJul.

RUBIACEAE

Dry open fields, mountain slopes. Guangxi (Shangsi).

37. Hedyotis longipetala Merrill, J. Arnold Arbor. 8: 18. 1927.

chang ban er cao
Oldenlandia longipetala (Merrill) Chun.
Subshrubs, erect, to 40 cm tall; stems glabrous, terete to 4angled with angles obtuse to acute or ridged. Leaves sessile to
petiolate; petiole to 8 mm, glabrous; blade drying stiffly leathery, narrowly lanceolate, narrowly elliptic, or linear-lanceolate,
38 0.41.2 cm, glabrous and shiny, base cuneate to usually
acute and decurrent, margins revolute at least when dry, apex
long acuminate; secondary veins indistinct; stipules interpetiolar, triangular, ovate, or oblong-ovate, 35 mm, glabrous, shiny
and hardened, acute to acuminate. Inflorescence terminal and
sometimes in uppermost leaf axils, congested-cymose to subcapitate, flattened-globose, 1.52 cm in diam., glabrous, several
to many flowered, sessile, subtended by 2 pairs of leaves or
leaflike bracts with inner (i.e., distal) pair ca. 1/2 as large as
outer (i.e., proximal), or axillary cymes few flowered and without leaflike bracts; floral bracts elliptic to triangular, 26 mm;
pedicels to 2 mm. Flowers sessile to shortly pedicellate. Calyx
glabrous, shiny; hypanthium portion obconic, ca. 1 mm; limb
lobed for ca. 1/2; lobes triangular, lanceolate, or spatulate, 24
mm. Corolla white, tubular; tube ca. 3.5 mm, pilosulous in
throat; lobes lanceolate, ca. 11 mm. Anthers ca. 2 mm. Fruit
capsular, ovoid or ellipsoid, 34 mm in diam., glabrous, cartilaginous to hard, septicidal then tardily loculicidal; seeds several. Fl. AprJun, fr. JulSep.
Broad-leaved forests on mountaintops, grasslands at roadsides;
ca. 800 m. Fujian, Guangdong.
The dry plants have a yellowish-brownish cast and a distinctive
shiny, hardened surface on all parts.

38. Hedyotis matthewii Dunn, J. Bot. 47: 376. 1909.

shu hua er cao
Hedyotis acuminatissima Merrill; Oldenlandia matthewii
(Dunn) Chun.
Herbs or subshrubs, erect, perennial, branched, to 75 cm
tall; stems terete to 4-angled, glabrous. Leaves subsessile to
petiolate; petiole to 8 mm, glabrous; blade drying papery to
subleathery, narrowly elliptic, elliptic, or lanceolate, 3.512
0.83 cm, glabrous, base acute to obtuse, apex acute to long
acuminate; secondary veins ca. 3 pairs but indistinct; stipules
fused to petiole bases, broadly triangular to ovate-triangular, 2
4.5 mm, glabrous, sometimes costate, marginally entire to glandular-serrulate, acute to acuminate and sometimes thickened at
apex. Inflorescences terminal and usually also in axils of uppermost leaves, cymose, paniculate, or racemiform, 612 1.57
cm, glabrous, several to many flowered, sessile and apparently
tripartite or pedunculate; peduncles 14 cm; axes mostly dichotomous; bracts linear-lanceolate, 17 mm; pedicels 0.54
mm. Flowers subsessile to pedicellate, distylous. Calyx glabrous; hypanthium turbinate or obovoid, 11.5 mm; limb 1.5
2.2 mm, lobed for 1/2 to deeply lobed; lobes lanceolate to
triangular. Corolla white or purplish, funnelform, outside gla-

165

brous; tube 3.54 mm, inside pubescent in throat; lobes triangular, 23 mm, inside puberulent. Anthers included or exserted,
0.81 mm. Stigmas included or exserted, 0.51 mm. Fruit capsular, ellipsoid, 22.5 mm, crustaceous to cartilaginous, smooth,
glabrous, septicidally then loculicidally dehiscent; seeds numerous, black, angled. Fl. MarNov, fr. MayJul.
Dense forests or thickets on mountains; 100300 m. Guangdong.
The circumscription here of this species and in particular its
separation from Hedyotis mellii differs somewhat from that of some
previous authors but provides a more consistent separation of it from
H. mellii and H. cantoniensis.

39. Hedyotis mellii Tutcher, Rep. Bot. Dept. Hong Kong 1914:
32. 1915.
cu mao er cao
Hedyotis speciosa Handel-Mazzetti; H. wulsinii Merrill;
Oldenlandia mellii (Tutcher) Chun.
Herbs, erect, perennial, to 90 cm tall; stems subterete to 4angled, densely to sparsely hirtellous or pilosulous to sometimes glabrescent. Leaves sessile; blade drying papery, ovatelanceolate, elliptic, lanceolate, or narrowly elliptic, 3.59.5
0.53.5 cm, adaxially sparsely to densely puberulent or hispidulous or often glabrous on lamina, abaxially sparsely to densely
hirtellous, pilosulous, or hispidulous to glabrous; secondary
veins 3 or 4 pairs; stipules fused to petiole bases, triangular to
broadly triangular, 13 mm, densely puberulent, hirtellous, or
hispidulous to glabrescent, marginally entire to glandular-serrate, acuminate or with 35 narrowly triangular to linear lobes
or bristles 0.22 mm. Inflorescences terminal and usually also
in axils of uppermost leaves, cymose to compound-cymose,
paniculate to often racemiform, 325 cm, several to many flowered, densely to sparsely pilosulous, puberulent, or hispidulous
to glabrous; peduncle 0.57 cm; bracts narrowly elliptic to narrowly lanceolate, 15 mm; pedicels 15 mm. Flowers pedicellate, apparently monomorphic. Calyx glabrous or sparsely to
densely hirtellous to pilosulous; hypanthium portion cupulate to
obconic or elliptic, ca. 1 mm; limb 11.5 mm, lobed for 2/3
4/5; lobes ovate-lanceolate to narrowly triangular. Corolla presumably white, funnelform, outside glabrous to densely hirtellous or pilosulous, inside densely villosulous or tomentulose in
throat and onto lobes; tube 22.5 mm; lobes lanceolate to spatulate, 44.5 mm, acute. Anthers shortly exserted, ca. 1.2 mm.
Stigma long exserted, ca. 0.2 mm. Fruit capsular, ellipsoid to
subglobose, 23 mm, sparsely to densely hirtellous, pilosulous,
or glabrous, crustaceous to thickly papery or cartilaginous, septicidally then loculicidally dehiscent; seeds several, black, angled. Fl. JunNov, fr. AugNov.
Jungles or thickets on mountains or mountain slopes; 4001100
m. Fujian, Guangdong, Guangxi, Hunan, Jiangxi.
This species is very similar to Hedyotis cantoniensis, H. matthewii, and H. tenuipes, and these are here separated somewhat differently from the circumscriptions of FRPS (71(1): 48, 51, 52, 54. 1999).
Hedyotis matthewii may be only a glabrous form of H. mellii, although
they do appear to be distinct.

166

RUBIACEAE

40. Hedyotis merguensis Bentham & J. D. Hooker, Gen. Pl. 2:

57. 1873.
he ye er cao
Hedyotis connata J. D. Hooker, nom. illeg. superfl.; H.
coronaria (Kurz) Craib; H. coronata Wallich ex J. D. Hooker &
B. D. Jackson, nom. illeg. superfl.; Oldenlandia connata K.
Schumann, nom. illeg. superfl.; O. coronata F. N. Williams,
nom. illeg. superfl.; Scleromitrion coronarium Kurz.
Herbs, perennial or perhaps annual, procumbent; stems to
30 cm, weakly 4-angled to terete, sometimes sulcate, densely
pilosulous or hispidulous just below nodes grading to glabrous
at bases of internodes. Leaves sessile or subsessile; blade drying
membranous or thinly papery, oblong-elliptic or oblong-lanceolate, 2.510 13 cm, glabrous except sometimes puberulent
along midrib abaxially and/or scaberulous along margins, base
obtuse to rounded, apex obtuse to acute; secondary veins 35
pairs; stipules fused to petiole bases, triangular to rounded, 3
4.5 mm, puberulent to glabrescent, ciliolate, with 57 linear
lobes or bristles 212 mm. Inflorescences terminal and/or pseudoaxillary on short lateral stems, capitate, 11.5 cm in diam.,
several flowered, sessile, enclosed by stipules and leaf bases,
subtended by 2 or usually 4 leaves; bracts not seen. Flowers
sessile or subsessile, apparently distylous. Calyx glabrous;
hypanthium portion ellipsoid, ca. 2 mm; limb deeply lobed;
lobes lanceolate, oblanceolate, or spatulate, 45 mm, distinctly
veined, ciliolate. Corolla white tinged with pink or lilac, funnelform, outside glabrous; tube 912 mm, pubescent in throat;
lobes lanceolate to triangular, ca. 3 mm. Anthers 11.5 mm, exserted or included. Stigma 11.5 mm, exserted or included.
Fruit indehiscent, ellipsoid to obovoid, 2.23 mm, papery, glabrous, with calyx lobes to 5.2 mm; seeds numerous, black,
angled or ridged. Fl. [AugOct in Thailand].
Bamboo forests, on rocks at streamsides; 6001000 m. Hainan,
Yunnan (Mengla) [India, Malaysia, Myanmar, Philippines, Thailand,
Vietnam].
The widely used name Hedyotis coronaria was synonymized with
H. merguensis by Bakhuizen f. (Fl. Java 2: 286. 1965); this synonymy
was accepted by Dutta and Deb (Taxon. Rev. Hedyotis. 2004) but overlooked by some other authors.

41. Hedyotis minutopuberula Merrill & F. P. Metcalf, J.

Arnold Arbor. 23: 229. 1942.
fen mao er cao
Herbs or subshrubs, to 50 cm tall; stems terete, puberulent.
Leaves petiolate; petiole 48 mm, puberulent; blade drying submembranous, ovate or elliptic-oblong, 4.55 0.72.5 cm,
adaxially glabrous, abaxially glabrous or farinose pubescent,
base cuneate or decurrent, apex acute or weakly acuminate;
secondary veins ca. 3 pairs but indistinct; stipules ovate or
lanceolate, 25 mm, marginally glandular-serrulate, long acuminate. Inflorescences terminal, cymose to paniculate, 1.54
24.5 cm; axes dichotomous to subumbelliform; bracts subulate, ca. 2 mm; pedicels ca. 1 mm. Flowers pedicellate. Calyx
puberulent; hypanthium portion obovoid, ca. 1 mm; lobes
lanceolate to narrowly triangular, 1.52.5 mm. Corolla purple,
tubular-salverform, outside glabrous; tube ca. 6 mm, bearded in

throat; lobes obovate to triangular, 11.2 mm. Anthers included.

Fruit capsular, subglobose or obovate, ca. 2 1.5 mm, puberulent or subglabrous, dehiscent; seeds 24, black, angled. Fl.
AprJun.
Jungles or thickets at low elevations or on slopes of low hills.
Hainan.

42. Hedyotis obliquinervis Merrill, Lingnan Sci. J. 14: 56.

1935.
pian mai er cao
Hedyotis hedyotidea (Candolle) Merrill var. obliquinervis
(Merrill) Fukuoka.
Shrubs or subshrubs, climbing to scandent, to 40 cm;
stems terete to 4-angled, sometimes sulcate, densely villous and
usually also hirtellous or villosulous. Leaves petiolate; petiole
15 mm, densely hirtellous or villosulous; blade drying papery,
lanceolate, oblong-lanceolate, or lanceolate-elliptic, 38 0.8
2.5 cm, adaxially sparsely to moderately hispidulous and sometimes villosulous or villous on lamina and densely hispidulous
on principal veins, abaxially moderately to densely hispidulous,
villosulous, or hirtellous with pubescence denser on principal veins, base cuneate to rounded, apex acute to acuminate;
secondary veins 35 pairs; stipules fused to petiole bases,
densely villous to villosulous, rounded, 14 mm, hispidulous,
with 37 narrowly triangular to linear lobes or bristles 27 mm.
Inflorescences terminal and sometimes also in axils of uppermost leaves, cymose to corymbiform-rounded, 2.59 416
cm, densely villous, villosulous, or hirtellous or sometimes glabrescent, pedunculate or sessile and tripartite; peduncles 1.55
cm; bracts linear-lanceolate to lanceolate, 0.54 mm; pedicels
1.53 mm. Flowers pedicellate, distylous. Calyx moderately to
densely hirtellous or villosulous; hypanthium portion obconical,
11.5 mm; limb lobed nearly to base; lobes lanceolate to spatulate, 22.5 mm, sometimes reflexed. Corolla white or greenish
white, funnelform, outside subglabrous or hispidulous at least
on upper part; tube 11.5 mm, bearded in throat; lobes lanceolate to spatulate, 22.5 mm. Anthers exserted or included, 0.81
mm. Stigmas included or exserted, 0.20.5 mm. Fruit capsular,
compressed globose, ca. 2 mm in diam., hirtellous or villosulous, cartilaginous, septicidally then loculicidally dehiscent,
beak rounded, ca. 0.6 mm high; seeds several, blackish brown,
angled. Fl. DecAug, young fr. JunJul.
Broad-leaved forests, thickets on hills; 100400 m. Hainan [S
Vietnam].
This species is very similar to Hedyotis ampliflora and may be
only a pubescent form of that species.

43. Hedyotis ovata Thunberg ex Maximowicz, Bull. Acad.

Imp. Sci. Saint-Ptersbourg 29: 161. 1883.
luan ye er cao
Oldenlandia ovata (Thunberg ex Maximowicz) Kuntze.
Herbs or subshrubs, erect, perennial, to 90 cm tall; stems
4-angled, glabrous. Leaves petiolate; petiole ca. 5 mm, glabrous; blade drying papery, ovate, 2.57 1.53.5 cm, glabrous, base acute, margins revolute, apex acute; secondary

RUBIACEAE

veins 3 or 4 pairs but indistinct; stipules triangular to broadly

triangular, ca. 2 mm, marginally glandular-serrulate. Inflorescences terminal and sometimes also in axils of uppermost
leaves, fascicled, several flowered, sessile or pedunculate; peduncles 0.92 cm; bracts triangular, ca. 1.5 mm; pedicels 0.51
mm. Flowers subsessile to pedicellate. Calyx with hypanthium
portion subglobose, 11.5 mm in diam.; lobes lanceolate, ca. 4
mm. Corolla purplish red, villosulous in throat and onto lobes;
tube 34.5 mm. Anthers included. Stigmas included. Fruit capsular, subglobose, 34 mm in diam., septicidally then loculicidally dehiscent; seeds 3 or 4, 4-angled, smooth. Fl. and fr.
AprJul.
On humid soil rich in humus in forests. Hainan (Sanya).
This species has been reported from Vietnam by the Kew Rubiaceae checklist (Govaerts et al., World Checkl. Rubiaceae; http://www.
kew.org/wcsp/rubiaceae/; accessed on May 2007); it was not included in
the Vietnam checklist.

44. Hedyotis ovatifolia Cavanilles, Icon. 6: 52. 1801.

ai xiao er cao
Gonotheca ovatifolia (Cavanilles) Santapau & Wagh;
Oldenlandia nudicaulis Roth; O. ovatifolia (Cavanilles) Candolle; Thecagonum ovatifolium (Cavanilles) Babu.
Herbs, erect, annual, to 15 cm tall; stems terete to 4-angled,
often sulcate, densely villosulous, pilosulous, or puberulent
with pubescence denser, longer, and apparently glandular on
upper (i.e., distal) parts of internodes. Leaves subsessile to
usually petiolate, distributed along stem or those at uppermost 2
nodes clustered or apparently 4-verticillate especially at base of
inflorescence; petiole 15 mm, densely puberulent to villosulous; blade drying membranous, elliptic or ovate, 16 0.73
cm, adaxially glabrous, abaxially puberulent or villosulous to
glabrescent, base cuneate to rounded, apex acute to rounded;
secondary veins 36 pairs; stipules fused to petiole bases,
truncate to broadly rounded, 12.5 mm, pilosulous to glabrescent, erose to glandular-lacerate and usually also with 13 linear lobes to 1 mm. Inflorescence terminal, cymose to paniculate,
310 cm, glabrous, several to many flowered, pedunculate;
peduncles 13, 1.54 cm, slender; axes mostly dichotomous;
bracts triangular, 0.10.5 mm; pedicels 112 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion subglobose, ca. 0.5
mm; limb lobed essentially to base; lobes triangular, 0.20.3
mm. Corolla white, tubular or salverform, outside glabrous;
tube 11.5 mm, barbate in throat; lobes lanceolate, ca. 1 mm.
Anthers exserted, ca. 0.2 mm. Stigma ca. 0.6 mm, exserted.
Fruit capsular, compressed globose, 23 23 mm, glabrous,
loculicidally dehiscent across top through beak, beak broadly
rounded, to 0.3 mm high; seeds numerous, smooth. Fl. and fr.
JulAug.
Broad-leaved forests or grasslands on mountain slopes, sometimes
on limestone rocks. Guizhou, Hainan, Taiwan, Yunnan [India, Malaysia,
Nepal, Pakistan, Philippines, Sri Lanka, Thailand].
The measurements of several structures, in particular calyx, corollas, and anthers, given by Dutta and Deb (Taxon. Rev. Hedyotis.
2004) and also W. C. Ko (in FRPS 71(1): 7172. 1999) do not agree
with the specimens studied, nor with Dutta and Debs own figure. The
measurements in the description here are taken from specimens, which

167

agree with Dutta and Debs figure. All authors have found both the
anthers and stigmas exserted (cf. Dutta & Deb figure; Cooray
69011901R, MO!) except Fl. Bhutan (2(2): 762763. 1999), who found
the anthers included.

45. Hedyotis paridifolia Dunn, Bull. Misc. Inform. Kew 1912:

366. 1912.
yan ling er cao
Oldenlandia paridifolia (Dunn) Chun.
Herbs or subshrubs, perennial, erect, to 30 cm tall; stems
green or purplish, flattened to subterete, usually sulcate along
each side, glabrous. Leaves subsessile to shortly petiolate, often
clustered and apparently verticillate at tops of stems; petiole to
3 mm, glabrous; blade drying papery, ovate, elliptic, or ellipticovate, 79(12) 2.54.5 cm, glabrous, base obtuse to rounded
and often decurrent, apex acute or shortly acuminate; secondary
veins 57 pairs; stipules fused to petiole bases, triangular to
broadly triangular, 13.2 mm, glabrous, sparsely glandular-serrulate or glandular-setulose, apex with aristate projection 12
mm. Inflorescences terminal, capitate or subcapitate, hemispherical, 1.52.5 cm in diam., glabrous, subsessile; bracts triangular, 12 mm; pedicels to 2.5 mm. Flowers shortly pedicellate to sessile, perhaps monomorphic. Calyx glabrous; hypanthium portion purple, turbinate, ca. 1 mm, puberulent; limb
lobed nearly to base; lobes on a few flowers rarely 2 or 3,
elliptic, elliptic-oblong, or lanceolate (i.e., narrowed at base),
24 mm, obtuse to acute. Corolla white, funnelform to tubular,
outside glabrous; tube 45 mm, inside pilosulous in upper part
and throat; lobes triangular, 11.2 mm. Anthers exserted, ca. 1
mm. Stigma exserted, ca. 1.2 mm. Fruit indehiscent, straw-yellow, obovoid or subellipsoid, 33.5 22.5 mm, glabrescent;
seeds numerous, black, angled, foveolate. Fl. MayNov, fr.
AprMay.
Forests or thickets at middle elevations; ca. 200 m. Hainan.
W. C. Ko (in FRPS 71(1): 35. 1999) described the leaf blade as up
to 12 cm, but this has not been seen on specimens studied and may
include plants that have been subsequently separated as Hedyotis baotingensis.

46. Hedyotis pinifolia Wallich ex G. Don, Gen. Hist. 3: 526.

1834.
song ye er cao
Oldenlandia pinifolia (Wallich ex G. Don) K. Schumann.
Herbs, annual or perennial, diffusely branched, to 25 cm
tall; stems wiry, sharply 4-angled to subterete, sometimes sulcate, glabrous. Leaves sessile or subsessile, sometimes borne on
very short axillary stems and appearing verticillate or clustered;
blade drying stiffly papery to leathery, linear to narrowly spatulate, 525 0.82 mm, adaxially glabrous to densely scaberulous, abaxially glabrous, base straight to acute, margins markedly revolute at least when dry, apex acute; secondary veins not
visible; stipules shortly fused to petiole bases, triangular to
rounded, 0.81.2 mm, puberulent to glabrous, with 15 linear
lobes or bristles 0.52.5 mm. Inflorescences terminal and pseudoaxillary on short axillary stems, capitate to shortly fasciculate, 47 mm in diam., (1 or)310-flowered, sessile and sub-

RUBIACEAE

168

tended by 1 or 2 pairs of somewhat reduced leaves; bracts lanceolate to setose, 0.54 mm, entire to ciliate; pedicels to 1 mm.
Flowers sessile, subsessile, or pedicellate, apparently homostylous. Calyx with hypanthium portion obconical to subglobose, 0.81.2 mm, glabrous to densely hispidulous; limb lobed
for more than 1/2; lobes subulate to narrowly triangular, 12
mm, glabrescent, entire to densely ciliolate. Corolla white
sometimes flushed with pink, tubular to funnelform, outside
glabrous; tube 34.2 mm, pubescent in throat; lobes spatulateoblong to elliptic, 1.82 mm. Anthers exserted, ca. 1 mm. Stigma 0.51 mm, exserted and positioned above anthers. Fruit capsular, ovoid to lanceoloid, 2.53 1.52 mm, cartilaginous to
stiff, loculicidal across top; seeds several to numerous, pale
brown, angled. Fl. MayNov, fr. AprNov.
Open fields on hills, sandy wastelands at seasides or riversides;
sea level to below 100 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan, Yunnan [India, Malaysia, Myanmar, Nepal, Thailand, Vietnam].
The plant illustrated in FRPS (71(1): 70, t. 13, f. 16. 1999) as
Hedyotis umbellata is actually H. pinifolia. The Fl. Bhutan (2(2):
763764. 1999) included this species noting that the report is based on a
published historical record that has not been re-confirmed.

47. Hedyotis platystipula Merrill, Philipp. J. Sci. 21: 510.

1922.
kuo tuo ye er cao
Oldenlandia platystipula (Merrill) Chun.
Herbs or subshrubs, erect to perhaps weak, to 40 cm tall;
stems flattened to 4-angled, sometimes sulcate, glabrous. Leaves
petiolate; petiole 318 mm, glabrous; blade drying membranous, elliptic, oblong-ovate, or oblong-lanceolate, 512 2.54
cm, adaxially glabrous or rarely puberulent along principal
veins, abaxially glabrous or papillose on principal veins, base
cuneate to rounded and usually shortly decurrent, apex acute to
shortly acuminate; secondary veins 46 pairs; stipules interpetiolar, reniform, ovate, or suborbicular, 510 mm, glabrous,
broadly rounded to broadly triangular, fringed-lacerate or with
numerous lobes or bristles 0.32 mm, usually glandular at apex.
Inflorescence axillary, glomerate, subglobose, 12.5 cm in
diam., glabrous, sessile; bracts not seen. Flowers sessile to subsessile, biology unknown. Calyx glabrous; hypanthium portion
turbinate to ellipsoid, 1.22 mm; limb lobed essentially to base;
lobes narrowly lanceolate to triangular, 46 mm, ciliate. Corolla
white, tubular, outside glabrous; tube 67 mm, inside apparently glabrous; lobes lanceolate to triangular, 1.52 mm. Anthers ca. 2 mm, partially exserted. Stigmas ca. 1.5 mm, included. Capsules oblong-ellipsoid to obconic, 23 1.52 mm,
somewhat flattened laterally, septicidally then loculicidally dehiscent; seeds ca. 10, black, scabrous. Fl. MayAug, fr. Sep.
Forests in valleys, on rocks at streamsides. Guangdong, Guangxi.

48. Hedyotis prostrata Blume, Catalogus, 40. 1823.

fei l bin er cao
Hedyotis congesta R. Brown ex G. Don; H. laevigata
(Candolle) Miquel; H. philippensis (Willdenow ex Sprengel)
Merrill ex C. B. Robinson; Metabolos laevigatus Candolle;

Oldenlandia congesta (R. Brown ex G. Don) Kuntze (1891),

not Baker (1877); O. prostrata (Blume) Kuntze; Spermacoce
philippensis Willdenow ex Sprengel.
Herbs or subshrubs, erect to perhaps clambering, to 60 cm
tall; stems 4-angled, flattened, or terete, sometimes sulcate, glabrous. Leaves petiolate; petiole 36 mm, glabrous; blade drying
papery or membranous, ovate, lanceolate, ovate-lanceolate, or
elliptic, 3.212 1.24.5 cm, glabrous, base cuneate, obtuse, or
rounded then often decurrent, apex acute or acuminate; secondary veins 46 pairs but usually not visible; stipules shortly fused
to petiole bases or around stem, broadly triangular, 24 mm,
glabrous or pilosulous to hirtellous, acute or usually acuminate
to aristate, entire or glandular-serrulate. Inflorescence axillary
and/or terminal, capitate or glomerulate, 510 mm in diam., 10
18-flowered, glabrous, sessile or subsessile; bracts lanceolate,
triangular, or linear, 0.54 mm. Flowers sessile to subsessile,
floral biology unknown. Calyx glabrous; hypanthium portion
obconic to campanulate, 0.81 mm; limb lobed essentially to
base; lobes triangular, 11.2 mm. Corolla white to violet, funnelform; tube 23 mm; lobes lanceolate, 11.2 mm. Fruit indehiscent or tardily septicidal, ellipsoid-oblong to subglobose, 24
1.53 mm, with pedicels to 2 mm; seeds 1012, black, foveolate. Fl. AprJun.
Thickets; 200400 m. Hainan (Xinglong) [India, Indonesia,
Philippines, Vietnam].
Plants from the Philippines have fruit that are markedly ellipsoid,
but the Chinese plants treated under this name have subglobose fruit
that are quite distinctively different in shape.
This species has long been treated under the name Hedyotis
philippensis, based on the conclusion that Blumes valid publication of
the name H. prostrata was in 1826 while the basionym of H. philippensis was published in 1825. However, Bakhuizen f. (Fl. Java 2: 284
288. 1965) noted that Blumes name was actually published in an earlier
work and, thus, is the oldest name for this species. Dutta and Deb (Taxon. Rev. Hedyotis. 2004) treated this species under the name H. congesta, without explanation.

49. Hedyotis pterita Blume, Bijdr. 972. 18261827.

chi guo er cao
Gonotheca blumei Candolle; Hedyotis alata Koenig ex
Wight & Arnott; Oldenlandia alata (Koenig ex Wight &
Arnott) J. D. Hooker; O. pterita (Blume) Miquel; Thecagonum
pteritum (Blume) Babu.
Herbs, annual or perennial, at least weakly erect, to
15(30) cm tall; stems subterete to 4-angled, sometimes sulcate
and/or winged, glabrous. Leaves subsessile to petiolate; petiole
to 10 mm, glabrous; blade drying membranous, elliptic-oblong,
lanceolate, or elliptic, 16 0.41.5 cm, glabrous, base cuneate
to obtuse then often abruptly decurrent, apex acute or obtuse;
secondary veins indistinct; stipules fused to petiole bases, triangular, 0.52.5 mm, glandular-puberulent to glabrescent, apically truncate or serrulate to prolonged and acute. Inflorescences terminal becoming displaced to pseudoaxillary in upper
leaf axils by subsequent stem growth, 1-flowered or 28-flowered and cymose to fasciculate, glabrous, subsessile or pedunculate; peduncle to 1.5 cm; bracts reduced, triangular, to 0.3
mm; pedicels to 1.5 mm, winged. Flowers subsessile to shortly

RUBIACEAE

pedicellate. Calyx glabrous; hypanthium portion cupular to turbinate, 12 mm, with 4 wings to 0.6 mm wide; limb lobed for
1/22/3 its length; lobes ovate, ca. 1 mm, reticulately veined,
dorsally winged to apex, sometimes ciliolate, acute to acuminate. Corolla white, outside glabrous; tube 11.5 mm, barbate
in throat; lobes triangular, 11.5 mm. Anthers ca. 0.5 mm, included and positioned near base of corolla tube. Fruit capsular,
turbinate to ellipsoid, 58 35 mm, glabrous, with 4 wings to
1 mm wide; seeds numerous, ca. 0.5 mm. Fl. JulOct.
Thickets, slightly shaded wastelands. Guangdong, Guangxi [India,
Malaysia, Philippines, Thailand, Vietnam].

50. Hedyotis pulcherrima Dunn, Bull. Misc. Inform. Kew,

Addit. Ser. 10: 127. 1912.
yan li er cao
Oldenlandia pulcherrima (Dunn) Chun.
Herbs or subshrubs, perennial, erect, to 30 cm tall; stems
terete to slightly flattened, densely farinose puberulent. Leaves
petiolate; petiole 0.150.6 cm, farinose puberulent; blade drying
papery, ovate, lanceolate-elliptic, or lanceolate, 0.85 0.52
cm, both surfaces glabrous or puberulent on midrib, base obtuse
to rounded, margins plane to revolute, apex obtuse to acute;
secondary veins 3 or 4 pairs but mostly indistinct; stipules fused
to petiole bases, broadly triangular to broadly rounded, 12
mm, densely farinose puberulent, ciliolate-serrulate, apically
sometimes with obtuse to shortly apiculate protuberance. Inflorescences terminal and in axils of uppermost leaves, 1-flowered
or 24-flowered and fasciculate, densely puberulent, pedunculate; peduncles 26 mm, often articulate above middle; bracteoles lanceolate or spatulate, 0.52 mm. Flowers pedunculate.
Calyx glabrous; hypanthium portion turbinate to ellipsoid, ca.
1.2 mm; limb lobed nearly to base; lobes narrowly lanceolate to
narrowly spatulate, 33.5 mm. Corolla pale purple, funnelform,
outside glabrous; tube 1213 mm, pubescent in throat; lobes
spatulate-oblong to ovate, 46 mm. Anthers in apparent shortstyled form included or partially exserted, ca. 2 mm. Stigmas
included, positioned near middle of corolla tube, ca. 2.3 mm.
Fruit capsular, ovoid or obovoid, ca. 2.5 mm, crustaceous to
cartilaginous, outside glabrescent, septicidally then loculicidally
dehiscent; seeds several. Fl. JunJul, fr. Apr.
Open fields at roadsides. Guangdong (Raoping).

51. Hedyotis scandens Roxburgh, Fl. Ind. 1: 369. 1820.

pan jing er cao
Oldenlandia scandens (Roxburgh) Kuntze.
Shrubs or herbs, perennial, lianescent, climbing, or scandent, to several meters; stems compressed terete to angled,
smooth or sulcate, glabrous or puberulent in lines. Leaves subsessile to petiolate; petiole to 5 mm, glabrous; blade drying papery to subleathery, pale abaxially, oblong-lanceolate, elliptic,
narrowly elliptic, or elliptic-oblong, 510 1.54 cm, glabrous,
base acute to cuneate then decurrent, apex long acuminate;
secondary veins 35 pairs but not evident; stipules fused to
petiole bases, truncate to triangular, 2.54 mm, glabrous to usually puberulent or strigillose, ciliolate to entire, apex acute to
acuminate or bilobed, sometimes glandular and/or with bristle

169

inserted below top of stipule. Inflorescences terminal and usually also in axils of uppermost leaves, cymose to compoundcymose, 215 cm, densely pilosulous or hirtellous, pedunculate;
peduncles 23 cm; bracts triangular to narrowly triangular, 0.3
6 mm; pedicels 13 mm. Flowers pedicellate, distylous. Calyx
glabrous; hypanthium portion obconical, ca. 1 mm, sometimes
ridged; limb 12 mm, lobed for 1/31/2, lobes triangular and
thickened, sometimes costate. Corolla white or yellow, tubularfunnelform to funnelform, outside glabrous [or puberulent in
India], inside densely villous in throat and throughout lobes;
tube 12 mm; lobes narrowly spatulate-oblong to narrowly triangular, 34 mm, acute. Anthers shortly to long exserted, ca.
1.2 mm. Stigmas 0.31 mm. Fruit capsular, subglobose, ellipsoid, or ovoid, 35 35 mm, glabrous, cartilaginous to stiffly
papery, loculicidally dehiscent across top then sometimes septicidal, beak rounded to conical, 1.22 mm; seeds several, black,
angled. Fl. JulSep, fr. AugSep.
Sparse forests, on humid soil in valleys; 10001800 m [as low as
ca. 700 m in India]. Yunnan [Bangladesh, Bhutan, India, Myanmar, Nepal, Vietnam].

52. Hedyotis shenzhenensis Tao Chen, Edinburgh J. Bot. 64:

331. 2007.
shen zhen er cao
Herbs, perennial, erect, to 40 cm tall; stems glabrous.
Leaves subsessile; blade drying subleathery, adaxially dark
green, abaxially pale green, elliptic, elliptic-oblong, or obovate,
8.515 59 cm, glabrous, base cuneate to rounded or cordulate, apex acute to obtuse; secondary veins 46 pairs but rather
indistinct; stipules fused to leaf bases or very shortly around
stem, triangular, 35 mm, shortly pubescent, ciliolate, acute.
Inflorescence terminal, paniculate, many flowered, branched to
several orders, glabrous, flushed with purple; peduncle 1018
cm; axes flattened to 4-angled; bracts narrowly triangular, 0.23
mm; pedicels to 2 mm. Flowers sessile and/or pedicellate, distylous. Calyx glabrous; hypanthium portion obconic, ca. 1 mm;
limb deeply lobed; lobes narrowly triangular to ovate, ca. 1.5
mm. Corolla white, tubular-urceolate, glabrous outside; tube ca.
3 mm, villous from middle into throat; lobes ovate to elliptictriangular, 1.82.5 mm. Anthers included or exserted, 0.60.8
mm. Stigma included or exserted, ca. 1 mm. Fruit capsular, subglobose, ca. 2 mm, glabrous, weakly ribbed, septicidally dehiscent; seeds numerous, black, irregularly angled. Fl. AprJun,
fr. MayOct.
Understories of broad-leaved forests; 400700 m. Guangdong
(Shenzhen).
This species was treated by Chen (in T. L. Wu, Check List Hong
Kong Pl. 265266. 2002) under the name Hedyotis yangchunensis. The
flower sizes in the figure in the protologue, according to the scale in that
figure, do not precisely agree with the measurements given in the protologue description.

53. Hedyotis strigulosa (Bartling ex Candolle) Fosberg, Smithsonian Contr. Bot. 45: 28. 1980.
rou ye er cao
Oldenlandia strigulosa Bartling ex Candolle, Prodr. 4:
427. 1830; Hedyotis coreana H. Lveill; H. taiwanensis S. F.

170

RUBIACEAE

Huang & J. Murata [taiwanense]; Thecagonum strigulosum

(Bartling ex Candolle) Terrell & H. Robinson.
Herbs, annual or perennial, prostrate to erect, to 15(20)
cm tall; stems terete or weakly to sharply 4-angled, sometimes
sulcate, glabrous. Leaves sessile; blade very fleshy, drying leathery, oblong-obovate, elliptic-oblong, oblanceolate, or spatulate,
12.5 0.21 cm, glabrous, base acute to decurrent, apex
rounded, obtuse, or acute; secondary veins indistinct; stipules
fused to petiole bases, triangular, 12 mm, glabrous, with 2 central longitudinal lines of glandular trichomes or fleshy ornamentation, acute to acuminate. Inflorescences terminal and sometimes in axils of uppermost leaves, 1-flowered or 212-flowered and shortly cymose, fasciculate, or paniculate, glabrous,
subsessile to pedunculate; peduncles to 1.5 cm; bracts triangular, 0.12 mm; pedicels 112 mm. Flowers pedicellate. Calyx
glabrous; hypanthium portion turbinate to ellipsoid, 11.5 mm;
limb lobed for 1/22/3; lobes triangular to lanceolate, 0.82
mm. Corolla white, tubular, glabrous outside; tube 1.21.5 mm,
barbate in throat; lobes triangular to oblanceolate, 1.22 mm.
Anthers not seen. Stigmas ca. 0.8 mm, exserted. Fruit capsular,
subglobose, hemispherical, or compressed turbinate, 3.55 mm,
glabrous, loculicidal across top, sometimes with rudimentary
beak; seeds numerous, blackish brown, subglobose to angled,
foveolate. Fl. and fr. DecApr.
Sandy or muddy beaches, wastelands, on rocks near sea; sea level
to near sea level. Guangdong, Taiwan, Zhejiang [Japan, Korea; Micronesia].
Hedyotis taiwanensis was treated as a species separate from H.
coreana by W. C. Ko (in FRPS 71(1): 39, 73. 1999), and the name H.
strigulosa was not cited by her. However, H. coreana was synonymized
under H. strigulosa by Fosberg (loc. cit.), though provisionally. Hedyotis taiwanensis was synonymized with H. strigulosa by Fl. Taiwan (ed.
2, 4: 268. 1998). The Fl. Taiwan separated Hedyotis strigulosa from H.
biflora based on the stoutness of the pedicels and fleshiness of the
leaves, but different characters are used here and most of the specimens
included by them in H. strigulosa are here included in H. biflora.
The Fl. Taiwan and Fl. Japan (3a: 217. 1993) included the Chinese
plants in Hedyotis strigulosa var. parvifolia (Hooker & Arnott) T.
Yamazaki (J. Jap. Bot. 58: 284. 1983). This name was not mentioned by
W. C. Ko (loc. cit.) and was synonymized with H. strigulosa by the
Kew Rubiaceae checklist (Govaerts et al., World Checkl. Rubiaceae;
http://www.kew.org/wcsp/rubiaceae/; accessed on May 2007), though
without any source or explanation. The basionym of this name is H.
biflora var. parvifolia Hooker & Arnott (Bot. Beechey Voy. 264. 1838).
Hedyotis biflora var. parvifolia was reported from Zhejiang (Qiu &
Zhong, Fl. Zhejiang 6: 121122. 1986), but that report is here interpreted as H. strigulosa.

scaberulous along grooves and/or near nodes. Leaves sessile or

subsessile; petiole to 1 mm, glabrescent; blade drying thinly
leathery, linear, linear-lanceolate, narrowly elliptic-oblong, or
narrowly spatulate, 1.25 0.150.4 cm, adaxially glabrous or
scaberulous near margins, abaxially glabrous, both surfaces
sometimes apparently scaly due to collapsed large epidermal
cells, base cuneate, acute, or decurrent, margins frequently revolute, apex acute or acuminate; secondary veins not visible;
stipules fused to petiole bases, triangular to rounded, 12 mm,
puberulent, hispidulous, or glabrescent, with 25 linear or setiform lobes 14 mm. Inflorescences axillary, 1-flowered or 2- or
3-flowered in each axil, congested-cymose, glomerulate, or fasciculate, 48 mm in diam., sessile to subsessile; bracts acicular
to lanceolate, 12.5 mm, entire or marginally scaberulous; pedicels to 1 mm. Flowers sessile to subsessile, apparently homostylous. Calyx glabrous; hypanthium portion subglobose to
obovoid, ca. 1 mm; limb lobed nearly to base; lobes linear-lanceolate, triangular, or spatulate, 1.52 mm, ciliolate. Corolla
white, funnelform, outside glabrous; tube ca. 2 mm, pubescent
in throat; lobes narrowly spatulate-oblong, 12 mm. Anthers
exserted, ca. 1 mm. Stigma 0.20.3 mm. Fruit capsular, ovoid,
22.5 1.52 mm, cartilaginous to stiff, loculicidal across top;
seeds numerous. Fl. and fr. AprDec.
Slopes in valleys, ridges of fields; 1001400 m. Fujian, Guangdong, Guangxi, Hainan, Sichuan, Taiwan, Yunnan, Zhejiang [India,
Indonesia, Japan, Malaysia, Philippines, Thailand, Vietnam; Australia,
Melanesia].
This species has been treated by most authors as Hedyotis tenelliflora; however, Dutta and Deb (Taxon. Rev. Hedyotis. 2004) considered that a distinct species and treated these plants as H. angustifolia.
Unfortunately, they presented no discussion of this situation nor details
of the actual identity and range of true H. tenelliflora; consequently,
their conclusion is not yet accepted here.
W. C. Ko (in FRPS 71(1): 41. 1999) described the leaves as
densely rotund transparent scaly, which apparently refers to the relatively large epidermal cells that are visible under magnification and
sometimes apparently collapse. This character was noted in the Fl.
Bhutan (2(2): 757764. 1999) for several Hedyotis species.
The name Scleromitrion sinense is here provisionally placed in
synonymy with Hedyotis tenelliflora. As detailed by Sivarajan and Nair
(Taxon 35: 363369. 1986), the name Spermacoce stricta Linnaeus f.
has also been considered by some authors to apply to H. tenelliflora (in
which case it would have priority), but the identity of that Spermacoce
name is so unclear and controversial, as well as involving plants from
our flora area, that it is better left aside for now. Other authors have
suggested that S. stricta is actually a synonym of H. verticillata.

54. Hedyotis tenelliflora Blume, Bijdr. 971. 18261827.

55. Hedyotis tenuipes Hemsley, J. Linn. Soc., Bot. 23: 375.

1888.

xian hua er cao

xi geng er cao

Hedyotis angustifolia Chamisso & Schlechtendal; Oldenlandia angustifolia (Chamisso & Schlechtendal) Bentham;
O. tenelliflora (Blume) Kuntze; Scleromitrion angustifolium
(Chamisso & Schlechtendal) Bentham; S. sinense Miquel.
Herbs, annual or perennial, diffusely branched, to 40 cm
tall, often drying blackened; stems subterete and smooth or
weakly to sharply 4-angled and/or 2-sulcate, glabrous or often

Oldenlandia tenuipes (Hemsley) Kuntze.

Herbs or subshrubs, erect, perennial, to 50 cm tall; stems
terete to weakly 4-angled, often flexuous, puberulent to glabrous. Leaves subsessile to petiolate; petiole to 2(6) mm; blade
drying papery, usually clear green, narrowly lanceolate, lanceolate, ovate, or lanceolate-elliptic, 2.59 0.73 cm, both surfaces glabrous except densely puberulent on midrib adaxially,

RUBIACEAE

base cuneate to rounded, apex acute to acuminate; secondary

veins 3 or 4 pairs but usually indistinct; stipules fused to petiole
bases, triangular, ovate, or subtruncate, 13 mm, densely puberulent, acute to acuminate, marginally entire, serrulate, or
occasionally glandular-erose. Inflorescences terminal and often
axillary and/or pseudoaxillary in uppermost leaf axils, cymose
to paniculate, 418 cm, glabrous, several to many flowered,
pedunculate; peduncle 0.52 cm; axes slender, flexuous,
spreading at up to 90; bracts linear to narrowly triangular, 15
mm; pedicels 415 mm. Flowers all pedicellate, apparently
monomorphic. Calyx glabrous; hypanthium portion turbinate to
ellipsoid, 11.5 mm; limb lobed for 2/33/4; lobes narrowly
triangular to lanceolate, 11.5 mm. Corolla white, sometimes
tinged pink to purple, funnelform, outside glabrous, inside
densely barbate in throat and throughout lobes; tube 1.53 mm;
lobes lanceolate to spatulate, 34 mm, acute to acuminate. Anthers partially to fully exserted, ca. 1.5 mm. Stigma 0.10.2
mm, exserted by 34 mm. Fruit capsular, ovoid to ellipsoid, 2
2.5 mm, glabrous, smooth, septicidally then loculicidally dehiscent; seeds numerous, black, angled. Fl. JunAug, fr. JunNov.
Sandy lands, barren soil in sparse forests; 2001000 m. Fujian,
Guangdong, Hainan.
This species is similar to Hedyotis matthewii and H. mellii and is
perhaps separated here somewhat differently from other authors (e.g.,
W. C. Ko in FRPS 71(1): 52. 1999). The pedicel measurements here are
for flowers at anthesis and fruit; the pedicels subtending flower buds are
much shorter and apparently elongate rapidly as the buds mature.

56. Hedyotis terminaliflora Merrill & Chun, Sunyatsenia 2:

326. 1935.
ding hua er cao
Herbs or subshrubs, perennial, erect, to 100 cm tall; stems
terete, flattened or weakly 4-angled, 2- or 4-sulcate, glabrescent
or sparsely to densely strigillose to hirtellous throughout or in
lines along grooves. Leaves petiolate; petiole 14 mm, densely
puberulent, strigillose, or glabrescent; blade drying papery or
subleathery, a bit fragile, ovate, oblong-ovate, lanceolate, or oblong-lanceolate, 2.28 13 cm, adaxially glabrous and shiny,
abaxially sparsely to densely puberulent to strigillose, base obtuse to rounded, apex acute to shortly acuminate; secondary
veins 24 pairs; stipules shortly fused to petiole bases, ovate,
triangular, or narrowly triangular, 25 mm, glabrous to densely
puberulent or strigillose, laciniate or glandular-setose. Inflorescence terminal, subcapitate, hemispherical to subglobose, 1.5
1.8 cm in diam., glabrescent, sessile or subsessile; bracts narrowly lanceolate to narrowly triangular, 15 mm; pedicels to 2
mm. Flowers sessile to pedicellate. Calyx glabrescent; hypanthium portion cylindrical, ca. 1 mm; limb deeply lobed; lobes
narrowly spatulate-oblong, narrowly triangular, or narrowly
elliptic, 34 mm, sometimes unequal, usually ciliate. Corolla
funnelform, outside glabrous; tube ca. 3.5 mm, barbate in throat;
lobes triangular, ca. 2.5 mm. Anthers exserted, ca. 1 mm. Stigmas not seen. Fruit capsular, subglobose to ellipsoid, ca. 2.5
mm in diam., glabrescent, cartilaginous, septicidally dehiscent
then quickly loculicidal; seeds several, black, angled. Fl. Feb
Mar, fr. AugNov.
Broad-leaved forests on mountaintops; 6001100 m. Hainan.

171

Broader morphological variation is included in this species here

than in the original species circumscription, based on study of specimens. This is the first description of the corolla; it is based on Liu Qinqu
25932 (MO).

57. Hedyotis tetrangularis (Korthals) Walpers, Ann. Bot. Syst.

2: 769. 1852.
fang jing er cao
Diplophragma tetrangulare Korthals, Ned. Kruidk. Arch.
2: 149. 1851 [tetrangularis]; Hedyotis parryi Hance; H.
quadrangularis Miquel; Oldenlandia parryi (Hance) Kuntze;
O. quadrangularis (Miquel) Kuntze.
Herbs or subshrubs, erect, perennial or perhaps sometimes
annual, to 50 cm tall; stems subterete to 4-angled with angles
thickened to ridged, glabrous. Leaves sessile; blade drying papery, linear, linear-lanceolate, narrowly elliptic, or narrowly
elliptic-oblong, 13.5 0.20.55 cm, glabrous, base acute to
obtuse and usually decurrent, margins usually revolute, apex
acute; secondary veins not visible; stipules shortly fused to petiole bases, triangular to truncate, 11.5 mm, glabrous, rounded
to acute or usually with leaflike appendage, this narrowly elliptic to lanceolate, 210 mm, entire or shortly 2- or 3-lobed or
-setose. Inflorescences terminal and sometimes in axils of uppermost leaves, compound-cymose to paniculate, 413 cm, glabrous, many flowered, pedunculate; peduncle 15.5 cm; axes
mostly regularly dichotomous; bracts linear, 12.5 mm; pedicels (or ultimate inflorescence axes) 0.53 mm. Flowers subsessile or pedicellate, apparently distylous. Calyx glabrous; hypanthium portion turbinate, 0.81 mm; limb lobed nearly to base;
lobes narrowly lanceolate to narrowly triangular, 11.5 mm.
Corolla white, tubular or tubular-funnelform, outside glabrous,
inside tomentose at throat and onto lobes; tube ca. 2 mm; lobes
lanceolate-oblong, ca. 2 mm. Anthers included or exserted, 0.8
1 mm. Stigmas 0.81 mm, included or exserted. Fruit capsular,
subglobose, sometimes slightly didymous, ca. 2 2 mm, glabrous, septicidally then loculicidally dehiscent; seeds several,
black, angled. Fl. and fr. JulNov.
Open fields, grassy slopes, sometimes on ridges of farmlands; low
elevations. Guangdong, Guangxi [Borneo, Cambodia, Indonesia, Malaysia, Thailand, Vietnam].
Merrill and Metcalf (Lingnan Sci. J. 16(3): 400. 1937) noted that
Valeton provided a much-expanded and detailed description of this
species (Bot. Jahrb. Syst. 44: 543. 1910), and they formally synonymized Hedyotis parryi with H. tetrangularis, though as tetrangularia.

58. Hedyotis trinervia (Retzius) Roemer & Schultes, Syst.

Veg. 3: 197. 1818.
san mai er cao
Oldenlandia trinervia Retzius, Observ. Bot. 4: 23. 1786;
Hedyotis rotundifolia Sprengel, nom. illeg. superfl.
Herbs, annual, prostrate to weakly erect, to 15 cm tall;
stems 4-angled to 4-winged, hirsute to pilose. Leaves subsessile
to petiolate; petiole to 2 mm, pilose to hirsute; blade drying papery, ovate, elliptic, or suborbicular, 0.31.2 0.61 cm, adaxially glabrous, abaxially glabrous or sparsely strigillose to hispid, base obtuse and shortly decurrent, margins usually ciliate,

172

RUBIACEAE

apex obtuse and shortly acuminate; secondary veins 2 or 3

pairs; stipules fused to petiole bases, hispidulous to villosulous,
truncate, ca. 0.5 mm, irregularly erose and with 13 narrowly
triangular to linear lobes or bristles 0.31 mm. Inflorescences
axillary, 1-flowered to 24-flowered and cymose or fascicled,
pilose, subsessile; pedicels to 1.5 mm. Flowers subsessile to
pedicellate. Calyx pilosulous to hispidulous; hypanthium portion subglobose, ca. 1 mm; limb lobed nearly to base; lobes
triangular, 11.2 mm. Corolla white, rotate, apparently glabrous; tube ca. 0.5 mm; lobes 12 mm. Anthers exserted or included, 0.51 mm. Stigmas included. Fruit capsular, subglobose, somewhat didymous, 12 11.5 mm, pilosulous or hispidulous, loculicidally dehiscent across top; seeds several, black,
angled. Fl. and fr. OctMar.
Sparse forests on mountains. Hainan (Ledong) [India, Indonesia,
Malaysia, Sri Lanka, N Vietnam].
The descriptions seen suggest that this is a rather distinctive
species. The description here is compiled from Dutta and Deb (Taxon.
Rev. Hedyotis. 2004) and Augustine (J. Econ. Taxon. Bot. 26: 91.
2002).
The synonym Hedyotis rotundifolia was attributed by W. C. Ko
(in FRPS 71(1): 76. 1999) to Candolle, but Candolle clearly indicated
(Prodr. 4: 420. 1830) that Sprengel was the author of the name. W. C.
Ko gave the range of this species as including Tropical Africa, but
Verdcourt (Fl. Trop. E. Africa, Rub. (Pt. 1), 279280. 1976) noted that
throughout Africa this name was long incorrectly applied to Oldenlandia goreensis (Candolle) Summerhayes (= H. goreensis Candolle);
thus, H. trinervia is not currently known from Africa.

59. Hedyotis uncinella Hooker & Arnott, Bot. Beechey Voy.

192. 1833.
chang jie er cao
Hedyotis borrerioides Champion ex Bentham; H. kuraruensis Hayata; H. uncinella var. cephalophora Craib; Oldenlandia uncinella (Hooker & Arnott) Kuntze.
Herbs, perennial, erect, to 70 cm tall; stems 4-angled
and/or sulcate, angles becoming sharp and thickened or narrowly winged, sparsely to densely puberulent or hispidulous to
glabrescent. Leaves opposite or rarely ternate at 1 or 2 nodes,
subsessile to petiolate; petiole to 2.5 mm, puberulent to glabrescent; blade drying papery, ovate-oblong, oblong-lanceolate,
ovate, lanceolate, or elliptic, 1.57.5 0.43.8 cm, both surfaces sparsely to densely puberulent, hispidulous, or pilosulous,
base acute to rounded and usually shortly decurrent, apex acute
to acuminate; secondary veins 4 or 5 pairs; stipules interpetiolar, triangular to narrowly triangular, 23.5 mm, moderately to
densely pilosulous, pilosulous, or puberulent, acute, aristate, or
usually laciniate or with 25 linear lobes or bristles 0.52 mm.
Inflorescences terminal and axillary in axils of uppermost
leaves, capitate or glomerulate, 815 mm in diam., globose,
many flowered, sessile; bracts linear to lanceolate, 0.52 mm.
Flowers sessile or subsessile, distylous. Calyx moderately to
densely pilosulous to hispidulous; hypanthium portion subglobose to obconic, ca. 1 mm; limb lobed essentially to base; lobes
narrowly triangular, 2.54 mm, ciliolate. Corolla white or purple, funnelform or tubular-funnelform, outside puberulent or pilosulous at least on lobes; tube 33.5 mm, tomentose in throat;

lobes oblong-lanceolate, 1.52 mm. Anthers exserted or included, 0.81 mm. Stigma 0.31 mm, included or exserted.
Fruit capsular, subglobose to broadly ovoid, 1.52 1.82 mm,
often somewhat compressed, septicidally dehiscent then loculicidal, with stipes or pedicels to 1 mm; seeds several, pale
brown, angled. Fl. and fr. AprSep.
Dry open fields; 2001200 m. Fujian, Guangdong, Guizhou, Hainan, Hunan, Taiwan [India, Myanmar].
The name Hedyotis uncinella var. cephalophora has been cited as
a combination by some authors, but the basionym cited for it, H.
cephalophora R. Brown, is apparently a nomen nudum.
W. C. Ko (in FRPS 71(1): 60. 1999) described the anthers as ca. 3
mm, which has not been reported by any other authors nor seen on any
specimens.

60. Hedyotis vachellii Hooker & Arnott, Bot. Beechey Voy.

194. 1837.
xiang gang er cao
Oldenlandia vachellii (Hooker & Arnott) Kuntze.
Herbs or subshrubs, erect, to 40 cm tall; stems terete,
weakly angled, or flattened, glabrous. Leaves subsessile to petiolate; petiole to 8 mm, glabrous; blade drying papery to subleathery, oblong-lanceolate, elliptic, lanceolate, or elliptic-oblong, 48.5 13.5 cm, glabrous, base acute to usually cuneate
or obtuse, apex acute; secondary veins indistinct; stipules perhaps interpetiolar or shortly fused around stem, triangular to
ovate, 1.52 mm, marginally densely glandular-ciliolate, acute
to obtuse. Inflorescences terminal, paniculate, cymose, multichotomous, 48 3.58 cm, many flowered, glabrous, purple,
pedunculate; peduncles 24.5 cm; principal lateral axes 0.81.5
cm; bracts lanceolate to narrowly triangular, 0.53 mm; pedicels to 1.5 mm. Flowers subsessile to pedicellate, floral biology
unknown. Calyx glabrous, somewhat succulent; hypanthium
portion turbinate to ellipsoid, 0.91 mm; limb lobed essentially
to base; lobes lanceolate to narrowly triangular, 1.22 mm.
Corolla white to pale green, salverform, outside glabrous; tube
2.53 mm, sparsely pubescent inside; lobes ovate-triangular, ca.
2 mm. Anthers exserted, ca. 1.2 mm. Stigma included. Fruit
capsular, subglobose, ca. 1.8 mm in diam., dehiscent; seeds several, brown. Fl. AprAug.
Hong Kong (New Territories).

61. Hedyotis verticillata (Linnaeus) Lamarck, Tabl. Encycl. 1:

271. 1792.
cu ye er cao
Oldenlandia verticillata Linnaeus, Mant. Pl. 1: 40. 1767;
Hedyotis hispida Retzius; O. hispida (Retzius) Poiret.
Herbs, annual or perennial, diffusely branched, often decumbent, to 30 cm tall; stems flattened, subterete, or 4-angled
and/or usually 2-sulcate, sparsely to densely hirtellous, hispidulous, and/or scaberulous. Leaves sessile to petiolate; petiole to 2
mm, scaberulous or hispidulous to glabrescent; blade drying papery or thinly leathery, narrowly elliptic or linear-lanceolate,
2.56 0.31.3(2) cm, adaxially glabrous or scaberulous at
least near margins, abaxially glabrous to densely hirtellous or

RUBIACEAE

hispidulous at least on midrib, base acute to obtuse, margins often revolute at least when dry, apex acute or acuminate; secondary veins not visible; stipules shortly fused to petiole bases, triangular to subtruncate, 13 mm, sparsely to densely hispidulous
to hirtellous, with 59 linear lobes or bristles 18 mm. Inflorescences axillary, glomerulate to congested-cymose, 510 mm
in diam., several flowered, moderately to densely hispidulous,
sessile; bracts linear to lanceolate, 14 mm. Flowers sessile to
subsessile, apparently homostylous (e.g., Anon. 788, MO). Calyx densely hispidulous; hypanthium portion obconical to subglobose, ca. 1 mm; limb lobed essentially to base; lobes lanceolate to triangular, 12 mm, ciliolate. Corolla white, funnelform,
outside glabrous except lobes sometimes bearded at apex; tube
ca. 2 mm, glabrous inside; lobes lanceolate, 1.82 mm. Anthers
exserted, ca. 1 mm. Stigma ca. 0.3 mm. Fruit capsular, ovoid,
23 1.52 mm, loculicidal across top, with calyx lobes to 3
mm, with petioles to 1.5 mm; seeds numerous, pale brown,
angled. Fl. and fr. MarNov.
Tussocks or thickets on foothills, roadsides, sparse forests; 200
1600 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Zhejiang [Bangladesh, Bhutan, India, Indonesia, Japan (Ryukyu Islands),
Malaysia, Myanmar, Nepal, Philippines, Singapore, Thailand, Vietnam].
As noted under Hedyotis tenelliflora, Spermacoce stricta may be a
synonym of H. verticillata.
W. C. Ko (in FRPS 71(1): 42. 1999) described the leaves as up to
2 cm wide; this has not been reported by other authors nor seen on any
specimens.

62. Hedyotis vestita R. Brown ex G. Don, Gen. Hist. 3: 526.

1834.
mai er cao
Hedyotis capituliflora Miquel; H. costata (Roxburgh) Kurz
(1876), not R. Brown ex G. Don (1834); Metabolos lineatus
Bartling ex Candolle; Spermacoce costata Roxburgh.

173

bescent in throat; lobes narrowly elliptic to triangular, 0.81.2

mm. Anthers 0.51 mm, exserted. Stigma 0.30.6 mm, exserted. Fruit indehiscent, subglobose, 0.81.5 mm in diam., pilosulous; seeds 68, black, trigonous. Fl. and fr. JulNov.
Forest margins in valleys, open fields, grassy slopes; 4002000 m.
Guangdong, Guangxi, Hainan, Yunnan [India, Indochina, Indonesia,
Malaysia, Philippines, Thailand].
This species has been treated by many previous authors under the
name Hedyotis costata; the basionym of that name, Spermacoce costata
Roxburgh, is indeed the oldest name for this species, but that epithet
was previously occupied for another species in Hedyotis when Kurz
published his name and, thus, was blocked from transfer.

63. Hedyotis wangii R. J. Wang, Acta Phytotax. Sin. 41: 85.

2003.
qi wu er cao
Herbs, annual or perennial, to 6 cm tall; stems glabrous.
Leaves petiolate; petiole 36 mm; blade drying papery, ellipticoblong or ovate-elliptic, 35 1.52.3 cm, glabrous, base obtuse, apex acute; secondary veins ca. 5 pairs; stipules broadly
rounded, ca. 0.5 mm, with 3 or 4 linear to narrowly triangular
lobes or bristles ca. 2 mm. Inflorescence terminal, capitate or
subcapitate, pedunculate; peduncle ca. 1.7 cm; head solitary,
ovoid, ca. 0.5 cm, 4- or 5-flowered; bracts linear-lanceolate, ca.
3 mm. Flowers sessile. Calyx sparsely hirtellous or pilosulous
or perhaps glabrous; hypanthium portion ellipsoid to obconic,
0.81 mm, ribbed; limb lobed nearly to base; lobes lanceolate,
narrowly triangular, or linear, ca. 2 mm. Corolla white, tubularfunnelform, inside villosulous in upper part of tube and onto
lobes; tube ca. 7 mm; lobes lanceolate, ca. 1 mm. Anthers ca. 2
mm, apparently exserted. Stigmas ca. 0.8 mm, apparently exserted. Fruit capsular, subglobose, ca. 3 mm in diam., pilosulous
to perhaps glabrescent; seeds numerous, angled, reticulate. Fl.
and fr. Sep.
Mixed forests; ca. 1400 m. S Yunnan (Jinghong).

Herbs, annual or perennial, diffusely branched, weak to

clambering or prostrate, to 60 cm tall; stems flattened, 4-angled,
or subterete, sometimes sulcate, densely villous or hirsute with
trichomes drying golden yellow. Leaves subsessile to petiolate;
petiole 110 mm, densely hirtellous to villous; blade drying
membranous to papery, lanceolate, elliptic-lanceolate, or elliptic, 28 0.53 cm, adaxially sparsely to densely strigose
and/or strigillose to hirsute on lamina and moderately to densely
pilosulous on principal veins, abaxially sparsely strigillose to
glabrescent on lamina and densely strigose to hirsute on principal veins, base cuneate to obtuse and frequently decurrent,
apex acute to usually acuminate; secondary veins 46 pairs;
stipules fused to petiole bases, truncate to triangular, 24 mm,
moderately to densely hirtellous, hirsute, or villous, with 35
linear or setose lobes 25 mm. Inflorescences axillary, congested-cymose, capitate, or fasciculate, 520 mm, densely hirtellous to hirsute, several to many flowered, pedunculate; peduncles 13 per leaf axil, 212 mm; bracts subulate, to 1 mm.
Flowers sessile or subsessile, distylous. Calyx densely hirtellous to villosulous; hypanthium portion turbinate to obconic, ca.
0.5 mm; limb lobed nearly to base; lobes lanceolate to triangular, 0.31 mm. Corolla white or purple, tubular to funnelform, outside puberulent to pilosulous; tube 1.21.5 mm, pu-

The calyx (including presumably the hypanthium) was described

in the protologue as pubescent but both this and the fruit are shown in
the protologue figure as glabrous.

64. Hedyotis wuzhishanensis R. J. Wang, Novon 18: 266.

2008.
wu zhi shan er cao
Subshrubs or shrubs, erect, perennial, to 1 m tall; stems
terete, densely villous to hirtellous. Leaves petiolate; petiole 3
10 mm, densely hirtellous; blade drying papery, elliptic, ovate,
or lanceolate, 2.56.2 1.43.3 cm, adaxially sparsely to moderately strigose to hispidulous, abaxially densely to moderately
villosulous to hispidulous, base cuneate, obtuse, or rounded,
apex acute; secondary veins 4 or 5 pairs, prominulous abaxially; stipules persistent or perhaps tardily deciduous, interpetiolar, triangular, 12.5 mm, densely hirtellous, acute to aristate.
Inflorescences terminal, pseudoaxillary, and/or axillary in uppermost leaf axils, subcapitate, 0.71.2 cm in diam., densely
hirtellous, (2)615-flowered, subsessile, enclosed by lanceolate bracts 510 mm. Flowers sessile to subsessile, biology unknown. Calyx densely hirtellous; hypanthium portion ellipsoid,
ca. 1.8 mm; limb lobed essentially to base; lobes narrowly trian-

RUBIACEAE

174

gular, 1.53 mm. Corolla white, tubular-funnelform, outside

glabrescent; tube 5.56.5 mm, inside villosulous in upper part
and throat; lobes narrowly triangular to lanceolate, ca. 2.5 mm.
Anthers exserted, 1.52 mm. Stigmas included, 12 mm. Fruit
capsular, ovoid, ca. 3 2.5 mm, densely pilosulous or hirtellous, loculicidally dehiscent then septicidal; seeds ca. 15, black,
angled, ca. 0.5 mm. Fl. and fr. OctDec.
Forests; 6001600 m. Hainan.

65. Hedyotis xanthochroa Hance, J. Bot. 23: 324. 1885.

huang ye er cao
Oldenlandia xanthochroa (Hance) Kuntze.
Herbs, perennial, erect, to 40 cm tall; stems obtusely 4angled, shallowly sulcate, sparsely tomentulose, hispid, or pilosulous becoming glabrescent. Leaves sessile; blade drying
stiffly papery, elliptic-oblong to ovate, 45.5 0.62.6 cm,
adaxially sparsely to moderately scabrous to hispid on lamina
and densely hispidulous to scaberulous on principal veins,
abaxially hirtellous or hispidulous throughout with pubescence
denser on veins, black glandular-punctate under magnification
on both surfaces, base broadly rounded and somewhat amplexicaul, apex shortly acuminate; secondary veins ca. 4 pairs; stipules fused to petiole bases, triangular-lanceolate, 45 mm,
hispidulous on margins or throughout, marginally laciniate to
setose-lobed, apex acute to acuminate. Inflorescences terminal,
congested-cymose to subcapitate, 1.54.5 cm, sparsely to
densely hispid to pilosulous, several flowered, shortly pedunculate; peduncles 0.51.5 cm; bracts lanceolate, ovate, or narrowly elliptic, 48 mm; pedicels 14 mm. Flowers subsessile.
Calyx sparsely villous; limb lobed; lobes linear-lanceolate, ca. 6
mm. Corolla dark purple, tubular, outside villous; tube ca. 5.5
mm; lobes ca. 5.5 mm. Anthers exserted. Stigma exserted. Fruit
capsular, ellipsoid-oblong to obovoid, 3.54.5 mm, cartilaginous to papery, densely hispid to pilosulous, loculicidally and
septicidally dehiscent through apical portion then sometimes
splitting septicidally throughout, beak conical, ca. 1 mm, with
persistent calyx lobes narrowly triangular, 55.5 mm, with partially persistent septal structure spongy-cartilaginous, ellipsoid,
33.5 mm, deeply pitted where seeds were situated; seeds several, black, triangular to irregularly angled, ca. 1 mm. Fl. Jul
Aug, fr. Dec.
Mountain slopes in valleys at middle elevations. Guangdong.

66. Hedyotis yangchunensis W. C. Ko & Zhang, J. S. China

Agric. Univ. 16(4): 45. 1995.
yang chun er cao
Herbs or subshrubs, erect, to 50 cm tall; stems 4-angled to
subterete, densely scabrous to smooth. Leaves petiolate, often
crowded at tops of stems; petiole 1520 mm, glabrous; blade
drying thickly papery, narrowly elliptic, oblanceolate, or elliptic-oblong, (3)812 (1)34.5 cm, glabrous, base acute and
decurrent, apex obtuse to broadly obtuse; secondary veins 57
pairs; stipules broadly triangular, ca. 16 mm, lacerate or with
several narrow lobes. Inflorescences and flowers not seen. Infructescences terminal, in a panicle of congested-cymose or
capitate heads; heads 25, hemispherical to subglobose, 22.3
cm in diam., several flowered, pedunculate, enclosed by a pair
of ovate reduced leaves or leaflike bracts ca. 15 mm; peduncles
57 cm; floral bracts linear, 57 mm; pedicels 23 mm. Fruit
capsular, ellipsoid or subglobose, ca. 3 22.5 mm, membranous, septicidally dehiscent then loculicidal, with calyx lobes
lanceolate to elliptic, 33.5 mm, longitudinally veined, long
acuminate; seeds several, black, angled. Fl. AprJun.
Hills, sparse forests in valleys; ca. 400 m. Guangdong (Yangchun).

67. Hedyotis yazhouensis F. W. Xing & R. J. Wang, Acta

Phytotax. Sin. 41: 87. 2003.
ya zhou er cao
Herbs or subshrubs, perennial, erect, to 10 cm tall; stems
weakly 4-angled. Leaves sessile or subsessile, crowded at tops
of stems; blade drying papery, obovate or ovate-elliptic, 820
412 cm, base obtuse, apex obtuse to broadly obtuse; secondary
veins 58 pairs; stipules obtrapezoid to spatulate, 26 mm,
lacerate with lobes ca. 1 mm. Inflorescence terminal, capitate,
hemispherical, 35 cm in diam., several flowered, sessile to
subsessile; bracts linear-lanceolate, ca. 8 mm, acute. Flowers
sessile. Calyx densely puberulent; hypanthium portion ellipsoid
to obovoid, 34 mm; limb divided essentially to base; lobes
linear, ca. 35 1 mm. Corolla blue-purple, slenderly funnelform, outside sparsely puberulent; tube ca. 30 mm; lobes ovate,
ca. 1 mm. Stigmas included. Fruit indehiscent, ellipsoid to obovoid, ca. 3 mm, densely pilosulous; seeds 812, black, papillose. Fl. and fr. Oct.
Understories of tropical rain forests; 100300 m. S Hainan
(Sanya).

36. HIMALRANDIA T. Yamazaki, J. Jap. Bot. 45: 340. 1970.

xu mi qian shu shu
Chen Tao (); Charlotte M. Taylor
Shrubs, sometimes late deciduous, often with short shoots, unarmed [sometimes with spinescent short shoots]. Raphides absent.
Leaves opposite, often crowded at apices of short shoots, sometimes with domatia; stipules persistent, interpetiolar, triangular.
Inflorescences terminal usually on short shoots, 1-flowered, sessile or subsessile, enclosed by stipules or perhaps stipuliform bracts.
Flowers bisexual, monomorphic. Calyx limb 5-lobed. Corolla pale green to pale yellow, salverform, hirsute inside tube; lobes 5, convolute in bud. Stamens 5, inserted in corolla throat, partially to fully exserted; filaments short or reduced; anthers apparently
dorsifixed. Ovary 2-celled, ovules 2 or 3 in each cell on axile placentas; stigma fusiform, 2-lobed or 2-grooved, exserted. Fruit baccate, indehiscent, globose, apparently fleshy, color unknown, with calyx limb persistent; seeds 14, ellipsoid, medium-sized.
About three species: Afghanistan, Bhutan, China, India, Nepal, Pakistan; one species (endemic) in China.

RUBIACEAE

1. Himalrandia lichiangensis (W. W. Smith) Tirvengadum,

Nordic J. Bot. 3: 462. 1983.
xu mi qian shu
Randia lichiangensis W. W. Smith, Notes Roy. Bot. Gard.
Edinburgh 8: 200. 1914.
Shrubs, 0.63 m tall, many branched; branches stout,
weakly flattened to terete, rigid, pilosulous or strigillose to glabrescent. Leaves sessile or subsessile with petiole to 1 mm;
blade drying papery to thinly leathery and often black, obovate
or obovate-spatulate, 16.5 0.63.5 cm, both surfaces moderately to densely strigillose, base cuneate to obtuse, apex acute
to obtuse; secondary veins 35 pairs, sometimes with pilosulous

175

domatia in abaxial axils; stipules ovate to triangular, 2.55 mm,

densely pilosulous, acute to acuminate, often spreading to reflexed. Calyx pilosulous; hypanthium portion ellipsoid, ca. 2
mm; limb ca. 3 mm, deeply lobed; lobes triangular, ciliate,
acute. Corolla yellow; tube ca. 3 mm; lobes triangular to ovate,
ca. 5 mm, acute. Berry 56 mm in diam.; seeds 1 or 2, ca. 3 mm
in diam. Fl. May, fr. JulNov.
Forests or thickets at streamsides in valleys or on mountains;
14002400 m. Sichuan, N Yunnan.
The stamens are described as either partially exserted, according
to the description of W. C. Chen (in FRPS 71(1): 360. 1999), or fully
exserted, according to the accompanying figure (p. 359, t. 94).

37. HYMENODICTYON Wallich in Roxburgh, Fl. Ind. 2: 148. 1824, nom. cons.
tu lian qiao shu
Chen Tao (); Charlotte M. Taylor
Shrubs or trees [or sometimes epiphytic or lianescent], usually deciduous, unarmed; bark notably bitter. Raphides absent.
Leaves opposite, usually with domatia; margins rarely irregularly lobed and/or serrate; stipules deciduous, interpetiolar, triangular to
ligulate, entire or glandular-serrate, usually strongly reflexed. Inflorescence terminal and/or axillary, spiciform to racemiform or
sometimes paniculate with axes racemiform or spiciform, many flowered, erect to pendulous, pedunculate, bracteate with 14 leaflike to petaloid, stipitate, veined basal bracts and other bracts usually reduced [or sometimes well developed]. Flowers sessile to
shortly pedicellate, bisexual, monomorphic. Calyx limb deeply 5(or 6)-lobed. Corolla white, green, yellow, or red, funnelform or narrowly campanulate, inside glabrous; lobes 5, valvate in bud, apparently often ascending or remaining partially closed at anthesis.
Stamens 5, inserted in corolla tube below throat, included; filaments short, flattened; anthers basifixed, sagittate at base. Ovary 2celled, ovules several to numerous in each cell on axile placentas; stigma fusiform or capitate, well exserted. Infructescences with
pedicels and sometimes peduncles often elongating and/or becoming reflexed. Fruit capsular, ellipsoid-oblong to obovoid or ellipsoid, loculicidally dehiscent into 2 valves, woody to cartilaginous, with calyx limb deciduous; seeds numerous, medium-sized, flattened, with broad, membranous, shortly erose, basally 2-lobed wing; endosperm fleshy; embryo small; cotyledon oblong or orbicular.
Twenty-two species: Africa, tropical Asia, and Madagascar; two species in China.
The flowers apparently all open nearly simultaneously on a plant, probably within a very few days at most. The corolla lobes appear to remain
partly closed when the flower is mature and the stigma well exserted; Razafimandimbison and Bremer (Bot. J. Linn. Soc. 152: 335. 2006) reported
that the flowers are protandrous, so this may be a secondary position after the anthers have released their pollen and while the stigma is receptive.
Occasional irregularly lobed leaves, similar to those of plants of Hymenodictyon, are found in a few other Rubiaceae genera (e.g., Simira Aublet of the
Neotropics).
W. C. Chen (in FRPS 71(1): 227. 1999) described the corolla lobes as imbricate in bud and the anthers as dorsifixed, but Bridson and Verdcourt
(Fl. Trop. E. Africa, Rub. (Pt. 2), 452. 1988) and Razafimandimbison and Bremer (loc. cit.) described them as valvate and basifixed, respectively,
which corresponds to observations of Chinese specimens (Henry 12150, MO!).

1a. Inflorescences axillary, at each leaf axil simple, with 1 racemiform or spiciform axis; leaf blade glabrous to
glabrescent on both surfaces or sometimes pilosulous abaxially ............................................................................... 1. H. flaccidum
1b. Inflorescences axillary and sometimes also terminal, simple or at least part branched and paniculate with
several spiciform to racemiform axes; leaf blade pilosulous on both surfaces ............................................................. 2. H. orixense
1. Hymenodictyon flaccidum Wallich in Roxburgh, Fl. Ind. 2:
152. 1824.
tu lian qiao
Trees, deciduous, 620 m tall; bark gray, smooth; branches
rather stout, weakly flattened to terete, glabrous to puberulent.
Leaves often crowded at ends of branches; petiole 2.59 cm,
puberulent or pilosulous to glabrescent; blade drying papery or
thinly leathery, ovate, obovate, elliptic, or elliptic-oblong, 10
26 715 cm, glabrous to glabrescent on both surfaces or
sometimes pilosulous abaxially, base acute to obtuse, margins
entire or rarely serrate or lobed in distal part, apex acute to acu-

minate or rarely rounded; secondary veins 711 pairs, usually

with pilosulous domatia; stipules ovate-oblong or triangular, 5
20 mm, densely pilosulous to glabrescent, acute to obtuse or
bilobed for up to 1/2. Inflorescences axillary, simple, densely
racemiform to spiciform, 1030 cm, pilosulous to hirtellous,
often pendulous; peduncle 35 cm; basal bracts 1 or 2, with
blade leathery, ovate to elliptic-oblong or narrowly elliptic, 4
8.5 23 cm, pilosulous to glabrescent, on stipes 35.5 cm;
pedicels 0.52 mm. Calyx densely puberulent; ovary portion
ellipsoid, 1.21.5 mm; limb lobed essentially to base; lobes
triangular to elliptic, 11.5 mm, ciliolate, acute. Corolla yellow
to red, densely puberulent to glabrescent outside; tube ca. 2

RUBIACEAE

176

mm, slender and cylindrical in basal portion then abruptly inflated just below lobes; lobes ligulate to lanceolate or ovate, 2
3.5 mm, obtuse to acute. Style exserted for 25 mm. Fruiting
pedicels to 8 mm, reflexed. Capsules dark brown, 1.21.5
0.50.8 cm, woody, with several prominent whitened, ellipsoid
lenticels; seeds (including wing) ca. 10 5 mm. Fl. MayJul, fr.
AugDec.
Forests or thickets at streamsides or in valleys; 3003000 m.
Guangxi, Sichuan, Yunnan [Bhutan, N India, Nepal, N Vietnam].
The name Hymenodictyon yunnanense was written by Pitard on
a specimen of this species from Yunnan, China (Ducloux 6767, P!) but
never validly published (Razafimandimbison & Bremer, Bot. J. Linn.
Soc. 152: 370. 2006).

2. Hymenodictyon orixense (Roxburgh) Mabberley, Taxon 31:

66. 1982.
mao tu lian qiao
Cinchona orixensis Roxburgh, Bot. Descr. Swietenia, 21.
1793; C. excelsa Roxburgh; Hymenodictyon excelsum (Roxburgh) Wallich.
Trees, deciduous, to 25 m tall; bark smooth, gray;
branches rather stout, weakly flattened to terete, puberulent to
densely pilosulous or glabrescent. Leaves often grouped near
ends of branches; petiole 217 cm, pilosulous; blade drying papery or membranous, ovate-elliptic, elliptic, or broadly elliptic,
922 614 cm, both surfaces pilosulous with pubescence
sometimes denser abaxially, base acute to obtuse, margins en-

tire, apex shortly acuminate or acute; secondary veins 710

pairs, sometimes with pilosulous domatia; stipules ovate to
triangular or lanceolate, 520 mm, densely pilosulous, obtuse to
acute or bilobed for up to 1/2. Inflorescences terminal and in
axils of uppermost leaves, 1520 cm, simple or branched at
least in part, with axes densely spiciform to racemiform, pilosulous, usually pendulous; peduncles ca. 6 cm; basal bracts 2
4, with blade papery to leathery, ovate to elliptic or ellipticoblong, 917 25.5 cm, pilosulous, with stipe 38 cm. Flowers subsessile or with pedicels to 2 mm. Calyx densely puberulent to pilosulous; ovary portion subglobose to ellipsoid, 11.5
mm; limb lobed essentially to base; lobes triangular to elliptic,
11.5 mm. Corolla white or brown, outside densely puberulent
to pilosulous; tube 2.53.5 mm, slenderly cylindrical then
abruptly inflated at lobes; lobes ligulate to lanceolate, 22.5
mm, acute. Style exserted for 25 mm. Fruiting pedicels to 10
mm, reflexed. Capsules brown, 1.23 0.51.1 cm, woody,
with prominent whitened, elliptic lenticels; seeds (including
wing) 78 mm. Fl. MayJul, fr. MayDec.
Thickets or forests at riversides, at field edges, and in valleys;
1001700 m. Sichuan, Yunnan [Cambodia, India, Indonesia (Java),
Kashmir, Laos, Malaysia, Myanmar, Nepal, Philippines, Thailand, Vietnam].
Mabberley (loc. cit.) noted that Roxburgh studied this species in
India as one of several plants (in several families) for which the bitter
bark was used medicinally. In India this species is used for good
quality wood for furniture and small items, and its bark as a febrifuge
and a source of dye (color not noted; Razafimandimbison & Bremer,
Bot. J. Linn. Soc. 152: 375377. 2006).

38. HYPTIANTHERA Wight & Arnott, Prodr. Fl. Ind. Orient. 1: 399. 1834.
zang yao mu shu
Chen Tao (); Charlotte M. Taylor
Shrubs or small trees, unarmed. Raphides absent. Leaves opposite, decussate, apparently without domatia; stipules persistent,
interpetiolar, generally triangular with apices often twisted together at 180 in bud. Inflorescences axillary, glomerulate, several flowered, sessile, bracteate. Flowers sessile, apparently bisexual or reportedly at least sometimes unisexual, fragrant. Calyx limb 4- or 5lobed. Corolla white, shortly funnelform to subrotate, inside pubescent in throat; lobes 4 or 5, convolute in bud. Stamens 4 or 5,
inserted in corolla tube, included or partially exserted; filaments short or reduced; anthers dorsifixed, pubescent on base and outer
surface, with connective apically prolonged and widened. Ovary 2-celled, ovules 610 in each cell on axile placentas; stigma 2-lobed
with lobes oblong and pubescent, exserted. Fruit baccate, fleshy, ovoid or subglobose, at least sometimes black, with calyx limb persistent; seeds several, medium-sized, flattened, angular, with testa thickly fibrous striate, with hilum terminal and rather broad; endosperm fleshy; embryo small; cotyledons ovate, flat; radicle terete.
One or a few species: China, Bangladesh, Bhutan, India, Laos, Myanmar, Nepal, Thailand, Vietnam; one species in China.
Robbrecht and Puff (Bot. Jahrb. Syst. 108: 126127, table 8. 1986) described the flowers as unisexual, but the corresponding condition of the
plants (e.g., dioecious, polygamo-dioecious) is not noted and the specimens and figures seen seem to have bisexual flowers. W. C. Chen (in FRPS
71(1): 386. 1999) did not mention unisexual flowers for this species, and the accompanying figure apparently shows bisexual flowers (p. 387, t. 102).
He described the stipules as caducous, but this has not been seen on any specimens of Hyptianthera nor reported by other authors, and it contradicts
the figure. He also described the ovules as pendulous from the top of the cell, but Robbrecht and Puff (loc. cit.: 8691), in a detailed morphological
survey focused on gynoecium characters, described them as borne on axile placentas.

1. Hyptianthera stricta (Roxburgh) Wight & Arnott, Prodr. Fl.

Ind. Orient. 1: 399. 1834.
zang yao mu
Randia stricta Roxburgh, Fl. Ind. 2: 145. 1824.
Shrubs or small trees, 28 m tall; branches flattened be-

coming angled to subterete, glabrous. Petiole 0.41 cm, glabrous; leaf blade drying papery or thinly leathery and usually
reddish brown, oblong-lanceolate, elliptic-oblong, or lanceolate,
515 15 cm, adaxially glabrous and slightly shiny, abaxially
glabrous or sometimes hirtellous or pilosulous along principal
veins, base cuneate to acute, apex acuminate to long acuminate;
secondary veins 59 pairs; stipules triangular to ovate, 58 mm,

RUBIACEAE

glabrous, abaxially smooth to weakly keeled, acuminate to

shortly aristate. Inflorescences 610 mm in diam.; bracts triangular to ovate, 12.5 mm, adaxially white strigose, abaxially
glabrescent, marginally ciliate, acute to acuminate. Calyx with
ovary portion ellipsoid, ca. 0.5 mm, glabrescent; limb outside
(i.e., abaxially) strigillose to glabrescent, inside (i.e., adaxially)
densely strigillose, deeply lobed, with basal tubular portion 0.5
1 mm; lobes 5, lanceolate to narrowly triangular, 1.752 mm,

177

ciliate, shortly acuminate. Corolla abaxially glabrous; tube 12

mm; lobes 5, spatulate to subelliptic, 12 mm, adaxially white
strigose, obtuse or infrequently emarginate. Stigma lobes 12
mm. Berry yellowish green, 89 56 mm, puberulent to glabrous; seeds usually 8, ca. 5 3 mm. Fl. AprAug, fr. AugFeb.
Forests or thickets at streamsides or on mountains; 1001500 m.
Xizang (Mdog), Yunnan [Bangladesh, Bhutan, India, Laos, Myanmar,
Nepal, Thailand, Vietnam].

39. IXORA Linnaeus, Sp. Pl. 1: 110. 1753.

long chuan hua shu
Chen Tao (); Charlotte M. Taylor
Schetti Adanson; Tsiangia But, H. H. Hsue & P. T. Li.
Shrubs or small trees or occasionally perhaps climbing (Ixora hekouensis), unarmed. Raphides absent. Leaves opposite or rarely
ternate, decussate, without domatia; petioles articulate at base; stipules persistent to caducous, interpetiolar or shortly united around
stem, triangular, acute to usually aristate. Inflorescences terminal on principal stems [or sometimes terminal on reduced lateral stems
and appearing axillary], cymose to corymbiform or paniculiform, few to many flowered, sessile to pedunculate, bracteate or bracts
reduced; axes often articulate; bracteoles when present often fused in pairs. Flowers pedicellate or sessile, bisexual, monomorphic,
often fragrant. Calyx limb truncate or 4-lobed. Corolla yellow, orange, red, or white, sometimes becoming reddened when dry,
salverform with tube slender, inside glabrous or pubescent at throat; lobes 4 [rarely to 9], convolute in bud. Stamens 4, inserted at
corolla throat, partially to fully exserted; filaments short or reduced; anthers dorsifixed near base. Ovary 2-celled, ovules 1 in each
cell, pendulous from axile placentas attached in upper part of septum; style in upper portion fusiform or clavate; stigmas 2, linear,
recurved, exserted. Fruit black or red, drupaceous, leathery or fleshy, subglobose to ellipsoid or ovoid, with calyx limb persistent;
pyrenes 2, 1-celled, each with 1 seed, plano-convex or concavo-concave, smooth dorsally (i.e., abaxially), leathery, papery, or crustaceous; seeds medium-sized, ellipsoid to oblanceolate, grooved and concave ventrally (i.e., adaxially); testa membranous; endosperm
cartilaginous; radicle terete, hypogeous.
About 300400 species: widespread in tropical Africa, America, Asia, Madagascar, and Pacific islands; 18 species (nine endemic) in China.
Ixora has been studied in SE Asia only by Bremekamp; he published several articles treating the species of several regions there (cited by De
Block, Opera Bot. Belg. 9: 213. 1998) but not specifically treating the Chinese species, although his circumscription of the region Burma and the
Andaman Islands included some species from Xizang (Bremekamp, J. Bot. (London) 75: 108111, 169175, 260266, 295298, 318326. 1937).
Several species of Ixora are widely cultivated in tropical regions as ornamentals, notably I. casei Hance, I. coccinea Linnaeus, I. finlaysoniana,
and sometimes I. chinensis. In cultivation several of these have various forms with a wide range of flower color, flower and leaf size, and plant height,
and sometimes they do not set fruit. Ixora coccinea and I. casei are not treated in this current flora but are included in the key to species below for
identification. Reynolds and Forster (Austrobaileya 7(2): 253278. 2006) reported that I. coccinea is locally adventive in some parts of Australia,
which may be a possibility in China. The most commonly cultivated Ixora species were discussed in detail by Fosberg and Sachet (Baileya 23(2): 74
85. 1989).
Ixora foonchewii was described and illustrated as having consistently 5 corolla lobes and a stout, shortly bilobed stigma, so it does not seem to
belong to Ixora; it is provisionally included here in Tarenna.

1a. Calyx lobes 36 mm.

2a. Leaves relatively narrow, 14 cm wide, acute to acuminate at both ends, with secondary veins
1215 pairs; corolla tube 3040 mm, lobes 67 mm ....................................................................................... 18. I. yunnanensis
2b. Leaves not notably narrow, 39 cm wide, rounded to cuneate at base, obtuse, acute, or acuminate
at apex, with secondary veins 912 pairs; corolla tube 2030 mm, lobes 56 mm.
3a. Corolla lobes obtuse to rounded ............................................................................................................... 2. I. cephalophora
3b. Corolla lobes acute to subacute .................................................................................................................. 5. I. finlaysoniana
1b. Calyx lobes 0.11.5 mm.
4a. Leaves with secondary veins 5 or 6 pairs.
5a. Petioles 02 mm; corolla lobes 815 mm; flowers variously yellow to red; plants
cultivated ............................................................................................................................ I. coccinea (see comment above)
5b. Petioles 28 mm; corolla lobes 67 mm; flowers white; plants native ....................................................... 7. I. hainanensis
4b. Leaves with secondary veins 716 pairs.
6a. Petioles 1020 mm, at least some of them more than 10 mm.
7a. Leaves shallowly bullate with principal veins impressed on puffy upper surface; corolla white
to pink.
8a. Corolla tube 3040 mm, lobes 57 mm ............................................................................................... 12. I. nienkui
8b. Corolla tube 1518 mm, lobes ca. 5 mm ............................................................................................... 17. I. tsangii

RUBIACEAE

178

7b. Leaves plane, with principal veins flat to prominent on flat upper surface; corolla red, purple,
yellow-orange, pink, or white.
9a. Corolla red, tube ca. 10 mm, lobes ca. 5 mm ................................................................................. 13. I. paraopaca
9b. Corolla white, pink, yellow-orange, or red, tube 2540 mm, lobes 515 mm.
10a. Stems relatively stout, youngest sections 35 mm in diam.; plants cultivated
............................................................................................................................. I. casei (see comment above)
10b. Stems slender, youngest sections 13 mm in diam.; plants native.
11a. Corollas yellow-orange to red, with lobes 1012 mm; stipule body
(i.e., triangular portion) 23 mm .......................................................................................... 6. I. fulgens
11b. Corollas white to pink, with lobes 57 mm; stipule body 310 mm ................................ 12. I. nienkui
6b. Petioles 010 mm, at least some of them shorter than 10 mm.
12a. Leaves sessile or subsessile with petioles up to 4 mm, base cordate, cordulate, truncate,
cuneate, rounded, or obtuse.
13a. Corollas red-purple, tube 89 mm, lobes 45 mm .............................................................. 11. I. longshanensis
13b. Corollas white, red, or purple, tube 1345 mm, lobes 57.2 mm.
14a. Corolla lobes broadly obtuse to rounded .............................................................................. 3. I. chinensis
14b. Corolla lobes acute to obtuse.
15a. Leaves 1023 5.59 cm ......................................................................................... 1. I. auricularis
15b. Leaves 4.512 27 cm.
16a. Corolla lobes obtuse ................................................................................. 14. I. philippinensis
16b. Corolla lobes acute ......................................................................................... 15. I. subsessilis
12b. Leaves with petioles 210 mm, base rounded, cordulate, cuneate, obtuse, truncate, or acute.
17a. Peduncle ca. 14 cm ......................................................................................................................... 10. I. insignis
17b. Peduncle 0.45 cm.
18a. Inflorescences with tertiary and often also quaternary axes well developed and
spreading at 4590 or more; corolla tube 711 mm ............................................................... 4. I. effusa
18b. Inflorescences with tertiary and quaternary axes developed to reduced and
ascending at less than 45; corolla tube 1840 mm.
19a. Corolla lobes ovate, elliptic, or broadly elliptic, at apex broadly obtuse
to rounded .................................................................................................................... 3. I. chinensis
19b. Corolla lobes elliptic-oblong, narrowly elliptic-oblong, ovate-lanceolate,
narrowly spatulate, narrowly lanceolate, or oblong-lanceolate, at apex
obtuse to acute.
20a. Calyx puberulent or glabrescent; corolla tube 1820 mm, lobes 57 mm.
21a. Leaves 1018 cm, at base cuneate to obtuse, at apex acuminate to
acute; bracts 3.55 mm .......................................................................... 8. I. hekouensis
21b. Leaves 4.510 cm, at base truncate, rounded, or cordulate, at
apex obtuse or rounded and apiculate; bracts 1.52 mm ............... 14. I. philippinensis
20b. Calyx glabrous; corolla tube 2040 mm, lobes 57 mm.
22a. Leaves 1517 6.57.5 cm; stipules villous adaxially ............................ 16. I. tibetana
22b. Leaves 415 15 cm; stipules glabrescent adaxially.
23a. Low to medium-sized plants, flowering at 0.23 m tall; leaves
rounded, obtuse, or bluntly acute at apex; corollas with
tube 3040 mm, in bud acute to rather sharply acute ................. 7. I. hainanensis
23b. Medium-sized shrubs, flowering at 13 m tall; leaves sharply
acute to acuminate at apex; corollas with tube 2025 mm,
in bud sharply acute to acuminate ........................................................ 9. I. henryi
1. Ixora auricularis Chun & F. C. How ex W. C. Ko, Guihaia
19: 99. 1999.
er ye long chuan hua
Shrubs or small trees, to 6 m tall; branches glabrous.
Leaves opposite, sessile or subsessile; blade drying thinly papery, brown adaxially, pale abaxially, oblanceolate, oblongelliptic, or obovate-elliptic, 1023 5.59 cm, glabrous on
both surfaces, base cordate-auriculate, apex shortly acuminate;
secondary veins 1013 pairs; stipules ovate to broadly triangular, 718 mm, abruptly narrowed and aristate. Inflorescences

terminal, corymbose, 615 cm wide; peduncle 35 cm, articulate near base, at articulation with reduced leaves ca. 3 cm;
bracteoles linear-lanceolate, ca. 1 mm; pedicels 23 mm. Flowers sessile or pedicellate. Calyx with hypanthium obconic; limb
deeply lobed; lobes ligulate. Corolla purplish red; tube 2030
mm, glabrous at throat; lobes oblanceolate-oblong, 77.2 3
3.5 mm, acute. Drupe globose, 68 mm in diam. Fl. MayJun.
Broad-leaved forests or thickets at middle elevations; ca. 1100
m. Yunnan.
The protologue text described the calyx post anthesis as 22.5
mm, but the protologue figure illustrated it as 4 mm at anthesis.

RUBIACEAE

2. Ixora cephalophora Merrill, J. Arnold Arbor. 23: 194. 1942.

tuan hua long chuan hua
Shrubs, 12 m tall; branches glabrous. Leaves opposite;
petiole 12 cm, glabrous; blade drying olive-green adaxially,
paler abaxially, elliptic-oblong, oblanceolate, or oblong-lanceolate, 1025(30) 46(8) cm, base cuneate, apex obtuse to
broadly acuminate; secondary veins 9 or 10 pairs; stipule
broadly ligulate to ovate, 35 mm, obtuse to rounded with arista
ca. 3 mm. Inflorescences subsessile, congested-cymose to congested-corymbiform, ca. 17 9 cm; secondary axes 11.2 cm;
bracts oblong-lanceolate to lanceolate, 2.55 mm, acuminate;
pedicels 12 mm. Flowers sessile or pedicellate. Calyx with
hypanthium subglobose, 1.52 mm; limb lobed deeply; lobes
oblong-lanceolate to lanceolate, 45 1.52 mm, obtuse to
acute. Corolla white; tube 2025 mm, glabrous at throat; lobes
elliptic or elliptic-oblong, 56 2.54 mm, obtuse to rounded.
Drupe reddish yellow to red, subellipsoid and weakly compressed, ca. 11 9 mm. Fl. May, fr. Sep.
Thickets or shady broad-leaved forests at low elevations, or sometimes on open sandy lands. Guangxi, Hainan, Yunnan [Indochina, Philippines].

3. Ixora chinensis Lamarck, Encycl. 3: 344. 1789.

long chuan hua
Gaertnera hongkongensis Seemann; Ixora crocata
Lindley; I. stricta Roxburgh; I. stricta var. incarnata Bentham;
Pavetta kroneana Miquel; Sykesia hongkongensis (Seemann)
Kuntze; Tsiangia hongkongensis (Seemann) But, H. H. Hsue &
P. T. Li.
Shrubs, 0.82 m tall; branches glabrous. Leaves opposite,
sometimes apparently in whorls of 4 due to reduced stem internodes, sessile or petiolate; petiole to 5 mm, glabrous; blade
drying leathery, oblanceolate, oblong-oblanceolate, obovate,
elliptic-oblong, or lanceolate, 618 36 cm, glabrous on both
surfaces, base cuneate to shortly truncate or rounded, apex
obtuse or rounded to acute; secondary veins 79 pairs; stipules
persistent, united around stem to almost interpetiolar, triangular
to broadly triangular, 37 mm, glabrous to glabrescent, costate,
acute and with arista 210 mm. Inflorescence terminal, congested-cymose to congested-corymbiform, many flowered, puberulent to hirtellous, subsessile to pedunculate; peduncle to 1.5
cm, often subtended by 2 reduced leaves or leaflike bracts;
branched portion 14 15 cm (not including corollas); bracts
triangular, 0.21 mm; pedicels to 2 mm. Flowers subsessile to
pedicellate. Calyx glabrous; hypanthium obconic to ovoid, 1
1.5 mm; limb deeply lobed; lobes triangular to ligulate, 0.51
mm, acute or obtuse. Corolla red or reddish yellow, outside
glabrous; tube 2030 mm, glabrous in throat; lobes ovate, elliptic, or broadly elliptic, 57 45 mm, broadly obtuse to
rounded. Drupe reddish black, subglobose and shallowly didymous, 67 67 mm, glabrous. Fl. MayJul and Dec, fr. Sep
Oct.
Thickets, sparse forests; 200800 m. Fujian, Guangdong, Guangxi
[Indonesia, Malaysia, Philippines, Vietnam; widely cultivated in tropical
regions].

179

This is a commonly collected species of Ixora in China, apparently growing naturally as well as in cultivation. The occasional short
stem internodes, which sometimes produce congested groups of leaves,
appear to possibly be due to a change in growth pattern at the top of a
seasonal spurt that includes several internodes. The circumscription and
characters of this species were considered in some detail by Fosberg and
Sachet (Baileya 23(2): 77. 1989), who noted that it is sometimes
cultivated. Bridson (Kew Bull. 55: 10111012. 2000) studied the identity of Tsiangia, and formally synonymized its only species, T. hongkongensis, with I. chinensis.

4. Ixora effusa Chun & F. C. How ex W. C. Ko, Fl. Hainan. 3:

580. 1974.
san hua long chuan hua
Shrubs, ca. 1 m tall; branches glabrous. Leaves opposite;
petiole 0.51 cm, glabrous; blade drying thinly papery to papery and olive-greenish brown, oblanceolate to oblong, 1018
37 cm, both surfaces glabrous, base cuneate to obtuse, apex
acute or obtuse; secondary veins 8 or 9 pairs; stipules persistent,
shortly fused around stem to subinterpetiolar, triangular, 28
mm, glabrous, obtuse to acute with arista 29 mm. Inflorescence terminal, corymbiform, 711 cm wide, lax, with axes usually spreading at 4590, glabrous; peduncle 0.54 cm, articulate and with reduced leaves near base; bracts linear, narrowly
triangular, or ligulate, 0.55 mm; pedicels to 10 mm. Flowers
sessile to pedicellate. Calyx glabrous; hypanthium turbinate, ca.
1 mm; limb deeply lobed; lobes lanceolate to ligulate, 11.4
mm, acute to rounded. Corolla white or pale purple; tube 711
mm, glabrous at throat; lobes narrowly lanceolate, 59 ca. 2.3
mm, acute. Drupe subglobose, 89 mm in diam., glabrous. Fl.
AprMay.
Forests at middle elevations; ca. 500 m. Guangxi, Hainan [Vietnam].

5. Ixora finlaysoniana Wallich ex G. Don, Gen. Hist. 3: 572.

1834.
bao ye long chuan hua
Shrubs or small trees, to 56 m tall; branches glabrous.
Leaves opposite; petiole 410 mm, glabrous; blade drying rather leathery, elliptic-oblong, elliptic, oblanceolate, obovate, or
oblong-lanceolate, 1017(20) 36(9) cm, glabrous on both
surfaces, base cuneate to obtuse, apex obtuse to acute; secondary veins 812 pairs; stipules usually persistent, interpetiolar or
usually united around stem, broadly ovate to triangular, 36
mm, glabrous, acute or with arista 0.33.5 mm. Inflorescences
terminal, corymbiform to densely cymose, puberulent, subsessile to pedunculate; peduncle to 4.5 cm; branched portion 2.54
35 cm (not including corollas); bracts lanceolate, elliptic, or
narrowly ligulate, 810 mm, obtuse to acute. Flowers sessile or
subsessile. Calyx glabrous; hypanthium ellipsoid to subglobose,
11.5 mm; limb deeply lobed; lobes elliptic, narrowly elliptic,
or oblanceolate, 46 mm, acute. Corolla white, outside glabrous; tube 2030 mm, glabrous at throat; lobes narrowly elliptic to oblanceolate, 56 1.23 mm, acute to subacute. Drupe
subglobose and often weakly didymous, 68 mm in diam., glabrous. Fl. AprOct, fr. Sep.
Sparse forests at low elevations; 1001100 m. Guangdong, Hainan, Yunnan [NE India (Assam), Indochina, Philippines, Thailand;
widely cultivated in tropical zones worldwide].

RUBIACEAE

180

The name of this species is sometimes spelled incorrectly as

findlaysoniana. This species is native in our region and also widely
cultivated in warm tropical regions worldwide.

6. Ixora fulgens Roxburgh, Hort. Bengal. 10. 1814.

liang ye long chuan hua
Pavetta fulgens (Roxburgh) Miquel.
Shrubs; branches glabrous. Leaves opposite; petiole 1018
mm, glabrous; blade drying dark brown, shiny, oblong-lanceolate, lanceolate, elliptic, or narrowly elliptic, (9)1522 (2)
34 cm, both surfaces glabrous, base cuneate to obtuse, apex
acute to long acuminate; secondary veins 810 pairs; stipules
persistent, shortly united around stem, ovate, 23 mm, acute
with arista 35 mm. Inflorescences terminal, corymbose, 1217
cm wide, puberulent to hirtellous; peduncle ca. 1 cm; bracts
narrowly triangular, 11.5 mm; pedicels 0.53 mm. Flowers
pedicellate. Calyx glabrescent; hypanthium ellipsoid, ca. 1.5
mm; limb lobed for up to ca. 1/2; lobes subtriangular, 0.51
mm, obtuse. Corolla orange-yellow to dark red, outside glabrous; tube 2530 mm, in throat glabrous; lobes ovate, lanceolate, or elliptic, 1012 mm, acute. Drupe globose, 910 mm in
diam., glabrescent. Fl. JulSep.
On wet soil in sparse forests. Yunnan [India, Indonesia, Myanmar,
Philippines, Vietnam].
The application of this name has apparently varied among authors.
Bremekamp (J. Bot. (London) 75: 111. 1937) discussed the differing
usage, noting that the original description was based on a plant cultivated in the Calcutta Botanical Garden and of incorrectly attributed (in
his view) provenance. Plants from the Philippines have petioles as short
as 5 mm and corolla lobes up to 40 mm.
W. C. Ko (in FRPS 71(2): 42. 1999) described the branches as
sometimes with epidermis strips peeling off; this has not been reported
by others nor seen on specimens. Ko also described the secondary leaf
veins as 1825 pairs (p. 43); that count seems to include the often rather
well-developed intersecondary veins together with the secondary veins.

7. Ixora hainanensis Merrill, Lingnan Sci. J. 6: 287. 1930.

hai nan long chuan hua
Ixora pygmaea Merrill & F. P. Metcalf.
Shrubs, 0.23 m tall; branches glabrous. Leaves opposite;
petiole 28 mm, glabrous; blade drying papery to subleathery,
dry dark green, slightly shiny to matte, elliptic-oblong, elliptic,
or oblanceolate, 410(14) 15 cm, glabrous on both surfaces, base cuneate, rounded, or subcordate, apex rounded, obtuse, or bluntly acute; secondary veins 510 pairs; stipules persistent, shortly united around stem or subinterpetiolar, triangular
to ovate, 410 mm, glabrous, long acute with apical portion 3
10 mm, at least sometimes glandular at tip. Inflorescence terminal, corymbiform to cymose, many flowered, glabrous; peduncle 14 cm; branched portion 23 25 cm (not including corollas); bracts linear-oblong, 13.5 mm; pedicels 0.53 mm.
Flowers pedicellate. Calyx glabrous; hypanthium ellipsoid to
obconic, 11.5 mm; limb deeply lobed; lobes triangular to ovate
or lanceolate, 11.5 mm, acute to acuminate. Corolla white,
outside glabrous; tube 3040 mm, villous in throat; lobes narrowly spatulate to oblanceolate, 67 12 mm, acute. Drupe
red, subglobose to ellipsoid, weakly compressed and didymous,
ca. 6 68 mm, glabrous. Fl. MayDec, fr. Jan, OctDec.

On sandy soil at streamsides in dense forests; 1001100 m.

Guangdong, Hainan.
The name Ixora pygmaea was synonymized by W. C. Ko in FRPS
(71(2): 43. 1999); Merrill and Metcalf in fact suggested in the protologue that it might only be a depauperate form of I. hainanensis. The
leaves are rather variable in shape, usually on the same plant as noted in
the protologue.

8. Ixora hekouensis Tao Chen, nom. nov.

he kou long chuan hua
Replaced synonym: Ixora gracilis W. C. Ko, Guihaia 19:
103. 1999, not Ixora gracilis R. Brown ex Fawcett in H. O.
Forbes, Nat. Wand. E. Archip. 508. 1885, nor Ixora gracilis (A.
Richard ex Candolle) Kuntze, Revis. Gen. Pl. 1: 286287. 1891
[Pavetta gracilis A. Richard ex Candolle, Prodr. 4: 492. 1830].
Climbing shrubs. Stems glabrous. Leaves opposite; petiole
46 mm; blade drying papery and grayish brown, elliptic-oblong to obovate, 1018 37 cm, base cuneate to obtuse, apex
rather abruptly contracted and acute to acuminate; secondary
veins 79 pairs; stipules persistent, ovate, 67 mm, with arista
slightly longer than stipule. Inflorescences terminal, corymbose,
78 3.55 cm, several flowered, puberulent; peduncle 35
cm; bracts narrowly lanceolate, 3.55 mm; pedicels 23 mm.
Flowers pedicellate. Calyx puberulent; hypanthium subglobose,
ca. 2 mm; limb deeply lobed; lobes triangular, 11.2 mm. Corolla white; tube 1820 mm, glabrous at throat; lobes narrowly
elliptic-oblong, 56 mm, obtuse. Drupe unknown. Fl. JunAug.
Dense forests at riversides; ca. 200 m. Yunnan (Hekou).
The reported climbing habit of this species is highly unusual in
Ixora; however, W. C. Ko did not use that character in the protologue to
help distinguish this species.

9. Ixora henryi H. Lveill, Repert. Spec. Nov. Regni Veg.

13: 178. 1914.
bai hua long chuan hua
Shrubs, 13 m tall; branches glabrous. Leaves opposite;
petiole 37 mm, glabrous; blade drying papery, elliptic-oblong,
lanceolate, lanceolate-oblong, or elliptic, 515 1.54 cm, glabrous on both surfaces, base cuneate, obtuse, or rounded, apex
sharply acute to usually acuminate; secondary veins 7 or 8
pairs; stipules persistent or sometimes fragmenting, triangular
to ovate, 58 mm, glabrous, shortly obtuse to narrowed and
prolonged into arista 311 mm. Inflorescence terminal, corymbiform to congested-cymose, many flowered, glabrous; peduncle 0.41.5 cm; branched portion 1.54 27 cm (not including
corollas); bracts linear or linear-lanceolate, 0.83 mm; pedicels
to 2.5 mm. Flowers subsessile to pedicellate. Calyx glabrous;
hypanthium ellipsoid, 11.8 mm; limb deeply lobed; lobes triangular, 0.51 mm, acute. Corolla white sometimes becoming
dark red when dry, outside glabrous; tube 2025 mm, glabrous
in throat; lobes narrowly oblong to narrowly lanceolate, 56
1.62 mm, acute. Drupe subglobose, 810 mm in diam. Fl.
AprDec, fr. MayJul.
Broad-leaved forests, streamsides at forest margins; 2002000 m.
Guangdong, Guangxi, Guizhou, Hainan, Yunnan [Thailand, Vietnam].

RUBIACEAE

10. Ixora insignis Chun & F. C. How ex W. C. Ko, Guihaia 19:

97. 1999.
chang xu long chuan hua
Small trees, height not noted; branches glabrous. Leaves
opposite; petiole 710 mm; blade drying membranous, ellipticoblong to oblong-oblanceolate, 1014 45 cm, glabrous, base
cuneate, obtuse, or rounded, apex abruptly shortly acuminate;
secondary veins 1012 pairs; stipules triangular, 58 mm, with
arista almost as long as sheath. Inflorescence terminal, corymbiform, puberulent; peduncle ca. 14 cm, articulate below middle, at articulation with pair of reduced leaves (or leaflike
bracts) ovate to oblong-ovate, ca. 4 2.5 cm; branched portion
67 cm wide; secondary axes 1.51.8 cm; bracts linear-subulate, 0.53 mm; pedicels to 1.5 mm. Flowers sessile or pedicellate. Calyx puberulent; hypanthium obconic to ellipsoid, ca. 0.5
mm; limb deeply lobed; lobes narrowly elliptic-oblong, 1.31.5
mm, obtuse. Corolla in bud up to 2 cm with lobes up to 6 mm,
obtuse. Drupe not seen. Fl. May.
Mountains; ca. 1600 m. Yunnan (Hekou).
The reduced leaves borne on the peduncle that were mentioned
in the protologue to characterize this species are called by some authors
foliaceous bracts.

11. Ixora longshanensis Tao Chen, nom. nov.

long shan long chuan hua
Replaced synonym: Ixora amplexicaulis C. Y. Wu & W. C.
Ko, Guihaia 19: 98. 1999, not Ixora amplexicaulis Gillespie,
Bull. Bernice P. Bishop Mus. 74: 30. 1930.
Small trees, to 6 m tall; branches glabrous, drying brown.
Leaves opposite, sessile; blade drying thinly leathery or thickly
papery, brown, paler abaxially, elliptic to oblanceolate, (2)13
15 (1)56 cm, glabrous on both surfaces, base cordulate and
amplexicaul, apex acute to acuminate; secondary veins 1015
pairs; stipules subulate to triangular, ca. 7 mm, glabrous, acute
to acuminate. Inflorescence terminal, corymbiform, puberulent,
red to purple-red; peduncle 1.53.5 cm, often articulate near
base, at articulation bearing reduced leaves ca. 2 0.50.7 cm;
secondary axes 2.53 cm; bracts subulate, 1.52 mm. Flowers
subsessile to pedicellate. Calyx with hypanthium obconic, ca. 2
mm; limb deeply lobed; lobes triangular, obtuse. Corolla in bud
red-purple; tube 89 mm or longer, glabrous at throat; lobes
lanceolate, 45 mm or longer, cuneate. Drupe unknown.
Dense forests in valleys or at streamsides. S Yunnan.

12. Ixora nienkui Merrill & Chun, Sunyatsenia 2: 324. 1935.

pao ye long chuan hua
Shrubs, 13 m tall; branches glabrous. Leaves opposite;
petiole 1015 mm, glabrous; blade drying papery, oblong-lanceolate, elliptic-oblong, or oblong-oblanceolate, 1023 37
cm, both surfaces glabrous, sometimes shallowly bullate, base
rounded, obtuse, or cordulate, apex acuminate to rather long
acuminate; secondary veins 1015 pairs; stipules deciduous,
united around stem, 310 mm, glabrous, acute with arista 25
mm. Inflorescences terminal, corymbiform, many flowered, pu-

181

berulent, dark red, subsessile to shortly pedunculate; peduncle

to 0.6 cm; secondary axes 12.5 cm; bracts linear-lanceolate to
linear, 13 mm; pedicels 16 mm. Flowers pedicellate. Calyx
glabrous; hypanthium ellipsoid, tube 1.51.8 mm; limb lobed
nearly to base; lobes triangular, ca. 1 mm. Corolla white or
pink, outside glabrous; tube 3040 mm, glabrous at throat;
lobes lanceolate, 57 mm, acute. Drupe bright red, globose, 78
mm in diam., glabrous. Fl. JulOct, fr. JunOct.
Woods, forested ravines, streamsides; 4001000 m. Guangdong,
Guangxi, Hainan [Vietnam].

13. Ixora paraopaca W. C. Ko, Guihaia 19: 101. 1999 [paraopara].

ban na long chuan hua
Shrubs, to 2 m tall; branches glabrous, lenticellate, striate.
Leaves opposite; petiole 11.5 cm; blade drying thickly papery,
opaque, dark green, oblong-lanceolate, 1016 34.5 cm, base
obtuse, apex cuneate, acute, or acuminate; secondary veins 14
16 pairs; stipules triangular, ca. 10 mm, acute with arista ca. 3
mm. Inflorescence terminal, corymbiform-cymose, 89 910
cm, puberulent; peduncle 0.10.5 cm; bracts linear-lanceolate to
linear, 35 mm. Flowers sessile to subsessile. Calyx puberulent;
hypanthium turbinate, ca. 2 mm; limb deeply lobed; lobes
broadly triangular, ca. 1 mm, obtuse. Corolla red, outside glabrous; tube ca. 10 mm, glabrous inside; lobes narrowly oblong,
45 mm, obtuse to subrounded. Drupe unknown. Fl. AprJun.
Sparse forests on gentle hill slopes. Yunnan (Xishuangbanna).
This name was originally published with the spelling paraopara, a correctable typographical error as evidenced by the diagnosis,
which states that this new species is related to Ixora opaca R. Brown ex
G. Don. The protologue text described the corolla tube as 1 mm long,
but this seems to be shown as 1 cm in the protologue figure.

14. Ixora philippinensis Merrill, Philipp. J. Sci., C, 5: 238.

1910.
xiao xian long chuan hua
Shrubs or small trees; branches glabrous [to densely puberulent in Philippine plants]. Leaves opposite, sessile to
shortly petiolate; petiole to 0.4 cm, glabrous [to densely puberulent in Philippine plants]; blade drying thinly papery and
dark brown, elliptic, elliptic-oblong, or elliptic-ovate, 4.510
27 cm, glabrous [or puberulent along midrib abaxially in
Philippine plants], base truncate, rounded, or cordulate, apex
obtuse to rounded and usually apiculate; secondary veins 811
pairs; stipules deciduous, very shortly united around stem, lanceolate to broadly triangular, 25 mm, glabrous, acuminate. Inflorescences terminal, congested-cymose, 12 cm wide (not including corollas), few flowered, puberulent to glabrous; peduncle 11.5 cm, usually subtended by a pair of reduced, ovate or
subovate, cordate leaves 0.53 cm; bracts ovate-lanceolate to
narrowly triangular, 1.52 mm, acuminate. Flowers subsessile
to sessile. Calyx puberulent to glabrescent; hypanthium ellipsoid, ca. 1 mm; limb 0.51 mm, shallowly lobed; lobes dentiform. Corolla white to pink, outside puberulent; tube 1820
mm; lobes elliptic-oblong, ca. 7 3.5 mm, obtuse. Drupe
ovoid, didymous, weakly compressed, ca. 1 cm, red when dry.
Fl. JunAug.

RUBIACEAE

182

Broad-leaved forests. Taiwan [Philippines].

15. Ixora subsessilis Wallich ex G. Don, Gen. Hist. 3: 572.

1834.
nang guo long chuan hua
Shrubs or small trees, 22.5 m tall; branches glabrous.
Leaves opposite, subsessile; blade drying thinly leathery, lanceolate, elliptic, or oblanceolate, 1012 34 cm, glabrous on
both surfaces, base obtuse, apex long acuminate; stipules glabrous, long acuminate with apex subulate or linear, longer than
sheath. Inflorescences subsessile, corymbiform and trichotomous, several flowered; axes slender; bracts subulate or linear.
Calyx with hypanthium ca. 1 mm; lobes linear, 0.81.3 mm.
Corolla tube 1345 mm, glabrous in throat; lobes narrowly
elliptic-oblong, acute. Drupe saccate, 88.4 mm in diam.
Sparse forests; 12001500 m. Xizang (Mdog) [India, S Thailand].

Small shrubs, to 1 m tall; branches glabrous. Leaves opposite; petiole 11.8 cm, glabrous; blade olive-green when fresh,
drying papery and dark brown, oblong-lanceolate to elliptic, 7
20 2.56 cm, glabrous on both surfaces, shallowly bullate,
base cuneate to obtuse, apex acute to long acuminate; secondary veins 916 pairs; stipules deciduous through fragmentation,
interpetiolar or shortly united around stem, ovate to ligulatetriangular, 28 mm, glabrous, rounded with arista 26 mm. Inflorescences terminal, corymbiform, trichotomous, lax, sessile
to subsessile, sparsely to densely puberulent; branched portion
33.5 33.5 cm; bracts linear-lanceolate to narrowly triangular, 13 mm, acuminate; pedicels 35 mm. Flowers pedicellate. Calyx glabrous; hypanthium ellipsoid, ca. 1.5 mm; limb
deeply lobed; lobes ovate to triangular, ca. 1 mm, acute to acuminate. Corolla white, outside glabrous; tube 1518 mm, glabrous at throat; lobes elliptic, ca. 5 mm, obtuse to subrounded.
Drupe unknown. Fl. Sep.
Shady thickets. Guangxi (Shangsi).

W. C. Ko (in FRPS 71(2): 33. 1999) described this species as

highly variable, in particular in leaf shape, in the corolla tubes ranging
1345 mm, and in the stigma varying from bilobed to entire; this range
of variation is indeed rather unusual.

18. Ixora yunnanensis Hutchinson in Sargent, Pl. Wilson. 3:

412. 1916.

16. Ixora tibetana Bremekamp, J. Bot. 75: 261. 1937.

yun nan long chuan hua

xi zang long chuan hua

Shrubs, to 1 m tall, little branched; branches glabrous.

Leaves opposite or ternate; petiole 210 mm, glabrous; blade
drying papery, grayish green, narrowly oblanceolate or narrowly elliptic-oblong, 722 13(4) cm, glabrous on both
surfaces, base acute and decurrent, apex acute to long acuminate; secondary veins 1215 pairs; stipules persistent, shortly
united around stem, triangular to broadly ovate, 35 mm, glabrous, acute with arista 24.5 mm inserted below top of stipule,
sometimes shortly bilobed. Inflorescences terminal, subsessile,
corymbiform to congested-cymose, 24 2.54 cm (not including corollas), puberulent to glabrescent, subsessile to pedunculate; peduncle to 0.5 cm; bracts lanceolate to narrowly triangular, 26 mm, acute; pedicels to 2 mm. Flowers sessile to
shortly pedicellate. Calyx puberulent; hypanthium turbinate,
1.52 mm; limb deeply lobed; lobes linear-lanceolate, 34 mm,
subacute to acute. Corolla white; tube 3040 mm; lobes narrowly elliptic-oblong, 67 22.5 mm, obtuse. Drupe red,
ellipsoid-oblong, somewhat compressed and didymous, ca. 10
8 mm. Fl. May.

Shrubs or small trees; branches glabrous, 22.5 mm in

diam. Leaves opposite; petiole 0.650.8 cm, glabrous; blade
drying green or markedly discolorous, opaque, elliptic or obovate, 1517 6.57.5 cm, glabrous on both surfaces, base cuneate to acute, apex acuminate; secondary veins ca. 11 pairs; stipules shortly fused around stem, triangular, glabrescent except
adaxially villous, aristate. Inflorescence terminal, corymbiform,
tripartite, sparsely puberulent, 75100-flowered; peduncle 0.7
0.8 cm, with reduced leaves at base; secondary axes 4.55 cm;
bracts filiform, reduced; pedicels to 1 mm. Flowers sessile to
shortly pedicellate. Calyx glabrous; limb lobed to base; lobes
lanceolate, ca. 1.5 mm, subacute. Corolla white, glabrous
throughout; tube ca. 24 mm; lobes ovate-lanceolate, ca. 5.5
2.5 mm, acute. Drupe unknown.
12001500 m. SE Xizang.

17. Ixora tsangii Merrill ex H. L. Li, J. Arnold Arbor. 24: 456.

1943.
shang si long chuan hua

Dense forests at humid riversides. Yunnan.

40. KEENANIA J. D. Hooker, Fl. Brit. India 3: 101. 1880.

xi nan shu
Chen Tao (); Charlotte M. Taylor
Herbs or subshrubs, unarmed. Raphides apparently present. Leaves opposite, perhaps without domatia; stipules caducous or
usually persistent, interpetiolar, triangular, usually aristate, sometimes somewhat inflated. Inflorescences terminal or sometimes
pseudoaxillary, capitate, several to many flowered, pedunculate, bracteate with basalmost bracts usually involucrate and remaining
bracts often well developed and paired. Flowers sessile, presumably bisexual, biology unknown. Calyx limb 4-, 5-, or sometimes 6lobed with lobes sometimes unequal, sometimes imbricate in bud, sometimes gland-tipped. Corolla white or yellow, funnelform with
tube sometimes inflated, inside pubescent in upper part and throat; lobes 4, 5, or sometimes 6, ovate and acuminate, valvate in bud.
Stamens 5, inserted at corolla base, included or partially exserted; filaments short to developed; anthers dorsifixed near base. Ovary
2-celled, ovules numerous in each cell on globose, peltate, axile placentas; stigma 2-lobed, included or exserted. Fruit unknown.
About five species: Cambodia, S China, India, Laos, Myanmar, Thailand, Vietnam; two species (one endemic) in China.

RUBIACEAE

183

This genus is not well known or documented. Puff et al. (Rubiaceae of Thailand, 180. 2005) described the flowers as unisexual (?), though
they did not say whether the plants are dioecious, and suggested that the fruit of Keenania will eventually be found to be capsular. The absence of
raphides was implied by the placement of this genus in the Isertieae in FRPS (71(1): xiii. 1999), but raphides are clearly evident on specimens referred
to K. ophiorrhizoides Drake from Vietnam (MO!) and thus are provisionally cited for the genus.

1a. Peduncles less than 0.5 cm; involucral bracts narrowly lanceolate or linear, 1015 mm ................................................... 1. K. flava
1b. Peduncles 13.5 cm; involucral bracts orbicular, elliptic-oblong, or subovate, 67 mm ........................................ 2. K. tonkinensis
This name was previously published by H. S. Lo (loc. cit.) but not
validly so because no type was indicated (Vienna Code, Art. 37.1).

1. Keenania flava H. S. Lo, sp. nov.

huang xi nan
Type: China. Guangxi: Napo, Xiahua, Baikan, on slope,
21 Apr 1977, C. X. Luo 3-5491 (holotype, GXMI bar code
050450).
Validating Latin description: that of Koenania [sic!] flava
Lo (H. S. Lo, Bull. Bot. Res., Harbin 18: 282283. 1998).
Suffrutescent unbranched herbs, to 50 cm tall; stems terete, sparsely to densely villous. Petiole 0.82.5 cm, villous; leaf
blade drying papery, adaxially black, and abaxially pale, lanceolate to ovate, 512 2.54 cm, glabrous except pilose on
principal veins, base cuneate to obtuse, apex acuminate or
acute; secondary veins 59 pairs; stipules caducous, not seen.
Peduncle less than 0.5 cm; involucral bracts several, narrowly
lanceolate or linear, 1015 mm, acuminate; bracteoles narrowly
lanceolate or linear, 56 mm. Calyx with hypanthium portion
ca. 1 mm; lobes 5, narrowly lanceolate, 1.42 mm, slightly
unequal. Corolla in bud yellow, tubular, and with 5 sharp longitudinal ridges; tube ca. 3 mm; lobes 5, triangular, ca. 1 mm,
winged abaxially, apex rostrate-incurved. Fruit not seen. Fl.
Apr.
Valleys. Guangxi (Napo).

2. Keenania tonkinensis Drake, Bull. Mus. Hist. Nat. (Paris) 1:

118. 1895.
xi nan
Suffrutescent herbs, to 30 cm tall; stems villosulous. Petiole 11.5(4) cm, glabrescent; leaf blade drying thinly papery
and abaxially yellowed, elliptic-oblong or elliptic, 46(12)
22.5(5) cm, adaxially glabrous, abaxially glabrous or often
sparsely villosulous, base acuminate or cuneate, apex acute;
secondary veins 810 pairs; stipules triangular, ca. 8 mm, long
acuminate. Peduncle 13.5 cm; involucral bracts orbicular,
elliptic-oblong, or subovate, 67 mm. Calyx with hypanthium
portion broadly obconical, ca. 1 mm; lobes 5, triangular, 45
ca. 1.5 mm, veined. Corolla white; tube ca. 6 mm, slightly
swollen at base, slightly constricted in throat; lobes 5, ca. 1 mm.
Fruit not seen.
Dense forests in valleys. Guangxi (Longzhou) [N Vietnam].
This species name was published twice, first and validly in the
place cited above, then later by Drake (in J. Bot. (Morot) 9: 217. 1895);
this later work is sometimes incorrectly cited as the place of first publication.

41. KELLOGGIA Torrey ex Bentham & J. D. Hooker, Gen. Pl. 2: 137. 1873.
gou mao cao shu
Chen Tao (); Friedrich Ehrendorfer
Herbs, perennial, sometimes slightly woody at base; rootstock slender, with short rhizomatous and somewhat woody branches
and with ascending or erect stems. Raphides present. Leaves opposite, decussate, subsessile, without domatia; stipules persistent,
interpetiolar, hardly fused to petioles, triangular to linear or irregularly divided and often mutifid to fimbriate. Inflorescences
thyrsoid, with terminal and axillary branches at uppermost nodes, pedunculate cymes often with umbelliform flower groups, not
rarely with new axes developing and older axes elongating making inflorescences expansive, lax, few to many flowered and
bracteate; uppermost bracts often reduced to multifid or fimbriate stipules. Flowers pedicellate, bisexual, monomorphic. Calyx teeth
4 or 5, narrowly lanceolate, hardly fused at base. Corolla white to pink or red, funnelform, divided to ca. 1/2 into 4 or 5 lobes,
glabrous inside and valvate in bud. Stamens 4 or 5, inserted in corolla throat, finally slightly exserted; filaments flattened; anthers
dorsifixed near base. Ovary inferior, densely covered with hooked trichomes, 2-celled, ovules 1 in each cell, erect, basal; style with 2
short, linear stigmas, exserted. Fruit with calyx teeth persistent, schizocarpous, dividing into 2 oblong to ellipsoid, leathery and
indehiscent mericarps, densely covered with hooked trichomes; each mericarp with 1 medium-sized, ellipsoid and plano-convex
seed; endosperm fleshy; embryo large; cotyledons leaflike; radicle hypogeous.
Two species: disjunct, one in China and Bhutan, the other in W North America (Mexico, United States); one species in China.
In general aspect, Kelloggia resembles Galium, particularly with respect to the inferior ovary (hypanthium) developing into dry schizocarps
covered with hooked trichomes. Presumably, these fruit disperse similarly to those of Galium as stick-tights, by attaching to animals. In contrast to
Galium and other Rubiinae, Kelloggia has 3-colpate (and not polycolpate) pollen grains, calyx teeth, and not leaflike interpetiolar stipules, making
sterile plants resemble Nertera and Neanotis. In spite of these differences, Robbrecht and Manen (Syst. & Geogr. Pl. 76: 85146. 2006) have
transferred Kelloggia from the tribe Paederieae to the tribe Rubieae as a monotypic and basal subtribe Kelloggiinae. We concur with this transfer but
not with the inclusion of the totally different Theligonum into the Rubieae, which should be left in a separate tribe, Theligoneae (see also Bremer &
Eriksson, Int. J. Pl. Sci. 170: 766793. 2009). Kelloggia and all other Chinese Rubieae taxa have been briefly discussed and keyed out under Galium
in the present volume.

RUBIACEAE

184

All generic descriptions of Kelloggia give the number of calyx and corolla lobes as 4 or 5, implying that both conditions are equally common
as apparently is the case in the North American species. However, all the Asian specimens studied have 5 calyx and corolla lobes, as shown in the
FRPS illustration (71(2): 157, t. 41. 1999) and described by Springate et al. (Fl. Bhutan 2(2): 822. 1999).
The morphology, circumscription, biogeography, and molecular phylogeny of Kelloggia were studied by Nie et al. (Amer. J. Bot. 92: 642452.
2005). They concluded that the two species of the genus are most closely related to each other, that Kelloggia arrived in North America through longdistance dispersal from Asia, and that it occupies a basal position within Rubieae.

1. Kelloggia chinensis Franchet, J. Bot. (Morot) 6: 11. 1892.

yun nan gou mao cao
Galium aberrans W. W. Smith.
Herbs, perennial, to 30 cm tall. Stems flattened to subterete, puberulent to villosulous or hirtellous. Leaves subsessile or
with petiole to 1 mm; blade drying thinly papery and blackish,
narrowly lanceolate to narrowly elliptic, oblanceolate, or obovate, 515 25 mm, adaxially sparsely strigillose to villosulous, abaxially glabrous except sparsely to densely puberulent
to hirtellous along midrib, base cuneate to acute, apex obtuse to
acute; 1 main vein, secondary veins not or hardly visible;
stipules 1.54 mm, often irregularly 37-lobed, triangular to
linear, tomentulose or villosulous to glabrescent. Inflorescences

112 cm, axes strigillose to villosulous, 320-flowered; bracts

stipuliform, 12 mm; pedicels 23 mm. Calyx split essentially
to base, lobes 5, narrowly triangular, 0.51 mm. Corolla white
to pink or red, outside puberulent or hispidulous; tube 22.5
mm; lobes 5, lanceolate, 22.5 mm, acute. Ovary ellipsoid, 1
1.5 mm, densely hairy with hooked trichomes 0.51 mm. Mericarps ovoid, ca. 2.5 mm, densely covered with hooked trichomes of 0.51 mm. Fl. Jul, fr. JulSep.
Dry to wet mountain grasslands, along trails, forest and thicket
openings; 30003700 m. Sichuan, Xizang, Yunnan [Bhutan].
Springate et al. (Fl. Bhutan 2(2): 822. 1999) noted that this species
was collected once in Bhutan, at 3680 m, and that it is known from
Xizang also. We have seen no material or other citations from Xizang,
but this distribution seems reasonable and is added here.

42. KNOXIA Linnaeus, Sp. Pl. 1: 104. 1753.

hong ya da ji shu
Chen Tao (); Charlotte M. Taylor
Vissadali Adanson.
Annual or perennial herbs or subshrubs, unarmed. Raphides present. Leaves opposite or sometimes ternate, without domatia;
stipules persistent or deciduous, interpetiolar and fused to petioles and/or shortly united around stem, 26-lobed or -setose, with
apices of segments or setae usually glandular. Inflorescences terminal, thyrsiform, subcapitate, cymose, or often corymbiform with
axes sometimes elongating and becoming racemiform with age, several to many flowered, pedunculate or sessile, bracteate. Flowers
sessile or pedicellate, bisexual, distylous. Calyx limb 4-lobed; lobes sometimes unequal. Corolla white, pink, lilac, or violet, funnelform, salverform, or tubular, with shape sometimes differing between long-styled and short-styled forms, lanate inside tube; lobes 4,
valvate in bud. Stamens 4, inserted in corolla throat or near middle of corolla tube, included or exserted; filaments short; anthers
dorsifixed. Ovary 2-celled, ovules 1 in each cell, pendulous, apical; stigma 2-lobed, exserted or included. Fruit schizocarpous, ovoid
to ellipsoid, sometimes laterally compressed and/or didymous, dry, with calyx limb persistent; mericarps 2, ellipsoid, indehiscent,
with 1 seed, early to tardily separating from base upward and falling together with or separately from a carpophore, with carpophore
variously filiform and basal to comprising entire enlarged septum; seeds medium-sized, oblong-ellipsoid, compressed; testa thin;
endosperm fleshy; cotyledons thin; radicle ascending.
Seven to nine species: tropical Asia and Oceania; two species in China.
Knoxia was revised by Bhattacharjee and Deb (J. Econ. Taxon. Bot. 6(1): 7396. 1985), who recognized seven species. Then, it was reviewed
anecdotally by Puff and Robbrecht (Bot. Jahrb. Syst. 110: 511558. 1989), who circumscribed it differently and included nine species. The treatment
by W. C. Ko (in FRPS 71(2): 37. 1999) differed markedly from these others; it did not cite these authors so presumably Ko had not seen their works.
In particular, these other authors considered K. sumatrensis to be a wide-ranging, morphologically variable species and included as synonyms of var.
sumatrensis two species that were separated by Ko, K. corymbosa and K. mollis. Because it was based on geographically broader studies of more
literature and many more specimens from the type regions, Puff and Robbrechts taxonomy is used here.
W. C. Ko (loc. cit.: 4) described the fruit as capsules and the seeds as having a thick stipe, but the fruit of Knoxia have more often been described
within Rubiaceae as schizocarps with carpophores, as noted here.

1a. Herbs with rather large, fleshy, fusiform roots; leaves sessile or subsessile, blade lanceolate or oblong-lanceolate,
710 35 cm, with secondary veins 57 pairs; peduncles 312 cm; corolla tube 34 mm; fruit oblong-ellipsoid
and somewhat didymous ............................................................................................................................................. 1. K. roxburghii
1b. Herbs or subshrubs with slender fibrous roots; leaves subsessile to petiolate with petioles to 12 mm, blade
elliptic-oblong, elliptic, or lanceolate, 312 0.83.5 cm, with secondary veins 69 pairs; peduncles 0.32 cm;
corolla tube 1.52 mm; fruit ellipsoid ...................................................................................................................... 2. K. sumatrensis
1. Knoxia roxburghii (Sprengel) M. A. Rau, Bull. Bot. Surv.
India 10(Suppl. 2): 40. 1969.
hong da ji

Spermacoce roxburghii Sprengel, Syst. Veg. 1: 404. 1824;

Knoxia valerianoides Thorel ex Pitard.
Erect herbs, 3070 cm tall; root sometimes purple, rather

RUBIACEAE

enlarged, fleshy, fusiform; branches subquadrate, becoming

channeled, puberulent or villosulous becoming glabrescent.
Leaves sessile or subsessile, opposite or often ternate; blade
drying papery, lanceolate or oblong-lanceolate, 710 35 cm,
both surfaces glabrescent or usually hispidulous to pilosulous or
strigillose at least on principal veins, base acute to rounded,
apex obtuse to acuminate; secondary veins 57 pairs, usually
indistinct; stipules persistent, shortly fused to petioles or united
around stem, linear to narrowly triangular or sometimes shortly
to deeply 2- or 3-lobed, 810 mm, villosulous to glabrescent,
acute. Inflorescences congested-cymose to subcapitate, puberulent or villosulous to glabrescent, ebracteate; peduncles 312
cm; dense flower groups 11.5 12.5 cm, borne singly or 35
on cymose axes; pedicels 0.51 mm. Calyx glabrescent; ovary
portion ellipsoid, ca. 0.5 mm; limb deeply lobed; lobes triangular, 0.10.5 mm, often unequal on an individual flower.
Corolla red, purplish red, or white, salverform or tubular, externally densely villosulous or glabrous; tube 34 mm; lobes
triangular to ovate, ca. 1 mm. Schizocarps oblong-ellipsoid,
somewhat didymous, laterally somewhat flattened, ca. 1.5 mm;
mericarps separating from each other and carpophore; carpophore persistent, linear, ca. 1.3 mm.
Grasslands on mountain slopes. Fujian, Guangdong, Guangxi,
Hainan, Yunnan, Zhejiang [Cambodia, India, Myanmar, Nepal, Thailand].
Bhattacharjee and Deb (J. Econ. Taxon. Bot. 6(1): 85. 1985)
apparently included the Chinese plants within their circumscription of
Knoxia roxburghii var. brunonis (Wallich ex G. Don) R. Bhattacharjee
& Deb, which falls into K. roxburghii as circumscribed by Puff and
Robbrecht (Bot. Jahrb. Syst. 110: 511558. 1989). This name is based
on Spermacoce brunonis Wallich ex G. Don (Gen. Hist. 3: 621. 1834).
W. C. Ko (in FRPS 71(2): 4. 1999, as K. valerianoides) described the
corollas of this species as salverform and densely pubescent externally;
however, the corollas of Thai plants of this species are more tubular and
are glabrous externally.

185

2. Knoxia sumatrensis (Retzius) Candolle, Prodr. 4: 569. 1830.

hong ya da ji
Spermacoce sumatrensis Retzius, Observ. Bot. 4: 23. 1786;
Knoxia corymbosa Willdenow; K. mollis Wight & Arnott.
Herbs or subshrubs, 2090(100) cm tall; roots slender,
fibrous; branches terete to weakly quadrate, puberulent or villosulous to densely tomentulose. Leaves opposite, subsessile or
petiolate with petiole to 12 mm, puberulent or villosulous to
glabrescent; blade papery to membranous, elliptic, lanceolate,
or elliptic-oblong, 312 0.83.5 cm, both surfaces hispid-pilosulous to villosulous or sometimes glabrescent adaxially, base
acute to cuneate, apex acuminate to cuspidate; secondary veins
69 pairs; stipules persistent, shortly fused to petioles, deeply
35-lobed, 37 mm, villosulous to glabrescent, lobes sometimes setose. Inflorescences congested-cymose to corymbiform,
sometimes trichotomous, 16 cm, densely strigillose to villosulous, ebracteate; peduncle 0.32 cm; pedicels 0.11 mm. Calyx
puberulent or strigillose to glabrous; ovary portion ellipsoid,
0.50.8 mm; limb lobed essentially to base; lobes triangular,
0.10.3 mm, equal to subequal on an individual flower. Corolla
white or purplish red, funnelform, outside glabrous; tube 1.52
mm; lobes triangular, ca. 1 mm. Schizocarps ovoid to ellipsoid,
23 mm, weakly 48-ribbed, strigillose or puberulent to glabrescent; mericarps coherent at dispersal; carpophore persistent,
linear, ca. 2.5 mm. Fl. JulAug, fr. OctNov.
Thickets in open fields of low elevations. Fujian, Guangdong,
Guangxi, Guizhou, Hainan, Taiwan [India, Indonesia, Japan (Ryukyu
Islands), Malaysia, Myanmar, Nepal, New Guinea, Philippines, Thailand, Vietnam; Australia].
W. C. Ko (in FRPS 71(2): 5. 1999) reported several unusual features for Knoxia mollis in China: calyx lobes ca. 1 mm, corolla bright
green or rarely white, and corolla tube ca. 1 mm. These features have
not been confirmed on any specimens studied for this treatment nor
reported by other authors for Knoxia in SE Asia.

43. LASIANTHUS Jack, Trans. Linn. Soc. London 14: 125. 1823, nom. cons.,
not Adanson (1763).
cu ye mu shu
Zhu Hua (); Charlotte M. Taylor
Dasus Loureiro; Litosanthes Blume; Mephitidia Reinwardt ex Blume.
Erect subshrubs, shrubs, or rarely small trees, unarmed, with tissues sometimes fetid. Branches and branchlets terete, sometimes
compressed, rarely fistulous; lenticels inconspicuous or conspicuous. Raphides present. Leaves opposite, distichous, usually thinly
leathery or papery, base acute to rounded or cordate, apex acuminate, acute or cuspidate; midrib plane, depressed or slightly prominent adaxially, usually prominent abaxially; veins generally prominent abaxially, ascending at an angle of more than 45, curved to
margin or joining nerves above at margin; tertiary nervules parallel or reticulate; stipules caducous or usually persistent at least near
stem apex, interpetiolar, well developed or reduced, triangular, lanceolate, ovate, or oblong, acute or obtuse. Inflorescence axillary,
several flowered and glomerulate, capitate, cymose, or with flowers solitary, sessile or pedunculate, ebracteate or bracteate with
bracts persistent or not, well developed, sometimes fused. Flowers bisexual, small, sessile or pedicellate. Calyx with hypanthium
portion obovoid, ovoid, or campanulate; limb 36-dentate or lobed or rarely truncate. Corolla white, funnelform or salverform to
urceolate (Lasianthus biflorus), from several millimeters long up to 2.5 cm, glabrous or hairy outside, inside glabrous or usually
villous in throat; lobes 46, valvate or imbricate in bud. Stamens 46, inserted in corolla throat; filaments short; anthers linear or
oblong, dorsifixed, included or exserted. Style linear; stigma lobes 39, linear or lanceolate, included or exserted. Ovary 39-celled,
ovules 1 in each cell, basal, erect. Fruit blue or rarely white, black, or red, drupaceous, small, pulpy or fleshy, usually globose,
smooth or warty, rounded or ridged, with calyx limb persistent; pyrenes 39 (sometimes fewer than that developing), thick walled,

186

RUBIACEAE

smooth, warty or sulcate on abaxial face, usually triangular in transverse section, with preformed germination slits; seed black, with
abundant endosperm; embryo straight; cotyledon short, flattened; radicle long clavate.
About 184 species: 160 species in tropical Asia, ca. 20 in Africa, three in tropical America, and one in Australia; 33 species (seven endemic) in
China.
Lasianthus is commonly collected in China. This genus has been studied in detail by H. Zhu for both China (Acta Phytotax. Sin. 32: 4981.
1994; Syst. & Geogr. Pl. 72: 63110. 2002; Acta Bot. Yunnan. 30: 308314. 2008) and Thailand (Acta Phytotax. Sin. 39: 116150. 2001: 53 species
included). As Zhu detailed (loc. cit. 2002: 63), the characters that distinguish species of Lasianthus are mostly small and/or subtle, frequently
ephemeral, and often difficult to see; and, consequently, the taxonomy of this genus is complicated. Lasianthus can be confused with some species of
Damnacanthus, Diplospora, Prismatomeris, Saprosma, and Urophyllum, especially when collected only with young flower buds. Lasianthus (Chinese
species) is rather distinctive vegetatively in its combination of leaf blades that are completely glabrous adaxially, petioles that are usually densely
pubescent even when other parts of the plant are glabrescent or only sparsely pubescent, and small stipules that are usually persistent and also densely
pubescent.
The leaf venation of Lasianthus frequently has a characteristic regular pattern, comprising subparallel or exceptionally regularly oriented tertiary
veins (nervules). This is different, however, from the lineolate quaternary venation found in Antirhea, Timonius, and some other genera. A similar
arrangement is found in some species of Urophyllum. Cai et al. (Acta Bot. Yunnan. 29: 497512. 2007) studied leaf details of Lasianthus and
[broadly] related genera, and Cai et al. (J. Syst. Evol. 46(1): 6272. 2008) studied the pollen of a similar group.
The genus Litosanthes has been variously treated as a genus with one to several species, or included in Lasianthus. H. S. Lo (in FRPS 71(2):
106108. 1999) treated Litosanthes as a separate genus of one or two species, as did Puff et al. (Rubiaceae of Thailand, 102. 2005). However, H. Zhu
(loc. cit. 2002: 69) included it as a synonym of Lasianthus, as done here, based on morphological and molecular evidence.
Lasianthus verrucosus H. S. Lo (Bot. J. S. China 2: 2. 1993; type specimen: China. Hainan: Ledong, Q. Huang 820468, SCBI) was not seen.
From H. S. Los description and figure of this species, it has leaves with a looped venation, glabrous and fuscous when drying; inflorescence sessile or
subsessile; calyx limb with 4 small, broadly triangular lobes; and pyrenes 4, verrucose on the abaxial face. Many Lasianthus specimens from Hainan
were examined, but none was found matching the description of L. verrucosus. Consequently, this species is excluded from this account. From the
original description it appears to be most similar to Saprosma merrillii and thus would likely key out as that species.
The phylogenetic relationships among species of Lasianthus and related genera were studied based on molecular data by Xiao and Zhu (Bot.
Stud. (Taipei) 48: 227232. 2007). Their results supported the inclusion of Litosanthes biflora and Saprosma crassipes within Lasianthus. Litosanthes
is accordingly included here. However, S. crassipes has a 2-locular ovary and 2-pyrene drupes, which do not match the current circumscription of
Lasianthus. Saprosma merrillii seems similar to S. crassipes, and their systematic position needs further study.
Lasianthus cyanocarpus Jack was reported from China by the Flora of Japan. However, true L. cyanocarpus has a restricted distribution in S
Thailand and Malesia. This name is therefore misapplied in China and Japan, where the correct name for the species is L. hirsutus.
The following species was recorded from China but could not be treated here because no material was seen by the present authors: Lasianthus
areolatus Dunn (J. Bot. 47: 376. 1909), recorded from Guangdong by Merrill and Chun (Sunyatsenia 1(1): 49. 1930).

1a. Flowers in pedunculate congested cymes or subcapitate groups (though shortly pedunculate in L. chunii and
sometimes subsessile in L. japonicus).
2a. Peduncles slender, 0.53 cm.
3a. Leaves more than 3 cm; flowers 5-merous; pyrenes 5 ....................................................................................... 12. L. filipes
3b. Leaves less than 3 cm; flowers 4-merous; pyrenes 2 or 4 .................................................................................. 5. L. biflorus
2b. Peduncles short, 0.10.4 cm, or if more than 0.6 cm then robust.
4a. Peduncles very short, 12 mm.
5a. Branches and leaves abaxially appressed pubescent; leaves usually elliptic, acute or acuminate at apex,
with nervules conspicuously elevated abaxially; fruit with 5 or 6 longitudinally elevated angles or ribs,
strigillose ....................................................................................................................................................... 10. L. chunii
5b. Branches and leaves glabrous, or branches sparsely strigose when young and leaves abaxially strigose
or hirtellous; leaves usually lanceolate, long caudate at apex, with nervules slightly prominent
abaxially; fruit smooth, glabrous ............................................................................................................ 20. L. japonicus
4b. Peduncles conspicuous, 1.520 mm.
6a. Bracts numerous, linear, 612 mm ..................................................................................................... 26. L. rhinocerotis
6b. Bracts 2 and less than 1 cm, or reduced and apparently absent.
7a. Branches depressed pubescent or subglabrous; peduncles relatively robust, 520 mm; bracts
linear, 310 mm; calyx more than 5 mm with ovate-lanceolate lobes; corolla more than
1.5 cm ............................................................................................................................................... 4. L. biermannii
7b. Branches sparsely puberulent to glabrous; peduncles slender, 1.55 mm; bracts 2 mm or
shorter, usually inconspicuous; calyx less than 2.5 mm with triangular or subulate lobes;
corolla 8.5 mm or shorter ............................................................................................................... 24. L. micranthus
1b. Flowers solitary or in sessile fascicles or glomerules (though sometimes pedunculate in L. henryi).
8a. Bracts conspicuous.

RUBIACEAE

187

9a. Leaves slightly cordate to rounded and often oblique at base.

10a. Branches and leaves tomentose or setose; leaves 512 cm; stipules inconspicuous or to
ca. 3 mm ................................................................................................................................................ 1. L. attenuatus
10b. Branches and leaves glabrous, except very young branches subglabrous and leaves sparsely
hairy on nerves abaxially; leaves 1118 cm; stipules 56 mm ................................................................ 27. L. rigidus
9b. Leaves cuneate or subrounded and not oblique at base.
11a. Branches and leaves hirsute; bracts large, leaflike ................................................................................. 16. L. hirsutus
11b. Branches and leaves glabrous or villous or appressed pubescent; bracts not leaflike.
12a. Branches glabrous or thinly puberulent, leaves glabrous; bracts orbicular, leathery .................. 19. L. inodorus
12b. Branches and leaves abaxially tomentose, villous, or densely appressed pubescent; bracts
not orbicular, papery.
13a. Leaves usually no more than 10 cm; bracts small, subulate; fruit with 4 pyrenes ........... 28. L. schmidtii
13b. Leaves more than 12 cm; bracts ovate, lanceolate or linear; fruit with 5 or 6 pyrenes.
14a. Stipules not conspicuous; bracts linear to lanceolate, up to 2 cm; fruit
glabrous .................................................................................................................. 30. L. sikkimensis
14b. Stipules triangular, up to 8 mm; outer bracts ovate and inner ones lanceolate,
none more than 5 mm; fruit hairy.
15a. Pubescence appressed on branches, leaves abaxially, stipules, bracts,
and calyx, drupes subglabrous ................................................................... 9. L. chrysoneurus
15b. Tomentum dense and spreading on branches, leaves abaxially, stipules,
bracts, calyx, and drupes ................................................................................. 25. L. obscurus
8b. Bracts inconspicuous or absent.
16a. Calyx lobes linear, longer than 5 mm.
17a. Leaves 1116 3.55.5 cm; lateral veins more than 10 pairs; branches and nerves abaxially
densely yellow villous; calyx lobes equal in length and 914 mm .................................................... 7. L. chevalieri
17b. Leaves 711 2.53 cm; lateral veins 5 or 6 pairs; branches and nerves abaxially hirsute;
calyx lobes unequal in length, with longer ones up to 8 mm .................................................... 22. L. linearisepalus
16b. Calyx lobes not linear, no more than 5 mm.
18a. Calyx lobes longer than calyx tube or as long as tube.
19a. Leaves more than 12 cm.
20a. Branches, leaf nerves abaxially, calyx, corolla, and fruit strigillose to glabrescent;
calyx lobes linear-lanceolate ............................................................................................. 6. L. calycinus
20b. Branches, leaf nerves abaxially, and fruit tomentose with relatively long brown or
fuscous hairs, calyx and corolla densely brown tomentose; calyx lobes oblong ................ 33. L. wardii
19b. Leaves less than 12 cm.
21a. Branches and leaf nerves abaxially glabrous or sparsely strigose; fruit glabrous ............. 23. L. lucidus
21b. Branches and leaves abaxially very hairy; fruit pubescent.
22a. Calyx limb divided nearly to base, lobes lanceolate or oblong and much longer
than tube.
23a. Young branches and leaves abaxially appressed pubescent; calyx
lobes oblong to elliptic or oblanceolate, obtuse at apex ......................... 2. L. austrosinensis
23b. Branches and leaves abaxially densely spreading villous; calyx
lobes lanceolate, acuminate at apex .................................................................. 11. L. curtisii
22b. Calyx limb not divided to base, lobes linear-lanceolate, slightly longer
than or almost as long as tube.
24a. Branches and leaves abaxially densely spreading villous ........................ 14. L. formosensis
24b. Branches and leaves abaxially densely appressed pubescent ........................... 15. L. henryi
18b. Calyx lobes clearly shorter than tube.
25a. Calyx lobes or teeth small, ovate and reflexed; leaves usually with more than 10 pairs
of nerves, veinlets conspicuously reticulate ............................................................................... 8. L. chinensis
25b. Calyx lobes or teeth not ovate and not reflexed; leaves usually with less than 10 pairs
of nerves, veinlets parallel or subparallel or subreticulate.
26a. Calyx cupulate, with limb truncate or minutely dentate; fruit crowned by enlarged
calyx limb .................................................................................................................... 32. L. verticillatus
26b. Calyx with conspicuous lobes; fruit crowned by persistent calyx lobes but these
not enlarged.
27a. Stipules triangular-lanceolate to linear-lanceolate, equal to or more than
3 mm, strigillose or hirsute; young branches strigillose or hirsute.
28a. Stipules 35 mm; flowers 5-merous; fruit smooth, pyrenes 5 ......................... 29. L. simizui

RUBIACEAE

188

28b. Stipules usually more than 5 mm; flowers 4-merous; fruit

verrucose, pyrenes 4 .............................................................................. 31. L. trichophlebus
27b. Stipules triangular, less than 3 mm, puberulent; young branches puberulent
or pubescent.
29a. Leaves more than 12 cm, lanceolate or oblong, lateral veins 69 pairs.
30a. Leaves oblong, lateral veins 68 pairs, veinlets subparallel, conspicuously
elevated abaxially; branches densely appressed pubescent, leaves
abaxially at nerves and margins puberulent
or ciliate .................................................................................................. 18. L. hookeri
30b. Leaves lanceolate to elliptic-lanceolate, lateral veins ca. 9 pairs,
veinlets subreticulate with forks very slender; branches and leaf
nerves abaxially sparsely appressed puberulent ............................... 21. L. lancifolius
29b. Leaves less than 11 cm, ovate or ovate-oblong, lateral veins 46 pairs.
31a. Leaves 46 cm, less than 3 cm wide; branches and leaf
nerves abaxially densely appressed hirsute ............................ 3. L. austroyunnanensis
31b. Leaves 610 cm, as narrow as 2 cm but usually more than
3 cm wide; branches and leaf nerves abaxially glabrous
or pubescent or villous.
32a. Branches and leaf nerves abaxially glabrous or sparsely
appressed pubescent; calyx sparsely pubescent, with 5
triangular teeth; fruit glabrous .......................................................... 13. L. fordii
32b. Branches and leaves abaxially densely villous; calyx
densely strigillose, with 5 minute and broadly triangular
teeth; fruit strigillose ................................................................ 17. L. hispidulus
1. Lasianthus attenuatus Jack, Trans. Linn. Soc. London 14:
126. 1823.
xie ji cu ye mu
Lasianthus densifolius Miquel; L. setosus Craib; L. wallichii Wight; L. wallichii var. hispidocostatus H. Zhu.
Shrubs, 12 m tall; branches and branchlets densely tomentose to hirsute or sometimes setose. Petiole 13 mm, tomentose to hirsute; leaf blade leathery or subleathery, oblong,
elliptic-lanceolate, or oblong-ovate, 512 2.55 cm, glabrous
adaxially, strigillose to hirtellous or tomentose abaxially, base
slightly cordate or rarely rounded, slightly to markedly oblique,
apex acute to cuspidate-acuminate; lateral veins 68 pairs; nervules subparallel, conspicuous abaxially; stipules triangular or
ovate-lanceolate, 23 mm, tomentose to hirsute, apparently persistent. Inflorescence cymose or glomerulate, sessile, densely
hirsute to hirtellous; bracts subulate to lanceolate, 615 mm,
persistent. Flowers sessile. Calyx densely hirsute; hypanthium
portion campanulate, ca. 1 mm; limb 12 mm, with 5 triangular
teeth. Corolla sparsely hirtellous, puberulent to glabrescent outside, villous inside; tube 810 mm; lobes 5, spatulate, 45 mm.
Fruit globose or ovoid, 510 mm in diam., densely hirsute,
smooth; pyrenes 5 or 6. Fl. Apr, fr. AugSep.
Forests, shaded and wet places; 2001800 m. Fujian, Guangdong,
Guangxi, Hainan, Taiwan, Yunnan [Bhutan, Cambodia, NE India, Indonesia, S Japan, Laos, Malaysia, Myanmar, Nepal, Papua New Guinea,
Philippines, Thailand, Vietnam].

2. Lasianthus austrosinensis H. S. Lo, Bot. J. S. China 2: 4.

1993.
hua nan cu ye mu
Shrubs, 12 m tall; branches and branchlets densely
appressed strigillose. Petiole 38 mm, densely strigillose; leaf

blade papery, ovate, 58 2.53 cm, glabrous adaxially, appressed strigillose on midrib and nerves abaxially, base obtuse
or subrounded, apex shortly cuspidate; lateral veins ca. 5 pairs;
nervules subparallel; nerves and nervules elevated abaxially;
stipules generally persistent, triangular to broadly triangular, 1
1.5 mm, densely strigillose. Inflorescences glomerulate to subcapitate, sessile; bracts absent. Flowers sessile or subsessile.
Calyx densely strigillose; hypanthium portion obconic, ca. 1
mm; limb deeply lobed; lobes 5, oblong to elliptic or oblanceolate, 33.5 mm, nerved, obtuse. Corolla ca. 5 mm, densely hirsute outside, villous in upper half inside; lobes 5, triangular.
Fruit subglobose, 45 mm in diam., strigillose; pyrenes 5.
Forests, shaded and wet places; 300500 m. Guangdong, Guangxi, Hainan.
H. Zhu (Syst. & Geogr. Pl. 72: 85. 2002) noted that the relatively
well-developed calyx lobes with evident venation are distinctive for this
species; the calyx lobes are also distinctive in being widest near the
middle rather than at the base as in most other Lasianthus species.

3. Lasianthus austroyunnanensis H. Zhu, Syst. & Geogr. Pl.

72: 96. 2002.
dian nan cu ye mu
Shrubs, to 1 m tall; branchlets densely appressed hirsute.
Petiole 58 mm, hirsute; leaf blade papery to membranous,
ovate-elliptic, 46 23 cm, glabrous adaxially, appressed hirsute on midrib and nerves abaxially, base acute or obtuse, apex
cuspidate-acuminate or acuminate; lateral veins 4 or 5 pairs;
nervules subparallel; nerves and nervules elevated conspicuously abaxially; stipules ?persistent, triangular, minute, hirsute.
Inflorescences glomerulate, sessile; bracts minute, hirsute.
Flowers sessile. Calyx hirsute; hypanthium portion campanulate, ca. 1 mm; limb 11.5 mm, toothed; teeth 5. Corolla 810
mm, pubescent outside, villous inside. Fruit globose, 45 mm in
diam., subglabrous; pyrenes 5.

RUBIACEAE

Forests, shaded and wet places; 10001300 m. Taiwan, Yunnan.

This species has been often misidentified as Lasianthus microphyllus Elmer. After examining type specimens of L. microphyllus from
the Philippines (Elmer 13790, isotypes in A, CAS, E, K, L, MO), it is
clear that the Chinese populations differ from L. microphyllus by having
appressed-hirsute hairs on the branches, leaf midrib and nerves abaxially, and the outside of the calyx and corolla. The leaves are ovate-elliptic with a cuspidate-acuminate or acuminate apex. In contrast, L. microphyllus has subglabrous or glabrous branches, leaves, and flowers, and
the leaves are ovate with a caudate apex. Therefore, the Chinese taxon
represents a separate species. The name L. microphyllus has also been
sometimes misapplied in Taiwan to plants of L. micranthus.

4. Lasianthus biermannii King ex J. D. Hooker, Fl. Brit. India

3: 190. 1880.
geng hua cu ye mu
Shrubs, 25 m tall; branchlets sparsely pubescent to
densely appressed pubescent to strigose, or subglabrous. Petiole
714 mm, pubescent to strigose; leaf blade membranous or
papery, elliptic, elliptic-ovate, or elliptic-lanceolate, 820 35
cm, glabrous adaxially, sparsely pubescent to densely appressed
pubescent on midrib, lateral veins, and nervules abaxially, base
acute to obtuse or rounded, apex cuspidate-acuminate; lateral
veins 57 pairs; nervules reticulate, slender and inconspicuous
to parallel and conspicuously elevated abaxially; stipules triangular or lanceolate, 35 mm, pubescent to strigillose, apparently caducous. Inflorescences congested-cymose to subcapitate, densely strigillose, pedunculate; peduncles slender to
robust, 515 mm; bracts linear, 210 mm. Flowers sessile or
subsessile. Calyx purple when fresh, strigillose; hypanthium
portion turbinate, 11.5 mm; limb 45 mm; lobes 46, narrowly lanceolate to oblanceolate, 24 mm. Corolla light purple
when fresh, 78 mm, villous inside; tube 35 mm, glabrous
outside; lobes 46, oblong-ovate, 34 mm, pilosulous outside.
Fruit globose, 0.31.5 cm in diam., smooth or with ca. 5 ridges;
pyrenes 5. Fl. Apr, OctNov, fr. Jul.
Montane forests, shaded and wet places; 10002500 m. Guizhou,
Hainan, Xizang, Yunnan [Bhutan, NE India, Myanmar].
The well-developed calyx lobes that are usually broadest near the
apex are distinctive.

1a. Branches usually sparsely pubescent or

subglabrous and leaf nerves abaxially
sparsely pubescent; nervules reticulate,
slender and inconspicuous; calyx
lobes relatively narrow and short;
peduncles slender, longer or shorter
than petioles ..................................... 4a. subsp. biermannii
1b. Branches and leaf nerves abaxially
densely appressed strigose; leaf
nervules parallel and conspicuously
elevated abaxially; calyx lobes
relatively wide and long; peduncles
robust and usually longer than
petioles ................................ 4b. subsp. crassipedunculatus
4a. Lasianthus biermannii subsp. biermannii
() geng hua cu ye mu (yuan ya zhong)

189

Branches sparsely pubescent or subglabrous. Leaf blade

with nerves abaxially sparsely pubescent; nervules reticulate,
slender and inconspicuous. Peduncles slender, longer or shorter
than petioles. Calyx lobes relatively narrow and short.
Montane forests, shaded and wet places; 12002500 m. Yunnan
[Bhutan, NE India, Myanmar].

4b. Lasianthus biermannii subsp. crassipedunculatus C. Y.

Wu & H. Zhu, Acta Phytotax. Sin. 32: 75. 1994.
cu geng cu ye mu
Lasianthus esquirolii H. Lveill.
Branches densely appressed strigose. Leaf blade with
nerves densely appressed strigose abaxially; nervules parallel
and conspicuously elevated abaxially. Peduncles robust and usually longer than petioles. Calyx lobes relatively wide and long.
Forests, shaded and wet places; 10001700 m. Guizhou, Hainan, Yunnan.
Rehder (J. Arnold Arbor. 16: 323. 1935) considered that Henry
11148 from Yunnan province of China, cited by Hutchinson as Lasianthus biermannii, matched the type specimen of L. esquirolii (J.
Esquirol 648) from Guizhou province of China, so he reduced L.
esquirolii to a synonym of L. biermannii. However, Ferguson (Notes
Roy. Bot. Gard. Edinburgh 32(1): 109. 1972) contended that the type of
L. esquirolii did not sufficiently match the type of L. biermannii and
again recognized L. esquirolii as a separate species. Many specimens
from SE Yunnan and Hainan are closely related to L. biermannii but
clearly differ from it and have been treated as a geographic subspecies
of L. biermannii. When the type of L. esquirolii was checked, it was
concluded that L. esquirolii is the same as L. biermannii subsp. crassipedunculatus.

5. Lasianthus biflorus (Blume) M. Gangopadhyay & Chakrabarty, J. Econ. Taxon. Bot. 16: 338. 1992.
shi he mu
Litosanthes biflora Blume, Catalogus, 22. 1823; Lasianthus gracilis King & Gamble.
Shrubs, 12.5 m tall; branchlets terete, densely hirsute to
pilosulous. Petiole 0.52 mm, hirsute to pilosulous; leaf blade
thinly to thickly papery, subrhombic or elliptic-obovate, elliptic,
or subovate, 1.53 cm 715 mm, glabrous adaxially, hirtellous to pilosulous abaxially, base cuneate, margin flat, often undulate, apex acute, often mucronulate; midrib depressed adaxially, prominent abaxially, hirsute or pilosulous abaxially; lateral veins plane adaxially, prominent abaxially, 69 pairs, hirsute or pilosulous abaxially, extending to unite with margin;
nervules obscure on both surfaces; stipules inconspicuous,
narrowly triangular, 0.52 mm, densely pilosulous to hirtellous,
at apex acute sometimes with 24 very small lateral projections
or lobes. Inflorescence congested-cymose or with solitary flowers, pedunculate; peduncles 0.51.7 cm, sparsely hirsute to pilosulous; bracts inconspicuous, lanceolate to linear, 0.31.2
mm. Flowers pedicellate; pedicels 0.21 mm. Calyx glabrous to
sparsely hirtellous; hypanthium portion campanulate, 1.21.5
mm; limb deeply 5-toothed; teeth triangular, ca. 0.5 mm. Corolla 24 mm, glabrous outside, villous in throat and on inside
of lobes; lobes ovate, shorter than tube, rostriform-incurved at

RUBIACEAE

190

apex. Fruit ovoid or depressed globose, 35 mm in diam., glabrous, verrucose, 4-grooved; pyrenes 4.
Forests, shaded and wet places; 600700 m. Hainan, Taiwan, S
Yunnan [Indonesia, Malaysia, Philippines, Thailand, Vietnam].

6. Lasianthus calycinus Dunn, Bull. Misc. Inform. Kew 1912:

367. 1912.
huang guo cu ye mu
Shrubs, 1.53 m tall; branchlets appressed pubescent or
strigillose to glabrescent. Petiole 810 mm, appressed pubescent or strigillose to glabrescent; leaf blade thickly papery,
oblong-elliptic, 1315 46 cm, glabrous adaxially, sparsely
appressed pubescent to strigillose on nerves abaxially, base
broadly cuneate or subrounded, apex acute to shortly acuminate; lateral veins 79 pairs, extending to unite with margin;
nervules parallel; nerves and nervules conspicuous abaxially;
stipules generally persistent, broadly triangular, ca. 2 mm,
strigillose, acute. Inflorescences glomerulate to subcapitate, sessile, 46-flowered; bracts absent. Flowers subsessile. Calyx
strigillose; hypanthium portion 1.52 mm; limb deeply lobed;
lobes 5, linear-lanceolate to narrowly triangular, 2.55 mm,
acute. Corolla 79 mm, strigillose outside; lobes 5, ovate. Fruit
orange or red-orange, subglobose, 67 mm in diam., sparsely
strigillose to glabrescent, with longitudinal elevated angles. Fl.
AprMay, fr. SepOct, Dec.
Forests, shaded and wet places; 600700 m. Hainan.
H. Zhu (Syst. & Geogr. Pl. 72: 83. 2002) noted that this species is
similar to and may even be conspecific with the poorly known species
Lasianthus latifolius (Blume) Miquel of Borneo, Java, and Sumatra.

7. Lasianthus chevalieri Pitard in Lecomte, Fl. Indo-Chine 3:

384. 1924.
chang e cu ye mu
Lasianthus longisepalus Geddes; L. longisepalus var. jianfengensis H. S. Lo.
Shrubs, to 2 m tall; branches and branchlets densely yellow villous. Petiole 11.5 cm, long, yellow villous; leaf blade
subleathery, elliptic-oblong, 1116 3.55.5 cm, glabrous
adaxially, yellow villous abaxially, base cuneate, apex acuminate; lateral veins ca. 11 pairs, prominent abaxially; nervules
parallel, prominent abaxially; stipules generally persistent, narrowly triangular, ca. 3 mm, spreading yellow villous. Inflorescences glomerulate, sessile; bracts inconspicuous. Flowers sessile. Calyx densely pilose; hypanthium portion obconic, 1.53
mm; limb with tubular portion 1.53 mm; lobes 57, linear, 9
14 mm. Corolla tube 1213 mm; lobes 57, lanceolate, 78
mm, densely pilose. Fruit subglobose, 67 mm in diam., pilose;
pyrenes 57.
Forests, shaded and wet places; 8001500 m. Hainan [Thailand,
Vietnam].
The sessile flowers with relatively long calyx lobes are quite distinctive.

8. Lasianthus chinensis (Champion ex Bentham) Bentham, Fl.

Hongk. 160. 1861.

cu ye mu
Mephitidia chinensis Champion ex Bentham, Hookers J.
Bot. Kew Gard. Misc. 4: 196. 1852; Lasianthus dinhensis
Pierre ex Pitard; M. odajimae Masamune.
Shrubs, 14 m tall; branches and branchlets terete, densely
pubescent or puberulent to strigillose. Petiole 0.51.5 cm,
densely pubescent or puberulent to strigillose; leaf blade subleathery, oblong to elliptic, 1125 2.57 cm, glabrous adaxially, thinly to densely pubescent or puberulent to strigillose
abaxially, base acute or obtuse, margin plane or usually reflexed, apex acute or acuminate; lateral veins 915 pairs; nervules reticulate, slightly elevated abaxially; stipules generally
persistent, triangular to narrowly triangular, 1.53 mm, densely
pubescent or puberulent to strigillose. Inflorescences subcapitate to congested-cymose, sessile to subsessile; bracts reduced.
Flowers sessile or subsessile. Calyx puberulent to strigillose;
hypanthium portion globose to ellipsoid, 1.53 mm; limb 1.5
2.5 mm, lobed for 1/22/3; lobes 46, triangular, ovate, or orbicular, becoming reflexed. Corolla 1015 mm, densely strigillose to sericeous outside; lobes 5 or 6, lanceolate. Fruit globose
to depressed globose, 58 mm in diam., pubescent to strigillose,
with 5 or 6 conspicuous longitudinal angles; pyrenes 5 or 6. Fl.
MayJun, fr. SepOct.
Forests, shaded and wet places; below 100900 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan [Cambodia, Laos, Malaysia, Philippines, Thailand, Vietnam].
The relatively large flowers with the large globose to ellipsoid
hypanthium (i.e., ovary) portion is distinctive; Lasianthus verticillatus is
similar in these characters but can be distinguished by its truncate to
denticulate calyx limb and smooth fruit. The calyx limb often enlarges
as the fruit develops.

9. Lasianthus chrysoneurus (Korthals) Miquel, Fl. Ned. Ind.

2: 322. 1857.
ku zi cu ye mu
Mephitidia chrysoneura Korthals, Ned. Kruidk. Arch.
2(2): 222. 1851; Lasianthus hoaensis Pierre ex Pitard; L. kurzii
J. D. Hooker; L. kurzii var. howii H. S. Lo.
Shrubs, to 3 m tall; branchlets appressed pubescent to
densely velutinous. Petiole 510 mm, appressed pubescent to
densely velutinous; leaf blade membranous, elliptic-oblong to
elliptic-lanceolate, 1220 35 cm, glabrous adaxially, appressed pubescent to puberulent abaxially, base cuneate to
acute, apex acuminate; lateral veins 710 pairs; nervules parallel, inconspicuous; stipules deciduous, triangular or oblonglanceolate, 38 mm, pubescent to densely velutinous, strigillose
on margin. Inflorescences glomerulate to subcapitate, sessile;
bracts persistent, outer bracts ovate, inner bracts lanceolate, all
15 mm, pubescent to densely strigillose, especially on margin.
Flowers sessile or subsessile. Calyx densely strigillose; hypanthium portion campanulate, 11.5 mm; limb deeply lobed;
lobes 46, triangular, 1.52 mm. Corolla 56 mm, puberulent
outside on upper part, pubescent inside in throat. Fruit blueblack, subglobose, 46 mm in diam., pubescent to strigillose or
puberulent; pyrenes 5 or 6.
Forests, shaded and wet places; 5001200 m. S Yunnan [Cam-

RUBIACEAE

bodia, India, Indonesia (Java), Laos, Myanmar, Papua New Guinea,

Thailand, Vietnam].
H. Zhu (Syst. & Geogr. Pl. 72: 80. 2002) noted that this species is
very similar to Lasianthus obscurus.

10. Lasianthus chunii H. S. Lo, Bot. J. S. China 2: 10. 1993.

huan yong cu ye mu
Shrubs, 13 m tall; branchlets dark brown, densely hirsute
or strigose. Petiole 0.51 cm, densely hirsute to strigillose; leaf
blade subleathery or thickly papery, lanceolate or oblong-lanceolate, 815 25.5 cm, glabrous adaxially, hirsute or strigillose on midrib, nerves, and nervules abaxially, base cuneate or
obtuse to rounded, apex acuminate; lateral veins 7 or 8 pairs;
nervules subparallel; veins and nervules conspicuously elevated
abaxially; stipules 11.5 mm, densely hirsute to strigillose, apparently caducous. Inflorescence congested-cymose, shortly pedunculate, 24-flowered, densely strigillose; peduncle 12 mm;
bracts narrowly triangular, 12 mm. Flowers subsessile to pedicellate; pedicels to 1 mm. Calyx strigillose; hypanthium portion
campanulate, 11.5 mm; limb ca. 2 mm; lobes 6, subtriangular,
ca. 1 mm. Corolla ca. 1 cm, hirsute or strigillose outside, villous
in upper half inside; lobes 6, lanceolate. Fruit depressed globose, ca. 5 mm in diam., hirsute or strigillose, with 5 or 6 longitudinal elevated angles or ridges; pyrenes 5 or 6. Fl. Apr, fr.
JunJul, Sep.
Forests, shaded and wet places. SE Fujian, Guangdong, SE
Guangxi, S Jiangxi.

11. Lasianthus curtisii King & Gamble, J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist. 73: 128. 1904.
guang dong cu ye mu
Lasianthus condorensis Pierre ex Pitard; L. formosensis
Matsumura var. hirsutus Matsumura; L. kwangtungensis Merrill.
Shrubs, 12 m tall; branches and branchlets densely
spreading villous. Petiole 48 mm, densely villous; leaf blade
papery or subleathery, ovate-lanceolate or elliptic-lanceolate, 5
9 23 cm, glabrous adaxially, densely soft villous abaxially,
base cuneate or obtuse, apex cuspidate-acuminate; lateral veins
46 pairs; nervules subparallel; stipules generally persistent,
narrowly triangular, 11.5 mm, densely strigose. Inflorescences
glomerulate to subcapitate, sessile; bracts absent. Flowers sessile to subsessile. Calyx densely villous to hirsute; hypanthium
portion obconic, 11.5 mm; limb divided nearly to base; lobes
5, lanceolate to narrowly triangular, 25 mm, acuminate. Corolla 78 mm, outside hispid, inside villous; lobes 5. Fruit
ovoid-globose, 45 mm in diam., hirsute to villous; pyrenes 5.
Fl. AugSep, fr. JunDec.
Forests, shaded and wet places; 300900 m. Fujian, Guangdong,
Guangxi, Hainan, Taiwan [Indonesia (Sumatra), Japan (Kyushu, Ryukyu Islands), Malaysia, Thailand, Vietnam].

12. Lasianthus filipes Chun ex H. S. Lo, Bot. J. S. China 2: 8.

1993.
chang geng cu ye mu
Shrubs, 12 m tall; branchlets slender, densely strigose.
Petiole 15 mm, densely hirsute or strigillose; leaf blade pa-

191

pery, ovate or ovate-oblong, 58 23 cm, glabrous adaxially,

appressed pubescent to strigillose on midrib, veins, and nervules abaxially, base obtuse, rounded, or rarely subcordate,
apex shortly acuminate or cuspidate-acuminate; lateral veins 4
or 5 pairs; nervules subparallel, conspicuous abaxially; stipules
minute, densely appressed hirsute or strigose, apparently caducous. Inflorescence congested-cymose or subcapitate, pedunculate, few flowered; peduncles slender, 13 cm, densely hirsute
or strigose; bracts subulate, 12.5 mm, hirsute or strigose.
Flowers subsessile to shortly pedicellate. Calyx hirsute or
strigillose; hypanthium portion campanulate, ca. 1 mm; limb
1.52 mm; lobes 5, subulate, 11.5 mm. Corolla ca. 5 mm; tube
puberulent outside, densely villous in upper half inside; lobes 5,
ovate, ca. 1.5 mm, villous inside. Fruit globose, 78 mm in
diam., subglabrous; pyrenes 5. Fl. Apr.
Forests, shaded and wet places; 5001500 m. Fujian, Guangdong,
Guangxi, Hainan, Yunnan [N Vietnam].

13. Lasianthus fordii Hance, J. Bot. 23: 324. 1885.

luo fu cu ye mu
Lasianthus kamputensis Pierre ex Pitard; L. tashiroi Matsumura; L. zambalensis Elmer.
Shrubs, 12 m tall; branches and branchlets glabrous to
thinly appressed pubescent or strigose. Petiole 0.41 cm, subglabrous to hirsute or strigillose; leaf blade subleathery or papery, ovate-oblong, 610 25 cm, glabrous adaxially, thinly
appressed strigillose or subglabrous abaxially, base acute or obtuse, apex acuminate or cuspidate-acuminate; lateral veins 46
pairs; nervules subparallel, elevated abaxially; stipules generally persistent, triangular to narrowly triangular, 0.51.2 mm,
strigillose. Inflorescences glomerulate, sessile; bracts absent.
Flowers sessile. Calyx sparsely to densely strigillose; hypanthium portion campanulate to obconic, ca. 1 mm; limb 11.5
mm, shallowly toothed; teeth 5, triangular. Corolla 610 mm,
glabrous to puberulent outside, villous inside; lobes 5. Fruit globose or subglobose, 56 mm in diam., glabrous to strigillose;
pyrenes 5 or 6.
Forests, shaded and wet places; 2001000 m. Fujian, Guangdong,
Guangxi, Hainan, Taiwan, Yunnan [Cambodia, Indonesia (Java), Japan
(Kyushu, Ryukyu Islands, Yakushima), Papua New Guinea, Philippines,
Thailand, Vietnam].
Lasianthus fordii var. trichocladus H. S. Lo (Bot. J. S. China 2: 7.
1993) was described from SE China (Guangdong, Guangxi, and Hainan) but could not be treated here because no material was seen by the
present authors.

14. Lasianthus formosensis Matsumura, Bot. Mag. (Tokyo)

15: 17. 1901.
tai wan cu ye mu
Lasianthus kwangsiensis Merrill ex H. L. Li; L. tashiroi
Matsumura var. pubescens Matsumura; L. tenuicaudatus
Merrill.
Shrubs, 12 m tall; branches and branchlets densely villous or villosulous. Petiole 58 mm, villous to villosulous; leaf
blade papery or subleathery, oblong or ovate-elliptic, 712
2.55 cm, glabrous adaxially, villous or villosulous at least on

RUBIACEAE

192

veins abaxially, base acute or obtuse, apex acuminate or cuspidate-acuminate; lateral veins 6 or 7 pairs; nervules subparallel,
slightly elevated abaxially; stipules generally persistent, triangular, ca. 2 mm, villous to villosulous. Inflorescences glomerulate to congested-cymose, sessile to shortly pedunculate; bracts
absent. Flowers sessile. Calyx pilosulous to villosulous; hypanthium obconic to campanulate, ca. 1 mm; limb 34 mm, deeply
lobed; lobes 5, linear-lanceolate. Corolla 68 mm, villous outside, tomentose inside; lobes 5, oblong-lanceolate. Fruit ovoidglobose, ca. 5 mm in diam., subglabrous; pyrenes 5. Fl. Oct
Dec, fr. Apr.
Forests, shaded and wet places; 5001000 m. Guangdong, Guangxi, Hainan, Taiwan, Yunnan [Japan (Kyushu, Ryukyu Islands, Yakushima), Thailand, Vietnam].

15. Lasianthus henryi Hutchinson in Sargent, Pl. Wilson. 3:

401. 1916.
xi nan cu ye mu
Lasianthus appressihirtus Simizu; L. appressihirtus var.
maximus Simizu ex T. S. Liu & J. M. Chao; L. inconspicuus J.
D. Hooker var. hirtus Hutchinson.
Shrubs, 11.5 m tall; branches and branchlets densely
appressed pubescent or strigillose. Petiole 38 mm, appressed
strigillose; leaf blade subleathery or papery, oblong-lanceolate,
612 1.53.5 cm, glabrous adaxially, appressed pubescent
abaxially on nerves or strigillose on veins, base acute or cuneate, apex cuspidate-acuminate; lateral veins 7 or 8 pairs; nervules subparallel; nerves and nervules conspicuously elevated
abaxially; stipules generally persistent, triangular, 0.51.2 mm,
densely strigillose. Inflorescences glomerulate to subcapitate,
sessile or subsessile or very shortly pedunculate; bracts minute,
strigillose. Flowers sessile. Calyx densely strigillose; hypanthium portion campanulate to obconic, ca. 1 mm; limb 11.5 mm,
lobed for 1/33/4; lobes 5, linear-lanceolate. Corolla ca. 8 mm,
glabrous or strigillose on lobes outside, villous inside; lobes 5.
Fruit globose, ca. 5 mm in diam., strigillose; pyrenes 5. Fl. Jun,
fr. JulOct.
Forests, shaded and wet places; 2001900 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, E Xizang, Yunnan.

16. Lasianthus hirsutus (Roxburgh) Merrill, J. Arnold Arbor.

33: 229. 1952.
ji shi shu
Triosteum hirsutum Roxburgh, Fl. Ind. 2: 180. 1824.
Shrubs, 12 m tall; branchlets stiffly spreading rusty hirsute to glabrescent. Petiole 1015 mm, hirsute; leaf blade subleathery or leathery, oblong or oblanceolate-oblong, 1325 5
7.5 cm, glabrous adaxially, densely hirtellous or hirsute to strigose abaxially, base acute to obtuse, apex shortly acuminate;
lateral veins 710 pairs; nervules subparallel, elevated abaxially; stipules lanceolate to deltoid, 810 mm, hirsute, generally
persistent at least on apical nodes. Inflorescences glomerulate,
sessile, hirsute; bracts leaflike, numerous, persistent, outer ones
ovate-lanceolate, 23 cm, with distinct nerves, inner ones linear,
much smaller. Flowers sessile. Calyx densely hirsute; hypanthium portion obconical, ca. 1 mm; limb 5-lobed; lobes linear-

lanceolate, ca. 2.5 mm. Corolla 1115 mm; tube 812 mm, outside glabrous in lower half and hirsute in upper half, glabrous
inside; lobes 5, ovate, ca. 3 mm. Fruit ovoid-globose, 45 mm
in diam., hirsute to glabrescent, with ca. 5 rounded ridges; pyrenes 5. Fr. Nov.
Forests, shaded and wet places; 1001500 m. Guangdong, Guangxi, Hainan, Taiwan [Bangladesh, India (including Andaman and Nicobar Islands), Indonesia, Japan (Ryukyu Islands), Malaysia, Myanmar,
Papua New Guinea, Philippines, Thailand, Vietnam].
Lasianthus hirsutus is a widely distributed species. It was mistakenly called L. cyanocarpus by most authors until Merrill (loc. cit.)
showed the correct application of the name. Lasianthus cyanocarpus
has relatively smaller leaves with cordate and slightly oblique bases and
usually fewer and relatively smaller bracts; L. hirsutus differs from it in
relatively larger leaves with cuneate, slightly unequal bases, relatively
longer petioles, conspicuous triangular stipules, more numerous bracts
with the outer ones ovate-oblong and leaflike and the inner ones gradually narrower and smaller, as well as usually long rusty hirsute hairs on
most parts of the plants.

17. Lasianthus hispidulus (Drake) Pitard in Lecomte, Fl. IndoChine 3: 393. 1924.
wen shan cu ye mu
Mephitidia hispidula Drake, J. Bot. (Morot) 9: 239. 1895;
Lasianthus brevidens Craib; L. bunzanensis Simizu.
Shrubs, 11.5 m tall; branches and branchlets densely
villous or strigose. Petiole 47 mm, densely hirsute or strigose
to strigillose; leaf blade thinly leathery to papery, elliptic or
ovate-oblong, 710 35 cm, glabrous adaxially, densely villous abaxially, base acute or obtuse, apex cuspidate-acuminate
or acuminate; lateral veins 46 pairs; nervules subparallel;
midrib and veins elevated abaxially, nervules slightly elevated
abaxially, strigose to strigillose; stipules generally persistent,
0.51 mm, densely hirsute or strigillose. Inflorescences glomerulate, sessile; bracts absent. Flowers sessile. Calyx strigillose;
hypanthium campanulate, 11.5 mm; limb 11.5 mm, shortly
toothed; teeth 5, broadly triangular. Corolla 67 mm, pilose outside, villous inside; lobes 5. Fruit globose, 36 mm in diam.,
strigillose; pyrenes 5.
Forests, shaded and wet places; 300600 m. Guangdong, Guangxi, Hainan, Taiwan, Yunnan [Indonesia (Sumatra), Japan (Ryukyu
Islands), Malaysia (Borneo), Thailand, Vietnam].

18. Lasianthus hookeri C. B. Clarke ex J. D. Hooker, Fl. Brit.

India 3: 184. 1880.
hu ke cu ye mu
Shrubs, up to 5 m tall; branches and branchlets densely
appressed pubescent to sericeous. Petiole 510 mm, appressed
puberulent to strigose or sericeous; leaf blade papery or subleathery, oblong, 1224 36 cm, glabrous adaxially, puberulent or ciliate on nerves and margins abaxially, base obtuse or
cuneate, apex caudate-acuminate; lateral veins 68 pairs, outermost pair extending to tip of leaf; nervules subparallel, conspicuously elevated abaxially; stipules generally persistent, triangular to broadly triangular, 23 mm, appressed puberulent to
strigose or sericeous. Inflorescences glomerulate, sessile; bracts
absent or few, linear, 34 mm. Flowers sessile. Calyx subgla-

RUBIACEAE

brous to puberulent; hypanthium campanulate, 1.52 mm; limb

lobed for 1/23/4; lobes 46, lanceolate to narrowly triangular,
12 mm. Corolla 67 mm, puberulent outside; tube 45 mm;
lobes 46, ovate-lanceolate, ca. 2 mm. Fruit subglobose, 45
mm in diam., glabrous or puberulent, smooth; pyrenes 5.
Forests, shaded and wet places; 3001500 m. Guangxi, Guizhou,
Xizang, Yunnan [NE India, Myanmar, Thailand, Vietnam].

1a. Leaf margins densely ciliate; bracts

subulate; calyx limb plus hypanthium
ca. 2 mm, with minute subulate teeth
.............................................................. 18a. var. dunnianus
1b. Leaf margins sparsely ciliate; bracts
absent; calyx limb plus hypanthium
ca. 3 mm, with lanceolate, ca. 1 mm
lobes ......................................................... 18b. var. hookeri
18a. Lasianthus hookeri var. dunnianus (H. Lveill) H. Zhu,
Acta Phytotax. Sin. 32: 71. 1994 [dunniana].
jie mao hu ke cu ye mu
Lasianthus dunnianus H. Lveill, Repert. Spec. Nov.
Regni Veg. 11: 64. 1912 [dunniana].

193

20. Lasianthus japonicus Miquel, Ann. Mus. Bot. LugdunoBatavi 3: 110. 1867.
ri ben cu ye mu
Shrubs, 12 m tall; branches and branchlets glabrous or
subglabrous to sparsely strigose on young branches. Petiole 3
10 mm, sparsely strigillose to strigose or subglabrous; leaf
blade leathery or papery, lanceolate, lanceolate-oblong, or oblong, 915 23.5 cm, glabrous adaxially, glabrous to strigose
or hirtellous abaxially, especially on lateral veins and nervules,
base acute to obtuse, apex cuspidate or cuspidate-acuminate to
long caudate; lateral veins 57 pairs or numerous; nervules
reticulate; stipules generally persistent, triangular, 13 mm,
glabrous to minutely hirsute to strigillose. Inflorescences congested-cymose, pedunculate to subsessile, densely strigillose;
peduncle 12 mm; bracts 0.21 mm. Flowers sessile or subsessile. Calyx strigillose to glabrescent; hypanthium portion
campanulate, 11.5 mm; limb 12 mm, 4- or 5-dentate; teeth
short to well developed, linear. Corolla 1314 mm at anthesis,
glabrous outside or sometimes puberulent on lobes, villous inside; tube 910 mm; lobes 4 or 5, spatulate, 44.5 mm. Fruit
globose, 56 mm in diam., glabrous; pyrenes 4 or 5. Fl. Apr
May, fr. JunOct.

Leaf blade with margin densely ciliate. Bracts subulate.

Calyx limb plus hypanthium ca. 2 mm, with minute subulate
teeth.

Forests, shaded and wet places; 2002300 m. Anhui, Fujian,

Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [NE India, S Japan (Kyushu, Ryukyu
Islands, Shikoku), Laos, N Vietnam].

Forests, shaded and wet places; 3001500 m. Guangxi, Guizhou,

Yunnan [N Myanmar].

This species is commonly collected in China. The leaf margin is

often finely irregular, not crisped but actually undulate parallel to the
looping tertiary venation that is near it. The leaves generally flush in a
characteristic fashion, elongating but remaining rolled up and narrow,
and are covered with notable sericeous pubescence on the exposed
lower surfaces of the midrib and principal veins. This pubescence is
often deciduous, leaving the mature lower leaf surface glabrous.

18b. Lasianthus hookeri var. hookeri

() hu ke cu ye mu (yuan bian zhong)
Leaf blade with margin sparsely ciliate. Bracts absent. Calyx limb plus hypanthium ca. 3 mm; lobes lanceolate, ca. 1 mm.
Forests, shaded and wet places; 6001500 m. Xizang, Yunnan
[NE India, Myanmar, Thailand, Vietnam].

19. Lasianthus inodorus Blume, Bijdr. 998. 18261827.

ge ye cu ye mu
Lasianthus poilanei Pitard; L. tubiferus J. D. Hooker.
Shrubs, 23 m tall; branches and branchlets glabrous or
thinly puberulent. Petiole 610 mm, glabrous or sparsely puberulent; leaf blade leathery, elliptic to elliptic-lanceolate, 10
20 2.56 cm, glabrous adaxially, glabrous or sparsely puberulent abaxially, base acuminate, apex acuminate; lateral veins 6
or 7 pairs; nervules parallel; nerves and nervules conspicuous
abaxially; stipules persistent, triangular or ovate-lanceolate, 35
mm, glabrous, leathery. Inflorescences sessile, glomerulate;
bracts persistent, orbicular or ovate-orbicular, 36 mm, thickly
leathery, glabrous or pubescent. Flowers sessile. Calyx puberulent; hypanthium portion ca. 1 mm; limb 5-lobed; lobes ovatelanceolate. Corolla ca. 10 mm, puberulent outside, villous inside. Fruit red, ovoid, ca. 10 mm, glabrous; pyrenes 5.
Montane forests, shaded and wet places; 10001800 m. S Yunnan
[E Bangladesh, Cambodia, NE India, Indonesia (Java, Sumatra), Thailand, Vietnam].

Lasianthus japonicus is a widely distributed species from E Asia

to the Himalaya. Lasianthus japonicus subsp. japonicus occurs in SE
China to Japan, below 1800 m, and basically in the Sino-Japanese floristic region delineated by C. Y. Wu (Bull. Univ. Mus. Univ. Tokyo 37:
19. 1998). Lasianthus japonicus subsp. longicaudus occurs in SW
China and NE India in the Himalaya, at 10002300 m, i.e., in the SinoHimalayan floristic region of Wu. They are a typical vicariant pair of
subspecies. It may be surprising that L. lucidus var. caudisepalus from
Assam, NE India, is found to be conspecific with L. japonicus subsp.
japonicus. However, the geological history of E Asia gives an explanation. The formation of the vicarious distribution patterns of the SinoHimalayan and Sino-Japanese regions is supposed to be related to the
uplift of Himalaya in the Tertiary. Lasianthus japonicus subsp. longicaudus could have differentiated with the uplift of Himalaya becoming
an altitudinal vicariant taxon of L. japonicus.

1a. Leaves usually long caudate at apex,

lateral veins numerous, slender,
very similar to nervules or
subparallel secondary nerves;
calyx and corolla 4-merous;
calyx teeth reduced, limb almost
truncate at apex ............................ 20b. subsp. longicaudus
1b. Leaves cuspidate or cuspidateacuminate at apex, lateral veins
57 pairs, ascending at an
oblique angle, conspicuously

RUBIACEAE

194

different from reticulate nervules;

calyx and corolla 4-merous or
usually 5-merous; calyx with
well-developed triangular or
lanceolate teeth ................................. 20a. subsp. japonicus
20a. Lasianthus japonicus subsp. japonicus
() ri ben cu ye mu (yuan ya zhong)
Lasianthus caudatifolius Merrill; L. hartii Franchet; L.
japonicus var. satsumensis (Matsumura) Makino; L. lancilimbus Merrill; L. lucidus Blume var. caudisepalus Deb & M.
Gangopadhyay; L. satsumensis Matsumura.
Leaf blade cuspidate or cuspidate-acuminate at apex; lateral veins 57 pairs, ascending at an oblique angle, conspicuously different from reticulate nervules. Calyx 4- or usually 5lobed, with triangular or lanceolate teeth. Corolla 4- or usually
5-lobed.
Forests, shaded and wet places; 2001800 m. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [NE India, S Japan (Kyushu, Ryukyu Islands,
Shikoku)].
Two varieties are recognized within Lasianthus japonicus subsp.
japonicus; their morphological characters and geographic distribution
are presented below.
Lasianthus japonicus var. japonicus: Leaf blade lanceolate, less
than 3.5 cm wide. Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Zhejiang [NE India, S Japan
(Kyushu, Ryukyu Islands, Shikoku)].
Lasianthus japonicus var. latifolius H. Zhu (Acta Phytotax. Sin.
32: 79. 1994; kuan ye ri ben cu ye mu): Leaf blade
oblong-lanceolate, more than 3.5 cm wide. Guangxi, Guizhou, Sichuan,
Yunnan.

20b. Lasianthus japonicus subsp. longicaudus (J. D. Hooker)

C. Y. Wu & H. Zhu, Acta Phytotax. Sin. 32: 80. 1994.
yun guang cu ye mu
Lasianthus longicaudus J. D. Hooker, Fl. Brit. India 3:
190. 1880 [longicauda]; L. pseudojaponicus Masamune.
Leaf blade usually long caudate at apex; lateral veins numerous, slender, very similar to nervules or subparallel secondary nerves. Calyx 4-lobed, almost truncate at apex. Corolla 4lobed.
Montane forests, shaded and wet places; 10002300 m. Guangxi,
Guizhou, Sichuan, Xizang, Yunnan [NE India, Laos, N Vietnam].

21. Lasianthus lancifolius J. D. Hooker, Fl. Brit. India 3: 187.

1880.
mei mai cu ye mu
Shrubs, 1.53 m tall; branches and branchlets puberulent.
Petiole 510 mm, appressed pubescent; leaf blade papery, lanceolate to elliptic-lanceolate, 1320 2.54 cm, glabrous adaxially, puberulent on nerves and nervules as well as margin abaxially, base acute to obtuse, apex long acuminate; lateral veins
ca. 9 pairs, faint adaxially; nervules subreticulate, forked, very
slender; stipules generally persistent, triangular, 23 mm, pu-

bescent. Inflorescences glomerulate, sessile; bracts absent.

Flowers sessile. Calyx puberulent; hypanthium portion campanulate, 11.5 mm; limb 11.5 mm; teeth subulate. Corolla
1213 mm, puberulent outside, pubescent inside at upper half;
lobes 5, ovate, ca. 2 mm. Fruit globose, 45 mm in diam., glabrous or sparsely puberulent, smooth; pyrenes 5.
Forests, shaded and wet places; 5001700 m. Guangdong, Guangxi, Hainan, Yunnan [Bangladesh, Bhutan, NE India, NE Thailand, N
Vietnam].

22. Lasianthus linearisepalus C. Y. Wu & H. Zhu, Acta Phytotax. Sin. 32: 61. 1994.
xian e cu ye mu
Shrubs, ca. 1 m tall; branchlets hirsute to subglabrous.
Petiole ca. 5 mm, sparsely hirsute; leaf blade papery, oblong, 7
11 2.53 cm, glabrous adaxially, hirsute on nerves abaxially,
base unequally cuneate, apex caudate; lateral veins 5 or 6 pairs;
nervules subparallel; nerves and nervules elevated conspicuously abaxially; stipules ca. 1 mm, hirsute. Inflorescences
glomerulate, sessile, 13-flowered; bracts subulate, 23 mm,
hirsute. Flowers sessile. Calyx hirsute; lobes 5 or 6, linear,
unequal, longest to 8 mm. Corolla strigose-villous outside. Fruit
globose, sparsely hirsute; pyrenes 5.
Montane forests, shaded and wet places; 18002100 m. SW
Yunnan.

23. Lasianthus lucidus Blume, Bijdr. 997. 18261827.

wu bao cu ye mu
Shrubs, 11.5 m tall; branches and branchlets glabrous or
sparsely strigose on very young parts. Petiole 2.58 mm, strigillose or usually strigose; leaf blade membranous or subleathery,
ovate, elliptic, or ovate-elliptic, 49 1.53 cm, glabrous adaxially, sparsely strigose or hirsute on nerves and nervules abaxially, base cuneate to obtuse, apex acuminate or cuspidateacuminate; lateral veins 36 pairs, elevated abaxially; nervules
parallel, conspicuous abaxially; stipules generally persistent, triangular, 11.5 mm, strigose. Inflorescences glomerulate to subcapitate, sessile; bracts absent. Flowers sessile or subsessile.
Calyx with hypanthium portion obconic to widely campanulate,
11.5 mm, glabrescent; limb deeply lobed; lobes 5, narrowly
triangular, 23 mm, usually somewhat unequal on an individual
flower, strigose. Corolla 812 mm, glabrous or strigillose outside, villous at throat and on lobes inside; lobes 5, triangular, 2
2.5 mm. Fruit globose or ovoid, 45 mm in diam., glabrous;
pyrenes 5.
Montane forests, shaded and wet places; 9002400 m. Hainan,
Yunnan [Bangladesh, NE India, Indonesia, Myanmar, Philippines, Thailand, Vietnam].

1a. Leaves elliptic, with lateral veins

5 or 6 pairs ....................................... 23a. var. inconspicuus
1b. Leaves ovate or ovate-elliptic, with
lateral veins 3 or 4 pairs ........................... 23b. var. lucidus
23a. Lasianthus lucidus var. inconspicuus (J. D. Hooker) H.
Zhu, Acta Bot. Yunnan. 20: 154. 1998.
tuo yuan ye wu bao cu ye mu

RUBIACEAE

Lasianthus inconspicuus J. D. Hooker, Fl. Brit. India 3:

187. 1880.
Leaves elliptic; lateral veins 5 or 6 pairs.
Montane forests, shaded and wet places; 9001800 m. Yunnan
[Bangladesh, NE India, N Thailand].

23b. Lasianthus lucidus var. lucidus

() wu bao cu ye mu (yuan bian zhong)
Leaves ovate or ovate-elliptic; lateral veins 3 or 4 pairs.
Montane forests, shaded and wet places; 12002400 m. Hainan,
Yunnan [NE India, Indonesia, Myanmar, Philippines, Thailand, Vietnam].

24. Lasianthus micranthus J. D. Hooker, Fl. Brit. India 3: 190.

1880.
xiao hua cu ye mu
Lasianthus balansae (Drake) Pitard; L. microstachys Hayata; Mephitidia balansae Drake.
Shrubs, 12 m tall; branches and branchlets slender,
sparsely puberulent to glabrous, smooth. Petiole 49 mm, hirsute or strigillose; leaf blade papery or subleathery, ovate,
ovate-oblong, or oblong-lanceolate, 510 2.53.5 cm, glabrous adaxially, sparsely to densely hirsute or strigillose to strigose on midrib and nerves abaxially, base obtuse or rounded to
subacute, apex acute or acuminate; lateral veins 5 or 6 pairs;
nervules parallel or subparallel; stipules triangular or lanceolate,
11.5 mm, densely strigillose. Inflorescences congested-cymose to capitate, pedunculate, densely strigillose; peduncles
1.55 mm, slender; bract reduced or linear and up to 2 mm;
bracteoles minute. Flowers sessile. Calyx densely strigillose or
villous to glabrescent; hypanthium portion campanulate, ca. 1
mm; limb 11.5 mm; teeth 5, subulate or triangular. Corolla 5
6.5 mm, glabrous to puberulent outside, villous inside; tube 36
mm; lobes 5, spatulate, 22.5 mm. Fruit globose to depressed
globose, ca. 5 mm in diam., glabrous, smooth or with ca. 5 low
ridges; pyrenes 5. Fl. AugNov, fr. SepOct.
Forests, shaded and wet places; 1001800 m. Fujian, Guangdong,
Guangxi, Hainan, Taiwan, Xizang, Yunnan, Zhejiang [NE India, Thailand, N Vietnam].
The glabrous stems with strigillose rings at the nodes and the narrowly triangular, rather well-developed calyx lobes are distinctive.

25. Lasianthus obscurus (Blume ex Candolle) Miquel, Fl.

Ned. Ind. 2: 317. 1857.
lin sheng cu ye mu
Mephitidia obscura Blume ex Candolle, Prodr. 4: 453.
1830; Lasianthus kurzii J. D. Hooker var. fulvus C. Y. Wu & H.
Zhu; L. kurzii var. sylvicola H. S. Lo.
Shrubs, to 3 m tall; branchlets spreading tomentose or
densely pilosulous. Petiole 510 mm, spreading tomentose or
pilosulous to villous; leaf blade membranous, elliptic-oblong to
oblong, 1220 35 cm, glabrous adaxially, spreading tomentose or villous abaxially, base cuneate to acute, apex acuminate;
lateral veins 710 pairs; nervules parallel, distinct on both sur-

195

faces; stipules generally persistent, triangular or oblong-lanceolate, 48 mm, tomentose or pilosulous to strigillose. Inflorescences glomerulate or subcapitate, sessile; bracts persistent,
outer bracts ovate, inner bracts lanceolate, all 15 mm, tomentose or pilosulous to villous. Flowers sessile or subsessile. Calyx densely tomentose or strigose; hypanthium portion campanulate, 11.5 mm; limb deeply lobed; lobes 46, triangular, ca. 2
mm. Corolla 56 mm, puberulent outside on upper part, pubescent inside at throat. Fruit subglobose, 46 mm in diam., tomentose; pyrenes 5 or 6.
Forests, shaded and wet places; 3001200 m. Hainan, Yunnan [India (S Andaman Islands), Indonesia (Java, Sumatra), Myanmar, Thailand, Vietnam].
H. Zhu (Syst. & Geogr. Pl. 72: 80. 2002) noted that this species is
very similar to Lasianthus chrysoneurus.

26. Lasianthus rhinocerotis Blume, Bijdr. 996. 18261827

[rhinozerotis].
huang mao cu ye mu
Shrubs, 13 m tall; branches and branchlets densely villous. Petiole 510 mm, densely brown villous; leaf blade leathery, oblong-lanceolate or lanceolate, 1521 47 cm, glabrous
adaxially, densely villous abaxially, base broadly cuneate to
truncate, subrounded, rounded, or slightly cordate, apex caudate-acuminate; lateral veins 712 pairs; nervules parallel to
subreticulate; veins and nervules elevated conspicuously abaxially; stipules generally persistent, triangular, 2.54 mm. Inflorescences congested-cymose to subcapitate, shortly pedunculate, densely villous to strigose or pilose; peduncles 310[25]
mm; bracts numerous, narrowly triangular, linear, or filiform,
612 mm, densely brown villous. Flowers sessile. Calyx densely
strigose to pilose; hypanthium portion 1.52 mm; limb ca. 4
mm, usually 6-lobed; lobes linear-lanceolate, 1.53 mm. Corolla ca. 7 mm, densely strigose to pilose outside; tube ca. 4 mm;
lobes 5, oblong, ca. 3 mm. Fruit subglobose or obovoid, ca. 4
mm in diam., strigillose or subglabrous, slightly longitudinally
angled, crowned with calyx lobes; pyrenes 4 or 5. Fl. Apr, May.
Forests, shaded and wet places; 3001600 m. Guangxi, Hainan,
Yunnan [Indonesia (Java, Sumatra), Malaysia, Thailand, N Vietnam].
Lasianthus rhinocerotis subsp. rhinocerotis occurs in Indonesia
(Java, Sumatra), Malaysia, and Thailand. The description is as follows:
young branches and leaf abaxially densely hairy; leaf blade leathery,
base rounded or slightly cordate, lateral veins 912 pairs, nervules
densely anastomosing, parallel, stipules triangular, ca. 3 mm; bracts
slender, filiform.

1a. Leaf bases broadly cuneate to truncate

or subrounded; stipules inconspicuous;
bracts relatively thick, linear to narrowly
triangular; young branches and leaf
abaxially densely hairy; leaf blade
papery to leathery with 812 pairs
of lateral veins and with densely
anastomosing, parallel nervules
..................................................... 26a. subsp. pedunculatus
1b. Leaf bases rounded or slightly
cordate; stipules triangular, ca. 3
mm; bracts slender, filiform;
young branches and leaf

RUBIACEAE

196

abaxially sparsely hairy; leaf

blade papery with 7 or 8
pairs of lateral veins and
with loosely anastomosing,
subreticulate nervules ..... 26b. subsp. xishuangbannaensis
26a. Lasianthus rhinocerotis subsp. pedunculatus (Pitard) H.
Zhu, Acta Bot. Yunnan. 20: 154. 1998.
you geng cu ye mu
Lasianthus rhinocerotis var. pedunculatus Pitard in
Lecomte, Fl. Indo-Chine 3: 395. 1924; L. koi Merrill & Chun.
Young branches and leaf abaxially densely hairy. Leaf
blade papery to leathery, base broadly cuneate to truncate or
subrounded; lateral veins 812 pairs; nervules densely anastomosing, parallel; stipules inconspicuous. Bracts relatively
thick, linear to narrowly triangular.
Forests, shaded and wet places; 3001000 m. Guangxi, Hainan,
Yunnan [N Vietnam].

26b. Lasianthus rhinocerotis subsp. xishuangbannaensis H.

Zhu & H. Wang, Acta Phytotax. Sin. 38: 282. 2000.
ban na cu ye mu
Young branches and leaf abaxially sparsely hairy. Leaf
blade papery, base rounded or slightly cordate; lateral veins 7 or
8 pairs; nervules loosely anastomosing, subreticulate; stipules
triangular, ca. 3 mm. Bracts slender, filiform.
Montane forests, shaded and wet places; 15001600 m. S Yunnan
[Thailand].

27. Lasianthus rigidus Miquel, Fl. Ned. Ind. 2: 321. 1857.

da ye cu ye mu
Lasianthus humilis Elmer; L. tentaculatus J. D. Hooker.
Shrubs, height unknown; branchlets glabrous or subglabrous. Petiole 47 mm, sparsely hirsute; leaf blade papery or
leathery, obovate-oblong, obovate, or oblong, 1218 57 cm,
glabrous adaxially, hirsute on midrib, lateral veins, and nervules
abaxially, base oblique, obtuse, slightly cordate, or rounded,
margin thinly recurved, apex shortly cuspidate-acuminate; lateral veins 8 or 9 pairs, elevated abaxially; nervules parallel,
conspicuous abaxially; stipules persistent, triangular or subulate, 56 mm, hirsute. Inflorescences glomerulate, sessile, hirsute to hirtellous; bracts numerous, linear or linear-lanceolate to
lanceolate, 912 mm, hirsute. Flowers sessile. Calyx sparsely
hirsute; hypanthium portion campanulate, ca. 2 mm; limb ca. 2
mm; teeth 5, triangular, ca. 1 mm. Corolla ca. 1 cm, puberulent
outside, pubescent inside in upper part; lobes 5. Fruit globose,
45 mm in diam., glabrous; pyrenes 5.
Forests, shaded and wet places; 500700 m. Hainan, S Yunnan
[NE India, Indonesia (Java, Sumatra), Philippines].

28. Lasianthus schmidtii K. Schumann, Bot. Tidsskr. 24: 340.

1902.
tai bei cu ye mu
Lasianthus kerrii Craib.

Shrubs, 11.5 m tall; branches and branchlets densely tomentose to strigose. Petiole 49 mm, tomentose or strigose; leaf
blade papery or rigid-papery, oblanceolate, elliptic-oblanceolate, or oblong, 510 24 cm, glabrous adaxially, tomentose
or strigillose on midrib, nerves, and nervules abaxially, base
cuneate, obtuse, or rounded, apex shortly acuminate or acute;
lateral veins 57 pairs; nervules subreticulate; nerves and nervules elevated conspicuously abaxially; stipules subulate-triangular, 23 mm, tomentose or strigose. Inflorescence glomerulate, sessile; bracts subulate, 46 mm, tomentose or strigose.
Flowers sessile. Calyx tomentose or strigose; hypanthium portion obconic, 11.5 mm; limb deeply lobed; lobes 4, triangularsubulate to lanceolate, 1.52 mm. Corolla ca. 9 mm; tube ca. 7
mm, sparsely strigillose outside; lobes 4, ca. 2 mm. Fruit blue,
subglobose, 45 mm in diam., tomentose or strigillose; pyrenes
4.
Montane forests, shaded and wet places; 10001200 m. S Yunnan
[N Thailand].
H. Zhu (Syst. & Geogr. Pl. 72: 79. 2002) noted that this species is
rare and very similar to Lasianthus sikkimensis.

29. Lasianthus simizui (T. S. Liu & J. M. Chao) H. Zhu, Syst.

& Geogr. Pl. 72: 92. 2002 [shimizui].
qing shui shi ji shi shu
Lasianthus obliquinervis Merrill var. simizui T. S. Liu & J.
M. Chao, Taiwania 10: 143. 1964; L. trichophlebus Hemsley
var. simizui (T. S. Liu & J. M. Chao) H. Zhu.
Shrubs; branches and branchlets densely hirsute. Petiole
35 mm, hirsute; leaf blade leathery or subleathery, oblong or
elliptic-oblong, 610 2.54 cm, glabrous adaxially, hirsute on
midrib, nerves, and nervules abaxially, base acute, apex cuspidate or acute; lateral veins 68 pairs; nervules parallel; nerves
and nervules elevated abaxially; stipules triangular-lanceolate,
35 mm, hirsute. Inflorescences sessile; bracts subulate, reduced. Flowers sessile. Calyx hirsute; hypanthium portion campanulate; limb ca. 2 mm, toothed for ca. 1/2; teeth 5, ca. 1 mm.
Corolla not seen. Fruit subglobose, 56 mm in diam., subglabrous; pyrenes 5.
Forests, shaded and wet places. Taiwan.
This species is similar to Lasianthus trichophlebus and L. verticillatus. It has stipules longer than in L. verticillatus and shorter than in L.
trichophlebus, indumentum and a dentate calyx similar to L. trichophlebus, but 5-merous flowers and smooth fruit like L. verticillatus.

30. Lasianthus sikkimensis J. D. Hooker, Fl. Brit. India 3:

180. 1880.
xi jin cu ye mu
Lasianthus langkokensis Drake ex Pitard; L. sikkimensis
subsp. langkokensis (Drake ex Pitard) H. Zhu; L. tsangii Merrill
ex H. L. Li.
Shrubs, 13 m tall; branches and branchlets densely brown
tomentose or -villous. Petiole 614 mm, densely tomentose to
villous; leaf blade subleathery or thinly leathery, elliptic-lanceolate or elliptic, 1220 2.55 cm, glabrous adaxially, densely
to moderately brown tomentose or villous to hirtellous abaxially, base acute or obtuse, margin usually thinly recurved,

RUBIACEAE

apex cuspidate-acuminate; lateral veins 810 pairs; midrib and

nerves elevated conspicuously abaxially; nervules subparallel,
conspicuous abaxially; stipules generally persistent, triangular,
23 mm, densely tomentose to villous. Inflorescences glomerulate, sessile, 13-flowered, densely strigose to villous; bracts
persistent, linear to lanceolate, 0.32 cm. Flowers sessile or
subsessile. Calyx with hypanthium portion obconic, 12 mm,
glabrous; limb 12 mm, hirtellous to strigillose, 5-lobed; lobes
triangular. Corolla ca. 1 cm, puberulent outside. Fruit ellipsoid,
68 mm, glabrous, smooth or with ca. 5 low ridges; pyrenes 5.
Fr. Jun, Oct.
Forests, shaded and wet places; 3001600 m. Fujian, Guangdong,
Guangxi, Taiwan, Yunnan, Zhejiang [Bangladesh, NE India, Philippines, Thailand, N Vietnam].
The combination of the glabrous fruit body and densely pubescent
persisting calyx limb and the well-developed bracts on the sessile glomerulate inflorescences are distinctive for this species. H. Zhu (Syst. &
Geogr. Pl. 72: 80. 2002) noted that the two subspecies of Lasianthus
sikkimensis that were previously recognized have become difficult or
impossible to separate now that more collections are available. Zhu (loc.
cit.: 79) also noted that this species is very similar to L. schmidtii.

31. Lasianthus trichophlebus Hemsley, J. Linn. Soc., Bot. 23:

388. 1888.
zhong e cu ye mu
Shrubs, 12 m tall; branchlets strigillose to hirtellous. Petiole 58 mm, strigillose to hirsute; leaf blade leathery, oblong or
elliptic-oblong, 815 35 cm, glabrous adaxially, strigillose
or hirsute on midrib and nerves abaxially, base cuneate, apex
cuspidate or acute; lateral veins 79 pairs; nervules parallel;
nerves and nervules elevated abaxially; stipules triangular-lanceolate to narrowly triangular, 58 mm, densely strigillose. Inflorescences glomerulate, sessile; bracts absent. Flowers subsessile. Calyx strigillose to hirsute; hypanthium obconic to campanulate, ca. 2 mm; limb 11.5 mm, deeply 4-lobed; lobes narrowly triangular, ca. 1 mm. Corolla 68 mm, densely strigillose
in upper half outside; lobes 4. Fruit subglobose, 56 mm in
diam., verrucose, strigillose; pyrenes 4. Fl. AprMay, fr. Sep
Oct.
Forests, shaded and wet places; ca. 100 m. Guangdong, Hainan,
Taiwan [Indonesia, Malaysia (Peninsular), Philippines, Singapore, Thailand, Vietnam].
Lasianthus trichophlebus was often confused with both L. chinensis and L. verticillatus. However, it differs from L. chinensis by having
leathery leaves with 79 pairs of lateral veins and parallel veinlets;
stipules triangular-lanceolate, strigillose, 58 mm; flowers 4-merous;
and pyrenes 4. It differs from L. verticillatus by having strigillose hairs
on young branches and leaf nerves abaxially; stipules triangular-lanceolate, strigillose, 58 mm; flowers 4-merous; calyx with strigillose triangular lobes; and drupes with 4 pyrenes and verrucose on external surface.

1a. Branches densely hirsute, leaves

pubescent abaxially ................................ 31a. var. latifolius
1b. Branches sparsely strigillose; leaves
abaxially glabrous except strigillose
or hirsute on midrib and nerves ..... 31b. var. trichophlebus
31a. Lasianthus trichophlebus var. latifolius (Miquel) H.

197

Zhu, Syst. & Geogr. Pl. 72: 93. 2002.

qi lan zhong e cu ye mu
Lasianthus sylvestris Blume f. latifolius Miquel, Ann.
Mus. Bot. Lugduno-Batavi 4: 248. 1869 [latifolia]; L. hiiranensis Hayata; L. lei Merrill & F. P. Metcalf ex H. S. Lo.
Branches densely hirsute. Leaves pubescent abaxially.
Forests, shaded and wet places. Hainan, Taiwan [Indonesia (Java,
Sumatra), Malayasia (Peninsular), Philippines, Singapore, Thailand,
Vietnam].
This taxon was treated as Lasianthus tomentosus Blume by King
and Gamble (J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 73: 130. 1904),
Ridley (Fl. Malay Penin. 2: 161. 1923), Craib (Fl. Siam. 2(2): 219.
1934), and H. Zhu (Acta Bot. Yunnan. 20: 155. 1998), which may be
due to the authors following two misidentified sheets in K. These two
specimens, which are labeled as Horsfield s.n. [Rub. 50] from Java, are
indicated as L. tomentosus Blume with Miquels handwriting and also
were cited as such in print (Miquel, Fl. Ned. Ind. 2: 318. 1857). Miquel
(loc. cit. 1869) later changed his mind and considered that they were not
the same as L. tomentosus. The type materials of L. tomentosus are only
in Leiden and there are no overseas duplicates. After examining Malesian specimens and carefully comparing with the type of L. tomentosus,
it is recognized that the taxon that was mistaken for L. tomentosus has
conspicuous, strigose, linear-lanceolate stipules, 4-merous flowers, hirsute and verrucose drupes with 4 pyrenes, and is more closely related to
L. trichophlebus than L. tomentosus. The taxon differs from L. trichophlebus by having only dense hirsute hairs on the branches and pubescent hairs on the abaxial leaf surface. It is better, therefore, to treat this
taxon as a variety of L. trichophlebus.

31b. Lasianthus trichophlebus var. trichophlebus

() zhong e cu ye mu (yuan bian zhong)
Lasianthus barbellatus Ridley; L. cupreus Pierre ex Pitard.
Branches sparsely strigillose. Leaves glabrous except strigillose or hirsute on midrib and nerves abaxially.
Forests, shaded and wet places. Guangdong [Indonesia, Malaysia
(Peninsular), Philippines, Thailand, Vietnam].

32. Lasianthus verticillatus (Loureiro) Merrill, Trans. Amer.

Philos. Soc., n.s., 24: 372. 1935.
xie mai cu ye mu
Dasus verticillatus Loureiro, Fl. Cochinch. 1: 142. 1790;
Lasianthus andamanicus J. D. Hooker; L. taitoensis Simizu; L.
tamirensis Pierre ex Pitard.
Shrubs, 1.53 m tall; branches and branchlets appressed
pubescent or strigillose to glabrescent. Petiole 710 mm,
densely pubescent or strigillose to strigose; leaf blade leathery,
oblong to elliptic-oblong, 8.518 3.57 cm, glabrous adaxially, thinly pubescent or strigillose abaxially, base acute to
broadly obtuse, margins usually undulate, often thinly revolute,
apex acute or acuminate; lateral veins 79 pairs; nervules parallel; nerves and nervules elevated conspicuously abaxially; stipules generally persistent, triangular to narrowly triangular, 35
mm, densely puberulent to strigillose. Inflorescences subcapitate to congested-cymose, sessile; bracts absent or reduced.
Flowers sessile to subsessile. Calyx subglabrous or puberulent;

198

RUBIACEAE

hypanthium portion campanulate to subglobose, 2.53 mm;

limb 22.5 mm, truncate or minutely dentate. Corolla 1012
mm, hirtellous to villous outside, villous inside; lobes 5, ovate.
Fruit blue, ellipsoid, up to 10 mm in diam., strigillose to glabrescent, smooth; pyrenes 4 or 5. Fl. AprMay, fr. OctNov.
Forests, shaded and wet places; 1001000 m. Guangdong, Guangxi, Hainan, Taiwan, Yunnan [Cambodia, India (Andaman and Nicobar
Islands), Indonesia, Japan (Ryukyu Islands), Laos, Malaysia, Myanmar,
Philippines, Thailand, Vietnam].
The relatively large flowers with the large globose to ellipsoid
hypanthium (i.e., ovary) portion are distinctive; Lasianthus chinensis is
similar in these characters but can be distinguished by its deeply lobed
calyx limb and angled to ridged fruit.
Specimens from mainland SE China, Taiwan, and Ryukyu Islands
that have been treated as Lasianthus obliquinervis Merrill are clearly
conspecific with L. verticillatus. In Merrills description (Philipp. J. Sci.
1(Suppl. 1): 136. 1906), L. obliquinervis is described as having oblongovate stipules ca. 5 mm and fruit with 6 triquetrous pyrenes, which does
not match L. verticillatus. Lasianthus obliquinervis represents a different species with distribution in the Philippines and Papua New
Guinea. The so-called L. obliquinervis from China, Taiwan, and the

Ryukyus in Chinese and Japanese literature is a misidentification of the

species L. verticillatus.

33. Lasianthus wardii C. E. C. Fischer & Kaul, Bull. Misc.

Inform. Kew 1940: 292. 1941.
dian xi cu ye mu
Shrubs, height unknown; branchlets fuscous tomentose.
Petiole 510 mm, tomentose; leaf blade papery, dark olivaceous
adaxially, elliptic-oblong to oblanceolate-oblong, 1217 3.5
6 cm, glabrous adaxially, brown- or fuscous tomentose abaxially especially on nerves, base cuneate, apex caudate-acuminate; lateral veins ca. 7 pairs, elevated abaxially; nervules
subreticulate; stipules broadly ovate, cuspidate. Inflorescences
glomerulate, sessile; bracts absent. Flowers sessile. Calyx
brown tomentose; hypanthium portion campanulate, very short;
limb deeply lobed; lobes 5, oblong, ca. 3 mm, obtuse. Corolla
shortly brown tomentose outside; tube up to 1.6 cm, whitishor brownish pilose in upper half inside; lobes 5, triangularovate, ca. 4.5 mm. Fruit not seen.
Forests, shaded and wet places. Yunnan [Myanmar].
H. Zhu (Syst. & Geogr. Pl. 72: 8384. 2002) cited only three
specimens in total of this species, which suggests that it is rare.

44. LEPTODERMIS Wallich in Roxburgh, Fl. Ind. 2: 191. 1824.

ye ding xiang shu
Chen Tao ()
Shrubs, sometimes low or reduced, unarmed, usually with short shoots. Raphides present. Leaves opposite, apparently without
domatia; stipules persistent, interpetiolar, triangular, often aristate to spinescent. Inflorescences terminal on main stems and/or
axillary short shoots (and then appearing axillary), capitate to congested-fasciculate or -cymose and several flowered or sometimes
reduced to 1 flower, bracteate with bracts often fused in pairs. Flowers sessile to shortly pedicellate, bisexual, distylous. Calyx limb
(4 or)5(or 6)-lobed. Corolla white to pink or purple, funnelform, inside glabrous to pubescent; lobes (4 or)5(or 6), valvate or valvateinduplicate in bud. Stamens (4 or)5(or 6), inserted in corolla throat, exserted or included; filaments short; anthers dorsifixed. Ovary
5-celled, ovules 1 in each cell, basal, erect, anatropous; stigmas (2 or)35(or 6), linear, exserted or included. Fruit capsular, ellipsoid
to ovoid, opening through an apical operculum then splitting longitudinally into 5 valves, cartilaginous to woody, calyx limb
persistent; aril reticulate, free or adherent to seed; seeds few, medium-sized, ellipsoid, straight, with thin testa; cotyledons rounded;
radicle short, inferior.
About 40 species: Himalaya to Japan; 34 species (30 endemic) in China.

1a. Aril free from testa (fruit of L. brevisepala and L. dielsiana not seen).
2a. Bracteoles free.
3a. Stipules compressed triangular, shortly acuminate; bracteole longer than calyx ........................................... 5. L. dielsiana
3b. Stipules long triangular, acuminate; bracteole shorter than calyx or subequal in length ................................... 26. L. scissa
2b. Bracteoles connate.
4a. Bracteoles connate from base to middle.
5a. Bracteoles longer than calyces or nearly as long as calyces.
6a. Flowers mostly terminal on branches or laterally arranged; calyx lobes lanceolate-subulate,
acuminate or shortly acuminate, glabrous; corolla funnelform, lobes not incurved at apex;
stamens inserted in corolla tube throat ............................................................................................. 8. L. glomerata
6b. Flowers 3 terminal on branches; calyx lobes short or long triangular or subovate, very acute,
ciliate; corolla narrowly funnelform, lobes incurved at apex; stamens inserted below corolla
tube throat.
7a. Calyx lobes short or long triangular ........................................................................................ 14. L. limprichtii
7b. Calyx lobes subovate ..................................................................................................................... 21. L. pumila
5b. Bracteoles shorter than calyces.
8a. Leaves thickly papery, glabrous; stipules subtriangular; calyx lobes not ciliate ........................... 2. L. brevisepala
8b. Leaves papery, pilose; stipules broadly triangular, glandular on margins; calyx lobes ciliate.

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199

9a. Stipules with stiff acicular apiculus, sometimes with stipitate glands on margins; corolla
lobes triangular-oblate, apex obtuse; stamens inserted in corolla tube throat ...................... 29. L. velutiniflora
9b. Stipules with acicular apiculus, with 2 glandular teeth; corolla lobes ovate-triangular,
apex reflexed; stamens inserted below corolla tube throat ........................................................ 20. L. potaninii
4b. Bracteoles 1/22/3 connate.
10a. Bracteoles equal or subequal in length to calyces; corolla lobes inflexed at apex.
11a. Flowers with pedicels, 39 mm ................................................................................................. 28. L. umbellata
11b. Flowers sessile or subsessile.
12a. Corolla small, 1114 mm; free portion of bracteole triangular ........................................... 16. L. oblonga
12b. Corolla large, to ca. 16 mm; free portion of bracteole broadly triangular ............................. 9. L. gracilis
10b. Bracteoles longer than calyces; corolla lobes not inflexed at apex.
13a. Stigmas 3-lobed, filiform.
14a. Leaves thickly papery; bracteoles sparsely ciliate; calyx lobes shortly acuminate;
corolla purplish red, lobes 4 or 5, ovate-lanceolate; stamens inserted above corolla
tube throat ............................................................................................................................ 17. L. ordosica
14b. Leaves leathery; bracteoles pilose; calyx lobes obtuse; corolla white, lobes 5,
sublanceolate; stamens inserted below corolla tube throat ................................................ 24. L. scabrida
13b. Stigmas 5-lobed, linear.
15a. Leaves papery, 310 cm, petioles ca. 24 mm; calyx lobes triangular-ovate; corolla
lobes with 3 obtuse teeth, incurved .............................................................................. 12. L. kumaonensis
15b. Leaves subleathery or leathery, to 4 cm, petioles 47 mm; calyx lobes broadly
oblong to suborbicular; corolla lobes obtuse.
16a. Leaves ovate to broadly lanceolate, whitish or pale yellow abaxially when
dry, veinlets indistinct abaxially ........................................................................... 4. L. coriaceifolia
16b. Leaves ovate, ovate-oblong, elliptic, or suborbicular, pale green or iron-gray
abaxially when dry, veinlets conspicuous abaxially.
17a. Flowers 37 fascicled, terminal or in axils of upper 36 pairs of leaves,
subsessile or pedicels 0.32.6 mm ....................................................................... 30. L. vestita
17b. Flowers (1)35 fascicled, terminal on ultimate branches of
inflorescence, forming a thyrse, pedicels to 8 mm ................................ 33. L. yangshuoensis
1b. Aril adherent to testa.
18a. Bracteoles separated, scaly .......................................................................................................................... 1. L. beichuanensis
18b. Bracteoles connate.
19a. Bracteoles nearly as long as corolla.
20a. Stigmas 2-lobed .................................................................................................................................... 13. L. lanata
20b. Stigmas 5-lobed.
21a. Bracteoles tomentose.
22a. Bracteoles slightly longer than calyces; calyx lobes ovate, acuminate .................... 10. L. handeliana
22b. Bracteoles shorter than calyces; calyx lobes ovate-oblong, slightly obtuse .............. 27. L. tomentella
21b. Bracteoles pubescent or ciliate on upper portion.
23a. Leaves 14 ca. 2 cm, petioles to 1 cm; bracteoles subulate-acuminate at apex .... 23. L. rehderiana
23b. Leaves 49 1.252 mm, petioles very short or subsessile; bracteoles rounded
at apex, apiculate ......................................................................................................... 25. L. schneideri
19b. Bracteoles obviously longer or shorter than calyces.
24a. Bracteoles obviously longer than calyces.
25a. Stigmas usually 3-lobed ..................................................................................................... 32. L. xizangensis
25b. Stigmas usually 5-lobed.
26a. Leaves glabrous on both surfaces.
27a. Stipules triangular, glabrous; flowers subsessile; calyx lobes densely
fimbriate-ciliate; corolla pallid purple, narrowly funnelform, lobes
lanceolate; stamens inserted above corolla tube throat .......................................... 3. L. buxifolia
27b. Stipules ovate-triangular, covered with soft hairs; flowers with pedicel
35(8) mm; calyx lobes shortly ciliate; corolla purplish blue, funnelform,
lobes broadly ovate; stamens inserted below corolla tube throat ............................. 6. L. diffusa
26b. Leaves hairy on both surfaces, at least ciliate along margins.
28a. Stipules compressed triangular or triangular, acuminate, apiculus rarely
with 1 or 2 lateral glandular teeth; calyx lobes acuminate .............................. 10. L. handeliana
28b. Stipules broadly triangular, cuspidate-apiculate; calyx lobes obtuse to
truncate ..................................................................................................................... 19. L. pilosa

RUBIACEAE

200

24b. Bracteoles obviously shorter than calyces.

29a. Stigmas more than 3-lobed.
30a. Corolla lobes subovate, apex inflexed and beaked ................................................... 11. L. hirsutiflora
30b. Corolla lobes ovoid-lanceolate, apex mucronate or slightly obtuse.
31a. Leaves slightly thick; bracteoles abruptly acuminate at apex; calyx lobes
triangular-lanceolate; corolla lobes acute, margins revolute ............................. 18. L. parvifolia
31b. Leaves papery, margins obviously recurved; bracteoles abruptly acute;
calyx lobes oblong-ovate; corolla lobes slightly obtuse ..................................... 22. L. purdomii
29b. Stigmas 2- or 3-lobed.
32a. Corolla broadly funnelform; stamens inserted below corolla tube throat.
33a. Stipules apiculate at apex, sparsely pilose at base; bracteole lobes triangular;
corolla white, pallid pink, or purple, lobes apically inflexed and beaked ............ 15. L. ludlowii
33b. Stipules lacerate at apex, with glandular hairs along margins; bracteole
lobes lanceolate or ensiform-lanceolate; corolla purple, lobes shallowly
3-parted at apex, margins wavy ..................................................................................... 34. L. yui
32b. Corolla funnelform; stamens inserted in corolla tube throat.
34a. Stipules with fimbriate collaters along margins; corolla pallid blue or purplish
or purplish red; stigmas 2-lobed .............................................................................. 7. L. forrestii
34b. Stipules usually glandular along margins; corolla white or pallid red, stigmas
3-lobed ................................................................................................................... 31. L. wilsonii
1. Leptodermis beichuanensis H. S. Lo, J. Trop. Subtrop. Bot.
7(1): 19. 1999.
bei chuan ye ding xiang
Shrubs, ca. 2 m tall; branches slender, terete, gray or
grayish brown, with 2 bands of soft hairs. Petiole 13 mm; leaf
blade papery, olive-green when dry, narrowly lanceolate or oblong-lanceolate, sometimes subovate or elliptic, 0.83.5 0.31
cm, slightly scabrous adaxially, midrib strigose abaxially, base
cuneate, apex acuminate or acute; midrib and lateral veins
slightly prominent on both surfaces, lateral veins 3 or 4 pairs;
stipules small, subtriangular, ca. 1 mm, stiffly apiculate, usually caducous. Flowers 13 terminal, sessile; bracteoles scaly,
shorter than 1 mm, stiffly apiculate. Calyx tube ca. 1.8 mm;
lobes broadly triangular, width longer than length, ca. 0.6 mm,
sometimes margins denticulate. Immature fruit subobovate, ca.
3 mm; seed aril adherent to testa.
About 1400 m. NC Sichuan (Beichuan).

2. Leptodermis brevisepala H. S. Lo, J. Trop. Subtrop. Bot.

7(1): 21. 1999.
duan e ye ding xiang
Shrubs, ca. 40 cm tall; branchlets terete, erect and long,
ferruginous, pubescent or puberulent. Leaves sparsely arranged
or 4 fascicled at tips of branchlets; petiole 12 mm or sometimes leaves subsessile; blade thickly papery, nearly black adaxially and dark brown abaxially when dry, ovate or lanceolate, 6
10 35 mm, both surfaces glabrous, margins slightly revolute,
apex shortly acuminate or subacute, rarely obtuse; lateral veins
3 or 4 pairs, together with midrib conspicuously prominent
abaxially; stipules subtriangular, ca. 1.6 mm, subglabrous, with
a long stiff apiculus. Cymes almost sessile, terminal on short
branches, sometimes axillary, 57-flowered. Flowers subsessile
or shortly pedicellate; bracteoles transparent, connate below
middle portion, subequal to calyx tube in length, subglabrous or
puberulent, with a stiff acicular apiculus. Calyx tube black when
dry, 1.41.7 mm; lobes 5, ovate-orbicular or ovate-triangular,

length subequal to width, ca. 0.4 mm, not ciliate, obtuse or subacute. Corolla white, funnelform, 810 mm, puberulent outside,
densely white villous inside; lobes 5, suborbicular, margins
wavy or erose. Stamens 5, inserted below throat of corolla tube;
anthers linear, included. Style exserted; stigmas usually 3-lobed,
involute.
About 1800 m. Sichuan (Huili).

3. Leptodermis buxifolia H. S. Lo, J. Trop. Subtrop. Bot. 7(1):

20. 1999.
huang yang ye ye ding xiang
Leptodermis buxifolia f. strigosa H. S. Lo.
Shrubs, 0.52 m tall; branches terete, slightly stout, black
or dark gray, glabrous, branches slightly elongate or abbreviated. Leaves sparse or fascicled in axils of short branches;
petiole short or nearly absent; blade thickly leathery, ovate,
lanceolate, elliptic, or oblong, 58(12) 25(8) mm, both
surfaces glabrous, shiny, margins recurved, apex obtuse or
rounded, sometimes subacute; midrib raised abaxially, lateral
veins inconspicuous; stipules triangular, ca. 1.2 mm, glabrous.
Cymes terminal or axillary, usually 3-flowered, sometimes in
narrow thyrses. Flowers subsessile; bracteoles transparent, ca. 2
mm, ca. 3/4 connate, lobes short, with prominent tip, glabrous.
Calyx small; tube ca. 1 mm; lobes oval, 0.40.5 mm, densely
fimbriate-ciliate. Corolla light purple, narrowly funnelform, 9
10(14) mm; tube slender, pulverulent pubescent outside, pilose
inside, often pendulous; lobes 5, lanceolate, ca. 2 mm. Stamens
5, inserted above corolla tube throat; anthers filiform, exserted.
Style ca. 1/2 as long as corolla; stigmas (2 or)5-lobed, lobes
filiform, included. Fruit 44.5 mm; seed aril reticulate, adherent
to testa. Fl. JulAug, fr. AugSep.
Thickets, dry mountain slopes; 11002100 m. Gansu, SW
Shaanxi, Sichuan.

4. Leptodermis coriaceifolia Tao Chen, sp. nov.

ge ye ye ding xiang

RUBIACEAE

Type: China. Yunnan: Xichou Xian () [Si-chourhsien], Fadou () [Faa-doou], 1500 m, in open thickets
on rock hill, 26 Sep 1947, K. M. Feng 12065 (holotype, A;
isotypes, IBSC, KUN).
Haec species Leptodermidi vestitae Hemsley similis, sed
ab ea folii lamina ovata vel late lanceolata crassa in sicco coriacea ac abaxialiter albida vel pallide lutea, venis lateralibus
abaxialiter indistinctis vel parum prominentibus, venulis adaxialiter plerumque distinctis abaxialiter indistinctis, pedunculo
hirsuto vel glabrescente, pedicellis dense hirsutis, bracteolis
crassiusculis plerumque non hyalinis margine prope apicem
conspicue ciliato, corolla extus sparsim minuteque hirsuta trichomatibus sparsis longioribus prope alabastri apicem atque
stylo 3- usque 5-lobato differt.
Shrubs to 1.2(3) m tall. Stems dark brown, glabrescent,
lenticellate; branches many, young branches pale yellow when
dry, with 2 bands of sparse hairs decurrent from stipules; bark
gray, fissured and peeling off when old. Leaves decussate;
petiole short, to 0.7 mm, sparsely hairy to glabrous; blade dark
green adaxially, whitish or pale yellow abaxially when dry,
ovate to broadly lanceolate, 16 0.52.5 cm, thick, leathery
when dry, glabrous on both surfaces or sparsely hairy along
midvein, base attenuate, cuneate, margins sparsely hairy, apex
acuminate to acute; midvein impressed adaxially, prominent
abaxially, lateral veins distinct adaxially, indistinct or slightly
prominent abaxially, with 48 veins per side, veinlets usually
distinct adaxially, indistinct abaxially; stipules rigid, broadly
triangular, ca. 1 mm, hairy abaxially, cuspidate, mucronate at
apex. Inflorescences terminal on new shoots with all upper
nodes bearing 24 lateral shoots with axillary and terminal
clusters of cymules 39-flowered, forming a narrow thyrse with
an elongate central axis and short lateral branches; peduncle
hairy or glabrescent; bracts of cymules ligulate or leaflike, 1
4.5 mm or longer, with a midvein slightly prominent abaxially,
margins hairy, apex acuminate to acute. Flowers heterostylous,
subsessile or shortly pedicellate; pedicel densely hairy; bracteole pairs rigid, slightly thick, usually not transparent, obovate,
ca. 3 mm, longer than or subequal to calyx at anthesis, connate
for ca. 3/4 length, few veined, minutely short and sparsely hairy
abaxially, margins conspicuously ciliate near apex, apex acute
to obtuse, shortly mucronulate. Calyx tube absent; lobes 5,
ovate, ca. 1 mm, margins long ciliate, apex acute. Corolla purplish white; tube narrowly funnelform, ca. 10 mm, outside
sparsely minutely hairy, with sparse longer hairs near apex of
flower buds, inside villous above middle of tube; lobes 5, ovateoblong, ca. 3 mm, veined, lower part with thinner margins, upper part triangular, apex shortly acuminate, hooked. Longstyled flower: stamens inserted at throat ca. 1.5 mm from sinus,
filaments attached to lower 1/3 of anthers, free part shorter than
lower portion of anthers, anthers ca. 1.5 mm, introrse; style ca.
12 mm, glabrous, 3(5)-lobed, lobes ca. 1 mm, papillose. Capsule ca. 6 mm; seeds with reticulate aril free from testa. Fl.
SepOct, fr. NovDec.
Open thickets on limestone hills; 3001700 m. Guangxi, Yunnan
[Vietnam].

5. Leptodermis dielsiana H. J. P. Winkler, Repert. Spec. Nov.

Regni Veg. 18: 151. 1922.

201

li jiang ye ding xiang

Small shrubs, 850 cm tall, with rootlike flagella; young
branches not lignified, red, puberulent, old branches glabrescent, with gray bark, short branches 310 mm, sometimes
longer, with 2 or 3 pairs of leaves. Leaves sparsely arranged on
long branches, crowded on lateral short branches, smaller on
lower part of both short and long branches; petiole 26 mm,
pubescent; blade sometimes suborbicular, 615 412 mm, on
upper part of branches ovate or oblong, 1030 515 mm,
covered with conical hairs except pubescent on midrib adaxially, glabrous or sparsely hairy on nerves abaxially, base attenuate to petiole, apex acute or slightly obtuse, apiculate; lateral
veins 35 pairs, confluent in arcs; stipules membranous, ca. 1.5
mm, with 2 dominate veins parallel to margins and confluent at
apex, stipules on upper part, especially on inflorescences, compressed triangular, shortly acuminate, with 2 glandular teeth
near apex, 2 others slightly apart, stipules on lower part usually
without glandular teeth, ciliate at base, very shortly puberulent.
Flowers 13 terminal on very short branches, sometimes 1 or 2
flowers axillary in axils of upper leaf pair; pedicels covered
with papillose scabrid hairs; bracteoles 2, free, sessile, transparent, ovate-oblong or oblong, 2.53.5 mm, longer than calyx,
puberulent, ciliate on upper part, with veins percurrent, apiculate. Calyx lobes 5(or 6), submembranous, oblong, veined, ciliate, apiculate or slightly obtuse. Corolla large, 1517 mm; tube
funnelform, shortly papillose near base outside, villous, glabrous at base inside; lobes 34 shorter than corolla tube,
ovate-orbicular, 45 mm, glabrous outside, villous inside, not
reflexed, margins wavy. Stamens inserted in corolla tube throat;
filaments attached to middle of anthers, slightly longer than
base of anthers; anthers linear, included or semi-exserted. Style
5-lobed, glabrous, lobes relatively long, with long papilla, included or exserted. Fl. May.
NW Yunnan (Dali, Lijiang, Yongning).

6. Leptodermis diffusa Batalin, Trudy Imp. S.-Peterburgsk.

Bot. Sada 13: 373. 1894.
wen shui ye ding xiang
Shrubs, 11.5 m tall; branchlets often diffuse, together
with peduncles and pedicels puberulent, terete. Petiole to 4 mm;
leaf blade silver-gray or grayish yellow adaxially when dry,
pale abaxially, ovate-lanceolate to elliptic, 1840 815 mm,
both surfaces glabrous, base attenuate or cuneate, apex acute or
obtuse; midrib prominent below, lateral veins 4 pairs, slightly
conspicuous; stipules ovate-triangular, ca. 1.5 mm, with soft
hairs. Flowers terminal and in axils of upper leaves, subumbellate, usually many in a terminal, large, much-flowered panicle;
pedicels usually 35(8) mm; bracts purplish blue, turning
brown on margins, longer than calyces, 33.5 mm, glabrous or
hairy, membranous, connate to above middle, free portion very
shortly bilabiate, labellum short, apiculate. Flowers dimorphic,
heterostylous. Calyx lobes orbicular, shortly ciliate. Corolla purplish blue, funnelform, 1618 mm, puberulent outside, sparsely
pubescent inside; lobes more than 3 shorter than tubes,
broadly ovate, ca. 3.5 3.2 mm, abruptly acute, spreading. Anthers ca. 2.5 mm, slightly exserted in short-styled flowers; filaments as long as anthers, inserted in upper portion of calyx

202

RUBIACEAE

tube. Style exserted in long-styled flowers, filiform, glabrous;

stigmas 5-lobed, linear. Immature capsule broader and longer
than bracts. Fl. Aug, fr. SepOct.
Rocks at streamsides; 6001300 m. S Gansu (Wenxian), S
Sichuan.

7. Leptodermis forrestii Diels, Notes Roy. Bot. Gard. Edinburgh 5: 274. 1912.
gao shan ye ding xiang
Shrubs, 0.61.2 m tall, much branched; old branches gray
or slightly reddish, bark peeling off, young branches slender,
with 2 opposite shallow longitudinal channels, densely covered
with short soft hairs. Leaves petiolate or sometimes subsessile;
petiole shorter than 2 mm; blade membranous-papery, black
or sometimes brownish when dry, ovate or lanceolate, rarely
oblong or broadly ovate, 13 0.61.5 cm, dispersedly strigose
adaxially, glabrous or with crinkled long soft hairs on midrib
and lateral veins abaxially, rarely entirely hairy, base usually
sharply contracted, attenuate into short stalk, apex acute to subacuminate; lateral veins 46 pairs, slender, obvious abaxially;
stipules triangular or compressed triangular, 22.5 mm, pubescent outside, margins with fimbriate collaters, with 2 nerves
arching upward and joining at apiculate apex. Flowers usually
solitary, terminal, sessile, dimorphic, heterostylous; short-styled
flowers: bracts 2, small, ca. 1.5 mm, usually connate, awned
apiculate. Calyx black when dry, shiny; lobes 5, narrowly and
long triangular, 3.84 mm, slightly thick, glabrous or sparsely
ciliate on upper portion, equal to calyx in length or slightly
shorter. Corolla light blue or slightly reddish, funnelform, 20
22 mm, glabrous outside, white villous inside; limb broad,
spreading; lobes 5, induplicate, elliptic, ca. 6 mm, with an incrassate area in center, lateral areas thin, soft, equal to incrassate area in length, with branched venation, margins with
erose teeth, apex ?apiculate. Stamens 5, inserted in throat of
corolla tube; filaments ca. 1.5 mm; anthers linear, ca. 2.7 mm,
apex slightly exserted from tube. Style ca. 10 mm; stigmas 2lobed, lobes linear, ca. 2.7 mm. Capsules ca. 5 mm; seeds black;
aril reticulate, adherent to testa.
Forests; 32003400 m. Sichuan, SE Xizang (Bomi, Nyingchi),
NW Yunnan (Lijiang).

8. Leptodermis glomerata Hutchinson in Sargent, Pl. Wilson.

3: 406. 1916.
ju hua ye ding xiang
Small shrubs, up to 60 cm or slightly taller; stems
branched at base, branches erect, slender, usually purplish red,
with hairs in 2 opposite bands, internode 34 cm or longer.
Leaves opposite; petiole 12 mm or longer; blade thinly papery, lanceolate or ovate-lanceolate, sometimes ovate, 15
0.62 cm, glabrous or scabrid at margins adaxially, subglabrous abaxially, reticulately veined, base very pointed, apex
acute or acuminate; lateral veins 46 pairs, arched, prominent
abaxially; stipules broad at base, long cuspidate upward, 2.5
4 mm, sparsely hairy near base or subglabrous outside. Flowers congested at tips of branches, sessile or conspicuously petiolate for lateral flowers, dimorphic, heterostylous; long-styled
flowers with bracteoles membranous, opposite, connate at

base, 2.5 mm or slightly shorter, glabrous, abruptly subulateacuminate. Calyx turning black when dry; tube 22.2 mm, glabrous; lobes 5, lanceolate-subulate, ca. 2 mm, stiff, glabrous,
usually not ciliate, acuminate or acute. Corolla tube slightly
curved, ca. 1 cm, dilated upward, pubescent outside, sparsely
pubescent on throat; lobes 5, ovate, ca. 3.5 2.5 mm, obtuse
and apiculate. Stamens 5, inserted in throat of corolla tube;
filaments ca. 2.8 mm; anthers ca. 3 mm, slightly exserted. Style
glabrous, 1011 mm; stigmas 5-lobed, lobes ca. 1 mm; shortstyled flowers: style ca. 5 mm, stigmas 35-lobed, lobes linear,
2.55 mm. Capsule narrowly ellipsoidal, persistent calyx lobes
ca. 9 mm, 5-ridged, pale brown, glabrous, 5-valved when mature, with 5 utricle-like pyrenes; seeds linear, embedded in
reticulate aril; aril free from testa.
Sparse forests, hill slopes; 18002500 m. Yunnan.

9. Leptodermis gracilis C. E. C. Fischer, Bull. Misc. Inform.

Kew 1940: 293. 1941.
rou zhi ye ding xiang
Shrubs, 1.52 m tall; branches slender, usually nodding,
grayish white, pubescent. Leaves opposite; petiole 26 mm;
blade papery, silver-gray adaxially, slightly pallid abaxially,
elliptic to lanceolate, 1.43.5 513 mm, adaxially glabrous
except sometimes hispid on midrib and margin, abaxially usually hairy on midrib and lateral veins, margins hispidulous ciliate, base cuneate, decurrent, apex acute; stipules interpetiolar,
broad at base, long cuspidate upward, ca. 3 mm, puberulent,
ciliate. Cymes terminal and axillary on upper portion of branchlets; peduncle ca. 3.5 mm; flowers 3 fascicled, sessile; bracts
narrowly elliptic-lanceolate, ca. 4 mm, ciliate, acute; bracteoles
2, connate into a sheath ca. 2.5 mm, free portion broadly ovate,
11.5 mm, 3-nerved, apiculate. Calyx tube turbinate, 5-ridged,
1.21.5 mm; lobes 5, oblong, ca. 1.2 mm, ciliate, rounded or
subtruncate. Corolla white or purple, narrowly funnelform; tube
slender, ca. 8 mm, pilose above base inside; lobes 5, gladiate,
ca. 1.5 mm, subacute. Stamens 5, inserted in corolla tube throat;
anthers sessile, linear, ca. 1.5 mm, included. Style slender; stigmas 3, filiform, 11.5 mm, papillose, slightly exserted. Capsules linear-oblong, ca. 5 mm, dehiscent in 5 valves; seeds not
seen.
Thickets, hill slopes; 10002700 m. NC Sichuan, SE Xizang.

1a. Flowers smaller; corolla ca. 9.5 mm .......... 9a. var. gracilis
1b. Flowers larger; corolla ca. 16 mm ......... 9b. var. longiflora
9a. Leptodermis gracilis var. gracilis
() rou zhi ye ding xiang (yuan bian zhong)
Flowers smaller; corolla ca. 9.5 mm. Fl. Jul.
Hill slopes; 10002400 m. SE Xizang (Bomi, Mdog, Zay).

9b. Leptodermis gracilis var. longiflora H. S. Lo, var. nov.

chang hua ye ding xiang
Type: China. Sichuan: Barkam (Maerkang, ),
Zhoukeji (), 2650 m, 30 Jul 1957, Z. Y. Zhang & H. F.
Zhou 23393 (holotype, SZ; isotype, KUN).

RUBIACEAE

Validating Latin diagnosis: that of Leptodermis gracilis

C. E. C. Fischer var. longiflora Lo (H. S. Lo in W. C. Chen, Fl.
Reipubl. Popularis Sin. 71(2): 338. 1999).
Flowers larger, corolla ca. 16 mm. Fl. Jul.
Thickets; 26002700 m. NC Sichuan (Barkam).
This name was previously published by H. S. Lo (loc. cit.) but not
validly so because no type was indicated (Vienna Code, Art. 37.1).

10. Leptodermis handeliana H. J. P. Winkler in Handel-Mazzetti, Symb. Sin. 7: 1025. 1936.

chuan nan ye ding xiang
Shrubs; young branches with 2 bands of short woolly
hairs, old branches glabrescent, bark gray, longitudinally
lacerate. Leaves sparsely arranged on branches 310 cm, with
sparse and long internodes, very rarely crowded on short
branches; blade slightly thick, broadly ovate, elliptic, or broadly
elliptic, 718 411 mm, adaxially tomentose on midrib,
abaxially with scattered bristles on lateral veins, very shortly
setiform ciliate on margins, base abruptly contracted and
attenuate into petiole, apex acute or abruptly shortly acuminate; lateral veins 46 pairs, ascending; stipules membranous-leathery, triangular on lower part, compressed triangular
on upper part, acuminate, apiculus seldom with 1 or 2 lateral
glandular teeth, 12 mm, 2 veins joining at tip, dorsal surface
densely tomentulose, ventral surface densely covered with
glandular appendages. Flowers 57 terminal on branches, subsessile, highest node with 1 or 3 flowers in axils of leaves, next
node with 1 or 2 lateral branches terminated dichotomously
with flowers; bracts especially of flowers in center somewhat
leaflike, 45 mm, slightly longer than calyx, with midrib and
reticulate veins, tomentose, connate below middle part, acuminate. Calyx ca. 3 mm; lobes 5, ovate, ciliate, acuminate. Young
corolla hispidulous outside. Stigmas 5-lobed. Fl. Jun.
S Sichuan (Yanyuan).

11. Leptodermis hirsutiflora H. S. Lo, J. Trop. Subtrop. Bot.

7(1): 19. 1999.
la sa ye ding xiang
Shrubs; branchlets terete, long branches with conspicuous
internodes, short branches usually ca. 1 cm, young branches
shortly pilose, glabrescent when old. Leaves subsessile or petiole ca. 0.5 mm; blade oblong or elliptic, sometimes lanceolate
or ovate, 79 mm on long branches, 23 mm on short branches,
glabrous on both surfaces, base attenuate, apex obtuse or
slightly rounded; lateral veins inconspicuous; stipules broadly
triangular, ca. 1 mm, apiculate. Flowers sessile, terminal or
axillary on branchlets, usually densely flowered; bracteoles 2,
membranous, obovate, ca. 3.5 mm, connate below middle portion or one side free, with brown veins, densely hirsute outside, apex apiculate. Long-styled flower: calyx tube obovate,
ca. 2 mm, glabrous, lobes 5, narrowly triangular-lanceolate, ca.
2 mm, densely hirsute; corolla blue, narrowly funnelform,
densely hirsute outside, tube 89 mm, villous inside, base ca.
2.5 mm in diam., lobes 5, subovate, ca. 2.5 mm, apex incurved,
with a beak ca. 0.5 mm; stamens 5, inserted at base of throat,
filaments very short or absent, anthers linear, ca. 3 mm, nearly

203

included; style 1112 mm, stigmas 4- or 5-lobed, lobes linear,

exserted. Short-styled flowers similar to long-styled flowers;
calyx tube ca. 2 mm, lobes ca. 1.4 mm, acute; corolla tube
very shortly pilose outside; stamens inserted above corolla tube
throat, filaments ca. 0.8 mm, anthers exserted; style ca. 5.5 mm,
stigmas 3- or 4-lobed, included. Fruit ca. 5 mm, with persistent
calyx lobes 2.53 mm, hirsute; seeds ca. 4 mm; aril reticulate,
adherent to testa. Fl. Jul.
Thickets; 40004100 m. Xizang.

1a. Calyx lobes glabrous except ciliate on

margins; corolla pubescent ......................... 11a. var. ciliata
1b. Calyx lobes densely hirsute; corolla
densely hirsute outside, villous
inside .................................................. 11b. var. hirsutiflora
11a. Leptodermis hirsutiflora var. ciliata H. S. Lo, J. Trop.
Subtrop. Bot. 7(1): 20. 1999.
guang e ye ding xiang
Calyx lobes glabrous except ciliate on margins; corolla pubescent.
Thickets; ca. 4100 m. SE Xizang (Gyaca).

11b. Leptodermis hirsutiflora var. hirsutiflora

() la sa ye ding xiang (yuan bian zhong)
Calyx lobes densely hirsute; corolla densely hirsute outside, villous inside.
Thickets on desolate mountains; ca. 4000 m. Xizang.

12. Leptodermis kumaonensis R. Parker, Indian Forester 48:

576. 1922.
ji long ye ding xiang
Shrubs, 12 m tall, sometimes to 3 m tall; branchlets terete, usually purple, glandular pilose. Petiole 24 mm, somewhat
pilose; leaf blade papery, dark gray adaxially and grayish brown
or pale abaxially when dry, oblong-lanceolate or ovate-lanceolate, 310 12.5 cm on long branches, usually shorter than 1.5
cm on short branches, strigose pilose adaxially, pilose on midrib
and lateral veins abaxially, base broadly cuneate, margins entire, usually ciliate, apex subacuminate; lateral veins 69 pairs,
ascending in an arch, midrib compressed abaxially, veinlets
dark brown, inconspicuous adaxially, scrobiculate abaxially;
stipules triangular, 44.5 mm on upper part of branchlets,
shorter on lower part, pilose outside, abruptly contracted acuminate. Flowers 35 fascicled on ends of lateral short branches,
sessile or subsessile; bracts 2, ca. 7 mm or slightly longer, ca.
2/3 connate, free portion cuspidate, ciliate, often pilose near
midrib outside. Calyx tube 3.54 mm; lobes 5, triangular-ovate,
ca. 1.5 mm, ciliate, acute or slightly obtuse. Corolla funnelform,
1315 mm, pilose or subglabrous outside, white villous below
throat inside; lobes 5, with sinus ca. 3 mm in depth, apex obtusely 3-dentate, central portion slightly thickened, ca. 2 mm
wide, margins of both sides thin, 11.2 mm wide, inflexed in
bud. Long-styled flowers: stamens 5, inserted below throat, filaments almost absent, anthers dorsifixed, linear, ca. 3 mm, included; style ca. 9 mm, stigmas 5- or 6-lobed, lobes linear, 4

RUBIACEAE

204

5 mm, slightly exserted. Short-styled flowers: stamens with

filaments ca. 1.5 mm, anthers slightly exserted; style ca. 4.5
mm, stigmas 5 or 6, linear, ca. 2 mm, extending to below throat.
Capsule not seen, subcylindric, ca. 5.6 mm (according to record); seeds linear, covered in fibers; aril not adherent to testa.
Thickets, forest margins; 28003000 m. S Xizang (Gyirong)
[Bhutan, N India (Uttarakhand), Nepal].
Leptodermis parkeri Dunn (Bull. Misc. Inform. Kew 1920: 206.
1920), described from N India, was erroneously recorded from S Xizang (Gyirong) in FRPS (71(2): 139. 1999), based on the gathering
Qinghai-Xizang Expedition 6957, which is in fact referable to L.
kumaonensis.

22.5 mm, equal to calyx in length or slightly longer, 1/61/4

connate, shiny, 1-veined, not ciliate, long subulate-acuminate.
Calyx lobes usually 5, sometimes 4, long or shortly triangular,
ciliate, very apiculate. Corolla yellowish white, medium-sized;
tube narrowly funnelform, 89 mm, slightly curved, hispidulous outside, villous around middle inside; lobes ovate, ca. 2
mm, central portion thick and broader than transparent, wavy
margins, villous at base inside, apex incurved. Stamens in longstyled flowers inserted at middle of corolla tube; filaments very
short; anthers ca. 2 mm. Style usually (4 or)5-lobed.
Thickets or open fields on hill slopes; 10001500 m. C Sichuan
(Tianquan).

13. Leptodermis lanata H. S. Lo, J. Trop. Subtrop. Bot. 7(1):

19. 1999.

15. Leptodermis ludlowii Springate, Edinburgh J. Bot. 53:

102. 1996.

mian mao ye ding xiang

guan e ye ding xiang

Shrubs, 0.51 m tall; branches stout, terete, purplish red,

subglabrous, short branches densely covered with many persistent scaly stipules at base. Petiole ca. 1 mm or leaves subsessile;
leaf blade thickly papery, elliptic or obovate, sometimes oblong,
510 24 mm, tomentulose adaxially, grayish white lanate
abaxially, base cuneate, margins revolute, apex acute; lateral
veins inconspicuous; stipules long triangular, 1.52 mm, acuminate. Flowers sessile, usually 3 fascicled at tips of branchlets or
axillary near tips of short branches; bracteoles 2, scarious, lower
portion connate, longer than calyx or subequal, tomentulose,
apex cuspidate. Calyx tube ca. 2 mm, subglabrous; lobes 5, oblong-lanceolate, slightly shorter than calyx tube, puberulent and
shortly ciliate, shortly acuminate. Corolla pallid red, funnelform, 1214 mm, tomentulose outside; tube slender, decurved;
lobes oblong-lanceolate or oblong-ovate, ca. 2 mm. Long-styled
flowers: stamens 5, inserted below throat of corolla tube,
anthers linear, included; style slightly exserted, stigmas 2-lobed,
lobes filiform. Short-styled flowers similar to long styled; stamens inserted above throat of corolla tube, anthers slightly exserted. Fruit ca. 6 mm; seeds covered with reticulate aril adherent to testa. Fl. May.
Pinus forests, roadsides; 23002500 m. NW Yunnan (Ninglang).

14. Leptodermis limprichtii H. J. P. Winkler, Repert. Spec.

Nov. Regni Veg. Beih. 12: 490. 1922.
tian quan ye ding xiang
Small shrubs; young branches not lignified, with 2 bands
of soft hairs, old branches slender, glabrous, with gray bark.
Leaves sparsely borne on young abbreviated branches, subsessile; blade oblong-lanceolate, 12.5 0.30.9 cm, adaxially
with scattered hairs, tomentulose on midrib, abaxially glabrous,
base attenuate, margins with dense bristles, apex acute, apiculate; lateral veins 35 pairs, inconspicuous, reticulate veins
conspicuous; stipules triangular, subulate acuminate and with 2
glandular teeth near apex, another 2 glandular teeth remotely
arranged or lacerate, central portion triangular, membranous-papery, villous, with transparent broad margins, not ciliate. Flowers usually 3 terminal on branchlets, subsessile, sometimes lateral flowers pedicellate, sometimes with a pair of flowers in
axils of upper nodes; bracteoles thin, transparent, ovate-oblong,

Leptodermis tubicalyx H. S. Lo.

Shrubs, 0.62(3) m tall; branchlets slender, terete, with 2
bands of hairs decurrent from base of stipules. Petiole 0.20.8
cm, pilose adaxially; leaf blade membranous-papery or submembranous, turning black or iron-gray when dry, ovate to
ovate-oblong or broadly lanceolate, usually 15 0.51.5(2)
cm, subglabrous to puberulent on both surfaces, base attenuate
or slightly decurrent, margins ciliate, apex acute or acuminate;
lateral veins slender, (3 or)4 or 5 pairs, inconspicuous adaxially,
slightly prominent abaxially; stipules subtriangular or long
triangular, 1.52(4) mm, glandular-denticulate or laciniate,
sparsely pubescent at base, cuspidate. Inflorescences terminating short new growths, small, with uppermost pair of nodes
approximate, few cymules and 35(12) flowers; bract pairs
leaflike and petiolate, rarely ligulate on subsidiary 1-flowered
cymules; stipules as those of vegetative growth or reduced, hyaline, and laciniate. Flowers sessile; bracteole pairs ca. 1.5 mm,
ca. 1/3 connate at base, lobes triangular, obtuse or subacute, glabrous, usually ciliate, mucronulate, central pairs in a cymule
often more elongate and cuspidate. Calyx glabrous; hypanthium
ca. 3.4 mm; tube black when dry, obovate or tubular, 1.11.4
mm above ovary; lobes 5, triangular, rarely orbicular, 0.70.8
mm, hairy or papillate in bud or around margins, rarely 6 lobes
or 2 deeper clefts in tube. Corolla white, light pink, or purple,
sometimes bicolored, black when dry, broadly funnelform, with
white stiff hairs at apex in bud, exterior finely mammillate,
more prominently so toward apex, very rarely with few short
hairs decurrent from sinuses of lobes; tube 1011.5 mm, with
long fine hairs within; lobes 5, broadly ovate-triangular, 24
mm, margins thin, inflexed, ca. 1 mm wide, central portion ca.
2 mm wide, apex inflexed in a beak, with few short stout
smooth hairs on inner surface. Stamens 5. Long-styled flowers:
filaments ca. 0.3 mm, anthers 1.72 mm, attached near base or
rarely inserted in middle of corolla tube, included; stigmas
completely exserted. Short-styled flowers: filaments 11.7 mm,
anthers 1.92.7 mm, inserted below middle of corolla tube,
partly exserted; stigmas immediately below anthers. Stigmas 2
4-lobed, lobes linear, 22.5 mm, with hairs sometimes extending onto upper style. Capsule narrow, oblong, ca. 7 mm;
seeds oblong; aril reticulate, adherent to testa. Fl. JunSep.
Forests; ca. 2800 m. Xizang [Bhutan, NE India (Sikkim)].

RUBIACEAE

16. Leptodermis oblonga Bunge, Enum. Pl. Chin. Bor. 34.

1833.
bao pi mu
Hamiltonia oblonga (Bunge) Franchet; Leptodermis chanetii H. Lveill; L. huashanica H. S. Lo; L. oblonga var. leptophylla H. J. P. Winkler.
Shrubs, 0.21 m tall or slightly taller; branches slender,
gray to pale brown, puberulent, bark thin, usually peeling off
when old. Petiole short, 13 mm; leaf blade papery or thinly
leathery, lanceolate or oblong, sometimes elliptic or subovate,
usually 0.72.5(3) 0.31(1.7) cm, adaxially scabrid or
irregularly wrinkled and sparsely strigose, abaxially subglabrous or pubescent or hispid along midrib, base attenuate or
cuneate, margins revolute, apex acuminate, subacute, or obtuse;
lateral veins ca. 3 pairs, inconspicuous on both surfaces; stipules broadly triangular, 1.52 mm, cuspidate with a stiff apiculus. Flowers sessile, usually 3 fascicled and terminal on
branches, rarely axillary on upper part of branchlets; bracteoles
hyaline, ovate, 34.5 mm, 1/22/3 connate, pilose outside, lobes
subtriangular, with a stiff apiculus at apex, subequal to calyx in
length. Calyx tube ca. 2.5 mm; lobes subovate, ca. (0.7)1.3
1.5 mm, densely shortly ciliate, obtuse or shortly acuminate.
Corolla purplish red, funnelform, (9)1114(20) mm; tube
slender, often curved, puberulent outside; lobes subovate, narrowly triangular, or lanceolate, 24 mm, apex incurved. Shortstyled flowers: stamens slightly exserted, anthers linear; style
included. Long-styled flowers: stamens included, anthers linearoblong; style slightly exserted. Style with 4 or 5 linear lobes of
stigma. Seed aril reticulate, free from testa. Fl. JunAug, fr. Oct.
Sunny hill slopes, roadsides, thickets. Gansu, Hebei, N Henan,
Ningxia, Shaanxi, Shanxi, Sichuan [?Mongolia].

17. Leptodermis ordosica H. C. Fu & E. W. Ma, Fl. Intramongol. 5: 413. 1981.

nei meng ye ding xiang
Small shrubs, much branched, 2040 cm tall; branches
slightly stout, often twisted, dark gray, with thin cracks; branchlets slender and straight, sometimes acicular, gray, puberulent.
Petiole short or leaves subsessile; leaf blade thickly papery,
oblong to elliptic, sometimes broadly elliptic, 310 25 mm,
both surfaces subglabrous, slightly shiny adaxially, base cuneate or acuminate, margins often slightly revolute, apex acute or
slightly obtuse; midrib prominent abaxially, lateral veins very
inconspicuous; stipules triangular-ovate or ovate-lanceolate,
slightly longer than petioles, denticulate or entire, ciliate, apiculate. Flowers subsessile, 13 fascicled on tips of branches or in
axil near tips of branches; bracteoles 2, 1/22/3 connate, free
part bilabiate, 34 mm, transparent, lobes sparsely ciliate, caudate-acuminate at apex. Calyx 22.5 mm; lobes 5, oblong-lanceolate, subequal to calyx tube in length or slightly shorter, ciliate, shortly acuminate. Corolla purplish red, fragrant, funnelform, 1114 mm, puberulent outside, villous inside; lobes 4 or
5, ovate-lanceolate, ca. 3 mm. Stamens 4 or 5, inserted above
throat of corolla tube; anthers linear, slightly exserted. Style ca.
1/2 length of corolla tube; stigmas 3, filiform. Fruit 33.5 mm;
seeds surrounded by reticulate aril free from testa. Fl. and fr.
JulAug.

205

Rock crevices; ca. 1600 m. Nei Mongol.

18. Leptodermis parvifolia Hutchinson in Sargent, Pl. Wilson.

3: 404. 1916.
wa shan ye ding xiang
Erect or procumbent shrubs, 0.51(1.5) m tall; branchlets
short or slightly long, gray or reddish when young, puberulent,
soon glabrescent. Petiole short; leaf blade slightly thick, spatulate-ovate or oblanceolate, sometimes subovate or elliptic, 5
10 25 mm, base attenuate, apex obtuse or subacute; lateral
veins ca. 3 pairs, usually inconspicuous; stipules subulate-acuminate, ca. 1 mm, shorter than calyx, glabrous, abruptly acuminate. Calyx tube with 10 ridges, 5 ridges extending to
sinus, glabrous; lobes 5, triangular-lanceolate, ca. 1.25 mm,
shortly ciliate, acute. Corolla tube ca. 1.2 cm, upper portion
enlarged, puberulent outside, sparsely pubescent inside; lobes 5,
ovate-lanceolate, ca. 2.5 mm, with revolute margins, acute. Anthers ca. 1.25 mm, slightly exserted. Style equal to corolla tube
in length, glabrous, 5-lobed, lobes slender. Capsules 45 mm;
seed aril reticulate, adherent to testa or free at one end. Fl. Aug
Sep.
Thickets or forest margins on sunny mountain slopes; 1500
3000 m. Sichuan.

19. Leptodermis pilosa Diels, Notes Roy. Bot. Gard. Edinburgh 5: 276. 1912.
chuan dian ye ding xiang
Shrubs, 0.72(3) m tall; branches subterete, young
branches tomentulose or pubescent, old branches glabrescent,
with thin, lamellar, longitudinally fissured bark. Petiole 15 mm,
hairy; leaf blade papery, occasionally thinly leathery, shape
and size often variable, broadly ovate, ovate, oblong, elliptic, or
lanceolate, 0.52.5 ca. 1.5 cm, both surfaces sparsely to
densely pilose or subglabrous, base cuneate or attenuate, margins usually ciliate, apex acute, obtuse, or sometimes rounded;
lateral veins 35 pairs, slightly prominent or inconspicuous
abaxially; stipules broadly triangular, pilose or tomentose,
cuspidate-apiculate, usually 12 mm. Cymes terminal or axillary near tips of branches, usually 3(7)-flowered; flowers sessile or shortly pedicellate; bracteoles scarious, transparent,
longer than calyx, hairy, 2/33/4 connate, free portion subulate-acuminate, apiculate, veined, ciliate. Calyx tube ca. 2 mm;
lobes 5, 11.2 mm, ciliate, obtuse or subtruncate. Corolla funnelform; tube 910(13) mm, densely tomentulose outside, villous inside; lobes 5, broadly ovate, 22.5 mm, margins narrow
and thin, inflexed, apex incurved. Stamens 5, inserted in corolla
tube throat; filaments short; anthers linear, slightly exserted in
short-styled flowers, included in long-styled flowers. Style usually with (3)5 filiform stigmas, exserted in long-styled flowers,
included in short-styled flowers. Fruit 4.55 mm; seed aril
reticulate, adherent to testa. Fl. Jun, fr. SepOct.
Thickets; 6003800 m. S Gansu, W Hubei, S Shaanxi, Sichuan,
Xizang, Yunnan.

1a. Branches spiny ............................... 19a. var. acanthoclada

1b. Branches not spiny.
2a. Cymes in spikelike compound
inflorescences ......................... 19d. var. spicatiformis

RUBIACEAE

206

2b. Cymes not in spikelike

compound inflorescences.
3a. Shrubs with branches
tomentulose or pubescent
when young, glabrescent
when old .................................... 19c. var. pilosa
3b. Shrubs sparsely pilose ...... 19b. var. glabrescens
19a. Leptodermis pilosa var. acanthoclada H. S. Lo ex X. Y.
Wen & Q. Lin, Acta Phytotax. Sin. 45: 410. 2007.
ci zhi ye ding xiang
Shrubs dwarf, much branched; branches spiny.
Thickets. W Sichuan (Xiangcheng, Xiaojin), Xizang.

19b. Leptodermis pilosa var. glabrescens H. J. P. Winkler,

Repert. Spec. Nov. Regni Veg. 18: 160. 1922.
guang ye ye ding xiang
Leptodermis mairei H. Lveill.
Shrubs sparsely pilose.
S Sichuan (Dechang, Ningyuan, Xichang), Yunnan (Baiyanjing,
Dongchuan, Lijiang).

19c. Leptodermis pilosa var. pilosa

() chuan dian ye ding xiang (yuan bian
zhong)
Leptodermis fusca H. J. P. Winkler; L. microphylla (H. J. P.
Winkler) H. J. P. Winkler; L. pilosa var. microphylla H. J. P.
Winkler.
Shrubs with branches tomentulose or pubescent when
young, glabrescent when old.
Thickets on roadsides or on sunny slopes; 6003800 m. W
Hubei, S Shaanxi (Hanzhong, Hua Shan), Sichuan, SE Xizang, C and
NW Yunnan (Kunming, Lijiang).

19d. Leptodermis pilosa var. spicatiformis H. S. Lo ex X. Y.

Wen & Q. Lin, Acta Phytotax. Sin. 45: 411. 2007.
sui hua ye ding xiang
Leaves to 1.7 cm. Cymes often in spikelike compound
inflorescences.
Thickets on roadsides on mountain slopes; ca. 800 m. S Gansu
(Huixian, Liangdang, Tianshui), SW Shaanxi (Baoji, Fengxian, Leyang).

20. Leptodermis potaninii Batalin, Trudy Imp. S.-Peterburgsk.

Bot. Sada 14: [319]. 1898 [potanini].
ye ding xiang
Shrubs, 0.52 m tall or taller; branches pale gray, young
branches usually reddish, with 2 bands of soft hairs. Leaves
sparsely arranged or slightly crowded; petiole short; blade thin,
ovate or lanceolate, sometimes oblong or elliptic, both surfaces
white pubescent, tomentose, subglabrous, or glabrous, base cuneate, margins entire, apex obtuse to subrounded, apiculate; lateral veins 3 or 4 pairs, prominent abaxially, reticulate veins con-

spicuous; stipules membranous, broadly triangular, acuminate,

with acicular apiculus. Cymes terminal, sessile, 3-flowered,
rarely reduced to 1 or 2 flowers, central flowers sessile, lateral
ones pedicellate; pedicels red, with 2 bands of stiff or soft hairs;
bracteoles 2, longer than calyx tube, connate, densely hispid
or pilose outside, acute to aciculate. Calyx tube narrowly obconical, densely hispid or pilose on upper portion and calyx
lobes; lobes 5 or 6, narrowly triangular, length 3 size of
width, ciliate, acute. Corolla funnelform, to 1.5 cm; tube pilose or subglabrous outside, densely hispid on upper portion
and throat inside, limb spreading, 3 shorter than corolla tube;
lobes 5 or 6, with valvate aestivation, transparent, glabrous,
margins membranous, apex rounded. Stamens 5 or 6, attached
to upper portion of corolla tube, glabrous; filaments longer than
anthers; anthers semi-exserted, linear-oblong. Pistils ca. 1/2 as
long as corolla; ovary 3-celled; stigmas 3 or 4. Capsule 5-valvate from apex to base; valves crowned by persistent calyx
lobes. Fl. May, fr. autumn and winter.
Thickets on hill slopes or at streamsides, mountains; 8002700
m. W Guizhou, W Hubei, S Shaanxi, W Sichuan, Yunnan.

1a. Leaves hairy.

2a. Leaves white pubescent ................ 20c. var. potaninii
2b. Leaves tomentose or long
hirsute .......................................... 20d. var. tomentosa
1b. Leaves glabrous.
3a. Leaves glaucous green ...................... 20b. var. glauca
3b. Leaves brown when dry ............ 20a. var. angustifolia
20a. Leptodermis potaninii var. angustifolia H. S. Lo, var.
nov.
xia ye ye ding xiang
Type: China. Yunnan: Luquan Xian (), 5th District,
Wumeng () town, A-sha-hei, 2000 m, 9 Jun 1952, P. I.
Mao 1185 (holotype, KUN; isotype, KUN).
Validating Latin diagnosis: that of Leptodermis potanini
[sic!] Batalin var. angustifolia Lo (H. S. Lo in W. C. Chen, Fl.
Reipubl. Popularis Sin. 71(2): 338. 1999).
Leaves narrowly lanceolate, brown when dry, both surfaces glabrous.
Thickets at streamsides; ca. 2000 m. NC Yunnan (Luquan).
This name was previously published by H. S. Lo (loc. cit.) but not
validly so because no type was indicated (Vienna Code, Art. 37.1).

20b. Leptodermis potaninii var. glauca (Diels) H. J. P.

Winkler, Repert. Spec. Nov. Regni Veg. Beih. 12: 490. 1922.
fen l ye ding xiang
Leptodermis glauca Diels, Notes Roy. Bot. Gard. Edinburgh 5: 275. 1912; L. esquirolii H. Lveill; L. motsouensis H.
Lveill.
Leaves glaucous green abaxially. Corolla and bracteoles
glabrous.
Mountains; 8002700 m. SW Guizhou (Xingyi), SW Sichuan,
Yunnan.

RUBIACEAE

20c. Leptodermis potaninii var. potaninii

() ye ding xiang (yuan bian zhong)
Leptodermis nigricans H. J. P. Winkler.
Leaves white pubescent on both surfaces, though usually
subglabrous and pale abaxially.
Thickets on hill slopes; 8002400 m. W Guizhou, W Hubei
(Shennongjia, Xingshan), S Shaanxi (Pingli), W Sichuan (Kangding),
Yunnan.

207

810 mm; tube slender, slightly curved, throat slightly dilated,

densely pilose outside, sparsely villous inside; lobes 5, ovatelanceolate, ca. 2 mm, sparsely pilose, subobtuse. Stamens 5, inserted in corolla tube throat; anthers linear, ca. 2 mm, included
in long-styled flowers, slightly exserted in short-styled flowers.
Style slender, exserted in long-styled flowers, reaching middle of tube in short-styled flowers; stigmas 5-lobed, lobes linear. Capsules ca. 5 mm; seed aril reticulate, adherent to testa.
Fl. JulAug, fr. SepOct.
Mountain slopes; 8001000 m. S Gansu, N Sichuan (Songpan).

20d. Leptodermis potaninii var. tomentosa H. J. P. Winkler,

Repert. Spec. Nov. Regni Veg. 18: 153. 1922.

23. Leptodermis rehderiana H. J. P. Winkler, Repert. Spec.

Nov. Regni Veg. 18: 157. 1922.

rong mao ye ding xiang

bai mao ye ding xiang

Leptodermis potaninii var. rufa H. Winkler; L. tongchouanensis H. Lveill.

Shrubs, 12 m tall, very much branched, white villous or

pilose; branchlets slender. Petiole to 1 cm; leaf blade papery,
brownish gray when dry, lanceolate-oblong or ovate-oblong,
sometimes subovate, 14 ca. 2 cm, base attenuate, apex acute
or shortly acuminate, sometimes obtuse; lateral veins 25 pairs,
slightly prominent abaxially; stipules stiff, ca. 2 mm, acicularacuminate on lower portion, shortly acuminate on upper portion. Flowers terminal on foliate branches, 57 flowers arranged
in a line; central flowers sessile, lateral ones conspicuously
pedicellate, branchlets usually with 2 or 3 flowers axillary on
upper 4 or 5 nodes, inflorescence-like on lower 1 or 2 nodes,
with peduncle; bracteoles transparent, oblong, 2.53 mm,
slightly longer than calyx, ca. 2/3 connate, upper portion very
shortly pilose, other parts glabrous, veined, subulate-acuminate.
Calyx 2.52.8 mm; lobes 5, very short, ciliate, with longitudinal veins. Corolla tube very narrowly funnelform, 1213 mm,
slightly curved, densely shortly tomentose outside; lobes 5, subovate, ca. 2 mm, central portion and narrow margins not differentiated conspicuously, apex incurved. Stamens of shortstyled flowers slightly exserted; filaments inserted at 1/41/3
of anthers, free portion very short. Stigmas 5-lobed. Capsules
black, 67 mm; seed aril reticulate, adherent to testa. Fl. Sep, fr.
OctNov.

Branches, leaves, stipules, bracteoles, and flowers tomentose or long hirsute.

Mountains; ca. 2600 m. SW Sichuan, C, NE, and NW Yunnan.

21. Leptodermis pumila H. S. Lo, J. Trop. Subtrop. Bot. 7(1):

18. 1999.
ai xiao ye ding xiang
Dwarf shrubs, 510 cm tall; taproots stout, lignose, brown;
branches relatively short, pubescent, with many nodes. Leaves
crowded on upper portion of short branches; petiole 12 mm;
blade papery, elliptic or oblong, sometimes subobovate, 515
37 mm, pilose on midrib and lateral veins on both surfaces,
base cuneate, apex acute; lateral veins 3 or 4 pairs, slightly
prominent abaxially; stipules thick, triangular, 1.52 mm,
densely pilose and ciliolate. Flowers sessile, 3 fascicled at tips
of short branches; bracteoles 2, connate below middle portion,
upper portion free, subtriangular, slightly shorter than or subequal to calyx in length, pilose. Calyx tube obconical, ca. 1.8
mm; lobes 5, subovate, ca. 1 mm, ciliolate, acute. Corolla pale
red, funnelform; tube 1.11.2 cm, densely pilose outside, villous inside; lobes 5, ca. 3.5 mm, with thin and broad margins.
Stamens 5, inserted below throat of corolla tube; filaments
short; anthers linear, ca. 3 mm, slightly exserted. Style 78 mm;
stigmas 3, filiform, ca. 2 mm. Fl. May.
Grassy slopes; ca. 3000 m. NW Yunnan (Lijiang).

22. Leptodermis purdomii Hutchinson in Sargent, Pl. Wilson.

3: 405. 1916.
gan su ye ding xiang
Shrubs, 12 m tall; branchlets slender, puberulent when
young, soon glabrescent. Leaves fascicled, papery, linear-oblanceolate, 0.51 1.53.5 mm, both surfaces glabrous, base
attenuate, margin obviously recurved, apex obtuse; nerves inconspicuous; stipules ovate, ca. 1.5 mm. Flowers sessile or subsessile, fascicled on apex of branches; bracteoles 2, opposite,
transparent, membranous, ovate, ca. 1.5 mm, shorter than calyx, connate at base, subglabrous (with white acicular lines?),
with carina on dorsal surface, abruptly acute at apex. Calyx
tube glabrous; lobes 5, oblong-ovate, ca. 1.5 0.75 mm, leathery, ciliate, triangular-acute. Corolla pink, narrowly funnelform,

Thickets, jungles; 16002400 m. NW Yunnan (Zhongdian).

24. Leptodermis scabrida J. D. Hooker, Fl. Brit. India 3: 199.

1881.
cao ye ye ding xiang
Shrubs, 11.5 m tall; branches stout, brown or dark gray,
short branches often with dense persistent scaly stipules, pilose.
Petiole short or leaves subsessile; leaf blade oblong-lanceolate
or oblong, sometimes narrowly lanceolate, 13(4) 410 mm,
both surfaces ?strigose or only midrib and lateral veins hairy
abaxially, base cuneate or decurrent, apex acute to acuminate;
midrib prominent abaxially; lateral veins 3 or 4 pairs, inconspicuous; stipules subtriangular, pilose, with a long and excurved stiff apiculus. Flowers subsessile, 35 terminal or axillary near tips of branches; bracteoles transparent, 34 mm, pilose, ca. 2/3 connate, free portion triangular-ovate, with a stiff
apiculus at apex. Calyx tube ca. 2 mm; lobes 0.50.7 mm,
width greater than length or subequal, densely ciliate, obtuse.
Corolla white, funnelform, 919 mm, pilose outside; lobes 5,
sublanceolate, 22.5 mm. Short-styled flowers with stamens

RUBIACEAE

208

inserted below throat of corolla tube, anthers linear, included;

style slightly exserted, stigmas 3-lobed, filiform, excurved. Capsule 56 mm; seed aril free from testa (according to literature).
Fl. Jun.
Forests, hill slopes; 24002600 m. S Xizang (Nyalam, Zay) [NE
India (Assam)].

25. Leptodermis schneideri H. J. P. Winkler, Repert. Spec.

Nov. Regni Veg. 18: 156. 1922.
xian zhi ye ding xiang
Leptodermis schneideri var. hutchinsonii H. J. P. Winkler.
Shrubs, 0.21.3 m, branches slender; young branches
tomentulose, soon glabrescent, old branches with gray longitudinally fissured bark. Leaves sparsely arranged on long
branches, very crowded on very short branches in axils of long
branches, appearing as if leaves whorled; petiole very short
or leaves subsessile; blade oblong or ovate, 49 1.252 mm,
glabrous on both surfaces, base attenuate, apex acute or slightly
obtuse or apiculate; midrib impressed adaxially and prominent
abaxially, lateral veins inconspicuous; stipules membranousleathery, small, ca. 1 mm, pilose and ciliate, upper portion triangular, lower part acicular acuminate, with 2 callose nerves near
margins extending to apex. Flowers terminal on lateral foliate
branches, upper 2 or 3 nodes with 13 flowers axillary; peduncles present or almost absent; bracteoles transparent, oblong, ca.
2 mm, slightly longer than calyx, connate to or above middle
portion, margins of upper portion hairy, apex rounded, apiculate. Calyx lobes 5, length slightly greater than width, ciliate,
rounded. Corolla small; tube narrowly funnelform, ca. 5 mm,
curved, covered with conical woolly hairs outside, densely covered with long hairs on throat inside, glabrous at base; lobes oblong, ca. 2 mm, covered with short papilla outside, villous at
base inside, with thin, soft, narrow margins, apex incurved. Stamens inserted in throat of corolla tube; long-styled flowers with
filaments short, anthers included; short-styled flowers with filaments long, anther exserted. Stigmas 5-lobed, conspicuously
exserted in long-styled flowers, extending below throat in shortstyled flowers. Capsules ca. 4 mm; seed aril adherent to testa.
Sichuan, Xizang, Yunnan.

26. Leptodermis scissa H. J. P. Winkler in Handel-Mazzetti,

Symb. Sin. 7: 1024. 1936.
si lie ye ding xiang
Shrubs, 2050 cm tall, occasionally to 1.5 m tall; branchlets gray or brown, tomentulose. Leaves crowded on branchlets,
blade papery, grayish when dry, oblong-ovate or broadly
ovate, 58 2.54 mm on lower branches, 1523 713 mm
on upper branches, adaxially densely scabrid on midvein,
slightly scabrid on lateral veins and margins, abaxially glabrous, base attenuate, apex apiculate; lateral veins 3 or 4 pairs,
straight; stipules submembranous, tomentulose outside, with
glandular appendages inside, long triangular on lower portion,
shortly triangular on upper portion, glandular acuminate upward, sometimes with 1 or 2 glandular teeth, with 2 veins confluent at apex, uppermost stipules short and broad, abruptly
acuminate, without veins. Flowers often 3 terminal on short

branches, with short pedicels or longer at each side, or 13

flowers axillary in axils of leaves at nodes; bracteoles ovate,
2.83 mm, free almost to base, midrib nearly brownish, shortly
hairy, other portions subglabrous, margins imbricate, ciliate,
apex acuminate, with apiculus almost as long as calyx or shorter.
Calyx 33.2 mm; lobes 5, triangular, usually scattered pilose,
ciliate. Corolla tube funnelform, ca. 11 mm, slightly curved,
glabrous or puberulent outside, long villous inside, glabrous
near base; lobes ovate-orbicular, ca. 2 mm, glabrous outside.
Stamens 5 in short-styled flowers, inserted in throat of corolla
tube; anthers linear, semi-exserted, covered with glandular
long soft hairs inside. Style 5-lobed, included, glabrous. Fruit
ovoid, ca. 4 mm; seed aril conspicuously free from testa. Fl.
JulAug, fr. Oct.
Thickets; 15002500 m. Sichuan, Yunnan.

27. Leptodermis tomentella H. J. P. Winkler, Repert. Spec.

Nov. Regni Veg. 18: 159. 1922.
meng zi ye ding xiang
Shrubs; stems, branches, and both surfaces of leaves tomentulose; branchlets stout, terete, upper branchlets short,
lower branchlets longer, secondary branchlets very short. Petiole short and thick; leaf blade thickly papery, brownish when
dry, ovate or elliptic, 519 2.510 mm, base broadly cuneate,
apex acute, with a short stiff apiculus; lateral veins 2 or 3(or 4)
pairs, inconspicuous adaxially, slightly prominent abaxially;
stipules triangular, 11.5 mm, with stiff acicular apiculus. Flowers 37 fascicled at tips of branchlets, subsessile; bracteoles
ovate, ca. 3 mm, ca. 1/3 connate, tomentose, with acicular apiculus slightly shorter than calyx lobes. Calyx tubes black when
dry; lobes pale brown, ovate-oblong, ca. 1.5 mm, ciliate,
slightly obtuse. Corolla purple or white, funnelform, 1.21.3
cm, tomentose outside, throat villous; lobes with margins broad,
soft and thin. Stamens 5, inserted in throat of corolla tube; filaments very short; anthers linear, ca. 2.5 mm. Style exserted; stigmas 5-lobed, lobes linear. Capsules ca. 5 mm; seed aril reticulate, adherent to testa. Fl. MayJun, fr. Sep.
Mountain slopes; 15002000 m. C and S Yunnan (Chengjiang,
Mengzi).

28. Leptodermis umbellata Batalin, Trudy Imp. S.-Peterburgsk. Bot. Sada 13: 374. 1894.
san hua ye ding xiang
Shrubs, to 2 m, much branched; young branches, peduncles, and pedicels covered with white stiff hairs. Leaves sparsely
arranged; petiole usually 46 mm; blade papery, black adaxially
and usually pale or brownish abaxially when dry, elliptic or
oblong, sometimes obovate, 1035 516 mm, both surfaces
shortly hispid, base cuneate, attenuate, apex acute or shortly
acuminate; lateral veins sparse, usually 2 or 3 pairs, inconspicuous; stipules broadly triangular, hispid. Flowers with pedicels straight, 39 mm, arranged in umbelliform groups, rarely
13 flowers axillary; peduncles to 1 cm or longer; bracteoles ca.
3 mm, subequal to calyx tube in length or slightly shorter, ca.
2/3 connate, upper portion bilabiate dehiscent, hispid. Calyx
lobes suborbicular, width slightly greater than length, ciliate,
apex slightly rounded or subtruncate. Corolla purple, narrowly

RUBIACEAE

funnelform, ca. 12 cm, densely hispid outside, slightly hairy inside; lobes 5, oblong, ca. 3 mm, with incurved abrupt apiculus.
Long-styled flowers: stamens inserted above throat of corolla
tube, filaments ca. 1.5 mm, anthers linear, ca. 2.5 mm, exserted.
Short-styled flowers: stigmas 5-lobed, lobes linear, included.
Capsule 45 mm; seed aril reticulate, free from testa or adnate
with testa on ventral side. Fl. SepOct.
Hill slopes; 500700 m. S Gansu (Bikou, Wenxian), N Sichuan
(Qingchuan).

29. Leptodermis velutiniflora H. S. Lo, J. Trop. Subtrop. Bot.

7(1): 17. 1999.
mao hua ye ding xiang
Shrubs, 0.51 m tall; branchlets terete, slender and firm,
densely pubescent. Petiole 36 mm, densely pilose; leaf blade
papery or thinly papery, when dry turning black adaxially, pale
brown abaxially, elliptic to oblong or ovate to lanceolate, 1040
520 mm, scattered with sparse appressed pubescence adaxially, villous on midrib and lateral veins abaxially, base cuneate
to broadly cuneate, apex acute or acuminate; lateral veins 4 or 5
pairs, slender, conspicuous abaxially; stipules ca. 1.5 mm,
broadly triangular at base, sometimes with stipitate glands on
margins, with a stiff acicular apiculus at apex. Short-styled
flowers 25 fascicled on tips or axillary on upper portion of
branchlets; pedicels usually 26 mm, sometimes up to 8 mm,
pilose; bracteoles 2, small, ca. 1.2 mm, ca. 1/3 of lower portion
connate, lobes triangular, ca. 7 mm, pubescent. Calyx tube ca. 2
mm, glabrous; lobes 5, triangular, ca. 0.4 mm, margins hirsute,
apex acute. Corolla funnelform or broadly funnelform, ca. 0.4
mm, brown tomentose outside, white villous on throat inside, or
glabrous on both sides; lobes 5, triangular-oblate, 33.5 ca. 4
mm, margins thin and broad, apex obtuse. Stamens 5, inserted
in throat of corolla tube; filaments ca. 2 mm; anthers ca. 2.5
mm, exserted. Style extended to middle of corolla tube; stigmas
4-lobed, lobes 0.50.6 mm. Fl. MayJun.
Forests, forest margins; 28003100 m. Sichuan, SE Xizang,
NW Yunnan.

1a. Leaves thinly papery; corolla glabrous ...... 29a. var. tenera
1b. Leaves papery; corolla brown
tomentose outside, white villous
on throat inside ................................. 29b. var. velutiniflora
29a. Leptodermis velutiniflora var. tenera H. S. Lo, J. Trop.
Subtrop. Bot. 7(1): 17. 1999.
bao ye ye ding xiang
Leaves thinly papery. Corolla glabrous. Fl. MayJun, fr.
Aug.
Forests, forest margins; 28003100 m. SW Sichuan (Muli, Yanyuan), SE Xizang (Bomi, Mainling, Nyingchi), NW Yunnan (Binchuan,
Weixi).

29b. Leptodermis velutiniflora var. velutiniflora

() mao hua ye ding xiang (yuan bian
zhong)
Leaves papery. Corolla brown tomentose outside, white
villous on throat inside.

209

Sichuan.

30. Leptodermis vestita Hemsley, J. Linn. Soc., Bot. 23: 390.

1888.
guang dong ye ding xiang
Leptodermis ovata H. J. P. Winkler.
Shrubs, with a few ascending branches, to 0.6 m tall;
young branches and leaves slightly to densely squarrose hirsute,
glabrescent. Petiole stout, 0.40.7 cm, slightly to densely hairy,
sometimes leaves sessile; leaf blade subleathery, iron-gray abaxially when dry, ovate, ovate-oblong, elliptic, or sometimes suborbicular, 1.54 12.2 cm, adaxially glabrous or hirsutulous
on midrib, abaxially hirsutulous on midrib and lateral veins or
glabrous, base attenuate or cuneate, margin slightly revolute,
apex shortly acuminate, acute, or obtuse; lateral veins 47 pairs,
together with veinlets adaxially inconspicuous, abaxially conspicuous; stipules broadly triangular, 0.81.5 mm, sometimes
inconspicuous, apex apiculate. Flowers 37 fascicled, terminal
or in axils of upper 36 pairs of leaves, subsessile or pedicel
0.32.6 mm; bracteoles rigid, hyaline, oblong, 2.53.5 mm,
longer than calyx or subequal at anthesis, ca. 2/3 portion connate, veined, subulate-acuminate. Calyx tube 1.33.3 mm, glabrous or slightly hairy outside; lobes 5, ca. 0.5 mm, width
greater than length, longitudinally veined, margins slightly imbricate, ciliate, apex subrounded. Corolla white, purple, or red,
narrowly funnelform, ca. 1.5 cm, slightly curved, tomentose to
sparsely pubescent; lobes 5, oblong, 1.41.8 mm, central portion thicker than narrow margins, apex incurved. Stamens 5.
Short-styled flowers: stamens inserted above throat of corolla
tube, filaments 1.33.2 mm, anthers 1.22 mm, slightly exserted; style 6.59.7 mm, stigma 0.31.8 mm, 36-lobed. Longstyled flowers: stamens inserted at throat of corolla tube,
filaments ca. 0.7 mm, anthers 1.62 mm, included; ovary 5celled, ovule 1 per cell; style 11.215.3 mm, stigma 0.61.8
mm, 35-lobed, slightly exserted. Capsule obovoid, 5.27.6
mm; seeds narrowly ellipsoid to ellipsoid or narrowly obovoid
to obovoid, 3.35.5 mm; aril reticulate, free from testa. Fl. Jun
Dec, fr. AugJan.
N Guangdong (Lianxian, Ruyuan), Guangxi.

31. Leptodermis wilsonii Diels, Notes Roy. Bot. Gard. Edinburgh 5: 275. 1912.
da guo ye ding xiang
Shrubs, usually 0.51 m tall; branches slightly stout,
with brown or grayish white bark and 2 longitudinal grooves,
grooves pilose. Petiole 24 mm or slightly longer; leaf blade
papery, black or dark gray adaxially when dry, light brown
abaxially, ovate or ovate-elliptic, sometimes lanceolate or oblong-lanceolate, 13.5(3.9) 0.51.8(2) cm, glabrous on
both surfaces except midrib and margins pilose adaxially,
slightly smooth, base broadly cuneate, apex acute or obtuse;
lateral veins 3 or 4 pairs, inconspicuous adaxially, not or
slightly prominent abaxially; stipules long triangular, ca. 2
mm, with stiff apiculus, margins usually glandular. Flowers
subsessile, usually 3 flowers terminal on branches, occasionally
axillary near tips of branches; bracteoles 2, subscarious, lanceolate or ovate-triangular, 22.5 mm, usually slightly shorter than

RUBIACEAE

210

sinus of calyx limb, glabrous or pilose, acuminate. Calyx black

when dry; tube ca. 2.5 mm; lobes narrowly triangular, ca. 1.8
mm, shortly ciliate, acuminate, slightly spreading. Corolla white
or pallid red, fragrant, funnelform, smooth outside, glabrous;
tube 1214 mm, villous inside; lobes spreading, suborbicular,
46 mm in diam., central portion relatively thick, margins broad
and thin, erose, apex callose acute. Stamens 5, inserted in throat
of corolla tube in short-styled flowers; filaments ca. 3 mm; anthers linear, ca. 4 mm, slightly exserted. Style dark brown or
black when dry, linear, 78 mm, glabrous. Seed aril reticulate,
adherent to testa. Fl. Jun, fr. OctNov.
Jungles, thickets; 18003000 m. W Sichuan (Shimian), NW
Yunnan (Dali, Heqing, Lijiang).

32. Leptodermis xizangensis H. S. Lo, J. Trop. Subtrop. Bot.

7(1): 16. 1999.
xi zang ye ding xiang
Shrubs, 5080 cm tall; branches terete, slightly stout, purplish black, glabrous or pubescent, short branch usually ca. 1
cm, with dense nodes and short internodes. Leaves arranged on
short branches; petiole short or leaves subsessile; blade thickly
papery or slightly fleshy, black adaxially and pale brown abaxially when dry, ovate-elliptic to lanceolate-oblong, 0.71.2
46 mm, both surfaces glabrous except margin ciliate, base
broadly cuneate, apex acute or obtuse; midrib conspicuous
abaxially, lateral veins sparse, inconspicuous; stipules long triangular, 22.5 mm, cuspidate-caudate, sometimes both sides
adnate to petiole. Flowers sessile, usually 3 flowers terminal on
tips of short branches; bracteoles 2, scarious, transparent, ca.
4.5 mm, connate to middle portion, apex with an acicular apiculus. Calyx tube ca. 2.5 mm; lobes 5, triangular, ca. 0.6 mm,
densely ciliate, acute. Corolla purple, turning black when dry,
broadly funnelform, ca. 1.6 mm, glabrous outside, throat expanded, ca. 5 mm in diam., sparsely villous or subglabrous inside; lobes 5, broadly ovate-suborbicular, ca. 4 mm, apex shallowly 3-lobed or cuspidate, apiculus obtuse. Stamens 5, inserted
below corolla tube throat; filaments very short; anthers lanceolate-linear, ca. 2 mm. Style ca. 12 mm; stigmas 3-lobed, ca. 2
mm, linear.
Thickets; ca. 3400 m. S Xizang (Gyirong).

33. Leptodermis yangshuoensis Tao Chen, sp. nov.

yang shuo ye ding xiang
Type: China. Guangxi: Yangshuo Xian (), Bilianfeng (), between rocks at roadsides or forest margins on
limestone hilltop, ca. 300 m, 16 Sep 1999, T. Chen & Y. K. Li
990916001 (holotype, SZG; isotypes, A, MO).
Haec species Leptodermidi vestitae Hemsley similis, sed
ab ea inflorescentiis ad ramos hornotinos dispositis terminalibus vel raro in axillis foliorum superiorum, pedunculo elongato atque pedicello manifesto usque ad 8 mm longo differt.
Paratype: China. Guangxi: Yangshuo Xian (),
Matang (), Longtouzhai (), 380 m, on limestone
hill, 20 Oct 1963, Z. Z. Chen () 53104 (IBK bar code
00100138).
Shrubs to 1 m high, with raphides distinctly present on

leaves, stipules, bracts, bracteoles, calyx, and corolla. Current

years branches subquadrangular, glabrous, ?straw-yellow when
dry; previous years branches with grayish white bark peeling
off; old branches dark brown. Leaves opposite and sometimes
in clusters on lateral short branches; petiole to 12 mm, grooved
adaxially; blade subleathery, slightly shiny when dry, adaxially
dark green, abaxially pale green, ovate to ovate-oblong, to 50
25 mm, glabrous on both surfaces, base obtuse, slightly decurrent to petiole, margins entire, slightly ciliate near apex, apex
acute or shortly acuminate; midrib impressed adaxially when
dry, lateral veins 36 pairs, slender, arcuate ascending, confluent along leaf margin, adaxially depressed or slightly impressed, abaxially prominent, veinlets abaxially conspicuous;
stipules broadly triangular, apiculate. Inflorescences thyrsoid,
borne on current years branches, terminal or rarely in axils of
upper leaves; peduncle purplish brown, elongate, slightly angular when dry, slender or slightly stout; bracts leaflike to subulate. Flowers (1)35 fascicled, terminal on ultimate branches
of inflorescence; pedicels purplish brown, to 8 mm; bracteoles
connate, membranous, ca. 3 mm, equal to or slightly longer
than calyx, distinctly veined, with raphides; lobes 2, triangular,
ca. 1.2 mm, slightly bilobed at apex. Calyx with hypanthium ca.
1.5 mm; tube ca. 0.3 mm; lobes 5, ovate-oblong, ca. 1.2 mm,
white ciliate. Corolla white or pale purple, narrowly funnelform, ca. 12 mm; tube narrow, 910 mm, densely hairy at
throat, with hairs becoming sparse toward base; lobes 5, ovatetriangular, ca. 3 mm, reticulately veined. Stamens 5, inserted at
throat; filaments very short; anthers ca. 1.5 mm. Ovary 5-celled,
each cell with 1 ovule, basal; style slender; stigma 5-lobed,
lobes linear, slender, revolute, adaxially hairy. Fruit a capsule,
ovoid, ca. 4 mm, septicidally dehiscing into 5 valves, 5-seeded;
valves persistent, with persistent calyx lobes at apex; seeds
black, ca. 2.5 mm; aril reticulate, free. Fl. AugOct, fr. Oct
Nov.
Limestone hills; 300400 m. NE Guangxi (Yangshuo).

34. Leptodermis yui H. S. Lo, J. Trop. Subtrop. Bot. 7(1): 15.

1999.
de jun ye ding xiang
Shrubs, 0.51.5 m tall; branches and branchlets slender,
old branches purplish red, glabrous, young branches with short,
soft hairs in 2 bands. Leaves sparsely arranged; petiole 0.30.6
cm; blade membranous, adaxially dark brown and abaxially
slightly pale when dry, broadly ovate to lanceolate, 1.54.5
0.72 cm, adaxially dispersedly strigose, abaxially glabrous or
with sparse stiff hairs on midrib, base broadly cuneate or cuneate, margins fimbriate ciliate, apex obtuse or subacute, sometimes rounded; midrib very compressed abaxially, lateral veins
slender, 4 or 5 pairs, conspicuous abaxially; stipules slightly
short on upper part of branchlets, ca. 1 mm, apex lacerate, glandular, triangular on lower portion of branchlets, ca. 2 mm,
margins with glandular hairs. Flowers 13 terminal, sessile;
bracteoles 2, base connate (ca. 1 mm), free portion lanceolate or
ensiform-lanceolate, ca. 2 mm, sparsely hispid, margins sometimes glandular. Calyx tube ca. 2 mm; lobes 5, narrowly triangular, longer than tube, hispid ciliate, acuminate. Corolla purple, broadly funnelform, apex with tufted long soft hairs; tube

RUBIACEAE

910 mm, glabrous outside, covered with white long soft hairs
inside; lobes 5, broad, oblate, ca. 5 mm wide, margin crisped,
both surfaces villous, apex shallowly 3-lobed. Stamens 5, inserted below throat of corolla tube; filaments short; anthers ob-

211

long-linear, ca. 2 mm, included. Style slightly exserted, stigmas

2-lobed, lobes filiform.
Forest margins; ca. 2500 m. SW Sichuan (Muli).

45. LEPTOMISCHUS Drake, Bull. Mus. Hist. Nat. (Paris) 1: 117. 1895.
bao chun qian shu
Chen Tao (); Charlotte M. Taylor
Indopolysolenia Bennet; Polysolen Rauschert, nom. illeg. superfl.; Polysolenia J. D. Hooker (1873), not Ehrenberg ex Ktzing
(1849) [Fungi].
Perennial herbs or subshrubs, unarmed, presumably rather succulent. Raphides present. Leaves opposite, sometimes crowded or
rosulate, perhaps without domatia, with margins sometimes crisped and/or reportedly serrulate; stipules persistent to deciduous, interpetiolar or shortly united around stem, triangular to suborbicular (Leptomischus funingensis), often well developed and veined, entire
or 2- or 3-lobed (L. wallichii), glabrous or glabrescent adaxially. Inflorescences terminal and apparently sometimes pseudoaxillary,
capitate to congested-cymose or umbelliform, several to many flowered, subsessile to pedunculate, bracteate with bracts often well
developed to involucral. Flowers sessile to pedicellate, bisexual, distylous and sometimes markedly dimorphic. Calyx limb 5-lobed.
Corolla white or yellow, funnelform, salverform, or sometimes inflated, inside pubescent in upper part or most of tube; lobes 5,
valvate in bud. Stamens 5, inserted below middle of corolla tube in long-styled flowers, inserted in corolla throat in short-styled
flowers, included or partially exserted; filaments usually short; anthers basifixed or perhaps dorsifixed near base. Ovary 2-celled,
ovules numerous in each cell borne on stipitate placentas apparently near base of septum; stigma 2-lobed, included or exserted. Fruit
capsular, subglobose, dehiscent through apical portion or operculum inside calyx limb, perhaps papery, with septum reportedly
mostly disintegrating leaving 1 incomplete cell; seeds numerous, small, reticulate or areolate.
Seven species: S China, NE India, Myanmar, Vietnam; five species (three endemic) in China.
Leptomischus seems to be similar to Keenania and Mouretia; all of these genera are rather poorly known. Several authors (e.g., Deb & Rout,
Kew Bull. 45(2): 339341. 1990; H. S. Lo in FRPS 71(1): 184189. 1999) have reported marked floral dimorphism in species of Leptomischus.

1a. Leaf blade shallowly cordate or subtruncate at base; stems well developed with leaves distributed along them;
calyx with multicellular long trichomes .................................................................................................................. 3. L. guangxiensis
1b. Leaf blade obtuse to acute at base; stems short to well developed with leaves distributed along them or
congested at ends; calyx glabrous or with unicellular or multicellular trichomes.
2a. Flowers smaller, corolla 66.5 mm; calyx and corolla glabrous outside ............................................................ 4. L. parviflorus
2b. Flowers larger, corolla longer than 10 mm; calyx and corolla glabrous to variously pubescent outside.
3a. Stems short, with leaves crowded and rosulate; calyx and corolla unicellular pilosulous outside ............ 5. L. primuloides
3b. Stems developed, with leaves distributed along them; calyx and corolla densely multicellular
villous, -tomentose, or -pilosulous outside.
4a. Stipules ovate to lanceolate, 89 mm ........................................................................................................ 1. L. erianthus
4b. Stipules suborbicular, 45 mm ............................................................................................................... 2. L. funingensis
1. Leptomischus erianthus H. S. Lo, Bull. Bot. Res., Harbin
18: 277. 1998.
mao hua bao chun qian
Herbs, weak to erect, 12 m tall; stems quadrangular,
densely hirtellous. Leaves opposite, distributed along stems;
petiole usually 0.51.5 cm, densely hirtellous; blade drying papery, black adaxially, dark brown abaxially, narrowly elliptic to
lanceolate-elliptic, 412 1.54 cm, adaxially sparsely strigose
to glabrescent, abaxially glabrescent except multicellular villous along principal veins, base cuneate, margins densely multicellular-ciliate, apex acuminate or caudate-cuspidate; secondary
veins 912 pairs; stipules persistent, ovate or lanceolate, 89
mm, densely striate veined. Inflorescences terminal, subcapitate, densely multicellular villous with trichomes drying brown;
peduncle shorter than 1 cm. Flowers subsessile. Calyx multicellular villous; hypanthium portion obconic, ca. 1.5 mm; limb
lobed for ca. 1/2; lobes narrowly triangular, ca. 4 mm, acute.

Corolla white, tubular, densely multicellular tomentose or -pilosulous outside with trichomes drying brown; tube 1516 cm,
inside upper part white villous; lobes triangular-lanceolate, 45
mm, acute. Capsules obconic, 56 mm, with persistent calyx
lobes to 3 mm. Fl. May, fr. Jul.
Dense forests in moist valleys; 15001700 m. Yunnan.

2. Leptomischus funingensis H. S. Lo, Bull. Bot. Res., Harbin

18: 278. 1998.
fu ning bao chun qian
Herbs, unbranched; stems terete to quadrangular, pilosulous. Leaves opposite, distributed along stems; petiole 13 cm;
blade drying black-brown, ovate, narrowly elliptic, or rarely
obovate, 815 2.54.5 cm, adaxially glabrescent to pilose,
abaxially villous, base cuneate, margins entire or serrulate-ciliolate, apex acuminate; secondary veins 57(or 8) pairs; stipules
suborbicular, 45 mm, usually reflexed, villous abaxially

RUBIACEAE

212

and marginally. Inflorescences terminal, cymose, densely many

flowered; peduncles shorter than 1 cm; pedicels 33.5 mm.
Flowers pedicellate. Calyx densely multicellular villous; hypanthium portion obconic, 3.54 mm; limb deeply lobed; lobes
subtriangular, ca. 1.5 mm, obtuse. Corolla white or bright yellow, tubular-salverform, densely multicellular villous outside;
tube 1416 mm, white villous inside; lobes subelliptic, 46
mm, obtuse. Capsules obconic or subglobose, pilose, weakly
shiny. Fl. summer.
Streamsides in dense forests; ca. 1000 m. Yunnan (Funing).

3. Leptomischus guangxiensis H. S. Lo, Bull. Bot. Res., Harbin 18: 278. 1998.

414 mm, veined, glabrescent, acute to obtuse. Inflorescences

terminal, capitate, several flowered, puberulent to glabrous; peduncle 0.45 cm; bracts obovate to oblong-oblanceolate or narrowly oblanceolate, 310 mm, entire or sparsely ciliate. Flowers subsessile. Calyx glabrous; hypanthium portion obovoid,
smooth to 5-ribbed, ca. 1.5 mm; limb lobed for ca. 1/2; lobes
ovate to ligulate, 11.2 mm, obtuse. Corolla white, funnelform,
66.5 mm, glabrous outside; tube densely white villous in
throat; lobes subtriangular, margins thickened, apex rostrate.
Capsules subglobose, 2.53.5 34.5 mm, glabrous; seeds 0.2
0.4 mm. Fl. JulAug, fr. Aug.
Dense forests. Hainan (Sanya), Yunnan (Hekou) [N Vietnam].

xin ye bao chun qian

5. Leptomischus primuloides Drake, Bull. Mus. Hist. Nat.

(Paris) 1: 117. 1895.

Herbs; stems densely multicellular tomentose with trichomes drying grayish brown. Leaves opposite, distributed
along stems; petiole 2.53.5 cm, densely pilosulous; blade
drying papery and brownish gray adaxially, ovate or oblongovate, 58.5 2.54.5 cm, sparsely strigose adaxially, pilosulous except multicellular tomentose along midrib abaxially,
base shallowly cordate to subtruncate, margins entire, apex
acute to obtuse; secondary veins 57 pairs; stipules subovate,
79 mm, villous. Inflorescences terminal, cymose, densely
multicellular tomentose; peduncle ca. 1 cm. Flowers not seen.
Capsules obconic, multicellular villous, together with persistent calyx lobes 910 mm. Fl. May, fr. Apr, Jan.

bao chun qian

Shady grasslands near water. Guangxi.

4. Leptomischus parviflorus H. S. Lo, Bull. Bot. Res., Harbin

6(4): 49. 1986.
xiao hua bao chun qian
Herbs, 1030 m tall; stems terete to 4-angled, densely villosulous. Leaves opposite, distributed along stems or in clusters
near stem apices; petiole 410(20) mm, puberulent to villosulous; blade drying papery, dark green adaxially, greenish yellow
abaxially, elliptic, narrowly elliptic, or oblanceolate, 414 14
cm, glabrous adaxially, glabrous on lamina and densely puberulent to villosulous on veins abaxially, base acute to obtuse, margins entire, apex obtuse to shortly acuminate; secondary veins
1216 pairs; stipules persistent, rather leaflike, ovate to ligulate,

Indopolysolenia burmanica Deb & Rout.

Perennial herbs or subshrubs, 1030 cm tall; stems stout,
subterete to quadrangular, pilosulous. Leaves opposite, crowded
and rosulate; petiole 0.44 cm, pilose; blade drying papery,
obovate, broadly obovate, or elliptic, (7)1525 (2)510
cm, adaxially glabrous, abaxially glabrescent except puberulent
along principal veins, base cuneate to acute, margins entire,
apex acute or obtuse; secondary veins 1419 pairs; stipules
deciduous, elliptic, lanceolate, or ovate, 618 mm, pilosulous,
acute or acuminate. Inflorescences terminal and/or pseudoaxillary in uppermost axils, capitate to subcapitate, several flowered, glabrescent; peduncle 2.512 cm; heads 25 cm wide;
bracts ovate, 614 mm, pilose, acuminate or acute. Flowers
subsessile. Calyx unicellular pilosulous; hypanthium portion
ellipsoid-obovoid, 11.5 mm; limb lobed for ca. 1/2; lobes lanceolate, (1)2.5(4) mm, ciliate, acute or acuminate. Corolla
funnelform or tubular to inflated, outside pilosulous; tube 16
18(25) mm, villous in throat; lobes triangular, ca. 2 mm, acute
or obtuse. Capsules obovoid, ca. 3 4 mm. Fl. early winter.
Forests; [ca. 200 m in Myanmar]. Yunnan (Hekou) [Myanmar,
Vietnam].
The measurements above include some from the protologue of
Indopolysolenia burmanica, which was placed by Lo (Acta Phytotax.
Sin. 31: 275. 1993) in synonymy with Leptomischus primuloides.

46. LEPTUNIS Steven, Bull. Soc. Imp. Naturalistes Moscou 29(2): 366. 1857.
le tu cao shu
Chen Tao (); Friedrich Ehrendorfer
Herbs, annual, slender. Raphides present. Stems subterete. Leaves and leaflike stipules in whorls of 816, sessile, linear to filiform, without domatia. Inflorescences terminal, thyrsoid, below with dichasial and above with monochasial cymes, few flowered
when young but usually continuing to grow and becoming many flowered and diffuse; peduncles and pedicels filiform with leaflike
bracts and reduced bracteoles. Flowers pedicellate to sessile, bisexual, monomorphic. Calyx limb obsolete. Corolla salmon-orange to
pale greenish, tubular to funnelform, glabrous inside; lobes 4, valvate in bud. Stamens 4, inserted in upper part of corolla tube,
included or partially exserted; filaments short; anthers dorsifixed, black. Ovary inferior, 2-celled, ovules 1 in each cell, basal; stigmas
2, globose, partially exserted. Fruit schizocarpous; mericarps obovoid, with 1 seed, indehiscent, upper part somewhat incurved, dry,
with appressed short and curved hairs.
One species: from the SE Caucasus through Afghanistan and C Asia to NW China.
Ehrendorfer et al. (Fl. Iranica 176: 1287. 2005) included the monotypic genus Leptunis in Asperula sect. Trichodes Boissier, based on the fol-

RUBIACEAE

213

lowing arguments. Segregation of Leptunis as a monotypic genus from Asperula was based only on its ripe mericarps being rather obovoid and
incurved than ellipsoid and straight. Furthermore, Leptunis appears closely linked to Asperula by A. seticornis Boissier. This local species from S Iran
has long been recognized as having fruit that are intermediate in shape between Leptunis and typical Asperula. Nevertheless, considering the still
incomplete DNA-analytical study of SW-Asiatic taxa of Asperula and the recent discovery of Leptunis in China (Abdusalih et al., Acta Bot. Boreal.Occid. Sin. 23(4): 674. 2003), the genus is provisionally retained here. It is inserted in the key to Rubieae genera and species found in the present
volume under Galium.

1. Leptunis trichodes (J. Gay ex Candolle) Schischkin, Fl.

URSS 23: 285. 1958 [trichoides].
le tu cao
Asperula trichodes J. Gay ex Candolle, Prodr. 4: 582.
1830; Leptunis tenuis Steven.
Herbs, annual, erect, 1040 cm tall; stems and branches
subterete, glabrous. Leaf blade drying stiffly papery, linear, 10
35 0.31 mm, glabrous to sparsely scaberulous, base acute,
margins usually revolute, apex acute; 1 main vein as midrib,
secondary veins not visible. Inflorescence 115 120 cm,

glabrous; bracts leaflike, 13.5 0.10.2 mm; pedicels lacking

or up to 20 mm, thickened toward flowers. Corolla puberulent
outside, 1.52 mm; lobes elliptic to ovate, 0.51 mm, obtuse.
Ovary inferior, obovoid, ca. 1 1 mm. Mericarps 12 mm, with
appressed short and curved hairs. Fl. JunJul, fr. JulAug.
Open ground and grasslands on mountains, riversides, sunny rocky
slopes; 9001500 m. Xinjiang (rmqi) [Afghanistan, Kazakhstan,
Kyrgyzstan, Tajikistan, Turkmenistan, Uzbekistan; SW Asia (Azerbaijan, Iran)].
Leptunis trichodes was first documented from China by Abdusalih
et al. (Acta Bot. Boreal.-Occid. Sin. 23(4): 674. 2003).

47. LERCHEA Linnaeus, Mant. Pl. 2: 155, 256. 1771, nom. cons., not Lerchia
Haller ex Zinn (1757).
duo lun cao shu
Chen Tao (); Charlotte M. Taylor
Codaria Linnaeus ex Kuntze.
Subshrubs or perennial herbs, unarmed, sometimes unbranched. Raphides present. Leaves opposite, often grouped at stem
apices, perhaps without domatia; stipules caducous or persistent, interpetiolar, triangular or generally ligulate to obovate. Inflorescences terminal or pseudo-axillary, cymose, corymbiform, or racemiform with axes scorpioid, spiciform, or bearing small heads, several to many flowered, pedunculate, bracteate or bracts reduced. Flowers pedicellate to sessile, bisexual, distylous. Calyx limb shallowly to deeply 5-lobed, inside with well-developed colleters. Corolla white or yellowish green, tubular or funnelform, inside with
pubescent ring in throat; lobes 5, often cucullate, valvate in bud. Stamens 5, inserted near or above middle of corolla tube, exserted or
included; filaments developed; anthers dorsifixed, sometimes pubescent at one or both ends. Ovary 2-celled, ovules numerous in
each locule on peltate axile placentas; stigma 2-lobed, stout, sometimes scabrous, exserted or included. Fruit baccate, fleshy except
with bony endocarp, subglobose, with calyx limb persistent; seeds numerous, brown, small, angled.
About ten species: SE Asia; two species (one endemic) in China.
This genus was reviewed in detail in Sumatra and Java by Axelius (Blumea 32: 91114. 1987). She noted among other observations that the
plants are quite infrequently encountered in the field. The genus was apparently first reported from China by H. S. Lo (Bull. Bot. Res., Harbin 18:
275283. 1998), who transferred one described species of Ophiorrhiza and Xanthophytum into Lerchea.

1a. Inflorescences with secondary axes mostly scorpioid; leaves with secondary veins 1218 pairs; stipules
caducous, not seen ....................................................................................................................................................... 1. L. micrantha
1b. Inflorescences with secondary axes mostly dichasial; leaves with secondary veins 2025 pairs; stipules
persistent, 2530 mm ......................................................................................................................................................... 2. L. sinica
1. Lerchea micrantha (Drake) H. S. Lo, Bull. Bot. Res., Harbin 18: 275. 1998.
duo lun cao
Ophiorrhiza micrantha Drake, J. Bot. (Morot) 9: 214.
1895; Notodontia micrantha (Drake) Pierre ex Pitard; Spiradiclis micrantha (Drake) H. S. Lo.
Herbs, rather fleshy; stems procumbent and rooting on
nodes with apical parts ascending. Petiole densely pilosuloushirtellous; leaf blade drying membranous, elliptic, ovate-elliptic, or lanceolate-oblong, 518 2.58 cm, base cuneate to
rounded and often decurrent along petiole, apex acuminate or

obtuse; secondary veins 1218 pairs; stipules caducous, not

seen. Inflorescence terminal, ferruginous tomentose, 1018 cm,
many branched; secondary branches arcuate to recurved, mostly
scorpioid, with flowers rather densely grouped; bracts reduced.
Calyx with hypanthium portion weakly 5-ridged, 0.50.7 mm.
Corolla white, shortly tubular; tube swollen in lower part; lobes
obtuse. Berries subglobose, ca. 1.5 mm in diam.
Shady wet streamsides in forests. Yunnan (Hekou) [N Vietnam].
This species was incorrectly called Polyura geminata J. D. Hooker by Wu (Acta Phytotax. Sin. 6: 294. 1957) in its first report from China, but that was a misidentification. Polyura J. D. Hooker is an accepted
genus of Rubiaceae but is not (yet) known from China.

RUBIACEAE

214

2. Lerchea sinica (H. S. Lo) H. S. Lo, Bull. Bot. Res., Harbin

18: 275. 1998.
hua duo lun cao
Xanthophytum sinicum H. S. Lo, Guihaia 11: 97. 1991.
Suffrutescent herbs, rather fleshy, to 1 m tall; stems ascending, angled, densely villous. Petiole 36 cm, villous to hirsute; leaf blade drying papery, oblanceolate to elliptic, 1827
710 cm, glabrous on both surfaces or pilose along midrib
abaxially, base cuneate, apex acute or abruptly acute; secondary

veins 2025 pairs; stipules persistent, obovate, 2530 mm, parallel nerved, obtuse. Inflorescence terminal, paniculiform, lax,
ca. 15 cm, multicellular villous; axes spreading to reflexed,
mostly dichotomous with flowers rather closely grouped; bracts
reduced; pedicels 0.51 mm. Calyx with hypanthium portion
subturbinate to globose, ca. 2 mm, multicellular villous; lobes
narrowly triangular, ca. 3.5 mm, glabrescent. Corolla yellowish
green, tubular, glabrous outside; tube 33.5 mm; lobes triangular-ovate, ca. 1.5 mm. Fl. Aug.
Dense forests. Yunnan (Hekou).

48. LUCULIA Sweet, Brit. Fl. Gard. 2: t. 145. 1826.

dian ding xiang shu
Chen Tao (); Charlotte M. Taylor
Shrubs or trees, unarmed. Raphides absent. Leaves opposite, sometimes with domatia; stipules caducous, interpetiolar, triangular, acute. Inflorescences terminal and sometimes in axils of uppermost leaves giving tripartite appearance, cymose and usually
rounded-corymbiform, pedunculate, bracteate with bracts caducous. Flowers pedicellate, bisexual, distylous, generally showy. Calyx
limb deeply 5-lobed, lobes well developed to subleaflike, usually deciduous shortly after anthesis and often before corolla. Corolla
red to pink or white, salverform; tube prolonged and in long-styled form expanded in throat to enclose anthers, with upper part of
throat spreading with lobes at anthesis, glabrous inside; lobes 5, imbricate (and apparently quincuncial) in bud, rounded, margins
usually crisped and slightly irregular, sometimes with a lamellate basal appendage on each side. Stamens 5, inserted in upper part of
corolla tube, included in long-styled form or partially to completely exserted in short-styled form; filaments short; anthers dorsifixed.
Ovary 2-celled, ovules numerous in each cell on axile placentas; stigmas 2, linear, included in short-styled form or exserted in longstyled form. Fruit capsular, obovoid to ellipsoid, smooth, septicidally dehiscent from apex for half or completely, woody, becoming
deflexed, with calyx limb caducous; seeds numerous, small, fusiform, flattened, prolonged into narrow wings at each end; endosperm
fleshy; embryo minute, slightly clavate; cotyledons obtuse.
About five species: Bhutan, China, India, Myanmar, Nepal, Thailand, Vietnam; three species (one endemic) in China.
The relationships of Luculia have not been clear until recently, when molecular studies (e.g., Bremer et al., Syst. Biol. 48: 413435. 1999) found
it to be basal in the Rubiaceae. Additional references regarding Luculias biology and classification, not listed here, were cited by Rydin et al. (Pl. Syst.
Evol. 278: 101120. 2009).
The treatment here summarizes the existing taxonomy, as a basis for future work; it seems possible that there are only two species in China, but
evaluation of this is beyond the scope of this study and the available specimens. Hutchinson (in Sargent, Pl. Wilson. 3: 408. 1916) gave taxonomic
significance at the species level to the presence vs. absence of striking flap-like processes upon and continuous between the lobes of the corolla for
separating species of Luculia; these structures have not been much studied or detailed by other authors.

1a. Corolla tube 3050 mm; corolla lobes 1215 mm wide, suborbicular to broadly elliptic, without lamellate
appendages at their bases ............................................................................................................................................ 1. L. gratissima
1b. Corolla tube 2532 mm; corolla lobes 915 mm wide, obovate to suborbicular, usually with a lamellate
appendage at each side of base inside.
2a. Inflorescence axes, hypanthium portion of calyx, and fruit glabrous or sparsely hirtellous or pilosulous,
surfaces of structures easily visible between trichomes ......................................................................................... 2. L. pinceana
2b. Inflorescence axes, hypanthium portion of calyx, and fruit densely tomentose, surfaces of structures
completely covered by pubescence ................................................................................................................... 3. L. yunnanensis
1. Luculia gratissima (Wallich) Sweet, Brit. Fl. Gard. 2: t. 145.
1826.
fu yu dian ding xiang
Cinchona gratissima Wallich in Roxburgh, Fl. Ind. 2: 154.
1824; Mussaenda luculia Buchanan-Hamilton ex D. Don, nom.
illeg. superfl.
Shrubs or small trees, to 5 m tall, with thin pale brown
bark; branches somewhat flattened to subterete, pilosulous [to
glabrous], sometimes with sparse elliptic lenticels. Petiole 0.82
cm, pilosulous to glabrous; leaf blade drying papery or thinly

leathery, elliptic, lanceolate-elliptic, lanceolate, or elliptic-oblong, 515 26 cm, adaxially glabrous, abaxially pilosulous at
least along principal veins, base cuneate or acute, apex acute to
acuminate; secondary veins 812 pairs, sometimes with pilosulous domatia; stipules lanceolate to narrowly lanceolate, 68
mm, glabrous to strigillose, acute to acuminate. Inflorescence
corymbiform, many flowered, villosulous to pilosulous; peduncle 23 cm; branched portion 35 68 cm (not including corollas); bracts linear to narrowly ligulate, 47 mm, acute; pedicels 45 mm. Calyx glabrous to densely hirtellous; ovary portion obovoid, 34 mm; lobes lanceolate, oblanceolate, or narrowly ligulate, 1016 mm, acute. Corolla red, outside glabrous;

RUBIACEAE

tube 3050 mm, slenderly cylindrical; lobes suborbicular to

broadly elliptic, 1112 1215 mm, rounded. Capsule 12
11.5 cm, pilosulous to glabrous; seeds 23 mm, spongy-reticulate. Fl. and fr. AprNov.
Forests or thickets on mountains; 8002400 m. Xizang (Mdog),
Yunnan [Bhutan, NE India, Myanmar, Nepal, Thailand, Vietnam].
W. C. Chen (in FRPS 71(1): 239. 1999) distinguished this from
other species of Luculia in part by its villous pubescence on the calyx
and hypanthium, but some individual specimens seen vary from densely
pubescent to glabrous. The pollination biology of this species was
studied by Murray (Ann. Bot. 65: 691698. 1990).

2. Luculia pinceana Hooker in Curtis, Bot. Mag. 71: t. 4132.

1845.
dian ding xiang
Shrubs or trees, 210 m tall; branches flattened to subterete, with usually rather dense elliptic lenticels, hirtellous to
glabrous. Petiole 13.5 cm, glabrous or hirtellous; leaf blade
drying stiffly papery to thinly leathery and paler below, elliptic,
elliptic-oblong, or oblanceolate, 522 28 cm, adaxially glabrous, abaxially glabrous or hirtellous to strigillose on principal
veins to throughout, base cuneate, acuminate, or obtuse, apex
acuminate; secondary veins 914 pairs, often with pilosulous
domatia; stipules triangular to narrowly triangular, 519 mm,
glabrous, long acuminate. Inflorescence corymbiform, many
flowered, glabrous; peduncle 28 cm; branched portion 26
38 cm (not including corollas); bracts stipuliform, linear-lanceolate, 1215 mm, glabrous, acute; pedicels 26 mm. Calyx
glabrous or sparsely hirtellous; ovary portion cylindrical-obovoid, 45 mm; lobes lanceolate to narrowly oblanceolate, 818
26 mm, often ciliate, acute. Corolla red or rarely white, glabrous; tube slenderly cylindrical, 3032 mm; lobes suborbicular, 1522 1115 mm, at base with a lamellate appendage on
each side inside, apex rounded. Capsules obovoid to ellipsoidobovate, 1525 510 mm, weakly longitudinally ridged, glabrous or sparsely hirtellous; seeds ca. 4 mm. Fl. and fr. Mar
Dec.
Forests or thickets on mountain slopes, streamsides in valleys;
6003000 m. Guangxi, Guizhou, Xizang, Yunnan [India, Myanmar, Nepal, Vietnam].
The varieties below were recognized by W. C. Chen in FRPS
(71(1): 239241. 1999) and are included here for reference.

1a. Branchlets and peduncles glabrous ......... 2a. var. pinceana

1b. Branchlets and peduncles pubescent .... 2b. var. pubescens

215

2a. Luculia pinceana var. pinceana

() dian ding xiang (yuan bian zhong)
Luculia intermedia Hutchinson.
Branchlets and peduncles glabrous. Fl. and fr. MarNov.
Forests or thickets on mountain slopes, streamsides in valleys;
6003000 m. Guangxi, Guizhou, Xizang, Yunnan [India, Myanmar, Nepal, Vietnam].

2b. Luculia pinceana var. pubescens (W. C. Chen) W. C.

Chen, Fl. Reipubl. Popularis Sin. 71(1): 241. 1999.
mao dian ding xiang
Luculia intermedia var. pubescens W. C. Chen, Acta Phytotax. Sin. 22: 139. 1984.
Branchlets and peduncles pubescent. Fl. and fr. AprDec.
Forests or thickets on mountain slopes or at streamsides; 600
1800 m. Guangxi (Napo), Xizang (Mdog), Yunnan.

3. Luculia yunnanensis S. Y. Hu, J. Arnold Arbor. 32: 398.

1951.
ji guan dian ding xiang
Shrubs and trees, 3.5(10) m tall; branches weakly flattened to terete, densely tomentose to hirtellous, with dense
elliptic lenticels. Petiole 0.82 cm, sparsely pilosulous to glabrous; leaf blade drying leathery or stiffly papery, oblanceolate,
oblanceolate-oblong, or elliptic-oblong, 5.518 25.5 cm,
adaxially glabrous, abaxially glabrous except strigose to pilose
along principal veins, base cuneate or acute, apex acute to
acuminate; secondary vein 914 pairs, usually with pilosulous
domatia; stipules lanceolate to narrowly triangular, 414 mm,
glabrous, acute to acuminate. Inflorescence corymbiform, many
flowered, densely tomentose; peduncle 1.53.5 cm; branched
portion 510 615 cm (not including corollas); bracts linear
to narrowly spatulate, 515 mm; pedicels 310 mm. Calyx with
ovary portion obconic, 56 mm, densely tomentose; lobes lanceolate to narrowly ligulate or narrowly oblanceolate, 1220
33.5 mm, glabrous except strigose along veins and margins,
apex acute to shortly acuminate. Corolla red, outside glabrous;
tube 2532 mm, cylindrical to slightly funnelform; lobes suborbicular to obovate, 1318 912 mm, with 2 lamellate appendages or crests at each side of base inside, apex rounded.
Capsules obovoid, 1.52.5 0.50.8 mm, smooth to weakly
ridged, densely tomentose; seeds 24 mm. Fl. and fr. MarNov.
Forests or thickets on mountains; 12003200 m. Yunnan.

49. METADINA Bakhuizen f., Taxon 19: 472. 1970.

huang mian mu shu
Chen Tao (); Charlotte M. Taylor
Trees, unarmed; buds pyramidal to conical with stipules obvolute. Raphides absent. Leaves opposite, sometimes with domatia;
stipules caducous, interpetiolar, generally triangular. Inflorescences terminal and in axils of uppermost leaves, capitate with several
globose heads in fascicles or cymes, many flowered, pedunculate, bracteate; peduncles articulate; bracts often subtending heads;
bracteoles filiform to filiform-clavate. Flowers sessile, bisexual, monomorphic. Calyx limb 5-lobed. Corolla color not noted, salverform to narrowly funnelform, inside glabrous; lobes 5, in bud valvate except apices subimbricate. Stamens 5, inserted in upper part of
corolla tube, partially exserted; filaments short; anthers basifixed. Ovary 2-celled, ovules 412 in each cell, pendulous on axile pla-

RUBIACEAE

216

centas attached in upper third of septum; stigma globose to clavate, smooth, exserted. Fruit capsular, obconic, septicidally then sometimes loculicidally dehiscent into 2 or 4 valves from base to apex, woody to stiffly papery, with calyx limb persistent on persistent
septum; seeds several, small, subglobose to trigonous or ellipsoid, unwinged.
One species: Cambodia, China, India, Indonesia, Laos, Malaysia, Myanmar, Philippines, Thailand, Vietnam.
The taxonomy of this genus here follows Ridsdale (Blumea 24: 350351. 1979), with a broader morphological variation as found in more
recently collected specimens, in particular in inflorescence morphology. This taxonomy is complicated by varied application of the name Adina
polycephala: Ridsdale treated this as a synonym of Metadina trichotoma, but older Chinese authors (in herb.) have long applied it to the plants treated
here as A. pubicostata.

1. Metadina trichotoma (Zollinger & Moritzi) Bakhuizen f.,

Taxon 19: 472. 1970.
huang mian mu
Nauclea trichotoma Zollinger & Moritzi, Syst. Verz. 61.
1846; Adina polycephala Bentham; A. trichotoma (Zollinger &
Moritzi) Bentham & J. D. Hooker ex B. D. Jackson; A. zschokkei Elmer; N. polycephala Wallich ex G. Don (1834), not A.
Richard ex Candolle (1830).
Trees, apparently evergreen, flowering at 510 m tall;
branches compressed becoming terete, brown to gray, puberulent to glabrous, usually densely lenticellate with elliptic white
raised lenticels. Petiole 310 mm, glabrous; leaf blade drying
stiffly papery to subleathery, lanceolate, elliptic-lanceolate, or
ovate-oblong, 620 27 cm, adaxially glabrous and rather
shiny, abaxially glabrous to puberulent or tomentulose at least
along veins, base acute to obtuse, apex acute to caudate-acumi-

nate; secondary veins 812 pairs, sometimes with small pilosulous domatia in axils; stipules deltoid to narrowly triangular,
58 mm, acute to perhaps obtuse. Inflorescence glabrous to
densely puberulent or tomentulose; peduncles 1.53 cm, usually articulate and with 4 caducous bracts 13 mm near middle;
flowering heads 67 mm in diam. across calyces, ca. 12 mm in
diam. across corollas; bracteoles ca. 2 mm. Calyx with ovary
portion obconic, 0.51 mm, pilosulous to glabrescent, surrounded at base by a ring of pilose trichomes ca. 0.5 mm; limb
lobed essentially to base; lobes 12 mm, narrowly elliptic-oblong, obtuse. Corolla outside glabrous; tube 33.5 mm; lobes
triangular-spatulate, ca. 1 mm. Stigmas obconic, ca. 1 mm, exserted for ca. 5 mm. Fruiting head 810 mm in diam. Capsules
obovoid to obconic, ca. 1.5 mm, pilosulous at least on apical
portion. Fl. and fr. AprDec.
Forests at streamsides in valleys; 3001400 m. Guangdong,
Guangxi, Hunan, Yunnan [Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Philippines, Thailand, Vietnam].

50. MICROPHYSA Schrenk, Bull. Cl. Phys.-Math. Acad. Imp. Sci.

Saint-Ptersbourg 2: 115. 1844.
pao guo qian cao shu
Chen Tao (); Friedrich Ehrendorfer
Herbs, perennial, rhizomatous. Raphides present. Stems rough. Leaves and leaflike stipules of middle stem region in whorls of
4, without domatia, sessile or subsessile. Inflorescences terminal, thyrsoid, corymbose, cymes pedunculate, several to many flowered, bracteate but bracteoles strongly reduced. Flowers sessile to pedicellate, remarkably small, bisexual, monomorphic. Calyx limb
obsolete. Corolla white, funnel-shaped, glabrous; lobes 4, valvate in bud. Stamens 4, inserted on upper part of corolla tube, exserted;
filaments short; anthers dorsifixed. Ovary (hypanthium) with 2 cells, each with 1 erect ovule inserted at base of placenta; stigma 2lobed, exserted. Fruit indehiscent or tardily schizocarpous, dry, with pericarp leathery and inflated (i.e., bladderlike) and usually
including both seeds at dispersal; seeds small, ellipsoid-oblong or plano-convex, grooved on ventral (i.e., adaxial) side; testa membranous; endosperm corneous; embryo curved; cotyledons leaflike; radicle terete, hypogynous.
One species: NW China, Kazakhstan, Uzbekistan.
Microphysa was originally described as Asperula elongata. Its subsequent generic separation by Schrenk (Bull. Cl. Phys.-Math. Acad. Imp. Sci.
Saint-Ptersbourg 2: 116. 1844) was justified by its peculiar fruit structure. Only much later, Ehrendorfer et al. (Fl. Iranica 176: 161258. 2005)
postulated that Microphysa (as well as A. platygalium and A. maximoviczii: see under Galium), in spite of their funnel-shaped corollas, phylogenetically belong to G. sect. Platygalium (usually with rotate corollas), a section to which they correspond in all remaining essential characters. In the
aberrant fruit of Microphysa the two parts of the inferior ovary do not develop into separate mericarps, each with its own pericarp, but remain united
within a inflated leathery and common pericarp. Nevertheless, this peculiar development is not limited to Microphysa but, according to personal
observations (F. Ehrendorfer), also occurs sporadically within populations of the G. boreale-G. rubioides group (corresponding to the series G. ser.
Rubioidea Pobedimova and G. ser. Borealia Pobedimova within G. sect. Platygalium; cf. Pobedimova et al., Fl. URSS 23: 345354. 1958). Thus, on
the basis of available evidence, the single species of Microphysa should be placed into G. sect. Platygalium.
Considering that Microphysa was separated as a monotypic genus by Pobedimova (loc. cit.) and by H. S. Lo (in FRPS 71(2): 318. 1999), and
that DNA-analytical proof for its transfer to Galium sect. Platygalium is still lacking, the genus is maintained here provisionally. Furthermore, its
possible future transfer will make a new species name necessary, because elongata has been used already for another taxon within Galium.

1. Microphysa elongata (Schrenk) Pobedimova in Schischkin,

Fl. URSS 23: 286. 1958.
pao guo qian cao

Asperula elongata Schrenk in Fischer & C. A. Meyer,

Enum. Pl. Nov. 1: 58. 1841; Microphysa galioides Schrenk.
Herbs, perennial, erect, rhizomatous. Stems 3050(70)

RUBIACEAE

cm tall, with 4 scabrous angles. Leaf blade drying subleathery,

linear-lanceolate, lanceolate, or narrowly oblong, 3060 3
5(12) mm, glabrous except antrorsely aculeolate along margins and veins abaxially, base acute, margins thinly revolute,
apex acute or obtuse; vein 1, distinct, secondary veins not
visible. Inflorescence corymbose, with scabrous axes, inconspicuously bracteate. Ovary ellipsoid, 1.52 mm, glabrous. Co-

217

rolla funnel-shaped, 2.53 3.54 mm, lobed to ca. 1/2 length

or slightly more; tube 1.21.5 mm; lobes elliptic-oblong,
shortly acuminate. Fruit 34 mm in diam., vesicular, smooth to
granulate; seeds ca. 1 1 mm. Fl. and fr. MayJul.
Meadows, banks of rivers and lakes, foothills and lower mountains. Xinjiang (Chabuchaer) [Kazakhstan, Uzbekistan].

51. MITCHELLA Linnaeus, Sp. Pl. 1: 111. 1753.

man hu ci shu
Chen Tao (); Charlotte M. Taylor
Chamaedaphne Mitchell; Perdicesca Provancher.
Herbs, perennial, unarmed, creeping, rooting at nodes. Raphides present. Leaves opposite, without domatia; stipules generally
persistent, interpetiolar, triangular, entire to deeply 3(5)-lobed, often glandular at apex. Inflorescences terminal or pseudoaxillary
near stem apices, 2-flowered, pedunculate, ebracteate. Flowers sessile, bisexual, distylous, fused in pairs by their ovaries. Calyx with
ovary portion of individual flowers subglobose, with structure formed by fused ovaries oblate to dicoccous, with limb of individual
flowers (3 or)4-lobed. Corolla white, funnelform, pilose in throat and onto lobes; lobes (3 or)4, valvate in bud. Stamens (3 or)4,
inserted in corolla throat, exserted in short-styled flowers, included in long-styled flowers; filaments stout; anthers basifixed. Ovary
of individual flowers 4-celled, ovules 1 in each cell, axile; stigmas 4, linear, exserted in long-styled flowers, included in short-styled
flowers. Fruit multiple, orange to red, drupaceous, subglobose to oblate, fleshy, with calyx limbs 2, persistent; pyrenes 8, 1-celled,
each with 1 seed, angled, 3-ridged; seeds medium-sized, ellipsoid; endosperm corneous; embryo small; radicle hypogynous.
Two species: one in E Asia (China, Japan, Korea), the other in Central America (Guatemala) and E North America (Canada, Mexico, United
States); one species in China.
Y. Z. Ruan (in FRPS 71(2): 159. 1999) gave the number of calyx lobes, corolla lobes, and stamens as 3 or 4; in general, the flowers of Mitchella
are considered 4-merous, although throughout Rubiaceae occasional individual flowers vary from the characteristic in having fewer or more calyx
lobes, corolla lobes, and infrequently also stamens.

1. Mitchella undulata Siebold & Zuccarini, Abh. Math.-Phys.

Cl. Knigl. Bayer. Akad. Wiss. 4(3): 175. 1846.
man hu ci
Mitchella repens Linnaeus var. undulata (Siebold & Zuccarini) Makino.
Plants with stems quadrate, to 30 cm, glabrous or subglabrous. Petiole 0.31.1 cm, glabrous or subglabrous; leaf
blade drying papery, triangular-ovate or ovate, 0.22.1 0.2
1.5 cm, both surfaces glabrous, base truncate or cordulate to
rounded, margins sometimes undulate, apex acute to rounded;
secondary veins 2 or 3 pairs; stipules 11.5 mm. Flowers with
peduncles 112 mm, glabrous. Calyx glabrous; ovary portion

subglobose to turbinate, ca. 2 mm; limb deeply lobed; lobes

narrowly to broadly triangular, 0.51 mm. Corolla glabrous
outside; tube 910 mm; lobes lanceolate to triangular, 45 mm,
acute. Multiple fruit subglobose, 68 mm in diam., glabrous;
pyrenes ca. 2.5 mm. Fl. autumn, fr. winter.
Understories of wet forests. Taiwan, Zhejiang [Japan, Korea].
In Fl. Taiwan (ed. 2, 4: 297. 1998) Liu and Yang reported the
plants there are evergreen. Y. Z. Ruan (in FRPS 71(2): 159. 1999)
described the leaf blades as unequal and gave measurements for large
ones and small ones without further explanation. All other authors
reported this species to be generally isophyllous, which corresponds to
specimens seen. The leaf description by Ruan may apply to the variable
leaf sizes along the stems of some plants, rather than consistent size
differences between the two leaves at a single stem node.

52. MITRACARPUS Zuccarini in Schultes & J. H. Schultes, Mant. 3: 210

[Mitracarpum], 399. 1827.
gai lie guo shu
Chen Tao (); Charlotte M. Taylor
Herbs [to small shrubs], annual or perennial, unarmed. Raphides present. Leaves opposite, subsessile or sessile, without domatia; stipules persistent, interpetiolar and fused to leaf bases or petioles, truncate to rounded, setose. Inflorescences terminal and/or
axillary, glomerulate or capitate, several to many flowered, sessile, sometimes immediately subtended by 1 or 2 pairs of leaves,
bracteate. Flowers sessile, bisexual, monomorphic. Calyx limb deeply 4(or 5)-lobed; lobes usually unequal in pairs, usually with
hyaline margins. Corolla white, salverform or funnelform, inside glabrous or pubescent in throat; lobes 4, valvate in bud. Stamens 4,
inserted in corolla throat, included or exserted; filaments developed; anthers dorsifixed. Ovary 2-celled, ovules 1 in each cell on
peltate axile placentas attached at middle of septum; stigmas 2, linear, usually exserted. Fruit capsular, subglobose to somewhat
dicoccous, with dehiscence circumscissile around equator, with apical valve or lid deciduous and basal portion persistent, papery to

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cartilaginous, with calyx limb persistent or deciduous on lid; seeds 2, medium-sized, oblate to rounded, on ventral (i.e., adaxial) face
with cruciform (i.e., X-shaped) scar; endosperm fleshy; cotyledon leaflike; radicle hypogeous.
About 30 species: widespread in tropical and subtropical Central, North, and South America and the Antilles, with one species widely naturalized in tropical Africa, Asia, Australia, and Pacific islands; one species (introduced) in China.
This genus is frequently overlooked even in its native range, but the circumscissile capsules, seeds with a distinctive cruciform scar, leaves
scabrous to the touch on the upper surface, and well-developed calyx lobes with hyaline erose margins are distinctive. W. C. Ko (in FRPS 71(2): 210.
1999) described the ovary as sometimes 3-celled; this condition is not otherwise known in Mitracarpus and has not been reconfirmed, although it does
characterize the very similar, likewise adventive genus Richardia.

1. Mitracarpus hirtus (Linnaeus) Candolle, Prodr. 4: 572.

1830.
gai lie guo
Spermacoce hirta Linnaeus, Sp. Pl., ed. 2, 1: 148. 1762;
Mitracarpus scaber Zuccarini; M. senegalensis Candolle; M.
verticillatus (Schumacher & Thonning) Vatke; M. villosus
(Swartz) Candolle; S. villosa Swartz; Staurospermum verticillatum Schumacher & Thonning.
Herbs, annual, branched, 4080 cm tall; branches flattened
to subterete or 4-angled, sometimes becoming woody in lower
part, sparsely hirsute to villous. Leaves sessile; blade drying
thinly papery, elliptic-oblong or lanceolate, 34.5 0.71.5 cm,
adaxially scaberulous and usually also sparsely villosulous or
pilosulous, abaxially moderately to densely villous or pilose,
base acute to obtuse or rounded, apex acute; secondary veins 3
6 pairs; stipule sheaths 14 mm, villosulous or pilosulous to
glabrescent, with 19 setae 15 mm. Inflorescences 520 mm
in diam. (not including subtending leaves), villosulous or pilosulous; bracts linear, 12 mm. Calyx sparsely to densely puberulent or strigillose; ovary portion subglobose to ellipsoid, ca.

0.5 mm; limb deeply lobed; lobes unequal, triangular to lanceolate, 2 larger 1.82 mm, 2 smaller 0.81.2 mm, ciliate. Corolla
funnelform, outside puberulent to glabrous; tube 11.5 mm,
glabrous inside; lobes triangular to ovate, 0.51 mm, obtuse to
acute. Capsules subglobose, ca. 1 mm in diam., scaberulous or
sparsely puberulent; seeds dark brown, oblate-suboblong, ca.
0.8 mm. Fl. and fr. AprNov.
Wastelands at highway sides; near sea level to 800 m. Hainan
(Wanning), Hong Kong, Yunnan [native to the Antilles and Central,
North, and South America; naturalized in tropical Africa, Asia, Australia, and Pacific islands].
The correct name for this species, whether Mitracarpus hirtus or
M. villosus, has been controversial (for additional comments, see Taylor
et al., Fl. Venez. Guayana 8: 497847. 2004); most authors working
with neotropical Rubiaceae today use the name M. hirtus. The capsules
of all the Chinese specimens studied are smaller than those of neotropical plants. Fruit of similar size are found in plants of India (Sebastine & Ramamurthy, Bull. Bot. Surv. India 9: 291292. 1967, see in particular f. 17) and may characterize several adventive populations. W.
C. Ko (in FRPS 71(2): 212. 1999, as M. villosus) suggested that this
species might be distylous, but Mitracarpus is monomorphic so far as
known.

53. MITRAGYNA Korthals, Observ. Naucl. Indic. 19. 1839, nom. cons.,
not Mitragyne R. Brown (1810).
mao rui mu shu
Chen Tao (); Charlotte M. Taylor
Paradina Pierre ex Pitard; Stephegyne Korthals.
Trees, unarmed; buds flattened, with stipules erect and pressed together. Raphides absent. Leaves opposite, sometimes with
domatia; stipules caducous, interpetiolar, generally ovate to obovate, sometimes keeled, entire, often well developed. Inflorescences
terminal on main stems and axillary branches and often accompanied by reduced, petaloid, and/or bracteate leaves, capitate with
globose heads in fascicles, cymes, umbels, or thyrses, sessile to shortly pedunculate, bracteate; bracteoles spatulate to obpyramidal.
Flowers sessile, bisexual, monomorphic. Calyx limb truncate to 5-lobed. Corolla cream to yellow-green, funnelform or narrowly
salverform, inside glabrous to variously pubescent; lobes 5, valvate in bud. Stamens 5, inserted near corolla throat, exserted or
included; filaments short; anthers basifixed, partially to fully exserted. Ovary 2-celled, ovules numerous in each cell on fleshy,
pendulous, axile placentas attached in upper third of septum; stigma clavate to mitriform (i.e., upside-down cupular), exserted. Fruit
capsular, obovoid to ellipsoid, septicidally then loculicidally dehiscent, cartilaginous to woody, with calyx limb persistent or deciduous; seeds numerous, small, somewhat flattened, fusiform to lanceolate, shortly winged at both ends with basal wing sometimes bifid
or notched.
About seven species: one species in Africa, six species in Asia and Malesia; three species in China.
Ridsdale reviewed this genus in detail (Blumea 24: 4668. 1978) and excluded the African species. H. H. Hsue and H. Wu (in FRPS 71(1): 245.
1999) reported only Mitragyna rotundifolia from China; Ridsdale (loc. cit.: 65) reported only M. diversifolia from China; and Wu (Acta Phytotax. Sin.
6: 293. 1957) reported a third species, M. hirsuta, in a report that has been overlooked. Several other species of Mitragyna are found widely in
Thailand and Myanmar, as well as cultivated for lumber, and should be expected in China (in particular, see comments under M. diversifolia). The
leaves of M. speciosa (Korthals) Haviland are the source of kratom and used for tea, chewing, smoking, and as medicine in Thailand and Malaysia;
the main active ingredient here is the alkaloid mitragynine, known only from this species and said to be stimulating at low doses but narcotic at high
doses.

RUBIACEAE

219

1a. Calyx limb deeply lobed, with lobes 1.52.5 mm, spatulate to oblanceolate, usually quite evident on fruit ............... 2. M. hirsuta
1b. Calyx limb subtruncate or lobed for up to 1/2, with lobes up to 1 mm, triangular, deciduous or at least
hardly evident on fruit.
2a. Branch leaves 614 39 cm, with secondary veins strongly ascending (i.e., departing midrib at 5575);
corolla tube ca. 3 mm, lobes ca. 2.5 mm ............................................................................................................ 1. M. diversifolia
2b. Branch leaves 925 620 cm, with secondary veins spreading (i.e., departing midrib at 3560);
corolla tube 23 mm, lobes 45 mm ................................................................................................................. 3. M. rotundifolia
1. Mitragyna diversifolia (Wallich ex G. Don) Haviland, J.
Linn. Soc., Bot. 33: 71. 1897.
yi ye mao rui mu
Nauclea diversifolia Wallich ex G. Don, Gen. Hist. 3: 467.
1834; Stephegyne diversifolia (Wallich ex G. Don) J. D. Hooker.
Trees, perhaps deciduous, to 15 m tall; branches angled
becoming terete, pilosulous to glabrescent. Petiole 515 mm,
glabrous, puberulent, or pilosulous; leaf blade drying papery,
ovate-oblong to elliptic-ovate, 614 39 cm, adaxially glabrous, abaxially sparsely to densely pilosulous or tomentulose,
base rounded to cordulate, apex obtuse to shortly acuminate;
secondary veins 8 or 9 pairs, strongly ascending, sometimes
with pilosulous domatia in abaxial axils; stipules elliptic-oblong
to ovate, ca. 2.5 cm, strigillose to glabrous, abaxially weakly to
strongly keeled and pilosulous, apex obtuse to rounded. Inflorescence densely pilosulous or strigillose to glabrescent; peduncles 13 mm (i.e., portion above articulation of subtending
leaves but not including entire growth of branch); flowering
heads 3 to numerous, 810 mm in diam. across calyces, 1320
mm in diam. across corollas; bracteoles linear-spatulate, ca. 3
mm, glabrous to sparsely ciliolate. Calyx glabrous; ovary portion obconic, ca. 1.5 mm; limb ca. 1.5 mm, subtruncate to lobed
for up to ca. 1/2; lobes triangular, obtuse. Corolla yellowish
white, outside glabrous, inside densely pilosulous in throat and
on lobes; tube ca. 3 mm; lobes triangular, ca. 2.5 mm, acute.
Fruiting heads 810 mm in diam. Capsules 34 mm, with persistent calyx limb markedly thickened; seeds 12 mm. Fl. [Feb,
Jul, Aug], fr. [Jan, Mar, Dec].
Forests; [300400 m]. Yunnan [Cambodia, Indonesia, Laos, Malaysia, Myanmar, Philippines, Thailand, Vietnam].
This species seems to be similar to and at least sometimes confused with the commonly collected Mitragyna parvifolia (Roxburgh)
Korthals; these species differ at least in their corollas, with the tube 56
mm and substantially longer than the lobes ca. 2 mm in M. parvifolia,
vs. the tube ca. 3 mm and less than twice as long as the lobes ca. 2.5
mm in M. diversifolia. Ridsdale (Blumea 24: 6365. 1978) reported a
difference in the distribution of these, with M. parvifolia primarily
found in India and Sri Lanka and M. diversifolia to the north and east of
this, including China. Puff et al. (Rubiaceae of Thailand, 46. 2005)
noted that in Thailand M. diversifolia is most often found in deciduous
vegetation and is common in secondary vegetation, in particular in
edges along fields.

2. Mitragyna hirsuta Haviland, J. Linn. Soc., Bot. 33: 72.

1897.
mao mao rui mu
Paradina hirsuta (Haviland) Pitard.
Trees, deciduous, to 20 m tall; branches angled to terete,

densely pilosulous to glabrescent. Petiole 530 mm, glabrous to

densely pilosulous; leaf blade drying stiffly papery, suborbicular to broadly elliptic or ovate, 818(30) 212(20) cm,
adaxially glabrous, abaxially sparsely to densely pilosulous or
rarely glabrescent, base broadly obtuse to cordulate, apex
rounded to acute; secondary veins 612 pairs, spreading, sometimes with pilosulous domatia in abaxial axils; stipules ellipticoblong to ovate, 1020 815 mm, pilosulous and weakly
keeled, apex obtuse to rounded. Inflorescences densely puberulent to pilosulous; flowering heads sessile, 7 to numerous, 10
12 mm in diam. across calyces, 2025 mm in diam. across
corollas; bracteoles linear-spatulate, 2.53.5 mm, glabrous to
sparsely pubescent and/or ciliolate. Calyx glabrous; ovary portion obconic, 1.22 mm; limb deeply lobed; lobes oblanceolate
to spatulate, 1.52.5 mm, entire to ciliolate. Corolla yellow, outside glabrous, inside densely hairy; tube 56 mm; lobes narrowly elliptic, 22.5 mm, acute. Fruiting heads 1520 mm in
diam. Capsules 58 mm, weakly ridged; seeds ca. 1 mm. Fl.
[JunJul, Dec], fr. [Apr, Dec].
Forests; [1001500 m]. Yunnan [Cambodia, Laos, Myanmar,
Thailand, Vietnam].
This species was illustrated by Ridsdale (Blumea 24: 60, f. 6.
1978).

3. Mitragyna rotundifolia (Roxburgh) Kuntze, Revis. Gen. Pl.

1: 289. 1891.
mao rui mu
Nauclea rotundifolia Roxburgh, Fl. Ind. 2: 124. 1824;
Mitragyna brunonis (Wallich ex G. Don) Craib; N. brunonis
Wallich ex G. Don.
Trees, perhaps deciduous, to 30 m tall; branches angled to
subterete, glabrous to glabrescent. Petiole 1560 mm, glabrous
to densely pilosulous; leaf blade drying papery, suborbicular to
broadly elliptic or ovate, 925 620 cm (to 75 cm on seedlings and sprouts), adaxially glabrous to puberulent, abaxially
sparsely to densely pilosulous or tomentulose, base rounded to
cordate, apex rounded to obtuse; secondary veins 57 pairs,
spreading, sometimes with pilosulous domatia in abaxial axils;
stipules elliptic-oblong to ovate, 1350 530 mm, pilosulous,
keeled, apex obtuse to rounded. Inflorescences densely puberulent to pilosulous; peduncles 13 mm (i.e., portion above articulation of subtending leaves but not including internode below
node bearing inflorescence); flowering heads 15, 710 mm in
diam. across calyces, 1520 mm in diam. across corollas; bracteoles linear-spatulate, 11.5 mm, glabrous or sparsely pubescent. Calyx glabrous; ovary portion 1.53 mm; limb ca. 0.5
mm, subtruncate to lobed for ca. 1/2; lobes triangular, obtuse.
Corolla yellowish white, outside glabrous, densely hairy inside;
tube 23 mm; lobes narrowly oblanceolate, 45 mm, acute.
Fruiting heads 1016 mm in diam. Capsules 35 mm, weakly

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220

ridged, with persistent calyx thickened; seeds ca. 1 mm. Fl.

[AugNov], fr. Sep, Dec.
Dense forests; ca. 1000 m. S Yunnan [Bangladesh, India, Laos,
Myanmar, Thailand].

This species was illustrated by Ridsdale (Blumea 24: 66, f. 8.

1978). Puff et al. (Rubiaceae of Thailand, 46. 2005) noted that in Thailand this species is most often found in deciduous vegetation and is
common in secondary vegetation.

54. MORINDA Linnaeus, Sp. Pl. 1: 176. 1753.

ba ji tian shu
Chen Tao (); Charlotte M. Taylor
Rojoc Adanson.
Lianas, climbing shrubs, erect shrubs, or small trees, rarely dioecious, sometimes with spines; branching sometimes sylleptic
with growth continued from an axillary or subapical node, with lateral branches sometimes surrounded at base by persistent leafless
stipules. Raphides present. Leaves opposite, rarely ternate, or sometimes anisophyllous and apparently 1 at flowering nodes, sometimes with domatia in axils of secondary and occasionally tertiary veins, margins rarely sinuate-undulate or lyrate; stipules persistent
or infrequently caducous, interpetiolar, united around stem, or fused to petioles, triangular, entire. Inflorescences terminal, axillary, or
leaf-opposed, capitate with 1 to several hemispherical to subglobose heads, these fasciculate or cymose, few to many flowered,
pedunculate or sessile, bracteate or bracts reduced. Flowers sessile, shortly to fully fused by their ovaries [to free or nearly so], bisexual and distylous, rarely bisexual and monomorphic, or rarely dioecious. Calyx limb truncate to sinuate or rarely in 13 flowers of an
inflorescence with 1 petaloid calycophyll (Morinda citrifolia). Corolla white or pink, funnelform, salverform, or campanulate, inside
glabrous or pubescent in throat, [tube sometimes fenestrate]; lobes 37, valvate in bud. Stamens 37, inserted in corolla throat or
tube, exserted or included; filaments short; anthers dorsifixed, sometimes with connective prolonged into an apical appendage. Ovary
2-celled with ovules 2 in each cell, or incompletely to completely 4-celled due to secondarily formed false septa with ovules 1 in
each cell, ovules attached to septum near base; stigmas 2, linear, exserted or included. Fruit multiple with entire fruiting heads comprising one fruit (i.e., drupecetum) [sometimes fruit simple]; individual fruit drupaceous, fleshy, generally obovoid, blue to black,
with calyx limb persistent; pyrenes 24, 1-locular, with 1 seed, cartilaginous or bony, subtrigonous to plano-convex, adaxially (i.e.,
ventrally) flat or sulcate; seeds medium-sized, subtrigonous or ellipsoid; endosperm abundant, corneous; embryo small; cotyledons
oblong; radicle inferior.
About 80100 species: widespread in tropical and subtropical regions worldwide; 27 species (18 endemic) in China.
Morinda includes a notable range of breeding systems (Johansson, Opera Bot. 122: 1167. 1994), but most of the species are apparently distylous, with the anthers and stigmas separated and their positions reciprocal between the short-styled and long-styled form of the same species; however,
this biology has been sometimes overlooked. Also, as noted by Johansson (loc. cit.), the position of the inflorescences, in particular terminal vs. leafopposed, deserves careful observation and aids identification of species. In particular, the leaf-opposed inflorescences often are produced on the
terminal node, then later displaced by subsequent growth from the axil of that leaf, and can be confused with true terminal inflorescences that have
two subtending leaves. The inflorescences with fasciculate to umbellate peduncles actually appear to be condensed cymes or racemes, with the
peduncles arising from a very shortly prolonged structure at the stem apex that also bears several stipuliform bracts, usually one above the other. The
twining Asian species apparently share having their lateral branches surrounded at the base (i.e., at the divergence from main stem) by persistent
leafless stipules. Some species of Morinda have petaloid bracts or possibly calyx lobes; this character appears to vary within some individual species.
Y. Z. Ruans (in FRPS 71(2): 179202. 1999) taxonomy of Morinda distinguished species based on different characters, in particular pubescence, leaf
shape, peduncle length compared across developmental stages, drying color and texture, pattern of tertiary leaf veins on dried specimens, and degree
of fusion of flowers, than used by many other authors (e.g., Johansson, loc. cit.; Springate et al., Fl. Bhutan 2(2): 804. 1999).

1a. Erect trees or shrubs; inflorescences terminal, axillary, or leaf-opposed, 1 or 2 per node.
2a. Plants of seashores and low elevations, 050 m; leaves with 57 pairs of secondary veins; fruit 2.55 cm
in diam. .................................................................................................................................................................... 6. M. citrifolia
2b. Plants of terrestrial inland habitats, 5001400 m (exact elevation unknown in M. leiantha).
3a. Inflorescences terminal and/or axillary.
4a. Stems hirtellous, hispidulous, or subglabrous; leaves with secondary veins 58 pairs; peduncles
0.20.5 cm; corollas with tubes ca. 20 mm and lobes ca. 4 mm ............................................................. 13. M. leiantha
4b. Stems glabrous or subglabrous; leaves with secondary veins 810 pairs; peduncles ca. 2 cm;
corollas with tubes ca. 32 mm and lobes ca. 13 mm .......................................................................... 15. M. longissima
3b. Inflorescences leaf-opposed.
5a. Stems glabrous to pubescent; leaves pubescent abaxially, with secondary veins 710 pairs;
peduncles 0.10.3 cm .......................................................................................................................... 19. M. persicifolia
5b. Stems glabrous; leaves glabrous abaxially, with secondary veins 924 pairs; peduncles 16 cm.
6a. Leaves 45.5 cm wide, with secondary veins 1224 pairs; corollas with tubes 1518 mm,
lobes 34 mm ................................................................................................................................... 26. M. undulata
6b. Leaves 511 cm wide, with secondary veins 914 pairs; corollas with tubes 1633 mm,
lobes 415 mm.

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221

7a. Petioles 0.51 cm; corollas with tubes 1633 mm, lobes 415 mm; fruit ca. 2.5 cm
in diam. .................................................................................................................................... 1. M. angustifolia
7b. Petioles 24 cm; corollas with tubes ca. 22 mm, lobes ca. 4 mm; fruit ca. 1 cm in diam. ....... 21. M. rosiflora
1b. Lianas, twiners, or subshrubs; inflorescences terminal, with heads solitary or in groups of 211.
8a. Calyx limb developed, 12.5 mm; leaves acute to obtuse, rounded, truncate, or cordulate at base.
9a. Stems and leaves glabrous ............................................................................................................................. 10. M. howiana
9b. Stems and leaves glabrous to densely hirtellous or pilosulous, with pubescence present at least on veins
abaxially on young leaves.
10a. Stems and leaves sparsely to moderately puberulent to glabrescent; corollas with tubes and
lobes equal, tube 34 mm, lobes 34 mm; fruit 0.51.1 cm in diam. ............................................ 17. M. officinalis
10b. Stems and leaves abaxially densely hirtellous or pilosulous; corollas with lobes longer than tube,
tube 12 mm, lobes 34.5 mm; fruit 12 cm in diam.
11a. Leaves with 710 pairs of secondary veins; corollas with tubes 1.52 mm and lobes
44.5 mm; calyx lobes narrowly triangular, acute ............................................................ 8. M. cochinchinensis
11b. Leaves with 1013 pairs of secondary veins; corollas with tubes 11.2 mm and lobes
34 mm; calyx lobes rounded to obtuse ........................................................................................ 27. M. villosa
8b. Calyx limb reduced to shortly developed, less than 1 mm; leaves acute to cuneate, obtuse, or rounded
at base.
12a. Leaves rugulose; corolla tube ca. 10 mm ................................................................................................... 22. M. rugulosa
12b. Leaves flat, smooth; corolla tube 13 mm or 1225 mm.
13a. Corolla tube 1225 mm.
14a. Corolla tube 1220 mm.
15a. Young stems densely ferruginous hirtellous when dry; leaves drying leathery
and abaxially whitened ....................................................................................... 5. M. cinnamomifoliata
15b. Young stems densely puberulent, strigillose, or hirtellous with pubescence
drying colorless; leaves drying papery to membranous and abaxially golden
yellow or brownish yellow ................................................................................................... 7. M. citrina
14b. Corolla tube ca. 25 mm.
16a. Leaves glabrous throughout .............................................................................................. 3. M. brevipes
16b. Leaves sparsely to densely pubescent at least abaxially on veins.
17a. Stems puberulent with trichomes drying colorless, to glabrescent .............................. 2. M. badia
17b. Stems densely hirtellous with trichomes drying golden brown ..................... 4. M. callicarpifolia
13b. Corolla tube 13 mm.
18a. Corollas with tubes and lobes equal in length, tube 22.5(3) mm, lobes 22.5(3) mm.
19a. Stems and leaves glabrous .............................................................................................. 12. M. lacunosa
19b. Stems and leaves puberulent to hirtellous at least when young.
20a. Leaves drying with gray to black cast .................................................................. 14. M. litseifolia
20b. Leaves drying with brown or yellow cast.
21a. Leaves drying with brown cast; corollas with tubes ca. 3 mm,
lobes ca. 3 mm .................................................................................. 24. M. shuanghuaensis
21b. Leaves drying with brown, grayish brown, or yellowish brown
cast; corollas with tubes 22.5 mm, lobes 22.5 mm.
22a. Petioles 0.40.8 cm; leaves adaxially sparsely hispidulous to
glabrescent ....................................................................................... 11. M. hupehensis
22b. Petioles 0.51.5 cm; leaves adaxially sparsely hirtellous,
subglabrous, or glabrous .................................................................. 23. M. scabrifolia
18b. Corollas with lobes longer than tubes by 50% or more, tube 12 mm, lobes 24 mm.
23a. Leaves drying with gray to black cast; stems densely yellowish, clear, or
ferruginous strigillose or hirtellous .............................................................................. 9. M. hainanensis
23b. Leaves drying with green, brown, yellow, or reddish cast; stems moderately to
sparsely hirtellous, hispidulous, or puberulent with colorless trichomes, to
subglabrous.
24a. Leaves drying with yellow or reddish cast.
25a. Leaves 27 0.33 cm, cuneate to acute at base; corolla lobes
34 mm ....................................................................................................... 18. M. parvifolia
25b. Leaves 713 24 cm, cuneate to obtuse or rounded at base;
corolla lobes ca. 2 mm ......................................................................... 20. M. pubiofficinalis
24b. Leaves drying with brown, grayish brown, or greenish cast.
26a. Leaves with secondary veins 4 or 5 pairs; peduncles 0.41.1 cm;
fruit 0.40.8 cm in diam. ....................................................................... 16. M. nanlingensis

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RUBIACEAE

26b. Leaves with secondary veins 57 pairs; peduncles 11.5 cm; fruit
0.71.2 cm in diam. .................................................................................... 25. M. umbellata
1. Morinda angustifolia Roxburgh, Pl. Coromandel 3: 32.
1815.

apically thickened and rostrate. Drupecetum subglobose, 0.5

0.8 cm in diam. Drupes fused, orange. Fl. Jun, fr. Oct.

huang mu ba ji

Forests on mountains, thickets at watersides. Guangdong

(Guangzhou), Guangxi, Hainan, Hunan.

Erect shrubs or small trees, ca. 6 m tall; branches quadrangular, glabrous. Leaves opposite, or solitary opposite an inflorescence; petiole 0.51 cm, glabrous; blade drying papery,
matte on both surfaces, brownish green, elliptic-oblong, elliptic,
oblong-lanceolate, or oblanceolate, 1530 610 cm, adaxially
glabrous, abaxially scabrous to glabrous or sometimes sparsely
hirtellous on veins, base acute to attenuate, apex acute to acuminate; secondary veins 913 pairs; stipules interpetiolar, free
or shortly united to petioles, triangular, 4.55 mm, acuminate or
acute. Inflorescence solitary and leaf-opposed; peduncle 1.54
cm; flowering head 1, subglobose to cylindrical, many flowered, 0.82.5 cm (not including corollas); bracteoles subulate.
Flowers fused only shortly at base, distylous. Calyx glabrous;
limb truncate to denticulate, ca. 1 mm. Corolla white, salverform, outside glabrous; tube cylindrical to slenderly funnelform, 1633 mm; lobes 5, ovate-lanceolate, 415 mm, acute.
Ovary 4-celled. Drupecetum mulberry-shaped, subglobose, or
ellipsoid-oblong, to 2.5 cm in diam. Drupes partially fused to
nearly separate, white or black, obovoid to subglobose, 1015
mm. Fl. AprMay, fr. summerautumn.
Forests; 5001400 m. S Yunnan [Bhutan, India, Laos, Myanmar,
Nepal, Thailand].
The inflorescences are indeterminate and begin flowering with a
dozen developed buds; the flowers continue to be produced through
growth at the top of the spike while the fruit are developing on the lower
parts, thus the older inflorescences become cylindrical or oblong in
shape. Y. Z. Ruan (in FRPS 71(2): 185. 1999) described the inflorescence position as consistently leaf-opposed at the terminal node; however, inflorescences can also be found along the stem well below its
apex (e.g., Tsi Zhanhou 92-130, MO), and the infructescences are
usually found along the stem well below the apex (e.g., Puff et al.,
Rubiaceae of Thailand, 113. 2005).

2. Morinda badia Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 327. 1999.
li se ba ji
Lianas; branches at base with persistent leafless stipules,
when young puberulent and terete, becoming glabrescent,
angled, dark brown or purplish blue. Leaves opposite; petiole
48 mm, densely hirtellous; blade drying papery, adaxially
brownish black, abaxially ferruginous, elliptic-oblong, ellipticlanceolate, or oblanceolate, 712 24 cm, adaxially shiny and
hirtellous or glabrescent, abaxially hirtellous at least along
veins, base cuneate to attenuate, apex acuminate; secondary
veins 46 pairs; stipules fused into a tube, 25 mm, truncate.
Inflorescence terminal; peduncles 35, umbellate, 518 mm,
hirtellous; heads 1 per peduncle, 310-flowered. Flowers fused
at base, biology not noted. Calyx limb reduced, truncate or
often with 13 teeth in outermost flowers of head. Corolla yellowish white; tube ca. 2.5 mm; lobes 4 or 5, narrowly oblong,

3. Morinda brevipes S. Y. Hu, J. Arnold Arbor. 32: 399. 1951.

duan bing ji yan teng
Lianas; young branches densely hirtellous or pilosulous,
becoming glabrous, brown. Leaves opposite; petiole 310 mm,
puberulent or hirtellous; blade drying papery or subleathery,
adaxially straw-yellow or brownish black, abaxially olivegreen, brownish yellow, or brownish red, obovate-oblong, obovate, oblanceolate, lanceolate, narrowly lanceolate, or linearlanceolate, 510(13) 0.73(4) cm, glabrous on both surfaces, shiny adaxially, matte abaxially, base cuneate to acute,
apex acute or acuminate; secondary veins 57 pairs, with
foveolate or pubescent domatia; stipules fused into a tube or
spathe, 23 mm, puberulent, hispidulous, or hirtellous, truncate,
on each side with 2 bristles 0.31 mm. Inflorescence terminal;
peduncles 49, fasciculate to shortly racemiform, 410 mm,
densely puberulent, as a group sometimes subtended by 24
stipuliform bracts; heads 1 per peduncle, conical or cylindrical
to subglobose, 46 mm, 616-flowered. Flowers fused for ca.
1/2 length of hypanthium, biology not noted. Calyx puberulent
to glabrescent; limb 0.30.5 mm, truncate to denticulate. Corolla white, campanulate, outside glabrous; tube ca. 2.5 mm; lobes
4 or 5, narrowly oblong, ca. 3 mm, apically thickened and rostrate. Drupecetum subglobose to cylindrical, 11.2 cm in diam.
Drupes almost fully fused, orange, subglobose, 58 mm. Fl.
AprMay, fr. JulDec.
Hills, mountains; 200800 m. Hainan.

1a. Leaf blade obovate-oblong, obovate,

oblanceolate, or lanceolate, 510(13)
23(4) cm ............................................... 3a. var. brevipes
1b. Leaf blade narrowly lanceolate to
lanceolate-linear, 711.5
0.71.4 cm ........................................... 3b. var. stenophylla
3a. Morinda brevipes var. brevipes
() duan bing ji yan teng (yuan bian zhong)
Leaf blade obovate-oblong, obovate, oblanceolate, or lanceolate, 510(13) 23(4) cm. Fl. AprMay, fr. JulDec.
Hills, mountains; 200800 m. Hainan.

3b. Morinda brevipes var. stenophylla Chun & F. C. How ex

W. C. Ko, Fl. Hainan. 3: 581. 1974.
xia ye ji yan teng
Leaf blade narrowly lanceolate to lanceolate-linear, 711.5
0.71.4 cm. Fl. May.
Wet places in forests on hills. Hainan.

RUBIACEAE

4. Morinda callicarpifolia Y. Z. Ruan, Fl. Reipubl. Popularis

Sin. 71(2): 325. 1999 [callicarpaefolia].
zi zhu ye ba ji
Lianas or subshrubs; branches at base with persistent leafless stipules, when young densely golden hirtellous, becoming
sparsely pubescent or subglabrous, terete or angled, brown or
purplish black. Leaves opposite; petiole 710 mm, densely
golden hirtellous; blade drying papery, adaxially brownish
black, abaxially brown, obovate-oblong, oblong-lanceolate, or
elliptic, 714 2.54 cm, adaxially sparsely hirtellous or hispidulous, abaxially sparsely hispidulous or hirtellous, or glabrous, on both surfaces pubescence denser along midrib, base
cuneate to acute, apex acuminate, long acuminate, or obtuse
then abruptly mucronate; secondary veins 57 pairs, with pilosulous domatia; stipules fused into a tube, 34 mm, densely
golden hispidulous or -hirtellous, subtruncate, on each side with
1 or 2 bristles. Inflorescence terminal; peduncles 47, umbellate
or fasciculate, 615 mm, densely hirtellous, as a group often
subtended by 1 subulate bract; heads 1 per peduncle, 28-flowered. Flowers fused at base, biology not noted. Calyx puberulent; limb reduced, truncate or sometimes with 1 subulate tooth.
Corolla white, tubular; tube ca. 2.5 mm, densely villous inside;
lobes 4, lanceolate. Drupecetum globose, 48 mm in diam.
Drupes fused. Fl. JunJul, fr. winter.
Forests on mountains, thickets at roadsides, ditch sides, hill
slopes. Guizhou, Sichuan, Yunnan.

5. Morinda cinnamomifoliata Y. Z. Ruan, Fl. Reipubl. Popularis Sin. 71(2): 336. 1999.
zhang ye ba ji
Lianas or subshrubs; branches at base with persistent leafless stipules, when young densely ferruginous hirtellous or -pilosulous, becoming glabrescent, angled, gray or indigo. Leaves
opposite; petiole 510 mm, densely hirtellous; blade drying
leathery, adaxially grayish black, abaxially whitened, narrowly
elliptic-oblong, oblong-lanceolate, or obovate-oblanceolate, 8
11 2.53.5 cm, adaxially ferruginous pilosulous, abaxially
hispidulous to hirtellous, base cuneate or rounded, apex acuminate or obtuse then abruptly mucronulate; secondary veins 4 or
5(or 6) pairs; stipules united into a tube. Inflorescence terminal;
peduncles 610, umbellate or fasciculate, 510 mm, ferruginous pilosulous; heads 1 per peduncle, 57-flowered. Flowers
fused at base, biology not noted. Calyx limb reduced, truncate
or denticulate. Corolla white, outside glabrescent; tube ca. 2
mm, inside densely villous; lobes 4, linear-oblanceolate, ca. 3
mm. Drupecetum subglobose. Drupes fully fused. Fl. Jul, fr.
winter.
Thickets on slopes near villages. SE Guangxi.

6. Morinda citrifolia Linnaeus, Sp. Pl. 1: 176. 1753.

hai bin mu ba ji
Morinda bracteata Roxburgh.
Evergreen shrubs or small trees, to 5 m tall, often fleshy;
branches subquadrangular, glabrous. Leaves opposite or solitary
opposite an inflorescence; petiole 520 mm, glabrous; blade
fleshy, drying papery, elliptic-oblong, elliptic, or ovate, 1025

223

513 cm, glabrous and shiny on both surfaces, base acute or

acuminate, apex acute to obtuse; secondary veins 57 pairs,
with pubescent domatia; stipules interpetiolar, free or shortly
fused to petioles, broadly triangular to ovate, 416 mm, obtuse
or rounded. Inflorescence solitary and leaf-opposed; peduncle
11.5 cm; head 1, oblong to subglobose, 510 mm in diam.,
many flowered; bracts absent. Flowers with hypanthia partially
fused, distylous. Calyx glabrous or puberulent; limb subtruncate
to truncate, 0.20.5 mm, sometimes in 1 to numerous flowers of
a head with 1(3) calycophylls, these white, narrowly elliptic to
oblanceolate, 516 mm, obtuse to acute. Corolla white, funnelform, outside glabrous; tube ca. 15 mm, densely villous in
throat; lobes 5, ovate-lanceolate, ca. 6 mm. Drupecetum white,
irregularly ovoid to subglobose, 2.55 cm. Drupes not distinguishable individually. Fl. and fr. year-round.
Flat land on seashores, sparse forests; below 100 m. Guangdong,
Hainan, Taiwan [?Cambodia, India, Indonesia, Japan (Bonin and Ryukyu Islands), Malaysia, Myanmar, Papua New Guinea, Philippines, Sri
Lanka, Thailand, Vietnam; N Australia, Solomon Islands; introduced in
tropical America and Pacific islands].
The distinctive form called Morinda bracteata has well-developed
white calycophylls that give the plants a markedly different appearance
and may function in pollination, but these two forms have generally
been considered conspecific. Nelson and Elevitch (Noni, 4243. 2006)
noted that plants with bracteate inflorescences produce smaller fruit and
that the cultivated plants with variegated leaves are called M. citrifolia
Potteri. Both of these forms are found in Taiwan (e.g., Yang & Chuang
11410, MO, citifolia form; Yang & Chuang 12060, MO, bracteata
form). The fruit of this species are edible (though not particularly palatable) and said to have medicinal and/or tonic value; they are sold by
natural food vendors under the name noni or nona. This species is
increasingly widely cultivated, as detailed by Nelson and Elevitch (loc.
cit.).

7. Morinda citrina Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 331. 1999.
jin ye ba ji
Lianas; branches at base with persistent leafless stipules,
when young densely puberulent, strigillose, or hirtellous, becoming glabrescent, angled, brown, purplish blue, or purplish
black. Leaves opposite; petiole 26 mm, densely puberulent,
hirtellous, or pilosulous; blade drying membranous to papery,
adaxially gray, abaxially golden yellow or brownish yellow,
matte on both surfaces, ovate-lanceolate, oblong-linear, or elliptic, 512 1.53 cm, adaxially sparsely to moderately puberulent, strigillose, hispidulous, or hirtellous, abaxially moderately
to sparsely strigillose, puberulent, or hirtellous especially along
principal veins, base cuneate to acute, apex acuminate or acute;
secondary veins 46 pairs, with pilosulous domatia; stipules
fused into a tube or spathe, 2.54 mm, moderately to densely
strigillose to pilosulous, truncate, on each side with 1 or 2 bristles 0.31 mm. Inflorescence terminal; peduncles 25, fasciculate or umbellate, 68 mm, densely puberulent to pilosulous, as
a group subtended by 1 or 2 stipuliform bracts; heads 1 per peduncle, hemispherical, 45 mm in diam., 15-flowered. Flowers fused at base or for up to half of hypanthium, biology not
noted. Calyx strigillose, puberulent, or glabrescent; limb 0.3
0.5 mm, truncate. Corolla white, salverform, outside densely
puberulent; tube 1.22 mm, inside villous; lobes 4(or 5), narrowly oblong to lanceolate, 2.53 mm, apically thickened and

RUBIACEAE

224

rostrate. Drupecetum subglobose to oblate, 812 mm in diam.

Drupes fully fused, orange, subglobose, 35 mm. Fl. AprJul,
fr. OctNov.
Forests or thickets on mountains; 5001300 m. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hunan, Jiangxi, Zhejiang.

1a. Older leaves usually drying greenish,

sometimes brown, or rarely yellow,
usually not hirsutulous but rather thin
and sparsely hairy along midrib ................ 7a. var. chlorina
1b. Older leaves usually drying golden
yellow or brownish yellow, hirtellous
especially along midrib ............................... 7b. var. citrina
7a. Morinda citrina var. chlorina Y. Z. Ruan, Fl. Reipubl.
Popularis Sin. 71(2): 332. 1999.
bai rui ba ji
Older leaves usually drying greenish, sometimes brown,
or rarely yellow, usually not hirsutulous but rather thin and
sparsely hairy along midrib.
Forests or thickets on mountains. Anhui, Fujian, Guangxi, Guizhou, Hunan, Jiangxi, Zhejiang.

7b. Morinda citrina var. citrina

() jin ye ba ji (yuan bian zhong)
Older leaves usually drying golden yellow or brownish
yellow, hirtellous especially along midrib. Fl. AprJul, fr. Oct
Nov.
Sparse or dense forests on mountains; 5001300 m. Guangdong, Guangxi, Guizhou, Hunan.

8. Morinda cochinchinensis Candolle, Prodr. 4: 449. 1830.

da guo ba ji
Morinda trichophylla Merrill.
Lianas; branches at base with persistent leafless stipules,
when young densely ferruginous- or yellow villosulous, terete
to weakly quadrangular. Leaves opposite; petiole 310 mm,
densely villosulous; blade drying papery, yellowed, matte to
shiny adaxially, matte abaxially, elliptic, elliptic-oblong, obovate-oblong, or oblanceolate, 5.514 26 cm, adaxially
sparsely strigose to strigillose, abaxially densely ferruginous- or
yellow hirtellous to villosulous with pubescence denser along
veins, base rounded to truncate, subcordate, or cordulate, apex
caudate-acuminate or shortly acuminate; secondary veins 710
pairs, with pilosulous domatia; stipules fused into a tube or
spathe, 59 mm, densely hispidulous to hispid, broadly triangular to truncate, on each side with 2 bristles 14 mm, usually
quickly deciduous. Inflorescence terminal; peduncles 312, fascicled or umbellate, 13 cm, densely ferruginous- or yellow hirtellous, as a group subtended by 2 to several bracts 13 mm, 2to several lobed; heads 1 per peduncle, subglobose, 56 mm in
diam., 515-flowered; bracteoles linear, 0.21 mm. Flowers
with hypanthia partially fused, biology not noted. Calyx with
hypanthium portion densely strigose to strigillose; limb 12.5
mm, puberulent to strigillose, lobed for ca. half, lobes 4 or 5,
narrowly triangular, 12 mm, sometimes unequal on an in-

dividual flower, often reflexed. Corolla white, rotate to salverform, outside hirtellous, pilosulous, or glabrescent, inside
densely villous throughout tube and onto lobes; tube 1.52 mm;
lobes 4 or 5, narrowly oblong to lanceolate, 44.5 mm, apically
thickened and rostrate. Drupecetum subglobose, oblong-globose, or irregular, 12 cm in diam., with peduncles elongating
to 4 cm. Drupes mostly fused, orange-yellow to orange-red,
subglobose, 46 mm. Fl. MayJul, fr. JulNov.
Forests or thickets on mountain slopes, in valleys, or at streamsides or roadsides; 1001200 m. Fujian, Guangdong, Guangxi, Hainan
[Vietnam].
The names Morinda umbellata and M. villosa were synonymized
with M. cochinchinensis by Merrill and Chun (Sunyatsenia 1(1): 8081.
1930) but were separated by Y. Z. Ruan (in FRPS 71(2): 187, 190.
1999) as provisionally treated here.

9. Morinda hainanensis Merrill & F. C. How, Sunyatsenia 5:

188. 1940.
hai nan ba ji
Lianas; branches at base with persistent leafless stipules,
when young densely yellow-, clear-, or ferruginous strigillose to
-hirtellous, becoming scabrous to glabrescent, brownish gray.
Leaves opposite; petiole 511 mm, densely villosulous to hirtellous; blade drying papery, matte and gray on both surfaces,
narrowly elliptic, elliptic-oblong, oblong-lanceolate, or linearlanceolate, 59 1.22.5 cm, adaxially sparsely strigillose to
hirtellous, abaxially densely villosulous, pilosulous, or sericeous, base acute or attenuate, apex acute to acuminate; secondary
veins 69 pairs, with pilosulous domatia; stipules fused into a
tube or spathe, 2.56 mm, pilosulous to hirtellous, on each side
with 2 bristles 0.21 mm. Inflorescence terminal; peduncles 3
9, fasciculate to umbellate, 510 mm, densely sericeous to
strigillose, as a group sometimes subtended by 1 or 2 linear
bracts 0.51 mm; heads 1 per peduncle, hemispherical, 34 mm
in diam., 38-flowered. Flowers fused at base, apparently
distylous. Calyx with hypanthium puberulent to densely strigillose; limb 0.50.8 mm, glabrescent, truncate. Corolla salverform, outside puberulent to pilosulous, inside densely villous in
upper part of tube and onto lobes; tube ca. 1 mm; lobes 4,
narrowly lanceolate, ca. 2 mm, apically thickened and rostrate.
Drupecetum subglobose or compressed globose, 68 mm in
diam. Drupes fully fused, subglobose, ca. 3 mm. Fl. May, fr.
May.
Wet places under dense forests on hills; ca. 900 m. Hainan.

10. Morinda howiana S. Y. Hu, J. Arnold Arbor. 32: 400.

1951.
kang teng
Lianas; branches at base with persistent leafless stipules,
glabrous, terete, drying brown. Leaves opposite; petiole 610
mm, glabrous to puberulent; blade drying thinly to thickly papery, adaxially shiny and reddish dark brown, abaxially matte to
somewhat shiny and reddish brown, elliptic, elliptic-lanceolate,
or oblong, 614 26 cm, glabrous, base rounded, cuneate, or
acute, apex acute to acuminate; secondary veins 69 pairs, with
pilosulous domatia; stipules fused into a tube, 515 mm, puberulent to glabrous, truncate, on each side with 2 bristles 0.21

RUBIACEAE

225

mm. Inflorescence terminal; peduncles 510, fasciculate to umbellate, 818 mm, puberulent, as a group subtended by several
triangular to bifid bracts 12 mm; heads 1 per peduncle, subglobose to hemispherical, 46 mm in diam., 412-flowered.
Flowers fused only shortly at base, biology not noted. Calyx
puberulent to glabrescent; limb ca. 1 mm, truncate, sinuate, or
shallowly lobed; lobes rounded, ciliolate. Corolla subcampanulate, puberulent outside; tube ca. 2 mm, inside densely villous
in upper part and throat; lobes 4 or 5, narrowly oblong to lanceolate, ca. 2 mm, apically thickened and rostrate. Drupecetum
subglobose to oblate, 814 mm in diam. Drupes fully fused,
subglobose, 45 mm. Fl. AprMay, fr. JulOct.

veins 6 or 7 pairs, with pilosulous foveolate domatia; stipules

caducous, interpetiolar or fused into a spathe or tube, 317 mm,
glabrous, obtuse, rounded, or truncate. Inflorescence terminal,
pilosulous [to perhaps glabrous]; peduncles 13, 0.58 cm,
terminating in 1 head or 620 rays 0.82 cm and subtended by
linear bracts 13 mm; heads several, subglobose, 810 mm in
diam., 620-flowered. Flowers fused for half or more of hypanthium, biology not noted. Calyx glabrous; limb reduced, truncate to shallowly 4-dentate. Corolla white, subcampanulate;
tube 22.5 mm, villous in upper part inside; lobes 4, 22.5 mm.
Drupecetum subglobose to oblate, 411 mm in diam. Fl. Jun, fr.
Dec.

Forests in valleys and at streamsides, thickets at roadsides or on

hill slopes; 300700 m. Guangdong (Yangjiang), Hainan.

Shady places in forests at streamsides or roadsides; 10001100 m

[to only ca. 700 m in Thailand]. Yunnan [Malaysia, Thailand].

The protologue commented that the presence of bristles on the

glabrous capitulum is also very characteristic, but these bristles are
apparently small bracts, which are now also known from several other
Chinese species of Morinda.

13. Morinda leiantha Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 41: 313. 1872.

11. Morinda hupehensis S. Y. Hu, J. Arnold Arbor. 32: 400.

1951.

Erect shrubs or small trees; branches hirtellous or subglabrous, angled, dark brown. Leaves opposite; petiole 0.51(3)
cm; blade drying thinly papery, matte and dark brown on both
surfaces, oblong-lanceolate, narrowly lanceolate, rhombic-lanceolate, or oblanceolate, 814(20) 25 cm, adaxially glabrous or pilosulous, abaxially scabrous, glabrous, or pilosulous
along veins, base acute to attenuate, apex acuminate; secondary
veins 58 pairs; stipules ovate-triangular to ovate-lanceolate,
acute or 2-lobed. Inflorescence terminal or axillary; peduncle
25 mm, often with 24 branches, each with 1 capitulum; capitulum ellipsoid-oblong, many flowered. Flowers partially fused,
each with 35 bracts in a whorl, biology not noted. Calyx limb
truncate. Corolla white, salverform, outside glabrous; tube ca.
20 mm, inside glabrous; lobes 5, lanceolate, ca. 4 cm. Young
drupecetum mulberry-shaped. Fl. and fr. year-round.

hu bei ba ji
Lianas; branches at base surrounded by persistent leafless
stipules, when young densely white hirtellous, white puberulent, ferruginous hirtellous, or glabrescent, becoming angled,
purplish blue. Leaves opposite; petiole 48 mm, densely hirtellous; blade drying papery, matte on both surfaces, adaxially
greenish brown, abaxially coffee-colored or brownish black,
oblanceolate, elliptic-lanceolate, elliptic-oblong, or linear-oblong, 59 1.53 cm, adaxially sparsely hispidulous to glabrescent, abaxially ferruginous hirtellous, yellow hirtellous, or
brownish gray hirtellous, base cuneate, obtuse, or acute, apex
acuminate or sometimes obtuse then abruptly mucronulate;
secondary veins 57 pairs, with pilosulous domatia; stipules
frequently deciduous through fragmentation, fused into a tube
or spathe, 2.55 mm, pilosulous to hirtellous, truncate, on each
side with 2 bristles 0.20.5 mm. Inflorescence terminal; peduncles 49, fasciculate to umbellate, 515 mm, densely puberulent
to pilosulous, as a group subtended at base by 14 stipuliform
bracts; heads 1 per peduncle, 47-flowered. Flowers with hypanthia partially fused, biology not noted. Calyx puberulent to
glabrescent; limb reduced, truncate or sinuate-undulate. Corolla
white; tube ca. 2.5 mm, densely villous inside; lobes 4 or 5, ca.
2.5 mm. Drupecetum subglobose, 410 mm in diam. Drupes
fully fused, red, subglobose, 45 mm. Fl. JulAug, fr. OctNov.
Forests, thickets at forest margins; 4001000 m. Fujian, Guangxi, Guizhou, Hubei, Hunan, Sichuan.

12. Morinda lacunosa King & Gamble, J. Asiat. Soc. Bengal,

Pt. 2, Nat. Hist. 73: 87. 1903.
chang xu yang jiao teng
Lianas, to 20 m tall; branches when young terete, glabrous, becoming angled, brownish gray. Leaves opposite; petiole 820 mm, glabrous; blade drying thinly papery or subleathery, adaxially brownish black, abaxially dark brownish red,
elliptic, elliptic-lanceolate, or ovate-oblong, 1016 37 cm,
glabrous, base cuneate, apex acuminate or acute; secondary

ding hua mu ba ji

Forests on hill slopes, shady thickets. S Yunnan [Myanmar].

14. Morinda litseifolia Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 335. 1999.
mu jiang ye ba ji
Lianas or subshrubs; branches at base surrounded by persistent leafless stipules, when young sparsely puberulent, becoming glabrescent, purplish black. Leaves opposite; petiole 4
6 mm, hirtellous; blade drying membranous or papery, adaxially gray, grayish green, or grayish black, abaxially brownish
gray or greenish red, linear, oblong-linear, lanceolate-linear, oblanceolate, or elliptic-oblanceolate, 511 1.52(3) cm, adaxially glabrous or sparsely pilosulous, abaxially usually glabrous
except hirtellous along veins, base cuneate to acute, apex acuminate or acute; secondary veins 46; stipules fused into a
spathe or tube, ca. 4 mm. Inflorescences terminal; peduncles
35, fasciculate or umbellate, ca. 7 mm, puberulent; heads 1
per peduncle, subglobose, 57-flowered. Flowers fused at base,
biology not noted. Calyx limb reduced, truncate. Corolla yellowish white; tube ca. 2 mm, densely villous inside; lobes 4
or 5, ca. 2 mm. Drupecetum subglobose, 28 mm in diam.
Drupes almost fully fused. Fl. Jul, fr. OctNov.
Sparse or dense forests on mountains; 7001300 m. Fujian,
Guangxi, Hunan, Jiangxi, Sichuan.

RUBIACEAE

226

15. Morinda longissima Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 323. 1999.
da hua mu ba ji
Erect shrubs, to 2 m tall; branches glabrous or subglabrous, subquadrangular, green or pale green. Leaves opposite,
sometimes crowded at ends of branches; petiole 23 cm, glabrous or subglabrous; blade drying submembranous to membranous, pale green and matte on both surfaces, oblong-oblanceolate, oblong-lanceolate, or oblanceolate, 1323 610 cm,
glabrous or pilosulous abaxially, base cuneate to attenuate, margin sometimes weakly repand or sinuate, apex acuminate or
obtuse then abruptly acuminate; secondary veins 810 pairs,
apparently without domatia; stipules interpetiolar, ovate, semicircular, or subcordate, acute or 2-lobed. Inflorescence terminal
or axillary; peduncle 1, ca. 2 cm, glabrous; heads 1 per peduncle, subglobose, many flowered; bracts encircling flowers in
1 or 2 whorls, 36 per whorl, subulate. Flowers shortly fused at
base, biology not noted. Calyx limb truncate to sinuate. Corolla
white, slenderly funnelform to salverform, glabrous outside;
tube ca. 32 mm, glabrous inside; lobes 5, lanceolate to ligulate,
ca. 13 mm. Infructescences and fruit unknown. Fl. AprMay.
Sparse forests, shady thickets; ca. 700 m. S Yunnan.
The bracts arranged in whorls subtending the flowers are notable
and probably distinctive for the species; however, these bracts are not
shown in the illustrations of this species in the protologue nor in the
additional figure presented in the FRPS Morinda treatment (Y. Z. Ruan,
loc. cit.: 184, t. 46, f. 5).

16. Morinda nanlingensis Y. Z. Ruan, Fl. Reipubl. Popularis

Sin. 71(2): 329. 1999.
nan ling ji yan teng
Lianas or subshrubs; branches surrounded at base by persistent leafless stipules, when young hirtellous or hispidulous,
becoming glabrescent, angled, purplish blue. Leaves opposite;
petiole 35 mm, hirtellous; blade drying papery, adaxially
greenish brown or dark brown, abaxially brownish gray, obovate, elliptic-oblanceolate, elliptic, or rhombic-lanceolate, rarely
dimorphic, 49(12) 1.53.5 cm, adaxially pilosulous to glabrescent, abaxially glabrous to densely hirtellous at least on
principal veins, base cuneate, apex acute, acuminate, or obtuse
then abruptly mucronulate; secondary veins 57 pairs; stipules
fused into a tube, truncate. Inflorescence terminal; peduncles 7
10, umbellate or fasciculate, 1015 mm, pilosulous; heads 1 per
peduncle, 311-flowered. Flowers fused at base, biology not
noted. Calyx limb reduced, truncate or perhaps sometimes with
1 calycophyll. Corolla campanulate, mealy puberulent outside;
tube ca. 2 mm, internally densely bearded; lobes 4 or 5, sublanceolate, ca. 3 mm. Drupecetum subglobose, 48 mm in diam.
Drupes orange or dark. Fl. Jun, fr. Oct.
Shady places, forests, thickets on mountains, hills. Guangdong,
Guangxi, Hunan, Yunnan, Zhejiang.

1a. Flowering heads with 3 or 4(7) flowers;

leaf blade rhombic-lanceolate, 47
1.52 cm .............................................. 16b. var. pauciflora
1b. Flowering heads with 511 flowers;
leaf blade obovate, elliptic-oblanceolate,
or elliptic, 712 23.5 cm.

2a. Leaf blade glabrous abaxially

.................................................. 16a. var. nanlingensis
2b. Leaf blade densely pilosulous
abaxially ....................................... 16c. var. pilophora
16a. Morinda nanlingensis var. nanlingensis
() nan ling ji yan teng (yuan bian zhong)
Leaf blade obovate, elliptic-oblanceolate, or elliptic, 79(
12) 23.5 cm, glabrous abaxially. Flowering heads 511flowered. Fl. Jun, fr. Oct.
Shady places at streamsides in forests, thickets on mountains.
Guangdong, Guangxi, Hunan, Yunnan.

16b. Morinda nanlingensis var. pauciflora Y. Z. Ruan, var.

nov.
shao hua ji yan teng
Type: China. Zhejiang: Hangzhou, X. Y. He 20528 (holotype, IBSC).
Validating Latin diagnosis: that of Morinda nanlingensis
Y. Z. Ruan va. [sic!] pauciflora Y. Z. Ruan (Y. Z. Ruan in W.
C. Chen, Fl. Reipubl. Popularis Sin. 71(2): 331. 1999).
Leaf blade rhombic-lanceolate, 47 1.52 cm. Flowering
heads 3- or 4(7)-flowered.
Forests on hills. S Zhejiang.
This name was previously published by Y. Z. Ruan (loc. cit.) but
not validly so because the herbarium in which the type is conserved was
not specified (Vienna Code, Art. 37.7).

16c. Morinda nanlingensis var. pilophora Y. Z. Ruan, Fl.

Reipubl. Popularis Sin. 71(2): 331. 1999.
mao bei ji yan teng
Leaf blade obovate, elliptic-oblanceolate, or elliptic, 79(
12) 23.5 cm, abaxially densely pilosulous. Flowering heads
511-flowered.
Forests, shady thickets on mountains. Guangxi, Hunan.

17. Morinda officinalis F. C. How, Acta Phytotax. Sin. 7: 326.

1958.
ba ji tian
Lianas; branches surrounded at base by persistent leafless
stipules, when young strigillose, hirtellous, or pilose, becoming
glabrescent and scabrous, angled, brown or bluish black. Leaves
opposite; petiole 411 mm, densely puberulent, strigillose, hirtellous, or hirsute to glabrescent; blade drying papery, on both
surfaces brown to yellow-brown, shiny to matte adaxially,
matte abaxially, ovate-oblong, obovate-oblong, or elliptic, 613
36 cm, adaxially sparsely strigillose, hirtellous, or hirsute to
glabrescent, abaxially glabrous or sparsely hirtellous along principal veins, base obtuse, rounded, cuneate, or acute, apex acute,
obtuse, or rounded and abruptly mucronulate; secondary veins
(4 or)57 pairs, with small pilosulous domatia; stipules fused
into a spathe or tube, 35 mm, membranous, puberulent to hirtellous, truncate, on each side 2-denticulate. Inflorescence terminal; peduncles 17 or 1525, umbellate or fasciculate, 0.11

RUBIACEAE

cm, densely hirtellous to strigillose, as a group usually subtended by 1 or 2 stipuliform bracts; heads 1 per peduncle, subglobose to hemispherical, 57 mm in diam., 13- or 410-flowered. Flowers fused for ca. half of hypanthium, biology not
noted. Calyx puberulent to glabrous; limb 11.5 mm, lobed for
ca. 1/2; lobes 24, triangular, sometimes markedly unequal on
an individual flower, obtuse to acute. Corolla white, campanulate or urceolate, outside puberulent, hirtellous, or glabrescent;
tube 34 mm, inside densely villosulous from middle of tube to
throat; lobes (2)4, lanceolate or narrowly oblong, 34 mm,
apically thickened and rostrate. Drupecetum globose to oblate,
511 mm in diam. Drupes fully fused, red, subglobose, 45
mm. Fl. MayJul, fr. OctNov.
Sparse or dense forests and thickets on mountains, also cultivated; 100500 m. Fujian, Guangdong, Guangxi, Hainan.
According to the protologue, the species is apparently cultivated
and used medicinally. The roots were described by Y. Z. Ruan (in FRPS
71(2): 199. 1999) as fleshy, irregularly intestine-like constricted, slightly
purplish red, purplish blue when dry; they are illustrated in the protologue figure. The roots of almost no other Morinda species have been
described by Y. Z. Ruan or any other authors seen.

1a. Flowering heads 1525, each with 1(3)

flowers; peduncles ca. 2 mm; calyx
lobes often becoming white ........................ 17c. Uniflora
1b. Flowering heads 110, each with 410
flowers; peduncles 110 mm; calyx
lobes green.
2a. Young branches and leaf blades
abaxially sparsely hirtellous,
puberulent, or hirsute to
glabrescent ................................... 17b. var. officinalis
2b. Young branches and leaf blades
densely transparent villous ............... 17a. var. hirsuta
17a. Morinda officinalis var. hirsuta F. C. How, Acta Phytotax. Sin. 7: 328. 1958.
mao ba ji tian
Young branches and leaf blade densely transparent villous.
Flowering heads 110, 410-flowered; peduncles 110 mm.
Calyx lobes green.
Forests on mountains. Hainan.

17b. Morinda officinalis var. officinalis

() ba ji tian (yuan bian zhong)
Young branches and leaf blades adaxially sparsely hirtellous, puberulent, or hirsute to glabrescent. Flowering heads 1
10, 410-flowered; peduncles 110 mm. Calyx lobes green. Fl.
MayJul, fr. OctNov.
Sparse or dense forests and thickets on mountains. Fujian,
Guangdong, Guangxi, Hainan.

17c. Morinda officinalis Uniflora

mi geng ba ji tian
Flowering heads 1525, 1(3)-flowered; peduncles ca. 2
mm. Calyx lobes often becoming white.

227

Cultivated in Guangdong.
This taxon was originally published as a cultivar (Y. Z. Ruan in W.
C. Chen, Fl. Reipubl. Popularis Sin. 71(2): 337. 1999), but it was not
made clear if it is a well-known form that is deliberately selected and
propagated or a wild-encountered form.

18. Morinda parvifolia Bartling ex Candolle, Prodr. 4: 449.

1830.
ji yan teng
Lianas, climbing, twining, or prostrate; branches surrounded at base by persistent leafless stipules, when young glabrous, puberulent, or densely hispidulous-hirtellous, becoming
weakly angled, brown to slightly purplish blue. Leaves opposite; petiole 38 mm, glabrous, puberulent, or hispidulous-hirtellous; blade drying papery, on both surfaces matte and yellowish green, yellowish brown, or yellowish gray, obovate, linear-oblanceolate, sublanceolate, obovate-oblanceolate, oblanceolate, or obovate-oblong, 25(7) 0.33 cm, both surfaces
glabrous or infrequently moderately to sparsely hirsute or hirtellous, base cuneate to acute, apex acute, obtuse, or rounded
and shortly abruptly acuminate or mucronulate; secondary veins
3 or 4(6) pairs, with pilosulous domatia; stipules fused into a
spathe or tube, 24 mm, membranous, glabrous, puberulent, or
hispidulous, truncate and on each side with 1 or 2, sometimes
caducous bristles 0.51 mm. Inflorescence terminal; peduncles
(2 or)39, umbellate to fasciculate, 0.62 cm, densely puberulent to hispidulous, as a group often subtended by 1 to several
stipuliform bracts; heads 1 per peduncle, subglobose, oblate,
conical, or rarely cylindrical, 58 mm in diam., 315(17)-flowered, often with 1 to several linear bracts 0.51 mm. Flowers
fused at base, biology not noted. Calyx puberulent to glabrescent; limb 0.50.8 mm, truncate to sinuate or denticulate. Corolla white, campanulate, outside puberulent to glabrous; tube
1.52 mm, inside densely villous from middle to throat; lobes 4
or 5, narrowly oblong, 34 mm, apically thickened and rostrate.
Drupecetum subglobose to oblate, 615 mm in diam. Drupes
fully fused, orange, subglobose, 45 mm. Fl. AprJun, fr. Jun
Aug.
Thickets at roadsides or ditch sides, prostrate on bare land, thickets or forests on hills; sea level to 400 m. Fujian, Guangdong, Guangxi,
Hainan, Jiangxi, Taiwan [Philippines, Vietnam].
Y. Z. Ruan (in FRPS 71(2): 194. 1999) commented that leaf shape
in this species varies based to some degree on ecology, with obovate
and dimorphic leaves in dry, sunny, bare habitat; linear-oblanceolate or
sublanceolate leaves in shady, dry, bare habitat; and obovate-oblanceolate, oblanceolate, or obovate-oblong leaves when climbing on shrubs.

19. Morinda persicifolia Buchanan-Hamilton, Trans. Linn.

Soc. London 13: 535. 1822.
duan geng mu ba ji
Shrubs or small trees, to 11 m tall; branches quadrangular,
pubescent or glabrous. Leaves opposite or solitary opposite an
inflorescence; petiole 12 cm; blade drying papery, adaxially
gray, abaxially pale green, oblong-lanceolate, lanceolate, oblanceolate, or linear-lanceolate, 618 212 cm, glabrous or
sparsely pilosulous, base acute to attenuate, apex acute or
shortly acuminate; secondary veins 710 pairs; stipules inter-

RUBIACEAE

228

petiolar, triangular to broadly triangular, acute to subulate acuminate. Inflorescence solitary and leaf-opposed; peduncle 1, 1
3 mm; head 1, many flowered; bracteoles subulate. Flower biology not noted. Calyx limb truncate. Corolla white, salverform;
tube ca. 16 mm, somewhat curved, inside glabrous; lobes 5, ca.
4 mm. Drupecetum mulberry-shaped or conical-oblong, 22.5
1.52 cm. Fl. and fr. summerautumn.
Sparse forests on mountains. S Yunnan [Cambodia, NE India,
Indonesia, Laos, Malaysia, Myanmar, Vietnam].

20. Morinda pubiofficinalis Y. Z. Ruan, Fl. Reipubl. Popularis

Sin. 71(2): 334. 1999.
xi mao ba ji
Lianas or subshrubs, 210 m tall; branches when young
sparsely hirtellous, becoming subglabrous, weakly angled,
brown or pale brownish purple. Leaves opposite; petiole ca. 4
mm, hirtellous or pilosulous; blade drying papery, adaxially
shiny and dark brownish red or yellowish brown, abaxially pale
brownish red or brownish yellow, oblong-lanceolate, linearlanceolate, or ovate-oblong, 713 24 cm, adaxially glabrous
or sometimes sparsely hirtellous when young, abaxially glabrous, base cuneate or rounded, apex acute to acuminate; secondary veins 5 or 6 pairs; stipules fused into a tube, ca. 4 mm,
subtruncate. Inflorescence terminal; peduncles 25, umbellate,
711 mm; heads 1 per peduncle, 3- to many flowered. Flowers
fused at base, biology not noted. Calyx limb reduced, truncate.
Corolla white, campanulate to weakly urceolate, outside glabrous; tube ca. 1.2 mm, inside densely bearded; lobes 4 or 5,
linear, ca. 2 mm. Drupecetum subglobose, 510 mm in diam.
Drupes fully fused, red. Fl. MayJun, fr. OctNov.
Forests in valleys or on mountains, thickets at watersides.
Guangdong, Guizhou, Hunan.
The roots were described by Y. Z. Ruan as irregularly intestinelike constricted, with fleshy cortex.

21. Morinda rosiflora Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 324. 1999.
hong mu ba ji
Erect shrubs, ca. 2.5 m tall; branches glabrous, quadrangular, purplish black. Leaves opposite or solitary opposite an inflorescence; petiole 24 cm; blade matte on both surfaces,
adaxially purplish black or purplish red, abaxially dark brown,
elliptic-oblong, elliptic, or oblong-oblanceolate, 1322 511
cm, adaxially glabrous, abaxially scabrous, base acute to attenuate, margin sometimes weakly sinuate, apex acuminate or
mucronulate; secondary veins 1014 pairs; stipules interpetiolar, triangular, ovate, or ovate-lanceolate, acute, acuminate, or
bifid. Inflorescence leaf-opposed; peduncle 1(or 2), 12 cm,
glabrous; heads 1 per peduncle, globose, many flowered; bracteoles 35, subulate. Flowers fused at base, biology not noted.
Calyx limb truncate. Corolla pink, salverform, outside glabrous;
tube ca. 22 mm, somewhat curved, glabrous inside; lobes 4 or
5, lanceolate, ca. 4 mm. Young drupecetum globose, ca. 1 cm in
diam. Young drupes dark red. Fl. summer, fr. autumn.
Sparse forests on hill slopes; 500800 m. S Yunnan.
The protologue gave the number of corolla lobes as 5, but the protologue figure shows 4 corolla lobes.

22. Morinda rugulosa Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 328. 1999.
zhou mian ji yan teng
Lianas or subshrubs, to 6 m tall; branches when young
pilosulous, becoming glabrescent. Leaves opposite; petiole 69
mm, hirtellous; blade drying papery, adaxially brownish black,
abaxially ferruginous, obovate or obovate-lanceolate, 79
2.54 cm, adaxially rugulose and sparsely pilosulous to glabrescent, abaxially glabrous except hirtellous along midrib, base cuneate or attenuate, apex rounded, obtuse, or acute; secondary
veins 5 or 6 pairs, with pilosulous domatia; stipules fused into a
tube, 34 mm, truncate. Inflorescence terminal; peduncles 48,
umbellate, 715 mm, pilosulous; heads 1 per peduncle, 512flowered. Flowers fused in lower half of hypanthium, biology
not noted. Calyx limb reduced. Corolla white, campanulate;
tube ca. 1 mm, inside densely bearded; lobes 4, narrowly oblong, ca. 3 mm, apically rostrate. Drupecetum subglobose, 410
mm in diam. Drupes fully fused, red. Fl. Jul, fr. Dec.
Forests at riversides and roadsides, thickets. N Guangxi, SW
Hunan.

23. Morinda scabrifolia Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 332. 1999.
xi nan ba ji
Lianas or shrubs; branches when young densely hirtellous,
becoming glabrescent, angled, whitened, pale purplish blue,
or brown. Leaves opposite; petiole 515 mm, hirtellous; blade
drying papery or leathery, adaxially brownish yellow or
brownish gray, abaxially pale brown, elliptic-oblong, oblonglanceolate, or oblong-linear, 713 24 cm, adaxially sparsely
hirtellous, subglabrous, or glabrous, abaxially glabrous or strigillose-pilosulous at least on principal veins, base cuneate or
rounded, apex acute, acuminate, or obtuse then abruptly mucronulate; secondary veins 57 pairs; stipules 35 mm. Inflorescence terminal; peduncles 310, fasciculate or umbellate, 520
mm, puberulent, as a group sometimes subtended by 1 linear
bract; heads 1 per peduncle, 510-flowered. Flowers fused at
base, biology not noted. Calyx limb reduced, truncate. Corolla
white, urceolate to campanulate, outside puberulent; tube ca.
2.5 mm, inside densely bearded; lobes 4 or 5, linear, ca. 2.5
mm. Drupecetum subglobose, ca. 1 cm in diam. Fl. Jun, fr.
SepOct.
Forests or thickets on mountains, shady rock sides. Guangxi,
Jiangxi, Sichuan, Yunnan.

24. Morinda shuanghuaensis C. Y. Chen & M. S. Huang, J.

Trop. Subtrop. Bot. 16: 578. 2008.
jia ba ji
Lianas; branches when young hirtellous, becoming glabrescent, angled, bluish black. Leaves opposite; petiole 35
mm, hirtellous; blade drying papery or thinly papery and brown,
elliptic, elliptic-oblong, or obovate-oblong, 410(13) 2.55
cm, adaxially hirsute to pubescent or sparsely hispidulous,
abaxially glabrous or sparsely pilosulous along midrib, base
rounded or cuneate, apex acute or mucronulate; secondary veins
5 or 6 pairs; stipules 35 mm, pilosulous, truncate. Inflores-

RUBIACEAE

cence terminal; peduncles 48, umbellate, 315 mm, pubescent;

heads 1 per peduncle, 38-flowered. Flowers fused at base, biology not noted. Calyx limb reduced, truncate or with 2 or 3
undulate teeth. Corolla white, urceolate, outside puberulent; tube
ca. 3 mm, inside densely bearded; lobes 3 or 4, narrowly oblong, ca. 3 mm, apically thickened and rostrate. Drupecetum
compressed globose. Drupes red. Fl. MayJul, fr. Oct.
Forests on mountains. Fujian, Guangdong.
Y. Z. Ruan (in FRPS 71(2): 201. 1999) described the roots as
irregularly intestine-like constricted, with cortex more or less fleshy.

25. Morinda umbellata Linnaeus, Sp. Pl. 1: 176. 1753.

yin du yang jiao teng
Lianas, climbing or twining; branches glabrous to puberulent or hispidulous, becoming weakly angled, often channeled,
bluish black to reddish brown. Leaves opposite; petiole 46
mm, glabrous, puberulent, or sparsely hirsute; blade drying
papery, leathery, or rigid-membranous, adaxially shiny and
greenish, pale brown, or brownish black, abaxially matte,
greenish, pale brown, or straw-colored, ovate, obovate-lanceolate, obovate-oblong, lanceolate, or linear-lanceolate, 69 2
3.5 cm, both surfaces glabrous or sometimes hirsute, hispidulous, or puberulent along principal veins, base acute or cuneate,
apex acuminate or mucronulate; secondary veins 4 or 5 pairs,
usually with pilosulous domatia; stipules fused into a tube, 26
mm, scarious to membranous, puberulent, broadly rounded to
truncate, on each side with 2 bristles 0.51 mm, often caducous.
Inflorescence terminal; peduncles 311, fasciculate, umbellate,
or shortly racemiform, 411 mm, puberulent to glabrescent,
subtended by 1 to several linear caducous bracts 12 mm; heads
1 per peduncle, subglobose to oblate, 110 mm in diam., 612flowered. Flowers fused at base or for up to half of hypanthium,
biology not noted. Calyx glabrous; limb 0.20.8 mm, truncate
to denticulate. Corolla white, campanulate, outside glabrous to
puberulent; tube 1.22 mm, inside densely villous from middle
to throat; lobes 4 or 5, narrowly oblong to ligulate, 2.23 mm,
apically thickened and hooked. Drupecetum subglobose or
compressed globose, 712 mm in diam., glabrescent. Drupes
mostly to fully fused, red, subglobose, 45 mm. Fl. JunJul, fr.
OctNov.
Forests on mountains, sparse or dense thickets at streamsides and
roadsides; 3001200 m. Anhui, Fujian, Guangdong, Guangxi, Hainan,
Hunan, Jiangsu, Jiangxi, Taiwan, Zhejiang [India, Japan, Sri Lanka, S
Thailand].
This is the most commonly collected species of Morinda in China.
This species is here circumscribed broadly and a bit differently from
FRPS (71(2): 190191. 1999); in particular, the species is restricted to
glabrous plants there but pubescent plants are included here. There
seems to be no other difference among these two sets of plants, and on
specimens there is continuous variation in pubescence that cannot be
clearly partitioned into separate states.

1a. Leaf blade drying papery or leathery,

ovate, obovate-lanceolate, or
obovate-oblong; petioles puberulent
to sparsely hirsute ............................... 25a. subsp. obovata

229

1b. Leaf blade rigid-membranous,

lanceolate to linear-lanceolate;
petioles glabrous .............................. 25b. subsp. umbellata
25a. Morinda umbellata subsp. obovata Y. Z. Ruan, Fl. Reipubl. Popularis Sin. 71(2): 325. 1999.
yang jiao teng
Petiole puberulent to sparsely hirsute; leaf blade drying papery or leathery, ovate, obovate-lanceolate, or obovate-oblong.
Fl. JunJul, fr. OctNov.
Forests on mountains, sparse or dense thickets at streamsides
and roadsides; 3001200 m. Anhui, Fujian, Guangdong, Guangxi, Hainan, Hunan, Jiangsu, Jiangxi, Taiwan, Zhejiang.

25b. Morinda umbellata subsp. umbellata

() yin du yang jiao teng (yuan ya zhong)
Petiole glabrous; leaf blade drying rigid-membranous, lanceolate to linear-lanceolate. Fl. JunJul, fr. OctNov.
Forests on mountains, sparse or dense thickets at streamsides and
roadsides; 3001200 m. Anhui, Fujian, Guangdong, Guangxi, Hainan,
Hunan, Jiangsu, Jiangxi, Taiwan, Zhejiang [India, Japan, Sri Lanka].

26. Morinda undulata Y. Z. Ruan, Fl. Reipubl. Popularis Sin.

71(2): 321. 1999.
bo ye mu ba ji
Small trees, to 20 m tall; branches glabrous, subquadrangular, smooth, reddish brown. Leaves opposite or solitary opposite an inflorescence; petiole 12 cm, glabrous; blade drying
yellowish brown, linear-oblong, linear-oblanceolate, or slightly
lyrate, 1820 45.5 cm, glabrous and shiny on both surfaces,
base cuneate, margin irregularly undulate to lyrate, apex acute
or acuminate; secondary veins 1224 pairs; stipules interpetiolar, triangular to broadly triangular, acute to obtuse. Inflorescence solitary and leaf-opposed; peduncle 1, 16 cm, glabrous;
heads 1 per peduncle, ellipsoid-oblong to cylindrical, many
flowered; bracts absent or subulate. Flowers with hypanthia
fully fused, biology not noted. Calyx limb truncate. Corolla
white, salverform; tube 1518 mm, densely villous in upper
part and throat; lobes 4 or 5, triangular to ligulate, 34 mm,
acute. Drupecetum mulberry-shaped to cylindrical or oblong,
11.5 0.60.8 cm. Fl. autumn, fr. winter.
About 900 m. S Yunnan (also cultivated in villages).
The protologue described the corolla lobes as 4, but the protologue figure shows 5 lobes.

27. Morinda villosa J. D. Hooker, Fl. Brit. India 3: 158. 1880.

xu mi ba ji
Woody lianas; branches surrounded at base by persistent
leafless stipules, densely ferruginous- or yellow villous, weakly
quadrangular. Leaves opposite; petiole 0.51.5 cm, shorter at
upper nodes, densely ferruginous- or yellow hirtellous; blade
drying papery, darkened, narrowly elliptic-oblong, obovate-lanceolate, or lanceolate, 812 26 cm, adaxially sparsely stri-

RUBIACEAE

230

gose or strigillose with pubescence denser along veins, abaxially densely ferruginous- or yellow hirtellous, base rounded,
slightly oblique, or sometimes cordulate, apex caudate-acuminate or shortly acuminate; secondary veins 1013 pairs, with pilosulous domatia; stipules interpetiolar or partially fused into a
tube, ovate to elliptic, 712 mm, pilosulous or hirtellous, acute,
obtuse, or truncate. Inflorescences terminal; peduncles 210,
fasciculate or umbellate, 0.92.2 cm, villosulous, as a group
subtended by 2 to several bracts 46 mm with 2 to several
subulate lobes; heads 1 per peduncle, subglobose, 56 mm in

diam., 410-flowered; bracteoles linear, 0.20.5 mm. Flowers

with hypanthia partially fused, biology not noted. Calyx villous,
strigillose, or hirtellous; limb ca. 1 mm, with 4 or 5 rounded to
obtuse teeth. Corolla greenish white, salverform, outside puberulent; tube 1012 mm, densely barbate in throat; lobes 4 or
5, narrowly oblong, 34 mm, apically rostrate. Drupecetum
compressed globose to subglobose, 11.5 cm in diam. Drupes
fused at base, orange, obovoid, ca. 5 mm. Fl. May, fr. JulSep.
Forests or thickets in valleys, at watersides, or at roadsides; 800
900 m. S Yunnan [N India, Thailand, Vietnam].

55. MOURETIA Pitard in Lecomte, Fl. Indo-Chine 3: 71. 1922.

du li cao shu
Chen Tao (); Charlotte M. Taylor
Perennial herbs or subshrubs, unarmed; stems sometimes becoming corky. Raphides present. Leaves opposite, sometimes anisophyllous, apparently without domatia; stipules persistent, interpetiolar, triangular to leaflike, often reflexed, acute to bifid. Inflorescences terminal, pseudoaxillary [or sometimes axillary, Mouretia vietnamensis Tange], capitate, subcapitate, or congested-cymose,
several to many flowered, subsessile to pedunculate, bracteate. Flowers sessile to pedicellate, bisexual, distylous, sometimes fused by
their ovaries. Calyx limb 5-lobed. Corolla white, yellow, or pink, tubular-funnelform to salverform, inside densely pubescent at
middle and sometimes through throat; lobes 5, valvate in bud. Stamens 5, inserted near middle of corolla tube, included with anthers
positioned near middle of corolla tube in long-styled form, exserted in short-styled form; filaments short or reduced in long-styled
form, well developed in short-styled form; anthers dorsifixed near base. Ovary 2-celled, ovules numerous in each cell on axile placentas; disk puberulent; stigma 2-lobed, exserted in long-styled form, included in short-styled form. Fruit capsular, obconic to subglobose, dehiscent through apical portion or operculum inside persistent calyx limb, thinly to thickly papery; seeds numerous, small,
angled.
Five species: E and SE Asia; one species in China.
This genus was revised by Tange (Nordic J. Bot. 17: 123132. 1997), who recognized four species from Indochina but documented each with
few specimens. Mouretia was studied in China by H. S. Lo (Bull. Bot. Res., Harbin 6(4): 48. 1986; Bull. Bot. Res., Harbin 18(3): 282. 1998), who
apparently did not have communication with Tange: they independently described and named the same Chinese species, which are here synonymized
(Tanges publication has priority). The known species of Mouretia are distylous (Tange, loc. cit.; Puff et al., Rubiaceae of Thailand, 182. 2005). The
breeding biology of the Chinese plants has not been reported by Chinese authors and is not evident from the material available to us but is presumably
similar. Lo (in FRPS 71(1): 183. 1999) described the stipules as caducous, but they are persistent on the specimens seen and according to Tange.

1. Mouretia inaequalis (H. S. Lo) Tange, Nordic J. Bot. 17:

127. 1997.
guang dong du li cao
Mouretia tonkinensis Pitard var. inaequalis H. S. Lo, Bull.
Bot. Res., Harbin 6(4): 48. 1986; M. guangdongensis H. S. Lo.
Weak herbs, 3040 cm tall; stems angled to subterete,
densely puberulent or villosulous-tomententulose to glabrescent. Leaves subequal to distinctly unequal, varied along stems;
petiole 515 mm, densely villosulous; blade drying thinly papery and olive-green, elliptic-oblong, elliptic, elliptic-ovate, or
obovate, larger blades 49 24 cm, smallest blades 1.52
0.81 cm, adaxially glabrous, abaxially glabrescent on lamina
and villosulous to strigillose-villous along principal veins, base
cuneate, obtuse, or rounded, apex acute to obtuse and shortly
acuminate; secondary veins 58 pairs; stipules leaflike, shortly
stipitate with blade portion subovate to obovate or subreniform, (1)2.55 mm, glabrescent, obtuse to rounded. Inflorescences congested-cymose, terminal and/or pseudoaxillary, several flowered, densely puberulent-tomentulose to strigillose;

peduncle 25 mm; bracts lanceolate to narrowly triangular, 12

mm; pedicels to 1.5 mm. Flowers free, subsessile to pedicellate.
Calyx puberulent-hispidulous; hypanthium portion obovoid, 1
1.5 mm; limb lobed nearly to base; lobes lanceolate to narrowly
triangular, 1.72(2.5) mm, entire to ciliolate. Corolla yellow,
white, or pink, tubular-salverform, densely hispidulous-puberulent with stout trichomes outside; tube 22.5 mm, internally
with ring of trichomes near middle; lobes triangular, ca. 2 mm,
abaxially appendaged near apex. Capsules obconical-subglobose, ca. 3 mm, glabrescent to puberulent, with persistent calyx
limbs sometimes elongating to 3 mm; seeds 0.40.5 mm. Fl.
Aug, fr. May.
On rocks in dense forests. Fujian, Guangdong, Guangxi (Qinzhou)
[Vietnam].
The description above includes measurements from Tange based
on specimens from Vietnam, including near the Chinese border. This
species is illustrated by Tange (loc. cit.: 126, t. 2, f. D). Tange separated
Mouretia tonkinensis from the Chinese plants by its consistently isophyllous leaves, fully capitate inflorescences, calyx lobes with tips
quickly becoming scarious and whitened, smaller corollas (with tube ca.
2.5 and lobes 1.1 mm), and smaller fruit (ca. 1.5 mm).

RUBIACEAE

231

56. MUSSAENDA Linnaeus, Sp. Pl. 1: 177. 1753.

yu ye jin hua shu
Chen Tao (); Charlotte M. Taylor
Belilla Adanson.
Trees, shrubs, or clambering or twining lianas, rarely dioecious, unarmed. Raphides absent. Leaves opposite or occasionally in
whorls of 3, with or usually without domatia; stipules persistent or caducous, interpetiolar, entire or 2-lobed. Inflorescences terminal
and sometimes also in axils of uppermost leaves, cymose, paniculate, or thyrsiform, several to many flowered, sessile to pedunculate,
bracteate. Flowers sessile to pedicellate, bisexual and usually distylous or rarely unisexual. Calyx limb 5-lobed nearly to base, frequently some or all flowers of an inflorescence with 1(5) white to colored, petaloid, persistent or deciduous, membranous, stipitate
calycophyll(s) with 37 longitudinal veins. Corolla yellow, red, orange, white, or rarely blue (Mussaenda multinervis), salverform
with tube usually slender then abruptly inflated around anthers, or rarely constricted at throat (M. hirsuta), inside variously pubescent
but usually densely yellow clavate villous in throat; lobes 5, valvate-reduplicate in bud, often long acuminate. Stamens 5, inserted in
middle to upper part of corolla tube, included; filaments short or reduced; anthers basifixed. Ovary 2-celled, ovules numerous in each
cell, inserted on oblong, fleshy, peltate, axile placentas; stigmas 2-lobed, lobes linear, included or exserted. Fruit purple to black,
baccate or perhaps rarely capsular (M. decipiens), fleshy, globose to ellipsoid, often conspicuously lenticellate, with calyx limb persistent or caducous often leaving a conspicuous scar; seeds numerous, small, angled to flattened; testa foveolate-striate; endosperm
abundant, fleshy.
About 200 species: widespread in tropical Africa, Asia, Madagascar, and Pacific islands; 29 species (18 endemic, one introduced) in China.
The characteristic large, petaloid calycophylls of Mussaenda are frequently but mistakenly considered to be bracts. Mussaenda is frequently
confused with Schizomussaenda; Schizomussaenda can be recognized by its large shrub habit with relatively large leaves and calycophylls, its
capsular fruit, and its flower buds with the corollas clavate and rather flat-topped with the acuminate ends of the corolla lobes then bent upward to
form an erect appressed group of filaments on the top of the bud.
Several species of Mussaenda are frequently cultivated as ornamentals in tropical regions. Mussaenda philippica A. Richard is apparently the
most commonly cultivated species and has numerous cultivar forms; Puff et al. (Rubiaceae of Thailand, 215. 2005) presented photos of several of the
cultivated forms. This species is pilosulous to villous throughout, with white to pink flowers and calycophylls and with calyx lobes variously 1 to all
modified into calycophylls. Also frequently cultivated are M. frondosa of our flora and M. erythrophylla Schumacher & Thonning of Africa; the latter
has pilose to villous pubescence on all organs, including the corollas, and deep red inflorescence axes and flowers, including the calycophylls and the
outside of the corolla but excepting the corolla limb, which is creamy white.
In addition to the species treated here, Hooker and Arnott (Bot. Beechey Voy. 265. 1838) reported Mussaenda glabra Vahl from Loo Choo in
China. Mussaenda glabra was described from the Himalaya and has medium-sized leaves, a climbing habit, petioles 315 mm, a corymbiform
branched inflorescence, calyx lobes 14 mm and deciduous in fruit, corolla tubes 1318 mm, and corolla lobes 33.5 mm; this species is otherwise
known from 3001300 m in India and Bhutan and has not been confidently recorded from China. The name M. glabra has been frequently applied
in herb. to Chinese specimens of both M. erosa and M. frondosa. Also Henry 8279 from Hainan was treated by C. E. C. Fisher (Bull. Misc. Inform.
Kew 1928: 274. 1928) as M. parryorum C. E. C. Fischer, a species otherwise known only from Assam, NE India. This identification seems unlikely,
especially given that M. parryorum has not otherwise been reported or treated from China or Indochina, so the Henry specimen will need reexamination. Mussaenda parryorum is included for reference in the key below, based on its protologue, but the species is not otherwise treated here.
H. H. Hsue and H. Wu (in FRPS 71(1): 283306. 1999), Hutchinson (in Sargent, Pl. Wilson. 3: 395400. 1916), and some other authors have
distinguished Mussaenda species in large part based on leaf shape, ovary length, calyx lobe length, and corolla size, but the consistency and
distinctiveness of these characters within species of Mussaenda have been questioned by some other authors. In particular, corolla size is apparently
sometimes widely variable among living plants in a population (see comments by Wood, Fl. Bhutan 2(2): 781. 1999), and the flower buds of
Mussaenda apparently often open prematurely when collected and thus are shorter than flowers at anthesis (pers. obs.). The calyx lobes and the ovary
often elongate after fertilization of the ovary, and the calyx lobes then fall off; thus, fruiting plants are very difficult to identify. Also Hutchinson
misidentified several widely distributed sets of Chinese Mussaenda specimens, generating confusion in this group (e.g., he identified Henry 12157 as
M. pubescens in spite of its pedicellate flowers with corolla tubes ca. 32 mm, vs. sessile and 1120 mm in M. pubescens); and he described only the
typical leaf size and shape of several species although he saw material with more variability. The weakness in some cases of Hutchinsons characters
and species concepts is shown by his simultaneous description in his Mussaenda treatment of two new species that were separated primarily on
vegetative features and actually are both based on different specimens of a single distinctive species, Schizomussaenda dehiscens.
As noted below, Mussaenda multinervis and M. decipiens were both described as having capsular fruit and most likely belong in other genera,
but which other genus or genera is not yet clear so they are maintained here. The measurements below of inflorescence size do not include the corollas
and calycophylls.

1a. Individual flowers with all 5 calyx lobes enlarged into petaloid calycophylls.
2a. Petiole 0.30.8 cm, leaf blade 29 14 cm; calyx lobes linear ......................................................................... 25. M. pubescens
2b. Petiole 1.53.5 cm, leaf blade 620 3.513 cm; calyx lobes lanceolate to ovate, 22.5 mm wide ................ 27. M. shikokiana
1b. Individual flowers with only 1 or 2 calyx lobes enlarged into a petaloid calycophyll, or without calycophylls.
3a. Calyx lobes (i.e., not calycophylls) subleaflike, 1.55 mm wide, lanceolate, oblanceolate, oblong-lanceolate,
ligulate, or broadly triangular.

232

RUBIACEAE

4a. Corolla tube 2025 mm, sericeous with appressed indument, sericeous with apices of trichomes
spreading, or villous.
5a. Stems sericeous with appressed indument; stipules 58 45 mm; calyx lobes 411 mm ...................... 19. M. macrophylla
5b. Stems villous or hirsute with widely spreading trichomes; stipules 710 612 mm; calyx
lobes 718 mm ................................................................................................................................................... 29. M. treutleri
4b. Corolla tube 914 mm, sericeous.
6a. Stipules triangular-ovate to broadly ovate, acute to cuspidate; corolla tube 1011 mm; berries ellipsoid ...... 9. M. emeiensis
6b. Stipules triangular to ovate-lanceolate, acute; corolla tube 914 mm; berries ellipsoid to subglobose ...... 27. M. shikokiana
3b. Calyx lobes less than 1.5 mm wide, linear to narrowly triangular, not at all leaflike.
7a. Stipules 1320 mm, lobed for less than half their length; corolla yellow or pale blue.
8a. Corolla pale blue; leaves 8.512 cm wide; calyx lobes ca. 2 mm; fruit baccate .......................................... 22. M. multinervis
8b. Corolla yellow; leaves 2.57.5 cm wide; calyx lobes 14 mm; fruit capsular.
9a. Calyx lobes 34 mm; leaves 67.5 cm wide .................................................................................................. 5. M. decipiens
9b. Calyx lobes 12 mm; leaves 2.56 cm wide ............................................................................. Schizomussaenda dehiscens
7b. Stipules 28 mm, deeply 2-parted; corolla white, yellow, or orange.
10a. Leaves sessile, subsessile, or shortly petiolate with petioles to 3 mm, obtuse, rounded, truncate,
or cordulate and then sometimes decurrent at base, if petioles more than 2 mm then leaves
truncate or cordulate.
11a. Leaf blade obtuse to rounded then abruptly acuminate at apex; corolla tube outside densely villous
with pubescence drying reddened; stems densely villous ....................................................................... 26. M. sessilifolia
11b. Leaf blade tapered to acuminate apex; corolla tube outside with sparse to dense villosulous,
pilosulous, tomentulose, strigose, or strigillose pubescence drying clear, whitened, or
reddened; stems strigillose, villous, villosulous, or tomentulose.
12a. Corolla tube 2026 mm; inflorescences shortly branched, flowers with pedicels ca. 1 mm ..................... 14. M. hossei
12b. Corolla tube 1120 mm; inflorescences subcapitate to congested, flowers mostly or all
sessile to subsessile.
13a. Calyx lobes ca. 2 mm ............................................................................................................................. 2. M. breviloba
13b. Calyx lobes 36 mm .......................................................................................................................... 25. M. pubescens
10b. Leaves distinctly petiolate with petioles 215 mm, truncate, obtuse, acute, or attenuate at base, if
petioles 3 mm or shorter then leaves acute.
14a. Leaves glabrous throughout or sparsely strigose on principal veins, drying thickly papery and
usually brown, with tertiary venation sparsely visible abaxially, widely spaced and subparallel,
not areolate; stipules caducous exposing a persistent, reddish brown fringe of trichomes ............................. 10. M. erosa
14b. Leaves glabrous or pubescent at least sparsely on veins abaxially, drying thinly to thickly
papery or membranous and green to brown, with tertiary venation regularly visible abaxially,
subparallel or areolate; stipules persistent or deciduous, without persistent fringe of trichomes
or this present and clear or whitened.
15a. Corolla tube constricted at throat ............................................................................................................ 13. M. hirsutula
15b. Corolla tube uniformly cylindrical or inflated just below or at throat.
16a. Calyx lobes 2530 mm, as long as or longer than corolla tube ................................................... 15. M. kwangsiensis
16b. Calyx lobes 115 mm, shorter than corolla tube.
17a. Corolla tube 58 mm; fruit stipitate or pedicellate, pedicels to 12 mm ........................................ 23. M. parviflora
17b. Corolla tube 540 mm; fruit sessile or subsessile to pedicellate, pedicels to 8 mm.
18a. Flowers sessile in 15 capitate groups; bracts and calyx lobes linear, 815 mm, and so
densely hirsute as to obscure inflorescence morphology .............................................................. 6. M. densiflora
18b. Flowers sessile to pedicellate in cymes with axes at least shortly developed; bracts and
calyx lobes narrowly triangular to lanceolate, 115 mm, glabrous to variously pubescent,
inflorescence morphology obscured or evident.
19a. Longest calyx lobes 615 mm on flowers at anthesis, longer than hypanthium portion.
20a. Corolla lobes 68 mm.
21a. Cultivated plants; leaves strigose, strigillose, or glabrescent abaxially .............................. 11. M. frondosa
21b. Native plants; leaves hirsute, subappressed villous, pilose, or strigose abaxially.
22a. Calyx densely villous, with pubescence spreading ....................................................... 3. M. caudatiloba
22b. Calyx hirsute ..................................................................................................................... 6. M. densiflora
20b. Corolla lobes 2.56 mm, at least some shorter than 6 mm.
23a. Corolla tubes 2630 mm.
24a. Inflorescences densely congested; flowers sessile or subsessile .................................... 13. M. hirsutula
24b. Inflorescences congested to laxly cymose; flowers with pedicels 15 mm .................... 17. M. laxiflora
23b. Corolla tubes 1125 mm.

RUBIACEAE

233

25a. Fruit ellipsoid to ellipsoid-oblong, 1820 1112 mm, densely lenticellate,

somewhat woody; stems and leaves abaxially densely villosulous to
tomentulose; Hainan .................................................................................................... 12. M. hainanensis
25b. Fruit subglobose to ellipsoid, 510 410 mm, smooth, fleshy or stiffly papery;
stems and leaves abaxially glabrescent to variously pubescent; widespread.
26a. Corolla with tube ca. 14 mm, lobes ca. 2.5 mm; Yunnan, elevation not noted ... 24. M. pingbianensis
26b. Corolla with tube 1123 mm, lobes 2.55 mm; widespread, below
1001600 m.
27a. Leaves 211.5 14.5 cm; flowers sessile to pedicellate, white to yellow;
below 100900 m, throughout China ..................................................................... 25. M. pubescens
27b. Leaves 620 310 cm; flowers sessile, mixed sessile and pedicellate, or all
pedicellate with pedicels to 3 mm, yellow-orange; S China, 3001400 m.
28a. Corolla lobes rounded ....................................................................................... 28. M. simpliciloba
28b. Corolla lobes acute or acuminate.
29a. Climbers; leaves 813 36 cm ......................................................................... 6. M. densiflora
29b. Erect shrubs; leaves 620 310 cm ..................................................................... 8. M. elliptica
19b. Longest calyx lobes 15.9 mm, shorter than to longer than hypanthium portion.
30a. Corolla tube ca. 40 mm ............................................................................................... 16. M. kwangtungensis
30b. Corolla tube 537 mm.
31a. Stems densely tomentose.
32a. Erect shrubs; calyx lobes 34 mm; corolla lobes acuminate ....................................... 21. M. mollissima
32b. Climbers; calyx lobes 57 mm; corolla lobes rounded .............................................. 28. M. simpliciloba
31b. Stems appressed pubescent to glabrescent.
33a. Most or all calyx lobes 12 mm.
34a. Leaves with 46 pairs of lateral veins; Hainan.
35a. Corolla tube 2531 mm; calycophylls elliptic to ovate ............................................... 1. M. antiloga
35b. Corolla tube 1315 mm; calycophylls oblong-elliptic or oblanceolate ............... 18. M. lotungensis
34b. Leaves with 68 pairs of lateral veins; mainland.
36a. Corolla tube ca. 12.8 mm ............................................................................................ 2. M. breviloba
36b. Corolla tube 1821 mm.
37a. Leaves 5.57.5 34.7 cm; petioles 0.50.6 cm ........................................................ 4. M. chingii
37b. Leaves 1017 2.56 cm; petioles 0.51.6 cm ................................. Schizomussaenda dehiscens
33b. Most or all calyx lobes 2.35.9 mm.
38a. Corolla with tube ca. 32 mm and lobes ca. 4 mm; Hainan ................................. 20. M. membranifolia
38b. Corolla with tube 1125 mm and lobes 26 mm; widespread, including Hainan.
39a. Corolla lobes ca. 2 mm.
40a. Corolla tube ca. 22 mm; Assam (not known from China,
included here for reference) ................................................... M. parryorum (see comment above)
40b. Corolla tube 1315 mm; Hainan ....................................................................... 18. M. lotungensis
39b. Corolla lobes 3.56 mm; widespread.
41a. Inflorescences subcapitate, mostly unbranched; flowers 320, sessile to
subsessile (though fruit may be stipitate).
42a. Corolla tube 2225 mm; stems generally without axillary short shoots ............ 6. M. densiflora
42b. Corolla tube 1120 mm; stems often with axillary short shoots with
small leaves ....................................................................................................... 25. M. pubescens
41b. Inflorescences branched, at least to one order, i.e., tripartite; flowers 5 to
numerous, sessile to long pedicellate.
43a. Stems strigose; near sea level to 1400 m ........................................................... 7. M. divaricata
43b. Plants tomentose to glabrescent; 12001400 m ............................................ 28. M. simpliciloba
1. Mussaenda antiloga Y. H. Chun & W. C. Ko, Fl. Hainan. 3:
582. 1974.
zhuang li yu ye jin hua
Climbing shrubs; branches terete, sparsely strigillose to
glabrescent. Leaves opposite; petiole 310 mm, densely strigillose to glabrescent; blade drying membranous, elliptic-oblong, elliptic, or oblanceolate, 711(18) 25.5 cm, adaxially
sparsely strigillose to glabrescent, abaxially moderately to

sparsely strigillose with pubescence denser on veins, base acute

to obtuse, apex acuminate; secondary veins 46 pairs, tertiary
venation visible and reticulate; stipules deciduous, triangular to
ovate, 36 mm, densely strigillose, deeply 2-lobed, lobes narrowly triangular to linear. Inflorescence laxly cymose, usually
regularly dichotomous, ca. 5 45 cm, strigillose to glabrescent, pedunculate; peduncles 3.54 cm; bracts subulate, 13
mm; pedicels 15 mm. Flowers pedicellate (or on higher order
inflorescence axes). Calyx with hypanthium portion ellipsoid to

RUBIACEAE

234

turbinate, 23 mm, glabrous; lobes narrowly triangular to linear,

12 mm, strigillose, 1 lobe on 1 flower of most inflorescences
enlarged into calycophyll, blade elliptic or ovate, 35 23 cm,
sparsely puberulent to strigillose especially on veins, base obtuse, stipe 1524 mm, apex acute to obtuse. Corolla yellow,
outside sparsely to moderately strigillose; tube 2231 mm, inside with clavate hairs at inflated throat; lobes broadly ovate, 3
4 mm, inside densely yellow papillose. Berry subglobose or
ellipsoid, 810 57 mm, glabrous, with calyx limb caducous.
Fl. MayDec.

pedunculate; peduncle 0.51 cm; bracts linear, ca. 8 mm. Flowers sessile. Calyx densely spreading villous; hypanthium portion turbinate, ca. 2.5 mm; lobes linear, 1013 mm, acute, 1
lobe on 1 or 2 flowers per inflorescence expanded into white
calycophyll, blade ovate, 56.7 35.3 cm, appressed villous,
base rounded, stipe 1825 mm, apex acute. Corolla outside
spreading grayish villous; tube ca. 25 mm, inside densely
clavate puberulent in upper part; lobes triangular-lanceolate, ca.
8 mm, caudate. Berry unknown. Fl. May.
Thickets. Guangxi.

Wet sites in dense forests; ca. 900 m. S Hainan.

The pubescence descriptions here are from the protologue.
H. H. Hsue and H. Wu (in FRPS 71(1): 296. 1999) gave the corolla tube length of this species as 2225 mm, but the corolla tube of the
type specimen is ca. 31 mm; this report is added here.

4. Mussaenda chingii C. Y. Wu ex H. H. Hsue & H. Wu, Acta

Phytotax. Sin. 24: 236. 1986.

2. Mussaenda breviloba S. Moore, J. Bot. 43: 137. 1905.

ren chang yu ye jin hua

duan lie yu ye jin hua

Shrubs, ca. 1.5 m tall; branches terete, appressed pubescent. Leaves opposite; petiole 56 mm, sparsely appressed pubescent; blade drying papery, adaxially shiny green, abaxially
glaucous, broadly ovate-elliptic or broadly ovate, 5.57.5 3
4.7 cm, both surfaces sparsely appressed pubescent with pubescence denser along veins, base rounded or obtuse, apex cuspidate, acute, or acuminate; secondary veins 6 or 7 pairs; stipules
deciduous, lanceolate or ovate, ca. 6 mm, densely pubescent,
deeply 2-lobed, lobes narrowly triangular. Inflorescences congested-cymose to somewhat lax, ca. 4 4.5 cm, densely pubescent, pedunculate or subsessile and tripartite; peduncle 11.5
cm; bracts linear, ca. 3.5 mm. Flowers shortly pedicellate. Calyx sparsely to densely pubescent; hypanthium portion cylindrical to ellipsoid, ca. 2.5 mm; lobes lanceolate-triangular, 1.5
2 mm, acuminate, 1 lobe on 13 flowers per inflorescence
expanded into calycophyll, blade elliptic to obovate, ca. 2.4 cm
9 mm, pubescent along veins on both surfaces, base cuneate,
stipe ca. 10 mm, apex acute. Corolla yellow to orange, outside
sparsely to densely appressed pubescent; tube ca. 21 mm, inside
densely yellow clavate puberulent in upper part; lobes orbicular-ovate, ca. 2.5 mm, inside yellow papillose, acute to apiculate. Berry unknown. Fl. May.

Shrubs to 1.5 m tall; branches terete, ferruginous tomentose. Leaves opposite; petiole 15 mm, tomentulose; blade
drying stiffly papery, long elliptic to broadly elliptic, 5.510
3.55 cm, adaxially sparsely strigillose with pubescence denser
along veins, abaxially pale tomentose along veins with indument grayish white and brownish, base obtuse, apex acuminate
or acute; secondary veins 7 or 8 pairs; stipules triangular-ovate,
45 mm, deeply 2-lobed, lobes triangular-subulate. Inflorescence cymose to corymbiform, densely flowered to rather lax,
ca. 4.5 78 cm, brown tomentose, pedunculate; bracts subulate, 35 mm, acute; pedicels 12 mm. Flowers sessile or pedicellate. Calyx tomentose; hypanthium portion ellipsoid to turbinate, 1.52.5 mm; lobes lanceolate or triangular-linear, ca. 2
mm, sometimes 1 lobe on 1 flower per inflorescence expanded
into calycophyll, blade elliptic, 34 12 cm, both surfaces
pubescent with pubescence denser along veins, base acute, stipe
ca. 6 mm, apex acute or obtuse. Corolla pale yellow, externally
densely tomentulose to strigose; tube ca. 12.8 mm, with clavate
hairs in throat; lobes triangular, ca. 2.2 mm, acuminate. Berry
unknown. Fl. Apr.
Roadsides in mountain forests; ca. 1300 m. Yunnan [Thailand].
This species was described from Thailand, and characters from
there are added to the description here. H. H. Hsue and H. Wu (in FRPS
71(1): 291. 1999) described the corolla tubes as 2023 mm, which
would make this species very similar to Mussaenda laxiflora, but the
protologue described them as ca. 12.8 mm.

3. Mussaenda caudatiloba D. Fang, Acta Phytotax. Sin. 40:

156. 2002.
wei lie yu ye jin hua
Shrubs, erect to climbing, size not reported; branches terete, retrorsely to spreading villous. Leaves opposite; petiole 5
10 mm, villous; blade drying thinly papery, dark green adaxially, pale abaxially, ovate to lanceolate, 411 25 cm, both
surfaces subappressed villous with pubescence denser on principal veins, base rounded to rarely cordate, apex acuminate to
subcaudate; secondary veins 69 pairs; stipules persistent,
deeply 2-lobed, villous, lobes linear-subulate, 710 mm. Inflorescences congested-cymose, several flowered, densely villous,

Thickets. Guangxi.

5. Mussaenda decipiens H. Li, Acta Phytotax. Sin. 18: 117.

1980.
mo tuo yu ye jin hua
Shrubs, 12 m tall; branches villous, terete, drying dark
green. Leaves opposite; petiole 15 cm, densely villous; blade
drying papery, elliptic, 1315 67.5 cm, adaxially subglabrous, abaxially hirsute with pubescence denser along veins,
base narrowly cuneate and decurrent on petiole, apex acuminate; secondary veins 79 pairs, tertiary venation indistinct;
stipules apparently persistent, linear-lanceolate to lanceolate,
1520 mm, densely villous, shortly 2-lobed. Inflorescence laxly
to rather congested-cymose, generally dichotomous, ca. 10 cm
wide, branched 3 or 4 times, villous becoming glabrescent,
pedunculate. Flowers subsessile, biology not noted. Calyx with
hypanthium portion turbinate, ca. 5 mm, hirtellous; lobes linear,
34 mm, sparsely villous, with 1 lobe on ca. 5 flowers per inflorescence expanded into white calycophyll, blade elliptic, ca. 7.5

RUBIACEAE

4 cm, base obtuse to cuneate, apex shortly acuminate. Corolla

yellow, tubular-salverform, outside densely pale yellow strigose; tube ca. 18 mm; lobes ovate-lanceolate, ca. 6 3 mm,
abaxially carinate, adaxially densely orange-yellow papillose,
long caudate. Capsule or berry globose, ca. 6 mm in diam.,
smooth, glabrous, with calyx limb persistent. Fl. Aug.
Broad-leaved forests or thickets on mountain slopes; 3001700
m. Xizang (Mdog), Yunnan (Gongshan).
The protologue described the fruit of this species as capsular,
which would be anomalous in Mussaenda, although H. H. Hsue and H.
Wu (in FRPS 71(1): 293. 1999) described it as a berry.

6. Mussaenda densiflora H. L. Li, J. Arnold Arbor. 24: 455.

1943.
mi hua yu ye jin hua
Climbers, ca. 2 m tall; branches terete, densely yellowish
brown villous or hirsute. Leaves opposite; petiole 57 mm,
densely hirsute; blade drying membranous or thinly papery,
broadly oblong-lanceolate or narrowly oblong-elliptic to usually ovate, 813 36 cm, adaxially moderately to sparsely
strigose, hirsute, or hirsutulous, abaxially moderately to densely
hirsute, pilose, or strigose with pubescence denser on principal
veins, base obtuse to rounded and sometimes abruptly attenuate, apex acute or acuminate; secondary veins 810 pairs, tertiary venation visible and reticulate; stipules persistent, 810
mm, densely villous to hirsute, deeply 2-lobed, lobes lanceolate,
narrowly triangular, or linear. Inflorescences capitate or subcapitate with 25 subglobose heads, 26 26 cm, densely villous, subsessile to pedunculate; peduncle to 3.5 cm; bracts lanceolate, narrowly triangular, narrowly elliptic, or linear, 1015
mm. Flowers sessile, biology not noted. Calyx densely hirsute;
hypanthium portion urceolate to ellipsoid, 23 mm; lobes linear-lanceolate to usually linear, 815 0.22 mm, sometimes 1
lobe of 1 or 2 flowers per inflorescence expanded into white
calycophyll, blade ovate, 56 34 cm, moderately to densely
strigose to pilose on both surfaces, base obtuse to cordulate,
stipe 815 mm, apex acute to acuminate. Corolla yellow, outside moderately to densely hirsute; tube 2225 mm, sometimes
contracted around anthers; lobes narrowly lanceolate or orbicular-lanceolate, 68 mm, inside densely yellow papillose, long
acuminate. Berry ovoid, 89 67 mm, glabrous, with calyx
limb deciduous. Fl. May.
Sparse thickets; 300800 m. Guangxi [Vietnam (Tsang 29049,
P!)].
This species is said in the protologue to be similar to Mussaenda
subsessilis Pierre, and it may be that with more specimens the distinctions between them (smaller calycophylls and longer narrower corolla
lobes in M. densiflora) may turn out to be endpoints of continuous variation. H. H. Hsue and H. Wu (in FRPS 71(1): 300. 1999) described the
stipules as caducous, but this has not been seen on specimens studied.
The protologue described the corolla tube as 33.5 cm, but this does not
match specimens seen, including the type; this may have been the
length of the entire corolla that was labeled incorrectly in the final
article.

7. Mussaenda divaricata Hutchinson in Sargent, Pl. Wilson. 3:

394. 1916.
zhan zhi yu ye jin hua

235

Climbing or suberect shrubs; branchlets terete, sparsely or

moderately strigose to densely mixed strigose, -strigillose, and
-hirtellous, sometimes becoming glabrescent, sometimes with
congested groups of leaves borne on reduced axillary buds.
Leaves opposite; petiole 0.51 cm, densely strigose; blade
drying thinly papery, adaxially pale to bright green or brown,
abaxially pale gray to bright green, elliptic, lanceolate, ellipticoblong, or ovate-elliptic, 3.512 1.57 cm, adaxially sparsely
strigillose on lamina and densely strigillose on principal veins,
abaxially strigillose to glabrous on lamina and densely strigillose along veins, base obtuse to acute, apex acute to abruptly
acuminate; secondary veins 511 pairs, tertiary venation reticulate to subparallel; stipules usually deciduous, ovate to triangular, 37 mm, moderately to densely strigose, deeply 2-lobed,
lobes narrowly triangular to subulate. Inflorescence congested
to somewhat laxly cymose, 12 34 cm, densely strigose, tripartite and sessile or pedunculate; peduncle 0.33.5 cm; bracts
narrowly triangular to linear, 17 cm, strigose to glabrescent;
pedicels to 1.5 mm. Flowers subsessile to pedicellate, biology
not noted. Calyx with hypanthium portion ellipsoid to turbinate,
1.52 mm, sparsely strigose; lobes subulate, 2.25 mm, densely
strigillose to strigose, with 1 lobe on 13 flowers per inflorescence expanded into white calycophyll, blade broadly elliptic or
ovate, 3.56 35 cm, sparsely strigillose on lamina and
densely strigillose along veins on both surfaces, base cuneate to
truncate, stipe 1025 mm, apex obtuse to shortly acuminate.
Corolla yellow, salverform, densely strigillose outside; tube 18
25 mm, densely yellow clavate pubescent inside upper part;
lobes ovate, 3.54 mm, adaxially densely yellow papillose,
acute to shortly acuminate. Berry ellipsoid, 11.2 0.40.6 cm,
sparsely strigillose, on pedicels or stipes to 6 mm. Fl. MaySep,
fr. Oct.
Thickets at riversides, in fields, or in valleys; near sea level to
1400 m. Guangdong, Guangxi, Guizhou, Hubei, Sichuan, Yunnan [Vietnam].
The illustration of this species presented by H. H. Hsue and H. Wu
(in FRPS 71(1): 304, t. 80, f. 13. 1999) is difficult to separate from the
plants included here in Mussaenda frondosa.

1a. Branches and abaxial surface of leaf

blades sparsely pubescent ..................... 7a. var. divaricata
1b. Branches and abaxial surface of leaf
blades densely pubescent ............................. 7b. var. mollis
7a. Mussaenda divaricata var. divaricata
() zhan zhi yu ye jin hua (yuan bian
zhong)
Branches sparsely pubescent. Leaf blade sparsely pubescent abaxially. Fl. JunSep.
Thickets at riversides or in fields; near sea level to 1200 m.
Guangdong, Guangxi, Guizhou, Hubei, Sichuan, Yunnan.

7b. Mussaenda divaricata var. mollis Hutchinson in Sargent,

Pl. Wilson. 3: 398. 1916.
rou mao yu ye jin hua
Branches densely pubescent. Leaf blade densely pubescent
abaxially. Fl. May, fr. Oct.

RUBIACEAE

236

Thickets in valleys; ca. 1400 m. Yunnan [Vietnam].

Mixed forests on slopes; 700900 m. Sichuan (Emei Shan).

8. Mussaenda elliptica Hutchinson in Sargent, Pl. Wilson. 3:

395. 1916.

10. Mussaenda erosa Champion ex Bentham, Hookers J. Bot.

Kew Gard. Misc. 4: 193. 1852.

tuo yuan yu ye jin hua

nan teng

Shrubs, 12 m tall; branchlets terete, sparsely strigillose.

Leaves opposite; petiole 0.71 cm, strigillose to strigose; blade
drying thinly papery, elliptic, 620 310 cm, both surfaces
glabrous to strigillose at least on veins, base rounded to cuneate,
apex acuminate; secondary veins 6 or 7 pairs, tertiary venation
apparently reticulate; stipules deciduous, ovate-triangular, ca. 8
mm, sparsely puberulent, deeply 2-lobed, lobes narrowly triangular, acuminate. Inflorescence congested-cymose, ca. 1.5
1.5 cm, strigillose or strigose, subsessile to pedunculate; bracts
linear-subulate, to 8 mm. Flowers pedicellate, biology not
noted. Calyx strigillose; hypanthium portion turbinate, ca. 2
mm; lobes subulate, 67 mm, with 2 lobes on 25 flowers per
inflorescence expanded into white calycophyll, blade broadly
ovate, 57.5 3.55 cm, strigillose along veins, base obtuse to
truncate, stipe 715 mm, apex acute to shortly acuminate. Corolla yellow, salverform, outside strigose; tube ca. 20 mm,
densely yellow clavate pubescent inside upper part; lobes triangular-ovate, ca. 3 mm, mucronate with appendage ca. 1 mm.
Berry unknown. Fl. MayJun.

Climbing shrubs, to 5 m tall; branches glabrous, terete,

sparsely lenticellate. Leaves opposite; petiole 0.31.5 cm, glabrous; blade drying thickly papery, brown, ovate, oblanceolate,
lanceolate, or oblong-elliptic, 5.514 2.46 cm, both surfaces
glabrous or sparsely strigose on principal veins, base acute, cuneate, or occasionally obtuse, apex acute to acuminate; secondary veins 46 pairs, without domatia, tertiary venation only
sparsely visible abaxially; stipules caducous exposing a fringe
of persistent reddish brown trichomes, narrowly triangular, 38
mm, glabrous, deeply 2-parted, lobes acute to acuminate. Inflorescence compound-cymose to compound-corymbiform, 28
212 cm, congested when young becoming lax with age, glabrous, pedunculate or sessile and 3- or 5-partite; peduncles 1
2.5 cm; bracts triangular to linear-lanceolate, 1.57 mm,
sparsely strigillose to glabrous; pedicels 15 mm. Flowers pedicellate, biology not noted. Calyx with hypanthium portion ellipsoid, 2.53.5 mm, glabrous; lobes linear-lanceolate to narrowly
triangular, 24.5 mm, often unequal on an individual flower,
strigillose or hispidulous, with 1 lobe on 16 flowers per inflorescence expanded into white calycophyll, blade broadly elliptic to ovate, 3.56.5 35 cm, glabrous except sparsely strigillose on principal veins, base cuneate, rounded, or truncate, stipe
920 mm, apex rounded to acute. Corolla yellow to orangeyellow, outside densely strigillose to strigose; tube 2224 mm;
lobes ovate, 45 mm, adaxially yellow papillose, rounded then
abruptly shortly acuminate. Berry ellipsoid to subglobose, 10
13 810 mm, glabrous, smooth or sparsely lenticellate, calyx
limb deciduous. Fl. AprJul, fr. SepDec.

Forests in valleys, forest margins; 6001000 m. Guangxi, Sichuan, Yunnan.

It is implied in the protologue that the stipules are entire, but on an
isotype specimen (E. H. Wilson 4604, US bar code 00137848, Web!)
they are deeply bilobed; also the protologue described the inflorescences as pedunculate, but they are subsessile on this isotype. In FRPS
(71(1): 302. 1999), the leaves were described as sparsely pubescent and
the calyx lobes as 4.55.5 mm, but in the protologue they were described as glabrous and 67 mm, respectively.

9. Mussaenda emeiensis Z. Y. Zhu & S. J. Zhu, Bull. Bot. Res.,

Harbin 28: 257. 2008.
e mei yu ye jin hua
Weak shrubs, to 5 m tall; branches compressed, shortly pubescent. Leaves opposite; petiole 23.5 cm, shortly pubescent;
blade drying thickly papery, green adaxially, pale abaxially, 7
18(20) 4.512(15) cm, both surfaces pilose on veins to
glabrescent, base obtuse to acute, apex acute to acuminate; secondary veins 79 pairs; stipules triangular-ovate to broadly
ovate, 67 mm, shortly pubescent, acute to cuspidate or 2lobed. Inflorescence cymose or fasciculate, ca. 1 1.5 cm,
many flowered, densely pubescent, sessile; bracts ovate-lanceolate, 410 mm, acuminate; pedicels ca. 2 mm. Flowers pedicellate, biology not noted. Calyx obovoid, ca. 4 mm, pilosulous;
lobes oblong-lanceolate to lanceolate, 810 22.5 mm, pilosulous, acute, with 1 lobe in ca. 2 flowers per inflorescence
expanded into calycophyll, blade ovate to elliptic, 2.55.5 cm,
pubescent to glabrous, base obtuse, stipe 810 mm, apex obtuse
to acute. Corolla yellow or orange, tubular, outside densely pubescent; tube 1011 mm, inside densely clavate pilose; lobes
broadly ovate, 34 mm, inside verrucose, acute to acuminate.
Berry ellipsoid, ca. 10 8 mm, pilosulous to glabrescent, calyx
limb deciduous.

Sparse evergreen forests, streamsides, along roads; 300800 m.

Fujian, Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan, Yunnan [Japan (Ryukyu Islands), Vietnam].

11. Mussaenda frondosa Linnaeus, Sp. Pl. 1: 177. 1753.

yang yu ye jin hua
Climbing shrubs; branches terete to compressed, moderately to densely pale red sericeous to hirsute becoming glabrescent, red-brown or brown, rather densely lenticellate. Leaves
opposite; petiole 410 mm, densely sericeous; blade drying
thinly papery or leathery, adaxially dark green to brownish
green, abaxially pale green to yellowed, broadly elliptic, elliptic-oblong, ovate, or oblanceolate, 815 38 cm, both surfaces sparsely strigillose on lamina and strigose to sericeous on
principal veins, base acute, obtuse, or rounded, apex acute to
caudate-acuminate; secondary veins 710 pairs, tertiary venation visible and reticulate; stipules persistent, triangular, 510
mm, densely strigose to pilose, deeply 2-lobed, lobes lanceolate
to narrowly triangular, acute to acuminate. Inflorescences congested-cymose becoming lax with lateral axes elongating markedly, 48 820 cm with lateral axes to 8 cm, sparsely to
moderately pubescent with 2 types of pubescence, mixed strigillose to appressed puberulent and hirsute to villous, pedunculate or sessile and 3-partite; peduncles 13 cm; bracts triangular

RUBIACEAE

or elliptic, 410 mm, usually 2- to multifid, acute to acuminate;

pedicels 16 mm. Flowers pedicellate, biology not noted. Calyx
with hypanthium portion ellipsoid, 34 mm, strigose to sericeous; lobes narrowly triangular to narrowly ligulate, 712 mm,
hirsute, acute to acuminate, with 1 lobe of 14 flowers per inflorescence sometimes expanded into calycophyll, blade elliptic-oblong to ovate, 68 2.55 cm, both surfaces glabrescent
on lamina and strigillose on principal veins, base acute to cuneate, stipe 1030 mm, apex acute or acuminate. Corolla salverform, outside hirsute; tube 2225 mm; lobes ovate, 67 mm,
acuminate. Berry ovoid or ellipsoid, ca. 10 7 mm, strigose to
glabrescent, calyx limb deciduous. Fl. AprMay.
Cultivated in Guangdong and Hainan [native to Cambodia, India,
Indonesia, Sri Lanka, and Vietnam].

12. Mussaenda hainanensis Merrill, Lingnan Sci. J. 14: 58.

1935.
hai nan yu ye jin hua
Climbing shrubs; branches terete, densely ferruginous- or
gray villosulous to tomentulose. Leaves opposite; petiole 2
5 mm, densely velutinous to hirtellous; blade drying papery,
adaxially dark green, abaxially pale, oblong-elliptic, obovate, or
lanceolate, 312 1.54 cm, adaxially sparsely to densely villosulous to strigillose or hispid, abaxially densely villosulous or
tomentulose, both surfaces with pubescence denser along principal veins, base cuneate to acute, apex acute to shortly acuminate; secondary veins 7 or 8 pairs, tertiary venation visible
and reticulate; stipules persistent, triangular to ovate, 37 mm,
densely pilosulous to villosulous, entire or usually deeply 2lobed, lobes lanceolate to linear, acuminate. Inflorescence subcapitate to congested-cymose, 35 35 cm, densely pilosulous to villosulous, pedunculate or sessile and 3-partite; peduncle 13 cm; bracts linear-lanceolate, 36 mm. Flowers sessile or
subsessile, biology not noted. Calyx densely strigose to sericeous or pilose; hypanthium portion ellipsoid to turbinate, 1.52
mm; lobes linear-lanceolate to narrowly elliptic, 68 mm, acute
to acuminate, with 1 lobe on 1 or 2 flowers per inflorescence
expanded into calycophyll, blade broadly elliptic to ovate, 1.5
4 13.5 cm, moderately to sparsely strigillose or sericeous,
base obtuse, truncate, or cordulate, stipe 1016 mm, apex acute.
Corolla yellow, salverform, densely strigose outside; tube 20
25 mm; lobes triangular-ovate, 35 mm, adaxially densely yellow papillose, acuminate. Berry ellipsoid to ellipsoid-oblong,
1820 1112 mm, pilosulous to hirtellous, densely lenticellate, somewhat woody, with calyx limb deciduous, with pedicels sometimes elongating, to 4 mm. Fl. MarJun, fr. JulAug.
Forests at middle elevations; 300800 m. Hainan.
The protologue and H. H. Hsue and H. Wu (in FRPS 71(1): 297.
1999) described the calyx tube as 34 mm, but this has not been seen
on specimens studied.

13. Mussaenda hirsutula Miquel, J. Bot. Nerl. 1: 109. 1861.

cu mao yu ye jin hua
Mussaenda inflata H. S. Hsue & H. Wu.
Climbing shrubs; branches terete, densely ferruginous- or
gray villosulous, hirtellous, or tomentulose. Leaves opposite;

237

petiole (2)315 mm, moderately to densely villosulous, tomentulose, or velutinous; blade drying papery to subleathery,
abaxially pale, oblong-elliptic, lanceolate, or ovate, (4)713
(2)2.54 cm, adaxially sparsely to moderately hirtellous or
hispid with pubescence usually denser along principal veins,
abaxially moderately to densely villosulous, hirtellous, or strigose-hirsute, base acute, cuneate, or rounded, apex acute or
acuminate; secondary veins 58 pairs, tertiary venation visible
and reticulate; stipules persistent (or in M. inflata form deciduous), triangular, (4)510 mm, densely strigose, villosulous, or
pilose, deeply 2-lobed, lobes lanceolate, narrowly triangular, or
linear. Inflorescence subcapitate to congested-cymose, (1)1.5
4 1.54 cm, densely villosulous to hirsute, pedunculate; peduncle 0.31.5 cm; bracts linear-lanceolate, 46 mm. Flowers
subsessile or sessile, biology not noted. Calyx sparsely to
densely villosulous or villous; hypanthium portion ellipsoid
to subglobose, 1.52 mm; lobes linear to narrowly triangular,
(4)710(13) mm, sometimes 1 lobe on 13 flowers per inflorescence expanded into calycophyll, blade broadly elliptic to
ovate, (2.5)46 (2)35 cm, sparsely strigillose to villosulous with pubescence usually denser on veins, base cuneate to
rounded, stipe 1014 mm, apex rounded or acute. Corolla yellow to orange-yellow, salverform, outside densely strigose to
sericeous; tube cylindrical and 2628 mm (or constricted in
throat and 1819 mm in M. inflata form); lobes elliptic to lanceolate, 46 mm, adaxially densely yellow papillose, acute to
acuminate. Berry ellipsoid or subglobose, 1420 912 mm,
lenticellate, with calyx limb persistent, with pedicels sometimes
elongating, to 4 mm. Fl. AprJun (Nov in M. inflata form), fr.
JulJan of following year.
Thickets in fields, at streamsides, or in valleys, often on tree
crowns; 300800 m. Guangdong, Guizhou, Hainan, Hunan, Yunnan.
The name Mussaenda inflata was based on a single specimen
distinguished primarily by a notably swollen corolla. Deng and Zhang
(Acta Phytotax. Sin. 44: 608609. 2006) studied this and concluded that
M. inflata is a synonym of M. hirsutula and was described based on a
specimen with malformed corollas. In the description above, in general,
the measurements in parentheses apply only to the plants previously included in M. inflata.

14. Mussaenda hossei Craib, Beih. Bot. Centralbl. 28(2): 444,

457. 1911.
hong mao yu ye jin hua
Mussaenda rehderiana Hutchinson.
Shrubs, to 2 m tall; branches densely whitened, reddish, or
brownish villosulous, tomentulose, or villous, terete, sometimes
becoming glabrescent. Leaves opposite; petiole 0.53 mm, villosulous to villous; blade drying thickly papery, oblanceolate,
oblong-oblanceolate, elliptic, lanceolate, or ovate, 314 1.54
cm, adaxially sparsely to densely pilosulous to villosulous,
abaxially densely hirtellous, villosulous, or villous, base obtuse,
subcordate, or rounded, apex acuminate or acute; secondary
veins 810 pairs, tertiary venation not readily visible; stipules
usually persistent, lanceolate to triangular, 57 23 mm,
densely strigose, villosulous, or villous, entire or usually 2-lobed
for up to 1/2, segments acute to acuminate. Inflorescence congested-cymose, densely tomentose to villous, pedunculate or

RUBIACEAE

238

sessile and 3(or 5)-partite; peduncle 15 cm; bracts lanceolate,

narrowly triangular, or linear, 110 mm; pedicels to 1 mm.
Flowers subsessile to shortly pedicellate, biology not noted. Calyx densely villosulous to hirtellous; hypanthium portion subellipsoid to oblanceoloid, 22.5 mm; lobes lanceolate to narrowly triangular, 22.2 mm, acute, with 2 lobes on 46 flowers
of each inflorescence expanded into white calycophyll, blade
elliptic to suborbicular, 25.5 24 cm, base cuneate to cordulate, stipe 1215 mm, apex obtuse or rounded and shortly
acuminate. Corolla orange-yellow, salverform, outside densely
pilosulous, villosulous, or tomentulose; tube 2026 mm; lobes
suborbicular, 2.53 mm, obtuse then abruptly acuminate or
acute. Berry oblong-ellipsoid, ca. 18 8 mm, calyx lobes persistent. Fl. NovMar.
Forests; 6001600 m. S Yunnan [Laos, Myanmar, Thailand, Vietnam].
In the protologue, Craib noted variation in density of the pubescence among the few specimens he studied; continuous variation in
density and length of the trichomes is evident among the specimens collected since then.

15. Mussaenda kwangsiensis H. L. Li, J. Arnold Arbor. 24:

455. 1943.
guang xi yu ye jin hua
Climbing shrubs; branches terete, densely strigillose, sometimes with congested groups of leaves borne on reduced axillary buds. Leaves opposite; petiole 58 mm, appressed pubescent; blade drying thinly papery, oblong-lanceolate or lanceolate, 811 2.54 cm, adaxially sparsely strigillose with pubescence denser along veins, abaxially densely villous, sparsely
villosulous, or hirtellous, base attenuate or cuneate, apex acuminate or shortly acuminate; secondary veins 68 pairs, with tertiary venation not readily visible; stipules caducous, deeply 2parted, lobes linear, ca. 6 mm. Inflorescence congested-cymose,
ca. 4 cm, moderately to densely strigillose; bracts linear, 515
mm. Flowers densely grouped, sessile, biology not noted. Calyx
with hypanthium portion ellipsoid, ca. 5 mm, densely pubescent; lobes linear, 2530 12 mm, sparsely pilose, with 1 lobe
on 1(or ?more) flower in each inflorescence enlarged into white
calycophyll, blade ovate, ca. 6 cm, sparsely pubescent on both
surfaces, base rounded, stipe ca. 8.5 mm, apex acuminate. Corolla yellow, slenderly salverform, outside gray pubescent; tube
2025 mm; lobes ovate, ca. 3 1.5 mm, adaxially densely yellow papillose, acuminate, acute, or mucronate. Berry not seen.
Fl. SepJan.
Sparse forests at streamsides in valleys. Guangxi.

ondary veins 36 pairs, apparently without domatia, tertiary venation not visible; stipules caducous, 1.53 mm, densely strigillose, deeply 2-lobed, lobes linear, 1.52 mm. Inflorescence
compact-cymose to subcapitate, ca. 1 11.5 cm, few to several flowered, densely strigillose, pedunculate; peduncles 210
mm; bracts linear, ca. 1 mm. Flowers subsessile, biology not
noted. Calyx strigillose to glabrescent; hypanthium portion ellipsoid, 23 mm; lobes linear, 2.53.5 mm, with 1 lobe on 24
flowers per inflorescence sometimes expanded into calycophyll,
blade oblong-ovate, elliptic, or elliptic-ovate, 3.55 1.52.5
cm, strigillose, base cuneate, stipe 1215 mm, apex acute to obtuse. Corolla yellow, salverform, outside strigose to strigillose;
tube (31)3942 mm; lobes ovate, 4.55 mm, adaxially densely
yellow papillose, acuminate. Berry not seen. Fl. MaySep.
Thickets on mountains. S Guangdong.

17. Mussaenda laxiflora Hutchinson in Sargent, Pl. Wilson. 3:

399. 1916.
shu hua yu ye jin hua
Shrubs, to 3 m tall; branches terete, densely strigillose,
strigose, or appressed tomentose. Leaves opposite; petiole 410
mm, densely strigillose or appressed tomentulose; blade drying
papery, elliptic, obovate, or obovate-oblanceolate, 6.514
2.55 cm, adaxially sparsely strigillose especially along veins,
abaxially densely strigillose to strigose along veins and glabrescent on lamina, base cuneate, apex caudate to acuminate; secondary veins 69 pairs, apparently without domatia, tertiary
venation visible and subparallel; stipules deciduous, 48 mm,
densely strigillose, deeply 2-lobed, lobes narrowly triangular,
acute. Inflorescences congested to laxly cymose, 25 37 cm,
densely strigose to strigillose, pedunculate or sessile and tripartite; peduncle 2025 mm; bracts linear, 110, usually caducous;
pedicels 15 mm. Flowers pedicellate in umbelliform groups of
2 or 3, biology not noted. Calyx moderately to densely strigose;
hypanthium portion narrowly ellipsoid to turbinate, ca. 3 mm;
lobes linear to narrowly triangular, 57 mm, with 1 lobe in 13
flowers per inflorescence sometimes expanded into calycophyll,
blade lanceolate or ovate, 3.57 1.52.5 cm, sparsely strigillose with pubescence usually denser on veins, base cuneate to
obtuse, stipe 1012 mm, apex acute to subacuminate. Corolla
yellow or orange-yellow, salverform, outside densely sericeous;
tube 2630 mm, inside densely yellow clavate pubescent in upper part; lobes triangular-lanceolate, 34 mm, adaxially densely
yellow papillose, acuminate to long acuminate. Berry ellipsoid,
ca. 1 cm, sparsely pubescent. Fl. JunJul, fr. AugDec.
Forests, thickets; ca. 1600 m. Yunnan.

16. Mussaenda kwangtungensis H. L. Li, J. Arnold Arbor. 25:

427. 1944.

18. Mussaenda lotungensis Y. H. Chun & W. C. Ko, Fl. Hainan. 3: 581. 1974.

guang dong yu ye jin hua

le dong yu ye jin hua

Climbing shrubs, 12.5 m tall; branches terete, brown,

densely strigillose becoming glabrescent. Leaves opposite; petiole 35 mm, strigillose; blade drying thinly papery, lanceolateelliptic to elliptic-oblong, 39 13 cm, adaxially sparsely
strigillose to glabrescent, abaxially sparsely to densely strigose
or strigillose with pubescence denser along principal veins, base
acute to obtuse, apex acute to acuminate with tips to 1 cm; sec-

Climbing shrubs; branches terete, sparsely strigillose to

glabrescent. Leaves opposite; petiole 412 mm, moderately to
densely strigillose; blade drying papery, lanceolate or oblonglanceolate, 510 1.53 cm, adaxially glabrous or sparsely
strigillose along principal veins, abaxially sparsely strigillose
to glabrous with pubescence denser along veins, base cuneate
or obtuse, apex acuminate; secondary veins 4 or 5 pairs, tertiary

RUBIACEAE

venation subparallel; stipules deciduous, triangular, 36 mm,

moderately strigillose, deeply 2-lobed, lobes narrowly triangular to subulate. Inflorescences terminal on main stems and
sometimes short lateral stems, congested to somewhat laxly
cymose, 13 1.53 cm, moderately to densely strigillose and
strigose, pedunculate; peduncles 1.52 cm; bracts triangular, 3
3.5 mm; pedicels 13 mm. Flowers pedicellate, biology not
noted. Calyx with hypanthium portion ellipsoid to subturbinate,
22.5 mm, sparsely strigillose; lobes narrowly triangular, 1.5
2.5 mm, densely strigillose, with 1 lobe on 13 flowers per
inflorescence expanded into calycophyll, blade narrowly oblong-elliptic to oblanceolate, 34 0.81.2 cm, both surfaces
sparsely to moderately strigillose, base acute, stipe 516 mm,
apex obtuse or acute. Corolla yellow, salverform, outside moderately to densely strigillose; tube 1315 mm, with yellow clavate pubescence in throat; lobes triangular to ovate, ca. 2 mm,
adaxially densely yellow papillose, acuminate. Berry unknown.
Fl. AprJun.
Wet soil in dense forests. Hainan (Ledong).

19. Mussaenda macrophylla Wallich in Roxburgh, Fl. Ind. 2:

228. 1824.
da ye yu ye jin hua
Mussaenda hispida D. Don; M. kotoensis Hayata.
Erect or climbing shrubs; branches terete to quadrangular,
sparsely to moderately sericeous. Leaves opposite; petiole 435
mm, sparsely hirsute to glabrescent; blade drying membranous
to papery, green to brownish, paler below when specimen well
preserved, elliptic-oblong, elliptic, or ovate, 1221 811 cm,
both surfaces sparsely strigillose to pilose on lamina and moderately hirsute along principal veins, base cuneate to obtuse,
apex acute to acuminate; secondary veins 68 pairs, without
domatia, tertiary venation reticulate; stipules deciduous, ovate
to triangular, 58 mm, sparsely brown hirsute to glabrescent,
deeply 2-lobed, lobes acute to acuminate. Inflorescences laxly
cymose, 615 cm, hirsute, sessile with arching lateral axes;
bracts lanceolate or 2- or 3-parted, 510 mm, obtuse to acuminate. Flowers subsessile, biology not noted. Calyx with hypanthium portion campanulate to obconic, 34 mm, densely
brown strigose to sericeous; lobes lanceolate, ligulate, or oblanceolate, 411 1.53 mm, often unequal on an individual
flower, densely to moderately strigillose to strigose, acute, with
1 lobe on 1 to several flowers in each inflorescence expanded
into white calycophyll, blade broadly ovate or rhombic, 512
cm, both surfaces sparsely hirsute to glabrescent on lamina and
moderately to densely pilosulous to puberulent on principal
veins, base obtuse to truncate, stipe 1837 mm, apex obtuse to
shortly acuminate. Corolla orange-yellow to golden yellow, salverform, outside densely sericeous; tube 2025 mm; lobes
ovate, 710 mm, acute to acuminate. Berry ellipsoid, 1015
mm, strigose, lenticellate, calyx limb deciduous. Fl. JunJul, fr.
AugNov.
Thickets or forests on mountains; sea level to 1300 m. Guangdong, Guangxi, Taiwan [Indonesia, Malaysia, Philippines].
This species is circumscribed somewhat differently here than by
H. H. Hsue and H. Wu (in FRPS 71(1): 287289. 1999); see comments
under Mussaenda treutleri.

239

20. Mussaenda membranifolia Merrill, Philipp. J. Sci. 23: 267.

1923.
mo ye yu ye jin hua
Climbing shrubs; branches terete, glabrous. Leaves opposite; petiole 330 mm, hispid to strigillose; blade drying membranous, oblanceolate, elliptic, or elliptic-oblong, 915 2.5
4.5 cm, both surfaces sparsely strigose with pubescence denser
along veins abaxially, base obtuse to acute, apex long acuminate; secondary veins 69 pairs, tertiary venation visible, reticulate; stipules generally deciduous, triangular, 35 mm, strigose, deeply 2-lobed, lobes linear. Inflorescences laxly dichotomous, 1012 1013 cm, strigillose to glabrescent, pedunculate or apparently sessile and tripartite; peduncles 4.56 cm;
bracts linear, ca. 3 mm; pedicels 0.51 mm. Flowers pedicellate
(or borne on higher order inflorescence axes), biology not
noted. Calyx glabrous; hypanthium portion subellipsoid, 22.5
mm; lobes linear, 34 mm, with 1 lobe on 1 flower per inflorescence expanded into calycophyll, blade ovate to elliptic, 3
7.5 25 cm, strigose along veins on both surfaces, base cuneate to obtuse, stipe ca. 22 mm, apex acute to subacuminate.
Corolla white, slenderly salverform, outside sparsely strigose;
tube ca. 32 mm, densely yellow clavate pubescent in throat;
lobes lanceolate or ovate-lanceolate, ca. 4 mm, adaxially yellow
papillose, acute to acuminate. Berry subglobose or ellipsoid,
1013 57 mm, glabrous, calyx lobes caducous. Fl. AprJun,
fr. AugOct.
Humid sites in forests. Hainan.

21. Mussaenda mollissima C. Y. Wu ex H. H. Hsue & H. Wu,

Acta Phytotax. Sin. 24: 235. 1986.
duo mao yu ye jin hua
Shrubs, 17 m tall; branches terete, densely pale yellowto brown tomentose, -hirtellous, or -hirsute. Leaves opposite;
petiole 912 mm, sparsely to moderately strigose, hirtellous, or
hirsute; blade drying subleathery, elliptic, broadly elliptic, or
broadly ovate, 811 47.2 cm, both surfaces densely pale
yellow tomentose to -hirtellous, base cuneate or obtuse, apex
acute to shortly acuminate; secondary veins 9 or 10 pairs, apparently without domatia, tertiary venation reticulate to subparallel; stipules deciduous, ovate, ca. 6 mm, densely tomentose, deeply 2-lobed, lobes linear, ca. 3.5 mm. Inflorescence
congested-cymose to subcapitate, ca. 4 4 cm, densely tomentose to hirtellous; peduncles 34 cm; bracts lanceolate, ca. 6
mm. Flowers subsessile, biology not noted. Calyx densely tomentose to hirtellous; hypanthium cylindrical to ellipsoid, 2.5
3.5 mm; lobes narrowly lanceolate to narrowly triangular, 34
mm, with 1 lobe in 14 flowers per inflorescence sometimes
expanded into calycophyll, blade elliptic to obovate, ca. 5.5
2.8 cm, densely tomentose, base acute, stipe ca. 14 mm, apex
obtuse to acute. Corolla orange, salverform, outside densely
yellowish brown villous; tube ca. 22.5 mm, inside densely yellow clavate pubescent in upper part; lobes ovate, ca. 4 mm,
adaxially densely yellow papillose, acuminate. Berry ellipsoid,
ca. 12 9 mm, sparsely tomentose to hirtellous. Fl. May, fr.
Jun.
Forest margins, roadsides. S Yunnan.

RUBIACEAE

240

22. Mussaenda multinervis C. Y. Wu ex H. H. Hsue & H. Wu,

Acta Phytotax. Sin. 24: 237. 1986.
duo mai yu ye jin hua
Shrubs, 23 m tall; branches terete to angled, densely
white pubescent to glabrescent. Leaves opposite; petiole ca. 3
cm, sparsely pubescent; blade drying membranous, adaxially
drying pale green, abaxially gray, broadly elliptic or broadly
ovate, 1622 8.512 cm, adaxially sparsely appressed pubescent with pubescence denser on principal veins, abaxially glabrescent except densely appressed pubescent on principal veins,
base acute, apex acute; secondary veins 11 or 12 pairs; stipules
ovate-lanceolate, ca. 13 mm, acuminate and shortly 2-lobed. Inflorescence congested-cymose, ca. 4 8 cm, densely pale gray
pilose, pedunculate or sessile and tripartite; bracts lanceolate,
ca. 9 mm; pedicels ca. 1 mm. Flowers pedicellate, biology not
noted. Calyx sparsely pubescent; hypanthium portion urceolate
to ellipsoid, ca. 2.5 mm; lobes lanceolate, ca. 2 mm, with 1 lobe
in ca. 3 flowers per inflorescence expanded into calycophyll,
blade ovate, ca. 4 2.1 cm, densely pubescent along veins on
both surfaces, base broadly cuneate, stipe ca. 19 mm, apex
acute. Corolla pale blue, salverform, outside densely pubescent;
tube ca. 23 mm, inside densely yellow clavate puberulent at upper part; lobes orbicular-ovate, ca. 2.5 mm, adaxially sparsely
yellow papillose, mucronate. Berry globose, ca. 5 5 mm. Fl.
May.
Thickets, jungles; ca. 1500 m. S Yunnan.
The blue flowers, the lack of barbate pubescence in the throat of
the tubular corolla (according to the protologue figure), and the apparently exserted stigmas (all these features are illustrated or described in
the protologue) are quite anomalous features in Mussaenda.

23. Mussaenda parviflora Miquel, Ann. Mus. Bot. LugdunoBatavi 3: 110. 1867.
xiao yu ye jin hua
Mussaenda albiflora Hayata; M. parviflora var. formosana
Matsumura; M. taihokuensis Masamune.
Climbing shrubs or lianas; branches terete, sparsely to
densely strigillose or glabrescent. Leaves opposite; petiole 1
2.3 cm, sparsely to densely strigillose; blade drying thickly papery, ovate, elliptic, or lanceolate, 715 2.36 cm, adaxially
glabrous or strigillose on principal veins, abaxially sparsely to
densely strigillose to appressed villous especially along veins,
base acute to obtuse or subrounded, apex acuminate to caudate;
secondary veins 57 pairs, tertiary venation rather finely reticulate; stipules usually deciduous, triangular, 57 mm, moderately
to densely strigillose, deeply 2-lobed, lobes narrowly triangular
to linear. Inflorescences laxly cymose, 48 38 cm, terminal
and often in uppermost leaf axils, axes often somewhat scorpioid, sparsely to densely strigillose, pedunculate; peduncle 25
cm; bracts broadly triangular to linear, 17 mm, often caducous;
pedicels 25 mm. Flowers pedicellate (or borne on higher order
inflorescence axes), floral biology not noted. Calyx sparsely to
densely strigillose; hypanthium portion ellipsoid to turbinate,
1.55 mm; lobes linear to narrowly triangular, 36 mm, with 1
lobe on 13 flowers of some inflorescences expanded into
white calycophyll, blade broadly ovate or elliptic, 34.5 cm,

sparsely strigillose to glabrescent, base obtuse to rounded, stipe

410 mm, apex obtuse to shortly acuminate. Corolla yellow,
salverform to tubular-funnelform, outside puberulent to glabrous; tube 58 mm, densely yellow clavate pubescent in
throat; lobes ovate, 23 mm, abaxially densely yellow papillose, acute to cuspidate. Berry ellipsoid to subglobose, 1015
mm, glabrescent, calyx limb caducous, with pedicels sometimes
elongating, to 12 mm. Fl. MarMay, fr. AugDec and Jan of
following year.
Forests, thickets; 1001700 m. Guangdong, Taiwan [Japan].

24. Mussaenda pingbianensis C. Y. Wu ex H. H. Hsue & H.

Wu, Acta Phytotax. Sin. 24: 233. 1986.
ping bian yu ye jin hua
Shrubs, ca. 3 m tall; branchlets glabrous. Leaves opposite;
petiole ca. 6 mm, sparsely appressed pubescent; blade drying
papery, adaxially shiny green, abaxially pale, elliptic or ovate,
78.5 2.53.2 cm, adaxially glabrous, abaxially pubescent
along veins, base cuneate, apex acuminate; secondary veins ca.
7 pairs; stipules caducous to persistent, ovate, ca. 3.5 mm,
pubescent, deeply 2-lobed, lobes narrowly triangular. Inflorescences congested-cymose, ca. 6 10 cm, many flowered,
densely pubescent, sessile and tripartite or pedunculate; peduncles 12 cm; bracts lanceolate, 812 mm, densely pubescent.
Flowers subsessile, biology not noted. Calyx sparsely pubescent; hypanthium portion turbinate, ca. 3 mm; lobes linear to
narrowly triangular, ca. 8.5 mm, with 1 lobe in ca. 4 flowers per
inflorescence expanded into white calycophyll, blade ovate to
obovate, 5.56 cm, base obtuse to acute, stipe 1014 mm, apex
obtuse to shortly acuminate. Corolla orange to orange-red, salverform, outside sparsely to densely pubescent; tube ca. 14 mm,
inside densely yellow clavate pubescent in upper part; lobes
ovate, ca. 2.5 mm, adaxially densely yellow papillose, acuminate to mucronate. Berry unknown. Fl. May.
Valleys. SE Yunnan.
The protologue described the stipules as caducous, but the
protologue figure showed them as persistent. The specific epithet was
written as pingpienensis by H. H. Hsue and H. Wu (in FRPS 71(1):
299. 1999), but it was originally published with the spelling above.

25. Mussaenda pubescens W. T. Aiton, Hort. Kew., ed. 2, 1:

372. 1810.
yu ye jin hua
Mussaenda bodinieri H. Lveill & Vaniot; M. pubescens
var. alba X. F. Deng & D. X. Zhang; M. pubescens f. clematidiflora Chun ex H. H. Hsue & H. Wu.
Climbing shrubs, often extensively twining; branches terete, densely strigillose and sometimes also villosulous, sometimes with congested groups of leaves borne on reduced axillary buds. Leaves opposite or perhaps rarely whorled; petiole
38 mm, moderately to densely strigillose; blade drying membranous or thinly papery, ovate-oblong, ovate-lanceolate, elliptic, lanceolate, or oblanceolate, 29 14 cm, adaxially sparsely
strigillose to glabrescent on lamina and moderately to densely
strigillose on veins, abaxially sparsely to densely strigillose
with pubescence denser on veins, base acute to obtuse, apex

RUBIACEAE

acute to slightly acuminate; secondary veins 47 pairs, tertiary

venation reticulate; stipules usually deciduous, triangular, 37
mm, moderately to densely strigillose, deeply 2-lobed, lobes
narrowly triangular to subulate. Inflorescences terminal, subcapitate to congested-cymose, unbranched or sometimes tripartite, 13 13 cm, densely strigillose to villosulous, sessile to
pedunculate; peduncles 0.11.4 cm; bracts linear, 35 mm;
pedicels to 1 mm. Flowers sessile or infrequently pedicellate,
biology not noted. Calyx moderately to densely strigillose to
strigose; hypanthium portion ellipsoid to turbinate, 1.53 mm;
lobes linear to narrowly triangular, 36 mm, with 1(or 5) lobes
on 13 flowers per inflorescence of some plants expanded into
white calycophyll, blade elliptic, broadly elliptic, ovate, or lanceolate, (0.4)2.55 (0.2)23.5 cm, sparsely to moderately
puberulent or strigillose on both surfaces, base acute to
rounded, stipe 328 mm, apex obtuse to acute. Corolla white or
yellow, salverform, outside moderately to densely strigillose or
strigose; tube 1120 mm, densely clavate pubescent in throat;
lobes oblong-lanceolate to lanceolate, 2.54 mm, adaxially
densely golden yellow papillose, acuminate. Berry subglobose,
810 67.5 mm, sparsely strigillose to glabrescent, smooth or
finely lenticellate, calyx limb caducous, sometimes stipitate
with stipe to 5 mm. Fl. AprJul, fr. JunDec.
Thickets in ravines, on hill slopes, or at village margins or roadsides; below 100900 m. Fujian, Guangdong, Guangxi, Hainan, Hunan,
Jiangxi, Taiwan, Zhejiang [Vietnam].
This species is widespread, common, and morphologically variable. H. H. Hsue and H. Wu (in FRPS 71(1): 296. 1999) described the
leaf arrangement as opposite or whorled, but only opposite leaves have
been seen on the numerous specimens studied.
One plant from Guangdong (Gaoyao) has all five of the calyx
lobes petaloid and enlarged on each flower, though these structures are
smaller than calycophylls that are borne singly on a flower. These plants
have been separated as Mussaenda pubescens f. clematidiflora; this case
was studied by Deng and Zhang (Acta Phytotax. Sin. 44: 611. 2006),
who concluded that this plant is better regarded as developmentally abnormal and formally synonymized this name here. Plants with white corollas have been separated as M. pubescens var. alba; at least a third of
the specimens of M. pubescens studied belong to this group, and considering that this color variation is common in other Rubiaceae species
the variety is not recognized taxonomically here.

26. Mussaenda sessilifolia Hutchinson in Sargent, Pl. Wilson.

3: 397. 1916.
wu bing yu ye jin hua
Climbing shrubs; branches subterete, pale red- or red villous. Leaves opposite; petiole 13 mm, villous; blade drying
thickly papery, oblong-elliptic, 610 34.5 cm, densely red
villous on both surfaces with pubescence denser on principal
veins abaxially, base rounded to subtruncate, apex subacute
then abruptly narrowed and acuminate; secondary veins 9 or 10
pairs; stipules generally deciduous, ca. 8 mm, densely villous,
deeply 2-lobed, lobes linear-subulate. Inflorescence congestedcymose with developed principal axes, 1.57.5 1.54.5 cm,
densely villous, pedunculate; peduncles 0.752.2 cm; bracts
subulate, ca. 8 mm. Flowers subsessile, biology not noted. Calyx densely villous; hypanthium portion ca. 3 mm; lobes subulate, 23.5 mm, with 1 lobe of 15 flowers of each inflores-

241

cence expanded into calycophyll, blade broadly ovate, 58.5

36.5 cm, slightly pubescent, base rounded, stipe ca. 15 mm,
apex obtuse. Corolla yellow, salverform, outside densely villous; tube ca. 17 mm; lobes ovate, shortly acuminate. Berry
not seen. Fl. spring.
Forests; ca. 1300 m. S Yunnan.
The protologue described the calyx lobes as ca. 2 mm, while H. H.
Hsue and H. Wu (in FRPS 71(1): 289. 1999) described them as ca. 3.5
mm.

27. Mussaenda shikokiana Makino, Bot. Mag. (Tokyo) 18: 44.

1904.
da ye bai zhi shan
Mussaenda anomala H. L. Li; M. esquirolii H. Lveill;
M. taiwaniana Kanehira; M. wilsonii Hutchinson.
Erect or climbing shrubs, 13 m tall; branches terete,
densely strigillose and sometimes also strigose, villosulous, or
tomentulose to occasionally glabrescent. Leaves opposite; petiole 1.53.5 cm, moderately to densely strigillose; blade drying
thinly papery, adaxially green to pale green, abaxially pale gray
to whitened, broadly ovate, ovate, or broadly elliptic, 620
3.513 cm, both surfaces sparsely strigillose to glabrescent with
pubescence denser along veins, base cuneate or usually obtuse
to rounded, apex abruptly acuminate or acute; secondary veins
710 pairs, tertiary venation regularly areolate; stipules caducous, triangular to ovate-lanceolate, 610 mm, sparsely to
densely strigillose, often deeply 2-lobed, segments acute. Inflorescences subcapitate becoming laxly cymose, 25 27 cm,
densely strigillose and/or tomentulose, many flowered, sessile
and tripartite or pedunculate; peduncle 1.53 cm; bracts deciduous, ovate, lanceolate, or bilobed, 410 mm, sparsely to moderately strigose to strigillose; pedicels 14 mm. Flowers pedicellate, biology not noted. Calyx with hypanthium portion turbinate to ellipsoid, 2.55 mm, densely strigose; lobes subleaflike, white, lanceolate to ligulate, 510 22.5 mm, moderately
to densely strigillose to strigose, acute to long acuminate, with 1
lobe of 1 or few flowers on each inflorescence (or rarely all
lobes of all flowers, M. anomala) usually expanded into calycophyll, blade obovate, ovate, or elliptic, (2)34.5 cm, glabrescent on lamina and densely villosulous to strigillose on veins,
base cuneate, stipe 515(25) mm, apex acute to shortly acuminate. Corolla yellow, salverform-funnelform, outside densely
sericeous; tube 914 mm; lobes ovate, 23 mm, abruptly acuminate. Infructescences to 6 12 cm. Berry subglobose to ellipsoid, ca. 10 10 mm, moderately to densely strigose, sparsely
lenticellate, calyx limb deciduous. Fl. MayJul, fr. JulOct.
Roadsides or sparse forests on hills; 1001000 m. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan, Zhejiang [Japan].
The names Mussaenda shikokiana and M. taiwaniana were not
cited by H. H. Hsue and H. Wu (in FRPS 71(1): 283306. 1999). No
differences are apparent between M. shikokiana and M. esquirolii, as
noted by Japanese taxonomists (in herb.), and the latter are accordingly
synonymized here.
Mussaenda anomala was described based on one specimen that
appears to represent only an aberrant form of M. shikokiana with all its

RUBIACEAE

242

calyx lobes expanded and petaloid. This name was formally synonymized by Deng and Zhang (Acta Phytotax. Sin. 44: 609611. 2006,
under M. esquirolii), who reported visiting the type locality where they
were unable to find a similar plant.

28. Mussaenda simpliciloba Handel-Mazzetti, Anz. Akad.

Wiss. Wien, Math.-Naturwiss. Kl. 62: 147. 1925.
dan lie yu ye jin hua
Climbing shrubs; branches subterete, grayish brown tomentose becoming glabrescent. Leaves opposite; petiole to 4
cm, glabrous; blade drying papery, adaxially dark green, abaxially pale green, broadly ovate or elliptic-ovate, 615 37.5
cm, both surfaces densely pubescent and villous along veins,
base acute or rounded, apex acuminate; secondary veins 8 or
9 pairs; stipules triangular, ca. 6 mm, abaxially hispidulous,
adaxially densely hirsute, 2-lobed. Inflorescences laxly cymose;
bracts lanceolate; pedicels 23 mm. Flowers sessile or lateral
flowers pedicellate (or on expanded axes), biology not noted.
Calyx with hypanthium campanulate, ca. 4 mm, sparsely pubescent; lobes linear-lanceolate, 57 mm, densely pubescent,
with ?1 lobe on 1(or ?more) flower per inflorescence expanded
into calycophyll, blade ovate, ca. 6 56 cm, densely pubescent along veins, base obtuse, stipe ca. 2 mm, apex acute. Corolla orange-yellow, salverform, outside pubescent; tube ca. 23
mm, densely yellow clavate pubescent inside upper part; lobes
rounded, ca. 5 mm, adaxially yellow papillose. Berry globose,
89 89 mm, calyx limb deciduous. Fl. JunJul, fr. Aug.
Thickets in valleys or at riversides; 12001400 m. Guizhou, Sichuan, Yunnan.
The rounded corolla lobes described for this species are very
unusual in Mussaenda.

29. Mussaenda treutleri Stapf, Bot. Mag. 135: t. 8254. 1909.

gong shan yu ye jin hua
Erect or climbing shrubs; branches terete, moderately to
densely villous or hirsute. Leaves opposite; petiole 410 mm,
hirsute or villous; blade drying membranous to papery, green to
brownish, paler below when specimen well preserved, elliptic
to ovate, 2.521 411 cm, both surfaces moderately strigillose
to pilose on lamina and moderately to densely strigillose to
hirsute along principal veins, base cuneate to obtuse, truncate,

or rounded, apex acute to acuminate; secondary veins 68 pairs,

without domatia, tertiary venation reticulate; stipules generally
persistent, ovate, 712 612 mm, moderately to densely hirsute or pilose, entire to deeply 2-lobed, segments acute to
acuminate. Inflorescences subcapitate to congested-cymose or
sometimes with axes later elongating, 26 28 cm, hirsute,
sessile to subsessile; bracts lanceolate, 0.51 cm, obtuse to acuminate or 2- or 3-parted. Flowers subsessile, biology not noted.
Calyx with hypanthium portion campanulate to obconic, 34
mm, densely brown strigose to -sericeous; lobes lanceolate, ligulate, or oblanceolate, 718 34 mm, often markedly unequal
on an individual flower, densely brown strigillose to -strigose,
acute, with 1 lobe on 1 to several flowers on each inflorescence
sometimes expanded into white calycophyll, blade ovate or
rhombic, 57 3.54 cm, both surfaces sparsely hirsute to
glabrescent on lamina and moderately to densely pilosulous to
puberulent on principal veins, base obtuse to truncate, stipe 25
37 mm, apex obtuse to shortly acuminate. Corolla orange-yellow, salverform, outside densely spreading villous; tube ca. 22
mm; lobes ovate, 57 mm (to 10 mm in other regions), acute to
acuminate. Berry ellipsoid, 1012 1012 mm, strigose, lenticellate, calyx limb deciduous. Fl. JulSep.
Thickets or dense forests on mountains; [600]10001500[2000]
m. Yunnan [Bhutan, NE India, Nepal].
The description of the fruit here is based on specimens from outside our flora region.
This species is similar to Mussaenda macrophylla, and in the protologue M. treutleri was explicitly separated from plants commonly
called M. macrophylla, both in the wild and in cultivation; in fact, the
protologue noted that M. treutleri was already widely distributed in cultivation at that time incorrectly under the other name. Mussaenda treutleri subsequently has apparently still been confused with or combined
with M. macrophylla, including by recent authors (e.g., Springate et al.,
Fl. Bhutan 2(2): 783. 1999). Mussaenda treutleri is here circumscribed
somewhat differently than by H. H. Hsue and H. Wu (in FRPS 71(1):
301. 1999): the plants from montane Yunnan and adjacent regions with
broad, generally persistent stipules were separated by Stapf from M.
macrophylla and included in M. treutleri, and this distinction is followed here.
The illustration presented for Mussaenda treutleri by Fu et al.
(Higher Pl. China 10: 575. 2004) is incorrect; this figure shows a
species of Asteraceae.

57. MYCETIA Reinwardt, Syll. Pl. Nov. 2: 9. 1825.

xian e mu shu
Chen Tao (); Charlotte M. Taylor
Adenosacme Wallich ex Miquel, nom. illeg. superfl.
Small shrubs, unarmed, usually rather succulent; bark often straw-yellow to nearly white, soft, often corky. Raphides present.
Leaves opposite, sometimes weakly to markedly anisophyllous, frequently somewhat asymmetrical or falcate, apparently without
domatia, often with margins crisped; tertiary venation often closely reticulate and prominulous below and frequently also above;
stipules persistent with leaves or caducous, interpetiolar, triangular or sometimes leaflike, often well developed, sometimes with
glands. Inflorescences terminal, pseudoaxillary, cauline, or perhaps sometimes axillary, cymose to paniculiform or subcapitate, several to many flowered, pedunculate, bracteate or bracts reduced; bracts sometimes leaflike, sometimes glandular. Flowers pedicellate,
bisexual, at least usually distylous. Calyx limb deeply (4 or)5(or 6)-lobed; lobes frequently somewhat unequal, usually with sessile to
stalked marginal glands. Corolla yellow or white, tubular, funnelform, salverform, or campanulate, sometimes gibbous at base, inside
glabrous or usually pubescent; lobes (4 or)5(or 6), in bud valvate-induplicate. Stamens (4 or)5(or 6), inserted in throat or above mid-

RUBIACEAE

243

dle of corolla tube in short-styled flowers, inserted near base of corolla tube in long-styled flowers; filaments short or reduced; anthers apparently dorsifixed, included or partially exserted. Ovary 2(5)-celled, ovules numerous in each cell on fleshy axile placentas; stigmas 2(5), included in short-styled flowers or exserted in long-styled flowers. Fruit white or perhaps brown, baccate or perhaps capsular and irregularly dehiscent, leathery to fleshy or spongy, subglobose, with calyx limb persistent; seeds numerous, small,
angled, with testa somewhat granular.
About 45 species: tropical and subtropical Asia; 15 species (ten endemic) in China.
This genus was reviewed for Thailand by Fukuoka (Acta Phytotax. Geobot. 40: 107118. 1989), for the Indian subcontinent by Deb (Bull. Bot.
Surv. India 28(14): 114132. 1986), and for China by H. S. Lo (Guihaia 11: 105116. 1991). Mycetia is said to belong to Isertieae but also,
anomalously, to have raphides. H. S. Lo (in FRPS 71(1): 314. 1999) reported that the inflorescences may sometimes be axillary, but this has not been
noted by other authors; this description may be using the term axillary for the position elsewhere considered pseudoaxillary. Lo also reported that
the anthers may sometimes be partially exserted, which has not been reported by other authors nor seen on specimens studied by us.

1a. Pedicels mostly or all 920 mm.

2a. At least some nodes with markedly anisophyllous leaves, larger leaf 310 (or more) as long as smaller leaf;
calyx lobes 2.53 mm; corolla tube 1216 mm ....................................................................................................... 7. M. gracilis
2b. Leaves generally isophyllous, larger leaf at most 2 as long as smaller leaf; calyx lobes 0.86 mm; corolla
tube 510 mm (unknown in M. yunnanica).
3a. Calyx lobes 36 mm, markedly longer than length of hypanthium together with unlobed basal part
of limb ............................................................................................................................................................ 2. M. bracteata
3b. Calyx lobes 0.83 mm, shorter than or equal to length of hypanthium together with unlobed basal
part of limb.
4a. Inflorescences terminal and pseudoaxillary; calyx lobes 0.81 mm ................................................... 4. M. brevisepala
4b. Inflorescences pseudoaxillary or possibly axillary at lower stem nodes, often below leaves;
calyx lobes 2.53 mm ........................................................................................................................... 15. M. yunnanica
1b. Flowers subsessile or with pedicels mostly or all 8 mm or shorter.
5a. At least some calyx limbs and sometimes also bracts bearing few to numerous marginal glands on short
to well-developed flexuous stipes.
6a. Stipules elliptic-oblong, obovate, or suborbicular and narrowed to stipitate at base, at base less than
1/2 as wide as maximum width; calyx lobes with 13 pairs of glands .......................................................... 14. M. sinensis
6b. Stipules lanceolate, oblong-lanceolate, or ovate, at base straight or narrowed but more than 2/3 as
wide as maximum width; calyx lobes with numerous stipitate glands.
7a. Corolla with tube 46 mm; leaves moderately to densely hispidulous or hirtellous adaxially ..................... 9. M. hirta
7b. Corolla with tube 714 mm; leaves glabrous or sparsely strigillose or hispidulous adaxially.
8a. Petioles 115 mm; calyx lobes 11.5 mm; corolla tube 710 mm ................................................ 6. M. glandulosa
8b. Petioles 660 mm; calyx lobes 1.54 mm; corolla tube 1014 mm .............................................. 11. M. longifolia
5b. Calyx limbs and bracts without raised or stipitate marginal glands.
9a. Calyx lobes 1.52 mm; corolla tube 57 mm (mature corollas unknown in M. hainanensis).
10a. Calyx densely hirtellous; flowers sessile or subsessile; stipules ovate, not strongly contracted
at base ................................................................................................................................................. 8. M. hainanensis
10b. Calyx glabrous; flowers pedicellate; stipules elliptic-oblong, obovate, or suborbicular,
contracted to stipitate at base ................................................................................................................. 14. M. sinensis
9b. Calyx lobes 36 mm; corolla tube 717 mm (corolla unknown in M. macrocarpa).
11a. Stipules broadly elliptic, subovate, or suborbicular, 510 mm.
12a. Leaves glabrous on both surfaces; fruit ca. 6 mm; Yunnan .................................................. 12. M. macrocarpa
12b. Leaves strigillose to hirtellous abaxially; fruit 78 mm; Xizang ............................................ 13. M. nepalensis
11b. Stipules triangular, lanceolate, suborbicular, or ovate, 36 mm.
13a. Corolla tube 78 mm; leaves subsessile or with petioles up to 15 mm.
14a. Leaves subsessile; inflorescences several flowered (i.e., with 57 flowers) ...................... 3. M. brevipes
14b. Leaves with petioles 315 mm; inflorescences many flowered (i.e., with
940 flowers) ........................................................................................................................ 5. M. coriacea
13b. Corolla tube 1017 mm; leaves with petioles 1070 mm.
15a. Flowers sessile or subsessile; petioles 1020 mm .......................................................... 1. M. anlongensis
15b. Flowers pedicellate with pedicels 2.57 mm; petioles 2070 mm ................................. 10. M. longiflora
1. Mycetia anlongensis H. S. Lo, Guihaia 11: 108. 1991.
an long xian e mu
Shrubs, ca. 40 cm tall; branches glabrous or sparsely villosulous. Leaves generally isophyllous; petiole 12 cm, densely

villosulous (var. anlongensis) or villous and sometimes with

black glandular hairs at each side (var. multiciliata); blade
drying membranous, elliptic-oblong or long elliptic, 1217 5
7 cm, adaxially sparsely strigillose (var. anlongensis) or sparsely
hirsute (var. multiciliata), abaxially densely villosulous at least

RUBIACEAE

244

along midrib and lateral veins (var. anlongensis) or sparsely

hirsute (var. multiciliata), base cuneate, apex acute to shortly or
long acuminate; secondary veins 1418 pairs; stipules persistent, suborbicular or elliptic, 56 mm, obtuse. Inflorescences
terminal, congested-cymose, several to many flowered; peduncle ca. 5 mm. Flowers sessile to subsessile. Calyx glabrescent;
hypanthium portion obconic, ca. 2 mm; limb lobed nearly to
base; lobes narrowly triangular, ca. 6 mm. Corolla yellow, tubular, outside glabrous; tube 1013 mm (var. multiciliata) or 15
17 mm (var. anlongensis), sparsely villosulous inside; lobes
triangular, ca. 3 mm. Berries unknown. Fl. AprMay.
Streamsides in dense forests, sometimes on limestone hill
slopes; 12001700 m. Guangxi, Guizhou.
H. S. Lo (in FRPS 71(1): 316. 1999) intended to describe two
varieties of this species but did not provide a validating Latin description or diagnosis or designate a type for Mycetia anlongensis var. multiciliata. Chen et al. (J. Fairylake Bot. Gard. 7(2): 21. 2008) have re-evaluated these taxa with new collections; in particular, they noted that they
cannot confirm the description by H. S. Lo of black glandular trichomes
on the petioles in var. multiciliata, a character not otherwise reported
from this genus and considered questionable.

1a. Leaves shortly acuminate at apex; petioles

without black glandular hairs at base;
corolla 1820 mm ............................... 1a. var. anlongensis
1b. Leaves long acuminate at apex, with
tip 13 cm; petioles sometimes with
black glandular hairs at base; corolla
1315 mm ............................................ 1a. var. multiciliata
1a. Mycetia anlongensis var. anlongensis
() an long xian e mu (yuan bian zhong)
Petiole without black glandular hairs at base on each side;
leaf blade adaxially sparsely strigillose, abaxially densely villosulous at least along midrib and lateral veins, shortly acuminate at apex. Corolla 1820 mm. Fl. May.
Streamsides in dense forests; 12001700 m. Guizhou (Anlong).

1b. Mycetia anlongensis var. multiciliata H. S. Lo ex Tao

Chen, K. J. Yan & D. Fang, J. Fairylake Bot. Gard. 7(2): 21.
2008.
na po xian e mu
Petiole sometimes with black glandular hairs at each side;
leaf blade sparsely hirsute on both surfaces, long acuminate at
apex, with tip 13 cm. Corolla 1315 mm. Fl. Apr.
Limestone hill slopes; ca. 1200 m. Guangxi (Napo).

2. Mycetia bracteata Hutchinson in Sargent, Pl. Wilson. 3:

409. 1916.
chang bao xian e mu
Shrubs, ca. 1 m tall; branches puberulent becoming glabrescent. Leaves generally isophyllous; petiole 0.51 cm, puberulent; blade drying thinly leathery to membranous, long oblanceolate or lanceolate, 916 1.54 cm, both surfaces glabrous except puberulent along principal veins, base attenuate or
acute, apex acuminate; secondary veins 1620 pairs; stipules

persistent, broadly triangular-ovate, 710 mm, veined, stiffly

papery, puberulent, acute. Inflorescences terminal, laxly cymose, several flowered, puberulent; branched portion to 7 cm;
bracts persistent, lanceolate or leaflike, to 1.5 cm, acuminate or
shortly acuminate; pedicels 1015 mm. Flowers pedicellate.
Calyx glabrescent; hypanthium portion obconic, ca. 2 mm; limb
deeply lobed; lobes 5 or 6, filiform, 36 mm. Corolla yellow, in
bud glabrous, ca. 10 mm. Berries not seen.
About 1300 m. Yunnan (Simao).

3. Mycetia brevipes F. C. How ex S. Y. Jin & Y. L. Chen, Cat.

Type Spec. Herb. China (Suppl.), 188. 1999.
duan bing xian e mu
Shrubs, ca. 1 m tall. Branchlets glabrous. Leaves generally
isophyllous, subsessile; blade drying membranous and fuscous
or dark brown, elliptic or obovate, 1215 4.56.5 cm, both
surfaces glabrous or puberulent on principal veins, base cuneate
to obtuse, apex acuminate to caudate; secondary veins 811
pairs; stipules persistent, ovate, 45 mm, membranous. Inflorescences terminal, congested-cymose, 57-flowered; peduncle ca.
1.6 cm; bracts linear, ca. 3 mm; pedicels 13 mm. Flowers
pedicellate. Calyx puberulent; hypanthium portion obconic, ca.
1.8 mm; limb lobed nearly to base; lobes linear-lanceolate, ca. 3
mm. Corolla yellow, tubular, outside glabrous; tube ca. 7 mm,
sparsely villous inside; lobes broadly ovate-triangular, ca. 1.5
mm. Berries not seen. Fl. Sep.
Bamboo forests; ca. 1500 m. NW Yunnan.

4. Mycetia brevisepala H. S. Lo, Guihaia 11: 113. 1991.

duan e xian e mu
Shrubs, 0.51.5(2) m tall; branches densely puberulent or
strigillose to glabrous. Leaves generally isophyllous to slightly
anisophyllous; petiole 0.21 cm, puberulent; blade drying thinly
leathery to papery and grayish green, elliptic-oblong, elliptic,
obovate, or oblong-lanceolate, 618 2.56 cm, adaxially glabrous and rather shiny, abaxially glabrescent or densely puberulent to hispidulous on principal veins, base cuneate to obtuse,
apex acuminate; secondary veins 712 pairs; stipules quickly
deciduous, narrowly triangular, 36 mm, densely puberulent
to strigillose, acuminate. Inflorescences terminal becoming displaced to pseudoaxillary, laxly cymose, several flowered, puberulent to glabrous; peduncle 12 cm; branched portion 35
58 cm; bracts narrowly triangular to lanceolate, 13 mm; pedicels 920 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion turbinate to subglobose, 1.22 mm; limb deeply
lobed; lobes triangular to narrowly triangular, 0.81 mm. Corolla yellow, narrowly tubular, outside glabrous; tube ca. 5 mm,
inside sparsely pubescent; lobes subtriangular, ca. 2 mm. Berries
subglobose, 3.54 mm in diam. Fl. AugSep, fr. Dec.
Dense forests; 2001100 m. Yunnan [N Vietnam].

5. Mycetia coriacea (Dunn) Merrill, Philipp. J. Sci., C, 13: 159.

1918.
ge ye xian e mu
Adenosacme coriacea Dunn, Bull. Misc. Inform. Kew,
Addit. Ser. 10: 130. 1912.

RUBIACEAE

Shrubs, 12 m tall; branches glabrous. Leaves isophyllous

to slightly anisophyllous; petiole 315 cm, glabrous; blade
drying thinly leathery or subpapery, lanceolate, oblanceolate, or
obovate, 615 25.5 cm, glabrous on both surfaces, base acuminate to cuneate, apex acute or acuminate; secondary veins 8
11 pairs; stipules persistent, triangular to ovate, 35 mm, glabrous except sometimes ciliolate marginally, acute to acuminate. Inflorescences terminal and/or pseudoaxillary, cymose,
many flowered, glabrous; peduncle 0.83.5 cm; branched portion 26 37 cm; bracts reduced or triangular, 0.53 mm;
pedicels 0.53 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion turbinate to obconic, 1.52.5 mm; limb lobed
to base; lobes narrowly triangular to linear, 2.56 mm, sometimes unequal. Corolla pale yellow or white, tubular, outside
glabrescent; tube ca. 8 mm, villous at throat; lobes triangular,
ca. 2 mm. Fruit capsular, dry, ovoid to subglobose, 2.54 mm,
irregularly dehiscent or regularly loculicidal through disk portion, with calyx lobes elongating, up to 15 mm. Fl. Apr, fr. Sep.
Roadsides or streamsides in dense forests; 7001200 m. S Fujian, E Guangdong.

6. Mycetia glandulosa Craib, Bull. Misc. Inform. Kew 1914:

125. 1914.
xian e mu
Shrubs, ca. 1 m tall; branches villosulous to hirtellous becoming glabrescent. Leaves isophyllous or slightly anisophyllous; petiole 18 mm, moderately to densely villosulous or hirtellous; blade drying papery, oblanceolate, oblong-oblanceolate,
narrowly elliptic, or narrowly lanceolate, 722 25.5 cm,
adaxially sparsely hispidulous, abaxially moderately to densely
hirtellous, base cuneate to attenuate and often long decurrent,
apex acuminate; secondary veins (7)1223 pairs; stipules persistent, lanceolate, 410 mm, densely villosulous to glabrescent,
acuminate. Inflorescences terminal, congested to laxly cymose,
many flowered, hirtellous to glabrous; peduncles 0.41.4 cm;
branched portion 35 48 cm; bracts ovate to reniform, 13
mm, marginally ciliate or lacerate with stipitate glands; pedicels
1.55 mm. Flowers pedicellate. Calyx glabrous; hypanthium
portion hemispherical to subglobose, ca. 1.5 mm; limb deeply
lobed; lobes ovate to lanceolate or triangular, 11.5 mm, marginally fimbriate or ciliate with stipitate glands, these 0.20.6
mm. Corolla white or yellow, tubular, outside pilose or glabrous; tube 710 mm, villous inside; lobes triangular, 11.5
mm. Berries subglobose, ca. 5 mm in diam., subglabrous. Fl.
May, fr. autumn.
Forests; 9001500 m. Yunnan [N Thailand].

7. Mycetia gracilis Craib, Bull. Misc. Inform. Kew 1914: 125.

1914.
xian geng xian e mu
Shrubs, ca. 1.5 m tall; branches densely puberulent or strigillose becoming glabrescent. Leaves markedly anisophyllous;
petiole 0.10.5 cm, densely puberulent to strigillose; blade
drying thinly leathery, oblanceolate, elliptic, or narrowly lanceolate, larger 515 23.5 cm and smaller 14 0.51.5 cm,
glabrous throughout or puberulent abaxially on principal veins,
base acute to cuneate, apex acute to acuminate; secondary veins

245

814 pairs; stipules persistent, narrowly triangular to lanceolate,

48 mm, puberulent to glabrescent, acuminate. Inflorescences
terminal and/or pseudoaxillary, laxly cymose, several flowered,
puberulent to glabrescent; peduncle 12.5 cm; branched portion
29 28 cm; bracts linear, 1.53 mm; pedicels 915 mm.
Flowers pedicellate. Calyx glabrous; hypanthium portion turbinate to obovoid, 22.5 mm; limb lobed to base; lobes linear to
narrowly triangular, 2.53 mm. Corolla yellow, tubular, outside
glabrous; tube 1216 mm, pilose inside; lobes triangular to
ovate, ca. 3 mm. Berries globose or slightly compressed, 46
mm in diam., glabrous. Fl. AugSep, fr. NovDec.
Streamsides in forests; 6001300 m. S Yunnan [N Thailand, Vietnam].

8. Mycetia hainanensis H. S. Lo, Guihaia 11: 112. 1991.

hai nan xian e mu
Subshrubs, 1025 cm tall; branches hirtellous or villosulous. Leaves generally isophyllous; petiole 12.5 cm, villosulous to glabrescent; blade drying papery, adaxially grayish
brown or nearly grayish black, abaxially grayish brown, oblanceolate or narrowly elliptic, 512 34 cm, adaxially glabrous,
abaxially glabrescent on lamina and villosulous on veins, base
acute to attenuate, apex acute or weakly acuminate; secondary
veins 912 pairs; stipules persistent, ovate, 59 mm, puberulent
to glabrescent, obtuse. Inflorescences terminal, capitate or subcapitate, villosulous, sessile to pedunculate; peduncles when
present 1.53.5 cm, villosulous; head ca. 1.5 1.5 cm; bracts
apparently reduced. Flowers subsessile to sessile. Calyx densely
hirtellous; hypanthium portion obconical-ovoid, ca. 1.5 mm;
limb deeply lobed; lobes triangular-ovate, ca. 1.5 mm, obtuse.
Corolla in bud white, tubular, outside hirtellous; tube to ca. 5
mm, subglabrous inside; lobes ovate, to ca. 2 mm. Berries not
seen. Fl. Apr.
Dense forests on mountains; ca. 800 m. Hainan.

9. Mycetia hirta Hutchinson in Sargent, Pl. Wilson. 3: 410.

1916.
mao xian e mu
Shrubs, 12 m tall; branches densely villosulous, hirtellous, or tomentose sometimes becoming glabrescent. Leaves
isophyllous to slightly anisophyllous; petiole 0.73 cm, densely
villosulous to hirtellous; blade drying papery, oblong-elliptic to
elliptic or ovate, 825 3.59 cm, both surfaces moderately to
densely hispidulous to hirtellous, base obtuse to acute and often
long decurrent, apex acute to acuminate; secondary veins 1223
pairs; stipules usually persistent with leaves, oblong-lanceolate
to ovate, 0.82 cm, glabrous except villous along midrib and
sometimes margins, acute, acuminate, or shortly 2-lobed. Inflorescences terminal, congested- to laxly cymose, several to many
flowered, densely villosulous or hirtellous; peduncle 0.73 cm;
branched portion 1.56 2.59 cm; bracts ovate, lanceolate,
suborbicular, reniform, or stipuliform, 210 mm, often marginally sparsely to densely stipitate-glandular; pedicels 1.54
mm. Flowers pedicellate. Calyx densely hirtellous; hypanthium
portion subglobose-campanulate, ca. 2 mm; limb deeply lobed;
lobes broadly triangular, 22.5 mm, marginally densely stipitate-glandular and sometimes appearing lacerate, glands 0.30.6

RUBIACEAE

246

mm. Corolla yellow, tubular, outside sparsely to densely villosulous or hirtellous; tube 46 mm, sparsely villous inside;
lobes triangular, 11.8 mm. Berries capsular, subglobose, 3.5
4.5 mm in diam., densely hirtellous or villosulous. Fl. JunJul,
fr. SepOct.
Forests; 5001600 m. Hainan, Yunnan.

10. Mycetia longiflora F. C. How ex H. S. Lo, Guihaia 11: 107.

1991.
chang hua xian e mu
Mycetia longiflora f. howii H. S. Lo.
Shrubs, 0.62 m tall; branches puberulent becoming glabrescent. Leaves generally isophyllous; petiole 27 cm, densely
tomentulose or puberulent; blade drying thinly papery, ellipticoblong, obovate, or elliptic, 1025 410.5 cm, adaxially glabrous or sparsely puberulent along midrib, abaxially glabrous or
puberulent to tomentulose along principal veins, base cuneate to
acute and often long decurrent, apex acute to acuminate; secondary veins 1015 pairs; stipules persistent, triangular to
ovate, 35 mm, puberulent, obtuse, acute, or shortly 2-lobed.
Inflorescences terminal, laxly cymose, densely puberulent to
tomentulose, many flowered, often deflexed to pendulous;
peduncle 0.31 cm; branched portion 45 710 cm; bracts
reduced or narrowly triangular to lanceolate, 25 mm; pedicels
2.57 mm. Flowers pedicellate. Calyx densely puberulent,
tomentulose, or glabrescent; hypanthium portion subglobose to
obconic, 1.52.5 mm; limb lobed to base; lobes narrowly triangular to narrowly lanceolate, 46.5 mm. Corolla yellow, tubular, densely puberulent to glabrescent outside; tube 1416
mm, white villous above middle inside or in throat; lobes
broadly ovate-triangular, 22.5 mm. Berries subglobose, ca. 5
mm in diam., puberulent to glabrescent. Fl. JulAug, fr. Oct
Jan.
Dense forests; 6001700 m. Yunnan.
Two forms of this species were separated in the protologue and by
H. S. Lo in FRPS (71(1): 315. 1999). Lo distinguished Mycetia longiflora f. howii by its glabrous corollas and glabrous smaller leaves though
no measurements were given. Presumably f. longiflora thus comprised
plants with pubescent to subglabrous corollas and pubescent, larger
leaves. The characters used to distinguish these forms vary continuously
within most populations and species of Rubiaceae; accordingly, they are
not recognized here.

11. Mycetia longifolia (Wallich) Kuntze, Revis. Gen. Pl. 1:

289. 1891.
chang ye xian e mu
Rondeletia longifolia Wallich in Roxburgh, Fl. Ind. 2: 137.
1824; Adenosacme longifolia (Wallich) J. D. Hooker; Wendlandia longifolia (Wallich) Candolle.
Shrubs, to 2 m tall; branches hirtellous or villosulous becoming glabrescent. Leaves isophyllous or slightly anisophyllous; petiole 0.62.5(6) cm, hirtellous or villosulous; blade
drying papery, elliptic-lanceolate or elliptic, 518(35) 3
7(10) cm, adaxially sparsely strigillose, hispidulous, or glabrous, abaxially sparsely to densely puberulent or hirtellous to
glabrescent, base cuneate to acute and often decurrent, apex acu-

minate to caudate; secondary veins 1320 pairs; stipules usually persistent, oblong-lanceolate to ovate, 515 mm, villosulous or hirtellous, acute to acuminate and sometimes shortly 2lobed. Inflorescences terminal or sometimes pseudoaxillary,
laxly cymose, villosulous to glabrous, subsessile to pedunculate; peduncle 0.51.5 cm; branched portion 34 56 cm;
bracts elliptic to ovate, 13 mm, marginally sparsely stipitateglandular; pedicels 25 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion subglobose, 1.52 mm; limb deeply
lobed; lobes triangular, 1.54 mm, with 1 or 2 pairs of stipitate
glands on margins, these 0.20.5 mm. Corolla yellow, tubular,
outside glabrous to villosulous; tube 1014 mm, inside villous;
lobes broadly ovate, 11.5 mm. Berries subglobose, 45 mm in
diam., glabrescent. Fl. summer and autumn.
Forests. Xizang (Mdog), S Yunnan [Bangladesh, Bhutan, NE India, Malaysia, Myanmar, Nepal].
Springate et al. (Fl. Bhutan 2(2): 784. 1999) observed that this
species sometimes grows as an epiphyte.

12. Mycetia macrocarpa F. C. How ex H. S. Lo, Guihaia 11:

111. 1991.
da guo xian e mu
Shrubs; branches glabrous. Leaves generally isophyllous;
petiole 11.5 cm, glabrous; blade drying papery, narrowly elliptic-oblong or narrowly lanceolate, 814 23.5 cm, glabrous
on both surfaces, rather shiny adaxially, base cuneate, apex caudate-acuminate; secondary veins 1012 pairs; stipules persistent, suborbicular to broadly elliptic, 710 mm, parallel-veined,
glabrous, rounded. Inflorescences and flowers not seen. Infructescences terminal, cymose, with ca. 5 fruit. Fruit obovoid to
subglobose, ca. 6 mm, with persistent calyx lobes lanceolate, 5
6 mm. Fr. Jan.
Forests; ca. 100 m. Yunnan (Malipo).

13. Mycetia nepalensis H. Hara, J. Jap. Bot. 52: 198. 1977.

chui hua xian e mu
Shrubs, 0.52 m tall; branches strigillose or hirtellous becoming glabrescent. Leaves generally isophyllous; petiole 2
4.5 cm, strigillose to hirtellous; blade drying papery, elliptic,
obovate, or ovate-elliptic, 1025 3.510 cm, adaxially glabrous or hispidulous along veins, abaxially strigillose to hirtellous, base cuneate to acute or attenuate, apex acute to acuminate; secondary veins 1018 pairs; stipules persistent or deciduous, subovate to suborbicular, 510 mm, strigillose, acute and
shortly 2-lobed. Inflorescences terminal, often pendulous, laxly
cymose, many flowered, strigillose to glabrescent; peduncle 1
2 cm; branched portion 47 47 cm; bracts lanceolate to
triangular, 310 mm; pedicels 1.55 mm. Flowers pedicellate.
Calyx puberulent to hispidulous; hypanthium portion narrowly
turbinate, 23 mm; limb lobed to base; lobes narrowly triangular, 35.5 mm. Corolla yellow to deep yellow, tubular, glabrescent outside; tube 812 mm, villous inside; lobes lanceolate, 1.22 mm. Berries subglobose, 78 mm in diam., glabrescent. Fl. AprMay, fr. Aug.
Broad-leaved rain forests; ca. 1000 m. Xizang (Mdog) [NE India,
Nepal, ?Vietnam].
H. S. Lo (in FRPS 71(1): 320. 1999) reported that the calyx lobes
have glands on each side at their bases, but the protologue, Springate et

RUBIACEAE

al. (Fl. Bhutan 2(2): 785. 1999), and Deb (Bull. Bot. Surv. India 28(1
4): 124. 1986) said explicitly that this species does not have glands on
the calyx lobes. Adenosacme nepalensis (Wallich, Numer. List, no.
6281. 1832) belongs here but is a nomen nudum and was therefore not
validly published (Vienna Code, Art. 32.1(d)).

247

ovate, 1.52 mm. Berries subglobose, 44.5 mm in diam., glabrous. Fl. JulAug, fr. SepNov.
Streamsides or roadsides in dense forests; 2001000 m. S Fujian, Guangdong, Guangxi, Hainan, Hunan, Jiangxi, Yunnan.

14. Mycetia sinensis (Hemsley) Craib, Bull. Misc. Inform.

Kew 1914: 29. 1914.

This species is said to be locally common in the notes on several

specimens.

hua xian e mu

H. S. Lo (Guihaia 11: 115116. 1991) described three forms of

this species, which were recognized in FRPS (71(1): 322323. 1999).
Mycetia sinensis f. angustisepala was described from Guangxi and distinguished by its narrowly lanceolate calyx lobes that are as long as
the calyx tube (i.e., the hypanthium portion together with the unlobed
portion of the calyx limb); f. trichophylla (not a validly published
name: Vienna Code, Art. 37.6) was described from Guangdong and distinguished by its leaf blades that are sparsely villous throughout; and f.
sinensis presumably has broader and/or differently shaped calyx lobes
that are longer than or shorter than the calyx tube and glabrous or appressed pubescent leaves. The distinctions used to separate these forms
do not seem taxonomically meaningful; accordingly, these forms are not
recognized here.

Adenosacme longifolia (Wallich) J. D. Hooker var.

sinensis Hemsley, J. Linn. Soc., Bot. 23: 379. 1888; Mycetia
oligodonta Merrill; M. sinensis f. angustisepala H. S. Lo.
Shrubs or subshrubs, 0.20.5(1) m tall; branches densely
hirtellous or strigillose to glabrous. Leaves isophyllous or
slightly anisophyllous; petiole 0.22 cm, hirtellous or strigillose
to glabrous; blade drying submembranous, usually pale, and
slightly grayish green, oblong-lanceolate to elliptic-oblong,
ovate, or elliptic, 820 35 cm, adaxially sparsely hispid to
glabrous, abaxially glabrous or puberulent, hirtellous, or hispidulous at least on veins, base obtuse to cuneate then usually long
decurrent, apex acute to acuminate; secondary veins 520 pairs;
stipules usually persistent, elliptic-oblong, obovate, or suborbicular, markedly contracted to stipitate at base, 318 mm, sometimes veined, hispidulous, hirtellous, or glabrous, obtuse or
rounded. Inflorescences terminal, laxly cymose, many flowered, glabrous, deflexed to pendulous; peduncles 3.57 cm;
branched portion 27 216 cm; bracts elliptic, obovate, reniform, or stipuliform, sometimes fused in pairs, 13 mm, marginally entire or with stipitate glands, these 0.20.5 mm; pedicels 12.5 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion subglobose to obconic, 12 mm; limb deeply
lobed; lobes lanceolate, spatulate, or triangular, 12 mm, entire
or with 13 pairs of stipitate glands. Corolla white, tubular, outside glabrous; tube 57 mm, inside apparently glabrous; lobes

15. Mycetia yunnanica H. S. Lo, Guihaia 11: 116. 1991.

yun nan xian e mu
Shrubs or subshrubs, ca. 1.5 m tall; branches glabrous.
Leaves generally isophyllous; petiole 13 cm, subglabrous;
blade drying thinly leathery, elliptic, elliptic-oblong, or obovate,
1020 3.56 cm, adaxially glabrous, abaxially glabrescent or
hirtellous along midrib, base cuneate or attenuate, apex abruptly
acuminate; secondary veins 1218 pairs; stipules elliptic-oblong, 812 mm, glabrous, obtuse. Inflorescences and flowers
not seen. Infructescences pseudoaxillary or perhaps axillary,
often on lower leafless nodes, glabrous; peduncle ca. 3 cm.
Berries subglobose, ca. 4 mm in diam., with persistent calyx
lobes ovate-triangular, 2.53 22.5 mm, obtuse. Fr. Oct.
Streamsides in forests. Yunnan (Dehong).

58. MYRIONEURON R. Brown ex Bentham & J. D. Hooker, Gen. Pl. 2: 69. 1873.
mi mai mu shu
Chen Tao (); Charlotte M. Taylor
Small shrubs, subshrubs, or large herbs, unarmed, often succulent; bark often soft or spongy, usually grayish white. Raphides
absent. Leaves opposite, apparently without domatia, sometimes with crisped margins; stipules persistent or deciduous, interpetiolar,
generally triangular, often closely densely parallel- to palmately veined, entire or shortly bilobed. Inflorescences terminal and/or
pseudoaxillary, laxly cymose or usually congested-cymose to subcapitate, several to many flowered, sessile to pedunculate, bracteate
with bracts usually well developed, often densely veined to stipuliform, sometimes outermost (i.e., basalmost) 4 or 6 bracts shortly
fused at base into an involucre. Flowers sessile to pedicellate, bisexual, distylous. Calyx limb 5-lobed; lobes often densely parallelveined. Corolla white or yellow, tubular to salverform, villous in throat; lobes 5, valvate in bud. Stamens 5, inserted in corolla tube,
included or perhaps partially exserted; filaments short; anthers apparently dorsifixed, included. Ovary 2-celled, ovules numerous in
each cell on presumably axile placentas; stigma 2-lobed with lobes linear, included or shortly exserted. Fruit white, baccate, fleshy to
rather dry, ovoid to globose, with calyx limb persistent; seeds numerous, small, angled, with endosperm fleshy; embryo small; testa
areolate.
About 14 species: Bhutan, China, India, Nepal, Vietnam; four species (one endemic) in China.
H. S. Lo (in FRPS 71(1): 309. 1999) described the inflorescences as sometimes axillary, but this has not been noted elsewhere; the term here
may be used to include the position separated elsewhere as pseudoaxillary. Lo also reported that the anthers are sometimes partially exserted in
long-styled flowers of Myrioneuron faberi and M. effusum (loc. cit.: 310, 313), which is a new condition not previously noted for the genus.
The number of pairs of lateral leaf veins described by H. S. Lo (loc. cit.: 310313) for Myrioneuron species and used in part to distinguish
species does not correspond to the secondary veins on specimens cited and apparently includes both secondary and intersecondary veins; thus, the

RUBIACEAE

248

counts disagree with those of Wright (Fl. Bhutan 2(2): 786. 1999, M. nutans) and the illustrations of Fu and Hong (Higher Pl. China 10: 577578.
2004), which consider only the secondary veins as done by most Rubiaceae authors.

1a. Corolla with tube 56 mm; calyx lobes 1012 mm ......................................................................................................... 3. M. nutans
1b. Corolla with tube 7.510 mm; calyx lobes 614 mm.
2a. Shrubs; leaves with secondary veins 1525 pairs; calyx lobes 67 mm ................................................................ 1. M. effusum
2b. Herbs, sometimes becoming suffrutescent; leaves with secondary veins 918 pairs; calyx lobes 614 mm.
3a. Stipules 6.515 mm; inflorescences shortly pedunculate; corolla with tube 910 mm and lobes 1.52 mm ... 2. M. faberi
3b. Stipules 1225 mm; inflorescences subsessile; corolla with tube 7.58 mm and lobes 2.53 mm ............ 4. M. tonkinense
1. Myrioneuron effusum (Pitard) Merrill, J. Arnold Arbor. 23:
195. 1942.
da ye mi mai mu
Myrioneuron nutans Wallich ex Kurz var. effusum Pitard
in Lecomte, Fl. Indo-Chine 3: 192. 1923.
Shrubs, to 2 m tall; branches densely puberulent or often
papillose to glabrous. Petiole 0.54 cm, densely puberulent; leaf
blade in life adaxially dark green, abaxially pale white, drying
papery, elliptic-oblong, oblanceolate, or obovate, 1325 5.5
14 cm, adaxially glabrous, abaxially densely puberulent at least
on veins, base acute to cuneate, apex acute; secondary veins
1523 pairs; stipules ovate to lanceolate, 1.52.2 cm, densely
puberulent, distinctly palmately veined, acute to shortly bilobed. Inflorescences pseudoaxillary and/or infrequently terminal, congested-cymose, subglobose to ovoid, subsessile, 1.52
cm; bracts lanceolate or ovate, 810 mm, densely puberulent,
densely longitudinally veined, acuminate. Flowers subsessile.
Calyx puberulent to glabrous; hypanthium portion obconic,
1.53 mm; limb lobed to base; lobes lanceolate to narrowly
triangular or linear, 67 mm. Corolla tubular; tube ca. 10 mm;
lobes oblong-ovate, ca. 2.5 mm. Berries subglobose, 34 45
mm, glabrescent. Fl. JulAug, fr. OctNov.
On rocks in ravines; 500700 m. Guangxi (Longzhou) [N Vietnam].

2. Myrioneuron faberi Hemsley, J. Linn. Soc., Bot. 23: 380.

1888 [faberii].
mi mai mu
Myrioneuron oligoneuron Handel-Mazzetti.
Large to suffruticose herbs, 0.21 m tall; branches densely
puberulent, shortly pilosulous, or tomentulose. Leaves sometimes crowded at upper part of branches; petiole 0.71(2) cm,
densely puberulent to tomentulose; blade drying papery and
grayish green, pale abaxially, obovate, elliptic, or oblong-obovate, (10)1223 4.510 cm, adaxially glabrous, abaxially
densely puberulent to glabrescent on lamina and densely puberulent to tomentulose along veins, base acute to obtuse, apex
cuspidate or acute; secondary veins 915 pairs; stipules lanceolate-oblong to ovate or narrowly triangular, 6.515 mm, glabrous to densely puberulent, densely parallel-veined to palmately veined, acute to shortly bilobed. Inflorescences terminal,
globose; peduncle 310 mm; branched portion subcapitate to
congested-cymose, subglobose, 23.5 cm; bracts ovate to lanceolate or obovate, 820 mm, densely puberulent, densely parallel-veined to palmately veined, acute to acuminate. Flowers
subsessile. Calyx puberulent to glabrous; hypanthium portion
subglobose to obconic, 1.52 mm; limb divided to base; lobes

narrowly triangular to linear, 612 mm, densely longitudinally

veined. Corolla yellow, tubular, outside puberulent to glabrous;
tube 910 mm; lobes triangular to ovate, 1.52 mm. Berries
subglobose, ca. 3.5 mm in diam., glabrous to puberulent, with
pedicels often elongating, to 8 mm. Fl. Aug, fr. OctDec.
Forests, often by streams; 5001500 m. Guangxi, Guizhou, Hubei, Hunan, Sichuan, Yunnan.
This name was originally published as faberii and has been
cited thus by several authors; however, current guidelines for orthography require correction to the form used here.

3. Myrioneuron nutans Wallich ex Kurz, Forest Fl. Burma 2:

55. 1877.
chui hua mi mai mu
Small shrubs, ca. 1 m tall. Petiole 0.52 cm; leaf blade
drying leathery and green, broadly elliptic or obovate, 1025(
35) 614(18) cm, adaxially glabrous, abaxially puberulent at
least on veins, base acute to attenuate, apex abruptly acuminate;
secondary veins 1120 pairs; stipules triangular-lanceolate, 10
20 mm. Inflorescences terminal or pseudoaxillary, globose, 2.5
6 cm in diam., congested-cymose or rarely laxly cymose; peduncle short; bracts ovate or lanceolate, 815 mm, acuminate.
Calyx glabrous to scabridulous; limb lobed to base; lobes linear-lanceolate, 1012 mm. Corolla white or yellow, tubular, outside puberulent; tube 56 mm; lobes ovate, ca. 1.8 mm. Berries
fleshy to dry, 47 mm in diam., glabrescent. Fl. Mar.
Forests; ca. 700 m [to only ca. 300 m in Bhutan]. Xizang (Mdog), Yunnan (Mengzi) [Bangladesh, Bhutan, N India].

4. Myrioneuron tonkinense Pitard in Lecomte, Fl. Indo-Chine

3: 193. 1923.
yue nan mi mai mu
Herbs, sometimes suffruticose, to 1(2) m tall; branches
densely puberulent, papillose, or tomentulose. Petiole 0.83 cm,
densely puberulent to tomentulose; leaf blade drying papery
and grayish green, obovate, elliptic-oblong, or elliptic, 1228
411 cm, adaxially glabrous, abaxially densely puberulent, base
acute to cuneate, apex acute or rounded then abruptly acuminate; secondary veins 918 pairs; stipules ovate to oblong, 12
25 mm, densely puberulent, densely striate veined, acute or
shortly 2-lobed. Inflorescences terminal and/or usually pseudoaxillary, subsessile, congested-cymose to subcapitate to shortly
racemiform, subglobose to ovoid, 13 cm; bracts ovate or ovatelanceolate, 1025 mm, densely puberulent, densely longitudinally veined, acute to acuminate. Flowers subsessile or with
pedicels to 3 mm. Calyx densely puberulent; hypanthium portion subglobose to ovoid, ca. 2 mm; limb divided to base; lobes
linear-subulate, 614 mm. Corolla yellow, tubular; tube 7.58

RUBIACEAE

mm; lobes 2.53 mm. Berries subglobose, 34 mm in diam. Fl.

JunAug, fr. OctDec.
Dense forests; below 1001700 m. Guangdong, Guangxi, Hainan,
Yunnan [N Vietnam].

249

H. S. Lo (in FRPS 71(1): 312. 1999) recognized two forms of this

species: f. tonkinensis [sic!] and f. longipes Lo. However, Lo failed
to provide the latter name with a Latin description or diagnosis (Vienna
Code, Art. 36.1) and an indication of a type (Art. 37.1); therefore, neither name was validly published.

59. NAUCLEA Linnaeus, Sp. Pl., ed. 2, 1: 243. 1762.

wu tan shu
Chen Tao (); Charlotte M. Taylor
Bancalus Kuntze.
Trees, unarmed; buds strongly compressed with stipules erect and pressed together [rarely subconical]. Raphides absent. Leaves
opposite, usually with domatia; stipules caducous or persistent, interpetiolar, generally elliptic or obovate. Inflorescences terminal
and sometimes also axillary, capitate with heads 15, globose, and solitary to fasciculate, many flowered, pedunculate, bracteate; peduncles articulate and often bracteate near middle. Flowers sessile, fused by their ovaries, bisexual, monomorphic. Calyx limb 4- or
5-lobed. Corolla white to yellow, funnelform to salverform, inside glabrous; lobes 4 or 5, imbricate in bud. Stamens 4 or 5, inserted
in upper part of corolla tube, exserted; filaments short; anthers basifixed. Ovary 2-celled, ovules numerous in each cell, pendulous to
horizontal on Y-shaped axile placentas attached to upper third of septum; stigma fusiform, exserted. Infructescences with fruiting
heads each comprising a globose multiple fruit (i.e., syncarp). Fruit (i.e., fruitlets, arising from one flower) baccate, fleshy or fibrous,
with calyx limb deciduous; seeds numerous, small, ovoid or ellipsoid, sometimes slightly compressed, without aril.
About ten species: tropical Africa, Asia, and Australia; one species in China.
The wood of several Nauclea species is strong and used for lumber.
The S Asian species Nauclea orientalis (Linnaeus) Linnaeus is occasionally cultivated; this can be recognized by its elliptic-oblong leaves 612
cm wide, its flowering heads 1520 mm in diam. across the calyces, and its fruiting heads to 5 cm in diam. Both N. officinalis and N. orientalis species
are encountered and occasionally confused in cultivation with the more commonly cultivated Neolamarckia cadamba, which has free (i.e., distinct or
separate) flowers and free dehiscent fruit (see p. 255).

1. Nauclea officinalis (Pierre ex Pitard) Merrill & Chun, Sunyatsenia 5: 188. 1940.
wu tan
Sarcocephalus officinalis Pierre ex Pitard in Lecomte, Fl.
Indo-Chine 3: 26. 1922.
Trees, apparently evergreen, to 12 m tall; branchlets
angled and rather flattened becoming terete, puberulent to glabrescent. Petiole 1015[20] mm; leaf blade drying papery,
elliptic to broadly elliptic or infrequently ovate or obovate, 7
11[15] 3.57[10] cm, glabrous, adaxially dark brown and
matte to shiny, abaxially pale brown and matte, base cuneate to
obtuse, apex shortly acuminate with tip slightly blunt; secondary veins 57 pairs, usually with tiny foveolate domatia in
abaxial axils; stipules caducous, obovate to elliptic, 610 mm,

glabrous, smooth, rounded. Inflorescences terminal, puberulent to glabrous; peduncles 13.5[4.5] cm, in lower part
articulate, bearing caducous bracts 24 mm; flowering heads 1
3, 56 mm in diam. across calyces, ca. 15 mm in diam. across
corollas. Calyx limb deeply lobed; lobes 5, oblanceolate to spatulate, ca. 1 mm, glabrescent, fleshy, rounded to subtruncate.
Corolla color unknown, narrowly funnelform, glabrous outside
and apparently inside; tube 34 mm; lobes 5, ligulate to elliptic,
11.5 mm, obtuse to rounded. Stigmas 1.52 mm, exserted for
ca. 3 mm. Fruiting heads (i.e., multiple fruit) yellow, 915 mm
in diam., fleshy, with surface rough (i.e., pitted with calyx limb
scars); seeds ca. 1 mm, with testa shiny black, foveolate. Fl.
summer, fr. Jul, Sep.
Forests at middle elevations. Guangdong, Guangxi, Hainan [Borneo, Cambodia, Indonesia (Sumatra), Laos, Malaysia, ?Philippines,
Thailand, Vietnam].

60. NEANOTIS W. H. Lewis, Ann. Missouri Bot. Gard. 53: 34. 1966.
xin er cao shu
Chen Tao (); Charlotte M. Taylor
Herbs, annual or perennial, unarmed, often procumbent, often fetid when bruised, often fleshy, often drying blackened. Raphides present. Leaves opposite, without domatia; stipules persistent, interpetiolar and usually fused to petioles, truncate to triangular,
laciniate to setose, sometimes glandular. Inflorescences axillary and/or terminal and sometimes displaced to pseudoaxillary, laxly
cymose to capitate, few to many flowered, pedunculate to sessile, bracteate or bracts reduced. Flowers sessile to pedicellate, bisexual,
at least sometimes distylous. Calyx limb deeply 4(or 5)-lobed. Corolla white, pink, or purple, funnelform to tubular, inside glabrous
or villous in tube and/or throat; lobes 4(or 5), valvate in bud. Stamens 4(or 5), inserted usually in corolla throat, exserted or included;
filaments short to developed; anthers dorsifixed near base. Ovary 2[4]-celled, ovules several or rarely numerous or 1 in each cell on
ascending axile placenta attached to septum near base; stigmas 2(4), linear, included or exserted. Fruit capsular or rarely

RUBIACEAE

250

indehiscent, subglobose, turbinate, obconic, ovoid, or dicoccous, often laterally compressed, loculicidally dehiscent through apical
portion, this portion plane or sometimes shortly raised into a beak, leathery to membranous, with calyx limb persistent; seeds few to
numerous, small, disciform to plano-convex, rounded or rarely winged, scabrous foveolate; endosperm corneous; embryo clavate.
About 30 species: mainly in tropical Asia and Australia; eight species (two endemic) in China.
When proposing this generic name, Lewis showed that Anotis Candolle, as then circumscribed, included a broadly heterogeneous group of both
New and Old World species that shared only a herbaceous habit and fruit with few peltate seeds under an illegitimate name. Accordingly he transferred the Asian species to Neanotis, without descriptions or keys, although this name has sometimes been overlooked. The fruit and seed morphology
of several species was studied by Terrell and Robinson (J. Bot. Res. Inst. Texas 1(1): 373384. 2007), including N. calycina and N. hirsuta of our
flora. The description here of the placenta attachment position and the seeds of other species are all from W. C. Ko (in FRPS 71(1): 7786. 1999); no
other authors seen have described these features. The floral biology of Neanotis has not been described in the literature but the flowers appear to be
distylous in at least some Chinese species.
The treatment here differs from that of W. C. Ko (loc. cit.) in the circumscription of several species. In particular, the application of the name
Neanotis hirsuta is applied more narrowly, and many specimens previously included there are here treated as N. kwangtungensis, including one variety named in N. hirsuta.
Some plants from Sichuan that are shorter than 40 cm tall with leaf blades narrowly ovate and 1.52.5 0.71 cm have been called Neanotis
ingrata f. parvifolia How ex W. C. Ko (J. S. China Agric. Univ. 16(4): 46. 1995); no specimens of this have been seen, and no more information was
provided in the protologue. Due to the limited available information and the narrowed circumscription of species here, the identity of this name is not
clear, and it is neither accepted nor synonymized here.

1a. Inflorescences axillary at various nodes all below stem apex, flowers solitary to few, subsessile to shortly
fasciculate; at least older stems prostrate and regularly rooting at nodes .......................................................... 1. N. boerhaavioides
1b. Inflorescences terminal and/or axillary at least at some uppermost nodes, flowers solitary to numerous,
sessile to pedicellate and/or pedunculate; stems prostrate and rooting at nodes to erect.
2a. At least some flowers and fruit pedicellate with pedicels 210 mm.
3a. Flowers mixed pedicellate and subsessile, with pedicels of various lengths; corollas tubular, with
tube longer than lobes, tube ca. 2 mm and lobes ca. 1 mm .............................................................................. 2. N. calycina
3b. Flowers all pedicellate, with most pedicels well developed; corollas campanulate to rotate, with
tube shorter than lobes, tube 11.5 mm and lobes 2.53.5 mm ................................................................ 7. N. thwaitesiana
2b. Flowers and fruit subsessile to sessile in small glomerules or mixed sessile and pedicellate, glomerules
sometimes separated by developed inflorescence axes, pedicels when present to 1.5 mm.
4a. Plants mostly procumbent; inflorescences capitate or branched to 1 order, sessile or on peduncles
to 1 cm, flowers in heads or congested cymes; corolla with tube 12.5 mm and lobes 0.22 mm.
5a. Leaves 16.5 0.52 cm, with 39 pairs of secondary veins; corolla with tube 11.5 mm
and lobes 1.52 mm; fruit ca. 3 4 mm ........................................................................................ 6. N. kwangtungensis
5b. Leaves 0.52.5 0.31.8 cm, with 2 or 3 pairs of secondary veins; corolla with tube
1.32.5 mm and lobes 0.21.5 mm; fruit 1.52 1.52.5 mm .............................................................. 8. N. wightiana
4b. Plants weak to procumbent or erect; inflorescences congested to laxly cymose, branched for 24
orders, on peduncles 0.84.5 cm, flowers pedicellate or sessile in heads or glomerules; corolla
with tube 3.56 mm and lobes 2.23.2 mm.
6a. Leaves 411.5 14 cm; stipules with setae or lobes 3 to numerous per side, 315 mm with
at least 1 of them longer than 5 mm; corolla tube 46 mm ......................................................................... 5. N. ingrata
6b. Leaves 15.5 14 cm; stipules with setae or lobes 37 per side, 0.55 mm; corolla tube
3.55.5 mm.
7a. Flowers some or all pedicellate, pedicels 0.51.5 mm; Taiwan ..................................................... 3. N. formosana
7b. Flowers all sessile; widespread .............................................................................................................. 4. N. hirsuta
1. Neanotis boerhaavioides (Hance) W. H. Lewis, Ann. Missouri Bot. Gard. 53: 37. 1966.
juan mao xin er cao
Hedyotis boerhaavioides Hance, J. Bot. 8: 73. 1870;
Anotis boerhaavioides (Hance) Maximowicz.
Herbs, annual or perhaps perennial, procumbent, fleshy,
regularly rooting at nodes; stems subterete to 4-angled and
sulcate, moderately to densely pilosulous to hirtellous and/or
hirsute in lines. Leaves subsessile or petiolate; petiole to 3 mm,
pilosulous or hirtellous to glabrous; blade drying papery, ovate,
lanceolate, ovate-orbicular, or lanceolate-elliptic, 12 0.41.5

cm, adaxially hirtellous, pilosulous, and/or hispidulous near

margins, abaxially hirtellous to pilosulous, base cuneate to
rounded or truncate, apex obtuse to acute; secondary veins 2 or
3 pairs; stipules triangular to rounded, 13 mm, hirtellous or pilosulous, lobe or bristle 1, 22.5 mm, ciliate, sometimes with 2
lateral bristles to 1.5 mm. Inflorescences axillary at middle and
lower stem nodes, fasciculate, (1 or)25-flowered, hirtellous;
bracts linear, 0.52 mm; peduncles 0.53.5 mm. Calyx moderately to densely pilosulous to hirtellous; hypanthium portion
subglobose to obconic, 0.81 mm; limb divided essentially to
base; lobes 4, narrowly triangular, 1.63 mm, ciliolate, acute.
Corolla white to pale blue, shortly funnelform, outside glabrous; tube 23 mm, apparently pilose in throat; lobes narrowly

RUBIACEAE

ligulate to narrowly triangular, 34 mm. Capsule compressed

globose to subglobose, 1.52 mm, hirtellous, smooth to weakly
ridged. Fl. AprAug, fr. JulAug.
Sparse forests on mountain slopes at middle elevations; 100
600 m. Fujian, Guangdong, Jiangxi, Zhejiang.
This species appears to be distylous (long-styled, Zhang Shaoyao
5681, short-styled Zhang Shaoyao 2416, both MO!). It is very similar to
Hedyotis chrysotricha, which has calyx lobes ca. 2 mm and the corolla
lobes 2.53 mm.

251

pseudoaxillary, cymose, lax with flowers in groups of 25,

branched to 2 or 3 orders, glabrous; peduncle 0.82.5 cm;
bracts laciniate or stipuliform, 0.52 mm; pedicels 0.51.5 mm.
Calyx glabrous; hypanthium portion subcupuliform to obconic;
limb divided to base; lobes triangular, 1.52 mm, acute. Corolla white, funnelform, outside glabrous; tube 45.5 mm, puberulent-papillose to perhaps pubescent inside; lobes triangular,
33.2 mm. Capsule compressed globose, 22.5 33.5 mm,
glabrous. Fl. AprJul, fr. JunJul.
Mountain slopes, roadsides; 11001700 m. Taiwan [Japan, Malay-

2. Neanotis calycina (Wallich ex J. D. Hooker) W. H. Lewis,

Ann. Missouri Bot. Gard. 53: 37. 1966.

sia].

zi hua xin er cao

4. Neanotis hirsuta (Linnaeus f.) W. H. Lewis, Ann. Missouri

Bot. Gard. 53: 38. 1966.

Anotis calycina Wallich ex J. D. Hooker, Fl. Brit. India 3:

73. 1880.

bao ye xin er cao

Herbs, erect to ascending, annual or perhaps perennial;

stems 4-angled to subterete or sulcate, glabrous or sparsely puberulent. Leaves subsessile or petiolate; petiole to 3 mm, glabrous; blade drying papery, lanceolate to ovate-lanceolate or
lanceolate-elliptic, 13.5 0.51.5 cm, adaxially sparsely scaberulous to puberulent, abaxially glabrous or puberulent to scaberulous on principal veins, base cuneate to acute, margins ciliate
to scaberulous, apex acute to acuminate; secondary veins 3 or 4
pairs; stipules broadly triangular, 13 mm, glabrescent, subentire or with 1 or 3 bristles to 1 mm, entire or ciliate. Inflorescences axillary and/or terminal, 1-flowered or cymose, several
flowered, and dichotomous, glabrous; peduncles 0.52.5 cm;
pedicels to 7 mm. Flowers sessile to pedicellate. Calyx glabrous; hypanthium portion obconic, ca. 0.8 mm; limb divided
essentially to base; lobes triangular, 0.51.3 mm, entire to ciliolate, acute. Corolla white, pale pink, or pale purple, tubular to
tubular-funnelform, outside glabrous; tube 1.52 mm, apparently glabrous in throat; lobes triangular to lanceolate, 0.51
mm, obtuse to acute. Capsule compressed globose, 1.52 ca.
3 mm, glabrous. Fl. SepOct.
Streamsides, slightly shady mountain slopes, valleys; 1100
1700 m. Yunnan [Bhutan, India (Darjeeling), Nepal].
R. R. Mill in Fl. Bhutan (2(2): 770. 1999) described this as annual,
but it is keyed there (p. 767) as a perennial.

3. Neanotis formosana (Hayata) W. H. Lewis, Ann. Missouri

Bot. Gard. 53: 38. 1966.
tai wan xin er cao
Anotis formosana Hayata, Icon. Pl. Formosan. 9: 54. 1920.
Herbs, perhaps perennial, stems decumbent near base and
ascending in upper parts, to 30 cm; stems terete to angled and
sometimes sulcate, glabrous. Leaves subsessile or petiolate;
petiole to 3 mm, sparsely hirtellous to glabrous; blade drying
papery or membranous, ovate, ovate-oblong, or lanceolate, 13
0.51.6 cm, adaxially sparsely to moderately hirtellous, scaberulous, or hispid, abaxially glabrous or sparsely to moderately
hirtellous along principal veins, base obtuse to subrounded,
apex acute; secondary veins 3 or 4 pairs; stipules triangular, 12
mm, hirtellous, erose to pectinate or with 35 bristles or lobes
15 mm, sometimes glandular. Inflorescences terminal or rarely

Oldenlandia hirsuta Linnaeus f., Suppl. Pl. 127. 1782;

Anotis hirsuta (Linnaeus f.) Boerlage; Hedyotis hirsuta (Linnaeus f.) Smith (1811), not Lamarck (1789); H. stipulata R.
Brown ex J. D. Hooker, nom. illeg. superfl.; O. japonica Miquel.
Herbs, perennial, lower stems decumbent and upper stems
apparently ascending; stems terete to 4-angled, glabrous to hirtellous in lines or throughout, angles sometimes thickened.
Leaves petiolate; petiole 15 mm, glabrous to hirtellous or hispid; blade drying papery, ovate, lanceolate, or elliptic, 15.5
0.52 cm, adaxially glabrous or sparsely to moderately strigillose, scaberulous, or hirtellous, abaxially densely hirtellous or
glabrous except sparsely strigillose or puberulent on principal
veins, base rounded to cuneate, apex acute; secondary veins 35
pairs; stipules rounded to triangular, 12 mm, glabrous to
hirtellous, laciniate to setose, lobes or bristles 37, 0.55 mm,
glabrous to ciliate. Inflorescences terminal and/or pseudoaxillary, laxly cymose, branched to 2 or 3 orders often asymmetrically, with flowers borne separately or in glomerules of 25,
glabrous or hirtellous; peduncle 0.83 cm; bracts laciniate or
stipuliform, 0.52 mm. Flowers sessile or subsessile, floral biology unknown. Calyx glabrous to densely hirtellous; hypanthium portion obconic, ca. 1 mm; limb divided essentially to base;
lobes linear-lanceolate, 12 mm. Corolla white, funnelform,
outside glabrous; tube 3.54 mm, puberulent or perhaps pubescent inside; lobes ovate to triangular, 2.23 mm. Capsule compressed globose, 22.5 2.53 mm, glabrous to hirtellous. Fl.
and fr. JunOct.
Wet sites at streamsides or in forests; 5001000(1500) m.
Guangdong, Hainan, Jiangsu, Jiangxi, Taiwan, Yunnan, Zhejiang [?Bhutan, Cambodia, India, Japan, Korea, Laos, Myanmar, Nepal, Pakistan,
Thailand, Vietnam].
The application of this name is problematic, and it seems to have
been used in the literature and the herbarium for different species in
different regions (e.g., cf. Fl. Bhutan 2(2): 768. 1999; Fl. Japan 3a: 218
219. 1993). The confusion probably started with Hookers circumscription of this species (Fl. Brit. India 3: 63. 1880), which included
three other names in synonymy to circumscribe morphologically highly
varied group of plants from India through Java and Japan. Here Neanotis hirsuta is circumscribed more narrowly. The varieties of N. hirsuta
recognized by W. C. Ko (in FRPS 71(1): 8485. 1999) are not completely distinct morphologically and fall within N. kwangtungensis as
circumscribed here.

252

RUBIACEAE

5. Neanotis ingrata (Wallich ex J. D. Hooker) W. H. Lewis,

Ann. Missouri Bot. Gard. 53: 39. 1966.
chou wei xin er cao
Anotis ingrata Wallich ex J. D. Hooker, Fl. Brit. India 3:
71. 1880.
Herbs, perennial, erect to procumbent, to 1 m tall; stems
terete to flattened or ridged, often sulcate, glabrous or sometimes sparsely hirtellous to puberulent near nodes. Leaves sessile or petiolate; petiole to 12 mm; blade drying papery, lanceolate, elliptic, ovate-lanceolate, or rarely ovate, 411.5 14 cm,
adaxially glabrous or sparsely to moderately strigillose, pilosulous, pilose, or hispid, abaxially glabrous except usually densely
puberulent, hirtellous, or pilosulous on principal veins, base
obtuse to acute, apex acute to acuminate; secondary veins 49
pairs; stipules rounded to broadly triangular, 12.5 mm, puberulent, hirtellous, or glabrescent, with 3 to numerous setae or linear lobes 315 mm, glabrous to ciliolate, often inserted below
top of sheath. Inflorescences terminal, subterminal, or pseudoaxillary, cymose, lax, dichasial with axes often asymmetrical,
branched to 24 orders, glabrous; peduncle 14.5 cm; bracts
stipuliform to reduced and erose or ciliate, 0.21.5 mm; pedicels to 0.5 mm. Calyx glabrous; hypanthium portion obconic,
0.81 mm; limb divided to base; lobes narrowly triangular, linear, or narrowly ligulate, 1.22 mm, entire to ciliolate. Corolla
white, funnelform, outside glabrous; tube 46 mm, apparently
pubescent inside; lobes narrowly triangular to ligulate, 2.53
mm, often puberulent adaxially. Capsule compressed globose,
ca. 2 3 mm, slightly dicoccous, glabrous, with calyx lobes
often elongating, to 4 mm, with pedicels sometimes elongating,
to 2 mm. Fl. JunSep, fr. Jul.
Grassy slopes on riverbanks, forests on mountain slopes; 500
1500 m. Fujian, Guizhou, Hubei, Hunan, Jiangsu, Sichuan, Xizang,
Yunnan, Zhejiang [Bhutan, India, Nepal].
Several specimens from Sichuan have been suggested (in herb.) to
belong to Neanotis urophylla (Wallich ex Wight & Arnott) W. H. Lewis
(Ann. Missouri Bot. Gard. 53: 40. 1966; Hedyotis urophylla Wallich ex
Wight & Arnott, Prodr. Fl. Ind. Orient. 404. 1834), but they are provisionally included here pending further study, including clarification of
the identity of N. urophylla (W. P. Fang 2148, 2182, 3061, all P!).
The identity of the name Neanotis mairei (H. Lveill) Lauener
& D. K. Ferguson (Notes Roy. Bot. Gard. Edinburgh 32: 110. 1972;
Ophiorrhiza mairei H. Lveill, Repert. Spec. Nov. Regni Veg. 13: 177.
1914, from Yunnan) is not entirely clear. This name was found to belong
to Neanotis after Lewiss work and was said by Lauener to be similar to
N. urophylla and N. ingrata. The characters used by Lauener to distinguish N. mairei from N. ingrata fall within the circumscription of N.
ingrata here, so this name is probably a synonym; however, until authentic material is seen this cannot be conclusively synonymized.

6. Neanotis kwangtungensis (Merrill & F. P. Metcalf) W. H.

Lewis, Ann. Missouri Bot. Gard. 53: 39. 1966.
guang dong xin er cao
Anotis kwangtungensis Merrill & F. P. Metcalf, Lingnan
Sci. J. 16: 177. 1937; Hedyotis lindleyana Hooker ex Wight &
Arnott f. glabricalycina (Honda) S. S. Ying; H. lindleyana var.

glabricalycina (Honda) H. Hara; Neanotis hirsuta (Linnaeus f.)

W. H. Lewis var. glabricalycina (Honda) W. H. Lewis; Oldenlandia hirsuta Linnaeus f. var. glabricalycina Honda.
Herbs, perhaps perennial, procumbent or ascending in upper parts; stems subterete to angled or compressed, smooth to
ridged and/or sulcate, glabrous. Leaves subsessile or petiolate;
petiole to 10 mm, glabrous; blade elliptic, lanceolate, or ovate,
15 0.52 cm, glabrous or sparsely strigillose or scaberulous
adaxially at least near margins, abaxially glabrous or sometimes
puberulent on principal veins, base acute to obtuse, apex
acuminate to acute; secondary veins 39 pairs; stipules sheath
rounded to broadly triangular, 0.81.5 mm, glabrous to puberulent, with 27 bristles or linear lobes 0.33 mm, glabrous, often
glandular. Inflorescences terminal and/or pseudoaxillary at upper nodes on principal and short axillary stems, capitate to congested-cymose, 1 to usually several flowered, glabrous, sessile
or peduncle to 3 mm; bracts reduced, stipuliform, to 0.5 mm;
pedicels to 2 mm. Flowers subsessile to pedicellate. Calyx glabrous; hypanthium portion cupuliform to turbinate, 0.81.2
mm; limb lobed to base; lobes triangular or ligulate, 1.52.5
mm. Corolla white, shortly tubular to rotate, outside glabrous or
puberulent; tube 11.5 mm, apparently glabrous in throat; lobes
narrowly triangular-oblong to ligulate, 1.52 mm. Capsule compressed subglobose to obovoid, ca. 3 4 mm, slightly dicoccous, glabrous, smooth. Fl. and fr. JulOct.
Forests on gentle slopes or at streamsides; 200800 m. Guangdong, Guangxi, Jiangxi, Sichuan, Taiwan [Japan, Thailand].
This name is applied here more broadly than by previous authors;
many of the plants that now belong to this species were formerly included in a more broadly circumscribed Neanotis hirsuta. The varieties
of N. hirsuta recognized by W. C. Ko (in FRPS 71(1): 8485. 1999) are
not completely distinct from each other and belong to N. kwangtungensis as circumscribed here. The name Hedyotis lindleyana as used for
specimens from Japan appears to be a synonym of N. kwangtungensis
rather than of N. hirsuta, and the range of this species is here extended
to include Japan; N. kwangtungensis as treated here is apparently equivalent to N. hirsuta of the Fl. Japan (3a: 218219. 1993). The name
Hedyotis kwangtungensis (Merrill & Metcalf) Ko is annotated on
some specimens but does not appear to have been published.

7. Neanotis thwaitesiana (Hance) W. H. Lewis, Ann. Missouri

Bot. Gard. 53: 40. 1966.
xin er cao
Hedyotis thwaitesiana Hance, J. Bot. 6: 298. 1868; Anotis
thwaitesiana (Hance) Maximowicz.
Herbs, apparently perennial, weak to procumbent in lower
parts and ascending in upper parts of stems; stems terete to 4angled with angles often thickened, glabrous. Leaves sessile or
petiolate; petiole to 1.5 mm, puberulent to glabrous; blade
drying papery, ovate or ovate-lanceolate, 0.82.2 0.31.5 cm,
adaxially puberulent or scaberulous at least on midrib and margins, abaxially glabrous, base cuneate to rounded, apex obtuse
to acute; secondary veins 2 or 3 pairs or not evident; stipules
triangular to ligulate, 12 mm, puberulent to hirtellous or glabrous, erose or with 13 irregular lobes or bristles 0.53.5 mm,
often glandular. Inflorescences pseudoaxillary at upper stem
nodes and/or occasionally terminal, laxly cymose, several flow-

RUBIACEAE

ered, branched to 13 orders, glabrous; peduncles slender, 1.5

3 cm; bracts linear-lanceolate or stipuliform, 13 mm; pedicels
slender, 210 mm. Calyx glabrous; hypanthium portion cupular
to obconic, ca. 1 mm; limb lobed to base; lobes triangular, 1.2
2 mm, entire. Corolla white or pale red, campanulate to rotate,
outside glabrous; tube 11.5 mm, apparently pilosulous in
throat; lobes ligulate to narrowly triangular, 2.53.5 mm. Capsules compressed globose to compressed turbinate, ca. 2 3
mm, glabrous. Fl. FebMay, fr. May.
Wastelands or streamsides in valleys. Guangdong.
This species is clearly distylous. W. C. Ko (in FRPS 71(1): 82.
1999) described the corollas as soft hairy outside, but these features
have not been seen on the specimens studied.

8. Neanotis wightiana (Wallich ex Wight & Arnott) W. H.

Lewis, Ann. Missouri Bot. Gard. 53: 40. 1966.
xi nan xin er cao

253

(1.8) cm, adaxially sparsely to moderately puberulent, scaberulous, or scabrous at least on midrib and margins, abaxially
glabrous except hirtellous on midrib and sometimes principal
veins, base broadly cuneate to subrounded, apex acute to obtuse; secondary veins 2 or 3 pairs; stipules rounded to triangular, ca. 1 mm, hirtellous or pilosulous, erose to pectinate or
with 17 bristles or linear lobes 0.22 mm, often glandular.
Inflorescences capitate to congested-cymose, terminal or pseudoaxillary at upper nodes on principal stems and short lateral
stems, several flowered, sessile or with peduncle to 1 cm; bracts
reduced. Flowers sessile or subsessile. Calyx glabrous to puberulent; hypanthium portion obconic to turbinate, ca. 1 mm;
limb divided to base; lobes triangular, 11.5 mm, entire to ciliolate. Corolla white or rarely pale red, tubular to tubular-funnelform, outside glabrous; tube 1.32.5 mm, inside glabrous;
lobes 0.21.5 mm, obtuse to acute. Capsule compressed globose, 1.52 1.52.5 mm, often markedly dicoccous, glabrous,
smooth. Fl. MayJul, fr. JunOct.

Hedyotis wightiana Wallich ex Wight & Arnott, Prodr. Fl.

Ind. Orient. 1: 410. 1834; Anotis wightiana (Wallich ex Wight
& Arnott) J. D. Hooker.

Grassy slopes, roadsides, banks at streamsides; 9001900 m.

Guangxi (Damiao Shan), Guizhou, Sichuan, Yunnan (Pingbian) [Bhutan, India, Vietnam].

Herbs, perennial, mostly procumbent and rooting at nodes;

stems angled to subterete, sometimes sulcate, glabrous to hirtellous or hirsute at least near nodes. Leaves sessile or petiolate;
petiole to 4 mm, hirtellous to glabrescent; blade drying papery
to subleathery, ovate to lanceolate, 0.51.8(2.5) 0.31.2

The inflorescences were described by W. C. Ko (in FRPS 71(1):

84. 1999) as usually having two leaflike bracts, but here these structures
are considered leaves subtending the inflorescence, similarly to the
morphological interpretation by Mill (Fl. Bhutan 2(2): 770771. 1999);
these leaves are small when the flowers form but enlarge to the size of
the other vegetative leaves as the fruit develop.

61. NEOHYMENOPOGON Bennet, Indian Forester 107: 436. 1981.

shi ding xiang shu
Chen Tao (); Charlotte M. Taylor
Hymenopogon Wallich in Roxburgh, Fl. Ind. 2: 156. 1824, not Hymenopogum P. Beauvois (1804) [Musci].
Shrubs, usually epiphytic, unarmed, often deciduous, often rather succulent. Raphides presumably absent. Leaves opposite,
apparently without domatia; stipules persistent, interpetiolar or shortly united around stem, generally ovate, acute to rounded. Inflorescences terminal, corymbose-cymose, many flowered, pedunculate, bracteate; bracts subtending 2 or more cymes per inflorescence
enlarged, petaloid, stipitate (i.e., similar to a calycophyll). Flowers pedicellate, bisexual, monomorphic. Calyx limb 5-lobed. Corolla
white to pale green, salverform or salverform-funnelform with tube prolonged, inside reflexed villous in throat and on lobes; lobes 5,
valvate in bud. Stamens 5, inserted below corolla throat, included; filaments short; anthers dorsifixed, shortly bifid at base. Ovary 2celled, ovules numerous in each cell on peltate axile placentas; stigmas 2, linear, partially exserted to included. Fruit capsular,
oblong-ellipsoid, obovoid, or turbinate, apically prolonged into short beak, septicidally dehiscent through beak or sometimes splitting
deeply into 2 valves, papery to slightly woody, with calyx limb persistent; seeds numerous, medium-sized, fusiform, acute to caudate
at each end, with hilum lateral; testa membranous; endosperm rich; embryo minute; cotyledon ovate; radicle short.
About three species: Bhutan, China, India, Myanmar, Nepal, Thailand, Vietnam; two species (one endemic) in China.
The lack of raphides has not been specifically noted but is presumed here based on the classification of this genus in Cinchoneae in FRPS
(71(1): x. 1999). The enlarged petaloid bracts of the inflorescences resemble the calycophylls of many other Rubiaceae species, but in Neohymenopogon many of these structures are inserted below the base of the hypanthium and are thus actually considered bracts. Puff et al. (Rubiaceae of
Thailand, 172. 2005) noted that N. parasiticus grows in a variety of seasonal to evergreen epiphytic and epilithic [micro]habitats, that, not surprisingly,
it is very variable morphologically, probably in correlation with habitat, and that the petaloid bracts persist on the fruit and appear to function in seed
dispersal as well as in pollination. The length of the corollas of N. parasiticus for example is notably variable, by 300%, but there seems to be
continuous variation and no clearly separable subgroups. Raizada and Bennet (Indian Forester 107: 432437. 1981) noted that the name Hymenopogon, long used for these plants, was a later homonym of a moss genus and published a new name for the genus; their article contained no information about the plants apart from a summary of general geographic ranges, which were not entirely correct even then. The specific epithets of these
species have sometimes been spelled as parasiticum and oligocarpum, but the -us ending is correct (Vienna Code, Art. 62.2(a)).

1a. Leaves elliptic-oblong, oblanceolate, or elliptic, with apex acuminate, with secondary lateral veins to 11 pairs
and well separated, i.e., 816 mm apart at midrib; fruit pilosulous to glabrous ..................................................... 1. N. oligocarpus

RUBIACEAE

254

1b. Leaves elliptic-obovate, lanceolate, oblanceolate, or obovate, with apex obtuse to acute, with secondary veins
1528 pairs and closely set, i.e., 511 mm apart at midrib; fruit villosulous to pilosulous or strigillose ................. 2. N. parasiticus
1. Neohymenopogon oligocarpus (H. L. Li) Bennet, Indian
Forester 107: 436. 1981.

157. 1824; H. parasiticus var. longiflorus F. C. How ex W. C.

Chen.

shu guo shi ding xiang

Small shrubs, 0.32 m tall; branches stout, villosulous to

hirtellous sometimes becoming glabrescent with age. Leaves
sometimes grouped at apex of shortened branches; petiole 0.4
2 cm, villosulous to hirsutulous; blade drying papery or membranous, often grayish black, elliptic-obovate, lanceolate, oblanceolate, or obovate, 525 1.511 cm, adaxially puberulent to
hirtellous, strigillose, or glabrescent, abaxially strigillose to pilosulous at least on principal veins, base obtuse to acute, apex
obtuse, acute, or rarely acuminate; secondary veins 1528 pairs;
stipules ovate to suborbicular, 612 mm, strigillose to glabrous,
cuspidate or obtuse to rounded. Inflorescences tomentulose to
villosulous or villous, sessile and 3- or 5-partite, 418 424
cm (including petaloid bracts); bracts stipuliform, ovate, 515
mm, acuminate to 2-lobed, petaloid bracts white to cream, with
blade portion drying papery to stiffly papery, elliptic-oblong to
elliptic, 310 1.53.3 cm, strigillose to pilosulous or glabrescent, obtuse to acute, with stipe 2.54 cm; pedicels 0.81.2
cm. Calyx densely villosulous to tomentulose; ovary portion
obconic, ca. 3 mm; limb deeply lobed, strigillose to glabrous;
lobes lanceolate to narrowly triangular, 610 mm, acute. Corolla white [to pale green in Thailand], outside crisped villosulous
to strigillose or strigose; tube 2560 mm; lobes ovate-oblong to
ovate, 510 mm, acute to obtuse. Capsules ellipsoid-oblong to
ellipsoid, smooth or longitudinally weakly ridged, 1.53 0.6
1 cm, villosulous to pilosulous or strigillose; seeds 56 mm. Fl.
JunAug, fr. SepDec.

Hymenopogon oligocarpus H. L. Li, J. Arnold Arbor. 25:

316. 1944.
Shrubs, ca. 2 m tall; branches flattened to subterete, glabrous. Leaves often crowded at stem apices; petiole 0.63 cm,
pilosulous to strigillose; blade drying membranous, green adaxially, often whitened abaxially, elliptic-oblong, oblanceolate,
or elliptic, 1021 36 cm, strigillose to pilosulous on both
surfaces with pubescence denser on principal veins, base cuneate to obtuse, apex slightly to markedly acuminate; secondary
veins 711 pairs; stipules ovate, 35 mm, perhaps strigillose.
Inflorescences strigillose to pilosulous; peduncle 15 cm;
branched portion ca. 2 3 cm (not including petaloid bracts);
bracts triangular, ca. 2 mm, petaloid bracts with blade portion
elliptic-oblong to lanceolate, 23 0.51 cm, strigillose to pilosulous, with 3 principal veins, with stipe 13 cm; pedicels
0.51 cm. Flowers unknown. Capsules ellipsoid, ca. 1 0.5 cm,
glabrous to pilosulous, with persistent calyx lobes lanceolate,
ca. 5 mm, acute; seeds black, ca. 6 mm. Fr. Aug.
Forests on mountains; ca. 2400 m. W Yunnan.
The only description of the flowers was given by W. C. Chen (in
FRPS 71(1): 233. 1999), who listed the corolla as green-white and gave
the flowering period as Aug. Chen distinguished this species in part by
its glabrous fruit, but the accompanying illustration (p. 232, t. 57, f. 4)
shows pilosulous fruit, as added to the description here.

2. Neohymenopogon parasiticus (Wallich) Bennet, Indian

Forester 107: 436. 1981.
shi ding xiang
Hymenopogon parasiticus Wallich in Roxburgh, Fl. Ind. 2:

On trees or rocks in thickets or forests in valleys; 12002700 m.

Xizang, Yunnan [Bhutan, N India, Myanmar, Nepal, Thailand, Vietnam].
Hymenopogon parasiticus var. longiflorus was originally distinguished by its corolla length (57 cm vs. 2.54 cm in var. parasiticus). W. C. Chen (in FRPS 71(1): 233. 1999) synonymized these
with some commentary.

62. NEOLAMARCKIA Bosser, Bull. Mus. Natl. Hist. Nat., B, Adansonia 6: 247. 1985.
tuan hua shu
Chen Tao (); Charlotte M. Taylor
Trees, unarmed; buds conical. Raphides absent. Leaves opposite, usually with domatia; stipules caducous, interpetiolar, triangular. Inflorescences terminal, capitate with heads globose and solitary, many flowered, pedunculate, bracteate. Flowers sessile,
bisexual, monomorphic. Calyx limb 5-lobed; lobes sometime spatulate. Corolla yellow to white, salverform to funnelform, glabrous
inside; lobes 5, imbricate in bud. Stamens 5, inserted in upper part of corolla tube, partially exserted; filaments short; anthers basifixed. Ovary 2-celled in basal portion, 24-celled in upper portion, ovules several in each cell on axile, simple or 2-forked placentas
attached to upper third of septum; stigma cylindrical or fusiform, at apex bifid, exserted. Fruiting head with central axis becoming
markedly enlarged and fleshy to fibrous. Fruit drupaceous, indehiscent or tardily dehiscent into 4 indehiscent segments or perhaps
dehiscent valves, ellipsoid to cylindrical or obconic, with exocarp somewhat fleshy to membranous or papery, with endocarp cartilaginous to bony, with calyx limb persistent; seeds several, small, compressed, fusiform to angled, with testa membranous; endosperm fleshy; embryo small, cylindrical.
Two species: S and SE Asia, Australia, and New Guinea, one species occasionally cultivated for lumber worldwide; one species in China.
The flowers are tightly packed on the thickened axis of the inflorescence heads but not actually fused to each other as has sometimes been
(incorrectly) suggested.
The name of this species and its genus have long been confused. Ridsdale (Blumea 24: 307366. 1979) in his revision of the Naucleeae treated

RUBIACEAE

255

this species as Anthocephalus chinensis (Lamarck) A. Richard ex Walpers, as done by several other authors; however, the nomenclature here follows
the conclusions of Bosser (loc. cit.: 243248; Adansonia 21: 9395. 1999) and Razafimandimbison (Tribal Delimit. Naucleeae (Ph.D. Diss.): 7075.
2000).

1. Neolamarckia cadamba (Roxburgh) Bosser, Bull. Mus.

Natl. Hist. Nat., B, Adansonia 6: 247. 1985.
tuan hua
Nauclea cadamba Roxburgh, Fl. Ind. 2: 121. 1824; Anthocephalus cadamba (Roxburgh) Miquel; A. indicus A. Richard;
A. indicus var. glabrescens H. L. Li; Sarcocephalus cadamba
(Roxburgh) Kurz.
Trees, deciduous, to 30 m tall; trunk with small buttresses;
bark thin, grayish brown, fissured and scabrous when old;
branches horizontally spreading, flattened becoming subterete,
glabrescent. Petiole 2035 mm, glabrous; leaf blade drying
thinly leathery, elliptic or oblong-elliptic, on juvenile growth
5060 1530 cm, on adult growth 1525 712 cm, adaxially shiny and glabrous, abaxially glabrous to densely puberulent, base shallowly cordate on juvenile growth, rounded or

truncate on adult growth, apex acute; secondary veins 812

pairs, apparently without domatia; stipules lanceolate, 1220
mm, acute. Inflorescences with peduncle 24 cm, rather stout;
flowering heads 3545 mm across calyces, 4060 mm across
corollas. Calyx puberulent to pilosulous; ovary portion ellipsoid
to obovoid, ca. 1.5 mm; limb 34 mm, partially to deeply lobed;
lobes oblong to spatulate, obtuse to rounded. Corolla yellowish
white, funnelform, outside glabrous; tube ca. 10 mm; lobes lanceolate, ca. 2.5 mm. Fruiting heads yellowish green, 3040 mm
in diam., with peduncles markedly thickened. Fruit cylindrical
to ellipsoid or obovoid, 22.5 ca. 1 mm, glabrous; seeds 3angled, 0.50.7 mm. Fl. and fr. JunNov.
Broad-leaved forests, streamsides in valleys. Guangdong, Guangxi, Yunnan [Bhutan, India, Malaysia, Myanmar, Sri Lanka, Thailand,
Vietnam].
This species is occasionally cultivated for lumber in Asia, including probably in China, and in the Neotropics.

63. NEONAUCLEA Merrill, J. Wash. Acad. Sci. 5: 538. 1915.

xin wu tan shu
Chen Tao (); Charlotte M. Taylor
Nauclea Korthals, Observ. Naucl. Indic. 17. 1839, not Linnaeus (1762).
Trees or shrubs, unarmed; buds flattened with stipules erect and pressed together [to conical], [sometimes with fusiform
swellings housing ants in branches]. Raphides absent. Leaves opposite, sometimes with numerous foveolate domatia; stipules caducous [or rarely persistent], interpetiolar, generally ligulate, entire. Inflorescences terminal, capitate with 19 globose heads in fascicles or cymes, many flowered, pedunculate, bracteate; peduncles and axes articulate near middle; bracts enclosing heads involucrate and caducous; bracteoles spatulate to conical or sometimes absent. Flowers sessile, bisexual, monomorphic. Calyx limb deeply
5-lobed; lobes prolonged into a slender shaft bearing thickened to rhomboidal apical portion, this often ornamented and/or pubescent
differently from shaft and frequently deciduous before shaft. Corolla pale green to white or red, salverform to narrowly funnelform,
inside glabrous or glabrescent; lobes 5[or rarely 6], imbricate in bud. Stamens 5, inserted in corolla throat, partially to fully exserted;
filaments short; anthers basifixed. Ovary 2-celled, ovules numerous in each cell, pendulous on axile placentas attached in upper third
of septum; stigma globose to obovoid, exserted. Fruiting heads globose. Fruit capsular, obconic, septicidally then loculicidally
dehiscent into 4 valves from base to apex, with valves separating from septum, with septum persistent on inflorescence or later deciduous, woody to cartilaginous, with calyx limb deciduous with capsule valves; seeds numerous, small, fusiform, flattened, winged;
embryo ellipsoid, somewhat bilaterally compressed.
About 62 species: tropical Asia and Pacific islands, perhaps ten species in continental Asia; four species (one endemic) in China.
This genus was studied in some detail by Ridsdale (Blumea 24: 337342. 1979; Blumea 34: 177275. 1989). The unusual, usually quite elaborately developed calyx lobes are distinctive of the genus in general, and their form is distinctive for individual species; care must be taken to look for
the caducous apical part of the calyx lobes and to not confuse the persistent tubular calyx limb base with the entire calyx limb. Ridsdale (loc. cit. 1989)
developed a very detailed terminology, but did not suggest a function, for the calyx lobe morphology of this genus. He also noted that Neonauclea has
secondary pollen presentation. Ridsdale (loc. cit. 1989) and Zou (J. Arnold Arbor. 69: 7376. 1988) noted that some Neonauclea species are large trees
that produce very good lumber.

1a. Leaves sessile or subsessile, with petioles to 5 mm; leaf base obtuse to rounded, truncate, or cordulate.
2a. Leaf blade elliptic-oblong to elliptic or suborbicular, with 69 pairs of secondary veins, with domatia in
abaxial axils of secondary, tertiary, and often quaternary veins; corolla tube 56 mm ...................................... 2. N. sessilifolia
2b. Leaf blade obovate, broadly obovate, broadly elliptic, or elliptic-oblong, with 79 pairs of secondary veins,
abaxially with domatia only in axils of secondary veins; corolla tube 810 mm .................................................. 3. N. truncata
1b. Leaves petiolate with petioles 840 mm; leaf base acute to cuneate or broadly obtuse, sometimes decurrent.
3a. Petioles 820 mm; leaf base acute to cuneate and sometimes decurrent; domatia glabrous; calyx lobes
densely pilosulous on middle and lower portion and glabrous at apex, with apical portion fusiform ................... 1. N. griffithii
3b. Petioles 1540 mm; leaf base broadly obtuse to cuneate; domatia tomentulose or glabrous; calyx lobes
puberulent throughout except glabrous adaxially in basal portion, with apical portion spatulate to cucullate ....... 4. N. tsaiana

256

RUBIACEAE

1. Neonauclea griffithii (J. D. Hooker) Merrill, J. Wash. Acad.

Sci. 5: 540. 1915.
xin wu tan
Adina griffithii J. D. Hooker, Fl. Brit. India 3: 24. 1880;
Cephalanthus navillei H. Lveill; Nauclea griffithii (J. D.
Hooker) Haviland; Neonauclea navillei (H. Lveill) Rehder.
Trees, evergreen, to 20 m tall; trunk often buttressed at
base, sometimes with aerial roots; bark greenish gray, fissured
and cracked, sometimes warty, with inner bark yellow, pale
brown to pink; branchlets quadrangular becoming terete, grayish
brown, rugose or striate, white lenticellate, glabrescent. Petiole
820 mm, stout, glabrous; leaf blade drying thickly papery to
subleathery, obovate to elliptic, 822 415 cm, both surfaces
glabrous, base acute to cuneate and frequently decurrent, apex
rounded then abruptly cuspidate or acute; secondary veins 57
pairs, in abaxial axils with glabrous domatia; stipules obovate to
obovate-oblong, 510 38 mm, weakly keeled, glabrous,
apex obtuse. Inflorescences puberulent to glabrescent; peduncles 1 or 3, 26 cm; flowering heads solitary, 812 mm in diam.
across calyces, 2530 mm in diam. across corollas; involucral
bracts not seen; bracteoles 0.51 mm. Calyx with ovary portion
obconic, 11.5 mm, glabrous in lower 2/3, pilosulous in upper
1/3; limb partially lobed, with basal tubular portion ca. 1 mm;
lobes with shaft clavate, 3.54.5 mm, densely pilosulous, with
apical portion fusiform, deciduous before corollas open, pilosulous on lower portions, glabrous at apex. Corolla red (Henry
12676, MO!), narrowly funnelform to salverform, outside glabrous; tube 78 mm, inside glabrous to sparsely pubescent;
lobes elliptic-oblong, 23 mm, acute. Stigmas subglobose, ca. 1
mm, exserted for 56 mm. Fruiting heads ca. 20 mm in diam.
Capsules obconic, 56 mm, pilosulous at least at apex; seeds
not seen.
Dense forests in valleys or on humid slopes; 8001300 m. Guangxi, Guizhou, Yunnan [Bhutan, N India, Myanmar].

2. Neonauclea sessilifolia (Roxburgh) Merrill, J. Wash. Acad.

Sci. 5: 542. 1915.
wu bing xin wu tan
Nauclea sessilifolia Roxburgh, Fl. Ind. 2: 124. 1824;
Adina sessilifolia (Roxburgh) J. D. Hooker ex Brandis; N.
sericea Wallich ex G. Don.
Trees, leaf persistence unknown, 730 m tall; bark dark
gray, transversely fissured and cracked, with inner bark brown,
sometimes mottled; branches flattened to angled, glabrous.
Leaves sessile or subsessile; petiole to 5 mm, stout, glabrous;
blade drying papery to thinly leathery, elliptic to elliptic-oblong
or suborbicular, 530 315 cm, both surfaces glabrous, base
rounded to truncate or cordulate, apex obtuse; secondary veins
69 pairs, with small, glabrous to pilosulous, foveolate domatia
in abaxial axils of lateral, tertiary, and often quaternary veins;
stipules broadly elliptic to obovate, 1030 510 mm, glabrous, keeled in lower portion, apex obtuse to broadly rounded.
Inflorescences glabrescent; peduncles 1(or 3), 13(8) cm; flowering heads solitary, 2025 mm in diam. across calyces, 3545
mm in diam. across corollas; involucral bracts not seen; bracteoles linear, 12(3) mm. Calyx with ovary portion obconic,

11.5 mm, glabrous or apically tomentulose; limb partially

lobed, with basal tubular portion 0.50.8 mm; lobes densely
pilosulous, with shaft clavate, 57 mm, with apical portion pyramidal, deciduous before corollas open. Corolla color unknown;
tube 56 mm, inside glabrescent; lobes deltoid to elliptic, 23
mm, outside densely strigillose to sericeous, acute. Stigma subglobose, ca. 0.8 mm, exserted for 56 mm. Fruiting heads (15)
2535 mm in diam. Capsules obconic, 810 mm, glabrous except densely pilosulous at apex; seeds ca. 2 mm. Fl. Oct.
Thickets or broad-leaved forests on hills; 500800 m. Taiwan,
Yunnan [Cambodia, India, Laos, Myanmar, Thailand, Vietnam].
This is the first report of this species in Taiwan.

3. Neonauclea truncata (Hayata) Yamamoto, J. Soc. Trop.

Agric. 7: 149. 1935.
tai wan xin wu tan
Nauclea truncata Hayata, J. Coll. Sci. Imp. Univ. Tokyo
30(1): 140. 1911.
Trees, evergreen, large, height not noted; branches flattened to angled, gray, glabrous. Leaves sessile or subsessile;
petiole to 2 mm, glabrous; blade drying leathery, obovate,
broadly obovate, broadly elliptic, or elliptic-oblong, 13.526
9.919.8 cm, glabrous, base rounded to truncate, obtuse, or cordulate, apex obtuse to acute or shortly acuminate; secondary
veins 79 pairs, in abaxial axils with pilosulous or foveolate
domatia; stipules elliptic-oblong to elliptic, 1225 815 mm,
glabrous, smooth to weakly keeled, rounded to obtuse. Inflorescences densely strigillose to glabrescent; peduncles 13(5),
2.23.5 cm, stout; flowering heads solitary on peduncles, 1225
mm in diam. across calyces, 3445 mm in diam. across corollas; bracteoles reportedly sparse, not seen. Calyx with ovary
portion 0.81.5 mm, glabrous; limb partially lobed, with basal
tubular portion 11.5 mm, pilosulous at least on apical portion;
lobes deciduous before corollas open, with shaft linear, 1.52.5
mm, glabrous below and densely pilosulous in upper part, with
apical portion obconic to thickly fusiform, 22.5 mm, pilosulous to glabrescent. Corolla white, funnelform; tube 810 mm,
glabrous; lobes ligulate to lanceolate, 23 mm, puberulent, obtuse to acute. Stigma subglobose to fusiform, ca. 1 mm, exserted for 810 mm. Fruiting heads 3035 mm in diam. Capsules obconic, 810 mm, glabrescent. Fl. Jul.
Forests, on coral rocks. Taiwan [Philippines].
This species was reported from Taiwan by Chun (Fl. Taiwan 4:
313. 1978) and later by Liu et al. (Fl. Taiwan, ed. 2, 4: 304306. 1998)
under the name Neonauclea reticulata (Haviland) Merrill, with the
name N. truncata placed in synonymy there. However, Yamamoto (loc.
cit.) had already noted the differences that separate N. truncata and N.
reticulata and concluded that the name N. reticulata had been incorrectly applied to the Taiwanese plants. Ridsdale (Blumea 34: 213217.
1989) later recognized these as two distinct species and reported an
extended range for N. truncata, in N Philippines.

4. Neonauclea tsaiana S. Q. Zou, J. Arnold Arbor. 69: 73.

1988.
dian nan xin wu tan
Trees, to 3040 m tall and 1 m d.b.h.; trunk cylindrical,

RUBIACEAE

with buttresses; bark rough, with inner bark fibrous, yellow or

sometimes with pink; branches flattened to angled, densely
lenticellate, glabrescent. Petiole 1240 mm, stout, glabrous; leaf
blade drying leathery, elliptic or ovate-elliptic, 1222 613
cm, adaxially glabrous and rather shiny, abaxially glabrous and
matte, base broadly obtuse to cuneate, apex acute to shortly
acuminate; secondary veins 6 or 8 pairs, with tomentulose to
glabrous foveolate domatia in abaxial axils; stipules ovate, 12
24 814 mm, glabrous, weakly keeled in basal portion, apex
obtuse. Inflorescences glabrous; peduncles 13, 24.5 cm;
flowering heads 17(or 9), solitary on peduncles or 37 in
branched cymes, 1018 mm in diam. across calyces, 2530 mm
in diam. across corollas; involucral bracts not seen; bracteoles
conical, 0.71 mm. Calyx with ovary portion obconic, 11.5
mm, glabrous except puberulent at apex; limb partially lobed,
with basal tubular portion 0.81 mm; lobes deciduous before

257

corollas open, with shaft clavate, 34 mm, densely puberulent

except adaxially glabrous in basal portion, apical portion
rounded to cucullate, densely puberulent. Corolla pale yellow,
funnelform, glabrous; tube 56.5 mm; lobes elliptic-oblong, ca.
2.5 1.2 mm, apex acute. Stigmas subglobose, ca. 1 mm, exserted for 56 mm. Fruiting heads 1520 mm in diam. Capsules clavate, somewhat flattened, 67 mm, glabrous in basal
portion, puberulent or pilosulous at apex; seeds unknown. Fl.
SepOct, fr. MayJun.
Tropical rain forests at streamsides or in bottom of valleys; 500
1100 m. Yunnan.
This species was described almost simultaneously with the publication of Ridsdales revision of the genus (Blumea 34: 177275. 1989)
and was not mentioned by him presumably because he was unaware of
it.

64. NERTERA Banks ex Gaertner, Fruct. Sem. Pl. 1: 124. 1788, nom. cons.
bao zhu cao shu
Chen Tao (); Charlotte M. Taylor
Erythrodanum Thouars; Gomozia Mutis ex Linnaeus f.
Herbs, perennial, unarmed, sometimes fetid when bruised, often creeping and rooting at nodes. Raphides present. Leaves opposite, without domatia, marginally usually thickened and sometimes crisped; stipules persistent, interpetiolar and fused to petioles,
triangular or bidentate. Inflorescences terminal and/or pseudoaxillary, 1-flowered, sessile or shortly pedunculate, ebracteate or sometimes with small involucre of bracts fused in pairs (i.e., calyculate) or of reduced stipules and leaves. Flowers bisexual, homostylous.
Calyx limb truncate, 4-lobed, or reduced. Corolla greenish white, white, or pink, funnelform, glabrous inside; lobes 5, valvate in bud.
Stamens 4, inserted near base of corolla tube, exserted; filaments developed; anthers basifixed. Ovary 2- or 4-celled, ovules 1 in each
cell on axile placentas; stigmas 2 or 4, linear, exserted. Fruit orange, red, or black, drupaceous, ovoid or globose, fleshy, with calyx
limb persistent; pyrenes 2 or 4, 1-celled, each with 1 seed, plano-convex, cartilaginous; seeds medium-sized, ellipsoid to plano-convex; testa membranous; endosperm scanty; cotyledons leaflike; hypocotyl hypogenous.
About six species: Antilles, Australia, Central, North, and South America, China, Indonesia, Malaysia, Pacific islands, Papua New Guinea, Philippines, Subantarctic islands (Tristan da Cunha), Vietnam; three species (one endemic) in China.
Phylogeography of the most widespread species Nertera granadensis (as N. depressa) was studied by Jakubowksy et al. (Evolution of Nertera.
Poster presented at XVII IBC. 2005) using molecular data; they concluded that this species originated in New Zealand, where Nertera has its center of
diversity, and dispersed independently to Australia, the Philippines, then SE Asia and Hawaii, and then Central and South America and eastward. They
also suggested that N. nigricarpa may be better included within the circumscription N. granadensis, although species identity and circumscription
were not the primary focus of their work so their sampling many not be adequate to address this. Nertera nigricarpa is distinguished primarily by its
black rather than red mature fruit and was synonymized with N. granadensis by Liu and Yang (Fl. Taiwan, ed. 2, 4: 306. 1998), without comment;
however, these species were separated by W. C. Ko in FRPS (71(2): 162165. 1999). If these populations are treated as conspecific, this represents the
only example known in Nertera of such fruit color dimorphism, which is known but uncommon in Rubiaceae. Nertera nigricarpa is provisionally
separated here pending further study.
Liu and Yang (loc. cit.) and W. C. Ko (loc. cit.) described the flowers as bisexual or unisexual, but other authors reported the flowers of Nertera
to be bisexual (e.g., Fosberg, Acta Phytotax. Geobot. 33: 7383. 1982; Andersson, Fl. Ecuador 47: 1112. 1993).

1a. Leaf blade oblong-lanceolate to elliptic, cuneate to acute at base and apex, abaxially with surface uneven due to
swollen cells, on dry specimens appearing scurfy; fruit with 4 pyrenes ........................................................................ 3. N. sinensis
1b. Leaf blade ovate, broadly ovate, ovate-triangular, or ovate-reniform, acute to obtuse or broadly rounded at apex,
abaxially generally smooth; fruit with 2 pyrenes.
2a. Leaves acute to obtuse at apex; fruit red ........................................................................................................... 1. N. granadensis
2b. Leaves obtuse to broadly rounded at apex; fruit black ....................................................................................... 2. N. nigricarpa
1. Nertera granadensis (Mutis ex Linnaeus f.) Druce, Rep.
Bot. Soc. Exch. Club Brit. Isles 1916: 637. 1917.

129. 1782; Nertera depressa Banks & Solander ex Gaertner; N.

taiwaniana Masamune.

hong guo bao zhu cao

Creeping herbs; stems angled, glabrescent. Petiole slender,

24 mm, glabrescent; leaf blade drying papery and often pale
abaxially, ovate or ovate-triangular, 0.31 0.20.8 cm, gla-

Gomozia granadensis Mutis ex Linnaeus f., Suppl. Pl.

RUBIACEAE

258

brescent, generally smooth abaxially, base obtuse to shallowly

cordate, apex acute to obtuse; secondary veins 2 or 3 pairs; stipules ovate-triangular, 0.51 mm, glabrescent, apex acute and
often glandular. Flowers sessile or subsessile. Calyx glabrous;
ovary portion ellipsoid, ca. 1 mm; limb reduced. Corolla white
to pale green, glabrous; tube 0.51 mm; lobes ca. 1 mm. Drupes
red, subglobose, 35 mm in diam.; pyrenes 2.
Hillsides at middle elevations. Taiwan [Indonesia, Malaysia, Papua New Guinea, Philippines; Australia, Central America (Costa Rica,
El Salvador, Guatemala, Honduras, Nicaragua, Panama), North America
(Mexico), Pacific islands (Hawaii, New Zealand), South America
(Argentina, Bolivia, Chile, Colombia, Ecuador, Peru, Venezuela), Subantarctic islands (Tristan da Cunha)].
See comments under the genus regarding the distinction of this
species from Nertera nigricarpa. No specimens have been seen from
China during the preparation of this treatment that correspond to N.
granadensis, separately from N. nigricarpa; thus, its presence in China
remains to be confirmed and the placement of the name N. taiwaniana
remains to be confirmed. Puff et al. (Rubiaceae of Thailand, 210. 2005)
reported that N. granadensis has been recorded for Thailand, but they
found no specimens nor the source of that report.
This species has often been called Nertera depressa, the type of
the genus; however, Gomozia granadensis is an older name for this
species and thus has priority over the epithet depressa, so the correct
name is N. granadensis.

2. Nertera nigricarpa Hayata, J. Coll. Sci. Imp. Univ. Tokyo

25(19): 115. 1908.
hei guo bao zhu cao
Creeping herbs; stems quadrate, glabrous. Petiole 18 mm,
glabrous; leaf blade drying papery, broadly ovate or ovatereniform, 0.251 0.31.1 cm, glabrous, generally smooth
abaxially, base rounded to truncate or cordulate then abruptly

attenuate, apex obtuse to broadly rounded; secondary veins 2 or

3; stipules broadly triangular, 11.3 mm, glabrous, acute and
often glandular. Flowers sessile. Calyx glabrous; ovary portion
ovoid, ca. 1 mm; limb truncate to denticulate, 0.10.2 mm. Corolla probably pale green, urceolate-funnelform, glabrous; tube
ca. 1 mm; lobes triangular, ca. 0.4 mm. Drupes black, ellipsoid
to subglobose, 44.5 in diam., glabrous; pyrenes 2. Fl. FebJul,
fr. MarJan.
Sparse forests, open fields; 9002500 m. Fujian, Taiwan [Vietnam
(Averyanov et al. VH 427, MO!)].
See comments under the genus description regarding the relationship of this species to Nertera granadensis. This is apparently the first
report of this species from Vietnam.

3. Nertera sinensis Hemsley, J. Linn. Soc., Bot. 23: 391. 1888.

bao zhu cao
Low herbs, with main stems creeping and reproductive
stems erect, to 10 cm tall; stems angled, glabrous. Petiole 13
mm, glabrous; leaf blade drying papery, oblong-lanceolate to
elliptic, 0.71.6 0.350.5 cm, glabrous or sparsely hispidulous on both surfaces, abaxially rough due to enlarged thinwalled cells that dry with scurfy appearance, base cuneate to
acute, margins hispidulous, apex acute to cuneate; secondary
veins not visible; stipules narrowly triangular, 0.81 mm, obtuse to acute or aristate and usually glandular. Flowers sessile.
Calyx glabrous; ovary portion 11.5 mm; limb truncate, 0.10.2
mm. Corolla pale green, funnelform, glabrous; tube ca. 1 mm;
lobes triangular, ca. 0.5 mm, acute to obtuse. Drupes dark blue
to black, subglobose, 26 mm in diam.; pyrenes 4. Fl. JulAug,
fr. JulNov.
Mountain slopes, roadsides, ditch sides, rocks at riversides;
5001300 m. Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi,
Sichuan, Yunnan.

65. OPHIORRHIZA Linnaeus, Sp. Pl. 1: 150. 1753.

she gen cao shu
Chen Tao (); Charlotte M. Taylor
Hayataella Masamune; Mitreola Boehmer; Mungos Adanson.
Annual or perennial herbs or rarely subshrubs, unarmed, often fleshy or rather succulent, sometimes creeping. Raphides present.
Leaves opposite, decussate, sometimes anisophyllous, without domatia; margins sometimes undulate to denticulate; stipules persistent or caducous, interpetiolar, entire or bifid to fimbriate, sometimes glandular. Inflorescences terminal and/or pseudoaxillary or
rarely axillary (Ophiorrhiza oppositifolia), cymose to capitate, fasciculate, or paniculiform with axes often helicoid, few to many
flowered, sessile to pedunculate, bracteate or bracts absent; bracts caducous to persistent, sometimes involucral. Flowers pedicellate
to sessile, bisexual and distylous or monomorphic or occasionally cleistogamous. Calyx with ovary portion turbinate to obconical,
usually strongly compressed, longitudinally often 5- or 10-ribbed; limb reduced or 5(or 6)-lobed, lobed essentially to base or rarely
subtruncate (O. repandicalyx). Corolla white, yellow, orange, pink, purple, or brown, sometimes drying with markedly different
color, usually notably clavate in bud, at anthesis salverform, tubular, or funnelform with tube often swollen to gibbous at base, outside often longitudinally ridged or winged, inside glabrous to variously pubescent; lobes 5(or 6), valvate in bud, smooth or occasionally ridged, winged, and/or with hornlike appendages near apex, apex sometimes adaxially rostrate. Stamens 5(or 6), inserted
near throat to below middle of corolla tube, included or exserted; filaments reduced to well developed; anthers dorsifixed near middle
or base. Ovary 2-celled, ovules numerous in each cell on axile placentas attached from middle to base of septum; stigmas 2, linear to
subcapitate, included or exserted. Fruit capsular, obovoid to oblate, mitriform, or obcordate, strongly laterally compressed perpendicular to septum, sometimes with apical portion prolonged into a beak, loculicidally dehiscent across top and sometimes along
sides, papery, with calyx limb persistent; seeds numerous, small, angled to rhomboid, areolate to alveolate.

RUBIACEAE

259

About 200300 species: tropical and subtropical Asia, Australia, New Guinea, Pacific islands; 70 species (49 endemic, one of unconfirmed
occurrence) in China.
Ophiorrhiza is a notably species-rich, taxonomically complicated genus found in wet tropical forests of SE Asia (Darwin, Lyonia 1(2): 47102.
1976); it has been little studied and is particularly poorly known in SE Asia (I. Schanzer, pers. comm.). Ophiorrhiza was studied for China by H. S. Lo
(Bull. Bot. Res., Harbin 10(2): 182. 1990), who variously described 44 of the 68 Ophiorrhiza species recognized by Lo in FRPS (71(1): 110174.
1999). The genus was studied in India by Deb and Mondal (Bull. Bot. Surv. India 39(14): 1148. 1997), who recognized 47 species there. Darwin
(loc. cit.) reported that the plants are notably variable in many vegetative features, many of which were shown by him not to be informative to separate
species. H. S. Lo (loc. cit. 1999: 111) reported that the calyx and corolla lobes are occasionally 6, but this has not been noted by other authors nor seen
on specimens studied; this may be an occasional variation found in one or two flowers on unusual plants, as in many Rubiaceae species. H. S. Lo also
described the placentas as ascending from the septum base; however, Darwin (loc. cit.: 56) reported that in the Pacific Ophiorrhiza species the
placenta is inserted in the middle of the septum in the flower and then often becomes displaced to near the base of the septum in fruit, whereas Puff et
al. (Rubiaceae of Thailand, 190. 2005) gave the insertion of the placenta as being in the lower half of the septum. Puff et al. reported that the fruit
function as splash cups for seed dispersal and observed that, regardless of the orientation of the flowers, the fruit become erect with enlarged and
strengthened pedicels. Tan and Rao (Biotropica 13: 232233. 1981) reported vivipary in a species of Ophiorrhiza growing in Singapore, with the
seeds germinating within the capsules, pushing their cotyledons out through the suture where the valves open normally, and extending roots through
the capsule locules and tissues; a similar condition may be found in Chinese Ophiorrhiza.
Darwin (loc. cit.: 47102) noted that the presence of distyly in Ophiorrhiza has been controversial because the arrangement of the stigmas and
anthers in the first dimorphic species found here differs from that of classic distyly, with strictly reciprocal sizes and positions. However, since then the
recognition of distyly in Rubiaceae has expanded to include species that are at least a bit dimorphic and have intra-morph incompatibility, and
Ophiorrhiza clearly belongs in this group and has been considered distylous by subsequent authors (Deb & Mondal, loc. cit.; Kudoh et al., J. Trop.
Ecol. 17: 719728. 2001; Schanzer, Thai Forest Bull. 33: 140166. 2004). Also, some species with markedly dimorphic distylous flowers have
subsequently been discovered (e.g., O. aureolina and O. rufopunctata). Deb and Mondal (loc. cit.: 15, f. 7) illustrated some of the variation in stamen
and stigma position and internal corolla pubescence in distylous species of this genus. Homostyly has also been confirmed in the genus (Nakamura et
al., J. Jap. Bot. 81: 113120. 2006; J. Plant Res. 120: 501509. 2007) and some species also appear to be autogamous (Nakamura et al., loc. cit. 2006).
Schanzer (loc. cit.) noted that some species appear to vary in floral biology across their range, to include both homostylous and distylous populations;
this situation has been found elsewhere in some distylous Rubiaceae, which have variation in expression of distyly (e.g., Faivre & McDade, Amer. J.
Bot. 88: 841853. 2001), though in those cases, the flowers resemble one of the distylous forms while Schanzer described a distinct floral form in the
monomorphic plants. Schanzer also noted that some of these floral forms may be aberrant and cleistogamous rather than distylous. Nakamura et al.
(loc. cit. 2007) studied two supposedly conspecific varieties of O. japonica in Japan, one homostylous and the other distylous, and concluded that the
self-compatible homostylous plants differed in ploidy level, comprised a distinct lineage according to cpDNA sequences, and are better considered a
separate species. They also noted that floral biology is not correlated with ploidy in general in Ophiorrhiza. Observations and documentation of the
floral biology of Chinese Ophiorrhiza species are so far limited. Kudoh et al. (loc. cit.) presented a detailed analysis of the floral forms and possible
genetic controls of these in O. napoensis in Guangxi, China. Also notable in the floral morphology of this genus is the apparent wide variation in
corolla pubescence within a species, sometimes with the long-styled and short-styled flowers reportedly different (e.g., O. oppositiflora, floral forms
similar but corolla pubescence variable, Deb & Mondal, loc. cit.: 88, f. 39; O. austroyunnanensis, pubescence apparently correlated with floral form,
H. S. Lo, loc. cit. 1990: 31, f. 8).
Deb and Mondal (loc. cit.: 1) noted that the genus name alludes to the presumed healing properties of the root of these plants for snakebite and
that Ophiorrhiza mungos and O. japonica are used for such in the Indian subcontinent. They also noted that species of this genus are used as medicine
(for snakebites, stomach ulcers, skin eruptions, rheumatism, heart diseases), dye (red, for wool and hair), and food (the fruit), and list several
references detailing their ethnobotany and medical chemistry.
The monotypic genus Hayataella, endemic to Taiwan, was recognized by several authors (particularly Taiwanese authors) as distinct from
Ophiorrhiza, though it was synonymized by H. S. Lo (Bull. Bot. Res., Harbin 18: 276277. 1998). Its morphology and molecular systematics were
studied by Nakamura et al. (J. Plant Res. 119: 657661. 2006), who concluded based on molecular data that the species belongs to a relatively derived
clade of Ophiorrhiza and formally transferred the species, eliminating another of Taiwans few endemic genera.
No infrageneric classification has been recognized by recent authors (Darwin, loc. cit.: 47102; Deb & Mondal, loc. cit.: 1148).
H. S. Lo (loc. cit. 1990: 182) presented the definitive work on this genus in China. Most recently, Ophiorrhiza has been studied in China by
Duan and Lin (Acta Phytotax. Sin. 45: 870879. 2007), who synonymized several of Los species. Their species circumscription is relatively broad
compared to that of Lo, and a few of the species they synonymized are provisionally recognized here pending further study and a broader, consistent
review of Ophiorrhiza in China.
Overall, the treatment of Ophiorrhiza here is primarily an organization of the published information, rather than a revisionary work. A few other
species are keyed here even though their descriptions are incomplete; their placement is based in part on the key of H. S. Lo in FRPS (loc. cit. 1999:
112117). In the FRPS treatment, H. S. Lo described in some detail the arrangement and degree of surface development and visibility of the tertiary
venation on the abaxial leaf surface of many Ophiorrhiza species; however, this is incompletely described for the Chinese species, is variable within
species, and was not used by Lo to distinguish species nor by other authors and, therefore, is not detailed here. Details of the anthers and stigmas are
also incompletely described for Chinese Ophiorrhiza and mostly not used to separate species; the details available are summarized in the comments
following the species description.
Among the names published in Ophiorrhiza by H. S. Lo (loc. cit. 1990), eight lacked an acceptable indication of type and were therefore not
validly published under Art. 37 of the Vienna Code. In one case (O. chingii), two gatherings were cited but neither was indicated as the type, and in the
other seven cases only one gathering was cited, which under Art. 37.3 is acceptable as indication of the type, but under Art. 37.6, on or after 1 January

260

RUBIACEAE

1990, indication of the type must also include the word typus or holotypus or an equivalent in a modern language, and Lo did not include such
words. All but one of the eight names were validated by S. Y. Jin and Y. L. Chen (Cat. Type Spec. Herb. China (Suppl.), 189191. 1999) in each case
by reference to Los Latin description and by indication of a single gathering as the type (as T.). The one remaining name, O. longicornis, is validated here.

1a. Calyx lobes relatively well developed, longest ones 3.5 mm or longer (do not confuse bracts with calyx lobes).
2a. Calyx lobes strongly unequal, more than 50% different in length, longer ones 36 mm and shorter ones
0.83 mm .......................................................................................................................................................... 52. O. pingbienensis
2b. Calyx lobes subequal, at most 50% different in length, 3.58 mm.
3a. Corolla purple, tube 2629 mm; stipules 1.52 mm ...................................................................................... 64. O. sichuanensis
3b. Corolla white or yellow, tube 1319 mm (mature corolla unknown in O. hunanica); stipules 310 mm.
4a. Stems glabrescent; calyx lobes linear; corolla externally with ciliate wings .................................................. 27. O. hunanica
4b. Stems pilosulous or villous at least when young; calyx lobes linear, spatulate, or ovate; corolla
externally smooth to ridged or with narrow, glabrous to pubescent wings.
5a. Stipules ca. 10 mm, 2- or 3-lobed, lobes linear; bracts 1020 mm; calyx lobes linear ........................... 41. O. medogensis
5b. Stipules 39 mm, triangular or 2-lobed; bracts 57 mm; calyx lobes ligulate, narrowly spatulate,
ovate, or lanceolate.
6a. Corolla with tube 1314 mm and externally ridged or ribbed, lobes 22.5 mm; leaves in subequal
pairs .......................................................................................................................................................... 15. O. ensiformis
6b. Corolla with tube 1819 mm and externally ridged to winged, lobes ca. 4.5 mm; leaves in unequal
pairs ............................................................................................................................................................... 60. O. rufipilis
1b. Calyx lobes none (i.e., limb truncate) or longest lobes 3.4 mm or shorter (calyx lobe length unknown
in O. rarior).
7a. Inflorescences axillary, paired at nodes below apex ......................................................................................... 49. O. oppositiflora
7b. Inflorescences terminal and/or pseudoaxillary, solitary at each node.
8a. Stems densely lenticellate and moderately brown villous or -hirsute with multicellular trichomes
when young ............................................................................................................................................................... 56. O. rarior
8b. Stems sparsely lenticellate to smooth, glabrous to densely pubescent with trichomes of various colors
and forms when dry.
9a. Corollas relatively small, tube 5.5 mm or shorter (corollas unknown in O. salicifolia).
10a. Leaves lanceolate-linear, 6 or more as long as wide ............................................................................. 63. O. salicifolia
10b. Leaves variously shaped, 3 as long as wide or broader.
11a. Corolla lobes with well-developed dorsal horns or appendages, these 1.52 mm.
12a. Inflorescences congested-cymose, branched to 1 or 2 orders; corolla pubescent in throat but
glabrous below inside tube, pubescent externally ......................................................................................... 6. O. cana
12b. Inflorescences cymose to corymbose, branched to 3 or 4 orders; corolla pubescent below
middle inside tube, glabrous externally ............................................................................................ 35. O. longicornis
11b. Corolla lobes dorsally smooth, ribbed, winged, or with thickenings or short hornlike
protuberances, these up to 0.8 mm.
13a. Bracts and bracteoles well developed, lanceolate to spatulate or elliptic, 13 mm wide,
partially to fully enclosing buds and flowers.
14a. Stipules ovate ....................................................................................................................... 4. O. austroyunnanensis
14b. Stipules triangular at base, quickly narrowed to a linear apex or linear lobes ..................................... 38. O. lurida
13b. Bracts and bracteoles reduced or linear to triangular, 0.10.8 mm wide, not enclosing buds
or flowers.
15a. Plants creeping or at least relatively small, at most 30 cm tall and most plants much shorter.
16a. Stipules 13 mm, caducous and often not visible; fruit 57 mm wide ............................................ 53. O. pumila
16b. Stipules 410 mm, usually persisting on uppermost nodes; fruit 45 mm wide.
17a. Leaves strigillose to glabrous adaxially .................................................................................... 24. O. hispidula
17b. Leaves sparsely hispidulous adaxially .......................................................................................... 62. O. rugosa
15b. Plants mostly weakly ascending to erect, generally rather robust, most plants 30 cm tall or taller.
18a. Plants when dry covered with golden yellow pubescence, including on corollas ......................... 3. O. aureolina
18b. Plants glabrous or with pubescence drying white to brown, reddened, or clear.
19a. Plants to 80100 cm tall, with leaves 1522 610 cm, with 1119 pairs of secondary
veins; calyx limb undulate to shallowly to moderately lobed.
20a. Stipules deciduous after uppermost nodes, 48 mm; corolla tube 2.54 mm .......................... 43. O. mungos
20b. Stipules caducous, unknown; corolla tube ca. 5.5 mm .................................................... 57. O. repandicalyx
19b. Plants to 70 cm tall, with leaves 215 16 cm, with (4 or)511(19) pairs of
secondary veins; calyx limb shallowly to deeply lobed.
21a. Stipules 38 mm, deciduous after uppermost nodes; Taiwan ................................................ 29. O. kuroiwae

RUBIACEAE

261

21b. Stipules caducous and unknown, or persistent on uppermost nodes and 211 mm;
mainland (including Hainan).
22a. Inflorescences well developed, cymose to paniculate; secondary leaf veins
719 pairs.
23a. Stipules caducous; corollas 3.55 mm ............................................................................... 43. O. mungos
23b. Stipules mostly persistent; corollas 66.5 mm .......................................................... 49. O. oppositiflora
22b. Inflorescences somewhat reduced, congested-cymose to subcapitate; secondary
leaf veins (4 or)57(11) pairs.
24a. Corolla with pubescent ring inside tube, with lobes 1/41/3 as long as tube and
spreading at anthesis ............................................................................................................ 62. O. rugosa
24b. Corolla pubescent in throat and on upper part of tube but glabrous through most
of tube, with lobes 1/31/2 as long as tube and spreading to strongly reflexed
at anthesis.
25a. Stipules triangular, 24 mm; corolla tube 2.54.5 mm, lobes spreading at anthesis ....... 38. O. lurida
25b. Stipules subfiliform, ca. 6 mm; corolla tube 4.55 mm, lobes strongly reflexed
at anthesis ................................................................................................................................ 70. O. wui
9b. Corollas larger, tubes more than 5.5 mm (corollas unknown in O. hainanensis and O. salicifolia).
26a. Leaves relatively narrow, more than 3 as long as wide, often falcate, 4.511 0.62 cm.
27a. Stems with 2 discrete lines of pubescence; bracteoles 1.53 mm ............................................................. 34. O. lignosa
27b. Stems glabrescent; bracteoles 45 mm .................................................................................................. 63. O. salicifolia
26b. Leaves rather narrow to relatively broad, less than 3 as long as wide, 0.525 0.310 cm.
28a. Corolla lobes dorsally with well-developed hornlike appendages 0.82 mm.
29a. Plants robust, to 2.5 m tall, leaves 1020 47.5 cm, with 15 or 16 pairs of secondary veins ..... 44. O. mycetiifolia
29b. Plants smaller to somewhat robust, to 1 m tall, leaves 217 24.5 cm, with 613 pairs
of secondary veins.
30a. Corolla tube 2224 mm .......................................................................................................................... 25. O. howii
30b. Corolla tube 10.512 mm.
31a. Leaves with secondary veins 812 pairs; calyx lobes 0.40.5 mm ................................................ 19. O. gracilis
31b. Leaves with secondary veins 6 or 7 pairs; calyx lobes ca. 1.5 mm .................................... 55. O. purpureonervis
28b. Corolla lobes dorsally smooth, ridged, winged, and/or with dorsal thickenings on lobes, these
sometimes hornlike but 0.7 mm or shorter.
32a. Plants creeping to procumbent, with most internodes prostrate and/or most nodes rooting.
33a. Corolla with tube 1520 mm, lobes 56.5 mm.
34a. Corolla externally with 5 pubescent lines ............................................................................... 42. O. mitchelloides
34b. Corolla externally glabrous or uniformly pubescent ........................................................ 33. O. liangkwangensis
33b. Corolla with tube 712 mm, lobes 25 mm.
35a. Corolla lobes ca. 2 mm; stems glabrescent or pilosulous in lines .......................................... 30. O. kwangsiensis
35b. Corolla lobes 2.25 mm; stems generally uniformly villous, hirtellous, or pilosulous.
36a. Bracts well developed, 3.56 mm; leaf base regularly cordate ................................................... 11. O. cordata
36b. Bracts reduced, 12 mm; leaf base obtuse, truncate, or sometimes cordulate.
37a. Corolla lobes 2.53 mm ..................................................................................................... 14. O. dulongensis
37b. Corolla lobes 45 mm ................................................................................................... 26. O. huanjiangensis
32b. Plants erect to weak, with most internodes ascending and most nodes not rooting.
38a. Corolla with tube 1827 mm.
39a. Bracts and bracteoles well developed, enclosing buds and at least partially flowers,
ligulate, lanceolate, ovate, elliptic, or elliptic-oblong, 1018 mm.
40a. Stems villous; corolla funnelform at least in upper part.
41a. Corolla lobes not evidently veined, ca. 5 mm .......................................................... 20. O. grandibracteolata
41b. Corolla lobes pinnately veined, 68 mm ........................................................................... 58. O. rhodoneura
40b. Stems glabrous; corolla salverform to funnelform.
42a. Corolla funnelform, tube villous inside above middle ......................................................... 16. O. fangdingii
42b. Corolla salverform to funnelform, tube glabrous inside ....................................................... 46. O. napoensis
39b. Bracts and bracteoles reduced to developed, not enclosing or covering buds or flowers,
linear, narrowly triangular, or narrowly lanceolate, 5.5 mm or shorter.
43a. Stems villous, villosulous, hirsute, hispidulous, strigose, strigillose, or pilosulous.
44a. Calyx lobes 0.41.5 mm; flowers several to many.
45a. Peduncle 1.53.5 cm; corolla tube 1820 mm, pubescent inside ........................................ 9. O. chinensis
45b. Peduncle 11.5 cm; corolla tube 2327 mm, glabrous inside ............................................. 68. O. wallichii
44b. Calyx lobes 13 mm; flowers 15.

262

RUBIACEAE

46a. Corolla tube 2226 mm; leaves without gland dots abaxially .................................... 54. O. purpurascens
46b. Corolla tube 1822 mm; leaves with or without gland dots abaxially.
47a. Leaves without or usually with reddish gland dots abaxially; fruit 56
ca. 11 mm ................................................................................................................... 61. O. rufopunctata
47b. Leaves without gland dots abaxially; fruit ca. 3 8 mm ......................................... 69. O. wenshanensis
43b. Stems glabrous to puberulent.
48a. Leaves broadly ovate to broadly elliptic, abaxially with numerous small scales ............... 12. O. crassifolia
48b. Leaves elliptic, lanceolate, ovate, ovate-oblong, or elliptic-oblong, glabrous to
variously pubescent but without scales.
49a. Stipules generally persistent at least on uppermost nodes, 68 mm ................................ 39. O. macrantha
49b. Stipules generally caducous, unknown or perhaps reduced.
50a. Calyx lobes unequal, 0.72 mm .................................................................................... 45. O. nandanica
50b. Calyx lobes subequal, 0.41.5 mm.
51a. Fruit 1415 mm wide .................................................................................................... 68. O. wallichii
51b. Fruit 811 mm wide.
52a. Leaves 3.515 cm, with 9 or 10 pairs of secondary veins; corolla white to pale
purple-red; widespread ................................................................................................ 9. O. chinensis
52b. Leaves 915 cm, with 1014 pairs of secondary veins; corolla red to purplish
red; Xizang, Yunnan ................................................................................................. 67. O. umbricola
38b. Corolla with tube less than 18 mm (corolla unknown in O. hainanensis).
53a. Stipules generally well developed, 316 mm, and persistent at least on uppermost nodes
of flowering stems.
54a. Calyx lobes rather well developed, 1.52.5 mm with at least some longer than 1.5 mm.
55a. Leaves larger, 6.522 2.510 cm; bracts 79 mm .......................................................... 40. O. macrodonta
55b. Leaves smaller, 14 0.62.5 cm; bracts reduced, to ca. 1 mm ....................................... 14. O. dulongensis
54b. Calyx lobes smaller, 0.51.5 mm with at least some shorter than 1.5 mm.
56a. Bracts and bracteoles reduced, to 3 mm, mostly or all deciduous before anthesis.
57a. Leaves smaller, 211 15 cm; corollas pubescent inside, glabrous to pubescent
outside ..................................................................................................................................... 62. O. rugosa
57b. Leaves larger, 6.525 210 cm; corollas glabrous inside and puberulent to glabrous
outside.
58a. Corollas yellow to pale yellow; peduncles in flower 310 cm (these later elongating
in fruit) .......................................................................................................................... 48. O. ochroleuca
58b. Corollas purplish red; peduncles in flower 2.55 cm (these later elongating in fruit) .......... 59. O. rosea
56b. Bracts and bracteoles generally well developed, 312 mm, generally persistent at
least through anthesis.
59a. Leaves in markedly unequal pairs, larger ones 1.53 as long as shorter.
60a. Corolla pink or purplish red, tube ca. 11 mm, lobes ca. 1.5 mm and dorsally
with short hornlike appendage ...................................................................................... 5. O. brevidentata
60b. Corolla white, tube ca. 9 mm, lobes 23 mm and dorsally narrowly winged ..................... 47. O. nutans
59b. Leaves in subequal pairs.
61a. Leaves larger, at least some 1016 46.5 cm; peduncles in flower 18 cm.
62a. Peduncle in flower 1.52 cm; bracts 34 mm; corolla pubescent inside ....................... 23. O. hispida
62b. Peduncle in flower 18 cm; bracts 612 mm; corolla glabrous inside.
63a. Calyx with hypanthium portion 11.5 mm, lobes 11.2 mm ................................ 17. O. fasciculata
63b. Calyx with hypanthium portion ca. 3 mm, lobes ca. 0.5 mm ............................... 48. O. ochroleuca
61b. Leaves smaller, 0.813 0.54 cm; peduncles in flower 0.56 cm.
64a. Peduncle in flower 13 cm; bracts 36.5 mm; leaves obtuse at apex ....................... 50. O. pauciflora
64b. Peduncle in flower 0.56 cm; bracts 510 mm; leaves acute to cuspidate or
acuminate at apex.
65a. Plants viscid puberulent, trichomes unicellular and not drying particularly
dark; leaves with secondary veins 1517 pairs; Hainan ....................................... 21. O. hainanensis
65b. Plants villous, trichomes multicellular, drying reddish brown, not glandular;
leaves with secondary veins 915 pairs; mainland ........................................................ 47. O. nutans
53b. Stipules caducous and not seen, or 3 mm or shorter.
66a. Calyx and fruit densely tuberculate with peglike, flat- to round-topped protuberances;
Taiwan ........................................................................................................................................ 22. O. hayatana
66b. Calyx and fruit smooth, glabrous to pubescent with slender trichomes.
67a. Plants drying purple throughout (on live plants, see old and dying tissues).

RUBIACEAE

263

68a. Corolla with tube 914 mm; leaves 411 0.73.5 cm; bracts 16 mm ........................... 28. O. japonica
68b. Corolla with tube 710 mm; leaves 520 2.58 cm, at least some longer than
measurements in alternate lead; bracts 69 mm ............................................................... 66. O. succirubra
67b. Plants drying green, brown, yellowed, gray, blackened, or with parts flushed purple.
69a. Stems (but not necessarily inflorescences) villous to hispid with well-developed
spreading trichomes.
70a. Calyx lobes ca. 0.5 mm; corolla pale purple or white, with tube 1012 mm and
lobes ca. 1 mm .................................................................................................................... 23. O. hispida
70b. Calyx lobes 11.3 mm; corolla yellowish white or tinged with purple, with tube
ca. 16 mm and lobes 55.5 mm ......................................................................................... 36. O. longipes
69b. Stems glabrous or strigillose, puberulent, hispidulous, and/or villosulous with short,
appressed to spreading trichomes.
71a. Leaves relatively broad, less than 2 as long as wide, 1116 5.510 cm .............. 8. O. carnosicaulis
71b. Leaves moderately broad to rather narrow, 2 or more as long as wide,
1.520 0.77 cm.
72a. Leaves with secondary veins 1323 pairs, at least some leaves with more
than 13 pairs.
73a. Calyx puberulent; corolla lobes 1.84 mm, dorsally winged ................................. 7. O. cantonensis
73b. Calyx hispidulous; corolla lobes ca. 1.5 mm, with short hornlike appendage ............... 13. O. densa
72b. Leaves with secondary veins 415 pairs, at least some leaves with fewer
than 13 pairs.
74a. Bracteoles well developed, 712 mm ......................................................................... 2. O. alatiflora
74b. Bracteoles none, reduced, or moderately well developed, 6 mm or shorter.
75a. Bracts and bracteoles reduced or caducous, not or hardly visible.
76a. Leaves in unequal pairs, with secondary veins prominent adaxially .......... 37. O. luchuanensis
76b. Leaves in subequal pairs, with secondary veins flat to impressed
adaxially.
77a. Leaves with 5 or 6 pairs of secondary veins; stems puberulent to
glabrescent ....................................................................................................... 31. O. laevifolia
77b. Leaves with 715 pairs of secondary veins; stems glabrous, puberulent,
or hispidulous.
78a. Leaves rounded-obtuse at apex ............................................................ 65. O. subrubescens
78b. Leaves acute or acuminate at apex.
79a. Stems hispidulous ............................................................................... 49. O. oppositiflora
79b. Stems glabrous or puberulent.
80a. Corolla tube ca. 12 mm .................................................................................. 1. O. alata
80b. Corolla tube 48 mm ....................................................................... 49. O. oppositiflora
75b. Bracts and bracteoles developed, evident, persisting at least with flowers,
bracteoles 16 mm.
81a. Leaves mostly or all in markedly unequal pairs, longer ones 23 or
more as long as shorter.
82a. Leaf pairs with longer ones 3 or more as long as shorter; inflorescences
pilosulous or puberulent with slender trichomes ................................................ 10. O. chingii
82b. Leaf pairs with longer ones 22.5 as long as shorter; inflorescences
mealy puberulent ........................................................................................... 32. O. laoshanica
81b. Leaves in subequal pairs or some in unequal pairs, longer ones at most
1.5 as long as shorter.
83a. Stems and leaves abaxially mealy puberulent; corolla lobes dorsally
with short hornlike appendages .............................................................. 18. O. filibracteolata
83b. Stems and leaves abaxially glabrous to variously pubescent with
slender trichomes; corolla lobes dorsally smooth to winged and/or
with short hornlike appendages.
84a. Leaves elliptic to elliptic-oblong, 620 1.57 cm, at least some
leaves longer than measurements in alternate lead, with 915 pairs
of secondary veins, generally becoming yellowed or brownish
yellow when dry ......................................................................................... 7. O. cantonensis
84b. Leaves elliptic to lanceolate or ovate, 111 0.74 cm, at least
some leaves smaller than measurements in alternate lead, with
413 pairs of secondary veins, generally drying with green, gray,
blackish, or purplish cast.

RUBIACEAE

264

85a. Stigmas ovate to elliptic or lanceolate; widespread, common .................. 28. O. japonica
85b. Stigmas linear; known only from Yunnan, rare ...................................... 51. O. petrophila
1. Ophiorrhiza alata Craib, Pl. Siam. Enum. 2: 61. 1932.
you chi she gen cao
Herbs, ascending, to 1 m tall; stems drying compressed,
glabrous or puberulent. Leaves in subequal pairs; petiole 13
cm, subglabrous; blade drying submembranous, grayish green
adaxially, pale abaxially, ovate, elliptic, or lanceolate-ovate, 7
13 35.5 cm, glabrous on both surfaces, base subcuneate and
sometimes shortly decurrent, margins undulate, apex subacuminate; secondary veins 810 pairs; stipules caducous, not seen.
Inflorescence many flowered, pubescent; peduncle 1.55 cm;
axes helicoid, usually shorter than 1 cm; bracts reduced or absent. Flowers distylous, with pedicels ca. 2.5 mm. Calyx pubescent; hypanthium portion turbinate-campanulate, ca. 1.8 mm, 5ribbed; lobes subtriangular, ca. 0.7 mm. Corolla white, tubular
with swollen base, subglabrous outside; tube ca. 12 mm, with
white villous ring in throat; lobes triangular to broadly ovate,
ca. 1.5 mm, dorsally winged and with short horn at apex. Capsules ca. 2.5 6 mm. Fl. AprMay.
Dense forests; 500700 m. Yunnan [Thailand].
H. S. Lo (in FRPS 71(1): 173. 1999) described this species as
distylous but described only an apparently short-styled flower.

2. Ophiorrhiza alatiflora H. S. Lo, Bull. Bot. Res., Harbin

10(2): 62. 1990.
yan chi she gen cao
Herbs or subshrubs, suberect; stems stout, terete to slightly
compressed. Leaves in subequal pairs; petiole 11.5(4) cm, pilosulous; blade drying papery to thickly papery, black adaxially,
pale brown abaxially, ovate or oblong-ovate, 513 27 cm,
glabrous on both surfaces or hirtellous along principal veins
abaxially, base cuneate, rather inequilateral, margins subentire,
apex shortly acuminate or subacute; secondary veins 712 pairs;
stipules caducous, not seen. Inflorescence congested-cymose,
many flowered, densely pubescent; peduncle 14 cm; axes 0.5
4 cm, helicoid; bracteoles narrowly lanceolate, 712 mm, acute,
glabrous or sparsely ciliate. Flowers distylous, subsessile. Calyx
densely pilosulous to puberulent; hypanthium turbinate, ca. 1.5
mm, 5-ribbed; lobes subtriangular or triangular, 0.50.7 mm,
with 1 globose gland in each sinus. Corolla white striped with
purple, subtubular or slightly swollen at base, outside glabrous
and winged along entire length, inside with a ring of pubescence just below throat and/or in throat and onto lobes; tube
1215 mm; lobes subovate, 23 mm, dorsally with wing 0.81
mm wide, apex rostrate. Capsules mitriform, ca. 3 10 mm, pilosulous.
Forests. Yunnan.
The protologue described both of the floral forms in detail, noting
explicitly that the corollas are similar and the anthers and stigmas reciprocally placed, near the middle of the corolla tube and at or just below the throat, respectively. However, the protologue figure shows partially different patterns of internal corolla pubescence in the two floral
forms, with the pubescence restricted to a narrow ring well below the
throat in the long-styled flowers but located in the throat of the shortstyled flowers.

The varieties treated by H. S. Lo in FRPS (71(1): 160. 1999) are

treated here for reference. The protologue figure was not fully labeled
but apparently illustrated only var. alatiflora; these same figures were
reproduced in FRPS (p. 161, t. 41, f. 18) where they were explicitly
labeled as var. alatiflora.

1a. Leaf blade drying with thinner texture,

glabrous on both surfaces ........................ 2a. var. alatiflora
1b. Leaf blade drying with thicker texture,
crisped pubescent along principal
veins abaxially ..................................... 2b. var. trichoneura
2a. Ophiorrhiza alatiflora var. alatiflora
() yan chi she gen cao (yuan bian zhong)
Leaf blade drying with thinner texture, glabrous on both
surfaces.
Forests. Yunnan.

2b. Ophiorrhiza alatiflora var. trichoneura H. S. Lo, Bull.

Bot. Res., Harbin 10(2): 63. 1990.
mao mai she gen cao
Leaf blade drying with thicker texture, crisped pubescent
along principal veins abaxially.
Forests. Yunnan.

3. Ophiorrhiza aureolina H. S. Lo, Bull. Bot. Res., Harbin

10(2): 34. 1990.
jin huang she gen cao
Herbs, ascending or sometimes weak at base, to 30(45)
cm tall; stems terete to compressed, usually striate, densely
golden yellow pubescent. Leaves in subequal pairs; petiole 0.5
1.5 cm, densely golden yellow pubescent; blade drying papery,
yellow or grayish yellow, ovate, lanceolate, or elliptic-oblong,
1.56 0.82 cm, adaxially sparsely strigose or glabrescent to
pubescent, abaxially golden yellow pubescent to glabrescent at
least on principal veins, base cuneate to acute, apex acute or
shortly acuminate; secondary veins 710 pairs; stipules persistent on upper nodes, deeply 2-parted, lobes triangular, 38 mm,
filiform or acuminate at apex. Inflorescences cymose to somewhat congested-cymose, many flowered, golden yellow pubescent; peduncle 515 mm; axes short to well developed, helicoid; bracts none or reduced and caducous. Flowers distylous,
subsessile. Calyx densely golden yellow hispidulous or -pilosulous; hypanthium turbinate, 1.21.5 mm, 5-ribbed; lobes subtriangular, 0.50.7 mm. Corolla pale yellow, white, or sometimes
reddish, tubular to often somewhat inflated, hispidulous outside; tube 34 mm, inside with a white villous ring in upper
half; lobes subovate, 11.3 mm, apex incurved-rostrate, dorsally smooth. Capsules obcordate, ca. 2 45.5 mm, hirtellous
to pilosulous. Fl. Jul.
Forests; ca. 1800 m. Yunnan (Xishuangbanna).
H. S. Lo (loc. cit.: 3437) recognized two forms of this species,
f. aureolina and f. qiongyaensis H. S. Lo; the latter form was formally
synonymized with Ophiorrhiza pumila by Duan and Lin (Acta Phyto-

RUBIACEAE

tax. Sin. 45: 878. 2007), who are followed here. Lo described the longstyled and short-styled forms as similar in their corollas but differing in
the anthers (situated below the middle of the corolla tube in the longstyled form vs. in the throat in the short-styled form); the stigmas appear
to be reciprocal in position with the anthers.

4. Ophiorrhiza austroyunnanensis H. S. Lo, Bull. Bot. Res.,

Harbin 10(2): 30. 1990 [astroyunnanensis].
dian nan she gen cao
Herbs, generally procumbent; stems pilosulous or tomentose. Leaves in equal to somewhat unequal pairs; petiole 0.5
1.5 cm, densely pilosulous or tomentose; blade drying thinly
papery, usually grayish with veins brown abaxially, ovate, elliptic, elliptic-oblong, or lanceolate, 25(7) 0.82.2 cm, adaxially sparsely pilose, multicellular pilose along principal veins
abaxially, base cuneate to subrounded, margins sometimes
undulate, apex obtuse; secondary veins 79 pairs; stipules persistent, ovate, 57 mm, glabrescent, ciliate, long acuminate. Inflorescences congested-cymose to subcapitate, several flowered, glabrescent; peduncles 23 cm; bracts elliptic-oblong, 56
1.52 mm, obtuse. Flowers distylous, subsessile. Calyx glabrous; hypanthium subturbinate, 0.81 mm, 5-ribbed; lobes triangular, ca. 0.4 mm. Corolla tubular-salverform, outside glabrous; tube ca. 2.6 mm, with white villous ring in throat; lobes
subtriangular, ca. 1.3 mm, dorsally smooth, obtuse. Infructescences with axes expanded, lax. Capsules oblate-ellipsoid, ca.
1.5 3.5 mm, glabrous, shortly white striate. Fl. May.

265

and the stigmas positioned near the base of the corolla tube.

6. Ophiorrhiza cana H. S. Lo, Bull. Bot. Res., Harbin 10(2):

20. 1990.
hui ye she gen cao
Herbs, creeping; stems drying compressed, striate or shallowly sulcate, pilosulous. Leaves in subequal pairs; petiole 1
1.5 cm, pilosulous; blade drying thinly papery or membranouspapery, leaden gray or sometimes green adaxially, pale and
puncticulate abaxially, ovate, elliptic-oblong, or lanceolate, 3.5
11 1.84.5 cm, adaxially subglabrous or sparsely puberulent,
abaxially pilosulous along principal veins, base cuneate to rarely
subrounded, margins flat or undulate, apex acute or obtuse then
acuminate; secondary veins 1015 pairs; stipules persistent,
broadly triangular then abruptly contracted, 45 mm, caudateacuminate. Inflorescences congested-cymose, many flowered,
densely pilosulous, branched to 1 order; peduncle shorter than 1
cm; axes very short, helicoid; bracteoles quickly caducous, ca.
0.3 mm. Flowers distylous, subsessile or with pedicels to 1 mm.
Calyx with hypanthium subglobose to subturbinate, ca. 1.1
1.5 mm, 5-ribbed, hispidulous; lobes linear, 1.41.6 mm, with
line of stiff hairs. Corolla drying orange-yellow, tubular-funnelform, hirtellous outside; tube ca. 5 mm, with villous ring in
throat; lobes subovate, ca. 2 mm, acute, dorsally with horn ca.
1.5 mm. Capsules obcordate, ca. 2 5 mm, hispidulous.
SE Yunnan.

The protologue figure, reprinted in FRPS (71(1): 133, t. 32, f. 1

10. 1999), shows a remarkable dimorphism in the shape and internal pubescence of the corollas (salverform and densely barbate in the throat in
the long-styled form vs. funnelform and glabrous internally in the shortstyled form), but the protologue description said only that the corollas of
both forms are similar; thus, this figure may deserve re-checking.

The protologue described both the long-styled and short-styled

forms in detail, noting that the forms have similar calyces and corollas
but differ in their stamen and stigma position (i.e., stamens inserted in
the corolla tube ca. 0.5 mm from the base and stigmas positioned in the
throat in the long-styled form vs. stamens inserted in the throat and stigmas positioned just below the throat in the short-styled flowers). The
protologue also noted that this species is known only from the type,
which lacks locality or date.

5. Ophiorrhiza brevidentata H. S. Lo, Bull. Bot. Res., Harbin

10(2): 32. 1990.

7. Ophiorrhiza cantonensis Hance, Ann. Sci. Nat., Bot., sr. 4,

18: 222. 1862.

duan chi she gen cao

guang zhou she gen cao

Herbs, ascending; stems drying purplish red or purplish

brown, slender, terete, densely hirtellous. Leaves in markedly
unequal pairs; petiole ca. 1 cm, pilosulous; blade drying papery,
ovate, larger ones 26 cm, smaller ones 1/31/2 as large, strigose adaxially, villous abaxially at least along principal veins,
base rounded to obtuse, apex cuspidate; secondary veins 9
11(15) pairs; stipules lanceolate, 56 mm, villous, acuminate.
Inflorescences congested-cymose to subcapitate, few to many
flowered, densely pilosulous; peduncle 1.54.5 cm; axes helicoid; bracts oblong-lanceolate or elliptic-oblong, 57 mm, ciliate. Flowers with biology unknown. Calyx pilosulous; hypanthium ca. 1.2 mm; lobes subtriangular, 0.70.9 mm. Corolla
pink or purplish red, tubular, outside glabrous or in bud puberulent at apex; tube ca. 11 mm, inside villous above middle
and in throat; lobes ca. 1.5 mm, dorsally with very short horn.
Capsule ca. 2 6 mm, puberulent. Fl. AprMay, fr. Jun.

Ophiorrhiza bodinieri H. Lveill; O. japonica Blume var.

leiocarpa Handel-Mazzetti; O. longzhouensis H. S. Lo; O.
paniculiformis H. S. Lo; O. seguinii H. Lveill; O. violaceoflammea H. Lveill; O. yingtakensis Masamune.

Thickets; ca. 1500 m. Yunnan.

Streamsides in forests. Yunnan.

The flowers described in the protologue resemble the short-styled
form of distylous species, with the anthers shortly exserted in the throat

Herbs or subshrubs, weak to erect, to 1.2 m tall, often

brown to yellowish brown when dry; stems glabrous to densely
puberulent or villosulous. Leaves in subequal pairs; petiole 14
cm, glabrous to densely puberulent or villosulous; blade drying
papery to thickly papery, grayish brown or grayish green adaxially, pale green to yellowish brown abaxially, oblong-elliptic,
elliptic, ovate, ovate-oblong, or lanceolate-oblong, 620 1.5
7 cm, glabrous throughout or sometimes puberulent abaxially,
base obtuse to acute, margins entire, apex acute to acuminate;
secondary veins 715(to 18, O. paniculiformis) pairs; stipules
broadly triangular, 0.81.5 mm, puberulent to glabrous, caducous. Inflorescences paniculiform to corymbose, several to
many flowered, densely puberulent to pilosulous; peduncles
1.57 cm; axes helicoid; bracts linear, 16 mm. Flowers distylous, subsessile or pedicels to 2 mm. Calyx densely puberulent;

RUBIACEAE

266

hypanthium subglobose to turbinate, 11.3 mm, smooth to

weakly 5-ribbed; lobes triangular, 0.41 mm, sometimes with 1
gland in each sinus. Corolla white to pink, often drying yellow
or pale red, subtubular to tubular-funnelform, slightly swollen
at base, puberulent to glabrous outside, inside villous near middle of tube and scaly pubescent above middle and onto lobes;
tube 912(15) mm; lobes triangular, 1.83(4) mm, dorsally
with wing ca. 0.3 mm wide and often prolonged near apex,
apex rostrate. Capsules mitriform, 34 69 mm, densely puberulent to subglabrous. Fl. winter and spring, fr. spring and
summer.
Ravines and watersides in forests; 1001700 m. Guangdong,
Guangxi, Guizhou, Hainan, Sichuan (Pingshan), Yunnan.
Ophiorrhiza longzhouensis was synonymized formally by Duan
and Lin (Acta Phytotax. Sin. 45: 870. 2007) with O. cantonensis; H. S.
Los separation of O. longzhouensis (Bull. Bot. Res., Harbin 10(2): 79.
1990) seems to have been based on its having relatively small leaves.
Duan and Lin also synonymized O. paniculiformis with O. cantonensis.
In FRPS (71(1): 165166. 1999), H. S. Lo described this species as
distylous with the corollas of both forms apparently similar and the
reciprocally placed anthers and stigmas positioned near the middle of
the corolla tube and in its throat, respectively; Lo also described the
dried color of specimens of this species as ranging to reddish or red, but
those specimens here belong to the more broadly circumscribed O. succirubra.

8. Ophiorrhiza carnosicaulis H. S. Lo, Bull. Bot. Res., Harbin

10(2): 60. 1990.
rou jing she gen cao
Herbs, apparently ascending; stems stout, subglabrous.
Leaves in subequal pairs; petiole 1.57 cm, subglabrous; blade
drying thickly papery, brownish yellow with veins dark brown
abaxially, broadly ovate or broadly elliptic, 1116 5.510 cm,
glabrous on both surfaces, base subrounded, decurrent, inequilateral, margins entire, apex cuspidate; secondary veins 7 or
8 pairs; stipules caducous, not seen. Inflorescence slightly
sparse, many flowered, pubescent; peduncle ca. 3 cm; axes helicoid, 23 cm; bracteoles subulate-ensiform, 23 mm, costate,
obtuse. Flowers distylous, subsessile. Calyx pilosulous; hypanthium turbinate, 1.52 mm, 10-ribbed; lobes triangular, 0.60.7
mm, usually with 1 gland in each sinus. Corolla pale purple,
subtubular, glabrous outside; tube ca. 14 mm, pilose in basal
portion inside; lobes ovate-triangular, ca. 2 mm, dorsally with
very short horn, apex rostrate. Capsules dark purple, mitriform,
ca. 5 12 mm, pilosulous. Fl. JunJul.
Wet places in forests. Yunnan (Hekou).
The protologue described the flowers as distylous but noted that
long-styled flowers had not been seen; the putative short-styled flowers
were described as having the anthers positioned just below the throat
and the stigmas in the lower part of the corolla tube. The protologue
figure was reproduced but not exactly by FRPS (71(1): 159, t. 40, f. 1
6. 1999); in particular, the protologue figure shows the calyx as pilosulous and the corollas as pilose internally in the basal portion, but the
FRPS figure depicts the calyx and the corollas as glabrous.

9. Ophiorrhiza chinensis H. S. Lo, Bull. Bot. Res., Harbin

10(2): 70. 1990.
zhong hua she gen cao

Herbs or subshrubs, to 40(80) cm tall; stems drying

straw-yellow or purplish black, terete, subglabrous to pilosulous. Leaves in subequal pairs; petiole 14 cm; blade drying
papery, rather pale red, lanceolate to ovate, 3.512(15) cm,
usually glabrous or subglabrous on both surfaces, base cuneate,
obtuse, or rarely rounded, margins entire, apex acuminate;
secondary veins 9 or 10 pairs; stipules caducous, not seen. Inflorescences paniculiform to cymose, several to many flowered,
densely hirtellous to pilosulous; peduncle 1.53.5 cm; axes 1
3.5 cm, helicoid, deflexed, later becoming erect; bracteoles absent or reduced and caducous. Flowers distylous, on pedicels 1
2 mm. Calyx mealy puberulent; hypanthium subturbinate, 1.2
1.4 mm, 5-ribbed; lobes subtriangular, 0.40.5 mm. Corolla
white or pale purplish red, tubular-funnelform, subglabrous or
mealy puberulent outside, inside pilosulous to scaly hairy in
upper part onto lobes and with a white villous ring near middle
of tube; tube 1820 mm; lobes triangular-ovate, 2.53 mm,
dorsally with carinate narrow wing prolonged into very short
horn near apex, apex cucullate-rostrate. Infructescence axes
often becoming thickened, glabrescent, expanded, peduncle to 5
cm, axes to 6 cm, pedicels to 4 mm. Capsules obcordate-mitriform, 33.4 810 mm, subglabrous. Fl. winter and spring, fr.
spring and summer.
Fertile soil in broad-leaved forests; ca. 1300 m. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan.
The protologue detailed the long-styled and short-styled flowers,
which are said to be similar in corolla size and to differ reciprocally in
anther and stigma position, with one structure held in the middle of the
corolla tube and the other exserted from the throat, respectively.

10. Ophiorrhiza chingii H. S. Lo in S. Y. Jin & Y. L. Chen,

Cat. Type Spec. Herb. China (Suppl.), 189. 1999.
qin shi she gen cao
Herbs, weak; stems elongated, drying brownish red, glabrous or puberulent. Leaves in markedly unequal pairs; petiole
0.53 cm, puberulent; blade drying thinly papery, straw-yellow
except usually purple on midrib, lanceolate-oblong to elliptic,
larger ones 3.59.5 1.32.7 cm, smaller ones 1.56.5 0.7
1.8 cm, subglabrous on both surfaces or sparsely strigose or pilose adaxially and puberulent along principal veins abaxially,
base cuneate, margins subentire, apex caudate-acuminate; secondary veins 8 or 9 pairs; stipules caducous, not seen. Inflorescence congested-cymose, few flowered, puberulent to pilosulous; peduncle 11.5 cm; axes reduced or 2 or 3, helicoid, ca. 1
cm; bracteoles linear, 3.54 mm, obtuse. Flowers distylous,
subsessile. Calyx puberulent or pilosulous to glabrescent; hypanthium turbinate, ca. 1.2 mm, 5-ribbed; lobes subovate or
sublanceolate, ca. 0.5 mm. Corolla white, tubular-funnelform,
subglabrous outside, inside with white villous ring near middle
and sometimes scaly villosulous in upper part and onto lobes;
tube ca. 9 mm; lobes elliptic-oblong, ca. 3 mm, dorsally with
wing ca. 0.5 mm wide, apex subrostrate. Capsules not seen. Fl.
Apr.
Wet places in forests. Yunnan (Yangbi).
Although the original description (H. S. Lo, Bull. Bot. Res., Harbin 10(2): 65. 1990, not validly published) reported that the long-styled
flowers were not found, the figure in FRPS (72(1): 161, t. 41, f. 913.

RUBIACEAE

1999) illustrated these. That figure shows the corollas as similar in size
between the two forms but with the internal pubescence restricted to the
upper middle part of the tube in the short-styled form and from the
middle of the tube through the top of the lobes in the long-styled form;
also, the anthers and stigmas are reciprocally positioned near or just
below the middle of the corolla tube and in the throat, respectively.

11. Ophiorrhiza cordata W. L. Sha ex S. Y. Jin & Y. L. Chen,

Cat. Type Spec. Herb. China (Suppl.), 189. 1999.
xin ye she gen cao
Herbs, procumbent to creeping; stems densely brown villous with multicellular trichomes. Leaves in subequal pairs;
petiole 0.81.7 cm, densely brownish red villous; blade drying
papery, adaxially grayish black, abaxially brownish, broadly
ovate to suborbicular, 3.56 2.34 cm, subglabrous or sparsely
pilose adaxially, villous abaxially along principal veins, base
cordate, margins sparsely ciliate, apex obtuse then abruptly
shortly acute; secondary veins 57 pairs; stipules caducous, not
seen. Inflorescence congested-cymose, 48-flowered, brownish
red villous; peduncle ca. 1 cm; axes reduced; bracts ellipticoblong, 3.56 mm, ciliate. Flowers with breeding biology unknown, subsessile. Calyx subglabrous; hypanthium turbinate,
ca. 2 mm, 10-ribbed; lobes triangular, ca. 0.5 mm, obtuse or
perhaps acute. Corolla white, tubular-funnelform, glabrous outside; tube 1112 mm, inside glabrous except with villous ring at
middle; lobes triangular, ca. 2.2 mm, dorsally narrowly winged
and with very short horn, apex rostrate. Capsules unknown. Fl.
Apr.

267

13. Ophiorrhiza densa H. S. Lo, Bull. Bot. Res., Harbin 10(2):

57. 1990.
mi mai she gen cao
Herbs or subshrubs, ascending, to 1 m tall; stems terete,
drying brownish purple, brown pilosulous to glabrescent.
Leaves in subequal pairs; petiole 13 cm or longer, densely pilosulous or tomentose; blade drying thinly papery, adaxially
black, abaxially pale purple, ovate or lanceolate-ovate, usually
inequilateral, (5)815(18) 1.54 cm, sparsely pilosulous
adaxially, crisped tomentose along veins abaxially, base subcuneate, margins entire, apex caudate; secondary veins 13
18(23) pairs; stipules caducous, not seen. Inflorescence 48
cm, many flowered, brown tomentose; axes helicoid; bracteoles
linear, ca. 3 mm. Flowers distylous, sessile to subsessile. Calyx
hispidulous; hypanthium turbinate to subglobose, ca. 1.2 mm,
5-ribbed; lobes narrowly triangular, ca. 1 mm. Corolla pale
purple, subtubular to inflated, densely pilosulous outside, inside
pilosulous near middle of tube and sometimes onto lobes,
and/or densely glandular-pilose on lobes; tube ca. 10 mm; lobes
subdeltoid, ca. 1.5 mm, dorsally with very short horn, apex
rostrate. Capsules unknown. Fl. Nov.
Dense forests; 14001600 m. Yunnan (Malipo).

Neither the original description (H. S. Lo, Bull. Bot. Res., Harbin
10(2): 55. 1990, not validly published) nor FRPS (71(1): 154156.
1999) described the floral biology of this species; the flowers described
resemble the long-styled form of distylous species, with the anthers
positioned near the middle of the corolla tube and the stigmas in the
corolla throat.

In the protologue and FRPS (71(1): 158. 1999), H. S. Lo described the floral forms as having similar corollas and reciprocally
placed anthers and stigmas near the middle of the corolla tube and
shortly exserted, respectively. However, the protologue figure showed
differences between the floral forms in the internal pubescence of the
corollas, with the pubescence confined to the general area of the middle
of the tube in the short-styled flowers but distributed from the middle of
the tube to the tops of the lobes in the long-styled form; this internal
corolla pubescence was not described in the protologue text. The insides
of the corolla lobes were described as glanduloso-piloso in the protologue text, which is unusual and also not shown in one of the protologue
figures of the corolla interior.

12. Ophiorrhiza crassifolia H. S. Lo, Bull. Bot. Res., Harbin

10(2): 47. 1990.

14. Ophiorrhiza dulongensis H. S. Lo, Bull. Bot. Res., Harbin

10(2): 27. 1990.

hou ye she gen cao

du long she gen cao

Herbs, perhaps ascending, notably fleshy, to 30 cm tall;

stems flattened to terete, drying brown, puberulent to glabrous.
Petiole 0.53.5 cm; leaf blade drying thickly papery, purplish
red or yellowish brown abaxially, broadly ovate or broadly
elliptic, 512 3.57 cm, glabrous on both surfaces, abaxially
densely minutely orbicular scaly, base obtuse, subrounded,
or subcordate, usually oblique, margins entire, apex obtuse
to shortly cuspidate; secondary veins 5 or 6 pairs; stipules caducous, not seen. Inflorescences cymose or congested-cymose,
many flowered, puberulent; peduncle 2.55.5 cm; axes short,
helicoid; bracteoles narrowly spatulate, 55.5 mm, obtuse.
Flowers with biology unknown, with pedicels 23 mm. Calyx
puberulent; hypanthium turbinate, ca. 2 mm, 10-ribbed; lobes
subtriangular, ca. 0.6 mm, in sinus with 1 gland. Corolla purplish red, salverform-funnelform, glabrous outside; tube 2425
mm, pilose inside; lobes ovate-triangular, ca. 2.2 mm, dorsally
ridged with very short horn. Capsules rhomboid, ca. 5 15
mm, glabrous or subglabrous. Fl. Oct.

Herbs, weak to procumbent; stems drying purplish brown,

pilosulous. Petiole 0.31 cm, densely pilosulous or villosulous;
leaf blade drying membranous to papery, often green with veins
purplish brown abaxially, broadly ovate or ovate, 14 0.62.5
cm, scattered scabrous-strigillose adaxially, abaxially moderately pilose along principal veins, base obtuse to rounded, apex
acute; secondary veins 46 pairs; stipules persistent, subulate,
46 mm, glabrous. Inflorescence fasciculate, 3- or 4-flowered,
glabrescent; peduncle ca. 1 cm or slightly longer; bracts linear,
ca. 1 mm. Flowers distylous. Calyx puberulent to glabrescent;
hypanthium 11.5 mm, 5-ribbed; lobes narrowly lanceolate or
lanceolate, 1.52 mm. Corolla white, funnelform, subglabrous
outside; tube 77.5 mm, densely villous in throat; lobes triangular-ovate, 2.53 mm, dorsally ribbed at least in bud. Capsules
unknown. Fl. Jul.

Forests. Guangxi (Napo).

Rocks in forests. Guangxi (Longzhou).

Evergreen broad-leaved forests; 23002400 m. NW Yunnan.

In the protologue and FRPS (71(1): 127. 1999), H. S. Lo detailed
the floral forms, noting that their corollas are similar in size and the an-

RUBIACEAE

268

thers and stigmas are reciprocally placed near the middle of the corolla
tube and in its throat, respectively.

in having the stigmas exserted and the anthers positioned below them in
the upper part of the corolla tube.

15. Ophiorrhiza ensiformis H. S. Lo, Bull. Bot. Res., Harbin

10(2): 22. 1990.

17. Ophiorrhiza fasciculata D. Don, Prodr. Fl. Nepal. 136.

1825.

jian chi she gen cao

cu hua she gen cao

Herbs, ascending; stems pilosulous at least when young.

Leaves in subequal pairs; petiole 13 cm, pilose; blade drying
submembranous, brown or dark brown, elliptic, ovate, or elliptic-oblong, 512 2.55.5 cm, adaxially sparsely ferruginous
pilosulous, abaxially glabrous or subglabrous, base obtuse
then narrowed to decurrent, margins entire, apex acute to rather
abruptly acuminate; secondary veins 1012 pairs; stipules generally persistent, broadly triangular then abruptly narrowed, 35
mm, glabrescent, ciliate, long acuminate. Inflorescences congested-cymose, several to many flowered, ferruginous pilosulous; peduncle ca. 1.5 cm; axes rather short, helicoid; bracts linear-ensiform to narrowly spatulate, 57 mm, glabrescent, pinnately veined, ciliate, acute. Flowers with biology unknown,
with short pedicels. Calyx with hypanthium subturbinate, ca.
1.5 2.5 mm, 5-ribbed, with dense ferruginous long trichomes
mixed with unicellular trichomes; lobes linear-ensiform to spatulate, 57.5 mm, pinnately veined, glabrescent except ciliate,
acute. Corolla white, tubular-funnelform, outside 5-ribbed or 5ridged in upper portion, glabrous or sparsely hispidulous along
ribs; tube 1314 mm, inside with white villous ring in throat;
lobes triangular, 22.5 mm, dorsally narrowly winged, obtuse,
rostrate. Capsules rhomboid-pyramidal, ca. 4 8 mm, pilosulous. Fl. Jan.

Herbs or subshrubs, erect, to 0.5(2) m tall; stems pilosulous or puberulent to glabrescent. Leaves in subequal pairs;
petiole 11.5[5] cm, puberulent; blade drying papery, sometimes darkened adaxially, pale abaxially, elliptic to lanceolateelliptic, 812[16] 36 cm, glabrous or sparsely strigillose
adaxially, abaxially puberulent on veins, base obtuse to cuneate
then attenuate, apex acute to caudate-acuminate; secondary
veins 1013 pairs; stipules narrowly triangular, 410 mm, puberulent to glabrescent. Inflorescences congested-cymose to
subfasciculate, several flowered, densely pilosulous; peduncle
18 cm; branched portion 12 cm; bracts ligulate-lanceolate, 6
12 mm, persistent. Flowers with biology unknown, subsessile.
Calyx densely puberulent to hirtellous; hypanthium compressed
cylindrical, 11.5 mm; lobes ovate to deltoid, 11.2 mm. Corolla white sometimes flushed with pink, drying yellowed, tubular-funnelform, outside puberulent; tube 1015 mm, inside
glabrous; lobes ovate-oblong, 1.52.5 mm. Capsules compressed rhombic, 1.54.5 410.5 mm, puberulent or hirtellous. Fr. Aug.

Streamsides; ca. 2000 m. Yunnan (Longling).

16. Ophiorrhiza fangdingii H. S. Lo in S. Y. Jin & Y. L. Chen,

Cat. Type Spec. Herb. China (Suppl.), 190. 1999.
fang ding she gen cao
Herbs, weak at base, ascending above, to 40 cm tall; stems
subterete, drying brown, subglabrous. Leaves in unequal pairs;
petiole 0.52 cm, glabrous; blade drying papery, adaxially
leaden gray, abaxially pale with veins brown, oblong-ovate or
oblong-lanceolate, larger ones 57.5 cm, smaller ones 25 cm,
glabrous on both surfaces, base subcuneate and somewhat inequilateral, margins subentire, apex obtuse then acuminate; secondary veins 510 pairs; stipules caducous, not seen. Inflorescences 4- or 5-flowered; peduncle ca. 1 cm; axes short, helicoid; bracteoles linear or narrowly lanceolate, 1018 1.54
mm, obtuse, costate. Flowers reportedly distylous, with short
pedicels. Calyx glabrous; hypanthium turbinate, ca. 1.2 mm, 5ribbed; lobes linear-lanceolate, unequal, 23 mm, obtuse. Corolla white, funnelform, glabrous outside; tube ca. 23 mm, white
villous above middle inside; lobes dorsally narrowly winged,
wing extending into very short horn. Capsules not seen. Fl. Jan.
Wet places in forests on limestone; ca. 1200 m. Guangxi
(Napo).
In the original description (Bull. Bot. Res., Harbin 10(2): 40, 42.
1990, not validly published) and FRPS (71(1): 144. 1999), H. S. Lo
described this species as distylous but described only the putative longstyled flowers; these resemble the long-styled form of distylous species

Broad-leaved forests; ca. 1700 m. Xizang (Mdog) [Bhutan, India,

Myanmar, Nepal].
Measurements in brackets are taken from the description of this
species by Deb and Mondal (Bull. Bot. Surv. India 39(14): 4447.
1997) and may be expected in Chinese plants.

18. Ophiorrhiza filibracteolata H. S. Lo in S. Y. Jin & Y. L.

Chen, Cat. Type Spec. Herb. China (Suppl.), 190. 1999.
da qiao she gen cao
Herbs, apparently ascending, to 30 cm tall; stems mealy
puberulent. Leaves in subequal pairs; petiole 15 cm; blade
drying papery, dark brown, ovate to broadly ovate, 2.55.5
1.53.3 cm, glabrous, base obtuse or subrounded, margins entire, apex acute; secondary veins 7 or 8 pairs; stipules caducous,
not seen. Inflorescence somewhat congested-cymose, many
flowered; peduncle ca. 3 cm; axes up to 1 cm, helicoid; bracteoles filiform, fleshy, 2.53 mm. Flowers distylous, subsessile
or pedicels to 2 mm. Calyx pilosulous; hypanthium subturbinate, ca. 1.4 mm, shallowly 5-ribbed; lobes triangular to narrowly lanceolate, ca. 0.7 mm, with 1 gland in each sinus. Corolla white, tubular-funnelform, outside pilosulous, inside with
white villous ring at middle and scaly pilose from middle onto
lobes; tubes 1112 mm; lobes ovate-triangular, ca. 3 mm, dorsally narrowly winged and with very short horn. Capsules not
seen. Fl. Apr.
Forests. Guangdong (Ruyuan).
The original description (H. S. Lo, Bull. Bot. Res., Harbin 10(2):
5253. 1990, not validly published) described the flowers as distylous
with the short-styled flowers as unknown; the putative long-styled flowers were described as having the anthers positioned just below the middle of the corolla tube and the stigmas in the throat. In FRPS (71(1):
153. 1999), H. S. Lo gave the plant height as 20 cm, but the original

RUBIACEAE

description said 30 cm. Much of the information on pubescence details

here is taken from the figure in the original description.

19. Ophiorrhiza gracilis Kurz, J. Asiat. Soc. Bengal, Pt. 2,

Nat. Hist. 41: 311. 1872.
xian ruo she gen cao
Herbs, weak at base, suberect above, to 30 cm tall; stems
glabrous. Petiole glabrous; leaf blade drying membranous-papery, grayish on both surfaces, paler abaxially, lanceolate or
subovate, 612(17) 24.5 cm, glabrous on both surfaces,
base cuneate to attenuate and usually oblique, apex caudate to
long acuminate; secondary veins 812 pairs; stipules caducous,
not seen. Inflorescences many flowered, puberulent; peduncle
23 cm; bracteoles subulate, 0.51.2 mm, early caducous.
Flowers distylous, subsessile. Calyx with hypanthium turbinate,
ca. 1.3 mm, 5-ribbed; lobes ovate-triangular, 0.40.5 mm. Corolla white or reddish at apex, tubular; tube ca. 10.5 mm, white
villous in throat and at middle inside; lobes triangular-ovate,
1.21.3 mm, dorsally with horn 0.81 mm. Capsules not seen.
Fl. spring.
Dense forests. Yunnan [Myanmar].
H. S. Lo (in FRPS 71(1): 171. 1999) described the anthers as situated near the base of the corolla tube and the stigmas positioned just
below the throat in the long-styled form vs. the anthers situated above
the middle of the corolla tube and the stigmas near the base in the shortstyled form.

20. Ophiorrhiza grandibracteolata F. C. How ex H. S. Lo,

Bull. Bot. Res., Harbin 10(2): 43. 1990.
da bao she gen cao
Herbs or subshrubs, weak at base, ascending above, to 70
cm tall; stems subterete, densely villous with trichomes multicellular or sometimes also unicellular. Leaves in unequal pairs;
petiole 0.52(3) cm, stout, densely multicellular villous; blade
drying thinly papery, gray-black adaxially, reddish or pale abaxially, ovate, broadly ovate, or lanceolate-ovate, larger ones 4
12(15) 24.5(6) cm, smaller ones 24(6) 1.22.5 cm,
both surfaces glabrescent or multicellular villous on principal
veins, base obtuse to cuneate and sometimes shortly decurrent,
apex acute; secondary veins 710 pairs; stipules caducous, not
seen. Inflorescence congested-umbelliform-cymose to subcapitate, 5- to many flowered or rarely 1-flowered, densely multicellular villous; peduncle ca. 0.8 cm; axes reduced; bracts and
bracteoles ovate to lanceolate, 1015 mm, pinnately veined,
glabrescent except multicellular ciliate along margin and on
dorsal costa. Flowers distylous, subsessile or on stout pedicels
to 2 mm. Calyx multicellular villous; hypanthium broadly turbinate, 22.3 mm; lobes triangular, 11.2 mm. Corolla white or
reddened, drying purplish red, funnelform, outside with 5 strigose lines from middle of tube to apices of lobes; tube 2225
mm, inside with white villous ring above middle; lobes subovate, ca. 5 mm, sometimes weakly reticulate-veined, dorsally
narrowly winged. Capsules rhomboid, 44.5 ca. 11 mm, villous. Fl. Nov.
Wet places in forests; 12001500 m. Guangxi (Napo), Yunnan.
The protologue and H. S. Lo in FRPS (71(1): 146. 1999) described this species as distylous, with the anthers borne well above the

269

middle of the corolla tube and the stigmas positioned in the throat in one
floral form vs. the anthers positioned near the middle of the corolla tube
and the stigmas well exserted in the other.

21. Ophiorrhiza hainanensis Y. C. Tseng, Fl. Hainan. 3: 582.

1974.
hai nan she gen cao
Herbs, weak to erect, to 50 cm tall; stems viscous puberulent with unicellular trichomes, to glabrescent. Leaves in subequal pairs; petiole 12.5 cm, puberulent; blade drying thinly
papery, green on both surfaces, elliptic to ovate, 613 2.55
cm, adaxially glabrous, abaxially pilosulous along principal
veins, base obtuse to acute, margins entire, apex shortly acuminate; secondary veins 1517 pairs; stipules deciduous, triangular, 35 mm, aristate-acuminate. Inflorescence congested-cymose, several flowered, viscous puberulent; peduncles 0.51
mm, stout; bracts elliptic-oblong, 56 mm, viscid ciliate. Flowers with biology unknown. Calyx in bud sparsely puberulent;
hypanthium to 2 mm; lobes triangular, to ca. 1 mm. Corolla in
bud tubular. Capsules turbinate, ca. 4 9 mm, sparsely puberulent. Fl. (bud) and fr. Dec.
Dense forests, uncommon. Hainan (Baoting, Qiongzhong).
Ophiorrhiza hainanensis was synonymized with O. nutans by
Duan and Lin (Acta Phytotax. Sin. 45: 877. 2007). However, this
species is here recognized provisionally, based on the distinctions given
in the key to species.

22. Ophiorrhiza hayatana Ohwi, Repert. Spec. Nov. Regni

Veg. 36: 57. 1934.
liu guo she gen cao
Ophiorrhiza stenophylla Hayata, Icon. Pl. Formosan. 2:
91. Oct 1912, not Valeton (Feb 1912).
Herbs, erect, to 50 cm tall; stems subterete or weakly 4angled, glabrous to puberulent or strigillose. Leaves in subequal
pairs; petiole 0.31.5(2) cm, glabrous to puberulent; blade
drying papery, discolorous, narrowly elliptic-oblong, elliptic, or
lanceolate, 210(15) 0.82.5(3) cm, glabrous on both surfaces or often hispidulous near margins adaxially, base acute to
attenuate, apex acute to acuminate; secondary veins 48 pairs;
stipules deciduous, broadly triangular to rounded, 0.51 mm,
puberulent to glabrescent and/or glandular, sometimes erose. Inflorescence several to many flowered, cymose, densely puberulent to pilosulous sometimes in lines, fasciculate or peduncle
1.52(4) cm; axes subcapitate to helicoid; bracteoles linear to
narrowly triangular, 0.54 mm, glabrous, persistent or deciduous as fruit develop. Flowers with biology unknown, subsessile
or on pedicels to 1.5 mm, sometimes tuberculate. Calyx glabrous to puberulent; hypanthium suburceolate to oblate, ca.
1.2(2) mm, densely tuberculate with peglike structures on
lower part and also sometimes on sides, 5-ribbed; lobes linear
to spatulate, 11.3 mm. Corolla white, slenderly funnelform,
outside glabrous; tube (12)1315 mm, inside barbate in throat;
lobes triangular, 2.53(4) mm, ciliate, dorsally winged and
with very short horn near apex. Capsules obcordate, ca. 4 8
10 mm, tuberculate and sometimes pilosulous.
Broad-leaved forests; 500900 m. Taiwan.

RUBIACEAE

270

Some measurements given by H. S. Lo in FRPS (71(1): 129.

1999) have not been seen on specimens and are provisionally presented
in parentheses. The floral biology of this species has not been described;
the specimens seen appear to be monomorphic with the anthers positioned at or just below the corolla throat and the stigmas perhaps positioned at the same level.

23. Ophiorrhiza hispida J. D. Hooker, Fl. Brit. India 3: 83.

1880.
jian ye she gen cao
Herbs, ascending, to 1 m tall; stems moderately villous or
hispid. Leaves in subequal pairs; petiole 1.54(7) cm, densely
pilose or hispid; blade drying thinly papery, gray or olive-green,
ovate or broadly ovate, 717 3.57 cm, sparsely hispid on
both surfaces or moderately so abaxially, base obtuse, decurrent, often oblique, margins subentire, apex acuminate; secondary veins 914 pairs; stipules sublanceolate, markedly contracted above, pilose, ciliate, acuminate. Inflorescence congested-cymose, many flowered, villous; peduncle 1.52 cm;
bracts linear, 34 mm, ciliate, persistent. Flowers distylous,
subsessile. Calyx pilose; hypanthium subturbinate, ca. 1 mm, 5ribbed; lobes subtriangular, ca. 0.5 mm, usually with 1 gland in
each sinus. Corolla pale purple or white, slenderly tubular,
sparsely pilose or hispid outside; tube 1012 mm, inside with
white villous ring near middle; lobes ovate-triangular, ca. 1
mm. Capsules ca. 2 5 mm, pilose or hispid.
Forests. SW Yunnan [NE India].
H. S. Lo (in FRPS 71(1): 134. 1999) noted that the corollas of the
two floral forms are similar but that the flowers differ in the reciprocally
placed anthers and stigmas positioned near the middle of the corolla
tube or in the throat, respectively.
This species is circumscribed here following H. S. Lo (loc. cit.)
and does not correspond to the circumscription of this species by Deb
and Mondal (Bull. Bot. Surv. India 39(14): 5961. 1997). Deb and
Mondal described Ophiorrhiza hispida as having corollas with tubes 3
4 mm, leaves 215 14 cm, a plant stature of 35 cm or less, and calyx
lobes 0.81 mm; the plants treated as O. hispida by Lo key to O. fasciculata in Deb and Mondals treatment.

24. Ophiorrhiza hispidula Wallich ex G. Don, Gen. Hist. 3:

523. 1834.
ban na she gen cao
Herbs, procumbent at base, ascending above, to 15 cm tall;
stems glabrescent to densely villosulous. Leaves in subequal
pairs; petiole 0.31 cm, puberulent to glabrescent; blade drying
thinly papery, grayish, ovate, broadly ovate, elliptic, or sublanceolate, 1.54.5(7) 12.5 cm, adaxially glabrescent to
sparsely strigillose, abaxially glabrous except pilosulous to puberulent along principal veins, base cuneate to obtuse, margins
entire, apex acute; secondary veins 57 pairs; stipules persistent
on uppermost nodes, triangular becoming strongly narrowed,
410 mm, glabrescent, long acuminate to aristate. Inflorescences cymose, several flowered, puberulent to hispidulous;
peduncle 12.5(4) cm; axes developed, helicoid; bracts few,
narrowly triangular to linear, 0.52 mm. Flowers subsessile.
Calyx puberulent to pilosulous; hypanthium ellipsoid to subglobose, ca. 1 mm; lobes narrowly triangular, 0.81 mm. Corolla
white, tubular, outside puberulent; tube 45 mm, barbate in

throat and glabrescent to puberulent inside tube; lobes triangular-ovate, ca. 1.5 mm. Capsules obcordate, 1.52.5 45
mm, puberulent to villosulous. Fl. MaySep, fr. JulOct.
Dense forests. Yunnan (Xishuangbanna) [Bangladesh, India (including Andaman Islands), Indonesia, Malaysia, Thailand].
Ophiorrhiza hispidula was synonymized by Deb and Mondal
(Bull. Bot. Surv. India 39(14): 131133. 1997) under O. trichocarpa
Blume, without explanation and reportedly without having seen Blumes
type; the name O. hispidula is provisionally used here pending further
study.

25. Ophiorrhiza howii H. S. Lo, Bull. Bot. Res., Harbin 18:

277. 1998.
kuan zhao she gen cao
Ophiorrhiza longiflora F. C. How ex H. S. Lo, Bull. Bot.
Res., Harbin 10(2): 70. 1990, not Blume (1826).
Herbs, ascending, to ca. 1 m tall; stems terete to somewhat
flattened, rather stout, drying purplish brown to black, subglabrous. Leaves in subequal pairs; petiole 12 cm, densely pilosulous; blade narrowly ovate, narrowly elliptic, or ovate, 59
23 cm, abaxially usually pilosulous along principal veins,
base cuneate, apex acuminate; secondary veins 813 pairs; stipules caducous, unknown. Inflorescence cymose, several to
many flowered, densely pilosulous; peduncles 1.52 cm; axes
short, helicoid; bracts absent or minute and caducous. Flowers
with biology unknown, on pedicels 12 mm. Calyx with hypanthium portion subturbinate and slightly compressed, ca. 1.7
2.3 mm, 5-ribbed, mealy puberulent; lobes sublanceolate, ca.
0.5 mm, subglabrous. Corolla white, salverform, puberulent outside; tube 2224 mm, pilose or glabrous inside; lobes broadly
ovate, 34 mm, dorsally with horn 1.52 mm, apex incurved
rostrate. Capsules not seen. Fl. Oct.
Forests; 11001500 m. Yunnan (Maguan).
The protologue described the corolla tubes as pilose inside, but the
protologue figure showed the corolla tube to be glabrous inside.

26. Ophiorrhiza huanjiangensis D. Fang & Z. M. Xie, Acta

Phytotax. Sin. 40: 155. 2002.
huan jiang she gen cao
Herbs, procumbent to creeping; stems to 23 cm, densely to
moderately hirtellous or pilosulous, to sometimes glabrescent.
Leaves in subequal pairs; petiole 0.151 cm, hirtellous; blade
drying papery, ovate to elliptic-ovate, 0.51.85 0.31.2 cm,
both surfaces puberulent at least on principal veins to glabrescent, base obtuse to cordulate, apex obtuse to acute; secondary veins 3 or 4 pairs; stipules caducous, not seen. Inflorescence cymose to umbelliform, 13-flowered, hirtellous to glabrescent; peduncle and/or pedicel 510 mm; bracts linear, 12
mm, sparsely puberulent to glabrescent. Flowers apparently
distylous, pedunculate or pedicellate. Calyx puberulent to glabrous; hypanthium turbinate, 11.5 mm; lobes ovate to narrowly triangular, ca. 1 mm. Corolla white, funnelform, outside
glabrous; tube 1012 mm, puberulent inside; lobes ovate, 45
mm. Capsules unknown. Fl. Apr.
Dense forests in valleys; ca. 400 m. Hunan (Huitong).

RUBIACEAE

The protologue described the flowers as distylous and reported

that only the short-styled form has been seen.

27. Ophiorrhiza hunanica H. S. Lo, Bull. Bot. Res., Harbin

10(2): 24. 1990.
hu nan she gen cao
Herbs, procumbent at base, to 15 cm tall; stems drying
black, subglabrous. Leaves in unequal pairs; petiole 13 cm,
subglabrous; blade drying papery, purple on veins, obovate or
ovate, 618 36.5 cm, adaxially sparsely hirtellous-strigose,
abaxially subglabrous or villous on veins, base cuneate, apex
obtuse to subacute; secondary veins 1014 pairs; stipules often
persistent, ovate, 68 mm, ciliate, acuminate. Inflorescence cymose, many flowered, densely villous, pendulous; peduncle ca.
3 cm, arching; principal axes 2 or 3 pairs, helicoid; bracts linear,
815 11.4 mm, sparsely pinnately veined, sparsely ciliate.
Flowers with biology unknown, subsessile. Calyx with hypanthium compressed turbinate, ca. 2 mm, 5-ribbed, densely multicellular villous; lobes linear, 67 mm, hispidulous along costa.
Corolla in bud with color unknown, 5-winged, wings ciliate.
Capsules unknown. Fl. Nov.
Dense forests in valleys; ca. 400 m. Hunan (Huitong).
In the protologue the anthers and stigmas of the flower buds were
described, but these immature structures are not reliable indicators of
the size or arrangement of the mature structures.

28. Ophiorrhiza japonica Blume, Bijdr. 978. 18261827.

ri ben she gen cao
Ophiorrhiza acutiloba Hayata; O. cavaleriei H. Lveill;
O. eryei Champion; O. dimorphantha Hayata; O. dimorphantha
f. brevistigma Hayata; O. dimorphantha f. longistigma Hayata;
O. japonica var. acutiloba (Hayata) Ohwi; O. japonica var.
minor J. Krause; O. labordei H. Lveill; O. monticola Hayata;
O. monticola f. brevistigma Hayata; O. monticola f. longistigma
Hayata; O. nigricans H. S. Lo.
Herbs, weak to ascending, to 60 cm tall, often drying
flushed with purple or darkened; stems subterete to slightly
compressed, glabrous or with 2 hirtellous or pilosulous lines.
Leaves in subequal pairs; petiole 0.32(3) cm, glabrous to
hirtellous or puberulent; blade drying papery, ovate, ellipticovate, elliptic, elliptic-oblong, lanceolate, or narrowly lanceolate, 111 0.73.5 cm, glabrous to strigillose or hispidulous
adaxially, glabrous to puberulent, hirtellous, or villosulous
abaxially, base cuneate to obtuse, margins flat to crisped, apex
acute to acuminate or rarely obtuse (to rounded, Ophiorrhiza
nigricans); secondary veins 48 pairs; stipules triangular, 0.82
mm, glabrescent, acute to bifid, caducous sometimes leaving 1
to several thickened scars to 0.3 mm. Inflorescence congestedcymose to cymose, few to many flowered, puberulent to strigillose, pilosulous, or hirtellous; peduncle 0.55 cm (to 6 cm, O.
nigricans); axes congested-cymose becoming helicoid; bracts
lanceolate-linear, spatulate, or linear, 16 mm, sometimes glabrescent, persistent. Flowers distylous, on pedicels 12 mm.
Calyx glabrous to densely puberulent or pilosulous; hypanthium subturbinate to oblate or subglobose, 0.81 mm, 5-

271

ribbed; lobes triangular, 0.41.2 mm. Corolla white or pink,

funnelform to tubular-funnelform, outside glabrous to puberulent or pilosulous and longitudinally winged, inside pilose near
middle and pilosulous above middle through throat and sometimes onto lobes; tube 914 mm; lobes triangular to ovate, (1.5
1.8 mm, O. nigricans) 2.54 mm, dorsally with wing to 0.5
mm wide, apex rostrate. Capsules submitriform, 2.54 6.59
mm, pilosulous to glabrous. Fl. winter, fr. spring and summer.
Fertile soil of ravines in forests; 1002400 m. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, S
Shanxi?, Sichuan, Taiwan, NE Yunnan, Zhejiang [Japan, Vietnam].
Duan and Lin (Acta Phytotax. Sin. 45: 873. 2007) synonymized
Ophiorrhiza nigricans with O. japonica. They also synonymized O.
kwangsiensis with O. japonica, but that species is provisionally separated here based on the characters given in the key to species pending
further study. The reproductive biology of O. japonica was studied by
Nakamura et al. (J. Plant Res. 120: 501509. 2007), who found the Chinese plants sampled to be distylous and diploid. The corollas are apparently similar between the two floral forms, with the reciprocally placed
anthers and stigmas positioned near the middle of the corolla tube vs. at
or just above its throat, respectively.

29. Ophiorrhiza kuroiwae Makino, Bot. Mag. (Tokyo) 20: 5.

1906 [Kuroiwai].
xiao hua she gen cao
Ophiorrhiza japonica Blume var. kuroiwae (Makino) Ohwi; O. kotoensis Hatusima; O. liukiuensis Hayata; O. parviflora
Hayata (1912), not Reinwardt ex Korthals (1851).
Herbs, erect, to 60 cm tall; stem 4-angled, densely strigillose-tomentulose to glabrescent. Leaves in subequal pairs; petiole 0.52.3(3) cm, villosulous-tomentulose to glabrescent;
blade drying membranous to papery, red or pale, oblong-ovate,
ovate, or elliptic-oblong, 512(15) 26 cm, adaxially
sparsely scaberulous, abaxially puberulent at least on principal
veins, base cuneate to obtuse then often narrowed and shortly
attenuate, margins entire, apex acute or occasionally obtuse or
shortly acuminate; secondary veins 611 pairs; stipules deciduous, deeply 2-parted, 37 mm, puberulent, lobes cuspidate or
filiform. Inflorescences cymose to paniculate, several to many
flowered, hirtellous- or hispidulous-tomentulose; peduncle 3.5
5 cm; axes several, elongating as buds develop, helicoid; bracts
reduced, caducous. Flowers with biology unknown, subsessile
or on pedicels to 2 mm. Calyx puberulent to glabrous; hypanthium subglobose, 11.5 mm; lobes triangular, 0.30.5 mm,
dorsally keeled. Corolla white, tubular or inflated, outside glabrous to puberulent, inside pubescent in upper part of tube and
onto lobes; tube 2.53.5 mm; lobes ovate-triangular, ca. 1 mm,
dorsally ridged. Capsules reniform-oblate, 23.5 69 mm, 5ribbed, glabrescent. Fl. AprOct, fr. SepFeb.
Taiwan [Japan, Philippines].

30. Ophiorrhiza kwangsiensis Merrill ex H. L. Li, J. Arnold

Arbor. 24: 453. 1943.
guang xi she gen cao
Herbs, creeping to weakly ascending, to 18 cm tall; stems
drying yellowish brown, subglabrous or with puberulent to

RUBIACEAE

272

strigillose lines. Leaves in subequal pairs; petiole 0.51.5 cm,

glabrous; blade drying papery to membranous, leaden gray
or olive-green adaxially, pale green or yellowed abaxially,
cordiform to cordate-ovate, 0.82 0.71.5 cm, glabrous on
both surfaces or puberulent adaxially, base cordate to subtruncate, margins entire, apex acute to somewhat obtuse; secondary
veins 46 pairs; stipules caducous, not seen. Inflorescence
congested-cymose, few or several flowered, densely hirtellous
to pilosulous; peduncle 0.71.5 cm; axes reduced or shortly
helicoid; bracts linear to linear-lanceolate, 4.56 mm, glabrous
except sparsely ciliolate. Flowers reportedly distylous, subsessile, fragrant. Calyx puberulent; hypanthium turbinate, ca. 1
mm, weakly 5-ribbed; lobes oblong-lanceolate, in bud 12.5
mm and equal to unequal. Corolla pale yellow or reddish, tubular-funnelform, outside subglabrous; tube 910 mm, inside
sparsely pilose; lobes ovate-triangular, ca. 2 mm, dorsally costate and with very short horn. Capsules ?mitriform, 22.5 67
mm, puberulent. Fl. early spring or Sep.
Shady wet places in forests. Guangxi (Shangsi).
H. S. Lo (in FRPS 71(1): 145. 1999) reported this species as distylous but described only one floral form, similar to the short-styled flowers of distylous species. The protologue described only young flower
buds with no description of anther or stigma position. The protologue
described the calyx lobes as ca. 1 mm without any note about them
being unequal, while H. S. Lo (loc. cit.) described these as 1.52.2 mm
and usually unequal; both conditions are included here provisionally.
Ophiorrhiza kwangsiensis was synonymized with O. japonica by
Duan and Lin (Acta Phytotax. Sin. 45: 873. 2007), but a number of
characters seem to distinguish it; it is provisionally separated here
pending further study.

31. Ophiorrhiza laevifolia H. S. Lo, Bull. Bot. Res., Harbin

10(2): 80. 1990.
ping hua she gen cao
Herbs, suberect, to 80 cm tall; stems subterete to slightly
compressed, drying black on upper part, puberulent or subglabrous. Leaves in subequal pairs; petiole 0.5(1.5) cm; blade
drying papery, adaxially dark green, abaxially pale green, ovate
or lanceolate, inequilateral, 412 24.5 cm, glabrous on
both surfaces, base cuneate, margins undulate or flat, apex
acute, acuminate, or caudate; secondary veins 5 or 6 pairs; stipules caducous, not seen. Inflorescence congested-cymose, many
flowered, densely ferruginous hirtellous; peduncle reduced or
up to 1 cm; axes reduced to shortly helicoid; bracts absent or
minute. Flowers reportedly distylous, subsessile. Calyx glabrescent; hypanthium obovoid, ca. 2 mm, 5-ribbed; lobes narrowly
lanceolate, ca. 1 mm, with 1 gland in each sinus. Corolla pink,
subtubular, glabrous outside; tube ca. 13 mm, sparsely white
villous above middle inside; lobes subovate, 1.82 mm, dorsally with short horn 0.10.2 mm, apex rostrate. Capsules not
seen. Fl. Apr.
Semi-evergreen forests; 8001000 m. Xizang (Mdog).
The protologue reported this species as distylous but described
only one floral form, similar to the short-styled flowers of distylous
species; this was described as having the stigmas positioned not far below the middle of the corolla tube and the anthers exserted.

32. Ophiorrhiza laoshanica H. S. Lo, Bull. Bot. Res., Harbin

10(2): 65. 1990.
lao shan she gen cao
Herbs, suberect, to 50 cm tall; stems sulcate to subterete,
drying grayish yellow, subglabrous. Leaves in markedly unequal
pairs; petiole 0.31 cm, subglabrous; blade drying thinly papery, adaxially gray to blackened, abaxially grayish yellow,
ovate, larger ones 68(11) 23 cm, smaller ones 1.55 12
cm, glabrous on both surfaces, base obtuse to acute then decurrent, apex shortly acuminate; secondary veins 69 pairs; stipules caducous, not seen. Inflorescence ferruginous mealy puberulent; peduncle 12 cm; axes short, helicoid; bracteoles filiform-subulate, ca. 3 mm. Flowers reportedly distylous, subsessile. Calyx mealy puberulent; hypanthium obconic-turbinate,
ca. 1 mm, 5-ribbed; lobes triangular to deltoid, ca. 0.4 mm. Corolla white, tubular-funnelform, outside glabrous; tube 910
mm, inside near middle with villous fascicles mixed with scalelike pilose trichomes; lobes subovate, ca. 0.7 mm, dorsally with
a broad semiorbicular wing. Capsules not seen. Fl. NovJun of
following year.
Wet places in forests. Guangxi.
The protologue reported this species as distylous but described
only one floral form, similar to the short-styled flowers of distylous
species with the anthers positioned in or shortly above the corolla throat
and the stigmas situated near the middle of the corolla tube.

33. Ophiorrhiza liangkwangensis H. S. Lo, Bull. Bot. Res.,

Harbin 10(2): 39. 1990.
liang guang she gen cao
Herbs, procumbent to creeping, rooting at nodes, to 30 cm
tall; stems densely white- or pale yellow villous. Leaves usually
in unequal pairs; petiole 0.52(3) cm, densely hirtellous or villosulous; blade drying thinly papery or submembranous, broadly
ovate, ovate, or oblong-ovate, 1.58(11) 16 cm, sparsely
hirtellous to villous on both surfaces, base subrounded to obtuse, usually inequilateral, apex acute to obtuse; secondary
veins 611 pairs; stipules caducous, not seen. Inflorescence
congested-cymose, 2- or 3(6)-flowered, densely villosulous
to villous; peduncle 0.51.2 cm; bracts linear or filiform, 1.5
10 mm. Flowers distylous, subsessile or on pedicels to 3 mm.
Calyx villosulous; hypanthium subglobose, ca. 1.5 mm; lobes
filiform, ca. 2 mm, with 1 globose gland in each sinus. Corolla
white or pale purple, drying yellow, slenderly funnelform,
sparsely villous to glabrescent outside; tube 1820 mm,
sparsely pilose inside in basal 1/3; lobes ovate-triangular, ca. 6
mm, dorsally ribbed, apex rostrate. Capsules mitriform, ca. 3
7 mm, densely villous. Fl. Jun.
Roadsides at forest margins. Guangdong (Xinyi), Guangxi.
The protologue detailed both long-styled and short-styled flowers
and noted that their calyces and corollas are similar. H. S. Lo in FRPS
(71(1): 142. 1999) described this species as rarely many flowered, but
this has not been noted by other authors nor seen on specimens studied.

34. Ophiorrhiza lignosa Merrill, Brittonia 4: 176. 1941.

mu jing she gen cao

RUBIACEAE

Subshrubs, erect, to 50 cm tall; stems weakly 4-angled to

terete, glabrous except with 2 pilosulous to hirtellous lines.
Leaves in subequal pairs; petiole 0.62 cm, glabrous or villosulous; blade drying papery or submembranous, adaxially olivegreen, pale abaxially, lanceolate or narrowly lanceolate, 511
12 cm, glabrous on both surfaces or sometimes pilosulous
along principal veins abaxially, base long cuneate, margins
entire, apex acuminate; secondary veins 8 or 9 pairs; stipules
caducous, not seen. Inflorescence cymose to somewhat congested-cymose, many flowered, 34.5 cm, subferruginous pilosulous; peduncle 12 cm; axes helicoid, up to 1 cm; bracteoles ensiform-linear or linear and acuminate, 1.53 mm. Flowers with biology unknown, subsessile or on pedicels to 1 mm.
Calyx hispidulous; hypanthium compressed turbinate, ca. 1.3
mm, 5-ribbed; lobes ovate-triangular, 0.50.7 mm, usually with
1 gland in each sinus. Corolla purple, subtubular, glabrous outside, inside with white villous ring in upper part of tube and
onto lobes; tube 1011 mm; lobes broadly ovate or oblongovate, ca. 3 mm. Capsules unknown. Fl. Apr.
Forests; ca. 1100 m. Yunnan (Malipo) [Myanmar].
The protologue reported this species only from Myanmar, at ca.
1200 m, and did not detail the position of the anthers but did describe
the style as long enough to position the stigmas in the corolla throat; H.
S. Lo (in FRPS 71(1): 152153, t. 37, f. 4. 1999) described and illustrated the anthers as positioned near the middle of the corolla tube.
Thus, the flowers described resemble the long-styled flowers of distylous species; none of these authors have posited the floral biology of
this species.

35. Ophiorrhiza longicornis H. S. Lo, sp. nov.

chang jiao she gen cao
Type: China. Guangxi: Napo, Y. Lin 3-5324 (holotype,
GXMI).
Validating Latin description: that of Ophiorrhiza longicornis Lo (H. S. Lo, Bull. Bot. Res., Harbin 10(2): 57. 1990).
Herbs, apparently ascending; stems drying brownish red,
terete, glabrous. Leaves in somewhat unequal pairs; petiole 12
cm, glabrous; blade drying papery or rather thickly papery,
abaxially yellowish or greenish yellow with veins brown, narrowly elliptic-ovate to oblanceolate, 410 1.53.5 cm, glabrous on both surfaces, base cuneate, margins entire, apex obtuse then cuspidate or caudate; secondary veins 711 pairs;
stipules caducous, not seen. Inflorescences cymose to subcorymbose, many flowered, branched to several orders, puberulent; peduncles 1.52.5 cm; axes spiciform to helicoid, principal
ones ca. 1 cm; bracteoles turning purple when dry, linear-lanceolate, 35 mm, sharply acute. Flowers with biology unknown,
subsessile. Calyx with hypanthium compressed subglobose-turbinate, ca. 1.5 mm, 5- or 10-ribbed, puberulent; lobes triangular,
0.71 mm, subglabrous, with 1 gland in each sinus. Corolla
purplish red, tubular, glabrous and slenderly 5-ribbed outside;
tube ca. 4.5 mm, inside with white villous ring below middle;
lobes subtriangular, 0.40.5 mm, dorsally with horn 1.52 mm
and drying black, apex incurved-rostrate. Capsules not seen. Fl.
Apr.
Forests. Guangxi (Napo).

273

This name was previously published by H. S. Lo (loc. cit.) but not

validly so because the type was not indicated in accordance with Art.
37.6 of the Vienna Code.

36. Ophiorrhiza longipes H. S. Lo, Bull. Bot. Res., Harbin

10(2): 49. 1990.
chang geng she gen cao
Herbs, apparently ascending, to 100 cm tall; stems drying
flattened, densely pilosulous. Leaves in markedly unequal pairs;
petiole 0.51(3) cm, pilosulous; blade drying thinly papery,
adaxially grayish green, abaxially pale or purple, ovate or elliptic, larger ones 3.57.5 23.2 cm, smaller ones 25 1.53
cm, glabrous adaxially, abaxially glabrous or pilosulous along
veins, base cuneate, obtuse, or subrounded, inequilateral, margins subentire, apex obtuse; secondary veins 68 pairs; stipules
caducous, not seen. Inflorescence cymose, 59-flowered, often
rather lax, hispidulous to hirsute; peduncle 22.5 cm; axes helicoid, ca. 1 cm; bracteoles subulate, 23 mm. Flowers with biology unknown, subsessile. Calyx pilosulous or hispidulous; hypanthium turbinate, ca. 2.5 mm, 5-ribbed; lobes triangular, 1
1.3 mm, with 1 gland in each sinus. Corolla yellowish white or
tinged with purple, funnelform, glabrous outside; tube ca. 16
mm, with pilose or pilosulous ring above middle inside; lobes
ovate-triangular, 55.5 mm, dorsally thinly winged, apex rostrate. Capsules drying brownish red, mitriform-rhomboid, ca. 3
7 mm. Fl. Apr.
Rocks in wet and shady places in forests. Guangxi.
The figure of this species presented by H. S. Lo in FRPS (71(1):
150, t. 36, f. 16. 1999) seems to be redrawn based on the protologue
figure but differs a bit from it; in particular, the lower right-hand leaf is
shown in FRPS as acute or shortly acuminate, while in the protologue
figure this was shown as obtuse and there corresponded to the textual
description. Neither the protologue nor FRPS posited the floral biology
of this species; the flower described in the protologue resembles the
short-styled form of distylous species, with the anthers positioned in or
just below the corolla throat and the stigmas positioned just below the
middle of the corolla tube.

37. Ophiorrhiza luchuanensis H. S. Lo, Bull. Bot. Res., Harbin 10(2): 74. 1990.
l chun she gen cao
Herbs or subshrubs, weak to ascending, to 2 m tall; stems
densely hispid. Leaves in markedly unequal pairs; petiole 0.32
cm, densely hispid or pilose; blade drying papery, lanceolate to
narrowly elliptic, larger ones 515 2.54.5 cm, smaller ones
26(8) 12 cm, strigose adaxially, densely hirtellous or hirsute along principal veins abaxially, base acute to attenuate,
apex acuminate to caudate; secondary veins 712 pairs, prominent adaxially; stipules caducous. Inflorescence cymose, many
flowered, densely villous; peduncle 13 cm; axes several, helicoid; bracts absent or minute. Flowers distylous, subsessile.
Calyx glabrous or ferruginous mealy puberulent; hypanthium
rhombic-turbinate, ca. 1 mm, 5-ribbed; lobes subtriangular, ca.
0.4 mm. Corolla white, salverform-tubular and slightly contracted in middle, glabrous outside; tube 78 mm, barbate in
throat; lobes triangular, 1.82 mm, dorsally costate to narrowly
winged near apex. Capsules drying reddish brown, broadly
mitriform, ca. 1.5 5 mm, 5-ribbed, subglabrous. Fl. Oct.

RUBIACEAE

274

Shady wet places in forests; ca. 2000 m. S Yunnan (Lchun).

The protologue described the corollas of both floral forms as similar and noted that the anthers and stigmas are reciprocally placed, at or
just below the throat or at the bottom of the corolla tube, respectively.
The protologue figure was apparently redrawn for FRPS (71(1): 170, t.
43, f. 17. 1999) and differs from the first version: in particular, the
stems, petioles, and inflorescence are depicted as densely hispid or pilose in the protologue figure but apparently glabrous in the redrawn figure, and the redrawn figure shows the secund-helicoid flower arrangement on the inflorescence axes as significantly more regular than in the
protologue figure.

38. Ophiorrhiza lurida J. D. Hooker, Fl. Brit. India 3: 82.

1880.
huang he she gen cao
Herbs, procumbent to ascending, to 20 cm tall; stems puberulent to densely villosulous. Leaves in subequal pairs; petiole (0.1)0.22.8 cm, densely villosulous; blade drying papery,
adaxially green, abaxially pale green, ovate, ovate-lanceolate,
or elliptic, 0.65 0.62.5 cm, adaxially sparsely hispidulous to
hispid, abaxially puberulent or hispid along principal veins,
base obtuse, truncate, or subrounded, margins entire, apex obtuse to acute; secondary veins 4 or 5 pairs; stipules generally
persistent, triangular to ovate, 24 mm, glabrous to hirtellous,
acuminate or aristate with 13 bristles 27 mm. Inflorescences
congested-cymose to subcapitate, few to several flowered, glabrous; peduncles 0.54 cm; heads ca. 1 1 cm; bracts subulatelinear, lanceolate, elliptic-oblong, or spatulate, 38 mm, costate,
obtuse to acute. Flowers with biology unknown, subsessile or
on pedicels to 1.5 mm. Calyx glabrous; hypanthium subglobose, ca. 1.5 mm; lobes narrowly triangular, ca. 1 mm. Corolla
white, drying yellow, tubular to inflated, glabrous outside, inside
villous in upper part of tube and onto lobes; tube 2.54.5 mm;
lobes triangular-ovate, 11.5 mm, dorsally narrowly winged, obtuse. Capsules not seen in China. Fl. Aug.
Broad-leaved forests, Tsuga forests; [300]18002300 m. Xizang
(Mdog), NW Yunnan [India (Darjeeling, Sikkim)].
Deb and Mondal (Bull. Bot. Surv. India 39(14): 67. 1997) described the capsules of Indian plants as flattened obturbinate, 1.53 5
8 mm, and glabrous to puberulent.

39. Ophiorrhiza macrantha H. S. Lo, Bull. Bot. Res., Harbin

10(2): 28. 1990.
da hua she gen cao
Herbs, weak at base, ascending to erect above, to 50 cm
tall; stems drying orange, subglabrous or puberulent when
young. Leaves in unequal pairs; petiole 14 cm, glabrous or
puberulent; blade drying thinly papery or papery, adaxially
grayish green, pale or yellowish green abaxially, oblong-ovate,
subovate, or elliptic-oblong, 416 1.84.5 cm, glabrous on
both surfaces or abaxially puberulent along principal veins,
base obtuse to subrounded, margins usually undulate, apex caudate; secondary veins 79 pairs; stipules generally persistent,
narrowly triangular, 68 mm, with globose gland at apex and at
each side of base. Inflorescences cymose, many flowered, puberulent; peduncle 22.5 cm; axes helicoid, 0.52 cm; bracteoles linear-spatulate, ca. 2 mm, ciliolate. Flowers distylous, on

pedicels 1.53 mm. Calyx puberulent to glabrescent; hypanthium rhomboid-subglobose, ca. 2.5 2 mm, 10-ribbed; lobes
triangular, subequal or unequal, 1.52 mm. Corolla reddish or
pink, funnelform, glabrous outside, inside with sparsely white
villous ring at middle of tube and sparsely villous above this
and sometimes onto lobes; tube 2223 mm; lobes subovate, ca.
5.5 mm, dorsally with wing ca. 0.8 mm wide with very short
apical spur, apex rostrate. Immature capsules submitriform, ca.
3 77.5 mm, 10-ribbed. Fl. Mar.
Streamsides in dense forests; ca. 3000 m. Yunnan.
In the protologue and FRPS (71(1): 129131. 1999), H. S. Lo
noted that the long-styled flowers have the anthers positioned near the
middle of the corolla tube and the stigmas in its throat, while the shortstyled flowers have the anthers positioned in the throat and the stigmas
below them inside the upper part of the corolla tube.

40. Ophiorrhiza macrodonta H. S. Lo, Bull. Bot. Res., Harbin

10(2): 25. 1990.
da chi she gen cao
Large herbs or subshrubs, erect, to 2.5 m tall; stems drying
black to brownish red, ferruginous pilosulous to glabrescent.
Leaves in subequal pairs; petiole 12 cm or sometimes longer;
blade drying papery, adaxially grayish brown, abaxially pale,
oblong-elliptic, ovate-oblong, or ovate, 6.517 2.55.5 cm,
glabrescent on both surfaces or sparsely pilosulous adaxially
and pilosulous along principal veins abaxially, base cuneate
then decurrent, margins entire, apex acuminate; secondary veins
1416 pairs; stipules ovate to lanceolate-ovate, 516 mm, entire
or occasionally dentate, parallel-nerved, subglabrous, acuminate. Inflorescence cymose to congested-cymose, many flowered, pendulous then becoming erect, densely ferruginous
hirsute or -hirtellous; peduncle ca. 1 cm; bracts sublinear, 7
9 mm, ciliolate, glabrous or sparsely pilose, persistent. Flowers reportedly distylous, on pedicels 11.5 mm. Calyx hispidulous; hypanthium obrhombic, ca. 1 mm, weakly 5-ribbed; lobes
narrowly lanceolate, ca. 2.5 mm. Corolla greenish yellow in
bud, reddish at anthesis, tubular-funnelform, glabrous outside;
tube 1314 mm, villous inside; lobes subovate, ca. 2 mm, dorsally with wing to 0.6 mm wide, apex rostrate. Capsules obcordate, 2.53 7.59 mm, hispidulous. Fl. Sep.
Wet places in forests; ca. 1500 m. Yunnan.
In the protologue and FRPS (71(1): 126127. 1999), H. S. Lo
described the flowers as distylous with the long-styled flowers unknown;
the putative short-styled flowers were described as having the anthers
partially exserted in the throat and the stigmas situated below the middle
of the corolla tube.

41. Ophiorrhiza medogensis H. Li, Acta Phytotax. Sin. 18:

116. 1980.
chang e she gen cao
Herbs or subshrubs, ascending, to 60 cm tall; stems
densely brown villous with multicellular trichomes. Leaves in
unequal pairs; petiole 0.51.5(3.5) cm, densely villous; blade
drying thinly papery, pallid green on both surfaces or reddened
abaxially, lanceolate, oblong-lanceolate, or subovate, 614
2.25.5 cm, adaxially glabrous or sparsely hispid, abaxially

RUBIACEAE

densely villous along principal veins, base rounded or subcordate, usually inequilateral, margins subentire or undulate, ciliate, apex abruptly acuminate; secondary veins 816 pairs; stipules generally persistent, with 2 or 3 subulate to filiform lobes
ca. 1 cm, ciliate. Inflorescences congested-cymose, many flowered, densely villous; peduncles shorter than 1 cm; axes short,
helicoid; bracts filiform or subulate, 12 cm, ciliate. Flowers
with biology unknown, subsessile. Calyx densely villous; hypanthium subglobose, ca. 2 mm; lobes filiform, 78 mm, ciliate. Corolla white, tubular-funnelform, puberulent outside; tube
ca. 18 mm, in throat notably expanded, inside villous above
middle; lobes oblong-ovate, ca. 4 mm. Capsules obovate, ca.
5 10 mm. Fl. Sep.
Broad-leaved forests on mountain slopes; ca. 1700 m. Xizang
(Mdog).

275

cm, glabrous to puberulent; blade drying thinly papery, elliptic

or lanceolate-elliptic, 2.522 19 cm, glabrous adaxially, glabrous or puberulent on principal veins abaxially, base cuneate to
obtuse, apex acute to acuminate; secondary veins 1019 pairs;
stipules deciduous, narrowly triangular, 48 mm, sometimes 2lobed, glabrescent. Inflorescence cymose to paniculate, several
to many flowered, puberulent; peduncle 0.56 cm; axes helicoid; bracts reduced and caducous or none. Flowers apparently
distylous, subsessile. Calyx densely puberulent; hypanthium
compressed turbinate, ca. 1 mm; lobes triangular, 0.51 mm.
Corolla white, tubular or sometimes constricted near middle,
outside puberulent to glabrous; tube 2.54 mm, inside villosulous near middle; lobes triangular, ca. 1 mm, dorsally smooth to
costate. Capsules broadly mitriform, 1.53 39 mm, glabrous
to densely puberulent.

42. Ophiorrhiza mitchelloides (Masamune) H. S. Lo, Bull.

Bot. Res., Harbin 18: 277. 1998 [michelloides].

Expected in Yunnan [India, Myanmar, ?Philippines, Thailand to

Malaysia, ?Vietnam].

dong nan she gen cao

This species was not included for China by H. S. Lo (Bull. Bot.

Res., Harbin 10(2): 182. 1990; FRPS 71(1): 110174. 1999) but has
been reported from the surrounding regions (Deb & Mondal, Bull. Bot.
Surv. India 39(14): 6773. 1997), and some specimens from China
have been identified with this name by Chinese botanists (in herb.). The
description here is presented at least for reference and is based on that of
Deb and Mondal (excluding Ophiorrhiza mungos var. nemorosa
(Thwaites) J. D. Hooker). Deb and Mondal (loc. cit.) reported O. mungos to flower and fruit throughout the year and did not describe its floral
biology or individual floral forms but illustrated both short-styled and
long-styled flowers.

Hayataella mitchelloides Masamune, Trans. Nat. Hist. Soc.

Formosa 24: 206. 1934 [michelloides]; Geophila exigua H.
L. Li; Ophiorrhiza exigua (H. L. Li) H. S. Lo.
Herbs, procumbent to creeping, generally rooting at most
nodes; stems villous with multicellular trichomes. Leaves in
subequal to unequal pairs; petiole 0.31.3 cm, densely villous to
villosulous; blade drying papery, adaxially dark gray or nearly
black, abaxially purplish red to reddish or rarely pale, broadly
ovate, ovate, or suborbicular, 0.82.5 0.62 cm, sparsely pilose to villous along principal veins to throughout on both surfaces, base subtruncate to obtuse or rounded, apex acute to
rounded-obtuse; secondary veins 35 pairs; stipules usually
caducous, triangular to ligulate, 11.5 mm, glabrescent, acuminate to 24-lobed. Inflorescences fasciculate to cymose, 1- or
2(or 5)-flowered, villosulous; peduncles and/or pedicels 0.42
cm; bracts linear, 48 mm. Flowers distylous, pedunculate or
pedicellate. Calyx with hypanthium compressed globose, ca.
1.2 mm, densely villosulous; lobes linear, ca. 1.4 mm, glabrescent or ciliate. Corolla white, funnelform or salverform, outside
with 5 strigose or hispidulous lines; tube ca. 15 mm, inside with
white villous ring just above middle and otherwise villous or
glabrous except pubescent in throat; lobes broadly ovate, 56.5
mm. Capsules obcordate, ca. 3.5 910 mm, villous. Fl. Apr,
fr. Jun.
Broad-leaved forests or forest margins; 4001500 m. Fujian,
Guangdong, Hunan, Jiangxi, Taiwan.
This small plant was separated by some authors in a monotypic
genus endemic to Taiwan, Hayataella. However, recent authors have
noted a broader range for it and included it in Ophiorrhiza based on
morphological (H. S. Lo, Bull. Bot. Res., Harbin 10(2): 182. 1990, and
references cited therein) and molecular (Nakamura et al., J. Plant Res.
119: 657661. 2006) characters. H. S. Lo in FRPS (71(1): 141142.
1999) detailed both short-styled and long-styled flowers.

44. Ophiorrhiza mycetiifolia H. S. Lo, Guihaia 11: 103. 1991.

xian mu ye she gen cao
Herbs or subshrubs, erect, to 2.5 m tall; stems puberulent
to villosulous. Leaves in generally equal pairs; petiole 24 cm,
densely villosulous; blade drying thinly papery, olive-green,
elliptic-ovate or elliptic, 1020 47.5 cm, sometimes weakly
bullate, adaxially glabrous or sparsely strigillose to puberulent, abaxially glabrous to densely villosulous, base obtuse to
rounded then shortly decurrent, margins entire, apex acute to
shortly cuspidate; secondary veins 15 or 16 pairs; stipules caducous, unknown. Inflorescence congested-cymose, many flowered, 56 cm wide, densely villosulous-tomentulose; peduncle
ca. 2.5 cm; axes several, dichasial at lower axes then helicoid
above; bracts linear-lanceolate, 510 mm, acuminate. Flowers
with biology unknown, subsessile or on pedicels to 2 mm. Calyx densely hirtellous; hypanthium subglobose, ca. 1.5 mm, 5ribbed with ribs glabrescent; lobes triangular, 0.81 mm. Corolla white, tubular, outside glabrous to pilosulous, pubescent
throughout inside; tubes 1011 mm; lobes subtriangular, ca. 2
mm, reflexed, dorsally with horn 1.52 mm near apex. Capsules not seen. Fl. Dec.
Sparse forests in valleys; ca. 600 m. Guangxi (Longzhou).

43. Ophiorrhiza mungos Linnaeus, Sp. Pl. 1: 150. 1753.

45. Ophiorrhiza nandanica H. S. Lo, Bull. Bot. Res., Harbin

10(2): 63. 1990.

she gen cao

nan dan she gen cao

Herbs or subshrubs, ascending, to 100 cm tall; stems glabrescent to puberulent. Leaves in subequal pairs; petiole 1.55

Herbs, suberect, to 40 cm tall; stems drying black to

brownish yellow, terete, glabrous. Petiole 12.5 cm, glabrous;

RUBIACEAE

276

leaf blade drying papery, dark gray or blackish gray adaxially,

pale yellow or greenish yellow with veins brown abaxially,
ovate to lanceolate, 411 1.54.5 cm, glabrous, base obtuse,
apex acuminate; secondary veins 59 pairs; stipules caducous,
not seen. Inflorescences congested, several flowered, pubescence in 2 pilose longitudinal lines; peduncle 12 cm; axes
helicoid; bracts linear-lanceolate, 35 mm, weakly pinnately
veined. Flowers distylous, sessile or subsessile. Calyx with
hypanthium submitriform, ca. 1.5 mm, 5-ribbed; lobes triangular or narrowly triangular, markedly unequal, 1 or 2 larger
ones 12 mm, 3 or 4 smaller ones ca. 0.7 mm, both lobes and
sinuses glandular. Corolla pale yellow, subsalverform, glabrous
outside; tube ca. 18 mm, glabrous inside; lobes long narrowly
triangular, ca. 1.5 mm, dorsally with wing ca. 1 mm wide with
very short horn at top, apex rostrate. Capsules not seen. Fl. Oct.
Shady places in forests in limestone regions. Guangxi (Nandan).
In the protologue and FRPS (71(1): 162. 1999), H. S. Lo detailed
both long-styled and short-styled flowers, with similar corollas but differing in anther and stigma positions (in the middle of the corolla tube
vs. the throat, respectively and correspondingly).

46. Ophiorrhiza napoensis H. S. Lo, Bull. Bot. Res., Harbin

10(2): 48. 1990.
na po she gen cao
Herbs, ascending, to 30 cm or taller; stems glabrous.
Leaves in unequal pairs; petiole 0.31.4 cm, glabrous; blade
drying papery, grayish adaxially, pale yellow abaxially, narrowly lanceolate to subovate, 512 1.53.5 cm, glabrous on
both surfaces or pilosulous along midrib abaxially, base cuneate
to obtuse, margins entire, apex acute to acuminate; secondary
veins 712 pairs; stipules caducous, triangular, 0.51 mm, entire or glandular. Inflorescences congested-cymose, several
flowered, ferruginous- to golden yellow tomentulose or -pilosulous; peduncle 0.81 cm; axes short, helicoid; bracts lanceolatelinear, 1014 mm, weakly costate, glabrous. Flowers with biology unknown, subsessile or on pedicels to 1.5 mm. Calyx with
hypanthium turbinate-rhombic, 1.52 2.32.5 mm, 5-ribbed,
densely pilosulous; lobes narrowly triangular, unequal, largest
ones 1.82 mm, smallest ones 1.31.5 mm, glabrous, sinus
sometimes with 2 or 3 globose glands. Corolla white, drying
orange-yellow, salverform to funnelform, glabrous outside; tube
2022 mm, glabrous inside; lobes ligulate to ovate, 2.55 mm,
dorsally ribbed and with very short horn. Immature fruit obcordate. Fl. Oct.
Forests on hill slopes. Guangxi (Napo), Yunnan (Maguan).

47. Ophiorrhiza nutans C. B. Clarke ex J. D. Hooker, Fl. Brit.

India 3: 84. 1880.
chui hua she gen cao
Herbs, weak to erect, to 70 cm tall; stems densely reddish
brown villous with multicellular trichomes. Leaves in subequal
to unequal pairs; petiole 0.52 cm, densely pubescent; blade
drying papery, ovate, lanceolate, elliptic, or elliptic-oblong,
38(13) 1.54 cm, sparsely pilose or strigose adaxially, glabrescent except pubescent along principal veins abaxially, base
obtuse to subrounded then attenuate, margins ciliolate, entire or
undulate, apex acute to cuspidate; secondary veins 913[15]

pairs; stipules persistent, lanceolate or subovate, 810 mm, acuminate to 2-lobed. Inflorescence congested-cymose to corymbiform, many flowered, densely hirtellous; peduncle 13[6] cm;
bracts linear-oblong or lanceolate, 510 mm, ciliate, sometimes
fimbriate and/or with a linear lobe at each side. Flowers distylous, subsessile. Calyx densely hispidulous to glabrescent; hypanthium subturbinate, ca. 1.5 mm; lobes sublanceolate, ca. 1.2
mm. Corolla white, tubular-funnelform, glabrous or sparsely
hispid outside; tube [6.5]9[19] mm, inside sparsely pilosulous near middle and barbate in throat; lobes subtriangular or
subovate, 23 mm, dorsally narrowly winged. Capsules mitriform, ca. 2.5 67 mm, hispidulous to subglabrous.
Moist forests; 7002400 m. Xizang (Mdog), Yunnan [NE India,
Myanmar, Nepal].
Measurements in brackets are taken from the description of this
species by Deb and Mondal (Bull. Bot. Surv. India 39(14): 82. 1997)
and may be expected in Chinese plants. H. S. Lo (in FRPS 71(1): 131.
1999) and Deb and Mondal noted that this species is distylous, with the
corollas similar in size in both floral forms and the stigmas and anthers
reciprocally placed in the corolla throat and near the base of the corolla
tube, respectively. H. S. Lo described the corolla tubes as ca. 9 mm; the
Fl. Bhutan (2(2): 779. 1999) described them as 6.58.5 mm; and Deb
and Mondal described them as 1319 mm.
Ophiorrhiza hainanensis was synonymized with O. nutans by
Duan and Lin (Acta Phytotax. Sin. 45: 877. 2007). However, several
distinctions between these were cited by H. S. Lo (Bull. Bot. Res., Harbin 10(2): 30. 1990), and these species are provisionally separated here
pending further study.

48. Ophiorrhiza ochroleuca J. D Hooker, Fl. Brit. India 3: 78.

1880.
huang hua she gen cao
Herbs, annual or perennial, or subshrubs, erect, to
40[100] cm tall; stems [glabrous to] pubescent. Leaves in subequal pairs; petiole 14.5 cm, subglabrous; blade drying thinly
papery, adaxially pale green, abaxially paler, elliptic to lanceolate, [3.5]1315[22.5] [1.5]4.56[10] cm, subglabrous
except pilosulous abaxially along principal veins, base cuneate
to obtuse, margins entire, apex shortly acuminate to subacute;
secondary veins [8]1012[15] pairs; stipules deciduous, subovate or triangular, [4]5[15] mm, at apex with 1 gland. Inflorescence cymose, many flowered, branched to several orders,
pilosulous; peduncle ca. 3[10] cm; axes helicoid, 13[10]
cm; bracts minute, caducous. Flowers subsessile or on pedicels
to 3 mm. Calyx puberulent to pilosulous; hypanthium subturbinate, ca. 3 mm, 5-ribbed; lobes triangular, ca. 0.5 mm. Corolla
yellow to pale yellow, tubular, outside glabrous and 5-ribbed in
upper part; tube 5.56.5[12] mm, glabrous inside; lobes ca. 1
mm. Capsules mitriform, ca. 3 [5]7.5 mm, puberulent to glabrescent.
Wet places in forests; 3002000 m. Yunnan [Bhutan, NE India,
Myanmar].
Measurements included in brackets are from the description of
this species by Deb and Mondal (Bull. Bot. Surv. India 39(14): 8486.
1997). Their description conflicts with that of the Fl. Bhutan (2(2): 777.
1999), which distinguished this species by its leaves glabrous below
while Deb and Mondal considered the pubescent lower leaf veins distinctive.

RUBIACEAE

277

49. Ophiorrhiza oppositiflora J. D. Hooker, Fl. Brit. India 3:

80. 1880.

51. Ophiorrhiza petrophila H. S. Lo, Bull. Bot. Res., Harbin

10(2): 66. 1990.

dui sheng she gen cao

fa dou she gen cao

Herbs, weak to ascending, to 70 cm tall; stems generally

terete, hispidulous to glabrescent. Leaves in subequal pairs;
petiole 12 cm, subglabrous or puberulent; blade drying papery, brown, narrowly elliptic, ovate, or lanceolate, [3]815
[1]3.56 cm, subglabrous or sparsely strigillose adaxially,
pilose along principal veins abaxially, base cuneate then narrowed and shortly decurrent, apex shortly acuminate; secondary
veins 911(15) pairs; stipules caducous or persistent on upper
nodes, linear or bifid, 211 mm, puberulent. Inflorescence terminal and/or paired in upper stem nodes, cymose to paniculate,
many flowered, ferruginous puberulent; peduncle 24 cm; axes
helicoid; bracts minute, caducous. Flowers with biology unknown, subsessile. Calyx pubescent; hypanthium ellipsoid, ca.
1 mm; lobes triangular, ca. 0.3 mm. Corolla white, tubular-funnelform, outside glabrous; tube ca. 4[8] mm, inside glabrous
or densely barbate in throat; lobes oblong-triangular, ca. 2 mm,
dorsally ridged, apex rostrate. Capsules drying yellow, mitriform-obcordate, ca. 2 67 mm, subglabrous. Fl. winter and
spring.

Herbs, ascending, to 100 cm tall; stems drying dark brown

or purplish black and shiny, slightly compressed to subterete,
glabrous. Leaves in subequal pairs; petiole 0.51 cm, glabrous;
blade drying papery, grayish green adaxially, pale abaxially,
ovate or narrowly ovate, 27(9) 1.53(3.5) cm, glabrous,
base cuneate to obtuse, margins entire, apex cuspidate to shortly
caudate; secondary veins 79 pairs; stipules caducous, not seen.
Inflorescence 35 cm, many flowered, pilosulous to glabrescent; bracts filiform to subulate, 45 mm, obtuse, persistent.
Flowers reportedly distylous, subsessile or on pedicels to 1.5
mm. Calyx glabrous; hypanthium turbinate, ca. 1.5 mm, 5ribbed; lobes triangular, 0.71 mm, with 1 gland in each
sinus. Corolla reddened, tubular-funnelform, outside glabrous;
tube 1011 mm, sparsely villous to scabrous throughout inside;
lobes ovate-triangular, 1.51.8 mm, dorsally with broad lunate
wing. Capsules purple, ca. 2.5 8 mm. Fl. May.

Wet places in forests; [below 1001500 m in Myanmar]. Hainan,

Yunnan [NE India, Myanmar].
Deb and Mondal (Bull. Bot. Surv. India 39(14): 8687. 1997)
described this species as annual and the flowers as variously glabrous
inside or densely barbate in the throat. Measurements in brackets above
are taken from their description and may be expected in Chinese plants.

50. Ophiorrhiza pauciflora J. D. Hooker, Fl. Brit. India 3: 84.

1880.
shao hua she gen cao
Herbs, weak to erect, to 30 cm tall; stems pilosulous to
villosulous. Leaves in subequal pairs; petiole 0.52 cm, hirtellous; blade drying thinly papery, adaxially dark purple, abaxially purple or grayish, ovate to elliptic-ovate, 1.55.5 0.82.5
cm, adaxially strigose or sericeous, abaxially pubescent on principal veins, base obtuse to subrounded or sometimes shortly
decurrent, apex obtuse; secondary veins 59 pairs; stipules persistent, narrowly triangular or usually 2-lobed, 36 mm, lobes
narrowly triangular to linear, glabrescent. Inflorescence congested-cymose, several flowered, pubescent; peduncle 13 cm;
bracts linear, 36.5 mm, persistent. Flowers distylous, subsessile. Calyx puberulent to pilosulous or glabrescent; hypanthium
obovoid, ca. 1.5 mm; lobes sublanceolate, 0.81 mm. Corolla
white, tubular, outside hispid to glabrescent; tube 89 mm,
barbate in throat and upper part; lobes ovate, ca. 1 mm. Capsules obcordate, ca. 2 4.5 mm, puberulent to hirtellous or glabrescent. Fl. and fr. MayOct.
Dense forests; 6001600 m. Yunnan (Xishuangbanna) [NE India].
Deb and Mondal (Bull. Bot. Surv. India 39(14): 8993. 1997)
recognized two varieties, the pubescent var. pauciflora with corollas 6
9 mm and the glabrous var. glabra Deb and Mondal with corollas 1012
mm; the plants treated as Ophiorrhiza pauciflora by H. S. Lo (in FRPS
71(1): 127. 1999) generally fall within var. pauciflora but have corollas
that are intermediate in size and thus cannot be conclusively included in
either of the Indian varieties (suggesting that the Indian varieties may
not ultimately be separable).

Dense forests in limestone regions. Yunnan.

The protologue described this species as distylous but noted that
long-styled flowers had not been seen; the putative short-styled flowers
were said to have the anthers positioned in or shortly above the corolla
throat and the stigmas situated near the middle of the corolla tube.

52. Ophiorrhiza pingbienensis H. S. Lo, Bull. Bot. Res., Harbin 10(2): 20. 1990.
ping bian she gen cao
Herbs, ascending; stems drying reddish brown and usually
angled, densely reddish brown pilosulous at least when young.
Leaves in subequal pairs; petiole 0.52.5 cm, densely pilosulous; blade drying thinly papery, oblong-ovate, elliptic, oblonglanceolate, or ovate, 27.5 13 cm, adaxially sparsely puberulent, abaxially glabrescent to usually densely puberulent on
principal veins, base cuneate, margins entire, apex acute to obtuse; secondary veins 57 pairs; stipules persistent to deciduous, lanceolate to subovate, ca. 4 mm, entire or sparsely dentate, at base usually with 2 glands, acuminate and sometimes
with small globose gland at apex. Inflorescence congestedcymose, several flowered, densely reddish brown pilosulous;
peduncles 11.5 cm; axes relatively short, helicoid; bracts linear-lanceolate, spatulate, or lanceolate, 57 mm, often pinnately
veined, subglabrous or ciliate, obtuse to subacute. Flowers with
biology unknown, subsessile or on pedicels 1.53 mm. Calyx
with hypanthium turbinate, ca. 1.5 mm, 5-ribbed, densely pilosulous; lobes lanceolate, subovate, or spatulate, unequal, largest ones 36 mm, smallest ones 0.83 mm, pinnately veined,
ciliate or subglabrous. Corolla dark red, tubular-funnelform,
outside 5-winged in upper part in bud and glabrescent; tube 15
17.5 mm, inside with 5 small villous fascicles below throat and
just above anthers; lobes subovate to ligulate, 2.53 mm, inside
densely shortly scaly pubescent, dorsally with broad lunate
wing, apex rostrate. Immature capsule obcordate, ca. 2 5.5
mm, ferruginous pilosulous. Fl. Jul.
Streamsides; ca. 1400 m. Yunnan (Pingbian).

RUBIACEAE

278

This species was keyed by H. S. Lo (loc. cit.: 8, as qinbienensis)

based in part on its persistent, usually discernible stipule, although the
accompanying figure there shows no stipules, which suggests these may
be deciduous at least sometimes. As shown in the figure in the protologue, the pedicels described there could be considered inflorescence
axes producing subsessile flowers by others.

53. Ophiorrhiza pumila Champion ex Bentham, Hookers J.

Bot. Kew Gard. Misc. 4: 169. 1852.
duan xiao she gen cao
Ophiorrhiza aureolina H. S. Lo f. qiongyaensis H. S. Lo;
O. humilis Y. C. Tseng; O. inflata Maximowicz; O. pumila var.
inflata (Maximowicz) Masamune.
Herbs, weak to ascending, to 20(30) cm tall; stems drying
gray or grayish yellow, weakly rugose, densely tomentulosevillosulous. Leaves in subequal pairs; petiole 0.11.5 cm,
densely tomentulose; blade drying papery, adaxially green,
grayish green, or dark grayish brown, abaxially pale to red,
ovate, lanceolate, elliptic, or elliptic-oblong, (0.5)25.5(9)
(0.4)12.5 cm, adaxially subglabrous to sparsely strigillose or
hispidulous, abaxially densely puberulent to tomentulose or
sometimes glabrescent, base cuneate and generally decurrent,
margins entire to somewhat undulate, apex acute to obtuse, subacuminate, or rounded; secondary veins 58 pairs; stipules caducous, linear, 13 mm, puberulent. Inflorescences congestedcymose, several to many flowered, densely tomentulose-puberulent; peduncle 0.31.2 cm; branched portion 510 10
12 mm; axes becoming helicoid; bracts few, narrowly triangular
to linear, 0.32 mm. Flowers homostylous, subsessile or on
pedicels to 1.5 mm. Calyx with hypanthium subglobose, 0.8
1.2 mm, 5-ribbed, densely hispidulous except ribs glabrescent;
lobes subtriangular to linear, 0.30.6 mm, glabrescent. Corolla
white, tubular to inflated, outside puberulent to hispidulous;
tube 2.52.8 mm, with villous ring in throat to villosulous
throughout inside; lobes ovate-triangular, 1.21.5 mm, dorsally
weakly ribbed, acute to obtuse. Capsules drying brownish yellow, mitriform or somewhat obcordate, 22.5 57 mm, hispidulous. Fl. AprSep, fr. JunOct.
Shady places on wet lands, streamsides or riversides in forests;
200700 m. Fujian, Guangdong, Guangxi, Hainan, Jiangxi, Taiwan,
Yunnan [Japan, N Vietnam].
The floral biology of this species was studied by Nakamura et al.
(J. Jap. Bot. 81: 113120. 2006), who found the plants studied in the
Ryukyu Islands and Taiwan to be long-styled-monomorphic, not distylous as previously reported. They also observed very low pollination
rates in wild plants and concluded that probably at least much of the
reproduction in this species is through autogamy.
This species was reviewed recently by Duan and Lin (Acta Phytotax. Sin. 45: 878879. 2007), who newly synonymized several names
under Ophiorrhiza pumila but separated relatively smaller plants in O.
humilis. However, these smaller plants were included within the circumscription of O. pumila by H. S. Lo (in FRPS 71(1): 171. 1999), who is
followed here.

54. Ophiorrhiza purpurascens H. S. Lo in S. Y. Jin & Y. L.

Chen, Cat. Type Spec. Herb. China (Suppl.), 191. 1999.
zi mai she gen cao
Herbs, often procumbent at base, ascending above; stems

densely brown villous or -hirsute with multicellular trichomes.

Leaves in subequal pairs; petiole ca. 0.5 cm; blade drying papery, with veins purple, ovate, 13 0.51.7 cm, both surfaces
brown villous with pubescence denser along veins abaxially,
base rounded to obtuse and slightly oblique, margins entire,
apex acute to subobtuse; secondary veins 58 pairs; stipules
caducous, not seen. Inflorescences 1-flowered or fasciculate
and few flowered, densely pubescent; peduncles or pedicels
310 mm; bracts filiform, ca. 5 mm. Flowers with biology
unknown, pedicellate or pedunculate. Calyx with hypanthium
ca. 1.5 mm; lobes filiform, 2.53 mm. Corolla white, slenderly
subfunnelform, puberulent outside; tube 2223 mm, with white
villous ring at middle inside; lobes ovate-triangular, 34 mm,
obtuse, puberulent marginally, dorsally narrowly winged. Capsules not seen. Fl. Sep.
Shady places in ravines; ca. 1000 m. Sichuan (Xuyong).

55. Ophiorrhiza purpureonervis H. S. Lo, Bull. Bot. Res.,

Harbin 10(2): 42. 1990.
cang wu she gen cao
Herbs, apparently ascending, to 30 cm tall; stems glabrous.
Petiole 0.30.5(1) cm, glabrous; leaf blade drying papery,
adaxially leaden gray, abaxially straw-yellow with veins purple,
ovate, lanceolate, or broadly ovate, 27.5 14 cm, glabrous
on both surfaces, base rounded or obtuse, margins entire, apex
obtuse; secondary veins 6 or 7 pairs; stipules caducous, not
seen. Inflorescences many flowered, densely ferruginous villosulous; peduncle 1.52.5 cm; axes stout, helicoid; bracts filiform, 2.53 mm, ciliate or sparsely hispidulous. Flowers reportedly distylous, subsessile or on pedicels to 1 mm. Calyx
with hypanthium obovoid, ca. 1.5 mm, 5-ribbed, densely pubescent; lobes lanceolate, ca. 1.5 mm, ciliate. Corolla white or pale
yellow, slenderly funnelform, glabrous outside; tube ca. 10 mm,
inside with white villous ring at middle and puberulent above;
lobes sublanceolate, ca. 4 mm, dorsally with horn ca. 1 mm
near apex. Immature capsules obcordate, ca. 1.5 4 mm, pubescent. Fl. Jul.
Dense forests; 200500 m. Guangxi (Cangwu).
In the protologue and FRPS (71(1): 145. 1999), H. S. Lo noted
that this species is distylous but described only long-styled flowers, with
the stigmas borne near the corolla throat and the anthers positioned
below the middle of the corolla tube.

56. Ophiorrhiza rarior H. S. Lo in S. Y. Jin & Y. L. Chen, Cat.

Type Spec. Herb. China (Suppl.), 191. 1999.
mao guo she gen cao
Herbs, apparently ascending; stems moderately brown
villous or -hirsute with multicellular trichomes to glabrescent,
densely lenticellate when young. Petiole 46 cm, sparsely villous; leaf blade drying membranous-papery, adaxially olivegreen, abaxially pale, narrowly elliptic-oblong, 1321 4.55
cm, sparsely strigose adaxially, abaxially glabrescent except
sparsely hirsute along veins, base acute, margins subentire,
ciliate, apex acuminate or acute; secondary veins 1418 pairs;
stipules caducous, not seen. Inflorescences and flowers not
seen. Infructescences branched to several orders, villous; peduncle ca. 8.5 cm, 4-angled; axes 45.5 cm, helicoid; bracts

RUBIACEAE

lanceolate to linear-subulate, 48 mm, ciliate, persistent. Capsules mitriform, ca. 4 1011 mm, crisped villous. Fl. Jun.
Wet places in forests. Guangxi (Longzhou).

57. Ophiorrhiza repandicalyx H. S. Lo in S. Y. Jin & Y. L.

Chen, Cat. Type Spec. Herb. China (Suppl.), 191. 1999.
da ye she gen cao
Herbs, erect, to 100 cm tall; stems drying dark brown,
rather stout, glabrous. Leaves in subequal pairs; petiole 0.81.5
cm, glabrous; blade drying papery, adaxially grayish green,
abaxially pale green, broadly ovate, ovate, or elliptic, 1522
610 cm, glabrous on both surfaces, base obtuse then abruptly
narrowed, apex cuspidate; secondary veins 1116 pairs; stipules
caducous, not seen. Inflorescences congested-cymose to subcapitate, many flowered, ca. 3 cm in diam., mealy pubescent;
peduncle ca. 1.5 cm; axes helicoid; bracts absent. Flowers reportedly distylous. Calyx mealy pubescent; hypanthium broadly
turbinate, ca. 2.1 mm; limb tubular, ca. 1.5 mm, undulate to
shallowly 5-lobed. Corolla green, rather stoutly tubular, mealy
puberulent outside; tube ca. 5.5 mm, white villous above middle inside; lobes triangular-ovate, ca. 1.5 mm, dorsally broadly
winged and with horn ca. 0.5 mm near apex. Capsules unknown.
Fl. Nov.
Forests in ravines; ca. 1100 m. Yunnan (Jinghong).
The original description (H. S. Lo, Bull. Bot. Res., Harbin 10(2):
7980. 1990, not validly published) noted that the flowers are distylous
but described only a presumed long-styled form, with the anthers positioned near the middle of the corolla tube and the stigmas positioned at
or just above the corolla throat.

58. Ophiorrhiza rhodoneura H. S. Lo, Bull. Bot. Res., Harbin

10(2): 45. 1990.
hong mai she gen cao
Herbs, ascending, to 50 cm tall; stems subterete, rather
stout, densely ferruginous villous with multicellular trichomes.
Leaves in somewhat unequal pairs; petiole 14 cm, densely villous; blade drying subleathery-papery, with veins ferruginousred abaxially, ovate or elliptic, 5.512 2.56 cm, glabrous
adaxially, villous on principal veins abaxially, base subrounded,
apex acute; secondary veins 913 pairs; stipules caducous, not
seen. Inflorescence congested-cymose and rather umbelliform
to subcapitate, many flowered, densely villous; peduncles 33.5
cm; axes short, helicoid; bracts linear-spatulate, 1013(15)
mm. Flowers reportedly distylous, subsessile. Calyx densely
villosulous; hypanthium compressed turbinate; limb reduced,
denticulate. Corolla funnelform, outside pubescent; tube 24
27 mm, inside villous below middle; lobes subovate, 68 mm,
pinnately veined, dorsally with narrow ciliate wing, apex rostrate. Capsules obcordate-mitriform, ca. 4.5 810 mm, villosulous. Fl. Sep, fr. Nov.
Broad-leaved forests; ca. 1300 m. Guangxi (Napo).
In the protologue and FRPS (71(1): 148. 1999), H. S. Lo noted
that this species is distylous but described only putative short-styled
flowers, with the stigmas borne near the base of the corolla tube and the
anthers positioned just below the corolla throat.

279

59. Ophiorrhiza rosea J. D. Hooker, Fl. Brit. India 3: 78. 1880.

mei li she gen cao
Herbs or subshrubs, to 100[150] cm tall; stems drying
rugose, grayish yellow, pilosulous becoming glabrescent.
Leaves in subequal pairs; petiole 13(5) cm, hirtellous; blade
drying thinly papery, ovate, elliptic, or broadly ovate, [6.5]10
22[25] [2]510 cm, glabrous adaxially, densely hispidulous
along principal veins abaxially, base cuneate, obtuse, or subcordate, apex abruptly acute to acuminate; secondary veins 8
13(16) pairs; stipules deciduous, 2-lobed nearly to base, 57
mm, lobes subulate, glabrescent. Inflorescence cymose to paniculate, many flowered, puberulent; peduncle 2.85 cm; axes
helicoid; bracts reduced, caducous. Flowers with biology
unknown, subsessile. Calyx densely mealy puberulent; hypanthium compressed turbinate, ca. 1.5 mm, 5-ribbed; lobes subtriangular, ca. 0.5 mm. Corolla purplish red, tubular-funnelform, outside puberulent to glabrous, sometimes 5-ridged in
bud; tube 89 mm, inside glabrous; lobes subovate, ca. 1 mm.
Capsules broadly mitriform, ca. 3 8 mm, subglabrous or
mealy pubescent. Fl. OctDec.
Broad-leaved forests; 13002100 m. Xizang (Mdog), Yunnan
[Bhutan, NE India, Myanmar, Thailand].
Measurements in brackets are taken from the description of this
species by Deb and Mondal (Bull. Bot. Surv. India 39(14): 103105.
1997) and may be expected in Chinese plants.

60. Ophiorrhiza rufipilis H. S. Lo, Bull. Bot. Res., Harbin

10(2): 17. 1990.
hong mao she gen cao
Herbs, suberect to procumbent and rooting along lower
portion; stems drying striate, densely reddish brown villous
with multicellular moniliform trichomes. Leaves in markedly
unequal pairs; petiole 15 cm, densely villous; blade drying
papery, pallid green, oblong-lanceolate, lanceolate, elliptic, or
subovate, larger ones 310 14 cm, smaller ones 16 0.5
2.5 cm, adaxially sparsely strigose-villous, abaxially densely
pubescent on principal veins, base obtuse, subrounded, or occasionally subcordate, margins ciliate, apex acuminate or obtuse
and abruptly acuminate; secondary veins in larger leaves 1016
pairs, in smaller leaves 58 pairs; stipules generally persistent,
lanceolate-triangular or usually 2-lobed, 79 mm, sparsely ciliate, long acuminate. Inflorescence cymose, several to many
flowered, densely reddish brown villous; peduncle shorter than
1 cm; axes rather short, helicoid; bracts linear, ca. 5 mm, ciliate.
Flowers with biology unknown, subsessile or on pedicels to 1.5
mm. Calyx with hypanthium turbinate-subglobose, ca. 1.5 1.5
mm, densely villous; lobes subovate or lanceolate, 3.54 mm,
glabrescent, pinnately veined, hispid along margin and midrib,
acute to shortly rounded. Corolla white or pale yellow, funnelform, outside glabrescent with 5 hispid or villous lines on
ridges; tube 1819 mm, sparsely scaly pubescent inside; lobes
lanceolate to ovate, ca. 4.5 mm, dorsally with broad ciliate
wings, apex rostrate. Immature capsules subobcordate, ca. 3.5
7 mm, 5-ribbed, villosulous. Fl. Feb.
Dense forests; ca. 1200 m. SE Yunnan.

RUBIACEAE

280

61. Ophiorrhiza rufopunctata H. S. Lo, Bull. Bot. Res., Harbin 10(2): 15. 1990.

Evergreen forests; 17003400 m. Xizang, Yunnan (Gongshan)

[Bhutan, NE India, Nepal, Sri Lanka].

hong xian she gen cao

The description here is based mostly on the description by Deb

and Mondal (Bull. Bot. Surv. India 39(14): 107118. 1997); the elevational range is taken from Fl. Bhutan (2(2): 778. 1999). This species
was treated by Deb and Mondal as a widespread, morphologically
rather variable species; Fl. Bhutan noted that this species is variable and
not well circumscribed.

Herbs, weak at base, ascending above, to 15 cm tall; stems

brownish hispidulous or -strigose. Leaves in somewhat unequal
pairs; petiole 0.31.5 cm, pilosulous or hirsutulous; blade
drying papery, oblong-ovate, ovate, or broadly ovate, 38(10)
23.5 cm, adaxially glabrescent and usually with scattered
reddened gland dots, abaxially with red gland dots and glabrous
or hispidulous to strigose on veins, base obtuse to subrounded
or subcordate, often oblique, margins undulate, apex obtuse,
subrounded, or acute; secondary veins 4 or 5 pairs; stipules
persistent at least on upper nodes, subtriangular to ovate, 23
mm, obtuse and glandular at apex. Inflorescence 1-flowered or
usually cymose and 25-flowered, densely brown hispidulous;
peduncle ca. 1 cm; bracteoles linear-subulate, ca. 3 mm, usually
hirtellous and/or with reddish gland dots. Flowers distylous, on
short pedicels. Calyx hispidulous and/or with red gland dots;
hypanthium turbinate to ellipsoid, ca. 3 mm, 5-ribbed; lobes
linear, ca. 3 mm, each sinus with 1 gland. Corolla pale purple,
funnelform; tube 1822 mm, white villous or pilosulous above
middle and densely pilosulous at middle inside; lobes ovate, 4
5 mm, dorsally ridged and with very small horn. Capsules 56
ca. 11 mm, hispidulous. Fl. Nov.
Wet places in forests. Sichuan.
In the protologue and FRPS (71(1): 118. 1999), H. S. Lo noted
that the short-styled flowers have the anthers situated in the upper part
of the corolla tube and the stigmas situated near its middle, while the
long-styled flowers have the anthers situated near the middle of the
corolla tube and the stigmas in its throat.

62. Ophiorrhiza rugosa Wallich in Roxburgh, Fl. Ind. 2: 547.

1824.
pu di she gen cao
Ophiorrhiza harrisiana Heyne ex G. Don var. rugosa
(Wallich) J. D. Hooker; O. prostrata D. Don; O. prostrata var.
rugosa (Wallich) Panigrahi & S. K. Kar.
Herbs, sometimes annual, weak to erect, to 60 cm tall;
stems pilosulous to tomentulose. Leaves in subequal pairs; petiole 0.51.5(3) cm, puberulent to tomentulose; blade drying
thinly papery, ovate-lanceolate, lanceolate, or elliptic, 26(11)
13(5) cm, adaxially glabrous to sparsely strigillose or hispidulous, abaxially pilosulous or hispidulous along principal
veins, base cuneate to obtuse, apex caudate-acuminate, obtuse,
acute, or weakly acuminate; secondary veins 57(11) pairs;
stipules generally persistent on uppermost nodes, triangular
then contracted to linear, 410 mm, puberulent to glabrescent.
Inflorescence cymose to congested-cymose, several to many
flowered, pilosulous to glabrescent; peduncles 1.53 cm; axes
short to developed, becoming helicoid; bracts few, deciduous,
linear, 23 mm. Flowers with biology unknown, subsessile. Calyx puberulent; hypanthium ellipsoid, ca. 0.8 mm; lobes 0.21.5
mm. Corolla pink to white, tubular-funnelform, outside puberulent to glabrous; tube 36 mm, inside with pubescent ring near
or above middle; lobes triangular, 12 mm, dorsally smooth to
ridged. Capsules obconic, 22.5 45 mm, puberulent to glabrescent.

63. Ophiorrhiza salicifolia H. S. Lo, Bull. Bot. Res., Harbin

10(2): 50. 1990.
liu ye she gen cao
Subshrubs, apparently ascending, to 1 m tall; stems flattened to terete, subglabrous. Leaves in subequal pairs; petiole
0.51.5 cm, subglabrous; blade drying papery, red, lanceolatelinear, falcate, 4.511 0.61.5 cm, subglabrous on both
surfaces, base cuneate, apex subobtuse; secondary veins 911
pairs; stipules caducous, not seen. Inflorescences and flowers
not seen. Infructescences cymose-paniculate, 1012 ca. 8 cm,
branched to several orders, pubescent; peduncles 46.5 cm;
axes dichotomous to helicoid; bracteoles linear, 45 mm, subglabrous, persistent; pedicels 23 mm. Capsules mitriform, 2.5
3 78 mm, puberulent. Fr. May.
Wet fertile soil. Guangxi (Shangsi).

64. Ophiorrhiza sichuanensis H. S. Lo, Guihaia 11: 104. 1991.

si chuan she gen cao
Herbs, procumbent in lower portion, ascending above;
stems drying black, subglabrous or puberulent. Leaves in subequal pairs; petiole 0.82 cm, subglabrous; blade drying thickly
papery, adaxially leaden gray, abaxially dark brown, broadly
elliptic, 1.55 1.23.2 cm, glabrous on both surfaces, base
rounded to obtuse, margin irregularly dentate or rarely entire,
apex obtuse or acute; secondary veins 46 pairs; stipules generally persistent, subulate, 1.52 mm, acuminate. Inflorescences
1- or 2-flowered; bracts linear, 46 mm. Flowers with biology
unknown, on pedicels 68 mm. Calyx with hypanthium broadly
turbinate, ca. 2.5 mm, 5-ribbed; lobes linear, 3.54 mm, slightly
obtuse. Corolla purple, funnelform, outside glabrous; tube 26
29 mm, white villous inside; lobes ovate-triangular, ca. 6 mm,
apex rostrate. Capsules not seen. Fl. Apr.
On rocks; ca. 1200 m. Sichuan (Leibo).

65. Ophiorrhiza subrubescens Drake, J. Bot. (Morot) 9: 215.

1895.
bian hong she gen cao
Herbs, weak to suberect, to 60 cm tall; stems glabrous to
pubescent. Leaves in subequal pairs; petiole 0.52(4) cm,
densely pilose; blade drying thinly papery, red on both surfaces
or grayish green adaxially, lanceolate to ovate, 311 14 cm,
glabrous or sparsely strigose adaxially, ?pilose along principal
veins abaxially, base cuneate, margins entire or undulate, apex
rounded-obtuse to acuminate or acute; secondary veins 713
pairs; stipules sometimes caducous, broadly triangular then
strongly narrowed, 23 mm. Inflorescence congested-cymose,
many flowered, densely pubescent; peduncle 16 cm; axes heli-

RUBIACEAE

coid, up to 1 cm, elongating as fruit develop; bracts absent, minute, or caducous. Flowers reportedly distylous, subsessile. Calyx densely pubescent; hypanthium subobcordate, ca. 1 mm, 5ribbed; lobes subtriangular, ca. 0.5 mm. Corolla slenderly tubular, 68(12) mm, outside pubescent and 5-ribbed at least in
bud, inside with white villous ring at middle and scaly pubescent in throat and onto lobes; lobes triangular to subovate, dorsally narrowly winged and with very short horn. Capsules
drying purplish red, obcordate, ca. 3 78 mm, pubescent. Fl.
AprJul.
Shady and wet places in forests. Guangxi, Hainan, Yunnan [N
Vietnam].
The circumscription of this species here follows that of H. S. Lo in
FRPS (71(1): 139. 1999), which partly differs from that of other authors
and also conflicts with the protologue in some details. The protologue
described the plants as glabrous with filiform stipules and bracts that
fall before the fruit mature, while H. S. Lo said they are pubescent with
narrowly triangular stipules and lacking bracts. Lo described the flowers
as distylous with the corollas apparently similar in both floral forms but
unusual in anther position: the forms were described as differing in style
length, ca. 2 mm vs. ca. 6 mm, and anther size, ca. 2 mm vs. ca. 3 mm,
but with the anthers in both forms positioned near the base of the corolla, at the same height as the short-styled stigmas but well below the
long-styled stigmas. However, the figure in FRPS (p. 140, t. 33, f. 18)
differs from Los description in its depiction of well-developed bracts
that persist with the flowers, deeply bilobed stipules, corollas with only
1 ring of internal pubescence, glabrous fruit, and flower forms with the
anthers of the short-styled flowers exserted on well-developed filaments.

66. Ophiorrhiza succirubra King ex J. D. Hooker, Fl. Brit.

India 3: 82. 1880.
gao yuan she gen cao
Herbs to subshrubs, weak to erect, to 60[75] cm tall, usually drying partly to wholly red; stems glabrous or with pilosulous lines. Leaves in subequal pairs; petiole 0.52 cm, glabrous or subglabrous; blade drying thinly papery, lanceolateelliptic, ovate-elliptic, or elliptic-oblong, 511(20) 2.54(8)
cm, glabrous on both surfaces or sparsely pubescent adaxially,
base cuneate to obtuse, margins entire or undulate, apex acuminate to caudate; secondary veins 710 pairs; stipules caducous,
not seen. Inflorescence congested-cymose, many flowered, often
pendulous later becoming erect, glabrescent or axes sometimes
puberulent to pilosulous in lines; peduncle 11.5[3] cm; axes
short; bracts linear-lanceolate, 69 mm. Flowers perhaps homostylous, on pedicels 11.5 mm. Calyx glabrous to puberulent or
densely pilosulous; hypanthium submitriform, ca. 1.5 mm, 5(or
10)-ribbed; lobes ovate-triangular to lanceolate-triangular, usually slightly unequal, 1.41.8 mm, with 1 gland in each sinus.
Corolla pink or white, tubular-funnelform and swollen at base,
outside glabrous; tube 77.5[10] mm, inside glabrous or villous; lobes ovate, ca. 2.5 mm, dorsally narrowly keeled. Capsules mitriform, [23.5 69 mm], glabrous to puberulent or
pilosulous. Fl. JulOct.
Broad-leaved forests; ca. 2000 m or more. Guizhou, Xizang
(Mdog), Yunnan [Bhutan, NE India, Myanmar, Nepal].
This species has apparently been distinguished based primarily on
its distinctive purple-red drying color; it is characterized further in the
key to species here.

281

Measurements in brackets are taken from the description of this

species by Deb and Mondal (Bull. Bot. Surv. India 39(14): 120122.
1997), where this species is reported to flower throughout the year and
grow from middle elevations up to 2400 m. H. S. Lo (in FRPS 71(1):
145146. 1999) did not posit the floral biology but described the flowers similarly to Deb and Mondal as essentially homostylous, with the
anthers and stigmas both positioned near the middle of the corolla tube.

67. Ophiorrhiza umbricola W. W. Smith, Notes Roy. Bot.

Gard. Edinburgh 12: 217. 1920.
yin di she gen cao
Herbs, sometimes weak at base, ascending above, to 45(
100) cm tall; stems obtusely 4-angled, drying purple, glabrous
or subglabrous. Petiole 14 cm, subglabrous; leaf blade drying
membranous or thinly papery, adaxially dark green, abaxially
pale green or red, ovate, elliptic, or lanceolate, 915 36.5
cm, glabrous on both surfaces or sparsely strigillose adaxially,
base cuneate to obtuse, apex long acuminate or cuspidate; secondary veins 1014 pairs; stipules caducous, not seen. Inflorescences congested-cymose, many flowered, glabrous or reddish
brown hirtellous; peduncle 1.53 cm; axes helicoid. Flowers
reportedly distylous, subsessile or on pedicels to 1 mm. Calyx
glabrous; hypanthium submitriform, 1.52 mm; lobes with 1
gland in each sinus. Corolla red or purplish red, subtubular, glabrous outside; tube 2224 mm, white hirsute above middle
inside; lobes ovate-triangular, 3(6) mm. Capsules drying red,
mitriform, 1011 mm wide. Fl. Jun.
Dense forests; 20003000 m. Xizang (Mdog), Yunnan [Myanmar].
H. S. Lo (in FRPS 71(1): 151. 1999) described this species as
distylous but described only putatively long-styled flowers, with the
anthers positioned just above the middle of the corolla tube and the
stigmas apparently positioned just above them.

68. Ophiorrhiza wallichii J. D. Hooker, Fl. Brit. India 3: 79.

1880.
da guo she gen cao
Herbs, weak at base, ascending above, to 20[60] cm tall;
stems pilosulous [to glabrous]. Leaves in unequal [to subequal]
pairs; petiole [0.5]13 cm or longer, glabrous; blade drying
thickly papery [or leathery], adaxially grayish green, abaxially
greenish yellow, ovate, sublanceolate, or elliptic-oblong, 314[
15] 24.5[6] cm, subglabrous on both surfaces, base obtuse
and often shortly decurrent, distinctly inequilateral [or symmetrical], margins entire or undulate, apex acuminate or cuspidate; secondary veins 7 or 8(or 9) pairs; stipules caducous, reduced, subulate. Inflorescences corymbose to congested-cymose, several to many flowered, puberulent to subglabrous; peduncle 11.5 cm; axes short or 0.51 cm, helicoid; bracts linear,
23 mm, caducous. Flowers with biology unknown, on pedicels
12 mm. Calyx densely puberulent; hypanthium broadly compressed turbinate, ca. 1 mm, 5-ribbed; lobes triangular, ca. 0.4[
1.5] mm, each sinus with 1 gland. Corolla pale red, drying yellow, salverform, glabrous outside; tube 2325[27] mm, glabrous inside; lobes ovate-triangular, [2]5 mm, dorsally narrowly winged, apex rostrate. Capsules mitriform, 56 1415
mm, puberulent. Fl. AprJun.
Shady and wet places in forests. Yunnan [NE India, Myanmar].

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282

Measurements in brackets are taken from the description of this

species by Deb and Mondal (Bull. Bot. Surv. India 39(14): 135137.
1997) and may be expected in Chinese plants.

69. Ophiorrhiza wenshanensis H. S. Lo, Bull. Bot. Res., Harbin 10(2): 17. 1990.
wen shan she gen cao
Herbs, weak in lower part, ascending above, to 20 cm tall;
stems densely pilosulous to hispidulous. Petiole 0.51 cm, hispidulous; leaf blade drying papery, adaxially grayish green, pallid abaxially, ovate to lanceolate, 1.53.5(7) 0.82(3) cm,
subglabrous to sparsely hispidulous adaxially, subglabrous except pilosulous on veins abaxially, base cuneate to obtuse, apex
obtuse; secondary veins 5 or 6 pairs; stipules glabrous, 2-parted
almost to base, 2.53 mm, lobes subulate-triangular, with globose gland at apex and sometimes also stipitate glands at base.
Inflorescences 1- or 2-flowered, glabrous; pedicels 35 mm;
bracts 2, fused to hypanthium base, linear-subulate, ca. 5 mm.
Flowers reportedly distylous, pedicellate or pedunculate. Calyx
glabrous; hypanthium turbinate, ca. 1.5 mm, 68-ribbed; lobes
57, lanceolate-linear, sometimes unequal, 13 mm. Corolla
white, funnelform with throat markedly expanded, outside glabrous; tube 1820 mm, densely white villous inside through
throat; lobes 6 or 7, subovate, 34 mm, dorsally with very short
horn, apex rostrate. Capsules subobcordate, ca. 3 8 mm, with
persistent bracteoles. Fl. May.

Streamsides in forests. Yunnan (Wenshan).

In the protologue and FRPS (71(1): 120. 1999), H. S. Lo described the flowers as probably distylous, but only the putative shortstyled form has been documented.

70. Ophiorrhiza wui H. S. Lo, Bull. Bot. Res., Harbin 10(2):

33. 1990.
wu shi she gen cao
Herbs, procumbent at base, ascending above, to 30 cm tall;
stems drying reddish brown, subterete to striate-grooved, puberulent to densely pilosulous. Leaves in subequal pairs; petiole
12.5 cm, pilosulous; blade drying thinly papery, ovate or lanceolate-ovate, 511 1.84 cm, adaxially glabrous or sparsely
strigillose, abaxially villosulous or hirtellous along veins, base
cuneate then shortly decurrent, apex obtuse then abruptly acuminate; secondary veins 57 pairs; stipules subfiliform, ca. 6
mm. Inflorescences cymose, several flowered, branched to several orders, on peduncles and axes with 2 pilosulous or strigillose lines; peduncle 12.5 cm; axes helicoid; bracts absent or
reduced. Flowers with biology unknown, subsessile. Calyx with
hypanthium ca. 1.5 mm; lobes ca. 0.4 mm. Corolla white or
reddish, tubular, glabrous outside; tube 4.55 mm, densely pubescent in throat; lobes oblong-triangular, ca. 2 mm, dorsally
winged, markedly reflexed in anthesis. Capsules unknown. Fl.
Apr.
On wet limestone; ca. 1100 m. Yunnan (Luxi).

66. PAEDERIA Linnaeus, Syst. Nat., ed. 12, 2: 135, 189; Mant. Pl. 1: 7, 52. 1767, nom. cons.
ji shi teng shu
Chen Tao (); Charlotte M. Taylor
Daun-contu Adanson.
Shrubs, subshrubs, or vines, unarmed, usually extensively twining, usually with fetid odor when bruised. Raphides present.
Leaves opposite or infrequently in whorls of 3 or 4, without or sometimes with (Paederia foetida, P. spectatissima) pubescent
domatia; stipules caducous or persistent, interpetiolar, triangular to bilobed. Inflorescences axillary and/or terminal on main stem or
often on short lateral stems, thyrsiform, paniculate, cymose, or spiciform, several to many flowered, sessile to pedunculate, bracteate
with bracts sometimes enlarged and stipitate [to petaloid]. Flowers sessile to pedicellate, bisexual, monomorphic. Calyx limb (4
or)5(or 6)-lobed [sometimes with calycophylls]. Corolla white, pink, or purple, funnelform to salverform usually with tube very
slender then abruptly enlarged shortly above base, inside pubescent in tube and throat, sometimes fenestrate near base; lobes (4
or)5(or 6), induplicate-valvate in bud, with margins often crisped to irregular, rarely shortly trifid at apex. Stamens (4 or)5(or 6),
included, inserted at various levels near middle of corolla tube, included; filaments reduced [or sometimes developed]; anthers dorsifixed. Ovary 2(or 3)-celled, ovules 1 in each cell, erect, basal, anatropous; stigmas 2, filiform, included or exserted. Fruit characteristic: dry, drupaceous becoming schizocarpous, globose or compressed globose to compressed ellipsoid, with calyx limb persistent
and occasionally becoming enlarged; exocarp dry, membranous to papery, usually drying shiny, at maturity fragmenting; pyrenes
(diaspores) 2(or 3), indehiscent, membranous to leathery, hemispherical to flattened, oblong to ovate in outline, entire to winged,
rarely pubescent (P. yunnanensis), sometimes borne on a persistent carpophore; seeds with testa thin; cotyledons broadly cordate;
radicle short, hypogeous.
Thirty species: tropical and subtropical Africa, Asia, Madagascar, North America (Mexico), and South America; nine species (three endemic) in
China.
Paederia was studied in detail by Puff and collaborators (in Puff, Opera Bot. Belg. 3: 1376. 1991). They recognized three subgenera based on
corolla morphology and size, anther position, style length, the presence of petaloid bracts, and fruit morphology. Two of their subgenera are found in
China: P. subg. Paederia, which is restricted to SE Asia and includes P. cavaleriei, P. foetida, P. pertomentosa, and P. stenobotrya; and P. subg.
Alatopaederia Puff, which is found worldwide except continental Africa and includes the remaining Chinese species. Puff (loc. cit.: 207292)
presented a species-level taxonomy of Asian Paederia that differed significantly from that of other authors, including W. C. Ko (in FRPS 71(2): 110
119. 1999). In particular, he recognized fewer species, circumscribed P. foetida more widely, and accordingly synonymized several names. Puff also
applied the name P. foetida differently than previous authors, and his conclusions were not adopted in FRPS: he applied the name P. foetida to plants

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283

treated by W. C. Ko (loc. cit.: 118119) as P. scandens, and he included the plants treated as P. foetida by W. C. Ko (loc. cit.: 112113) in P.
cruddasiana. Puff (loc. cit.: 216220) discussed in detail the confusion of these species and the typification of P. foetida and synonymized P. scandens
under P. foetida. The treatment here follows Puff, which is well documented and internally consistent, and thus is distinct from traditional taxonomy of
SE Asian Paederia.
The fruit of Paederia are unusual in Rubiaceae: they are drupaceous in structure but dry and tardily schizocarpous with the exocarp fragmenting
to expose the two pyrenes, which are the dispersal unit or diaspores, sometimes simply enclosed in the fruit and sometimes borne on carpophores
(Puff, loc. cit.: 1376). Paederia species are best distinguished by fruit characters; determinations of flowering specimens are usually provisional. The
corollas of most species of Paederia have a notable size range, sometimes varying by 100200. Paederia foetida is by far the most commonly
collected Asian species of the genus and one of the most commonly collected species of Rubiaceae in China. The descriptions below follow Puff (loc.
cit.: 207292) in describing primarily what he termed the mid-stem region, i.e., the mature stems below the apical, young region. Inflorescence
morphology was used by Puff in part to distinguish species; however, these are indeterminate and in several species continue to grow. In particular,
their axes continue to elongate for some time during the flowering period; thus, if inflorescences of different ages are compared these characters can
be problematic to interpret. W. C. Ko (loc. cit.: 111) described the anthers as basifixed or dorsifixed, but Puff (loc. cit.) reported them as dorsifixed.

Key to fruiting material

1a. Fruit globose to subglobose, 47 mm in diam.; pyrenes hemispherical to concavo-convex or plano-convex.
2a. Inflorescences paniculate, thyrsiform, corymbiform, or cymose, usually branched to several orders and
with flowers in open cymose groups, dichotomous or frequently with higher order axes scorpioid;
stipules 1.56 mm ....................................................................................................................................................... 3. P. foetida
2b. Inflorescences narrowly paniculate, racemiform, or spiciform, branched to 1 or several orders and with
flowers in congested groups to small heads, axes when developed usually dichotomously branched;
stipules 212 mm.
3a. Stipules 23.5 mm; calyx lobes 0.61 mm ............................................................................................... 5. P. pertomentosa
3b. Stipules 412 mm; calyx lobes 0.42 mm.
4a. Calyx lobes 0.41 mm; leaf margins flat or often finely and extensively crisped; plants of
mainland and Taiwan ................................................................................................................................ 1. P. cavaleriei
4b. Calyx lobes 12 mm; leaf margins flat; plants of mainland and Hainan ............................................. 8. P. stenobotrya
1b. Fruit orbicular, ovoid, or ellipsoid, rounded to strongly flattened, 515.5 4.511 mm; pyrenes flattened,
orbicular, ovate, or elliptic in outline, sharply edged to marginally winged.
5a. Stipules 11.5 mm; fruit 1011 mm wide, flattened; pyrenes papery; calyx lobes 0.30.6 mm;
inflorescence branched to several orders, becoming diffuse with well-developed axes, these
usually ascending .............................................................................................................................................. 7. P. spectatissima
5b. Stipules 2.525 mm; fruit 4.510 mm wide, rounded to flattened (if stipules less than 4 mm then
fruit 4.58 mm wide); calyx lobes 0.52 mm.
6a. Stipules 925 mm.
7a. Fruit 57 mm; calyx lobes 0.50.8 mm .............................................................................................. 6. P. praetermissa
7b. Fruit 69 mm; calyx lobes 0.51.7 mm ............................................................................................... 9. P. yunnanensis
6b. Stipules 2.58.9 mm.
8a. Stipules 2.56 mm ................................................................................................................................ 2. P. cruddasiana
8b. Stipules 48.9 mm.
9a. Fruit 915 79 mm ....................................................................................................................... 4. P. lanuginosa
9b. Fruit 69 57 mm ....................................................................................................................... 9. P. yunnanensis
Key to flowering material
1a. Well-developed inflorescences paniculate, thyrsiform, corymbiform, or cymose, usually branched to several
orders (up to 10) and with flowers borne separated in open cymose groups, with axes dichotomous or
frequently higher order axes scorpioid; stipules 16 mm.
2a. Inflorescences becoming diffuse, axes mostly dichotomously branched, well developed, and ascending;
stipules 11.5 mm ............................................................................................................................................. 7. P. spectatissima
2b. Inflorescences small to extensive and spreading, with axes dichotomously branched and/or often highest
order axes markedly scorpioid; stipules 1.56 mm.
3a. Flowers generally borne in small rather congested groups; calyx lobes 0.42 mm; corolla lobes
24 mm; pyrenes somewhat flattened ........................................................................................................ 2. P. cruddasiana
3b. Flowers generally borne in branched cymules; calyx lobes 0.41 mm; corolla lobes 12 mm;
pyrenes hemispherical .......................................................................................................................................... 3. P. foetida
1b. Well-developed inflorescences paniculate, racemiform, or spiciform, branched to 1 to several orders and with
flowers usually congested in small groups or heads, with axes short and dichotomously branched or sometimes
unbranched; stipules 225 mm.
4a. Calyx limb with short tube and lobes deltoid and equal in length to tube; stipules 925 mm .................... 6. P. praetermissa

RUBIACEAE

284

4b. Calyx limb lobed to base or with very short tube, lobes linear, ovate, suborbicular, or triangular and
longer than tube; stipules 215 mm.
5a. Stipules 23.5 mm; leaves usually with sides generally parallel; inflorescences with secondary axes
often not developed ................................................................................................................................... 5. P. pertomentosa
5b. Stipules 4.515 mm; leaves generally with sides curved; inflorescences with secondary axes at
least shortly developed.
6a. Corolla tube 15.517.5 mm .................................................................................................................... 4. P. lanuginosa
6b. Corolla tube 410.5 mm.
7a. Plants of mainland and Hainan; leaves 617 37 cm; corolla tube 56 mm; fruit usually
drying orange-yellow; pyrenes plano-convex or concavo-convex ................................................ 8. P. stenobotrya
7b. Plants of mainland and Taiwan; leaves 622 2.513 cm; corolla tube 410.5 mm; fruit drying
straw-yellow to brown, gray, or black; pyrenes plano-convex to concavo-convex or flattened.
8a. Leaf margins flat or often finely crisped; stems densely hirsute, hirtellous, pilosulous,
and/or tomentose to glabrescent; calyx lobes 0.41 mm; pyrenes plano-convex to
concavo-convex ........................................................................................................................... 1. P. cavaleriei
8b. Leaf margins flat; stems densely tomentulose and/or hirtellous to glabrescent; calyx
lobes 0.51.7 mm; pyrenes flattened ...................................................................................... 9. P. yunnanensis
1. Paederia cavaleriei H. Lveill, Repert. Spec. Nov. Regni
Veg. 13: 179. 1914.
er ye ji shi teng
Vines, to 4 m; stems densely hirsute and/or hirtellous, pilosulous, or tomentulose to glabrescent, drying brown. Leaves
opposite; petiole 28(21) cm, densely hirsute or hirtellous to
glabrescent; blade drying submembranous to papery, ovate, oblong-ovate, lanceolate, rhombic-ovate, or oblong, 618(22)
2.510(13) cm, adaxially sparsely to densely strigillose, hirtellous, hirsute, or scabrous, abaxially sparsely to densely pilosulous to hirtellous with pubescence denser on veins, base
rounded or truncate to cordulate or usually cordate, margin flat
or usually densely finely crisped and often appearing denticulate, apex acute to long acuminate; secondary veins 510 pairs;
stipules generally persistent, triangular to lanceolate, 512 mm,
acute to acuminate. Inflorescences axillary and/or terminal,
paniculate to racemiform, cylindrical to narrowly pyramidal, 7
20(30) cm, branched to 24 orders, densely hirtellous, pilosulous, or hirsute, pedunculate; bracts triangular to linear, 13
mm; pedicels to 1 mm. Flowers sessile to pedicellate in congested cymules or small heads. Calyx puberulent to glabrous;
hypanthium portion ellipsoid to turbinate, 11.6 mm; limb
lobed nearly to base; lobes triangular, 0.41 mm. Corolla
pinkish gray, lilac gray, grayish white, or purplish green, tubular-funnelform, outside densely mealy tomentose or mealy puberulent; tube 410 2.54.5 mm, without slits; lobes broadly
triangular to broadly ovate, 12 mm, obtuse to acute. Fruit globose, 4.55 4.55 mm, puberulent to glabrous, drying strawyellow; pyrenes plano-convex to concavo-convex. Fl. Apr
Aug, fr. AugNov.
Thickets on mountains; 1003000 m. Guangdong, Guangxi,
Guizhou, Hubei, Hunan, ?Sichuan, Taiwan [?Laos].

2. Paederia cruddasiana Prain, J. Asiat. Soc. Bengal, Pt. 2,

Nat. Hist. 67: 295. 1898.
chou ji shi teng
Paederia cruddasiana subsp. microcarpa (Kurz) Puff; P.
foetida Linnaeus var. microcarpa Kurz.
Vines, to 10 m; stems moderately to sparsely hirtellous or

strigillose to glabrous, drying yellowish brown. Leaves opposite; petiole 13 cm, hirtellous or strigillose to glabrescent;
blade drying membranous, ovate, lanceolate, or narrowly lanceolate, 516 210.5 cm, adaxially glabrous or puberulent on
principal veins, abaxially sparsely hirtellous to glabrous on
blade and sparsely to moderately puberulent or hirtellous along
veins, base rounded to truncate or cordate, margins flat, apex
acute to weakly acuminate; secondary veins 48 pairs; stipules
generally persistent, ovate-lanceolate to broadly triangular, 2.5
6 mm, acute or bifid. Inflorescences axillary and/or terminal,
paniculate, pyramidal, 650 cm, branched to 35 orders, hirtellous to glabrescent, pedunculate; bracts triangular to linear, 0.5
2 mm; pedicels to 2 mm. Flowers subsessile to pedicellate in
cymules. Calyx puberulent to glabrescent; hypanthium portion
ellipsoid, ca. 1 mm; limb lobed nearly to base; lobes triangular,
0.42 mm. Corolla purplish blue, lilac, or pink, funnelform,
outside densely puberulent to tomentulose; tube 616 1.54.5
mm, without slits; lobes triangular, 24 mm. Fruit broadly ellipsoid to ovoid, somewhat laterally compressed, 611 4.58
mm, glabrescent, drying yellowish gray; pyrenes ovate to elliptic, somewhat flattened, marginally winged. Fl. MayJun, fr.
NovDec.
Open forests; 1001900 m. Yunnan [Bangladesh, Bhutan, India,
Myanmar, Nepal, Thailand, Vietnam].
This species has long been incorrectly treated by several previous
authors as Paederia foetida, including by W. C. Ko (in FRPS 71(2):
112113. 1999); see comments above in the genus discussion. Puff
(Opera Bot. Belg. 3: 251252. 1991) recognized two subspecies of P.
cruddasiana, distinguished by fruit and pyrene size and form, and included some Chinese plants in subsp. cruddasiana and others in subsp.
microcarpa. The distribution of these subspecies is apparently complementary (Puff, loc. cit.: 252, f. 11), with the plants in the SE range of
this species falling into subsp. microcarpa; however, the measurements
that separate these taxa are overlapping generalizations, based on study
of ten collections of subsp. cruddasiana and four collections of subsp.
microcarpa from China, with several collections of each subspecies
apparently found in the same relatively small region (e.g., Mengla
Exped. 34288, subsp. microcarpa, and Li Y. H. 317, subsp. cruddasiana,
both reported as 2130N 10125E; this latter collection apparently not
mapped in his f. 11). Given the large morphological variation documented within other species of Paederia, the few specimens studied by
Puff, and the geographic overlap, these varieties are provisionally not
recognized here pending further study.

RUBIACEAE

3. Paederia foetida Linnaeus, Syst. Nat., ed. 12, 2: 189; Mant.

Pl. 1: 52. 1767.
ji shi teng
Gentiana scandens Loureiro; Paederia chinensis Hance;
P. dunniana H. Lveill; P. esquirolii H. Lveill; P. laxiflora
Merrill ex H. L. Li; P. mairei H. Lveill; P. scandens (Loureiro) Merrill; P. scandens f. mairei (H. Lveill) Nakai; P.
scandens var. mairei (H. Lveill) H. Hara; P. scandens var.
tomentosa (Blume) Handel-Mazzetti; P. tomentosa Blume; P.
tomentosa var. glabra Kurz; P. tomentosa var. mairei (H. Lveill) H. Lveill; P. stenophylla Merrill.
Vines, to 5 m; stems glabrous to densely puberulent, hirtellous, or pilosulous often becoming glabrescent, drying gray
to brown. Leaves opposite or rarely ternate; petiole 0.59 cm,
glabrous to densely hirtellous or pilosulous; blade drying papery to subleathery, ovate, ovate-oblong, lanceolate, lanceolateelliptic, or elliptic, (1)59(21) 14(9) cm, adaxially glabrous to puberulent at least on principal veins, abaxially glabrous to puberulent, hirtellous, or strigillose at least on principal
veins, base cuneate, rounded, truncate, or sometimes cordulate
to cordate, margins flat, apex acute or acuminate; secondary
veins 46 pairs; stipules generally persistent, triangular to
ovate, 1.56 mm, obtuse to acute, acuminate, or rarely bifid. Inflorescences axillary and/or terminal, paniculate, thyrsiform,
corymbiform, or cymose, pyramidal to rounded, 5100 cm,
branched to 25 orders with higher order axes dichotomous or
often scorpioid, hirtellous, strigillose, or glabrous, pedunculate;
bracts lanceolate to triangular, 0.83 mm; pedicels to 1.5 mm.
Flowers sessile to pedicellate in dichotomous to scorpioid, lax
to somewhat congested cymules. Calyx glabrous to densely puberulent; hypanthium portion turbinate to ellipsoid, 0.82 mm;
limb deeply lobed; lobes triangular, 0.41 mm. Corolla pale
purple, grayish pink, lilac, or grayish white, funnelform, outside
densely mealy puberulent or mealy tomentulose; tube (5)7
10(17) 26 mm, without slits; lobes broadly triangular to
ovate, 12 mm, acute. Fruit globose, 47 47 mm, glabrescent, drying gray to yellow; pyrenes concavo-convex to planoconvex. Fl. MayOct, fr. JulDec.
Forests, forest margins, thickets in ravines and on mountain
slopes; 2002000 m. Anhui, Fujian, Gansu, Guangdong, Guangxi,
Guizhou, Hainan, Henan, Hubei, Jiangsu, Jiangxi, Shandong, Shanxi,
Sichuan, Taiwan, Yunnan, Zhejiang [Bangladesh, Bhutan, Borneo, Cambodia, India, Indonesia, Japan, N Korea, Laos, Malaysia, Myanmar, Nepal, Philippines, Thailand, Vietnam; also occasionally cultivated, and
naturalized in United States (Florida) and perhaps Sri Lanka].
Paederia foetida is naturalized in a few places around the world
and apparently quite weedy wherever it grows; in particular, it has
been reported as an adventive and sometimes a problem weed in
Florida, where it is called skunk vine.
Puff (Opera Bot. Belg. 3: 207292. 1991) substantially changed
and clarified the application of this name; see the discussion under the
genus above. Paederia foetida as circumscribed by Puff (and here) is
widespread and morphologically widely variable but with continuous
morphological variation, as documented in detail (loc. cit.: 223224, f.
4, f. 5). W. C. Ko (in FRPS 71(2): 118119. 1999) recognized two
varieties within this species (treated as P. scandens), with the stems and
leaves glabrous or subglabrous in var. scandens and subglabrous to
pubescent in var. tomentosa; Ko reported var. scandens from throughout

285

the range of this species in China and var. tomentosa only from
Jiangxi, Guangdong, Hong Kong, Hainan, Guangxi, Yunnan. Puff
synonymized these based on his conclusion that pubescence varies
continuously within this species and thus does not distinguish separate
evolutionary lineages.
Puff synonymized Paederia stenophylla with P. foetida, and this
name is provisionally listed in synonymy here. The name P. stenophylla
seems to have been applied by W. C. Ko (loc. cit.: 115) at least in part to
plants that Puff treated as P. pertomentosa rather than P. foetida, and
Puff reported that he did not see the type of P. stenophylla; this situation
may deserve re-consideration, but that work is outside the scope of this
current project.
The name Paederia dunniana from Guizhou was apparently overlooked by Puff; this was reported by Lauener and Ferguson (Notes Roy.
Bot. Gard. Edinburgh 32: 112. 1972) to be a synonym of P. scandens,
which corresponds to P. foetida here, and this name is accordingly
synonymized here provisionally.

4. Paederia lanuginosa Wallich, Pl. Asiat. Rar. 2: 52. 1831.

rong mao ji shi teng
Hondbesseion lanuginosum (Wallich) Kuntze; Paederia
macrocarpa Wallich.
Vines, to 12 m; stems densely tomentose to glabrescent,
drying dark brown. Leaves opposite; petiole 3.512 cm,
densely tomentose; blade elliptic to oblong-elliptic, 820 4.5
15 cm, adaxially sparsely to moderately strigillose with pubescence denser on midrib, abaxially densely tomentulose, base
cordate to subrounded, margins flat, apex acuminate; stipules
caducous often by fragmentation, triangular, 4.58.5 2.35.5
mm, acute to bifid. Inflorescences axillary on main stems
and/or terminal on lateral stems, paniculate, densely tomentulose, pedunculate. Flowers subsessile in small heads. Calyx
densely tomentulose; hypanthium portion turbinate to subglobose, 1.62.5 mm; limb deeply lobed; lobes triangular to suborbicular, 11.4 mm. Corolla pale green, dull white, pink, reddish
purple, or dark purple, funnelform, outside densely tomentulose; tube 1517.5 3.54.5 mm, with slits at base; lobes
broadly triangular, 2.74 mm, acuminate, marginally crisped.
Fruit oblong-elliptic, laterally flattened, 915 79 mm, subglabrous, drying brown; pyrenes elliptic to ovate in outline, flattened, marginal wing 12 mm wide. Fl. JunJul, fr. AugFeb.
Open forests or thickets, twining on other shrubs and small trees;
sea level to 1900 m. Yunnan [Myanmar, Thailand].
Puff (Opera Bot. Belg. 3: 207292. 1991) noted that this species
has the largest leaves of the genus, as well as notably large flowers and
fruit, and sometimes has conspicuously fissured bark that is also distinctive. W. C. Ko (in FRPS 71(2): 114. 1999) described the corolla
lobes as 3-lobed, but this has not been mentioned by any other authors.

5. Paederia pertomentosa Merrill ex H. L. Li, J. Arnold Arbor.

24: 458. 1943.
bai mao ji shi teng
Vines or clambering subshrubs, to 5 m; stems densely pilosulous to tomentulose, when dry dark straw-yellow. Leaves
opposite; petiole 15 cm, densely pilosulous or hirtellous and/or
tomentulose; blade drying papery, ovate-elliptic, lanceolateoblong, or oblong-elliptic, 515 26 cm, adaxially moder-

286

RUBIACEAE

ately to densely pilosulous to strigillose with pubescence denser

along midrib, abaxially densely white tomentulose, base obtuse,
rounded, truncate, or cordulate, margins flat, apex acute to acuminate; secondary veins 68 pairs; stipules generally persistent,
triangular, 23.5 mm, acute to weakly acuminate. Inflorescences axillary and/or terminal, spiciform to paniculate, cylindrical to narrowly pyramidal, 1550 cm, branched to 13 orders
with ultimate axes often very short, densely pilosulous to tomentulose, pedunculate; bracts triangular, 14 mm. Flowers
sessile or subsessile in congested cymules or small heads. Calyx densely pilosulous to tomentose; hypanthium portion turbinate to ellipsoid, ca. 1 mm; limb lobed nearly to base; lobes triangular, 0.61 mm. Corolla lilac, grayish pink, or grayish purple, tubular-funnelform, outside densely papillose to puberulent; tube 58 23 mm, without slits; lobes ovate to triangular,
11.5 mm, acute, margins flat to crisped. Fruit globose to subglobose, 47 47 mm, glabrous, drying straw-yellow; pyrenes
plano-convex or concavo-convex. Fl. MayAug, fr. JulNov.

gular, 11.5 mm, obtuse to shortly acuminate. Inflorescences

axillary and/or terminal, paniculate, pyramidal to cylindrical,
20100 cm, branched to 610 orders, papillose, puberulent, or
glabrescent, pedunculate; bracts triangular, 12 mm. Flowers
sessile or subsessile. Calyx densely puberulent; hypanthium
portion ellipsoid, 11.5 mm; limb 0.81 mm, lobed for ca.
1/2; lobes triangular, 0.30.6 mm. Corolla greenish to purplish
white, funnelform, densely papillose-puberulent to scabrid-papillose outside; tube 58 2.63.7 mm, with slits at base; lobes
triangular, 1.72.4 mm, acute, margins often crisped. Fruit
orbicular, flattened laterally, 1011 1011 mm, puberulent,
drying dark gray; pyrenes orbicular in outline, flattened, papery,
marginal wing 44.5 mm wide. Fl. and fr. JunOct.
Open forests; 8001000 m. Guangxi, Yunnan (Pingbian) [Vietnam].

6. Paederia praetermissa Puff, Opera Bot. Belg. 3: 273. 1991.

W. C. Ko (in FRPS 71(2): 112. 1999) attributed this name to Puff

(Opera Bot. Belg. 3: 285. 1991). However, Puff attributed this name to
Li in unpublished work and provided no Latin diagnosis or designation
of a type, so this name was not validly published by him. Li cited only
three collections in her article in Novon and none were specimens cited
by Puff, so the specimens cited by Puff do not actually seem to be paratypes as has been suggested.

qi yi ji shi teng

8. Paederia stenobotrya Merrill, Lingnan Sci. J. 11: 57. 1932.

Forests, thickets, often on limestone; 2001400 m. Fujian,

Guangdong, Guangxi, Hainan, Hunan, Jiangxi.

Vines, to 5 m; stems densely hirsute to hirtellous becoming glabrescent, drying brownish gray. Leaves opposite;
petiole 1.58 cm, moderately to densely hirsute, hirtellous, or
pilosulous; blade drying papery to subleathery, ovate-elliptic or
lanceolate, 622 3.512 cm, adaxially sparsely hirtellous or
hispidulous to glabrescent, abaxially densely pilose to hirtellous
or tomentose, base cordulate to cordate, margins flat, apex acute
to acuminate; secondary veins ca. 8 pairs; stipules caducous, triangular to lanceolate, 925 513 mm. Inflorescences terminal
on lateral stems and/or axillary on main stems, paniculate, pyramidal to cylindrical, 820 cm, branched to 24 orders, densely
hirsute to hirtellous or pilosulous, pedunculate; bracts linear to
triangular. Flowers subsessile. Calyx densely pilosulous to hirtellous; hypanthium portion ellipsoid, 11.4 mm; limb lobed for
ca. 1/2; lobes narrowly lanceolate or linear-lanceolate, 0.50.8
mm. Corolla grayish purple to pink, funnelform, outside densely
puberulent or scabridulous; tube 4.59 1.21.5 mm, without
slits; lobes ovate-elliptic, 1.52.5 mm, acute to minutely trifid,
margins crisped. Fruit brown, ellipsoid to ovoid, laterally somewhat compressed, 57 56 mm, sparsely to moderately pilosulous to hirtellous to glabrescent; pyrenes ovate to elliptic in
outline, flattened, marginal wing 0.81 mm wide. Fl. JunJul or
OctNov, fr. DecJan.
Sparse forests or thickets; 6001300 m. Yunnan [Myanmar, Thailand, ?Vietnam].

7. Paederia spectatissima H. Li, Novon 9: 220. 1999.

yun gui ji shi teng
Vines, twining, to 12 m; stems glabrous, drying dark red
then grayish brown. Leaves opposite; petiole 14 cm, glabrous;
blade drying leathery to papery, ovate-elliptic to elliptic, 612
36 cm, glabrous on both surfaces, base cuneate to obtuse, margins flat, apex acuminate with tip usually prolonged; secondary
veins 610 pairs; stipules persistent or caducous, broadly trian-

xia xu ji shi teng

Vines, to 3 m; stems densely hirtellous, hirsute-hirtellous,
or glabrescent, drying dark yellow. Leaves opposite; petiole
2.57 cm, densely hirtellous to tomentose-hirtellous; blade
drying papery, ovate, oblong-ovate, or elliptic-ovate, 617 3
11 cm, adaxially sparsely to moderately scabrid to hirtellous,
abaxially hirtellous, pilosulous, or pilose with pubescence
denser along veins, base cordate, cordulate, truncate, or rarely
acute, margins flat, apex acute to acuminate; secondary veins
58 pairs; stipules generally caducous, triangular, 410 mm,
acute. Inflorescences axillary and/or terminal, spiciform to narrowly paniculate, cylindrical to narrowly pyramidal, 730 cm,
branched to 13 orders, densely hirtellous to hirtellous-tomentose, pedunculate; bracts triangular, 12 mm. Flowers sessile in
congested cymules or small heads. Calyx densely hirtellous; hypanthium portion turbinate to ellipsoid, 11.5 mm; limb lobed
nearly to base; lobes subulate to narrowly triangular, 12 mm.
Corolla funnelform, outside densely hirtellous to tomentulose;
tube 56 2.53 mm, without slits; lobes ovate, 12 mm. Fruit
globose, 56 56 mm, glabrous, drying orange-yellow; pyrenes plano-convex to concavo-convex. Fl. Jun, fr. JunNov.
Broad-leaved forests on hill slopes; 400900 m. Fujian, Guangdong, Hainan.
Puff (Opera Bot. Belg. 3: 207292. 1991) reported this species
only from Hainan, but W. C. Ko (in FRPS 71(2): 117. 1999) reported it
also from Guangdong and Fl. Fujian. (5: 185. 1993) from Fujian.

9. Paederia yunnanensis (H. Lveill) Rehder, J. Arnold Arbor. 18: 249. 1937.
yun nan ji shi teng
Cynanchum yunnanense H. Lveill, Cat. Pl. Yun-Nan,
13. 1915; Paederia bodinieri H. Lveill (19141915), not H.

RUBIACEAE

Lveill (1914); P. rehderiana Handel-Mazzetti; P. tomentosa

Blume var. purpureocaerulea H. Lveill & Vaniot.
Vines, to 7 m; stems densely tomentulose and/or hirtellous
becoming glabrescent, drying brown. Leaves opposite; petiole
27.5 cm, densely tomentulose and/or pilosulous; blade drying
submembranous to papery, ovate, lanceolate, or lanceolateelliptic, 616 312 cm, adaxially densely strigillose, pilosulous, or scabrous, abaxially densely tomentose to hirtellous,
base cordulate to deeply cordate or rarely obtuse, margins flat,
apex acute or acuminate; secondary veins 68 pairs; stipules
generally persistent, lanceolate-triangular to narrowly triangular, 415 mm, acute to acuminate or rarely bifid. Inflorescences
axillary on main stems and/or terminal on lateral stems, paniculate, cylindrical to narrowly pyramidal, 625 cm, branched to
13 orders, densely tomentulose to hirtellous or pilosulous, pedunculate or sometimes apparently sessile with basal axes sub-

287

tended by reduced leaves (or leaflike bracts); bracts linear to

narrowly triangular or deeply bilobed, 1.56 mm. Flowers sessile and subsessile in congested to subcapitate cymules (sometimes appearing pedicellate when axes of cymes develop later).
Calyx with hypanthium portion ellipsoid, 11.5 mm, glabrous
to puberulent; limb lobed nearly to base; lobes oblong-lanceolate to narrowly triangular, 0.51.7 mm, sparsely to densely puberulent, strigillose, or hirtellous. Corolla pale green, pink, reddish purple, or dark purple, tubular-funnelform, outside densely
mealy puberulent; tube 510.5 2.55 mm, without slits; lobes
broadly triangular, 12.5 mm, obtuse to acute, margins crisped.
Fruit ovoid, laterally compressed, 69 57 mm, glabrescent,
drying brown; pyrenes ovate in outline, flattened, papillose-puberulent, marginal wing ca. 1 mm wide. Fl. JunOct, fr. Jul
Dec.
Forest margins in valleys; 3003000 m. Guangxi, Guizhou, Sichuan, Yunnan [Vietnam].

67. PAVETTA Linnaeus, Sp. Pl. 1: 110. 1753.

da sha ye shu
Chen Tao (); Charlotte M. Taylor
Pavate Adanson.
Shrubs, small trees, or infrequently subshrubs, unarmed. Raphides absent. Leaves opposite or rarely whorled, usually with bacterial nodules and/or domatia in abaxial vein axils; stipules generally persistent, shortly united around stem, triangular, often aristate,
often sericeous adaxially. Inflorescences terminal on principal stems, terminal on reduced lateral stems and appearing axillary, or
rarely truly axillary, cymose to corymbiform, many flowered, sessile to pedunculate, bracteate with bracts often fused in pairs.
Flowers pedicellate or sessile, bisexual, monomorphic, fragrant, with secondary pollen presentation. Calyx limb truncate or 4(or 5)lobed. Corolla white to cream [or rarely red], salverform with tube slender, inside glabrous or pubescent in throat; lobes 4, convolute
in bud. Stamens 4(or 5), inserted in corolla throat, exserted or included; filaments short; anthers dorsifixed near base, sometimes
becoming twisted with age. Ovary 2-celled, ovules 1(or 2) in each cell on axile placentas attached at top of septum; stigma restricted
to terminal portion of thickened style, very shortly 2-lobed with lobes erect, exserted. Fruit black and often shiny or infrequently
white, red, or blue, drupaceous, thinly fleshy, globose to ovoid, with calyx limb persistent or deciduous; pyrenes 2, 1-celled, each
with 1 seed, plano-convex or concavo-convex, papery; seeds medium-sized, ellipsoid, discoid, or plano-convex; testa membranous;
endosperm corneous; embryo dorsal, curved; cotyledons leaflike; radicle hypogeous.
About 400 species: paleotropical, widespread in Africa, tropical Asia, Australia, and Pacific islands, apparently absent from Madagascar; six
species (two endemic) in China.
Secondary pollen presentation is found in Pavetta (De Block, Opera Bot. Belg. 9: 1218. 1998; Rout & Deb, Bull. Bot. Surv. India 41: 1182.
1999). Rout and Deb (loc. cit.) reported synchronous flowering in the Indian species of Pavetta. W. C. Ko (in FRPS 71(2): 25. 1999) described the
filaments as ranging from short to prolonged, but that latter condition has not been reported by other authors in Pavetta or Ixora. Ko also described the
placentas as sometimes attached to the middle of the septum, but Bremekamp (Repert. Spec. Nov. Regni Veg. 37: 211. 1934) said this is incorrect
and an old mistake in Pavetta.
Bremekamp (loc. cit.; Repert. Spec. Nov. Regni Veg. 47: 1228. 1939) recognized three subgenera of Pavetta; the Chinese species all belong to
his P. subg. Pavetta, and the other two subgenera are restricted to Africa.
Bremekamp, in general, distinguished Pavetta species more narrowly than many other authors; for example, he (loc. cit. 1934; loc. cit. 1939)
reported 42 species for the Indian subcontinent, while Rout and Deb (loc. cit.) recognized only 25 species with the rest of Bremekamps names
synonymized. W. C. Ko (loc. cit.: 2530) followed Bremekamp (loc. cit. 1934) closely; in contrast, all the morphological variation found among the
Chinese species falls well within Rout and Debs (loc. cit.: 114136) circumscription of P. indica Linnaeus. Bremekamp (loc. cit. 1939) noted that the
E Chinese plants previously identified as P. indica were included by him in P. hongkongensis.
In particular, Bremekamp (loc. cit. 1934) considered stem characters to be informative taxonomically in Pavetta, particularly green vs. corkybarked stems, but Rout and Deb (loc. cit.) concluded that these represent different developmental stages rather than differences between species.
Bremekamp (loc. cit. 1934) considered the arrangement of the bacterial nodules in the leaves to be taxonomically informative for distinguishing
infrageneric groups and sometimes species, but Rout and Deb (loc. cit., as bacterial leaf-galls) found them to have no taxonomic value and
concluded that the nodules vary in shape and number among different leaves on a plant as well as between plants of the same and different species.
Bremekamp (loc. cit. 1934) distinguished several species based on leaf shape and size, but Rout and Deb (loc. cit.) included notable variation in leaf
size and shape, from relatively very narrow to quite broad, within individual species of Pavetta; Bridson and Verdcourt (Fl. Trop. E. Africa, Rub. (Pt.
2), 619686. 1988) circumscribed several species similarly to Rout and Deb. Bremekamp (loc. cit. 1934) considered several species of Pavetta to have

RUBIACEAE

288

axillary inflorescences, as did Rout and Deb (loc. cit.), but Bridson and Verdcourt (loc. cit.) considered the inflorescences of the African species at
least to be terminal on reduced lateral short shoots, as found in a number of Rubiaceae genera, and to appear axillary but not be truly axillary.
Bremekamp (loc. cit. 1934) gave much attention to the arrangement of the inflorescence bracts in Pavetta; his descriptions apply to bracts but not
bracteoles, so his characterizations may be misinterpreted if not observed carefully. Bremekamp noted (loc. cit. 1934) that occasional flowers with 5
calyx and corolla lobes are found in most Pavetta species, but the majority of flowers are always 4-merous and the genus best considered to be 4merous; a similar situation is found in other Rubiaceae genera.
The treatment here follows that of W. C. Ko (loc. cit.), for reference. The key here has been augmented with characters from the descriptions,
and the descriptions have been augmented with characters from the available specimens cited by Bremekamp.

1a. Flowering branches not green.

2a. Leaf blade elliptic-oblong to obovate-oblong, 918 33.5 cm, drying membranous, adaxially glabrous,
abaxially villosulous, with secondary veins 68 pairs ............................................................................................. 1. P. arenosa
2b. Leaf blade elliptic or elliptic-lanceolate, 815 49 cm, drying thickly papery, adaxially scabrous,
abaxially tomentose, with secondary veins 1012 pairs ....................................................................................... 6. P. tomentosa
1b. Flowering branches green or almost green, sometimes becoming black when dry.
3a. Leaf blade scabrous on both surfaces on midrib; style ca. 38 mm ...................................................................... 4. P. scabrifolia
3b. Leaf blade smooth, glabrous, or variously pubescent but not scabrous on both surfaces; style 35 mm or
shorter.
4a. Flowering branches and hypanthium portion of flower glabrous; leaf blade elliptic-oblong to
elliptic-obovate, adaxially glabrous, abaxially subglabrous or pubescent along midrib and
in axils of veins ....................................................................................................................................... 2. P. hongkongensis
4b. Flowering branches and hypanthium portion of flower puberulent to pilosulous or glabrescent;
leaf blade narrowly obovate, lanceolate, narrowly elliptic, or oblanceolate, adaxially glabrescent,
abaxially puberulent to pilosulous at least on veins.
5a. Leaf blade narrowly obovate or lanceolate, 913 3.54.7 cm, with secondary veins 68 pairs ......... 3. P. polyantha
5b. Leaf blade narrowly elliptic or oblanceolate, 1013 ca. 5 cm, with secondary lateral veins
6 or 7 pairs ............................................................................................................................................... 5. P. swatowica
1. Pavetta arenosa Loureiro, Fl. Cochinch. 1: 73. 1790, emend.
Bremekamp, Repert. Spec. Nov. Regni Veg. 47: 23. 1939.
da sha ye
?Pavetta sinica Miquel.
Shrubs, 13 m tall; branches compressed, puberulent to
glabrous. Petiole 520 mm, glabrous to sparsely pilosulous; leaf
blade drying membranous, elliptic-oblong to obovate-oblong,
918 33.5 cm, usually with bacterial nodules, adaxially glabrous and somewhat shiny, abaxially sparsely to densely villosulous, base cuneate to acute, apex acuminate; secondary
veins 68 pairs; stipules broadly ovate-triangular, 212 mm, pilosulous to glabrous, acute to obtuse. Inflorescence terminal, 9
11 ca. 15 cm, pilosulous to glabrescent; peduncle 2.54 cm;
pedicels 1012 mm. Flowers pedicellate. Calyx with hypanthium portion ellipsoid, ca. 1 mm, densely pilosulous; limb ca.
1 mm, sparsely pilosulous to glabrescent, lobed for up to 1/2.
Corolla white, outside glabrous; tube 1018 mm, bearded in
throat; lobes narrowly oblong, 35 mm, obtuse. Style 2530
mm. Drupe globose, 67 mm in diam., glabrous, calyx limb
persistent. Fl. AprMay, fr. OctNov.
Sparse forests at low elevations. Guangdong, Guangxi, Hainan
[Vietnam].
Plants with calyx tube glabrous from Guangxi have been called
Pavetta arenosa f. glabrituba Chun & F. C. How ex W. C. Ko (Fl. Hainan. 3: 583. 1974). This may be best synonymized here, pending further
study.
In his revision of Pavetta Bremekamp (Repert. Spec. Nov. Regni
Veg. 37: 104. 1934) described P. hongkongensis for the common Chinese plants and considered P. arenosa a dubious name that he tenta-

tively referred to Tarenna, but later (Bremekamp, Repert. Spec. Nov.

Regni Veg. 47: 1228. 1939) he reported that Merrill had differed with
his conclusion, had examined the type of P. arenosa, and had sent him
information that persuaded him to apply this latter name to Chinese
plants. The protologue provided very limited information, which was
significantly expanded by Bremekamp, thus the annotation here of this
as an emended description. Bremekamp here also stated, without citing
any individual specimens or contrasting the species, that both P. arenosa
and P. hongkongensis are found commonly in China. W. C. Ko (in
FRPS 71(2): 26. 1999) separated these based on the color of their flowering branches: green or almost green in P. hongkongensis vs. not green
in P. arenosa; however, this is a character that Rout and Deb (Bull. Bot.
Surv. India 41: 1182. 1999) tested empirically and found to be variable
within all Indian species of Pavetta and, thus, of questionable use there
and elsewhere.
W. C. Ko (loc. cit.: 28) cited the name Pavetta sinica as a synonym of P. arenosa, but the source for this synonymy is unknown to us.
Bremekamp specifically excluded this name from synonymy in his circumscription of P. arenosa, though apparently Merrill did synonymize it
here (Lingnan Sci. J. 15: 17. 1936). Bremekamp treated P. sinica as a
species distinct from P. arenosa and P. hongkongensis and considered it
most closely related to P. tomentosa. Bremekamp saw its type and described the corolla tube as 17 mm, which does not agree with the descriptions presented by W. C. Ko (loc. cit.: 2728) for either P. arenosa
or P. hongkongensis. However, the specimens cited for P. arenosa by
Bremekamp (loc. cit. 1939) have corolla tubes 1018 mm, so Bremekamps description of P. sinica is easily included in his circumscription
of P. arenosa.

2. Pavetta hongkongensis Bremekamp, Repert. Spec. Nov.

Regni Veg. 37: 104. 1934.
xiang gang da sha ye
Tarenna kwangsiensis Handel-Mazzetti.

RUBIACEAE

Shrubs or small trees, 14 m tall; branches compressed,

glabrous, green or almost green, often drying blackened. Petiole
12 cm, glabrous; leaf blade drying membranous, elliptic-oblong to elliptic-oblanceolate, 815 36.5 cm, often with bacterial nodules, adaxially glabrous, abaxially glabrous, glabrescent, or pubescent along midrib, base cuneate to acute, apex
acuminate to acute; secondary veins 6 or 7 pairs; stipules
broadly ovate-triangular, 13 mm, glabrous, acute to shortly
aristate. Inflorescences terminal on lateral branches, laxly corymbose, 79 715 cm, many flowered, glabrous; peduncle 1
2 cm; pedicels 36 mm. Flowers pedicellate. Calyx with hypanthium portion ellipsoid, ca. 1 mm, glabrous; limb 0.51.5 mm,
glabrous, shallowly lobed. Corolla white, outside glabrous; tube
1219 mm, inside pilose at throat; lobes narrowly triangular-oblong, 57 mm, acute to obtuse. Style ca. 35 mm. Drupes globose, 67 mm, pilosulous to glabrous, calyx limb persistent. Fl.
MarJul, fr. JulNov.
Thickets; 2001300 m. Guangdong, Guangxi, Hainan, Yunnan
[Vietnam].
Bremekamp noted in his description of this species that it comprises the Chinese plants previously treated as Pavetta indica by at least
some authors.

3. Pavetta polyantha (J. D. Hooker) R. Brown ex Bremekamp,

Repert. Spec. Nov. Regni Veg. 37: 103. 1934.
duo hua da sha ye
Pavetta indica Linnaeus var. polyantha J. D. Hooker, Fl.
Brit. India 3: 150. 1880.
Shrubs, 13 m tall; young branches compressed to subterete, glabrescent or puberulent. Petiole 1030 mm, puberulent;
leaf blade drying membranous, narrowly obovate or lanceolate,
913 3.54.7 cm, with several bacterial nodules, adaxially
glabrescent, abaxially puberulent at least along veins, base cuneate or acute, apex acuminate; secondary veins 68 pairs;
stipules ovate-triangular, 57 mm, puberulent or glabrescent,
shortly aristate. Inflorescences terminal on developed branches,
laxly corymbose, ca. 9 15 cm, many flowered, strigillose to
glabrescent; peduncle 1.52 cm; pedicels 35 mm. Flowers
pedicellate. Calyx with hypanthium portion ellipsoid, 11.2 mm,
densely strigillose; limb 11.5 mm, sparsely strigillose, lobed
for up to 1/2. Corolla white, outside glabrous; tube 1922 mm,
bearded in throat; lobes narrowly ligulate, 67 mm, obtuse to
rounded. Style ca. 30 mm. Drupe globose, ca. 8 mm, glabrous.
Fl. AprJun.
Sparse forests, streamsides; 9001200 m. Guangdong, Guangxi,
Guizhou, Yunnan [Bhutan, India, Indonesia, Myanmar, Philippines].
Rout and Deb (Bull. Bot. Surv. India 41: 122128. 1999) included
Pavetta polyantha within their circumscription of P. indica var. glabrescens (Kurz) Deb & Rout. The name P. polyantha is based on a Wallich specimen from Assam.

4. Pavetta scabrifolia Bremekamp, Repert. Spec. Nov. Regni

Veg. 37: 100. 1934.
cao ye da sha ye
Shrubs, height not noted; branches subcompressed to
angled, puberulent to pilosulous, green or almost green. Petiole

289

11.5 cm, pubescent; leaf blade drying membranous, lanceolate, 1316 3.24.4 cm, adaxially subglabrous except scabrous along midrib, abaxially scabrous, base cuneate, apex caudate; secondary veins 5 or 6 pairs; stipules broadly triangular,
cuspidate. Inflorescence terminal on branches with 1 long internode and several short internodes, laxly corymbose, pubescent.
Calyx subglabrous; limb ca. 1 mm, lobed for ca. 1/2. Corolla
white; tube ca. 17 mm, glabrous inside; lobes narrowly oblongtriangular, ca. 5.5 mm. Style ca. 38 mm. Drupes subglobose, ca.
5 mm in diam., glabrescent, calyx limb persistent. Fl. MayJun.
Sparse forests, ditch sides; 9001300 m. Yunnan.

5. Pavetta swatowica Bremekamp, Repert. Spec. Nov. Regni

Veg. 37: 104. 1934.
shan tou da sha ye
Shrubs, height not noted; branches compressed to terete,
pilosulous, green or almost green becoming grayish or blackened when dry. Petiole 12 cm, pilosulous; leaf blade drying
thinly papery or submembranous, narrowly elliptic or oblanceolate, 1013 ca. 5 cm, adaxially glabrescent, abaxially pilosulous, base cuneate, apex long acuminate; secondary veins 6 or 7
pairs; stipules ovate-triangular, ca. 5 mm, puberulent, shortly
cuspidate. Inflorescence terminal on developed branches, laxly
corymbose, many flowered, pubescent; peduncles 23 cm;
pedicels 46 mm. Flowers pedicellate. Calyx with hypanthium
subglobose, densely grayish pilosulous or sericeous; limb shallowly lobed. Corolla white, outside glabrous; tube ca. 14 mm,
sparsely tomentose inside; lobes narrowly oblong or oblonglanceolate, ca. 6 mm. Style ca. 35 mm. Drupes unknown.
Sparse forests, streamsides. Guangdong.
This name was incorrectly spelled by W. C. Ko (in FRPS 71(2):
2627. 1999) as Pavetta swatouica. Kos description, perhaps following that of Bremekamp, described the stipules as caducous,
although the stipules are otherwise considered persistent in Pavetta
by other authors, including Bremekamp.

6. Pavetta tomentosa Roxburgh ex Smith in Rees, Cycl. 26:

Pavetta no. 2. 1814.
rong mao da sha ye
Ixora tomentosa (Roxburgh ex Smith) Roxburgh; I. tomentosa var. roxburghii Kurz; Pavetta indica Linnaeus var. tomentosa (Roxburgh ex Smith) J. D. Hooker; P. tomentosa var. roxburghii (Kurz) Bremekamp.
Shrubs or small trees, height not noted; branches tomentose. Petiole 13 cm, tomentose; blade drying thickly papery
and blackening, elliptic or elliptic-lanceolate, 815 49 cm,
scabrous adaxially, tomentose abaxially, base acute, apex acute
or acuminate; secondary veins 1012, with foveolate and/or pubescent domatia; stipules ovate-triangular, 68 mm, tomentose,
aristate. Inflorescences terminal on branches with 1 or 2 internodes, laxly corymbose, 510 525 cm, densely tomentose,
pedunculate; peduncle 0.51.5 cm; pedicels 48 mm. Flowers
pedicellate. Calyx densely tomentose; hypanthium portion ellipsoid, ca. 1 mm; limb ca. 0.5 mm, denticulate to lobed. Corolla
white, outside glabrous; tube (6.2)812 mm, sparsely pilose
inside; lobes narrowly oblong, 4.67 mm, acute. Style 2433

RUBIACEAE

290

mm. Drupes globose, ca. 5 mm, tomentulose. Fl. and fr. Jul
Sep.
Tropical rain forests; ca. 1000 m. Yunnan (Menghai) [India,
Malaysia, Myanmar, Nepal, Pakistan, Thailand, Vietnam].

Rout and Deb (Bull. Bot. Surv. India 41: 128136. 1999) treated
this species as Pavetta indica var. tomentosa. Bremekamp (Repert.
Spec. Nov. Regni Veg. 37: 113114. 1934) and Rout and Deb described
the corolla tubes as 812 mm, but W. C. Ko (in FRPS 71(2): 28. 1999)
gave the minimum measurement as 6.2 mm.

68. PENTAS Bentham, Bot. Mag. 70: t. 4086. 1844.

wu xing hua shu
Chen Tao (); Charlotte M. Taylor
Perennial herbs or subshrubs, unarmed. Raphides present. Leaves opposite or whorled, without domatia; stipules persistent,
interpetiolar and sometimes fused to petioles, triangular to truncate, multifid or setaceous, with segments often gland-tipped. Inflorescences terminal, cymose, paniculate, or corymbiform, several to many flowered, subsessile to pedunculate, bracteate. Flowers
subsessile, bisexual, distylous [or rarely tristylous]. Calyx limb deeply (4 or)5-lobed; lobes sometimes slightly to strongly unequal on
an individual flower [sometimes with calycophylls]. Corolla white, pink, red, purple, or yellow, salverform to narrowly tubular with
tube prolonged and usually expanded at throat, inside villous in throat; lobes (4 or)5(or 6), valvate in bud. Stamens (4 or)5(or 6),
inserted in corolla just below throat, included in long-styled, mid-styled, and homostylous forms or exserted in short-styled form;
filaments short to well developed; anthers dorsifixed. Ovary 2-celled, ovules numerous in each cell on axile placentas; stigmas 2,
linear, included in short-styled and mid-styled forms or exserted in long-styled and monomorphic forms. Fruit capsular, subglobose
to ovoid or obovoid with apex often elongated into conical beak, membranous or leathery, loculicidally dehiscent through apical portion, with calyx limb persistent; seeds numerous, small, angled to subglobose.
About 50 species: widespread in Africa and Madagascar, one species cultivated worldwide; one species (introduced) in China.

1. Pentas lanceolata (Forsskl) Deflers, Voy. Yemen, 142.

1889.
wu xing hua
Ophiorrhiza lanceolata Forsskl, Fl. Aegypt.-Arab. 42.
1775; Pentas carnea Bentham.
Subshrubs, 3070 cm tall; branches quadrangular, moderately to densely pilosulous to villous often becoming glabrescent with age. Leaves opposite; petiole 0.53 cm, pilosulous to
villous; blade drying papery, oblong-lanceolate to ovate, 514
25.5 cm, adaxially scabrous or villous to glabrescent, abaxially
densely villous or hirtellous at least along principal veins, base
cuneate to obtuse, apex acute or shortly acuminate; secondary
veins 810 pairs; stipules truncate to broadly rounded, 1.52
mm, villous, bristles 15, 14 mm. Inflorescence densely pilosulous to villous; peduncle 312 mm; branched portion congested-cymose often becoming lax, 1.54 1.54 cm; bracts
narrowly triangular to linear, 0.51.5 mm. Flowers sessile or

subsessile, distylous. Calyx densely hirtellous or villous; ovary

portion subglobose to obovoid, ca. 1 mm; limb deeply lobed;
lobes narrowly oblanceolate to elliptic or narrowly spatulate, 2
8 mm, usually unequal on an individual flower with nearly this
entire size range found on some flowers, acute. Corolla pale
purple to pink, red, white, or yellow, salverform, sparsely hirtellous to glabrescent outside; tube slender except rather abruptly
swollen in throat in long-styled form (around stamens), 1720
mm, densely barbate in throat; lobes elliptic or oblong-lanceolate, 34 mm, acute to obtuse. Capsules obovoid, 46 46
mm, stiffly papery to woody, with beak 12 mm tall; seeds 0.5
1 mm. Fl. JulSep.
Cultivated in gardens in S China; sea level to 1500 m. Fujian,
Guangdong [native to Africa (Democratic Republic of Congo, Ethiopia, Kenya, Rwanda, Sudan, Tanzania, Uganda); commonly cultivated worldwide].
Cultivated plants of Pentas lanceolata are usually all long-styled
and usually do not set fruit. They are variable in flower color, and new
varieties and colors continue to be developed.

69. PERTUSADINA Ridsdale, Blumea 24: 353. 1979.

cao lie mu shu
Chen Tao (); Charlotte M. Taylor
Trees or shrubs, unarmed; trunk often fluted or latticed. Raphides absent. Leaves opposite, usually with domatia; stipules caducous, triangular, entire or shortly bifid. Inflorescences axillary or terminal, capitate with 15 globose heads in fascicles or cymes,
pedunculate, bracteate; bracteoles spatulate to linear-spatulate. Flowers sessile, bisexual, monomorphic. Calyx lobes 5, spatulate, obtuse. Corolla yellow, salverform to slenderly funnelform, inside glabrous; lobes 5, valvate or with apices subimbricate in bud. Stamens 5, inserted in upper part of corolla tube, exserted; filaments short; anthers basifixed. Ovary 2-celled, ovules 410 in each cell,
pendulous on axile placentas attached to upper third of septum; stigma globose to obovoidal, exserted. Fruiting heads globose. Fruit
capsular, obconic, septicidally then loculicidally dehiscent into 2 or 4 valves from base to apex with valves separating also from persistent or tardily deciduous septum, cartilaginous to ligneous, with calyx limb persistent on septum; seeds several, small, ovoid-trigonous, bilaterally compressed, winged at both ends.
Four species: China, Malay Peninsula, Moluccas, New Guinea, Philippines, Thailand; one species in China.

RUBIACEAE

1. Pertusadina metcalfii (Merrill ex H. L. Li) Y. F. Deng & C.

M. Hu, Blumea 51: 559. 2006.
hai nan cao lie mu
Adina metcalfii Merrill ex H. L. Li, J. Arnold Arbor. 24:
454. 1943; A. affinis F. C. How; A. hainanensis F. C. How; A.
polycephala Bentham var. glabra F. C. How; Pertusadina
hainanensis (F. C. How) Ridsdale.
Shrubs to large trees, perhaps evergreen, to 30 m tall;
branches angled to terete, reddish brown becoming gray, puberulent to glabrous, with scattered lenticels. Petiole 325 mm,
glabrous or puberulent; leaf blade drying thickly papery, elliptic
to elliptic-oblong or oblanceolate, 412 1.55 cm, adaxially
glabrous and rather shiny, abaxially glabrous to puberulent, base
cuneate to obtuse, apex acuminate; lateral veins 610 pairs, in
abaxial axils with foveolate and/or pilosulous domatia; stipules
linear-oblong to subulate, 46 12 mm, glabrous, apex acute
or occasionally emarginate to shortly bifid. Inflorescence puberulent to glabrescent; peduncles 16.5 cm; flowering heads

291

solitary on peduncles or occasionally 3 in a cyme, 68 mm in

diam. across calyces, ca. 15 mm in diam. across corollas; bracteoles linear-clavate to linear-spatulate, 0.51 mm, at apex ciliate. Calyx puberulent to pilosulous at least at junction of ovary
and limb; ovary portion obconic, 0.50.7 mm, surrounded at
base by ring of trichomes 0.20.5 mm; limb deeply lobed; lobes
linear-oblong to spatulate, 1.52 mm. Corolla with tube 22.5
mm, glabrous inside; lobes deltoid, 0.71 mm. Stigmas obovoid, ca. 0.3 mm, exserted for 78 mm. Fruiting head 1012
mm in diam. Capsules 24 mm, puberulent to strigose; seeds 1
2 mm. Fl. MayJun, fr. SepDec.
Dense forests; 100900 m. Fujian, Guangdong, Guangxi, Hainan,
Hunan, Zhejiang [Thailand].
Pertusadina metcalfii was illustrated by How (Sunyatsenia 6: 239,
f. 28; 241, f. 29. 1946, the first as Adina affinis and the second as
Adina hainanensis). Ridsdale (Blumea 24: 354. 1979) treated this
species as P. hainanensis, but, as noted by Deng and Hu (loc. cit.), the
names listed in synonymy by him included an older name that has
priority, A. metcalfii, confirmed by them as conspecific.

70. PHUOPSIS (Grisebach) Bentham & J. D. Hooker, Gen. Pl. 2: 151. 1873.
chang zhu cao shu
Chen Tao (); Friedrich Ehrendorfer
Asperula sect. Phuopsis Grisebach, Spic. Fl. Rumel. 2: 167. 1846.
Herbs, perennial, rhizomatous. Raphides present. Leaves and leaflike stipules in whorls of 610, sessile, without domatia, with
margins minutely antrorsely aculeolate. Inflorescences terminal, pedunculate, capitate, many flowered, enclosed by a whorl of free
leaflike bracts, flowers subtended by acuminate bracts. Flowers sessile, bisexual, monomorphic. Calyx limb obsolete. Corolla pink,
slender, salverform, inside glabrous; lobes 5, valvate in bud. Stamens 5, inserted in corolla tube, included to partially exserted;
filaments short; anthers dorsifixed. Ovary 2-celled, smooth, ovules 1 in each cell, erect, basal, anatropous; stigma clavate, shortly 2lobed at apex, strongly exserted. Fruit schizocarpous, 2 mericarps ellipsoid to obovoid, dry, rather hard, indehiscent, 1-celled, each
with 1 ellipsoid to curved, smooth to striate seed.
One species: SW Asia (Azerbaijan, NW Iran), occasionally cultivated as an ornamental in China.
Phuopsis belongs to the core genera of the tribe Rubieae (subtribe Rubiinae). Remote and DNA-supported affinities exist with Crucianella,
some groups of Asperula, and particularly with the widespread annual Sherardia. In the present volume Phuopsis is briefly discussed in the introduction to the genus Galium and keyed out there. Phuopsis is a monotypic relict genus restricted to the small Hyrcanian area of NW Iran and Talysh in
S Azerbaijan (Ehrendorfer et al., Fl. Iranica 176: 1287. 2005). Its single species has secondary pollen presentation and is butterfly-pollinated.

1. Phuopsis stylosa (Trinius) Bentham & J. D. Hooker ex B. D.

Jackson, Index Kew. 2: 505. 1894.
chang zhu hua
Crucianella stylosa Trinius, Mm. Acad. Imp. Sci. St.
Ptersbourg Hist. Acad. 6: 485. 1818.
Herbs, 2060(70) cm tall; branches quadrangular, glabrous to sparsely hairy, at least at nodes and along angles
retrorsely aculeolate. Leaves drying papery, narrowly lanceolate, narrowly elliptic, or narrowly oblanceolate, 1230 1.56
mm, both surfaces glabrous, base acute to attenuate, margin and

midrib abaxially thickened and antrorsely aculeolate, apex

acute to acuminate with acicular tip; secondary veins not
visible. Inflorescences hemispherical to subglobose, 1.53 cm
wide (not including involucral leaves); bracts leaflike, lanceolate, 812 mm, glabrescent, marginally and abaxially spinulose
with acute apex. Ovary ellipsoid, ca. 1 mm, glabrous. Corolla
glabrous outside; tube 912 mm; lobes 5, 12 mm, oblongovate, apex acute and somewhat thickened. Mericarps oblongobovate, ca. 1.5 mm. Fl. MayAug, fr. AugSep.
Cultivated in Shaanxi (Wugong) [native in deciduous forests of
SW Asia (Azerbaijan, NW Iran)].

71. PORTERANDIA Ridley, Bull. Misc. Inform. Kew 1939: 593. 1940.
juan guan qian shu
Chen Tao (); Charlotte M. Taylor
Randia sect. Anisophyllea J. D. Hooker, Fl. Brit. India 3: 113. 1880.
Shrubs or trees, [sometimes dioecious], unarmed. Raphides absent. Leaves opposite, sometimes anisophyllous, sometimes with

RUBIACEAE

292

domatia; stipules persistent, interpetiolar to shortly united around stem, triangular. Inflorescences pseudoaxillary [or terminal],
cymose to thyrsiform, few to several flowered, pedunculate, bracteate. Flowers sessile to pedicellate, bisexual [or unisexual]. Calyx
limb well developed, 5-lobed. Corolla white, salverform, outside densely sericeous, inside variously glabrous or pubescent; lobes
5(or 6), convolute in bud. Stamens 5, inserted in upper part of corolla tube, included or partially exserted; filaments very short or
absent; anthers dorsifixed. Ovary 2-celled, ovules numerous in each cell on axile placentas; stigma clavate or fusiform, bifid, striate,
included or perhaps exserted. Fruit perhaps yellow to brown, baccate, fleshy to woody, subglobose or obovoid, with calyx limb
tardily deciduous; seeds numerous, medium-sized, ellipsoid or reniform, flattened, embedded in pulp.
About 22 species: S and SE Asia, Pacific islands; one species (endemic) in China.
Formerly several African species were included in Porterandia, but these have separated as Aoranthe Somers (see Somers, Bull. Jard. Bot. Natl.
Belg. 58: 4775. 1988). Puff et al. (Rubiaceae of Thailand, 64. 2005) reported that the calyx limb is persistent in fruit in Porterandia, but their own
figure seems to show it deciduous; on herbarium specimens of various Porterandia species the calyx limb appears to fall as the fruit approach mature
size. W. C. Chen (in FRPS 71(1): 386. 1999) reported that the corolla lobes are rarely 6, but this number has not been reported by other authors; it is
not clear if this number is consistent or is a report of infrequent unusual flowers, which occurs periodically in many species of Rubiaceae. W. C. Chen
and Puff et al. described the stamens and stigmas as included, but they are shown as exserted in Chens figure (loc. cit.: 385, t. 101). W. C. Chen (loc.
cit.: 384) also described the ovary as incompletely 4-celled, but no other authors have reported this.

1. Porterandia sericantha (W. C. Chen) W. C. Chen, Fl. Reipubl. Popularis Sin. 71(1): 384. 1999.
juan guan qian
Randia sericantha W. C. Chen, Guihaia 7: 298. 1987.
Shrubs or trees, 18 m tall; branches compressed to terete,
moderately to densely ferruginous hirtellous. Petiole 315 mm,
moderately to densely pilosulous or hirtellous to glabrescent;
leaf blade drying papery, elliptic or oblanceolate-oblong, 5.516
25 cm, adaxially and sparsely strigillose, abaxially sparsely
to densely pilosulous or hirtellous, base acute to obtuse, apex
acute to usually shortly acuminate; secondary veins 812 pairs,
in abaxial axils with pilosulous domatia; stipules ovate to trian-

gular, 57 mm, moderately to densely strigillose to hirtellous or

glabrous. Inflorescences few to several flowered, cymose, 25.5
35 cm, densely pilosulous to hirtellous, sessile to pedunculate; peduncle to 0.5 cm; bracts triangular, lanceolate, or stipuliform, ca. 3 mm, acute to acuminate; pedicels 515 mm. Calyx
densely pilosulous and/or strigose; ovary portion obconic, ca. 2
mm; limb 56 mm, lobed partially to deeply; lobes ovate-lanceolate, acute. Corolla white, outside densely yellow sericeous;
tube ca. 30 3.5 mm, inside glabrous except with a villous ring
at middle; lobes ovate-elliptic, ca. 12.5 6 mm. Stigma ca. 1.5
mm. Berry subglobose, 815 mm in diam., pilosulous to strigillose; seeds ca. 4 mm. Fl. MayJun, fr. AugJan.
Forests or thickets at streamsides in valleys or on mountain
slopes; 3001500 m. Guangxi, Yunnan.

72. PRISMATOMERIS Thwaites, Hookers J. Bot. Kew Gard. Misc. 8: 268. 1856.
nan shan hua shu
Chen Tao (); Charlotte M. Taylor
Shrubs or small trees, unarmed; branches usually longitudinally ridged on each side and surrounded at base by persistent leafless stipules, with bark often yellowed and hardened. Raphides present. Leaves opposite, decussate on ascending branches and distichous on horizontal branches, without domatia; stipules persistent at least on younger nodes, interpetiolar, bilobed, often becoming
hardened with age. Inflorescences terminal on principal branches, or terminal on axillary short shoots and apparently axillary, umbelliform to fasciculate and several flowered or reduced to 1 flower, sessile to pedunculate, bracteate with bracts usually reduced. Flowers pedicellate or sessile, bisexual, usually distylous [occasionally fused by their ovaries]. Calyx limb truncate or 4 or 5(or 6)-lobed,
with lobes infrequently unequal with 1 larger than others. Corolla white, salverform, inside glabrous; lobes (4 or)5(or 6), valvate in
bud. Stamens (4 or)5(or 6), inserted at or above middle of corolla tube, included in long-styled flowers or partially exserted in shortstyled flowers; filaments short; anthers dorsifixed. Ovary 2-celled, ovules 1(or 2) in each cell, on axile placentas attached near or
above middle of septum; stigma 2-lobed, exserted in long-styled flowers, included in short-styled flowers. Infructescences occasionally displaced to pseudoaxillary by subsequent branch growth. Fruit simple [or infrequently multiple], purple-black to blueblack, drupaceous, fleshy, subglobose or globose, with calyx limb persistent; pyrenes 1 or 2, 1-celled, subglobose when solitary to
plano-convex when paired, thin-walled, with membranous preformed germination slits; seeds medium-sized, subglobose to planoconvex, on ventral face with deeply concave hilum; testa membranous; endosperm corneous; embryo small, with hypocotyl; radicle
hypogynous.
Fifteen species: Bangladesh, Borneo, Cambodia, China, India (including Andaman Islands), Indonesia, Laos, Malaysia, Myanmar, Philippines,
Singapore, Sri Lanka, Thailand, Vietnam; one variable species in China.
This genus was revised in a broad treatment by Johansson (Opera Bot. 94: 162. 1987), who reported that the flowers are strongly fragrant. This
genus was also treated for China almost simultaneously and apparently independently by Ruan (Acta Phytotax. Sin. 26: 443449. 1988), whose
conclusions differed markedly from those of Johansson.
Two species have been recognized in China, Prismatomeris tetrandra and P. connata, with two subspecies recognized for P. connata (Y. Z.
Ruan in FRPS 71(2): 178179. 1999); P. tetrandra subsp. multiflora included only plants from Yunnan, while plants of the rest of China were treated

RUBIACEAE

293

in P. connata, with plants from the mainland in subsp. connata and plants from Hainan in subsp. hainanensis. However, wide and continuous morphological variation was documented by Johansson (loc. cit.) and Puff et al. (Rubiaceae of Thailand, 118. 2005) within P. tetrandra in adjacent countries, both across the region and in local populations; and Johansson (in herb.) recognized one species and no infraspecific taxa in China.
Prismatomeris connata was distinguished originally by its connate rather than separate stigmas; however, this character was later said by Y. Z. Ruan to
vary within this species. Its subspecies were distinguished by the degree of lobing of the calyx limbs and the shape and degree of swelling of the stigmas, but as there appears on the specimens studied to be continuous variation in these characters and some plants from the mainland match the characters given for the Hainan subspecies these plants are not separated here.

1. Prismatomeris tetrandra (Roxburgh) K. Schumann in

Engler & Prantl, Nat. Pflanzenfam. 4(4): 138. 1891.
si rui san jiao ban hua
Coffea tetrandra Roxburgh, Fl. Ind. 2: 193. 1824;
Octotropis terminalis C. B. Clarke; Prismatomeris connata Y.
Z. Ruan; P. connata subsp. hainanensis Y. Z. Ruan; P. multiflora Ridley; P. tetrandra subsp. multiflora (Ridley) Y. Z. Ruan.
Shrubs or small trees, to 8 m tall; branches quadrangular
to subterete, glabrous. Petiole 415 mm, glabrous; leaf blade
drying leathery to stiffly papery and rather shiny, lanceolate,
elliptic, elliptic-oblong, ovate, obovate, or oblong-lanceolate,
418 26 cm, glabrous, base cuneate to acute, apex acuminate or acute to obtuse; secondary veins 59 pairs; stipules with
sheath portion 0.21 mm, truncate, persistent or deciduous by
fragmentation, bilobed, lobes linear to narrowly triangular, 0.1
0.5 mm, caducous. Peduncles 116 and fascicled or umbellate,
535 mm, simple or bearing an umbellate group of pedicels, at
base with stipuliform bracts; pedicels when present 515 mm.
Flowers pedunculate or pedicellate. Calyx glabrous or sparsely
glandular-puberulent; hypanthium portion hemispherical, 1.52
mm; limb 12 mm, truncate or 4- or 5-denticulate. Corolla

white or pale purple, salverform, glabrous outside; tube 1420

mm; lobes 4 or 5, lanceolate, 710 mm, abaxially (i.e., dorsally)
ridged at least in bud, acute to obtuse. Drupes subglobose, 812
mm in diam., glabrous, smooth. Fl. MaySep, fr. SepDec.
Forests, thickets; 3002400 m. Fujian, Guangdong, Guangxi, Hainan, S Yunnan [Cambodia, India, Thailand, Vietnam].
Johansson (Opera Bot. 94: 29. 1987) recognized two subspecies,
Prismatomeris tetrandra subsp. tetrandra and P. tetrandra subsp. malayana (Ridley) J. T. Johannson; he reported only subsp. tetrandra from
China. However, Puff et al. (Rubiaceae of Thailand, 118. 2005) noted
that further observation shows that even these taxa cannot be completely separated either morphologically or geographically. Y. Z. Ruan
(in FRPS 71(2): 178. 1999) included the Chinese plants in P. tetrandra
subsp. multiflora, which Johansson synonymized with subsp. tetrandra.
Ruan (loc. cit.) reported that the flowers of P. connata are occasionally
unisexual; this was not further documented there and has not been
reported by others nor confirmed with specimens. Ruan (loc. cit.: 179)
also described the calyx lobes as ca. 3 mm, which has not been reported
by other authors nor seen on the rather ample materials studied.
This species was newly reported from India by Ayappan and
Parthasarathy (J. Econ. Taxon. Bot. 29: 802804. 2005), and its distribution in Cambodia and Vietnam is here reported based on specimens at
P annotated by Johansson after the publication of his article.

73. PSEUDOPYXIS Miquel, Ann. Mus. Bot. Lugduno-Batavi 3: 189. 1867.

jia gai guo cao shu
Chen Tao (); Charlotte M. Taylor
Herbs, perennial, low, unarmed, rhizomatous, sometimes with fetid odor. Raphides present. Leaves opposite or sometimes
basalmost pairs reduced to prophylls, without domatia; stipules persistent, interpetiolar and fused to petioles, triangular or 3(or 5)lobed, often glandular-dentate. Inflorescences terminal and in uppermost leaf axils (or subtended by leaflike bracts), fasciculate and
2- to several flowered or sometimes 1-flowered, pedunculate or subsessile, bracteate or bracts reduced. Flowers pedunculate or
pedicellate, bisexual, apparently monomorphic. Calyx limb deeply 5-lobed, usually markedly reticulate-veined, with lobes often
unequal. Corolla white or purplish red, slenderly tubular-funnelform to salverform, inside glabrous or pubescent in throat; lobes 5,
valvate-induplicate in bud. Stamens 5, inserted at base of corolla tube, included to exserted; filaments short to developed; anthers
dorsifixed. Ovary 4- or 5-celled, ovules 1 in each cell, anatropous on basal placentas; stigmas 2-, 4- or 5-lobed, exserted. Fruit capsular, obconic to hemispherical, papery, dehiscent through apical lid or operculum within persistent calyx limb, with pedicels becoming reflexed; seeds 35, obovoid, longitudinally grooved (or with many longitudinal idioblasts containing raphides).
Three species: China, Japan; one species (endemic) in China.
H. S. Lo (in FRPS 71(2): 153155. 1999) reported one species from China, Pseudopyxis heterophylla Maximowicz, with morphological characters that include some characters of both P. heterophylla and P. depressa Miquel as treated by the Fl. Japan (3a: 229. 1993). These two species were
both reported there to be endemic to Japan: P. depressa with a height of 38 cm, uniformly distributed stem pubescence, purple corollas 2025 mm,
very short filaments, and 5 stigmas; and P. heterophylla with a height to 50 cm, stem pubescence arranged in lines, white corollas 67 mm, developed
filaments, and 2 stigmas.
Subsequently, Chen (Edinburgh J. Bot. 64: 303309. 2007) recognized three species in the genus, describing the Chinese plants as a new
species, Pseudopyxis monilirhizoma, mainly based on root and stem characters (i.e., including number of leaf pairs). He considered this the species
that was treated as P. heterophylla by H. S. Lo (in FRPS 71(2): 153155. 1999).
One of us (Taylor) does not fully agree with the species treatment here but has not had adequate access to all the materials and specimens cited
by Chen to evaluate the situation completely. The species circumscription and description given here do not seem to include or account for all the
specimens of Pseudopyxis from China.

RUBIACEAE

294

1. Pseudopyxis monilirhizoma Tao Chen, Edinburgh J. Bot.

64: 304. 2007.
zhang jie jia gai guo cao
Herbs, to 10 cm tall, with fetid odor; rhizomes sparsely
branched, with nodes tuberous and conspicuously enlarged, with
internodes usually more than 2 cm; stems quadrate, glabrous
except densely hirtellous or pilosulous in 2 interpetiolar rows.
Leaves 2 or 3 pairs; petiole 0.32 cm, glabrous or densely pilosulous abaxially; blade drying membranous, deltoid-ovate,
0.84.5 0.63.5 cm, sparsely hispidulous on both surfaces,
base cuneate to truncate, margins entire or ciliate, apex acute or

obtuse; secondary veins 4 or 5 pairs; stipules ca. 0.5 mm. Inflorescences 26-flowered; peduncles 24 mm, puberulent. Calyx
hispidulous in lines; hypanthium portion obconic, longitudinally ribbed, ca. 1 mm; lobes ovate to ovate-lanceolate, ca. 2
mm. Corolla white or pink, salverform, externally glabrous, internally pilosulous in tube and on lobes; tube ca. 5 mm; lobes
lanceolate or narrowly elliptic-oblong, ca. 4 mm, acute to subacute. Capsules obconic, size not noted; seeds ca. 1.2 mm. Fl.
JunAug, fr. AugOct.
Wet sites in rock crevices or on stream banks in forest understories; 14001600 m. Zhejiang (Longquan).

74. PSYCHOTRIA Linnaeus, Syst. Nat., ed. 10, 2: 906, 929, 1364. 1759, nom. cons.
jiu jie shu
Chen Tao (); Charlotte M. Taylor
Cephaelis Swartz.
Shrubs, small trees, or rarely vines twining and/or climbing by adventitious roots (Psychotria serpens), [infrequently dioecious
or polygamo-dioecious], unarmed, tissues and/or pubescence often drying dark gray or dark reddish brown. Raphides present. Leaves
opposite or rarely in whorls of 3 or 4, often with foveolate and/or pubescent domatia; stipules caducous or infrequently persistent,
interpetiolar or sometimes shortly united around stem, entire or 2-lobed, rarely with lobe glandular, inside (i.e., adaxially) at base
with well-developed colleters, these usually persistent after stipule falls, usually drying red-brown. Inflorescences terminal often becoming displaced to pseudoaxillary or rarely axillary, cymose, corymbose, paniculiform, glomerulate, or capitate, several to many
flowered, sessile to pedunculate, bracteate with bracts sometimes reduced or sometimes enlarged or involucrate. Flowers sessile to
pedicellate, bisexual, usually distylous [or infrequently unisexual]. Calyx limb (4 or)5(or 6)-lobed. Corolla white, yellow, or flushed
with pink, funnelform to tubular, inside glabrous or variously pubescent, lobes (4 or)5(or 6), valvate in bud, sometimes abaxially
with thickenings or horns near apex. Stamens (4 or)5(or 6), inserted in corolla tube or throat, usually included or partially exserted in
long-styled flowers and exserted in short-styled flowers; filaments short to developed; anthers dorsifixed near base. Ovary 2-celled,
ovules 1 in each cell, basal; stigmas 2, linear to subcapitate, usually exserted in long-styled flowers and included in short-styled flowers. Fruit red, orange, or infrequently white (P. serpens), purple (P. manillensis), or black (P. cephalophora, P. straminea), drupaceous, fleshy, ellipsoid, ovoid, or subglobose, with calyx limb persistent or infrequently deciduous, with pedicels or stipitate base
sometimes elongating; pyrenes 2, 1-celled, each with 1 seed, plano-convex, bony, on dorsal (i.e., abaxial) surface smooth or longitudinally ridged, on ventral surface smooth or longitudinally sulcate; seeds medium-sized, ellipsoid to plano-convex, with testa thin;
endosperm fleshy or corneous, sometimes ruminate; embryo small, basal; cotyledon flat.
About 8001500 species: tropical and subtropical Africa, America, Asia, Madagascar, and Pacific islands; 18 species (five endemic) in China.
The genus Cephaelis was separated from Psychotria by numerous authors in the 19th and first part of the 20th centuries, based on inflorescence
form: Cephaelis included species with capitate inflorescences with enlarged, often involucral bracts, vs. branched inflorescences with smaller bracts in
Psychotria. However, it is now clear that this inflorescence arrangement has arisen far more than once within this group and that Cephaelis actually
included a polyphyletic set of species that are more closely related to various other species of Psychotria than to each other. Consequently, recent
authors (e.g., Steyermark, Mem. New York Bot. Gard. 23: 443717. 1972; Taylor, Opera Bot. Belg. 7: 261270. 1996) have formally synonymized
Cephaelis with Psychotria.
A recent treatment of Psychotria in the Philippines (Sohmer & Davis, Sida, Bot. Misc. 27: 1247. 2007) does not consider any species or names
outside its study area but includes some Chinese species that occur in that region. However, these authors have a partially different species concept
and morphological interpretation of inflorescence characters from C. M. Taylor, so their work is not completely comparable to the treatment here.

1a. Climbing or creeping vines or lianas with adventitious roots, often on tree trunks or rocks; fruit white .................... 14. P. serpens
1b. Erect, self-supporting subshrubs, shrubs, and small trees growing on ground; fruit orange or red.
2a. Leaves strigose to hirsute in similar density on both sides; stipules 1525 mm, deeply bilobed .......................... 12. P. pilifera
2b. Leaves glabrous adaxially, or pubescent adaxially with distinctly different pubescence than on abaxial
surface; stipules 1.520 mm, entire to bilobed.
3a. Inflorescences capitate, pedunculate, and enclosed by a cupuliform involucre formed of fused bracts ................. 9. P. laui
3b. Inflorescences capitate to branched, sessile to pedunculate, and variously bracteate, bracts when
present free and not forming a single involucre.
4a. Calyx limb 1.33 mm, shallowly to deeply lobed.

RUBIACEAE

295

5a. Inflorescences thyrsiform to paniculate, branched to 2 or 3 orders and with well-developed

secondary axes ............................................................................................................................. 18. P. yunnanensis
5b. Inflorescences capitate, subcapitate, or congested-cymose, unbranched or branched to 1
or 2 orders but without well-developed secondary axes.
6a. Inflorescences capitate to densely congested-cymose, subglobose in outline, sessile
or with peduncle to 1 cm ................................................................................................................ 13. P. prainii
6b. Inflorescences capitate, subcapitate, or shortly congested-cymose, ellipsoid to ovoid
or pyramidal in outline, sessile to pedunculate with peduncles to 6 cm.
7a. Subshrubs, often rhizomatous; leaves with well-developed, generally straight
submarginal vein extending along most or all of length of blade ....................................... 2. P. calocarpa
7b. Shrubs or small trees; leaves without submarginal vein or with submarginal vein
incomplete or only weakly developed, extending for up to 1/22/3 of length
of blade.
8a. Inflorescences sessile or with peduncle up to 0.6 cm; stipules 312 mm; Hainan ... 7. P. hainanensis
8b. Inflorescences pedunculate, peduncle 0.56 cm; stipules 820 mm; Yunnan ........ 11. P. morindoides
4b. Calyx limb 0.51.2 mm, truncate to deeply lobed.
9a. Inflorescences capitate to subcapitate or congested-cymose, unbranched or branched but
without well-developed secondary axes, or with secondary axes but then primary axis
not developed.
10a. Flowers subsessile to pedicellate in a single head, with all flowers arising from one
axis ...................................................................................................................................... 3. P. cephalophora
10b. Flowers variously sessile to pedicellate in a subcapitate head or congested cyme,
flowers arising from more than one point or axis .......................................................................... 8. P. henryi
9b. Inflorescences thyrsiform, paniculate, corymbiform, or congested-cymose to laxly cymose,
branched, with both primary and secondary axes developed.
11a. Stipules 1015 mm with at least some of them more than 10 mm; leaves with
secondary veins 1218 pairs with 13 or more pairs on at least some leaves ................................. 4. P. densa
11b. Stipules 1.512 mm with at least some less than 10 mm; leaves with secondary veins
412 pairs with less than 12 pairs on at least some leaves.
12a. Inflorescences pyramidal, with primary axis developed and longer than
secondary axes .......................................................................................................... 16. P. symplocifolia
12b. Inflorescences rounded-corymbiform to broadly pyramidal, with primary axis
reduced to developed but not longer than secondary axes.
13a. Stipules fused around stem into a distinct sheath, mostly persistent with
leaves; leaves with secondary veins prominulous adaxially ............................... 15. P. straminea
13b. Stipules interpetiolar, caducous or deciduous, falling before some of leaves;
leaves with secondary veins flat or thinly impressed adaxially.
14a. Leaves narrowly elliptic, narrowly elliptic-oblong, narrowly lanceolate,
narrowly lanceolate-oblong, or oblanceolate, 415 14.5 cm;
stipules at least shortly 2-lobed.
15a. Leaves abaxially with distinctive thickened epidermis often mottled
when dry, secondary veins covered by epidermis or visible but flat
to only slightly thickened ..................................................................... 6. P. fluviatilis
15b. Leaves abaxially with epidermis normal, thin and not mottled
when dry, secondary veins visible, with different epidermal texture
from lamina, flat to prominulous.
16a. Plants usually drying dull green, grayish brown, yellowish green,
or reddish brown; inflorescences congested-cymose ...................... 8. P. henryi
16b. Plants usually drying reddish brown to dark brown;
inflorescences laxly cymose, corymbiform ................................. 17. P. tutcheri
14b. Leaves elliptic, broadly elliptic, broadly elliptic-oblong, ovate, lanceolateoblong, or obovate, 523.5 29 cm; stipules entire to 2-lobed.
17a. Bracts subtending flowers rather well developed, lanceolate to
ligulate or triangular, 1.53 mm; stipules 2-lobed ................................. 5. P. erratica
17b. Bracts subtending flowers developed to reduced, triangular,
0.21.2 mm; stipules obtuse, acute, rounded, or shallowly
emarginate.
18a. Inflorescences sessile or with peduncle to 0.3 cm; leaves
with secondary veins not or only weakly forming a
submarginal vein ............................................................................ 1. P. asiatica

296

RUBIACEAE

18b. Inflorescences pedunculate, peduncles 0.13.5 cm; leaves

with secondary veins free or usually forming a weak to
well-developed, looping submarginal vein ........................... 10. P. manillensis
1. Psychotria asiatica Linnaeus, Syst. Nat., ed. 10, 2: 929.
1759.

2. Psychotria calocarpa Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 41: 315. 1872.

jiu jie

mei guo jiu jie

Antherura rubra Loureiro; Psychotria esquirolii H.

Lveill; P. reevesii Wallich; P. reevesii var. pilosa Pitard; P.
rubra (Loureiro) Poiret; P. rubra var. pilosa (Pitard) W. C.
Chen; Uragoga rubra (Loureiro) Kuntze.

Subshrubs, 0.251 m tall, often rhizomatous; stems pilosulous to glabrous. Petiole 0.54 cm, glabrous or brown pilosulous; leaf blade drying papery, brownish red, gray, or dark
green, elliptic-oblong, elliptic, obovate-oblong, or elliptic-lanceolate, 9.517 2.57 cm, glabrous on both surfaces or
densely puberulent to hirtellous abaxially, base cuneate to acute,
margins flat, apex acute, acuminate, or rarely obtuse; secondary
veins 915 pairs, forming a generally straight submarginal vein,
without domatia; stipules caducous, ovate to suborbicular, interpetiolar, basal portion 410 mm, glabrous or puberulent to hirtellous, 2-lobed, lobes narrowly triangular, 26 mm, acuminate
to subulate. Inflorescences terminal or pseudoaxillary, congested-cymose, several to many flowered, hirtellous; peduncle
0.82 cm; branched portion corymbiform to subglobose, 13
24 cm, branched to 1 or weakly 2 orders; bracts lanceolate to
narrowly triangular, 0.85 mm; pedicels 16 mm. Flowers pedicellate. Calyx glabrous to puberulent; limb ca. 2 mm, deeply
lobed; lobes linear to linear-lanceolate, entire to ciliolate. Corolla white; tube ca. 2 mm, white villous inside; lobes elliptic-oblong, obtuse. Drupes red or orange, ellipsoid, 78.5 4.56
mm; pyrenes with 4 or 5 low ridges. Fl. MayJul, fr. AugFeb
of following year.

Shrubs or small trees, 0.55 m tall; stems puberulent to

glabrous. Petiole 0.75 cm, glabrous or rarely puberulent; leaf
blade reportedly rather shiny in life, drying papery to leathery,
dark red, brownish red, yellowish green, or gray-green, ellipticoblong, lanceolate-oblong, or rarely oblong-ovate, 523.5 29
cm, adaxially glabrous, abaxially glabrous or puberulent, base
acute to obtuse, margins flat to narrowly revolute, apex acute to
acuminate or obtuse then abruptly narrowed and acuminate;
secondary veins 58(11) pairs, free or weakly forming a
broadly looping, incomplete submarginal vein, usually with
foveolate and sometimes pilosulous domatia; stipules caducous
or sometimes persisting on 2 or 3 nodes, triangular to broadly
triangular or broadly ligulate, interpetiolar or shortly united
around stem, [3]68 mm, glabrous to puberulent, broadly
rounded to obtuse or shallowly emarginate. Inflorescences terminal or sometimes pseudoaxillary, cymose to paniculiform,
many flowered, glabrous or usually densely puberulent, sessile
and apparently tripartite to shortly pedunculate; peduncle to 0.3
mm; branched portion corymbiform-rounded to broadly pyramidal, 210 35 cm, with 13 pairs of developed secondary
axes; bracts triangular, 0.21 mm; pedicels to 2.5 mm. Flowers
subsessile to pedicellate in dichotomous cymules of 35. Calyx
densely puberulent; hypanthium portion turbinate, 0.81.2 mm;
limb 0.81 mm, subtruncate to denticulate. Corolla white, funnelform, glabrous outside; tube 23 mm, in throat white villous;
lobes triangular, 22.5 mm. Drupes red, subglobose to broadly
ellipsoid, 58 47 mm, with pedicels to 10 mm; pyrenes shallowly 35-ribbed. Fl. and fr. year-round.
Thickets or forests in ravines, on hill slopes, or at village margins;
near sea level to 1500 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Taiwan, Yunnan, Zhejiang, also cultivated in Zhejiang
[Cambodia, India, Japan, Laos (commonly collected), Malaysia, Thailand, Vietnam (very commonly collected)].
This species has long been called Psychotria rubra in China and
P. reevesii in Vietnam. However, recently Davis et al. (Bot. J. Linn. Soc.
135: 3452. 2001) matched the Chinese plants to the type specimen of
P. asiatica, which is thus the correct name for these plants. The objective of their work was limited, however, and they did not address the
whole circumscription and range of this species, nor the identity of P.
rubra. The Chinese specimens treated under this name here closely
match the specimens included by them in P. asiatica, and the range of
this species is here accordingly expanded to include this range. As noted
by Davis et al. (loc. cit.), all the flowers seen have the anthers exserted
and the stigmas included and positioned near the middle of the corolla
tube, thus resembling the short-styled form of distylous species of Psychotria.

Forests on mountain slopes; 8001700 m. Xizang, Yunnan [Bangladesh, Bhutan, India, Malaysia, Myanmar, Nepal, Thailand, Vietnam].
This species is used medicinally.
W. C. Chen (in FRPS 71(2): 51. 1999) described the calyx lobes
as 4, but they are 5 on all the specimens studied.

3. Psychotria cephalophora Merrill, Philipp. J. Sci., C, 3: 438.

1909.
lan yu jiu jie mu
Psychotria kotoensis Hayata.
Shrubs or small trees, height not noted; stems glabrous.
Petiole 1.26 cm, glabrous; leaf blade drying papery, pale reddish brown, dark brown, or green sometimes tinged with redpurple, elliptic, elliptic-oblong, or elliptic-ovate, 1016 3.5
7 cm, glabrous on both surfaces, base cuneate to acute, margins
flat, apex acute or acuminate; secondary veins 612 pairs, free
or forming a weak submarginal vein, without domatia; stipules
caducous, ovate, interpetiolar or shortly fused around stem, 7
10 mm, glabrous, ciliate, 2-lobed for ca. 1/5 of length, lobes ligulate to triangular. Inflorescences terminal, subcapitate to congested-cymose, several flowered, subsessile to sessile, glabrous
to densely hirtellous; bracts triangular to ligulate, 13 mm;
pedicels to 2 mm. Flowers subsessile to pedicellate. Calyx glabrous; hypanthium portion turbinate, ca. 1 mm; limb ca. 1 mm,
lobed for 1/31/2; lobes deltoid to narrowly triangular. Corolla

RUBIACEAE

in bud white, funnelform, glabrous outside; tube ca. 1.5 mm,

villous inside; lobes triangular-oblong, ca. 1.5 mm. Drupes becoming red then black, ellipsoid to obovoid, 810 68 mm,
with pedicels to 7 mm; pyrenes smooth or very shallowly 3- or
4-ribbed. Fl. Apr, fr. Aug.
Broad-leaved forests; below 100400 m. Taiwan (Lan Yu) [Philippines].
The stipules of these plants have been described by previous
authors as acute to obtuse, but these are bilobed on all the specimens
studied, although this is difficult to see in many cases because the lobes
are usually imbricate in bud.
This name is here provisionally applied to these plants. Sohmer
and Davis (Sida, Bot. Misc. 27: 6063. 2007) excluded the Taiwanese
plants from their circumscription of Psychotria cephalophora, without
providing another name for them (presumably P. kotoensis would be
available) or delimiting the differences between the two. They noted
that the endosperm of P. cephalophora is ruminate although the pyrenes
are smooth on the outer surface; the Taiwanese specimens studied do
appear to have ruminate endosperm.

4. Psychotria densa W. C. Chen, Acta Phytotax. Sin. 30: 270.

1992.
mi mai jiu jie
Shrubs, 13 m tall; stems densely dark hirtellous to tomentulose. Petiole 1.57.5 cm, densely hirtellous; leaf blade drying
papery or thinly leathery, dark brownish gray or greenish gray,
elliptic, elliptic-oblong, or oblong-oblanceolate, 1026 412.5
cm, glabrous adaxially, hirtellous abaxially with pubescence
denser along principal veins, base cuneate to obtuse, margins
flat, apex acute or shortly acuminate; secondary veins 1218
pairs, forming a weak to distinct submarginal vein, without
domatia; stipules caducous, broadly triangular to triangularovate, interpetiolar, 1015 mm, densely hirtellous to tomentulose, acute to acuminate or bilobed for 1/101/3, lobes ligulate
to deltoid, rounded to acute or acuminate. Inflorescences terminal, congested-cymose to thyrsiform, many flowered,
densely hirtellous to tomentulose, pedunculate; peduncle 13
cm; branched portion pyramidal to rounded, 37 2.57.5 cm;
bracts triangular to ovate, 14 mm, often glabrescent. Flowers
sessile or subsessile. Calyx glabrous; hypanthium portion obconic, ca. 2 mm; limb 0.51.2 mm, lobed for ca. 1/2; lobes
triangular. Corolla presumably white, tubular, glabrous outside;
tube ca. 4 mm, in throat white villous; lobes triangular-oblong,
ca. 2 mm. Drupes red, ellipsoid, 810 56 mm; pyrenes 3- or
4-ribbed. Fl. AprMay, fr. JunJan of following year.
Forests on mountains; 12001700 m. Yunnan (Hekou, Pingbian).
This name was validly published as cited above; the same article
was republished later in the same journal in the same year, with this
species treated on pp. 484486. The later treatment is sometimes cited
as the place of publication but is superfluous. The protologue described
the stipules as acuminate and this was illustrated in the figure, but a
paratype (X. Q. Liu 100347, MO!) has shortly bilobed stipules.

5. Psychotria erratica J. D. Hooker, Fl. Brit. India 3: 168.

1880.
xi zang jiu jie

297

Shrubs, 0.41 m tall; stems glabrous. Petiole 0.55 cm,

glabrous to puberulent; leaf blade drying papery to stiffly papery, dark green, greenish brown, or reddish brown, elliptic,
broadly elliptic, or obovate, 717 28 cm, glabrous adaxially,
glabrous to densely puberulent abaxially, base acute to cuneate,
margins flat, apex acute to acuminate; secondary veins 912
pairs, not forming a submarginal vein, without domatia; stipules
caducous, oblong-lanceolate to broadly ovate, interpetiolar, 5
12 mm, puberulent, strigillose, or hirtellous, 2-lobed for 1/4
1/2, lobes triangular to narrowly triangular sometimes with linear tip 29 mm, sometimes erose and glandular. Inflorescences
terminal or pseudoaxillary, cymose, sessile to pedunculate, puberulent to densely hirtellous with pubescence sometimes in
lines; peduncle to 6.5 cm; branched portion corymbiform, 2.5
4.5 46 cm, with 1 or 2 pairs of developed secondary axes;
bracts ligulate, lanceolate, or narrowly triangular, 0.53 mm,
sometimes 2-lobed, those subtending flowers 1.53 mm; pedicels to 1.5 mm. Flowers subsessile to pedicellate in dichasial
cymules of 311. Calyx glabrous to puberulent; hypanthium
portion obconic, 0.81.2 mm; limb 0.81.2 mm, lobed for 1/4
1/2; lobes triangular to broadly triangular. Corolla white to
pale green, tubular to tubular-funnelform, outside glabrous;
tube 3.54 mm, densely villous in throat; lobes triangular, ca.
2.5 mm, thickened at apex. Drupes red or yellow, ellipsoid to
obovoid, 68 56 mm, with pedicels to 5 mm; pyrenes shallowly 3- or 4-ridged. Fl. AprMay, fr. Aug, OctNov.
Subtropical evergreen broad-leaved forests on mountains; 1000
2400 m. Xizang (Mdog), Yunnan [Bhutan, India, Nepal].
This species, as represented by Indian specimens and circumscribed here, is quite variable in leaf size and shape, degree of stipule
lobing, and inflorescence form (in particular peduncle development).

6. Psychotria fluviatilis Chun ex W. C. Chen, Acta Phytotax.

Sin. 30: 268. 1992.
xi bian jiu jie
Shrubs, 0.43 m tall; stems glabrous. Petiole 0.51.8 cm,
glabrous; leaf blade drying papery to thinly leathery, grayish
olive-green, yellowish green, or occasionally dark brown
(Zhang Guicai 488, MO!), sometimes paler below, oblanceolate, narrowly elliptic, or narrowly elliptic-oblong, 511 13.7
cm, glabrous, abaxially with silvery or mottled thickened epidermis, base acute to cuneate, margins flat, apex deltoid, acute,
or acuminate; secondary veins 48 pairs, not forming a submarginal vein, without domatia; stipules caducous, lanceolate
or triangular, interpetiolar, 47 2.55.5 mm, glabrous, 2-lobed
for 1/101/4, lobes narrowly triangular to linear, sometimes
gland-tipped. Inflorescences terminal or often to pseudoaxillary,
cymose-paniculiform, few to several flowered, glabrous, sessile
to pedunculate; peduncle to 0.7 cm; branched portion corymbiform or broadly pyramidal, 13 11.5 cm, with 1 or 2 pairs
of developed secondary axes; bracts triangular to linear-lanceolate, 0.51.5 mm; pedicels to 2 mm. Flowers subsessile and
pedicellate in small dichotomous cymes. Calyx glabrous; hypanthium portion obconic, 11.5 mm; limb 0.51 mm, lobed
shallowly or for up to 1/5; lobes broadly triangular. Corolla
white, tubular, glabrous outside; tube 33.5 mm, in throat white
villous; lobes triangular-oblong, 11.7 mm. Drupes red, oblong-

RUBIACEAE

298

ellipsoid or subglobose, 67 36 mm, with pedicels to 10

mm; pyrenes shallowly 3- or 4-ribbed. Fl. AprSep, fr. Aug
Dec.
Forests along valley streams; 5001000 m. Guangdong, Guangxi.
This name was validly published as cited above; the same article
was republished later in the same journal in the same year, with this
species treated on pp. 482483. The later treatment is sometimes cited
as the place of publication but is superfluous.

7. Psychotria hainanensis H. L. Li, J. Arnold Arbor. 25: 213.

1944.
hai nan jiu jie
Shrubs, 0.53 m tall; stems glabrous. Petiole 0.43.5 cm,
glabrous; leaf blade drying papery, grayish brown, grayish
green, abaxially paler to silvery, elliptic, oblong-elliptic, lanceolate, or lanceolate-oblong, 4.516 26 cm, glabrous on both
surfaces or puberulent abaxially, base acute to cuneate, margins
flat, apex acute to acuminate; secondary veins 614 pairs,
forming a rather straight submarginal vein, without domatia;
stipules caducous, subtriangular to ovate, interpetiolar, 312
mm, glabrous to puberulent, 2-lobed for 1/31/2, lobes linearlanceolate to subulate. Inflorescences terminal, congested-cymose to subcapitate, few flowered, puberulent to glabrous, subsessile to pedunculate; peduncle to 0.6 cm; branched portion
subglobose, ca. 2 2 cm, without developed axes; bracts linearlanceolate, 27 mm; pedicels 0.54 mm. Flowers subsessile to
shortly pedicellate. Calyx glabrous; hypanthium portion turbinate, ca. 1 mm; limb 23 mm, deeply lobed; lobes linear-lanceolate to narrowly ligulate. Corolla white, glabrous outside;
tube ca. 4 mm, white villous in throat; lobes oblong-triangular,
1.52 mm, apex incurved. Drupes red, ovoid or elliptic, 711
4.57 mm, with pedicels to 8 mm; pyrenes 3- or 5-ribbed. Fl.
AprMay, fr. JunFeb of following year.

mm; pedicels to 1 mm. Flowers subsessile to pedicellate. Calyx

puberulent; hypanthium portion turbinate, ca. 1 mm; limb 0.8
1.2 mm, shallowly to deeply lobed; lobes triangular to narrowly
triangular. Corolla white, funnelform, glabrous outside; tube
1.52 mm, densely villous in throat; lobes triangular, 1.21.5
mm. Infructescences sometimes expanding, to 2 3.5 cm; pedicels to 2 mm. Drupes red, ovoid or globose, 47 35 mm; pyrenes shallowly 3- or 4-ridged. Fl. MayJun, fr. AugFeb of following year.
Forests; 11001500 m. Yunnan [Vietnam].

9. Psychotria laui Merrill & F. P. Metcalf, Lingnan Sci. J. 16:

403. 1937.
tou jiu jie
Cephaelis laui (Merrill & F. P. Metcalf) F. C. How & W.
C. Ko.
Shrubs, 12 m tall; stems glabrous. Petiole 0.30.6 cm,
glabrous; leaf blade drying papery, reddish brown, elliptic,
elliptic-oblong, or lanceolate, 511 1.53.5 cm, glabrous on
both surfaces, base acute to cuneate, margins flat or weakly
crisped, apex acute to shortly acuminate; secondary veins 68
pairs, not forming a submarginal vein, without domatia; stipules
caducous, triangular, interpetiolar or shortly united around stem,
25 mm, glabrous, acute. Inflorescences terminal, capitate, glabrous to puberulent, pedunculate; peduncle 13.5 cm; heads 1
or rarely 2, hemispherical, 12 cm in diam.; outermost bracts
connate into a cupuliform involucre 56 mm, marginally irregular. Flowers sessile. Calyx with hypanthium portion turbinate,
ca. 2 mm, glabrous; limb ca. 2 mm, deeply 5-lobed; lobes narrowly triangular, densely hirsute, ciliate. Corolla white, funnelform, glabrous outside; tube 23 mm, densely villous in throat
inside; lobes subelliptic to triangular, 11.2 mm, apex rostrate.
Drupes ellipsoid, oblong-ellipsoid, or narrowly ovoid, 68 mm,
color not noted; pyrenes longitudinally 4- or 5-ridged. Fl. Jul.

Forests along valley streams; 6001200 m. Hainan.

In the protologue the stipules were incorrectly described as apice
longe acuminatis. In fact, it is the individual lobes that are long acuminate, while the stipules are deeply bilobed.

8. Psychotria henryi H. Lveill, Repert. Spec. Nov. Regni

Veg. 13: 179. 1914.
dian nan jiu jie
Shrubs, 0.752 m tall; stems puberulent. Petiole 0.42 cm,
puberulent; leaf blade drying papery, reddish brown, grayish
brown, yellowish green, or greenish brown, often paler abaxially, narrowly elliptic or narrowly oblong-lanceolate, 414 1
4.5 cm, glabrous adaxially, puberulent to glabrous abaxially,
base acute to cuneate, margins flat, apex acute to acuminate;
secondary veins 610 pairs, free or forming a weak looping
submarginal vein, without domatia; stipules caducous, ovate to
triangular, interpetiolar, 1.54 mm, puberulent, 2-lobed for 1/5
1/2, lobes linear-lanceolate to subulate. Inflorescences terminal
or pseudoaxillary, subcapitate to congested-cymose, few to several flowered, puberulent to glabrous, sessile to shortly pedunculate; peduncle to 0.5 cm; branched portion subglobose to
corymbiform, 0.51.5 0.61.5 cm; bracts triangular, 11.5

Mountain forests. Hainan (Changjiang, Dongfang) [Vietnam (Poilane 21252 P!)].

The one developed flower seen resembles the long-styled form of
distylous Psychotria species (Lau 27455, MO!). Several Vietnam specimens at P clearly belong to P. laui, and this species is here reported from
that country as well.

10. Psychotria manillensis Bartling ex Candolle, Prodr. 4: 522.

1830.
liu qiu jiu jie mu
Shrubs to 2(6) m tall; stems glabrous. Petiole 0.53.6 cm,
glabrous; leaf blade drying thinly leathery, reddish brown to
reddish gray, elliptic-oblong, oblong-lanceolate, or obovateelliptic, 918.7[20.5] 38 cm, glabrous on both surfaces,
base acute to cuneate, margins flat to thinly revolute, apex acute
or shortly acuminate; secondary veins [6 or]712 pairs, free or
usually forming a looping submarginal vein, sometimes with
foveolate domatia; stipules caducous, triangular to ovate, interpetiolar, 36[10] mm, glabrous, acute to obtuse. Inflorescences
terminal or pseudoaxillary, cymose, glabrous [to puberulent],
pedunculate; peduncle [0.1]0.53.5 cm; branched portion cor-

RUBIACEAE

ymbiform to broadly pyramidal, 2.56 37 cm; bracts triangular to broadly triangular, 0.31.5[5.8] mm, those subtending
flowers 0.31.2 mm; pedicels to 2 mm. Flowers sessile to
pedicellate. Calyx glabrous; hypanthium portion obconic, 0.81
mm; limb 0.50.8 mm, lobed for ca. 1/2; lobes broadly triangular. Corolla white, tubular-funnelform, glabrous outside; tube
ca. 2[4] mm, densely villous in throat; lobes triangular, ca. 2
mm. Drupes red or purple, ellipsoid or ovoid-ellipsoid, 812[
15] 56 mm; pyrenes 3- or 4-ribbed. Fl. JunAug, fr. Aug.
Broad-leaved forests; near sea level [to 900 m in the Philippines].
Taiwan (Lan Yu) [Japan (Ryukyu Islands), Philippines].
Measurements included here in brackets were reported by Sohmer
and Davis (Sida, Bot. Misc. 27: 138142. 2007) for Philippine plants of
this species.

11. Psychotria morindoides Hutchinson in Sargent, Pl. Wilson.

3: 414. 1916.
ju guo jiu jie
Shrubs, 0.53 m tall; stems densely hirtellous to tomentulose-pilosulous or glabrescent. Petiole 1.26 cm, densely hirtellous to glabrescent; leaf blade drying papery, dark reddish
brown, gray, grayish brown, or greenish gray, oblanceolate,
obovate, elliptic-obovate, or elliptic-oblong, 830 311.5 cm,
glabrous adaxially, puberulent to densely hirtellous abaxially,
base acute to obtuse and sometimes oblique, margins flat, apex
acute to acuminate and sometimes curved; secondary veins 8
15 pairs, free or forming only a weak looping submarginal vein,
without domatia; stipules caducous, oblong-ovate, interpetiolar,
820 mm, puberulent to densely hirtellous, ciliolate, 2-lobed for
1/41/2, lobes narrowly triangular to linear. Inflorescences terminal sometimes becoming pseudoaxillary, capitate to subcapitate, densely hirtellous, many flowered, subsessile to pedunculate; peduncle to 6 cm; head or branched portion ovoid, ellipsoid, or pyramidal in outline, 26.5 1.54 cm; bracts linear,
ca. 5 mm. Flowers sessile. Calyx glabrous; hypanthium portion
obconic, ca. 1.25 mm; limb 2.53 mm, deeply lobed; lobes linear-lanceolate, entire to sparsely ciliate. Corolla white, funnelform to tubular-funnelform, outside glabrous; tube ca. 4 mm,
densely villous in throat; lobes lanceolate, 1.21.5 mm, apex
thickened. Drupes red, ellipsoid, 68 35 mm; pyrenes 3- or
5-ribbed. Fl. AprNov, fr. JulDec.
Forests along valley streams; 10002300 m. Yunnan [Laos, Thailand (Rock 1773, A!)].

12. Psychotria pilifera Hutchinson in Sargent, Pl. Wilson. 3:

415. 1916.
mao jiu jie
Shrubs, 0.751.75 m tall; stems densely villous to villosulous. Petiole 15 cm, densely villous; leaf blade drying papery
or membranous, dark reddish brown, elliptic, elliptic-obovate,
or elliptic-oblong, 823 311 cm, moderately to densely and
similarly strigose to hirsute on both surfaces, base acute or
obtuse to rounded, margins flat and ciliate, apex acute to shortly
acuminate with tip to 1 cm and sometimes curved; secondary
veins 515 pairs, not forming a submarginal vein, apparently
without domatia; stipules caducous to tardily deciduous, ovate-

299

triangular, interpetiolar, 1525 mm, densely hirsute to villosulous, 2-lobed for ca. 1/2, lobes narrowly triangular, acuminate
to caudate. Inflorescences terminal becoming pseudoaxillary,
congested-cymose to subcapitate, densely hirsute or villous, pedunculate; peduncle 36.5 cm; branched portion pyramidal to
subglobose, 23.5 25 cm; bracts linear-lanceolate, 410 mm,
ciliate, acuminate. Flowers subsessile. Calyx glabrescent to
sparsely villosulous; hypanthium portion obconic, ca. 1 mm;
limb 2.53.5 mm, lobed for ca. 2/3; lobes narrowly triangular to
linear-lanceolate. Corolla in bud funnelform, outside villosulous, to 4 mm. Drupes red, oblong-ellipsoid, 810 45 mm,
sparsely villosulous to glabrescent; pyrenes 3- or 4-ribbed. Fl.
Jul, fr. AugDec.
Forests in ravines; 13001700 m. Yunnan.
The protologue described the inflorescences as sometimes axillary, but this appears to be a description of the position separated here as
pseudoaxillary. W. C. Chen (in FRPS 71(2): 59. 1999) described the
petioles as becoming glabrescent with age and the secondary leaf veins
as impressed above, but these conditions have not been seen on any of
the specimens studied.

13. Psychotria prainii H. Lveill, Repert. Spec. Nov. Regni

Veg. 9: 324. 1911.
bo gu jiu jie
Cephalis siamica Craib; Psychotria siamica (Craib)
Hutchinson.
Shrubs, 0.52 m tall; stems densely hirtellous. Leaves
opposite but sometimes crowded at stem apices; petiole 0.22.2
cm, densely hirtellous; blade drying papery or thinly leathery,
dark reddish brown, grayish green, or brownish green, elliptic,
elliptic-oblong, lanceolate-oblong, obovate-oblong, or ovate, 3
15 1.36.5 cm, glabrous adaxially, densely hirtellous to hirsute throughout abaxially, base acute to obtuse, margins flat,
apex shortly obtuse and sometimes then shortly acuminate; secondary veins 611 pairs, free or forming a weak, looping, incomplete submarginal vein, without domatia; stipules caducous,
ovate, interpetiolar, 515 mm, densely hirtellous, 2-lobed for
1/31/2, lobes subulate. Inflorescences terminal or pseudoaxillary, capitate to densely congested-cymose, several flowered,
hirtellous, sessile to pedunculate; peduncle to 1 cm; head subglobose, 11.5 11.5 cm; bracts narrowly triangular, 37 mm.
Flowers sessile or subsessile. Calyx hirtellous; hypanthium portion obconic, ca. 1 mm; limb 1.53.5 mm, deeply lobed; lobes
narrowly lanceolate, spatulate, or narrowly elliptic, ciliolate.
Corolla white, funnelform, outside glabrous except pilosulous
on tips of lobes; tube ca. 3 mm, densely villous in throat; lobes
triangular-ovate, ca. 1.5 mm. Drupes red, ellipsoid or obovoid,
58 45 mm; pyrenes shallowly 4- or 5-ridged. Fl. MayAug,
fr. JulNov.
Rocky thickets, forests in ravines, mountain slopes; 10001700
m. Guangdong (Yangshan), W Guangxi, SW Guizhou, Yunnan [Laos,
Thailand, Vietnam].
This species is used medicinally.
The illustration of the corollas of this species in FRPS (71(2): 56,
t. 14, f. 59. 1999) does not agree with the specimens seen; in particular,
the tube illustrated is too short in relation to the lobes. No differences

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300

are evident between specimens of Psychotria siamica and of P. prainii,

so the former are here synonymized.

14. Psychotria serpens Linnaeus, Mant. Pl. 2: 204. 1771.

man jiu jie
Psychotria scandens Hooker & Arnott.
Climbing or creeping vines or lianas, to 6 m or more,
rather fleshy, juvenile stems appressed to substrate with adventitious roots, reproductive stems spreading at apex; stems glabrous. Petiole 110 mm, glabrous; leaf blade drying papery to
leathery, pale green, dark reddish green, reddish brown, or dark
brown, often paler below, ovate or obovate on juvenile stems
and elliptic, elliptic-oblong, lanceolate, oblanceolate, or obovate-oblong on reproductive stems, 0.79 0.53.8 cm, glabrous, base acute to obtuse, margins plane or sometimes thinly
revolute, apex acute, obtuse, or sharply acuminate; secondary
veins not visible or 310 pairs, not forming a submarginal vein,
without domatia; stipules caducous, triangular to ovate, interpetiolar or shortly united around stem, 23 mm, glabrous, acute
to rounded. Inflorescences terminal, cymose, glabrous to puberulent, few to many flowered, branched for 35 orders, pedunculate; peduncle 0.53 cm; branched portion corymbiform,
15 15.5(10) cm; bracts subtending secondary axes 12
mm and triangular or often leaflike and 36 mm, those subtending pedicels triangular, 0.21.5 mm; pedicels 0.51.5 mm.
Flowers pedicellate. Calyx glabrous to puberulent; hypanthium
portion obconic, 0.81 mm; limb flared, 0.51 mm, partially
lobed; lobes triangular, ca. 0.5 mm. Corolla white, funnelform,
outside glabrous to usually puberulent; tube 1.53 mm, in throat
densely villous; lobes ligulate-oblong, 1.52.5 mm. Drupes
white, subglobose or ellipsoid, 47 2.56 mm, with pedicels
to 5 mm; pyrenes shallowly 4- or 5-ribbed. Fl. AprJun, fr.
year-round.
Thickets or forests in ravines, mountains, hills, flat lands; below
1001400 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan, Zhejiang
[Cambodia, Japan, N Korea, Laos, Thailand, Vietnam].
This species is used medicinally.
W. C. Chen (in FRPS 71(2): 60. 1999) described the stems,
petioles (but not the leaf blades), inflorescence axes, and calyces as
spreading pubescent, but this has not been seen on any of the specimens
studied nor reported by other authors.
This species is commonly collected and morphologically quite
variable, including in drying color, leaf size and shape, inflorescence
size, and fruit size. The climbing habit and white fruit of this species are
unusual in the genus but similar to some other species of Asian
Psychotria (Sohmer & Davis, Sida, Bot. Misc. 27: 1247. 2007). The
name P. ixoroides Bartling ex Candolle has been considered a synonym
of P. serpens by some authors, notably Merrill, but Sohmer and Davis
(loc. cit.: 4145) considered P. ixoroides an accepted species in the
Philippines. However, they did not make any comparison there with P.
serpens. Psychotria serpens is also similar to P. sarmentosa Blume,
which was considered a distinct species by Sohmer (in Dassanayake,
Revis. Handb. Fl. Ceylon 6: 349352. 1987) again without any comparison to P. serpens.

15. Psychotria straminea Hutchinson in Sargent, Pl. Wilson.

3: 416. 1916.
huang mai jiu jie

Shrubs, 0.53 m tall; stems glabrous. Petiole 0.53.5 cm,

glabrous; leaf blade drying papery to membranous, yellowish
green, often pale below, elliptic, elliptic-oblong, oblanceolate,
or obovate, 5.529 0.810.5 cm, glabrous on both surfaces,
base cuneate to obtuse, margins flat, apex acuminate or acute;
secondary veins 510 pairs, thinly prominulous adaxially, free
or forming an incomplete looping submarginal vein, without
domatia; stipules persistent on apical nodes, triangular to ovate
or elliptic, shortly fused around stem, 2.56.7 mm (including
lobes), glabrous, 2-lobed, lobes linear, 0.51.2 mm, often tipped
with a caducous gland. Inflorescences terminal, few flowered,
cymose, glabrous, pedunculate; peduncle 12.5 cm; branched
portion corymbiform, 15 1.54 cm, branched to 13 orders;
bracts triangular, 0.21 mm; pedicels 1.54 mm. Flowers pedicellate in dichotomous cymules of 37. Calyx glabrous; hypanthium portion obconic, ca. 1 mm; limb ca. 1 mm, lobed shallowly or to 1/2; lobes triangular. Corolla white or pale green,
tubular-funnelform, glabrous outside; tube 1.52 mm, villous in
throat; lobes ovate-triangular, 1.52.5 mm. Drupes red or perhaps ultimately black, subglobose or ellipsoid, 713 49 mm,
with pedicels or stipes to 10 mm; pyrenes smooth or shallowly
3- or 4-ridged. Fl. JanJul, fr. JunJan of following year.
Forests in ravines, hill slopes; 1002700 m. Guangdong, Guangxi,
Hainan, Yunnan [Vietnam].
W. C. Chen (in FRPS 71(2): 52. 1999) described the fruit as black
at maturity, and the dried fruit seen have this color; but it remains to be
confirmed in living fruit by field observations. Some other Psychotria
species have fruit that are red at maturity but dry black, while some
species have fruit that turn red then ultimately black when ripe.

16. Psychotria symplocifolia Kurz, Forest Fl. Burma 2: 11.

1877.
shan fan ye jiu jie
Shrubs or small tees, 15 m tall; stems glabrous. Petiole
0.53.5 cm, glabrous; leaf blade drying papery to leathery, dark
reddish green, brownish green, or clear green, oblong-obovate,
elliptic-obovate, or elliptic-lanceolate, 612.7 26 cm, glabrous on both surfaces, base cuneate to acute, margins flat to
thinly revolute, apex shortly and abruptly acuminate; secondary
veins 612 pairs, free or forming a weak, incomplete, looping
submarginal vein, sometimes with foveolate domatia; stipules
caducous, ovate, interpetiolar or shortly fused around stem, 6
12 mm, glabrous outside (abaxially), inside (adaxially) densely
villous, acuminate, entire or 2-denticulate. Inflorescence terminal, cymose-paniculiform, puberulent to hirtellous with pubescence often in lines, often glabrescent with age, pedunculate;
peduncle 12.5 cm; branched portion pyramidal, 2.56 1.5
2.5 cm, with 2 or 3 pairs of developed secondary axes; bracts
triangular, 0.11 mm. Flowers sessile and subsessile in glomerulate groups. Calyx glabrous; hypanthium portion turbinate,
ca. 1 mm; limb ca. 0.5 mm, truncate to shallowly lobed; lobes
triangular. Corolla presumably white, salverform to funnelform,
glabrous outside; tube ca. 2 mm, in throat white villous; lobes
ovate-triangular, 1.52.2 mm. Drupes red, ellipsoid, 69 46
mm; pyrenes with 1 longitudinal ridge. Fl. FebApr, fr. Jun.
Forests on mountains; 12002300 m. Yunnan [India, Myanmar, N
Thailand].

RUBIACEAE

17. Psychotria tutcheri Dunn, J. Bot. 48: 324. 1910.

jia jiu jie
Shrubs, 0.54 m tall; stems puberulent to glabrous. Petiole
0.52 cm, puberulent to glabrous; leaf blade drying papery to
thinly leathery, pale or usually reddish brown to dark brown,
narrowly elliptic, narrowly lanceolate-oblong, narrowly lanceolate, or narrowly elliptic-oblong, 5.515 1.54 cm, glabrous
adaxially, glabrous to puberulent abaxially, base cuneate to
acute, margins flat, apex acuminate or caudate-acuminate; secondary veins 48 pairs, free or forming a weak, looping, submarginal vein, without or rarely with pilosulous domatia; stipules caducous, ovate-triangular, triangular, or lanceolate, interpetiolar, 28 mm, glabrous to puberulent, 2-lobed for 1/41/2,
lobes narrowly triangular to linear. Inflorescences terminal and
sometimes pseudoaxillary or rarely in uppermost leaf axils (He
Guosheng 6040, MO!), cymose, densely puberulent, sessile to
pedunculate; peduncle to 0.6 cm; branched portion corymbiform, 1.56 14 cm, with 1 or 2 pairs of developed secondary
axes; bracts lanceolate to triangular, 0.32 mm; pedicels to 0.5
mm. Flowers sessile to subsessile in congested small cymules.
Calyx puberulent to glabrescent; hypanthium portion obconic to
turbinate, 0.81 mm; limb 0.50.8 mm, lobed shallowly or
up to 1/2; lobes broadly triangular. Corolla white or greenish
white, tubular, glabrous outside; tube 23 mm, in throat white
villous; lobes oblong-lanceolate, 1.52 mm. Drupes subglobose, 57 46 mm, color unknown; pyrenes shallowly 3- or
4-ribbed to subsmooth. Fl. AprJul, fr. JunDec.

301

pairs, respectively; these measurement have not been seen on any specimens studied nor reported by any other authors.

18. Psychotria yunnanensis Hutchinson in Sargent, Pl. Wilson.

3: 414. 1916.
yun nan jiu jie
Psychotria kwangsiensis H. L. Li.

Thickets or forests in ravines or on hill slopes; 2001000 m. Fujian, Guangdong, Guangxi, Hainan, Yunnan [Vietnam].

Shrubs, 14 m tall; stems glabrous. Petiole 15.5 cm, glabrous; leaf blade drying papery to membranous, dark brown or
greenish brown, sometimes paler below, obovate-oblong, elliptic, ovate-oblong, or oblanceolate, 930.5 311 cm, glabrous
on both surfaces or puberulent to hirtellous along principal
veins abaxially, base acute to obtuse, margins flat, apex acuminate or acute; secondary veins 816 pairs, not forming a submarginal vein, without domatia; stipules caducous, ovate, interpetiolar, 615 mm, glabrous or hirtellous in basal portion, 2lobed for 1/41/3, lobes narrowly triangular, acuminate. Inflorescences terminal or pseudoaxillary, thyrsiform to paniculate,
many flowered, puberulent or hirtellous in lines often becoming
glabrescent; peduncle 16 cm; branched portion pyramidal to
broadly pyramidal, 310 2.56.5 cm, with 1 or 2 pairs of
well-developed secondary axes, with 311 glomerules or
cymules; bracts triangular to lanceolate, 110 mm, acuminate;
pedicels to 3 mm. Flowers subsessile to pedicellate in groups of
515. Calyx glabrous; hypanthium portion obconic, ca. 1 mm;
limb campanulate, 1.32.5 mm, lobed shallowly to deeply;
lobes deltoid to narrowly triangular or narrowly lanceolate. Corolla white, salverform, outside glabrous; tube ca. 5 mm, in
throat villous; lobes elliptic-oblong, ca. 3 mm. Drupes ellipsoid
or subglobose, 612 47 mm, color not noted; pyrenes 4- or
5-ridged. Fl. AprDec.

W. C. Chen (in FRPS 71(2): 54. 1999) described leaf sizes and
secondary vein numbers that are unusually large, to 22 6 cm and 13

Forests in ravines, hill slopes; 8002300 m. Guangxi (Napo),

Xizang (Mdog), Yunnan.

75. PSYDRAX Gaertner, Fruct. Sem. Pl. 1: 125. 1788.

jia yu gu mu shu
Chen Tao (); Charlotte M. Taylor, Henrik Lantz
Shrubs or small trees [or sometimes scandent], unarmed. Raphides absent. Leaves opposite [or sometimes ternate], sometimes
with domatia; stipules persistent, interpetiolar or shortly fused to petioles or united around stem, generally triangular to ovate.
Inflorescences axillary, cymose and several flowered [or rarely 1-flowered], sessile to pedunculate, bracteate or bracts reduced.
Flowers subsessile to pedicellate, bisexual, monomorphic. Calyx limb truncate or 4- or 5-dentate. Corolla white to yellow, tubular to
funnelform, inside variously pubescent; lobes 4 or 5, valvate in bud, markedly reflexed at anthesis. Stamens 4 or 5, inserted in corolla
throat, partially to fully exserted; filaments developed, reflexed at anthesis; anthers dorsifixed near base. Ovary 2-celled, ovules 1 in
each cell, pendulous from apical placentas; stigma exserted, ovoid to cylindrical, bifid, with style attachment recessed. Fruit
generally yellow, drupaceous, fleshy, subglobose to ellipsoid or sometimes dicoccous, with calyx limb persistent; pyrenes 2, 1-celled
with 1 seed, bony or cartilaginous; seeds medium-sized, ellipsoid, cylindrical, or plano-convex; testa membranous; endosperm
fleshy; radicle ascending.
About 100 species: tropical Africa and Asia; one species in China.
The species now treated in Psydrax were long included in Canthium, but they were separated out and Psydrax was revived as a genus by
Bridson (Kew Bull. 40: 687725. 1985), subsequently supported by Lantz and Bremer (Bot. J. Linn. Soc. 146: 272276. 2004). The type species was
published as Psydrax dicoccos, using for the epithet a Greek term that is presumably analogous to dicoccus or dicoccum (cf. Stearn, Botanical
Latin. 1983, Group II nouns and Group A adjectives, respectively). Bridson was the first author to accept Psydrax in almost two centuries, and she
clearly and explicitly gave feminine endings to all its species including the type, P. dicocca. Thus ICBN Art. 62.1 seems to apply here: the assignment
of gender to a genus is based on predominance of usage, not the original publication, and Psydrax is here treated as feminine and the epithet of our
species as adjectival following Bridson. Bridson also suggested that P. dicocca is restricted to Sri Lanka and S India and the Chinese and Malesian
plants that have long gone under that name are not conspecific; however, no differences are evident on the specimens studied, and further resolution of
this is outside the scope of the current work.

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302

1. Psydrax dicocca Gaertner, Fruct. Sem. Pl. 1: 125. 1788

[dicoccos].
jia yu gu mu
Shrubs to trees, to 15 m tall; branches compressed or
quadrangular becoming terete, glabrous. Petiole 615 mm, glabrous; leaf blade drying leathery, shiny on both surfaces, dark
green adaxially, and pale brown abaxially, ovate, elliptic, obovate, ovate-elliptic, or ovate-lanceolate, 410 1.54 cm, glabrous on both surfaces, base cuneate to obtuse, margins often
crisped, apex shortly to long acuminate, acute, or obtuse; secondary veins 35 pairs, in abaxial axils sometimes with small
foveolate domatia; stipules 35 mm, glabrous, acute to shortly
aristate. Inflorescences 23.5 cm, cymose, umbelliform, or fasciculate, puberulent; peduncle 38 mm; bracts reduced; pedicels 38 mm. Calyx glabrous; ovary portion obconic to cupuliform, 11.2 mm; limb ca. 0.5 mm, truncate or usually shallowly
5-dentate. Corolla greenish white or pale yellow, funnelform,
outside glabrous; tube ca. 3 mm, tomentose in throat; lobes
5, triangular to ligulate, 2.53 mm, acute. Stigma ca. 1 mm.
Drupes obovoid to ellipsoid or subglobose, often weakly dicoccous and/or somewhat flattened, 810 68 mm, glabrous;
pyrenes rugose. Fl. JanAug, fr. JunNov.
Sparse forests or thickets at low to middle elevations, broadleaved forests; 100600 m. Guangdong, Guangxi, Hainan, Xizang,
Yunnan [India, Indochina, Indonesia, Malaysia, Philippines, Sri Lanka,
Thailand; Australia].
The filaments are bent at ca. 180 at anthesis, so the anthers are
fully reflexed and held parallel to the corolla tube but upside-down. W.
C. Ko (in FRPS 71(2): 11. 1999, as Canthium dicoccum) described the
corolla lobes as slightly shorter than the tube, but on several specimens
studied these structures appear to be essentially equal in length. The
fruit color at maturity seems not to have been noted by any authors or
collectors.
The varieties recognized by W. C. Ko (loc. cit.: 1113) are treated
here for reference. In addition to the characters given in the key below,

W. C. Ko distinguished Canthium dicoccum var. dicoccum by its stigmas entire to bilobed and C. dicoccum var. obovatifolium by its stigmas
entire or often bifid or emarginate; however, the stigmas of all of these
plants are held together and appear entire when young, then spread and
become shallowly to markedly bifid at anthesis. Thus, the stigma distinctions may be developmental rather than population-level differences.

1a. Leaf blade ovate, elliptic, or ovatelanceolate; flowers in axillary

cymes; fruit obovoid to ellipsoid .............. 1a. var. dicocca
1b. Leaf blade obovate or ovate-elliptic;
flowers 46 in axillary fascicles;
fruit subglobose or ellipsoid ............... 1b. var. obovatifolia
1a. Psydrax dicocca var. dicocca
() jia yu gu mu (yuan bian zhong)
Canthium dicoccum (Gaertner) Merrill; C. didymum C. F.
Gaertner; Plectronica dicocca (Gaertner) Merrill; P. didyma (C.
F. Gaertner) Merrill.
Leaf blade ovate, elliptic, or ovate-lanceolate. Inflorescences cymose. Stigma entire. Fruit obovoid or obovoid-ellipsoid. Fl. JanAug, fr. JunNov.
Sparse forests or thickets at low to middle elevations; 100600 m.
Guangdong, Guangxi, Hainan, Xizang (Mdog), Yunnan [India, Indochina, Indonesia, Malaysia, Philippines, Sri Lanka; Australia].

1b. Psydrax dicocca var. obovatifolia (G. A. Fu) Lantz, comb.

nov.
dao luan ye jia yu gu mu
Basionym: Canthium dicoccum var. obovatifolium G. A.
Fu, Bull. Bot. Res., Harbin 5(1): 183. 1985.
Leaf blade obovate or ovate-elliptic. Inflorescences fasciculate, 46-flowered. Stigma entire or often bifid or emarginate.
Fruit subglobose or ellipsoid. Fl. MayJun.
Broad-leaved forests. Hainan (Tunchang).

76. RICHARDIA Linnaeus, Sp. Pl. 1: 330. 1753.

mo mu xu shu
Chen Tao (); Charlotte M. Taylor
Ricardia Adanson; Richardsonia Kunth.
Herbs, annual or perennial, unarmed. Raphides present. Leaves opposite, without domatia; stipules persistent, interpetiolar and
fused to petioles or leaf bases, truncate to rounded, setose. Inflorescences terminal, capitate, several to many flowered, pedunculate
and enclosed by paired leaflike bracts (or sessile with involucral leaves in other morphological interpretations), bracteate. Flowers
sessile, bisexual, monomorphic. Calyx with ovary portion turbinate to globose, limb deeply 48-lobed. Corolla white or pink, funnelform, inside glabrous or pubescent at throat; lobes 46, valvate in bud. Stamens 3, 4, or 6, inserted in corolla throat, exserted; filaments developed; anthers dorsifixed near middle. Ovary 3- or 4-celled, ovules 1 in each cell, axile and attached at middle of septum;
stigmas 3 or 4, linear or spatulate, exserted. Fruit schizocarpous, subglobose to obovoid or tricoccous, dry, bony, with calyx limb
deciduous; mericarps 3 or 4, indehiscent, 1-celled with 1 seed, ellipsoid to angled, usually papillose to muricate on dorsal surface
(i.e., abaxially) and with 1 or more grooves and sometimes papillose to muricate on ventral surface (i.e., adaxially); seeds mediumsized, ellipsoid to plano-convex; endosperm corneous; cotyledon leaflike; radicle cylindrical, hypogeous.
Fifteen species: widespread in the Antilles and North and South America, three species naturalized in the Old World tropics; two species (both
introduced) in China.
As noted by Chaw and Peng (J. Taiwan Mus. 40: 7183. 1987), Asian collections of Richardia have long been confused in herb. with various
other weedy Rubiaceae. Richardia was studied in detail by Lewis and Oliver (Brittonia 26: 271301. 1974). The synonymous name Richardsonia has

RUBIACEAE

303

frequently been used for this genus, including in older references about invasive weeds. H. S. Lo (in FRPS 71(2): 203. 1999) described the flowers as
sometimes polygamo-dioecious, but the origin of this description is unknown. Lewis and Oliver did not report this condition, although they did
mention that the plants frequently have both chasmogamous and cleistogamous flowers. H. S. Lo also described the anthers as dorsifixed near the
base, but other authors have all considered them to be dorsifixed near the middle, which agrees with specimens studied.
Richardia stellaris (Chamisso & Schlechtendal) Steudel is naturalized in Australia and perhaps may be expected in China; it can be recognized
by its narrowly triangular to narrowly elliptic, sharply acute leaves.

1a. Mature mericarps somewhat dorsiventrally flattened, with 2 broad parallel depressions along length of inner
(i.e., adaxial) face ...................................................................................................................................................... 1. R. brasiliensis
1b. Mature mericarps triangular to somewhat rounded, with 1 narrow grove along length of inner (i.e., adaxial) face ...... 2. R. scabra
1. Richardia brasiliensis Gomes, Mem. Ipecacuanha Brasil,
31. 1801.

Peng, J. Taiwan Mus. 40: 7183. 1987). It was not cited at all by H. Y.
Liu and T. Y. A. Yang (Fl. Taiwan, ed. 2, 4: 245340. 1998).

2. Richardia scabra Linnaeus, Sp. Pl. 1: 330. 1753.

ba xi mo mu xu
Herbs, annual, decumbent or suberect, to 80 cm or longer;
stems flattened to subterete, hispidulous or scaberulous and
hirsute. Petiole 510 mm, hispidulous to pilosulous; leaf blade
drying membranous to thickly papery, ovate, elliptic, or lanceolate, 15 0.53.5 cm, both surfaces scaberulous to glabrescent, base acute to cuneate, apex acute to obtuse; stipule sheaths
13 mm, pilose to pilosulous, with 311 setae 24 mm. Inflorescences ca. 1 cm in diam. (not including leaflike bracts or
subtending leaves). Calyx with ovary portion obovoid, 11.5
mm, densely papillose or hispidulous to smooth; lobes 6, lanceolate or narrowly lanceolate, 1.53.5 mm, glabrescent, margins ciliate, apex acute. Corolla white, glabrous inside and outside; tube 38 mm; lobes 6, 13 mm. Fruit with mericarps 3,
ellipsoid to obovoid, laterally somewhat dorsiventrally flattened, 23 mm, dorsally papillose to subsmooth, ventrally with
2 broad parallel grooves along length of face. Fl. and fr. Feb
Sep.
Wastelands; ca. 300 m. Naturalized in Guangdong and Taiwan
[native to South America; adventive and naturalized occasionally
throughout Old World tropics].
This species was reported as naturalized in Taiwan by Ou (Bull.
Exp. Forest Natl. Chung Hsing Univ. 8: 1130. 1987, article not seen,
cited by Wu et al., Taiwania 49: 1631. 2004) and later by Wu et al.
(loc. cit.) but not by other contemporaneous authors (e.g., Chaw &

mo mu xu
Richardia pilosa Ruiz & Pavon; Richardsonia scabra
(Linnaeus) A. Saint-Hilaire.
Herbs, annual, decumbent or suberect, to 80 cm or longer;
stems flattened to subterete, hirsute. Petiole 510 mm, hirsute to
glabrescent; leaf blade drying membranous to thickly papery,
ovate, elliptic, or lanceolate, 15 0.53.5 cm, both surfaces
scabrous to glabrescent, base acute to cuneate, apex acute to
obtuse; stipule sheaths 14 mm, pilose to pilosulous, with 315
setae 25 mm. Inflorescences ca. 1 cm in diam. (not including
leaflike bracts or subtending leaves). Calyx with ovary portion
obovoid, 11.5 mm, papillose to hispidulous; lobes 6, lanceolate or narrowly lanceolate, 1.53.5 mm, glabrescent, margins
ciliate, apex acute. Corolla white, glabrous inside and outside;
tube 28 mm; lobes 6, triangular, 13 mm. Fruit with mericarps
3, ellipsoid to obovoid, in cross-section triangular to somewhat
rounded, 23.5 mm, dorsally densely papillose to hispidulous,
ventrally with 1 narrow groove along length of face. Fl. and fr.
FebNov.
Wastelands; sea level to 200 m. Naturalized in Guangdong, Hainan, and Taiwan [native to the Antilles and North and South America;
adventive and naturalized occasionally throughout Old World tropics].
H. S. Lo (in FRPS 71(2): 203. 1999) noted that this species was
introduced to China in the 1980s.

77. RONDELETIA Linnaeus, Sp. Pl. 1: 172. 1753.

lang de mu shu
Chen Tao (); Charlotte M. Taylor
Shrubs or trees, unarmed. Raphides absent. Leaves opposite or rarely verticillate, sometimes with domatia; stipules persistent or
caducous, interpetiolar, generally triangular. Inflorescence terminal or infrequently axillary, congested-cymose to paniculate or corymbiform, several to many flowered, pedunculate, bracteate. Flowers sessile to pedicellate, bisexual, distylous. Calyx limb 4- or 5lobed, with lobes often unequal. Corolla white, yellow, orange, or red, funnelform or salverform with tube often slender and prolonged, inside glabrous or villous, with thickened annular ring at throat; lobes 4 or 5, imbricate in bud, with margins frequently
crisped. Stamens 4 or 5, inserted in corolla throat, included or exserted; filaments short or reduced; anthers dorsifixed. Ovary 2celled, ovules numerous in each cell on axile placentas; stigmas capitate or 2-lobed, included or exserted. Fruit capsular, globose to
depressed globose or dicoccous, loculicidally dehiscent into 2 valves with these valves often then splitting, woody to papery, with
calyx limb persistent; seeds numerous, small, fusiform or discoid, flattened, winged; endosperm fleshy; embryo small, clavate.
About 20 species: tropical America, one species widely cultivated in tropical regions; one species (introduced) in China.
This genus was formerly circumscribed broadly, to include as many as 140 species, but neotropical taxonomists now separate the species
formerly treated in Rondeletia s.l. into a number of smaller genera, notably Arachnothryx Planchon and Rogiera Planchon. The cultivated species
treated here belongs to Rondeletia s.s.

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304

1. Rondeletia odorata Jacquin, Enum. Syst. Pl. 16. 1760.

lang de mu
Shrubs, to 2 m tall; branches flattened to subterete, hirtellous or villosulous to glabrescent. Leaves opposite, decussate; petiole 12 mm, hirtellous to glabrescent; blade drying
stiffly leathery, ovate, elliptic, or elliptic-oblong, 25 13.5
cm, adaxially scabrous and often rugulose or bullate, abaxially
glabrescent to pilosulous or hirtellous at least on principal
veins, base obtuse to subcordate, margins thinly revolute, apex
broadly obtuse to acute; secondary veins 36 pairs, sometimes
with pilosulous to pilose domatia; stipules persistent, triangular,
45 mm, strigillose to pilosulous, acute. Inflorescences terminal, cymose, several to many flowered, 23 34.5 cm, hirtellous to hirsute; peduncle 0.71.5 cm; bracts narrowly triangular

to ovate or elliptic, 215 mm, acute; pedicels 1.55 mm. Calyx

with ovary portion subglobose, 1.52 mm, densely strigillose to
pilosulous; limb lobed essentially to base; lobes 5, narrowly
triangular to linear, 45 mm, sparsely strigillose to puberulent.
Corolla bright red with yellow throat, salverform, pilosulous to
strigillose outside; tube 1220 mm; lobes suborbicular, ca. 3.5
4 mm, crisped, obtuse to rounded. Capsules globose to depressed globose, 34 mm in diam., densely hirtellous or villosulous; seeds ca. 0.8 0.5 mm. Fl. JulSep.
Cultivated in Fujian, Guangzhou, and Hong Kong [native to
Cuba; cultivated sporadically in tropical regions worldwide].
The cultivated plants apparently do not produce fruit; the description of the fruit here is taken from plants in the native range of this
species. The few Chinese species seen are all long styled.

78. ROTHMANNIA Thunberg, Kongl. Vetensk. Acad. Handl. 37: 65. 1776.
ye zhi zi shu
Chen Tao (); Charlotte M. Taylor
Shrubs or large trees, unarmed. Raphides absent. Leaves opposite and generally isophyllous or frequently apparently verticillate
in whorls of 3 due to marked anisophylly grouping 2 leaves at a node plus an apparently single leaf on a short to reduced axillary
branch, sometimes with domatia; stipules persistent, interpetiolar, triangular. Inflorescences terminal or pseudoaxillary, congestedcymose and several flowered or reduced to 1 flower, sessile or pedunculate, bracteate or bracts reduced. Flowers sessile to pedicellate
or pedunculate, bisexual, monomorphic. Calyx limb velvety pubescent inside, truncate to 5-lobed. Corolla white to pale green with
red or purple to brown spots, narrowly to broadly funnelform or campanulate, often fleshy to leathery, glabrous inside; lobes 5(7),
convolute in bud to left or right depending on species. Stamens 5(7), inserted in corolla tube, included or partly exserted; filaments
short or reduced; anthers dorsifixed. Ovary partially to perhaps completely [1 or]2-celled, ovules numerous on 2 to several large
parietal or perhaps sometimes axile placentas; stigma clavate, with receptive surface confined to shortly bilobed apex, exserted or
included. Fruit baccate, thickly fleshy to leathery, globose to ellipsoid and sometimes relatively large, smooth or ridged, yellow to
brown, with calyx limb persistent; seeds numerous, large, angled to sublenticular, embedded in pulp.
At least 30 species: tropical Africa, Asia, and Madagascar; one species in China.
Rothmannia does not seem at all well known in Asia, as to species or generic limits. The distinctive growth form of Rothmannia, with some or
several nodes bearing apparently unequal and ternate leaves, is due to the development of one very short axillary branchlet that bears one leaf (the
other being reduced to absent). The flowers are often if not usually nocturnal. Rothmannia has been described by some authors as having a 1-locular
ovary with parietal placentas, at least in Africa (Bridson & Verdcourt, Fl. Trop. E. Africa, Rub. (Pt. 2), 511512. 1988), but the Asian species have 2celled ovaries with apparently axile placentas (Puff et al., Rubiaceae of Thailand, 60. 2005).
The African species Rothmannia longiflora Salisbury is occasionally cultivated in tropical Asia, including Hong Kong (S. Y. Hu & K. H. Yung
403, MO!). This species has elliptic leaves that are 611 35 cm and relatively large showy flowers, with the tubular calyx limb 510 mm and
subtruncate or with small lobes 14 mm, and a slenderly funnelform, relatively large corolla with tubes 1214 cm and lobes 14 cm.

1. Rothmannia daweishanensis Y. M. Shui & W. H. Chen,

Novon 13: 322. 2003.
da wei shan ye zhi zi
Trees, 1015 m tall; bark dark gray; branches glabrous,
lenticellate. Petiole ca. 3 mm, glabrous; leaf blade drying stiffly
papery, greenish brown, ovate to elliptic, 1014 45 cm, both
surfaces glabrous and shiny, base cuneate to acute and sometimes slightly decurrent, apex acuminate with tip then blunt;
secondary veins 5 or 6 pairs; stipules broadly triangular-ovate,
1.52 23 mm, acuminate. Inflorescence terminal or pseudoaxillary, 1-flowered, glabrous; peduncle articulate, basal portion
ca. 0.5 cm, apical portion (pedicel) 2025 mm; bracteoles triangular, ca. 1.5 mm, sericeous adaxially, glabrous abaxially. Calyx sparsely sericeous outside; ovary portion ellipsoid, ca. 3
mm; limb with basal tubular portion 22.5 mm, inside densely

villous and with short colleters; lobes linear-oblong to spatulate,

1416 0.70.9 mm, 1-veined, obtuse. Corolla campanulate,
glabrous throughout; tube white outside, inside striate and/or
speckled with purple, 5055 mm, at base with slender portion
67 ca. 5 mm, abruptly dilated above this, ca. 30 mm in diam.
at middle; lobes white outside, purple-speckled inside, overlapping to left, broadly ovate, 1820 2223 mm, obtuse. Ovary
with placentas parietal, apparently confluent; style ca. 50 mm;
stigmatic portion ca. 20 1 mm. Berry ellipsoid becoming subglobose with base swollen-stipitate, 45 cm in diam., color inside and out unknown, smooth; seeds ca. 12 8 4 mm. Fl.
OctDec, fr. DecMar.
Rocky crevices in limestone rain forests; 300600 m. Yunnan
(Maguan) [Vietnam].
Shui and Chen described the flowers of this species as being

RUBIACEAE

(paraphrased here) terminal on a reduced branch that is produced in an

axillary position. The inflorescence in other species of Rothmannia is a
congested or reduced cyme generally borne on a short peduncle produced in a terminal or pseudoaxillary position; thus their reduced

305

branch is here treated as being the peduncle (i.e., the structure directly
bearing the solitary flower. In general, these structures have been interpreted variously by different authors. Zhang et al. (Acta Phytotax. Sin.
45: 92. 2007) reported this species also from Vietnam.

79. RUBIA Linnaeus, Sp. Pl. 1: 109. 1753.

qian cao shu
Chen Tao (); Friedrich Ehrendorfer
Shrubs, subshrubs, or perennial herbs, not rarely clambering or climbing vines or rarely lianas, unarmed; stems often prickly
and/or longitudinally ribbed or winged. Raphides present. Leaves opposite and with interpetiolar, triangular or ovate, persistent to
caducous (Rubia siamensis) or reduced (R. tibetica) stipules or with leaflike stipules in whorls of 4, 6, to many in middle stem
regions; domatia none; main veins single or 35(or more) and then palmate, secondary veins lateral. Inflorescences thyrsoid, with
terminal and/or axillary cymes, usually paniculiform and often expanding from new axes developing with age; individual cymes few
to many flowered, pedunculate, bracteate. Flowers pedicellate or sessile, rather small, usually bisexual and monomorphic, rarely
polygamo-dioecious (R. cordifolia). Ovary inferior (hypanthium), ellipsoid, subglobose, 2-celled, ovules 1 in each cell, erect, axile.
Calyx limb reduced and obsolete. Corolla white to cream, yellow, greenish or red to purplish, often turning black when dried, mostly
rotate, but rarely also campanulate to funnel-shaped, inside glabrous or infrequently papillose; lobes predominantly 5 (rarely also less
or more), valvate in bud, often long acuminate. Stamens usually 5, inserted at corolla base (or tube), exserted; filaments developed to
reduced; anthers dorsifixed. Stigmas 2-lobed, included or exserted. Fruit developing into 2 separate or (by reduction) into only 1
subglobose, baccate, berrylike mericarp with fleshy meso- and endocarp, dark red, purple, black, or infrequently orange (R. cordifolia), glabrous or somewhat hairy; seeds (pyrenes) 2, ellipsoid, subglobose, or plano-convex, with membranous testa; endosperm
corneous; embryo subincurved; cotyledons leaflike; radicle prolonged, basiscopic.
About 80 species: extending from tropical and temperate Asia to Japan and Indonesia, through the Himalaya to SW Asia, E to S Africa, through
the Mediterranean to W Europe, Macaronesia, and the Azores; locally introduced and persisting from cultivation in Mexico, Chile, and elsewhere; 38
species (20 endemic) in China.
As already mentioned in the present volume under Galium, Rubia is the type genus of the family, the tribe Rubieae, and the subtribe Rubiinae.
As an Old World clade, Rubia is related to the Mesoamerican genus Didymaea and occupies a relative basal position within Rubiinae: its 5-lobed
corollas, fleshy fruit, and always perennial growth form apparently are plesiomorphic features. This and its clear separation from the somewhat more
apomorphic Sherardia-Asperula-Galium group is well documented by DNA data (Natali et al., Opera Bot. Belg. 7: 193203. 1996; Soza & Olmstead,
Taxon 59: 755771. 2010). Rubia is keyed out from among the other Chinese taxa of Rubieae under Galium on p. 107. Its best differential characters
are the dominantly 5-merous flowers combined with baccate, berrylike mericarps. The latter also occur independently among New World taxa of
Galium (and Relbunium).
Among the Rubieae tribe Rubia (after Galium and Asperula in their present circumscription) is the third largest and obviously monophyletic
genus. Nearly half of its recognized species occur in China. Because of excessive variability, the occurrence of hybridization and polyploidy as well as
the lack of detailed studies (particularly on material in the major herbaria of China and elsewhere), our knowledge of Rubia is limited and the present
treatment of the genus still quite provisional.
More recent taxonomic surveys of Rubia are available for the former Soviet Union (Pojarkova, Fl. URSS 23: 382417. 1958), India (Deb &
Malick, Bull. Bot. Surv. India 10(1): 116. 1968), Iran (Ehrendorfer et al., Fl. Iranica 176: 4872. 2005), Bhutan (Long, Edinburgh J. Bot. 53: 108
110. 1996; Long, Fl. Bhutan 2(2): 823825. 1999), and Taiwan (T. Y. A. Yang, Fl. Taiwan, ed. 2, 4: 321324. 1998). Particularly in the first of these
contributions, but also in the following two, the infrageneric taxonomy of Rubia is briefly considered. The majority of the Chinese species are characterized by 35(11) palmate veins in their relatively broad leaves. These taxa correspond to R. sect. Oligoneura Pojarkova (= R. [unranked] Cordifoliae Candolle; R. sect. Cordifoliae (Candolle) K. Schumann ex Deb & Malick). Within this section Pojarkova (loc. cit.) has recognized two series:
one predominantly climbing vines with long leaf petioles as R. ser. Cordifoliae (Candolle) Pojarkova (with 16 species in China; see under R. cordifolia), the other mostly erect perennial herbs with very short leaf petioles as R. ser. Chinenses (with three species in China; see under R. chinensis).
The latter is close to the informal R. mandersii group, where the leaves are sessile (three species in China; see under R. mandersii). Finally, there are
two species groups with vines. One is the R. sikkimensis group with sessile leaves and leaflike stipules (two Chinese species; see under R. tenuis), in
the other, the R. siamensis group, typical (not leaflike) interpetiolar stipules appear between the opposite leaves (four species in China; see under R.
siamensis).
The remaining ten Chinese species have leaves with only 1(3) main vein(s) and predominantly pinnate vein branching. In the present treatment
this refers to the species Rubia chitralensis, R. deserticola, R. dolichophylla, R. rezniczenkoana, R. schugnanica, R. tibetica, and R. tinctorum, here
provisionally accommodated in R. sect. Rubia s.l. (including R. sect. Meganthera Pojarkova (= R. sect. Rubia s.s.), R. sect. Campylanthera Pojarkova,
and R. sect. Chonanthe Pojarkova). The placement of the E Asiatic species R. haematantha (the R. haematantha group; see there) as well as R. pseudogalium and R. truppeliana (the R. truppeliana group; see there) is doubtful; they may belong to R. sect. Oligoneura in spite of their very narrow
leaves and only 1 main vein. The more detailed infrageneric subdivisions of Rubia by Pojarkova listed above have been based on size and form of
anthers and other flower details and are in need of more general and detailed study.
Relevant characters for the separation of Rubia taxa on the species level are growth habit, indumentum, number, shape and consistency of leaves
and stipules, presence or absence of petioles, inflorescence structures, color and morphology of corollas from rotate to funnel-shaped, fruit color, etc.
Particularly, leaf characters often vary excessively under different environmental and developmental conditions (e.g., in more widespread taxa as
Rubia cordifolia and its allies; see further comments there). These facts are difficult to evaluate, at least on dried specimens.

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RUBIACEAE

The ground-up rhizomes and roots of Rubia tinctorum, the type species of the genus, have long been the source of important red textile dyes
(madder red, alizarin, rose madder, alizarin crimson). This use was of course much more important before the invention of aniline dyes (e.g., madder
colored the red coats of the 18th-century British army). Nevertheless, R. tinctorum is still widely cultivated at a local scale and used, in particular, to
color wool for handmade oriental rugs in C and SW Asia (Murphy, Root of Wild Madder, 1297. 2005) but also in fine art painting. The worldwide
occurrence, cultivation, chemistry, and cultural role of this species was discussed in detail by Chenciner (Madder Red. 2000). The stems of R. manjith
are also used to produce a red dye (fide Long, loc. cit. 1999).
The key here generally follows that of H. S. Lo (in FRPS 71(2): 287290. 1999), with the measurements updated from the descriptions and
species added as appropriate.

1a. Leaves with only 1 evident midvein; lateral veins pinnate, when palmate weak and obscure.
2a. Leaves linear to narrowly lanceolate or oblong, mostly 3.530 as long as wide.
3a. Leaves with well-developed petioles; plants of forests.
4a. Leaves in whorls of up to 68, with petioles 635 mm; peduncles 1040 mm; corolla lobes ca. 2 mm;
Shandong ....................................................................................................................................................... 36. R. truppeliana
4b. Leaves in whorls of never more than 4, with petioles 36 mm; peduncles 36 mm; corolla lobes
1.21.5 mm; Yunnan .................................................................................................................................. 24. R. pseudogalium
3b. Leaves without distinct petioles, sessile to subsessile; plants of open habitats.
5a. Corolla dark red or perhaps sometimes white; leaves 68 per whorl, narrowly linear, less than
1 mm wide, midvein without evident lateral veins ..................................................................................... 12. R. haematantha
5b. Corolla yellow or white; leaves 46 per whorl, linear-lanceolate to lanceolate-oblong, wider than
1 mm.
6a. Leaves 514 mm wide, midvein with pinnate lateral veins; stems with aculeolate angles;
inflorescence cymes terminal and distributed along lower stem nodes; corolla lobes ca. 2 mm ............ 8. R. dolichophylla
6b. Leaves 2050 mm wide, lateral veins not evident; stems smooth; inflorescence cymes clustered
near stem apices; corolla lobes 2.32.7 mm ............................................................................................ 28. R. schugnanica
2b. Leaves broadly (ob)lanceolate, lanceolate-oblong, ligulate, elliptic, elliptic-oblong, lanceolate-ovate,
ovate, or broadly ovate, mostly 13.5 as long as wide.
7a. Corolla lobes with apex aristate, arista 0.50.8 mm; leaves 46 in a whorl, lanceolate or elliptic-oblong,
drying firmly leathery .......................................................................................................................................... 7. R. deserticola
7b. Corolla lobes obtuse, acute, or acuminate to mucronate, with arista up to 0.4 mm; leaves lanceolate to
broadly ovate, drying papery to leathery.
8a. Larger leaves longer and wider than 3 1.5 cm, dried papery to subleathery.
9a. Leaves dried papery; corolla limb 67.5 mm in diam.; anthers ca. 0.4 mm ............................................... 4. R. chitralensis
9b. Leaves dried papery to subleathery; corolla limb 34.5 mm in diam.; anthers ca. 0.6 mm ........................ 34. R. tinctorum
8b. Leaves 0.53 0.21.5 cm, dried leathery.
10a. Leaves and leaflike stipules similar, mostly up to 6 per whorl; lower nodes of older stems not
sheathed with old leaf bases; corolla yellow, lobes often 4, obtuse with short incurved cusp ........ 26. R. rezniczenkoana
10b. Leaves 2 with 2 smaller leaflike interpetiolar stipules in whorls of 4; lower nodes of older stems
shortly sheathed with membranous bases of old leaves; corolla whitish, lobes usually 5, acuminate ......... 33. R. tibetica
1b. Leaves with 311 evident principal and palmate veins (including midrib), arising from at or near base.
11a. Leaves opposite, with evident interpetiolar stipules; plants of moist forests.
12a. Leaves tuberculate-hispidulous; stipules leaflike but smaller than true leaves ............................................... 6. R. crassipes
12b. Leaves glabrous, scabrous, or hairy; stipules ovate to triangular, very different from true leaves.
13a. Leaf margins entire and smooth, petioles 0.32.5 cm; inflorescence axes slender and filiform .................. 11. R. filiformis
13b. Leaf margins aculeolate, petioles (1)24(8) cm; inflorescence axes stout.
14a. Ovary and fruit densely hairy ................................................................................................................... 10. R. falciformis
14b. Ovary and fruit glabrous.
15a. Stipules large, ovate, acuminate, (5)1235(60) (3)825(40) mm; dried leaves light green
and ferruginous, particularly below and on main veins ........................................................................... 15. R. magna
15b. Stipules small, triangular, 35(7) 23 mm; dried leaves dark green ................................................ 30. R. siamensis
11b. Leaves and leaflike stipules similar, in whorls of 412; plants of various habitats.
16a. Erect herbs, if climbing vines then leaves (sub)sessile; leaves and leaflike stipules 4 or sometimes
6 per whorl.
17a. Leaves markedly cordate at base.
18a. Older stems broadly 4-winged ............................................................................................................. 25. R. pterygocaulis
18b. Older stems quadrangular to narrowly 4-angled.
19a. Stem angles retrorsely aculeate; mountains of Taiwan ................................................................................... 2. R. argyi
19b. Stem angles smooth; Sichuan ................................................................................................................. 13. R. latipetala
17b. Leaves cuneate, obtuse, truncate, rounded, or shallowly cordulate at base.

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307

20a. Leaves sessile or subsessile (if flowers 4-merous then see Galium).
21a. Principal leaf veins 711; stems and leaves strongly hairy; fused basal part of corolla only
0.20.3 mm, lobes 11.2 mm ............................................................................................................... 23. R. polyphlebia
21b. Principal leaves veins 3 or 5; stems and leaves glabrescent to glabrous, scabrous; fused basal
part of corolla 0.50.6 mm.
22a. Slender vines; leaves drying papery; flowers ca. 3 mm in diam. ............................................................. 32. R. tenuis
22b. Erect to spreading herbs; leaves drying mostly leathery; flowers 35 mm in diam.
23a. Stems 8-ribbed; leaves lanceolate to lanceolate-elliptic, 47 cm ................................................... 9. R. edgeworthii
23b. Stems 4-angled; leaves broadly elliptic, linear-lanceolate, ovate, obovate, or elliptic-oblong,
15 cm.
24a. Stems and leaves glabrous to scabrous, latter drying papery; principal main veins 5; corolla
ca. 5 mm in diam. .......................................................................................................................... 16. R. mandersii
24b. Stems and leaves often hairy or scabrous, latter drying papery; principal veins 35;
corolla ca. 3 mm in diam. .......................................................................................................... 38. R. yunnanensis
20b. Leaves petiolate, petioles 0.39 cm.
25a. Stems clearly retrorsely aculeolate.
26a. Leaves suborbicular to ovate, length/breadth index 11.8, dried papery .................................................... 2. R. argyi
26b. Leaves lanceolate to ovate, length/breadth index 23, dried papery to subleathery .......................... 22. R. podantha
25b. Stems smooth to sparsely scaberulous.
27a. Leaves broadly elliptic, principal veins 57, dried thinly papery; fused part of corolla
0.20.6 mm ............................................................................................................................................. 3. R. chinensis
27b. Leaves broadly lanceolate to broadly ovate, principal veins 35, dried papery to subleathery;
fused part of corolla 0.82 mm.
28a. Leaves broadly ovate, length/breadth index 1.21.5, base cordulate to cordate, dried papery,
principal veins 35; fused part of corolla 1.52 mm ....................................................................... 13. R. latipetala
28b. Leaves broadly lanceolate to ovate, length/breadth index 1.52.5, base obtuse, rounded,
or cordulate, dried thickly papery to subleathery, principal veins often impressed; fused
part of corolla 0.81 mm .......................................................................................................... 29. R. schumanniana
16b. Vines or lianas, climbing to sprawling; leaves petiolate, petioles 0.112 cm.
29a. Fruit, stems, and leaves hirsute, strigillose, hirtellous, or villosulous ....................................................... 35. R. trichocarpa
29b. Fruit glabrous, stems and leaves glabrous or with diverse indumentum.
30a. Leaves 412 per whorl, at least middle stem nodes with 6 or more leaves.
31a. Leaves oblanceolate, base cuneate to acute; petioles 0.63.5 cm ....................................................... 36. R. truppeliana
31b. Leaves ovate to suborbicular or lanceolate to oblong-lanceolate, base truncate, rounded,
cordulate, or cordate; petioles 111 cm.
32a. Leaves ovate to suborbicular, largest mostly longer than 4 cm, length/breadth index 1.21.5,
base cordulate or cordate; petioles 211 cm ......................................................................................... 31. R. sylvatica
32b. Leaves lanceolate to oblong-lanceolate, largest mostly shorter than 4 cm, length/breadth index
2.54, base truncate, rounded, or cordulate to cordate; petioles 19 cm .............................................. 5. R. cordifolia
30b. Leaves 4, very rarely more per whorl.
33a. Stems, leaves abaxially, and/or outside of corolla moderately to densely hirtellous or hispidulous
with trichomes regularly hooked at apex; leaves small, 0.83.5 0.31.5 cm ................................... 19. R. oncotricha
33b. Stems, leaves, and corollas outside glabrous, or with diverse indumentum, but never with
regularly hooked hairs; leaves larger, 0.713 0.36.5 cm.
34a. Leaves drying thickly leathery, oblong-ovate to elliptic, apex obtuse ................................................... 6. R. crassipes
34b. Leaves drying papery to leathery, ovate, oblong-lanceolate, ovate-lanceolate, oblong-ovate,
cordiform, suborbicular, or lanceolate, apex subacute, acuminate, caudate, or obtuse and cuspidate.
35a. Dried plants flushed with red, particularly on lower leaf side; corolla rotate, purplish red, red, or
orange, with spreading lobes of 1.21.5 mm; mature fruit dark red .................................................. 17. R. manjith
35b. Dried plants green, gray, or yellowish (if rarely flushed with red then corollas campanulate with
reflexed lobes); corolla white, yellow, greenish, or red; mature fruit black, dark blue, or orange.
36a. Corollas (sub)campanulate, tube (0.5)0.81.2 mm, lobes reflexed, 1.21.5 mm.
37a. Stems 4-ridged to markedly winged; leaves lanceolate, length/breadth index more than 3 .............. 1. R. alata
37b. Stems quadrangular but never winged; leaves broader, length/breadth index less than 3.
38a. Leaves of main stems ovate-cordiform to suborbicular-cordiform, as long as wide or
slightly longer than wide, when dry adaxially mealy green or pale green ............................ 20. R. ovatifolia
38b. Leaves lanceolate, oblong-ovate, oblong-suborbicular, or ovate, 23 as long as wide,
when dry green, brownish green, black, or perhaps red ........................................................ 22. R. podantha

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308

36b. Corollas rotate, fused part 0.20.5 mm, lobes spreading.

39a. Corollas purplish red, with lobes 34 mm ......................................................................... 18. R. membranacea
39b. Corollas white, yellow, greenish yellow, or purplish red, with lobes 12.5 mm.
40a. Corollas pale yellow to white, with lobes 22.5 mm ................................................................ 21. R. pallida
40b. Corolla variously colored, with lobes 11.5 mm.
41a. Leaves linear to narrowly lanceolate, 510 as long as wide ........................................... 27. R. salicifolia
41b. Leaves broader, oblong-lanceolate, oblong-ovate, ovate, cordiform, or suborbicular,
not more than 4 as long as wide.
42a. Leaves ovate to suborbicular, length/breadth index 11.8, base cordate or cordulate.
43a. Largest leaves mostly shorter than 4 cm, usually cuspidate or apiculate toward
apex, length/breadth index 11.8, drying thickly papery, blackening, with
lateral veins usually impressed and tertiary venation visible; petioles 0.55 cm ............. 2. R. argyi
43b. Largest leaves mostly longer than 4 cm, usually attenuate toward apex,
length/breadth index 1.21.5, drying thinly papery, often remaining greenish,
lateral veins never impressed and tertiary venation less visible; petioles 211 cm ..... 31. R. sylvatica
42b. Leaves lanceolate to oblong-lanceolate, attenuate toward apex, length/breadth
index 2.54, lateral veins never impressed and tertiary venation less visible,
base truncate, rounded, or cordulate.
44a. Flowers white; Taiwan .......................................................................................................... 14. R. linii
44b. Flowers purplish red, greenish, yellowish, or white; mainland.
45a. Stems smooth or sparsely aculeolate; flowers purplish red, greenish yellowish,
or whitish; fruit 3.54 mm in diam., black at maturity .......................................... 37. R. wallichiana
45b. Stems rather markedly or sparsely aculeolate; flowers greenish, yellowish, or
whitish; fruit 46(7) mm in diam., orange at maturity ............................................. 5. R. cordifolia
1. Rubia alata Wallich in Roxburgh, Fl. Ind. 1: 384. 1820.
jin jian cao
Rubia cordifolia Linnaeus var. longifolia Handel-Mazzetti;
R. lanceolata Hayata.
Climbers and vines, herbaceous, perennial; stems to 4 m,
quadrangular, 4-ridged, or usually 4-winged at least when older
with wings to 1.5 mm wide, glabrous or hirtellous-puberulent at
nodes, retrorsely aculeolate. Leaves and leaflike stipules in
whorls of 4, often unequal (stipules smaller); petiole 0.210 cm,
on principal axes longer than on lateral ones, those of stipules
often shorter or even lacking; blade drying thinly leathery,
linear-lanceolate or narrowly lanceolate, 3.59 0.42 cm, with
length/breadth index above 3, glabrous and smooth or sometimes sparsely scaberulous, base rounded to cordulate, margin
thinly revolute and usually aculeolate, apex acute; principal
veins palmate, 3(or 5) with lateral veins sometimes weakly evident. Inflorescences thyrsoid, paniculate, with terminal and
axillary, many-flowered cymes; axes smooth to aculeolate,
ridged to thinly winged; bracts elliptic to lanceolate-elliptic,
0.83 mm; pedicels 14 mm. Ovary ca. 0.7 mm, smooth. Corolla white, pale yellow, or greenish, campanulate, fused base
0.51 mm, glabrous; lobes triangular to lanceolate, 1.21.5 mm,
apex caudate-acuminate. Mericarp berry black, 57 mm. Fl.
MayAug, fr. AugNov.
Forest margins on mountain slopes, thickets; 6002000 m. Provinces south of the Chang Jiang, east to Taiwan, west to Sichuan, north to
C Henan and S Shanxi: Anhui, Fujian, Gansu, Guangdong, Guangxi,
Guizhou, Henan, Hubei, Jiangxi, Shaanxi, Sichuan, Yunnan, Zhejiang
[Nepal].
Rubia alata, an obvious member of R. ser. Cordifoliae, is here
treated in the sense of H. S. Lo (in FRPS 71(2): 312, t. 70, f. 16. 1999),

which may or may not correspond to its original application and its
type. Accordingly, it is mainly characterized by relatively narrow leaves,
petioles frequently bent near the base of the blade, stem angles with thin
ridges to narrow or remarkably well-developed wings, and paniculate
inflorescences, small flowers, and black fruit similar to other species of
the R. cordifolia group (see additional comments under that species).
The protologue of R. alata does not address the shape of the leaves and
describes the stems as winged or not. This suggests the possibility that
that the types leaves are not markedly narrower and its stems not more
markedly winged than those of R. cordifolia and related species. Rubia
alata was not treated by Deb and Malick (Bull. Bot. Surv. India 10(1):
116. 1968) for India, nor by Long (Fl. Bhutan 2(2): 823825. 1999) for
Bhutan. The Kew Rubiaceae checklist (Govaerts et al., World Checkl.
Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on 15 Sep
2010) considers it to be a synonym of R. cordifolia but gives no source
for this conclusion. From the relatively abundant herbarium material
studied and the rather narrow species concept used in the present
treatment, we believe that it is justified to distinguish R. alata sensu H.
S. Lo and R. cordifolia s.s.
Rubia lanceolata from Taiwan is provisionally referred here as a
synonym to R. alata. Some of the named varieties of R. cordifolia may
also belong to this species but a clarification is impossible with our
present insufficient knowledge of R. ser. Cordifoliae. The herbarium
name R. cordifolia var. stenophylla Franchet does not appear to have
ever been published anywhere.
The numerous collections from the Biodiversity Survey of the
Gaoligong Shan area in Yunnan have revealed the common occurrence
of Rubia alata and the presence of rare R. siamensis together with many
populations, which link these two quite different taxa. These intermediates exhibit most varied differential character recombinations of the two
species with respect to leaf shape, from broadly lanceolate to cordate
(length/breadth index below 3) or from small triangular and sessile stipules to fully leaflike elements with long petioles. In addition, new characters appear, such as greenish to yellowish fruit colors. The suspicion
that all this is the result of hybridization needs support by further
studies.

RUBIACEAE

309

2. Rubia argyi (H. Lveill & Vaniot) H. Hara ex Lauener & D.

K. Ferguson, Notes Roy. Bot. Gard. Edinburgh 32: 114. 1972.

3. Rubia chinensis Regel & Maack in Regel, Tent. Fl. Ussur.

76. 1861.

dong nan qian cao

zhong guo qian cao

Galium argyi H. Lveill & Vaniot, Bull. Soc. Bot. France

55: 58. 1908; Rubia akane Nakai; R. akane var. erecta Masamune; R. chekiangensis Deb.

Herbs, perennial, rhizomatous; stems erect, to 60 cm tall,

solitary or usually grouped, unbranched or few branched, quadrangular, glabrous to pilosulous at least near nodes, ribs
rounded, smooth to sparsely scaberulous. Leaves in whorls of 4;
petiole (0.3)0.52 cm; blade drying thinly papery, ovate, oblong-ovate, or broadly elliptic, 310 1.24.5 cm, length/
breadth index 1.82.3, adaxially subglabrous and scaberulous,
abaxially glabrous to pilosulous, base obtuse, rounded, or cordulate, margin scaberulous to ciliate, apex acute or acuminate;
principal veins 5 or 7, palmate. Inflorescences thyrsoid, terminal and axillary in upper nodes, paniculate, many flowered, 5
30 cm, glabrous to pilosulous; axes scaberulous to smooth;
bracts lanceolate, 1.58 mm; pedicels 27 mm. Ovary ca. 0.8
mm, smooth to scaberulous. Corolla greenish white, rotate, 34
mm in diam., glabrous, with fused base 0.20.6 mm; lobes 5,
lanceolate, 1.72 mm, acute to caudate. Mericarp berry black,
ca. 4 mm in diam., smooth. Fl. MayJul, fr. SepOct.

Vines, herbaceous, perennial (rarely erect in mountains of

Taiwan: Rubia akane var. erecta); stems probably to 2 m or
more, quadrangular to narrowly 4-winged especially when
older, glabrous to pilosulous, retrorsely aculeolate sometimes
becoming smooth with age. Leaves nearly exclusively in whorls
of 4, equal or sometimes unequal; petiole 0.55 cm; blade
when drying often blackening, thickly papery, suborbicularcordiform, broadly ovate-cordiform, or oblong-suborbicular,
(1)24.5(5) (1)1.53.5(4) cm, length/breadth index 1
1.8, glabrous or sparsely to densely pilosulous or hirtellous,
scaberulous, base cordate to cordulate, margins sparsely to
densely retrorsely aculeolate, apex cuspidate to apiculate; principal veins palmate, 5 or 7, veins usually impressed and tertiary venation visible. Inflorescences thyrsoid, paniculate, with
terminal and axillary, many-flowered cymes, axes aculeolate,
glabrous to pilosulous; bracts lanceolate or lanceolate-elliptic,
14 mm; pedicels rather stout, 12.5 mm. Ovary ca. 0.8 mm,
smooth. Corolla yellowish-greenish to white, rotate to bellshaped, glabrescent, fused base 0.50.7 mm; lobes lanceolate,
1.31.4 mm, spreading to reflexed. Mericarp berry black,
subglobose 57(9) mm in diam. Fl. JulOct, fr. AugNov.
Forest margins, thickets, fences at village sides; 3003400 m.
Anhui, Fujian, Guangdong, Guangxi, Henan, Hubei, Hunan, Jiangsu,
Jiangxi, Shaanxi, Sichuan, Taiwan, Zhejiang [Japan, Korea].
Rubia akane is here treated as a synonym of R. argyi, following
the Kew Rubiaceae checklist (Govaerts et al., World Checkl. Rubiaceae;
http://www.kew.org/wcsp/rubiaceae/; accessed on 15 Sep 2010), Yamazaki (Fl. Japan 3a: 232. 1993), and H. S. Lo (in FRPS 71(2): 316. 1999).
The recent Taiwanese flora (T. Y. A. Yang, Fl. Taiwan, ed. 2, 4: 322.
1998), however, continues to regard R. akane as a separate species and
characterizes it by white flowers, whereas R. argyi generally is regarded
as yellowish-greenish flowering. Aside from corolla color, no differential characters are apparent between the two taxa. However, in addition
to the typical climbing and vinelike var. akane in Taiwanese localities
below 2000 m, at higher elevations and more open habitats of the Taiwanese mountains, there is an upright growing type called R. akane var.
erecta, which seems to be endemic. As a species it has been called R.
nankotaizana Masamune (Hokuriku J. Bot. 2: 40. 1953). If it is actually
conspecific, it should be transferred as a variety to R. argyi.
Rubia argyi belongs to R. ser. Cordifoliae and is well illustrated in
the FRPS treatment by H. S. Lo (loc. cit.: 317, t. 71, f. 16). With the
differential characters available (mainly the relatively broad and short
leaves with veins impressed), it can be separated rather clearly from the
very closely related members R. cordifolia s.l. (or better R. cordifolia
agg., see there). To lump it with R. cordifolia (e.g., Z. Ying Zhang, Fl.
Tsinling. 1(5): 14. 1985) does not appear justified. Within the R. cordifolia agg. R. argyi comes closest to R. sylvatica, from which it deviates
primarily by its larger leaves, partly in whorls of more than 4. Rubia
ovatifolia can be separated by its definitely campanulate flowers with a
fused base of 0.81 mm. For additional comments see under R. cordifolia.

Forests on mountains, forest margins, meadows; 2001400 m. E

and N China [Japan, Korea, Russia].
Rubia chinensis and a few related species with erect (not climbing)
stems and broad leaves with short petioles have been placed by
Pojarkova (Fl. URSS 23: 391392. 1958) into R. ser. Chinenses. Other
Chinese species of this group include the closely related R. latipetala
and R. schumanniana. The R. mandersii species group (see there) is
similar but has sessile leaves.
For Rubia chinensis H. S. Lo (in FRPS 71(2): 301, t. 66, f. 813.
1999) gives good drawings and differentiates two varieties (elsewhere
also treated as forms). They are listed here for reference but apparently
are not well marked in China. They were not included in the Fl. Hebei.
(2: 577. 1988), and in Fl. Japan (Yamazaki, 3a: 231232. 1993) they
were synonymized and said to be difficult to distinguish. Generally, it
appears that Japanese populations of R. chinensis have somewhat
smaller leaves than those from China.

1a. Leaves adaxially subglabrous or

hirtellous along principal veins,
pilosulous abaxially ................................. 3a. var. chinensis
1b. Leaves glabrous or aculeolate along
midvein ............................................... 3b. var. glabrescens
3a. Rubia chinensis var. chinensis
() zhong guo qian cao (yuan bian zhong)
Rubia mitis Miquel.
Leaves subglabrous or hirtellous adaxially along principal
veins, abaxially pilosulous, margin ciliate.
E and N China [Japan, Korea, Russia].

3b. Rubia chinensis var. glabrescens (Nakai) Kitagawa, Lin.

Fl. Manshur. 405. 1939.
wu mao da zhen cao
Rubia mitis f. glabrescens Nakai, J. Jap. Bot. 14: 115.
1938; R. chinensis f. glabrescens (Nakai) Kitagawa.

RUBIACEAE

310

Leaves glabrous except aculeolate adaxially along midvein, abaxially glabrous, margin scaberulous.
NE China [Japan, Korea].

4. Rubia chitralensis Ehrendorfer, Nytt Mag. Bot. 3: 228.

1954.
gao yuan qian cao
Herbs, perennial, with woody rootstock; stems up to 50
cm tall, clustered, subterete to slightly winged, glabrous,
smooth. Leaves in whorls of up to 6(or 7), sessile; blade drying
membranous, lanceolate-ovate to lanceolate, (2)45 0.71.5
cm, adaxially glabrous or sparsely antrorsely scaberulous, abaxially glabrous with midrib antrorsely aculeolate, margins retrorsely scaberulous, apex acute to acuminate; principal vein 1,
lateral veins 2, weak. Inflorescences terminal, pyramidal, leaflike, with 1- to several-flowered cymes; peduncles glabrous,
smooth; bracts elliptic to ovate, 67 mm; pedicels 26 mm.
Ovary 12 mm, glabrous. Corolla yellowish green to whitish
yellow, rotate, 67.5 mm in diam., fused basal part 0.50.7 mm;
lobes lanceolate, ca. 3 mm, acuminate to incurved-mucronate,
arista to ca. 0.8 mm. Mericarp berry black and shiny, 3.55 mm
in diam., glabrous. Fl. JunJul, fr. JulOct.
Stony slopes in Juniperus and subalpine belt; ca. 2900[4000] m.
Xinjiang [Afghanistan, Pakistan, Tajikistan, Uzbekistan].
Rubia chitralensis is distantly related to other SW to C Asiatic but
rather subshrubby taxa, as R. gedrosiaca Bornmller, R. laxiflora
Gontscharow, or R. tibetica. These taxa were placed by Pojarkova (Fl.
URSS 23: 398417. 1958) into several series of the heterogeneous R.
sect. Campylanthera, distributed from C Asia to the Mediterranean.

5. Rubia cordifolia Linnaeus, Syst. Nat., ed. 12, 3: 229. 1768.

qian cao
Rubia cordifolia var. coriacea Z. Ying Zhang; R. cordifolia
subsp. pratensis (Maximowicz) Kitamura; R. cordifolia var.
pratensis Maximowicz; R. cordifolia var. rotundifolia Franchet;
R. pratensis (Maximowicz) Nakai.
Vines, herbaceous, climbing or scrambling herbs, with red
rhizomatous base and roots; stems to 3.5 m, several to many
from base, often much branched, quadrangular, glabrous to
puberulent, with ribs rounded to thinly winged, sparsely to
densely retrorsely aculeolate. Leaves in whorls of 4 or more (up
to 8 or rarely 12); petiole (1)1.53(6) cm; blade drying papery to thickly papery, usually remaining greenish, lanceolate, oblong-lanceolate, ovate, or oblong-ovate, (1)1.54(7)
(0.3)0.51.5(2.5) cm, length/breadth index mostly 2.54,
glabrous to pilosulous or hirtellous, sparsely to densely scaberulous, base rounded, truncate, cordulate, or cordate, margin serrulate-aculeolate, apex obtuse and apiculate to acute or acuminate; principal veins 3 or 5, palmate. Inflorescences thyrsoid,
paniculate, with terminal and axillary, several- to many-flowered cymes; axes glabrous to puberulent or pilosulous, aculeolate; bracts linear-lanceolate to ligulate, 13 mm; pedicels 1
4 mm. Ovary 0.50.8 mm, smooth to scaberulous. Flowers
hermaphroditic (rarely polygamo-dioecious?). Corolla pale yellow or greenish yellow, rotate, glabrous, fused base 0.20.4
mm; lobes lanceolate, spreading to reflexed, 1.21.5 mm, cau-

date. Mericarp berry becoming orange then apparently black,

46 mm in diam. Fl. AugSep, fr. OctNov.
Sparse forests, forest margins, grasslands; 3002800 m. Anhui,
Gansu, Hebei, Hunan, Qinghai, Shandong, Shanxi, Sichuan, Xizang,
Yunnan [Japan, Korea, Mongolia, Russia (Far East); S and SE Asia to
Sri Lanka and Java, through the Himalaya to Afghanistan; (sub)tropical
Africa].
As noted by most previous authors (Pojarkova, Fl. URSS 23:
387391. 1958; Ehrendorfer et al., Fl. Iranica 176: 5253. 2005), the
plants included in Rubia cordifolia s.l. comprise a geographically very
widespread (from E and SE Asia to Afghanistan, from Sudan to S
Africa), morphologically extremely polymorphic, polyploid, and still
most insufficiently understood racial complex. Its populations, together
with related taxa, have been grouped into R. ser. Cordifoliae by Pojarkova (loc. cit.), characterized by their generally clambering to climbing
habit; leaves and leaflike stipules in whorls of 4 or more, petiolate,
palmately 37-veined; and corollas rotate to shortly campanulate, with
anthers ellipsoid, somewhat curved, and 46 shorter than the corolla
lobes. Depending on narrow or wider species concepts and differential
characters chosen, the elements of this series have been quite variously
treated. In the present flora the following 16 species are assembled in R.
ser. Cordifoliae: 1. R. alata, 2. R. argyi, 5. R. cordifolia, 6. R. crassipes,
14. R. linii, 17. R. manjith, 18. R. membranacea, 19. R. oncotricha, 20.
R. ovatifolia, 21. R. pallida, 22. R. podantha, 25. R. pterygocaulis, 27.
R. salicifolia, 31. R. sylvatica, 35. R. trichocarpa, and 37. R. wallichiana. Species 5, 14, 20, 31, and 37 are so close and linked by occasional
intermediates that they can be understood as R. cordifolia s.l. or R.
cordifolia agg. The above species description refers to R. cordifolia s.s.
The type specimen of Rubia cordifolia in the Linnaean Herbarium
(no. 131.7, LINN) has no flowers or fruit, but its distinct habit with
leaves in whorls of 4, oblong-cordate, acute, and with long petioles
corresponds to the above description of the species in a more narrow
sense and to the figure in H. S. Lo (in FRPS 71(2): 307, t. 68, f. 712.
1999). The complications in the typification of R. cordifolia have been
detailed by Jarvis (Order Out of Chaos, 800. 2007). The description by
Linnaeus was emended by Gaertner (Novi Comment. Acad. Petrop.
14(1): 541. 1770). The original reference to 4-merous flowers may
have been due to the occasional occurrence of 4- among the typical 5merous flowers or simply to a mistake. The fruit were originally described as unknown, but later their color was given as red. Pojarkova
(loc. cit.: 466467) noted for R. cordifolia and for R. ser. Cordifoliae as
a whole that the fruit were orange or brownish when immature and
black when fully mature and dry. Personal observations revealed a
group of distinctive Chinese specimens with vegetative parts drying
yellowed and the mature, or near-mature, fruit drying clear bright orange but evidently turning black at maturity (e.g., Fu Kunjun 10394,
MO!). Thus, fruit color may be of taxonomic relevance in Rubia but is
in need of more detailed studies.
Even with the present, rather narrow circumscription, there is still
much variation among the Chinese populations of Rubia cordifolia. This
refers to indumentum, consistency, shape and size of leaves, number of
leaves and leaflike stipules per whorl, flower shape, and fruit color. Leaf
indumentum does not seem to be correlated with that of the inflorescence axes. Instead, either may be glabrous or pubescent, apparently independently, which is unusual in Rubiaceae. In zones of contact, particularly with the closely related R. sylvatica and R. ovatifolia, one has
to expect transitional individuals. The status of R. wallichiana (see
there) and its separation from R. cordifolia is doubtful anyway.
The infraspecific synonymy of Rubia cordifolia listed above
follows H. S. Lo (loc. cit.: 315); it has not yet been checked in detail for
lack of any more authoritative treatment of R. ser. Cordifoliae. Rubia
cordifolia var. coriacea was not listed by H. S. Lo and is here synony-

RUBIACEAE

mized provisionally, as we have seen no authentic material. According

to its protologue, it differs from typical R. cordifolia in its subleathery
leaves, which are glabrous below. With respect to R. cordifolia var.
munjista (Roxburgh) Miquel see R. manjith.

6. Rubia crassipes Collett & Hemsley, J. Linn. Soc., Bot. 28:

68. 1890.
hou bing qian cao
Vines, herbaceous, climbing, to 3 m; stems quadrangular
becoming subterete, tuberculate-hispid, scabrous. Leaves in
whorls of 4, often unequal; petiole 0.51.8 cm or smallest
leaves sometimes subsessile; blade drying thickly leathery,
ovate-oblong to elliptic, (0.7)24.5 cm, tuberculate-hispid,
base rounded to cordulate, margin slightly revolute, apex obtuse; principal veins 5, palmate. Inflorescences thyrsoid, paniculate, 37 cm, with axillary and terminal cymes; axes rather
stout; bracteoles ovate or lanceolate, 2.53.5 mm. Flowers
unknown. Immature mericarp berry black when dry, ca. 4 mm
in diam. Fr. autumn.
Forest margins on mountain ridges; 14002400 m. Yunnan
[Myanmar, Thailand].
We have seen no authentic material of Rubia crassipes. The above
description corresponds to the protologue and to H. S. Lo (in FRPS
71(2): 313. 1999). The extended description and photo of plants from
Thailand regarded as this species by Puff (Fl. Thailand: Rubiaceae;
http://homepage.univie.ac.at/christian.puff/FTH-RUB/FTH-RUB_
HOME.htm; accessed on 5 Oct 2010) deviates by straight and soft
hairs in addition to the scabrous indumentum and considerably larger
leaves. Puff also added data on the shallowly campanulate, 5-lobed
flowers 2.44 mm in diam., and considered the taxon as part of the R.
cordifolia group. Only further extensive studies will clarify the doubtful delimitation and taxonomic placement of R. crassipes.

7. Rubia deserticola Pojarkova in Schischkin, Fl. URSS 23:

722. 1958.
sha sheng qian cao
Plants herbaceous, perennial, with woody rootstock; stems
suberect, to 1 m tall quadrangular, angles recurved prickly.
Leaves in whorls of 46, subsessile; blade drying firmly leathery, lanceolate to elliptic-oblong, aculeolate abaxially on midrib
and along thickened margins, base acute, apex acuminate; principal vein 1. Inflorescences terminal, many flowered, leafy and
bracteose. Ovary 12 mm, glabrous. Corolla yellowish white,
rotate, fused basal part ca. 0.3 mm; limb 3.54.5 mm in diam.;
lobes lanceolate, aristate. Anthers very small, ca. 0.3 mm. Mericarp berry black, 3.55 mm in diam. Fl. Jun, fr. Jul.
Sandy and salty semideserts. Xinjiang (Yili) [Kazakhstan].
We have seen no authentic material of Rubia deserticola. This
local C Asiatic semidesert species was regarded as close to the common
Mediterranean R. peregrina Linnaeus by Pojarkova (loc. cit.) and placed
into R. ser. Peregrinae Pojarkova of R. sect. Campylanthera.

8. Rubia dolichophylla Schrenk, Bull. Cl. Phys.-Math. Acad.

Imp. Sci. Saint-Ptersbourg 2: 115. 1844.
chang ye qian cao
Herbs, perennial, rhizomatous; stems erect, to 1 m tall,
glabrous, angled, retrorsely aculeolate. Leaves in whorls of 4,

311

sessile or subsessile; blade drying papery, linear to lanceolateoblong, 512 0.51.4 cm, glabrous, abaxially prickly on midrib and often on veins, base acute to cuneate, margins revolute
and antrorsely aculeolate, apex acute to acuminate; lateral veins
pinnate, 610 pairs. Inflorescences thyrsoid, with terminal and
axillary, several- to many-flowered cymes, nearly as long as or
shorter than subtending leaves; axes aculeolate; bracts linear, 2
5 mm; pedicels 26 mm. Ovary 11.2 mm in diam. Corolla
pale yellow, rotate, fused basal part ca. 0.6 mm; lobes ovate, ca.
2 mm, abruptly contracted with arista ca. 0.5 mm. Mericarp
berry black, 2.55 mm. Fl. MayAug, fr. JulSep.
Along rivers and among rocks; 19002100 m. Xinjiang [Afghanistan, Kazakhstan, Pakistan, Tajikistan; SW Asia (Iran)].
Rubia dolichophylla, well illustrated in H. S. Lo (in FRPS 71(2):
293, t. 62, f. 16. 1999), according to Pojarkova (Fl. URSS 23: 404
407. 1958) is one of the two species of R. ser. Dolichophyllae Pojarkova, both C Asiatic rhizome-forming herbs. Specimens of R. jesoensis
(Miquel) Miyabe & Miyaki from Japan with a similar growth form
might key out as R. dolichophylla but are readily separable by their
stems with vegetative apex and exclusively axillary inflorescences with
cymes borne along the lower stem portion and usually shorter than the
supporting leaves. In R. dolichophylla the partial inflorescences are
terminal and axillary. As already correctly shown by Pojarkova (loc.
cit.), R. jesoensis appears related to R. tatarica (Treviranus) F. W.
Schmidt from SE Russia and adjacent Siberia, and both belong to R.
ser. Tataricae Pojarkova.

9. Rubia edgeworthii J. D. Hooker, Fl. Brit. India 3: 203. 1881.

chuan dian qian cao
Herbs, scandent to climbing; stems 8-ribbed, scabrous.
Leaves in whorls of 4, subsessile; blade drying papery, lanceolate to lanceolate-elliptic, 47 1.22 cm, both surfaces scaberulous and sometimes hirtellous, base acute to obtuse, margins
entire, apex acute to acuminate; principal veins 3 or 5, palmate.
Inflorescences thyrsoid, paniculate, with terminal and axillary,
many-flowered cymes, usually much longer than subtending
leaves; axes hirtellous, scaberulous to glabrescent; bracts lanceolate or subovate, 25 mm; pedicels 25 mm. Ovary subglabrous to hirtellous. Corolla pale yellow, somewhat funnelshaped, glabrous or hirtellous outside, fused basal part ca. 0.5
mm; lobes 5, ovate to lanceolate, 11.2 mm, obtuse to acute.
Mericarp berry unknown. Fl. Sep.
Grassy slopes; ca. 2100 m. Guangxi (Longlin), Sichuan (Miyi),
Yunnan (Heqing) [N India].
We have not seen authentic material of Rubia edgeworthii, but
both Deb and Malick (Bull. Bot. Surv. India 10(1): 1112. 1968) and
H. S. Lo (in FRPS 71(2): 304, t. 67, f. 813. 1999) presented good
drawings. Nevertheless, nothing is known about the important underground organs of this species (rootstock or rhizome?). Judging from
other characters, R. edgeworthii may belong to the vines of the R. sikkimensis group with R. tenuis (see there), but one also has to consider
the upright, not climbing members of the R. mandersii group; both are
elements of R. sect. Oligoneura. H. S. Lo (loc. cit.: 308) noted that the
unpublished name Rubia lancilimba F. C. How has been written on
specimens of this species.

10. Rubia falciformis H. S. Lo, J. Trop. Subtrop. Bot. 7(1): 23.

1999.
lian ye qian cao

RUBIACEAE

312

Herbs, perennial, presumably scandent vines; stems quadrangular, scabrous. Leaves opposite; petiole 34 cm, rather
stout, retrorsely aculeolate; blade drying greenish, thickly leathery, falcate-lanceolate, 1115 cm, both surfaces scaberulous,
usually aculeolate along principal veins, base rounded or subcordate, margin revolute and aculeolate, apex acuminate; principal veins 5, palmate, impressed above, with higher order
reticulate veins immersed; stipules caducous, unknown. Inflorescences axillary, with many-flowered cymes. Flowers unknown. Mericarp berries dark orange-yellow when dry, 3.54
mm in diam., binary or solitary, densely villous with pubescence drying ferruginous. Fr. Oct.
Wet lands in forests; ca. 1100 m. Yunnan (Lianghe).
Authentic material of Rubia falciformis has not been available, but
H. S. Lo (in FRPS 71(2): 295, t. 63, f. 911. 1999) presented a good
drawing. With the exception of the strongly hairy fruit and the deciduous (and unknown) stipules, its description corresponds to R. siamensis,
also known from Yunnan, and thus belongs to its group within R. sect.
Oligoneura.

11. Rubia filiformis F. C. How ex H. S. Lo, J. Trop. Subtrop.

Bot. 7(1): 24. 1999.
si geng qian cao
Vines, herbaceous, to 23 m tall; stems quadrangular or
narrowly 4-winged, glabrous or subglabrous, retrorsely aculeolate on angles. Leaves opposite; petiole (2)315(25) mm;
blade drying thinly leathery or papery, ovate or rarely ovatelanceolate, 29 14.5 cm, both surfaces glabrous and smooth
or sparsely aculeolate along veins, base rounded, subcordate, or
obtuse, margins entire and smooth, apex acute to acuminate;
principal veins 3 or 5(or 7), palmate, adaxially impressed; stipules ovate, ca. 5 mm, usually caducous. Inflorescences thyrsoid,
paniculiform, with terminal and axillary, many-flowered cymes
and slender, subfiliform, smooth and glabrous axes. Flowers
unknown. Fruiting pedicels 714 mm; mericarp berries black,
45 mm in diam. Fr. in late winter.
Forests; 10001500 m. Yunnan (Malipo).

mm. Mericarp berry black, 34 mm in diam. Fl. summer

early autumn, fr. late autumnearly winter.
Dry and rocky meadows; 30003800 m. Sichuan, NW Yunnan.
Together with Rubia angustissima Wallich ex G. Don and R.
charifolia Wallich ex G. Don from the Himalaya, both with greenishyellowish flowers, the dark-red flowering R. haematantha forms an
aberrant, closely related species assembly, provisionally called R.
angustissima group. It occurs from the W Himalaya (Kashmir) to Nepal, Myanmar, Bhutan, and SW China. The group shares retrorsely
aculeate stems, linear to filiform leaves in whorls of up to 8(10), and
caudate corolla lobes. Species delimitation within this group is still
provisional. Whereas Deb and Malick (Bull. Bot. Surv. India 10(1): 5.
1968) unite R. angustissima and R. charifolia (R. charaefolia), the
two taxa are maintained by Long (Fl. Bhutan 2(2): 823825. 1999). The
reliability of the flower color as differential character of R. haematantha
also needs further study. If only one species is accepted, its name would
have to be R. angustissima.

13. Rubia latipetala H. S. Lo, J. Trop. Subtrop. Bot. 7(1): 23.

1999.
kuo ban qian cao
Herbs, perennial, with ?rootstock; stems erect, to 20 cm
tall, little branched, quadrangular, glabrous, smooth. Leaves in
whorls of 4; petiole 0.81.8 cm; blade drying greenish, papery,
broadly ovate, 1.53.5 13 cm, length/breadth index 1.2
1.5, glabrous except hispid on principal veins, base cordulate
to cordate, apex acute; principal veins 35, palmate. Inflorescences thyrsoid, with terminal and sometimes axillary, 25flowered and 0.51 cm long cymes; axes glabrous. Corolla
green, turning blackish when dry, bell-shaped, fused part 1.5
2 mm; lobes triangular, ca. 1.5 mm. Mericarp berries unknown.
Fl. Aug.
Forest margins; ca. 3400 m. Sichuan (Barkam).
No authentic material of Rubia latipetala was available to us.
From the description it belongs to R. ser. Chinenses. A completely glabrous and smooth specimen with larger, ovate-cordate leaves (to 5.5
3.5 cm) and petioles up to 3.5 cm (1800 m, 21 Jul 1988, PE sheet no.
1847036) may belong here.

We have seen no authentic specimens of Rubia filiformis, but

there is a good drawing in H. S. Lo (in FRPS 71(2): 297, t. 64, f. 15.
1999). The species evidently belongs to the R. siamensis group of R.
sect. Oligoneura.

14. Rubia linii J. M. Chao, Biol. Bull. Natl. Taiwan Norm.

Univ. 1: 48. 1966.

12. Rubia haematantha Airy Shaw, Bull. Misc. Inform. Kew

1931: 450. 1931.

Herbs, climbing or scrambling, to 1.5 m; stems subterete,

glabrous, smooth to sparsely aculeolate. Leaves in whorls of 4;
petiole 14.5 cm, glabrous; blade drying thinly papery, lanceolate, lanceolate-oblong, or oblong-ovate, 27.5 0.53 cm,
length/breadth index 2.54, glabrous, upper side sparsely
scaberulous, base truncate, rounded, or cordulate, margin flat to
thinly revolute and smooth, apex acute to acuminate; principal
veins 3(or 5), palmate. Inflorescences thyrsoid, paniculate, with
terminal and axillary, several- to many-flowered cymes; axes
glabrous and smooth; bracteoles lanceolate to narrowly elliptic,
13 mm; pedicels 14 mm. Ovary 0.30.5 mm, smooth. Corolla white, rotate, ca. 2.5 mm in diam., with lobes usually reflexed, outside glabrous, fused base ca. 0.5 mm; lobes 5, triangular, 11.2 mm, acute. Mericarp berry black, 47 mm in diam.
Fl. MayAug, fr. Apr, Jul.

hong hua qian cao

Herbs, perennial, erect to somewhat climbing, with woody
rootstock; stems 0.5 m or more, clustered, quadrangular, angles
retrorsely aculeolate. Leaves in whorls of up to 68(10),
narrowly linear, 2050 0.51 mm, base acute, margins revolute and antrorsely aculeolate, otherwise glabrous and smooth,
single vein forming a thick midrib ca. 1/2 of leaf breadth, apex
acute, with hyaline point. Inflorescence thyrsoid, with lax, fewflowered axillary and terminal cymes; bracts filiform; pedicels
ca. 5 mm. Ovary ca. 0.5 mm, glabrous. Corolla dark red (perhaps sometimes ?white), rotate, 34 mm in diam., glabrous;
lobes triangular, 3-veined, abruptly caudate with acumen ca. 1

lin shi qian cao

RUBIACEAE

Margins of evergreen forests; 5003000 m. Taiwan.

Rubia linii was accepted in the Fl. Taiwan (ed. 2, 4: 324. 1998),
where neither R. cordifolia nor R. wallichiana figure. In FRPS (71(2):
316. 1999) it was treated in a comment under R. wallichiana, as a possible misidentification of that species. Rubia linii is an obvious member
of the R. cordifolia complex (= R. cordifolia agg., see additional comments under that species). More detailed studies will have to demonstrate its doubtful specific status; it is obviously very close to R. cordifolia and R. wallichiana.

15. Rubia magna P. K. Hsiao, Acta Phytotax. Sin. 23: 390.

1985.
e mei qian cao
Vines, herbaceous, to 3 m; stems quadrangular or narrowly
4-winged, sparsely retrorsely aculeolate on angles. Leaves opposite; petiole 14(6) cm, retrorsely aculeolate and sometimes
ferruginous hirtellous; blade drying thickly papery to leathery,
light green and ferruginous, particularly below and on main
veins, ovate, lanceolate, or oblanceolate, larger leaves 412.5
1.55(7) cm, smaller 23 12 cm, glabrous to hirtellous, retrorsely aculeolate at least along veins on both surfaces, base
rounded to cordate, margins serrulate-aculeolate, apex acuminate; principal veins 3 or mostly 5; stipules conspicuous, persistent, broadly ovate, (5)1060 (4)840 mm, acuminate.
Inflorescences predominantly axillary, with several- to manyflowered cymes; axes glabrous to hirtellous, sparsely aculeolate; bracts ovate or sublanceolate, 15 mm; pedicels 14 mm.
Ovary ca. 1 mm, glabrous to glabrescent. Corolla yellow to
green, 5-merous, cup-shaped to campanulate, 3.54.5 mm in
diam., papillose to hairy outside; tube 0.81.3 mm; lobes triangular-ovate, 1.51.7 mm, caudate. Mericarp berry black, 45
mm in diam. Fl. JunJul, fr. AugOct.
Broad-leaved subtropical forests; 12001500 m. Sichuan, Yunnan.
The above description of Rubia magna combines data from the
protologue, H. S. Lo in FRPS (71(2): 295, t. 63, f. 18. 1999), and new
collections from the Gaoligong Shan region of Yunnan. The latter are
conspicuous by their lack of hairiness and their conspicuous stipules
(much larger than shown in FRPS). Certain discrepancies (protologue:
leaves sometimes sessile and opposite but sometimes petiolate and 35verticillate; H. S. Lo (loc. cit.: 294): corolla lobes 4) may be due to
mixed material and still need clarification.
Rubia magna is mainly characterized by its opposite leaves,
dried green and ferruginous blades, and large stipules. It belongs,
together with R. falciformis and R. filiformis, to the closely related R.
siamensis group from SW China and adjacent areas.
Originally, Rubia magna was described as a new species of
medicinal Rubia, and presumably its uses are detailed in that article.

16. Rubia mandersii Collett & Hemsley, J. Linn. Soc., Bot. 28:
68. 1890.
hei hua qian cao
Herbs, perennial, with slightly woody rootstock; stems 20
60 cm tall, erect, unbranched to diffusely branched, quadrangular to narrowly winged, glabrous, ribs retrorsely aculeolate and
scabrous to glabrous. Leaves in whorls of 4, sessile; blade
drying thickly papery, broadly elliptic-oblong, ovate, or subor-

313

bicular, 1.53.5 0.81.9 cm, glabrous but scaberulous at least

on upper surface, base rounded to cuneate and subpetiolate,
margin aculeolate, apex obtuse or acute; principal veins palmate, 35. Inflorescences thyrsoid, narrowly paniculate, with
terminal and axillary, many-flowered, long-pedunculate cymes;
axes glabrous; bracteoles reduced, narrowly ligulate to lanceolate; pedicels 24 mm. Ovary ca. 1.5 mm, glabrous. Corolla
greenish, yellowish, or whitish, rotate, ca. 5 mm in diam., glabrous, fused basal part 0.50.6 mm; lobes lanceolate or ovate,
1.62 mm, apex incurved. Immature mericarp berry ca. 3 mm
in diam. Fl. Aug, immature fr. Oct.
Dry rocky mountains, Pinus forests; 19003000 m. Sichuan,
Yunnan [Myanmar, Thailand].
Rubia mandersii, together with the closely related R. polyphlebia
and R. yunnanensis, forms a SW Chinese group of erect herbaceous
perennials with rootstock and sessile, lanceolate to broadly ovate, and
scabrous or hairy leaves in whorls of 4(6), which belongs to R. sect.
Oligoneura.

17. Rubia manjith Roxburgh ex Fleming, Asiat. Res. 11: 177.

1810.
fan qian cao
Rubia cordifolia Linnaeus var. khasiana Watt; R. cordifolia var. munjista (Roxburgh) Miquel; R. munjista Roxburgh.
Vines, herbaceous, drying with reddish cast; stems to 3 m,
quadrangular, glabrous, retrorsely aculeolate to smooth, with
red pith. Leaves in whorls of 4, equal or unequal; petiole 0.84
cm, sparsely aculeolate; blade drying papery, mostly greenish
adaxially and purplish red abaxially, oblong-lanceolate, ovatelanceolate, or ovate, (2.5)46(8.5) (0.8)1.82.5(4) cm,
length/breadth index 23, both surfaces glabrous and scaberulous, base rounded to cordate, margin flat to thinly revolute,
aculeolate, apex long acuminate or caudate; principal veins (3
or)5(or 7), palmate. Inflorescences thyrsoid, paniculate, with
terminal and axillary, many-flowered and 2.510 cm long
cymes; axes glabrous and smooth to sparsely aculeolate; bracteoles elliptic-oblong or lanceolate, 0.52 mm; pedicels 1.53.5
mm. Ovary ca. 0.5 mm, smooth. Corolla red, purplish red, or
orange, rotate, glabrous, fused basal part 0.50.6 mm; lobes 5,
lanceolate to triangular, 1.21.5 cm, acuminate. Mericarp berry
dark red, 3.55 mm in diam. Fl. JulAug, fr. Oct.
Broad-leaved forests, Pinus forests and thickets; 7003600 m.
Qinghai, Sichuan, Xizang, Yunnan [Bhutan, India, Nepal].
Rubia manjith belongs to R. ser. Cordifoliae. Among the taxa with
small rotate flowers (R. cordifolia agg.) it is mainly characterized by its
conspicuous reddish cast, particularly on lower leaf sides and flowers (see additional comments under R. cordifolia). A similar cast also
appears in the otherwise quite different R. podantha, a taxon with campanulate flowers.
Deb and Malick (Bull. Bot. Surv. India 10(1): 68. 1968), after a
lengthy discussion, treated Rubia manjith (R. munjista) as a synonym
of R. cordifolia only and identified it with R. cordifolia var. khasiana. In
contrast, Long (Fl. Bhutan 2(2): 823825. 1999) distinguished R. manjith from R. cordifolia largely by its red cast, both alive and dried, but
agreed with the inclusion of R. cordifolia var. khasiana as a synonym.
The same was maintained by H. S. Lo (in FRPS 71(2): 314. 1999), who
added R. cordifolia f. rubra Kitamura as a synonym of R. manjith and

314

RUBIACEAE

qualified it as nom. non rite publ. Both Deb and Malick (loc. cit.) and
Long (loc. cit.) mention the economic and historical importance of the
taxon as a source of an excellent red dye.

18. Rubia membranacea Diels, Notes Roy. Bot. Gard. Edinburgh 5: 279. 1912.
jin qian cao
Rubia membranacea var. caudata Z. Ying Zhang; R. membranacea var. incurvata Z. Ying Zhang.
Vines or climbing herbs; stems to 2 m, quadrangular, glabrous or hirtellous at nodes, scaberulous, retrorsely aculeolate,
or sometimes subsmooth. Leaves in whorls of 4; petiole 0.5
2.5(4) cm; blade drying membranous to papery, lanceolate to
subovate, 16(8) 0.52(4) cm, base rounded to cordate,
margins usually aculeolate, apex acuminate or shortly acuminate; principal veins 3 or 5, palmate. Inflorescences thyrsoid,
paniculate, with terminal and axillary, few- to many-flowered
cymes, 23 cm; axes glabrous and smooth; pedicels 25 mm;
bracts narrowly lanceolate, 15 mm. Ovary ca. 1.8 mm, glabrous. Corolla purplish red, rotate, fused basal part 0.20.6 mm;
lobes spreading, ovate-lanceolate, 23(4) mm, caudate. Mericarp berry dark blue or black, 59 mm in diam. Fl. MayJun, fr.
AugOct.
Sparse forests, forest margins, thickets, grasslands; 11003000
m. Hubei, Hunan, Sichuan, Yunnan.
We have seen no authentic material of Rubia membranacea,
which was well illustrated in H. S. Lo (in FRPS 71(2): 293, t. 62, f. 7
12. 1999). The short cymes indicated together with small leaves and
relatively large rotate purplish flowers might be distinctive. We have
seen only one corresponding collection (Sichuan: Mianning Xian,
Lamagetou Nature Reserve, D. E. Boufford et al. 32941), but it deviates
in habit and more loose cymes.
The two varieties of this species listed above were described and
figured by Z. Ying Zhang (Fl. Tsinling. 1(5): 17, 421. 1985) but not
cited by H. S. Lo (loc. cit.: 314315). They were distinguished from var.
membranacea by the orientation of the corolla lobes, said to be incurved in var. incurvata and long caudate and glabrous in var. caudata,
differences of doubtful taxonomic relevance.

19. Rubia oncotricha Handel-Mazzetti, Symb. Sin. 7: 1031.

1936.
gou mao qian cao
Herbs, climbing or scrambling; stems 0.51.5 m, quadrangular, densely hirtellous or hispidulous with trichomes usually
hooked, angles aculeolate. Leaves in whorls of 4; petiole 0.2
1.5 cm, hirtellous; blade drying rather thickly papery, lanceolate
to ovate, 0.82.5(3.5) 0.30.8(1.5) cm, adaxially densely
scaberulous and sometimes also hooked hirtellous, abaxially
moderately to densely hirtellous, base rounded to usually cordulate, margin thinly revolute and aculeolate, apex obtuse to
shortly acute; principal veins 3(or 5), palmate. Inflorescences
thyrsoid, paniculate, with terminal and axillary, several- to
many-flowered cymes; axes hirtellous or pilosulous, aculeolate;
bracteoles lanceolate to narrowly elliptic, 25 mm; pedicels
0.53(8) mm. Ovary 0.81 mm, smooth. Corolla white or yellow, cup-shaped, outside sparsely to densely hirtellous, fused
basal part 0.81 mm; lobes triangular-ovate, 1.82 mm, cau-

date-acuminate. Mericarp berry 33.5 mm in diam., color

unknown. Fl. JulSep, fr. SepNov.
Forest margins, sparse forests, and grasslands on mountain
slopes; 5003200 m. Guangxi, Guizhou, Sichuan, Yunnan.
Rubia oncotricha, apparently endemic to SW China, is aberrant by
its mostly hooked indumentum (H. S. Lo in FRPS 71(2): 300, t. 65, f.
1014. 1999). Nevertheless, it clearly belongs to R. ser. Cordifoliae (see
additional comments under R. cordifolia).

20. Rubia ovatifolia Z. Ying Zhang ex Q. Lin, J. Wuhan Bot.

Res. 24: 212. 2006.
luan ye qian cao
Vines, herbaceous, perennial, climbing; stems to 1.5 m,
slender, quadrangular, glabrous and smooth to sparsely aculeolate. Leaves in whorls of 4; petiole (1.5)2.55.5(13) cm;
blade drying thinly papery, adaxially green, abaxially pale green,
ovate-cordiform to suborbicular-cordiform, on lateral branches
sometimes ovate, (2)47(12) (1)25(6.5) cm, length/
breadth index 1.52, glabrous to scaberulous, base cordulate to
cordate, margins retrorsely ciliolate or smooth, apex caudateacuminate, rarely obtuse; principal veins 5, palmate. Inflorescence thyrsoid, leafy, with terminal and axillary, few- to manyflowered cymes; axes glabrous and smooth or sparsely aculeolate; bracts linear or lanceolate-linear, 13.5 mm; pedicels 13
mm. Ovary ca. 1 mm, glabrous. Corolla whitish or pale yellow,
subcampanulate, glabrous; tube 0.81 mm; lobes spreading and
somewhat bent, ovate to lanceolate-triangular, ca. 1.4 mm, caudate. Mericarp berry black at maturity, 45 mm in diam. Fl. Jul,
fr. OctNov.
Sparse forests or thickets on mountains; 17002200 m. Gansu,
Guizhou (Bijie), ?Hubei, Hunan, Shanxi, Sichuan, Yunnan, Zhejiang
(Changhua).
This name was first published by Z. Ying Zhang (Fl. Tsinling.
1(5): 15, 420. 1985) but not validly so because two gatherings were designated as types (Vienna Code, Art. 37.2). This was corrected in the
above citation from 2006.
We have not seen authentic material of Rubia ovatifolia, but there
are good drawings in the Tsinling flora and in H. S. Lo (in FRPS 71(2):
307, t. 68, f. 15. 1999). This and the description show that it belongs to
the taxa of R. ser. Cordifoliae and the group with campanulate flowers,
as R. alata or R. podantha. This differential character, so far rather
neglected, separates these taxa, e.g., from R. argyi. Nevertheless, one
has to expect intermediates that link R. ovatifolia with R. sylvatica and
R. cordifolia s.s. (see the latter for additional comments).
H. S. Lo (loc. cit.: 306) differentiated two varieties of doubtful
status: the ranges of petiole length indicated can be found on a single
specimen among the principal and the lateral stems; the descriptions do
not specify which leaves to measure. The so-called var. oligantha
may be a depauperate or very young specimen of this or some other
species. The name was not validly published because no type was indicated and because the name of the species to which it was assigned
was not validly published (Vienna Code, Art. 37.1 and Art. 43.1, respectively). Without further study it should not be validated.

21. Rubia pallida Diels, Notes Roy. Bot. Gard. Edinburgh 5:

277. 1912.
qian se qian cao

RUBIACEAE

Herbs, climbing to scandent; stems to 2 m, quadrangular,

glabrous or sometimes pilose, retrorsely aculeolate. Leaves in
whorls of 4(or 6); petiole 0.56 cm; blade drying papery, lanceolate or subovate, 0.63.5 0.31.5 cm, length/breadth index
ca. 2, subglabrous or adaxially sparsely hirtellous, abaxially
scaberulous, base rounded to cordulate, margin densely serrateaculeolate, apex acuminate; principal veins 3 or 5, palmate. Inflorescences thyrsoid, paniculate, with terminal or axillary, fewto several-flowered cymes; axes retrorsely aculeolate; bracts
lanceolate, 12 mm, subglabrous or hirtellous; pedicels 14
mm. Ovary ca. 0.7 mm, smooth. Corolla white or pale yellow,
rotate, glabrous or papillose, fused basal part ca. 0.5 mm; lobes
spreading, ovate-triangular or lanceolate-triangular, 22.5 mm,
acuminate. Mericarp berry black, ca. 4 mm in diam. Fl. JunJul,
fr. SepOct.
Thickets, roadsides; 26003100 m. NW Yunnan.
Rubia pallida belongs to R. ser. Cordifoliae. It is obviously close
to R. cordifolia but deviates by its larger flowers. See further comments
under that species.

22. Rubia podantha Diels, Notes Roy. Bot. Gard. Edinburgh 5:

277. 1912.
bing hua qian cao
Rubia nephrophylla Deb.
Plants herbaceous, perennial, erect or climbing, with
rootstock and rhizomes; stems to 1.2 m, quadrangular, glabrous
to strigose, with retrorsely aculeolate ribs or narrow wings.
Leaves in whorls of 4(or 6), sometimes unequal; petiole 15
cm; blade drying papery to subleathery, reddish abaxially or
brownish green, lanceolate, lanceolate-ovate, or oblong-ovate,
1.55 0.51.5 cm, length/breadth index 2.53.5, both surfaces
glabrous, strigillose, or strigose and sparsely to densely scaberulous, base truncate to cordate, margin serrulate-aculeolate
or ciliate, apex acute to acuminate; principal veins 3 or 5, palmate. Inflorescences thyrsoid, paniculate, with terminal and
axillary, many-flowered cymes, axes strigillose to glabrous;
aculeolate; bracts lanceolate, 15 mm; pedicels 1.54 mm.
Ovary ca. 0.8 mm, subglabrous. Corolla purplish red or yellowish white, campanulate, glabrous to sparsely hirsutulous;
fused part 0.81 mm; lobes ovate to lanceolate, 11.5 mm,
strongly reflexed, caudate to shortly acuminate. Mericarp berry
black at maturity, 45 mm in diam. Fl. AprJun, fr. JunSep.
Forest margins, sparse forests, grasslands; 7003000 m. W
Guangxi, W Sichuan, Yunnan.
The type specimens of Rubia podantha collected by Forrest from
the Lichiang Range, Yunnan, are described in the protologue as semiscandent, with leaves lanceolate, base cordate, reddish below, and somewhat campanulate purplish flowers. This corresponds well with the figure in H. S. Lo (in FRPS 71(2): 317, t. 71, f. 712. 1999) and with a
specimen collected from the type locality by Handel-Mazzetti in WU.
Thus, R. podantha belongs to R. ser. Cordifoliae and appears related to
other species with campanulate flowers, as R. alata or R. ovatifolia. It
shares the reddish hue with the always scandent R. manjith, which has
broader leaves and smaller, rotate flowers.

23. Rubia polyphlebia H. S. Lo, J. Trop. Subtrop. Bot. 7(1):

22. 1999.

315

duo mai qian cao

Herbs, perennial, with rootstock; stems erect and rather
stout, to 50 cm tall, quadrangular, sulcate, densely shortly hirsute to hispid. Leaves in whorls of 4, sessile; blade drying papery, broadly elliptic-oblong, ovate, or sometimes suborbicular,
24.5 13 cm, both surfaces densely hairy and scabrous,
base obtuse to subrounded, margins shortly aculeolate-ciliate,
apex acute to weakly obtuse; principal veins 711, palmate. Inflorescences thyrsoid, paniculate, with terminal and axillary,
many-flowered and to 10 cm long cymes; axes shortly hairy,
aculeolate to glabrescent; bracts leaflike or lanceolate to linearlanceolate, 12 mm; pedicels ca. 1 mm. Ovary 0.40.5 mm,
subglabrous. Corolla pale yellow, rotate, 2.53 mm in diam.,
hairy outside, glabrous or scabrous inside, fused basal part 0.2
0.3 mm; lobes ovate or lanceolate, 11.2 mm, apex recurved.
Immature mericarp berry 35 mm in diam.
Sichuan.
Rubia polyphlebia belongs to the R. mandersii group (see there) of
R. sect. Oligoneura. We have seen no material of this species, but there
is a good drawing in H. S. Lo (in FRPS 71(2): 304, t. 67, f. 17. 1999).

24. Rubia pseudogalium Ehrendorfer, Novon 20: 268. 2010.

gao li gong shan qian cao
Herbs, perennial, sprawling or twining; stems to 2 m,
branched, quadrangular, glabrous and smooth or sparsely retrorsely aculeolate. Leaves and leaflike stipules in whorls of
never more than 4; petiole (1)24(6) mm; blade when drying
blackening, subleathery, lower side somewhat brighter than
upper, linear-lanceolate, lanceolate, or lanceolate-oblong, often
somewhat falcate, 2040 28 mm, base cuneate to acute,
margins and vein adaxially somewhat retrorsely aculeolate or
smooth, apex acute or shortly acuminate; principal vein 1, 2
basal lateral veins very weak, often hardly visible. Inflorescences thyrsoid, with axillary, somewhat narrowly elongate,
many-flowered cymes; peduncles glabrous, 36 mm; bracts
small, linear-lanceolate, 0.53 mm; pedicels 16 mm. Ovary
inferior, obovoid, ca. 0.8 mm. Corolla yellowish green, white,
or purple, rotate, fused part 0.20.5 mm; lobes 5, ovate-triangular, 1.21.5 mm, shortly acuminate. Mericarp berry blackening, globose, 35 mm in diam. Fl. MayJun, fr. JulAug.
Subtropical montane evergreen broad-leaved forests; 2400
3000 m. Yunnan (Gaoligong Shan region).
This new species is strongly reminiscent of certain taxa of Galium
in habit. Its technical features (as well as DNA data) clearly place it into
Rubia. In spite of its narrow leaves with only 2 weak lateral veins, it
appears to belong to R. sect. Oligoneura. It forms an obviously related
group with R. truppeliana from the mountains of Shandong. Main differences are the leaf whorls never with more than 4 elements, the
shorter leaf petioles and peduncles, and the smaller flowers. There is
remarkable variation in leaf shape and flower color, as documented by
the numerous specimens available from the Gaoligong Shan region,
where R. pseudogalium may be endemic.

25. Rubia pterygocaulis H. S. Lo, J. Trop. Subtrop. Bot. 7(1):

22. 1999.
chi jing qian cao

RUBIACEAE

316

Herbs, perennials, with erect stems tufted from rootstock

and from subterranean rhizomes; lower stem portions quadrangular, broadly winged, upper with wings reduced, retrorsely
aculeolate. Leaves in whorls of 46; petiole 0.51 cm; blade
drying leathery, ovate, 12 0.51 cm, both surfaces sparsely
scaberulous to aculeolate, base cordate, margin revolute and
aculeolate, apex cuspidate-acuminate; principal veins 3, palmate, with lateral veins extending for ca. half of blade length.
Inflorescences thyrsoid, with terminal and axillary, several-flowered and 1.52.5 cm long cymes; axes retrorsely aculeolate with
reduced bracts; pedicels filiform, 12 mm. Ovary ca. 1 mm,
smooth. Corolla dried dark brownish, rotate, ca. 2.5 mm in
diam., glabrous; lobes lanceolate-ovate, 11.2 mm, cuspidate.
Mericarp berry unknown. Fl. Jun.
Forests, thickets; 3001000 m. Sichuan (Jiuzaigou).
No authentic material of Rubia pterygocaulis was available, but a
good drawing (including underground organs) is found in H. S. Lo (in
FRPS 71(2): 300, t. 65, f. 19. 1999). The local taxon evidently is a
somewhat xerophytic member of R. ser. Cordifoliae.

26. Rubia rezniczenkoana Litvinov, Trudy Bot. Muz. Imp.

Akad. Nauk 7: 75. 1910.
xiao ye qian cao
Subshrubs, with oblique rootstock; stems to 0.5 m,
clumped, quadrangular, glabrous, smooth. Leaves in whorls of
46(or 7), subsessile or sessile; blade drying stiffly papery,
elliptic-oblong, ligulate, oblanceolate, or lanceolate, 0.51.7
0.20.8 cm, glabrous, smooth, base obtuse to cuneate, margins
thickened and aculeolate, apex acute to obtuse with stiff short
cusp; principal vein 1. Inflorescences thyrsoid, with terminal
and axillary, several- to many-flowered cymes; axes glabrous
and smooth; bracts leaflike, 25 mm; pedicels ca. 1 mm. Ovary
ca. 0.8 mm, smooth. Corolla yellow, funnel-shaped, (2.5)3
3.5(4) mm; tube ca. 1.5 mm; lobes often only 4, oblong-lanceolate, ca. 2 mm, obtuse with short incurved cusp. Mericarp
berry black-purple when dry, 67 mm in diam. Fl. Jun, fr. Aug.
Sandy lands. Xinjiang [Kazakhstan, Mongolia].
Rubia rezniczenkoana is a rather isolated C Asiatic psammophyte,
for which Pojarkova (Fl. URSS 23: 397. 1958) has created the monotypic R. sect. Chonanthe, which we include provisionally into R. sect.
Rubia (see introduction to Rubia).

27. Rubia salicifolia H. S. Lo, J. Trop. Subtrop. Bot. 7(1): 23.

1999.
liu ye qian cao
Vines, herbaceous, or climbing herbs; stems to perhaps
1.5 m, quadrangular to 4-ridged, glabrous, ribs aculeolate.
Leaves in whorls of 4; petiole 312 cm, aculeolate, often pulvinate and strongly bent at base of blade; blade drying papery,
grayish brown, linear to narrowly lanceolate, 39 0.51 cm,
both surfaces glabrous, smooth or sparsely scaberulous, base
obtuse to rounded, margin flat and smooth to denticulate, apex
acute to acuminate; principal veins 3, palmate. Inflorescences
thyrsoid, paniculate, with terminal and axillary, few- to manyflowered cymes; axes glabrous, aculeolate; bracts linear-lanceolate, 0.53 mm; pedicels 34 mm. Ovary ca. 0.5 mm, scaberu-

lous. Corolla green, rotate, glabrous; fused basal part ca. 0.5
mm; lobes lanceolate-triangular, ca. 1 mm, cuspidate. Fruit
apparently black, 56 mm. Fl. JunJul, fr. Sep.
Moist valleys; ca. 2000 m. Guangdong, Sichuan (Jiulong),
?Yunnan.
This species belongs to Rubia ser. Cordifoliae and has been well
illustrated in H. S. Lo (in FRPS 71(2): 312, t. 70, f. 712. 1999). The
above description is based in part on provisionally identified specimens,
some of them from Guangdong. Affinities may exist with R. alata and
its transitional forms with R. cordifolia agg. (see additional comments
under these species).

28. Rubia schugnanica B. Fedtschenko ex Pojarkova in

Schischkin, Fl. URSS 23: 725. 1958.
si ye qian cao
Herbs, perennial, or subshrubs, with a rootstock; stems
numerous, erect, quadrangular, glabrous, smooth. Leaves in
whorls of 4(6), subsessile or sessile; blade drying subleathery,
narrowly lanceolate to linear-lanceolate, 2050 25 mm, both
surfaces glabrous, smooth, sessile or base narrowed into a very
short petiole, margins thickened and retrorsely aculeolate, apex
long acuminate; principal vein 1. Inflorescences thyrsoid, with
terminal and axillary, several-flowered cymes, shorter than or as
long as subtending leaves; peduncles glabrous, 310 mm; pedicels 25 mm; bracts none or linear, 13 mm. Ovary ca. 0.8 mm,
smooth. Corolla yellow, rotate; tube 0.20.7 mm; lobes lanceolate, 2.32.7 mm, acute to mucronate with cusp 0.40.6 mm.
Mericarp berry black, 34 mm in diam. Fl. Jul, fr. Aug.
Sandy lands; ca. 2500 m. Xinjiang [Tajikistan].
We have not seen authentic material of Rubia schugnanica. Pojarkova (loc. cit.) placed it into the small C Asiatic and mostly subshrubby
R. ser. Laxiflorae Pojarkova of R. sect. Campylanthera, which we provisionally include in R. sect. Rubia.

29. Rubia schumanniana E. Pritzel, Bot. Jahrb. Syst. 29: 583.

1901.
da ye qian cao
Rubia chinensis Regel & Maack var. esquirolii (H.
Lveill) H. Lveill; R. cordifolia Linnaeus var. maillardii (H.
Lveill & Vaniot) H. Lveill; R. esquirolii H. Lveill; R.
leiocaulis Diels; R. maillardii H. Lveill & Vaniot; R. schumanniana var. maillardii (H. Lveill & Vaniot) Handel-Mazzetti.
Herbs, perennial, erect (or rarely ?climbing), with rhizomatous base; stems to 1 m, quadrangular to subterete, sometimes shallowly ribbed and/or sulcate, glabrous or puberulent
near nodes, smooth or sometimes sparsely retrorsely aculeolate.
Leaves in whorls of 4; petiole equal to unequal, 0.51.5(3) cm;
blade drying thickly papery to subleathery, broadly lanceolate,
oblong-ovate, or ovate, 310 1.74 cm, length/breadth index
1.52.5, glabrous or usually hispidulous along principal veins
and scaberulous on lamina, base obtuse to rounded or cordulate,
margins thinly revolute and smooth to scaberulous, apex acuminate or subacute; principal veins 3 or 5, palmate, plane to impressed adaxially. Inflorescences thyrsoid-paniculate, cymes 5
12 cm, many flowered, terminal and from uppermost stem

RUBIACEAE

axils; peduncles scaberulous, puberulent, or glabrescent; bracteoles lanceolate, 34 mm, ciliate; pedicels 16 mm. Ovary ca.
1 mm, glabrous. Corolla white or greenish yellow, somewhat
campanulate, 45 mm in diam., outside glabrous, inside puberulent to scaberulous, fused base 0.81 mm; lobes lanceolate, 2
2.2 mm, acute to acuminate. Mericarp berry black, 57 mm in
diam. Fl. MayJul, Nov, fr. AugOct.
Forests; 8003000 m. Sichuan, Yunnan.
Rubia schumanniana (see H. S. Lo in FRPS 71(2): 301, t. 66, f. 1
7. 1999) belongs to R. ser. Chinenses, where it is close to R. chinensis
and R. latipetala, but also to taxa of the R. mandersii group with sessile
leaves. This plant is described as rarely climbing in H. S. Lo (loc. cit.:
299), but that may be due to misidentified specimens.

30. Rubia siamensis Craib, Bull. Misc. Inform. Kew 1911:

397. 1911.
dui ye qian cao
Vines, herbaceous, to 3 m; stems quadrangular, glabrous,
scaberulous to retrorsely aculeate along angles. Leaves opposite; petiole (1)24(8) cm, aculeolate; blade drying papery to
subleathery, ovate or ovate-lanceolate, 612 (1.5)36(7.5)
cm, glabrous on both surfaces, scaberulous and aculeolate on
principal veins and margins, base cordate or rounded, apex
acute or acuminate; principal veins 35(or 7), palmate, reticulate veinlets usually visible on both surfaces; stipules triangular, 35(7) 23 mm, persistent. Inflorescences thyrsoid,
paniculiform, with axillary, many-flowered cymes; bracts reduced. Ovary ca. 0.5 mm, glabrous. Corolla greenish, campanulate, ca. 3 mm in diam.; tube ca. 1.25 mm; lobes linear-lanceolate, ca. 2 mm, acute. Fruit of 2 subglobose mericarp berries,
1 sometimes aborted, each 45 mm in diam., shiny and black
when fresh. Fl. JunJul, fr. AugSep.
Evergreen moist forests; [900]22002500 m. SW Yunnan [N
Thailand].
Rubia siamensis has been illustrated by H. S. Lo (in FRPS 71(2):
297, t. 64, f. 69. 1999) and by Puff (Fl. Thailand: Rubiaceae; http://
homepage.univie.ac.at/christian.puff/FTH-RUB/FTH-RUB_HOME.
htm; accessed on 5 Oct 2010) who also gives an extended description.
Together with other tropical taxa from SW China and N Thailand (R.
magna, R. falciformis, and R. filiformis), all with opposite leaves and
true stipules, it forms a well characterized group within R. sect. Oligoneura, obviously not closely related to the polymorphic R. ser. Cordifoliae. Nevertheless, in the intensively studied Gaoligong Shan region
of Yunnan, where R. siamensis occurs sporadically, a remarkable series
of transitional specimens was documented, linking it with R. alata, a
member of R. ser. Cordifoliae (see under R. cordifolia).

31. Rubia sylvatica (Maximowicz) Nakai, J. Jap. Bot. 13: 783.

1937.
lin sheng qian cao
Rubia cordifolia Linnaeus var. sylvatica Maximowicz,
Mm. Acad. Imp. Sci. St.-Ptersbourg Divers Savans 9 [Prim.
Fl. Amur.]: 140. 1859.
Vines, herbaceous, perennial; stems to 3.5 m, quadrangular, glabrous, aculeolate on ribs. Leaves in whorls of 410(12);
petiole 211 cm, aculeolate; blade drying thinly membranous or
papery, brown-black or blackish green, ovate to suborbicular,
311 29 cm, length/breadth index 1.21.5, both surfaces

317

glabrous, scaberulous on lamina and aculeolate on principal

veins, base cordulate to cordate, margins aculeolate, apex acuminate to caudate-cuspidate; principal veins 5 or 7, palmate. Inflorescences thyrsoid, with terminal and axillary, several- to
many-flowered cymes; axes slender, glabrous, scaberulous;
bracts lacking or linear to lanceolate-linear, 15 mm; pedicels
17 mm. Ovary ca. 0.8 mm, smooth. Corolla greenish, rotate to
slightly patelliform, glabrous, fused basal part 0.40.6 mm;
lobes triangular, 11.5 mm, acuminate. Mericarp berry black,
510 mm in diam., with pedicels elongating, to 15 mm. Fl. Jul
Aug, fr. SepOct.
Moist forests or forest margins; 8003500 m. Throughout N
China, also in Sichuan [Russia].
Rubia sylvatica belongs to R. ser. Cordifoliae. It is apparently connected by transitional specimens with R. cordifolia s.s., R. ovatifolia,
and other closely related species, but quite well separated from the similar R. argyi. See these taxa for additional comments and the key for differential characters.
A critical taxon is Rubia hexaphylla (Makino) Makino (1927)
from Korea and Japan, of which we have not seen authentic specimens.
From its description (Yamazaki, Fl. Japan 3a: 232. 1993) one could
suspect it to be the same as R. sylvatica. If this is proven, the former
name has priority. The Kew Rubiaceae checklist (Govaerts et al., World
Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on
15 Sep 2010) treats R. sylvatica as a synonym of R. cordifolia subsp.
cordifolia but maintains R. hexaphylla.

32. Rubia tenuis H. S. Lo, J. Trop. Subtrop. Bot. 7(1): 24.

1999.
xian geng qian cao
Vines, herbaceous, perennial; stems quadrangular, retrorsely aculeolate on ribs. Leaves in whorls of 4, sessile; blade
drying thinly papery, broadly elliptic, 1.54 0.92.3 cm, both
surfaces subsmooth or scaberulous to aculeolate on principal
veins, base rounded to obtuse, margin sparsely aculeolate, apex
cuspidate; principal veins 5, palmate. Inflorescences thyrsoid,
with terminal and at lower stem nodes axillary, many-flowered
cymes to 14 cm; peduncles slender; bracteoles lanceolate, 2.54
mm. Corolla yellow, shallowly campanulate to subrotate, 45
mm in diam.; lobes lanceolate, ca. 2 mm, apex incurved, acuminate. Fruit unknown. Fl. Jul.
Forests, thickets. Sichuan (Luding).
We have not seen authentic material of Rubia tenuis, but there is a
good drawing in H. S. Lo (in FRPS 71(2): 310, t. 69, f. 14. 1999). With
respect to habit, indumentum, sessile leaves, inflorescences, and flowers, affinities are suggested with R. edgeworthii (see there) and R. sikkimensis Kurz. This latter species, well described and figured by Deb
and Malick (Bull. Bot. Surv. India 10(1): 12. 1968), is distributed from
NE India to Bhutan, but may also reach adjacent China. It mainly differs by larger leaves (513 25 cm) with a rather cuneate base and
more acuminate leaf apex. The three species can be assembled provisionally in a R. sikkimensis group within R. sect. Oligoneura.

33. Rubia tibetica J. D. Hooker, Fl. Brit. India 3: 204. 1881.

xi zang qian cao
Herbs, perennial, erect, or subshrubs, forming loose cushions from a massive woody rootstock; stems to 0.3 m, quadran-

318

RUBIACEAE

gular, glabrous or mostly with dense hirtellous indumentum,

on angles shortly aculeolate and hispidulous with partly hooked
trichomes, lower nodes usually shortly sheathed with membranous bases of old leaves. Leaves opposite and with interpetiolar
and leaflike stipules in whorls of 4(6), sessile or subsessile;
blade drying leathery, broadly to narrowly ovate, elliptic, elliptic-oblong, or lanceolate, 13 0.31.5 cm, both surfaces hirtellous to glabrous, base acute to obtuse, margins retrorsely
aculeolate, toward acute and often cuspidate apex usually antrorsely aculeolate; principal vein 1, sometimes with 2 weak
lateral veins. Inflorescences leafy and bracteose, with axillary
and terminal, 1- to few-flowered cymes; axes mostly glabrous;
pedicels (2)46(14) mm. Ovary 0.81.2 mm, sometimes hirtellous. Corolla yellow, rotate, 58 mm in diam., outside sometimes scaberulous; fused base ca. 0.5 mm; lobes lanceolate or
lanceolate-ovate, 22.5 mm, acuminate. Mericarp berry 34
mm in diam. Fl. Jun, fr. Aug.

late to subsessile; blade drying papery to subleathery, lanceolate, lanceolate-oblong, or elliptic-oblong, 310 0.53.5 cm,
glabrous or mostly along lower midrib and margins retrorsely
aculeolate, base acute, apex acute; lateral veins 3 or 4 pairs,
pinnate. Inflorescences thyrsoid, leaflike and many-flowered
cymes terminal and axillary from upper stem nodes; axes
retrorsely aculeolate; peduncles up to 50 mm with bracts narrowly elliptic, 25 mm; pedicels (0.75)1.58(12) mm. Ovary
ca. 0.8 mm, glabrous. Corolla yellow to greenish yellow, rotatefunnelform, glabrous; tube ca. 1 mm; limb 34.5 mm in diam.;
lobes lanceolate, ca. 1 mm, shortly acuminate. Anthers large,
0.50.6(0.8) mm, straight. Mericarp berry black, 3.54 4
4.5 mm. Fl. JunAug, fr. JulSep.

In gravel at river bottoms; [1700]3600[4400] m. Xinjiang, Xizang [Afghanistan, India (Punjab), Kashmir, Kyrgyzstan, Pakistan, Tajikistan].

Rubia tinctorum is the madder of commerce, cultivated for the

dye derived from its rhizomes and roots, and still used in textiles and
fine paints. Mainly because of its relatively large and straight anthers,
this and a few related taxa from C Asia have been placed into R. sect.
Meganthera by Pojarkova (Fl. URSS 23: 392397. 1958). As R. tinctorum is the type species of the genus, this section has to be called R.
sect. Rubia (Ehrendorfer et al., Fl. Iranica 176: 54. 2005).

Rubia tibetica is a relatively widespread SW to C Asiatic and typically montane to alpine pioneer species. It was illustrated by Deb and
Malick (Bull. Bot. Surv. India 10(1): 4, f. 3. 1968), who lectotypified its
name with a specimen from Tibet (i.e., Xinjiang). The species is notable for its shoot morphology: the well-developed vegetative leaves of
lower stem nodes often appear in a 4-verticillate arrangement with
leaflike stipules, whereas the basalmost first leaves produced by the
stems and also the leaves of reproductive nodes are generally paired and
exhibit typical interpetiolar stipules.
Deb and Malick (loc. cit.: 45) described Rubia aitchisonii Deb &
Malick from Bagdis, Afghanistan, and separated it by: Lamina ovate,
sub-orbicular, elliptic-ovate or lanceolate, 26 in a whorl in R. tibetica
vs. Lamina elliptic-lanceolate, 2 opposite in R. aitchisonii. Ehrendorfer and Schnbeck-Temesy (Fl. Iranica 176: 67. 2005) noted that the
only locality of R. aitchisonii lies within the area of R. tibetica and that
the suspected specific differences fall within the morphological variability of R. tibetica. Thus, future studies may show that R. aitchisonii is
better synonymized under R. tibetica.
Rubia tibetica was placed by Pojarkova (Fl. URSS 23: 401404.
1958) into R. ser. Tibeticae Pojarkova in R. sect. Campylanthera together with two other alpine, C Asiatic species: R. regelii Pojarkova and
R. komarovii Pojarkova. They differ from R. tibetica by leaves and leaflike stipules in whorls of up to 6 but have not been recorded yet from
N China. Rubia garrettii Craib from Thailand, also suspected to be a
member of this group, certainly does not belong here (see Puff, Fl.
Thailand: Rubiaceae; http://homepage.univie.ac.at/christian.puff/FTHRUB/FTH-RUB_HOME.htm; accessed on 5 Oct 2010). Whereas the R.
tibetica group is provisionally included in R. sect. Rubia, R. garrettii
obviously belongs to R. sect. Oligoneura.

34. Rubia tinctorum Linnaeus, Sp. Pl. 1: 109. 1753.

ran se qian cao
Rubia iberica (Fischer ex Candolle) K. Koch; R. tinctorum
var. iberica Fischer ex Candolle.
Herbs, sprawling to climbing, perennial, with extensive
stout, woody, and red rhizomes; stems to 12.5 m, often fascicled, quadrangular with sharp angles, somewhat retrorsely
aculeolate or glabrous. Leaves in whorls of 46, shortly petio-

Rather dry open ground; 4002300 m. Xinjiang [Afghanistan,

NW India, Kashmir, Kazakhstan, Pakistan, Turkmenistan; SW Asia
(Iran, Turkey); widely cultivated and escaped or weedy from Europe
and the Mediterranean throughout the world].

35. Rubia trichocarpa H. S. Lo, J. Trop. Subtrop. Bot. 7(1):

23. 1999.
mao guo qian cao
Vines, apparently climbing; stems quadrangular, hirsute at
least at nodes, retrorsely aculeolate along ribs. Leaves in whorls
of 4(6); petiole slightly shorter than blade, retrorsely aculeate;
blade drying thinly papery, grayish green, ovate to cordate, 4
12 2.57 cm, adaxially glabrous and scaberulous, abaxially
sparsely strigillose, hirsutulous, or villosulous, base deeply cordate, margins densely ciliate, apex acuminate; principal veins
35, palmate. Inflorescences thyrsoid, paniculate, axillary and
perhaps also terminal cymes many flowered and longer than
subtending leaves; axes slender, villosulous and retrorsely aculeolate; bracteoles lanceolate, 1.52 mm. Flowers not seen. Immature mericarp berry black and shiny when dry, densely ferruginous villous.
Sichuan (Lixian).
Rubia trichocarpa evidently belongs to R. ser. Cordifoliae, but no
authentic material was available to us. A specimen from Xizang, collected 17 Jul 1973 (PE), fits the above description, with its short hispidulous indumentum extending from stems and leaves to the inflorescence, flowers, and young fruit. It has cymes rather condensed; flowers
sessile or on pedicels up to 1.2 mm; and corollas dried purple, ca. 2 mm
in diam., campanulate with reflexed petals of ca. 0.8 mm.

36. Rubia truppeliana Loesener, Beih. Bot. Centralbl., Abt. 2,

37: 183. 1920.
shan dong qian cao
Herbs, perennial, sprawling or twining, to 2 m; stems
branched, quadrangular, glabrous or sparsely puberulent, retrorsely aculeolate on angles, sometimes sulcate-striate. Leaves
in whorls of up to 6 or 8; petiole 635 mm, aculeolate; blade
drying dark green, subleathery, oblanceolate, broadest toward

RUBIACEAE

base, 2035 46 mm, scaberulous to retrorsely aculeolate on

margins and vein on lower or on both sides, base cuneate to
acute, apex acute or shortly acuminate; principal vein 1, 2 lateral veins weaker and hardly reaching half of leaf length. Inflorescences thyrsoid, with terminal and axillary congested to subcapitate cymes; axes aculeolate; bracts lanceolate or linear-lanceolate, 0.53 mm; peduncles 1040 mm; pedicels 0.54 mm.
Ovary ca. 0.8 mm, smooth. Corolla rotate, glabrous, color not
recorded; fused basal part ca. 0.4 mm; lobes ovate-triangular,
ca. 2 mm, acuminate. Mericarp berry not seen. Fl. JulAug.
Forests, thickets; 100300 m. Shandong.
This local species is certainly close to the newly described Rubia
pseudogalium from the Gaoligong Shan region of Yunnan but clearly
separable (see there). In habit both are quite similar to Galium. In spite
of their very narrow leaves and only slightly palmate veins, this R. truppeliana group probably should be placed into R. sect. Oligoneura.

37. Rubia wallichiana Decaisne, Nouv. Mm. Acad. Roy. Sci.

Bruxelles 12: 61. 1837.
duo hua qian cao
Vines or climbing herbaceous perennials; stems to 3 m, 4angled, glabrous or puberulent at nodes, smooth or sparsely
aculeolate. Leaves in whorls of 4(6); petiole 0.66 cm; blade
drying thinly papery, lanceolate or ovate-lanceolate, 27 0.5
2.5 cm, glabrous and sparsely scaberulous, base rounded, truncate, or sometimes cordulate, margins denticulate-aculeolate to
smooth, apex acuminate; principal veins 3(or 5), palmate. Inflorescences thyrsoid, paniculate, with terminal and axillary, manyflowered cymes; axes glabrous, smooth to sparsely scaberulous;
bracts lanceolate to ligulate, 13.5 mm; pedicels 14 mm.
Ovary ca. 0.5 mm, smooth. Corolla purplish red, greenish yellow, or white, rotate, fused basal part 0.10.5 mm; lobes lanceolate to lanceolate-triangular, 1.31.5 mm, acuminate, glabrous.
Mericarp berry black, 3.54 mm in diam. Fl. AugOct, fr. Aug
Dec.
Forests, forest margins, thickets, open fields, village fences; 300
2600 m. Guangdong, Guangxi, Hainan, Hunan, Jiangxi, Sichuan, Yunnan [Bhutan, NE India, Nepal].
The above description and distribution data of Rubia wallichiana
have been taken over from H. S. Lo (in FRPS 71(2): 315316. 1999).
There are hardly any differential characters relative to R. cordifolia s.s.
aside from vague references to less prickly stems or differences in flower and fruit color (see key). Thus, this dubious taxon clearly belongs to

319

R. ser. Cordifoliae. Deb and Malick (Bull. Bot. Surv. India 10(1): 116.
1968) do not even mention R. wallichiana. In Fl. Bhutan (2(2): 823
825. 1999) the name is used in a wide sense and evidently includes what
is here treated as R. cordifolia, R. sylvatica, and possibly even R. argyi.
The Kew Rubiaceae checklist (Govaerts et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on 15 Sep 2010)
accepts R. wallichiana as a distinct species. We have hardly seen specimens from China that clearly correspond to R. wallichiana (and not to
other taxa of R. cordifolia agg.). In view of all this, we regard R. wallichiana as a possible synonym of R. cordifolia s.s. but maintain it as a
species in the present flora in order to stimulate its clarification.

38. Rubia yunnanensis Diels, Notes Roy. Bot. Gard. Edinburgh 5: 278. 1912.
zi shen
Rubia ustulata Diels.
Herbs, perennial, with rootstock and somewhat thickened
storage roots; stems usually clumped, suberect, to 0.5 m, quadrangular or narrowly 4-winged, hirsute at nodes to glabrescent,
smooth or rarely scabrid. Leaves in whorls of 4(6), subsessile;
blade drying papery, lanceolate, ovate, obovate, elliptic-oblong,
broadly elliptic, or suborbicular, 14(5) 0.32 cm, both surfaces hairy to scabrid, base cuneate to rounded, margins flat or
often revolute, apex acuminate, shortly cuspidate, or acute;
principal veins 3(or 5), palmate. Inflorescences thyrsoid, paniculate, terminal and axillary cymes usually longer than subtending leaves; axes subglabrous to sparsely hirsutulous; bracteoles lanceolate, 25 mm; pedicels 13 mm. Ovary 0.30.4 mm
in diam., glabrous. Corolla yellow or pale yellow, rotate, ca. 3
mm in diam., glabrous; fused basal part ca. 0.5 mm; lobes subovate, 1.21.5(2) mm, apex thickened, incurved, shortly rostrate. Mericarp berries not seen.
Thickets, grassy slopes, roadsides; 17003000 m. Sichuan,
Yunnan.
In his publication of Rubia yunnanensis Diels referred to an
unpublished herbarium name R. sikkimensis var. yunnanensis Franchet and commented that it is quite different from R. sikkimensis.
Rubia ustulata was published on the same page immediately after R.
yunnanensis and said to differ by its smaller size, smaller and more
equal leaves and leaflike stipules, and more cuspidate petals. In view of
the variability of these characters, its synonymization by H. S. Lo (in
FRPS 71(2): 303. 1999) is accepted. The species evidently has a considerable altitudinal range and consequently varies from quite elongated to considerably condensed. Within R. sect. Oligoneura, R. yunnanensis belongs to the R. mandersii group of SW China.

80. RUBOVIETNAMIA Tirvengadum, Biogeographica (Paris) 74(4): 166. 1998.

yue nan qian shu
Chen Tao (); Charlotte M. Taylor
Shrubs, unarmed, sometimes with some internodes markedly shortened. Raphides absent. Leaves opposite, sometimes appearing
ternate due to reduced internode and marked anisophylly grouping 2 leaves at one node plus an apparently solitary leaf at another
node, usually with domatia, at apex sometimes aristate; stipules caducous to late deciduous, shortly united around stem, triangular,
inside (i.e., adaxially) pubescent, sometimes aristate. Inflorescences terminal and/or pseudoaxillary, cymose and 28-flowered or
sometimes reduced to 1 flower, pedunculate, bracteate with bracts often stipuliform. Flowers pedunculate or pedicellate, bisexual,
monomorphic. Calyx limb deeply 5-lobed; lobes sometimes imbricate in bud. Corolla white turning yellow with age, salverform to
funnelform, inside glabrous except with a pilosulous ring near base; lobes 5, convolute in bud. Stamens 5, inserted near corolla
throat, partially exserted; filaments reduced; anthers dorsifixed near base, becoming twisted with age, shortly sagittate at base, with

RUBIACEAE

320

connective prolonged in an apical triangular appendage. Ovary 1-celled, ovules 24 on parietal placentas; stigmas 2, spatulate,
partially exserted. Fruit baccate, fleshy to leathery, subglobose, smooth, mature color unknown, with calyx limb tardily deciduous;
seeds few to several, medium-sized, ovoid to subglobose, smooth; endosperm abundant; embryo minute, basal, nail-shaped; radicle
slightly longer than cotyledons; cotyledons suborbicular.
Two species: China, Vietnam; two species (one endemic) in China.
The protologue figure is labeled Vidalasia aristata, but this species was treated in the text in Rubovietnamia; Vidalasia Tirvengadum was also
described in this same article, and presumably R. aristata was to be included in that genus initially. Vidalasia has several-flowered compound cymes,
erose-setose stipules and bracts, numerous ovules and seeds, and probably may be expected in China also.

1a. Leaf blade elliptic or oblanceolate, 412 1.54 cm, adaxially glabrescent to sparsely strigillose, abaxially
sparsely strigillose, apex acuminate and aristate; stipules on each side deltoid to ovate, caudate-acuminate;
corolla tube cylindrical, lobes glabrous inside, hairy outside; fruit strigillose to glabrescent ....................................... 1. R. aristata
1b. Leaf blade obovate, 818 35 cm, densely pubescent on both surfaces, apex acuminate; stipules terminating
with reduced interpetiolar limbs; corolla tube funnelform, lobes mostly glabrous inside, pubescent outside;
fruit tomentose ..................................................................................................................................................... 2. R. nonggangensis
1. Rubovietnamia aristata Tirvengadum, Biogeographica
(Paris) 74(4): 167. 1998.
chang guan yue nan qian
Shrubs or small trees, to 5 m tall; branches compressed to
terete, densely strigillose or hirtellous to glabrous. Leaves in
equal to somewhat unequal pairs except in penultimate pair of
each stem 1 leaf reduced to a stipulelike scale; petiole 0.41
cm, strigillose to glabrescent; blade drying thickly papery, elliptic or oblanceolate, 412 1.54 cm, adaxially glabrescent to
sparsely strigillose, abaxially sparsely strigillose throughout,
base cuneate and sometimes asymmetrical, apex acuminate and
aristate with tip 12 mm, formed by prolongation of midrib and
inserted 23 mm below apex from tip of blade tissue; secondary
veins 7 or 8 pairs, with pilosulous domatia in abaxial axils; stipules deciduous throughout or persistent on distalmost several
nodes, deltoid to ovate, 56 mm, sparsely strigillose, caudateacuminate, entire to shortly aristate. Inflorescences 14-flowered, densely strigillose; bracts triangular to 3-lobed, 34 mm;
pedicel and/or peduncle 1525 mm. Calyx densely strigillose to
villosulous; ovary portion ellipsoid to obovoid, 45 mm; limb
lobed essentially to base; lobes ovate to lanceolate, 710 35
mm, often somewhat unequal on an individual flower, ciliolate,
aristate. Corolla salverform, outside densely sericeous-strigose;
tube 2025 mm; lobes obovate to oblanceolate, 1015 36
mm, acute or shortly acuminate. Anthers ca. 10 mm, with triangular appendage ca. 0.5 mm. Stigmas ca. 5 mm. Berry 11.7
cm in diam., strigillose to glabrescent; seeds 57 mm. Fl. May
Jul, fr. Jul.
Thickets or forests on limestone hills; 2001400 m. Guangxi,
Yunnan [Vietnam].

2. Rubovietnamia nonggangensis F. J. Mou & D. X. Zhang,

Bot. Stud. (Taipei) 51: 123. 2010.
nong gang yue nan qian

Shrubs or trees, to 3 m tall, without spines; young

branches covered with hairs and densely hirsute under stipules.
Petiole 0.51 cm; leaves thickly papery, obovate, 818 35
cm, densely pubescent on both surfaces, base cuneate, apex
acuminate; main and lateral nerves visible adaxially, very
prominent abaxially, secondary veins 714 pairs; stipules ca. 5
mm, papery, outside coated with thick hairs, terminating with 2
minute interpetiolar limbs. Inflorescences 28-flowered cymes,
superaxillary on lateral branches, with both bracts and bracteoles triangular, 34 mm; pedicels to 25 mm in open flowers.
Flowers 5-merous, very fragrant. Calyx tube campanulate, to 8
mm, hispid, widening at apex with enlarged (well-developed)
lobes; lobes elliptic-lanceolate, to 10 5 mm, imbricate,
markedly veined, coated with dense hairs, apiculate, deciduous.
Corolla tubular, glossy, fleshy, waxy, cream white at anthesis,
turning yellow with age and without spots in tube; tube dilating
gradually to throat, to 35 mm, very narrowly tubular in lower
part and convex in middle, ca. 5 mm wide at middle, outside
hairy, inside largely glabrous except for a zone 11.5 cm above
base covered with hairs; lobes obovate to lanceolate, to 18 5.5
mm, contorted to left in bud stage, pubescent outside, mostly
glabrous inside, reflexed at anthesis. Stamens 5, inserted just at
corolla throat and between corolla lobes, sessile; anthers linear,
to 5 mm, adnate, 1/2 of length exserted medifixed, exserted
after anthesis. Ovary inferior, with 2 parietal placentas, numerous ovules each in 2 rows on an oblong-elliptic placenta fusing
to form a compact mass when mature; style 3.54 cm, with
glabrous columnar basal part and pubescent at 2 cm above base;
stigma ca. 4.5 3 mm, with 2 lobes initially cohering together,
fluted, wholly exserted. Floral disk annular. Berry to 2 1.5
cm, tomentose, crowned by remnants of calyx and a large yellow disk, lacking ribs; petiole to 2.5 cm; seeds to 6.
Rocky crevices in moist forests of limestone areas; 200400 m.
SW Guangxi [?Vietnam].

81. SAPROSMA Blume, Bijdr. 956. 18261827.

ran mu shu shu
Chen Tao (); Charlotte M. Taylor
Erect shrubs, unarmed, usually fetid when bruised. Raphides present. Leaves opposite or in whorls of 3 or 4, sometimes with
domatia; stipules caducous or persistent, interpetiolar, triangular, entire or 13-setose or multifid. Inflorescences axillary or terminal,

RUBIACEAE

321

fasciculate or cymose and several flowered or reduced to 1 flower, pedunculate to sessile, bracteate with bracts usually fused in pairs.
Flowers sessile or pedicellate, bisexual, apparently monomorphic. Calyx limb 46-lobed, sometimes funnelform, lobes sometimes
unequal. Corolla white, campanulate, tubular, or funnelform, inside villous in throat; lobes 4(6), valvate-induplicate in bud, with
margins sometimes crisped. Stamens 4(6), inserted in corolla throat, usually partially exserted; filaments short or reduced; anthers
dorsifixed near base. Ovary 2-celled, ovules 1 in each cell, basal, erect; style 2-lobed, included or exserted. Fruit blue or purplish
black, drupaceous, fleshy, ellipsoid, with calyx limb persistent; pyrenes 1 or 2, 1-celled, with 1 seed, crustaceous to papery; seeds
medium-sized, plano-convex, subobovoid, or ellipsoid, abaxially (i.e., dorsally) smooth to verrucose, endosperm fleshy; cotyledon tiny, leaflike; radicle slender, close to hilum.
About 30 species: tropical Asia; five species (four endemic) in China.
Saprosma is not well known. Analyses of relationships among Lasianthus and related genera support the transfer of S. crassipes into that genus,
although the authors did not publish a formal nomenclatural transfer (Xiao & Zhu, Bot. Stud. (Taipei) 48: 227232. 2007).

1a. Flowers sessile in sessile glomerules or heads.

2a. Flowers mostly in terminal heads, sometimes also borne in uppermost leaf axils; petioles 13 mm .............. 2. S. hainanensis
2b. Flowers in glomerules borne along much of length of stem, sometimes also in a terminal head; petioles
1012 mm ................................................................................................................................................................. 4. S. merrillii
1b. Flowers sessile to pedicellate in pedunculate and/or branched inflorescences.
3a. Flowers on pedicels 210 mm, in branched cymes borne on peduncles 1040 mm; leaves opposite or more
often ternate; pyrenes smooth abaxially ..................................................................................................................... 5. S. ternata
3b. Flowers sessile in heads, borne on peduncles 520 mm; leaves opposite; pyrenes verrucose abaxially.
4a. Peduncles 58 mm; secondary leaf veins 710 pairs; corolla tube 78 mm .................................................. 1. S. crassipes
4b. Peduncles 1020 mm; secondary leaf veins 46 pairs; corolla tube ca. 10 mm .................................................. 3. S. henryi
1. Saprosma crassipes H. S. Lo, Bot. J. S. China 2: 15. 1993.
hou geng ran mu shu
Shrubs 12 m tall or occasionally small trees to 5 m tall;
branches weakly flattened to subterete, glabrous. Leaves opposite; petiole 310 mm, glabrous to sparsely strigillose; blade
drying thinly leathery, dark to pale green, elliptic-ovate or oblong-ovate, 816 ca. 6 cm, both surfaces glabrous, base obtuse or rounded, apex abruptly acuminate; secondary veins 7
10 pairs, joining in a submarginal vein, without domatia; stipules caducous, triangular, ca. 1.5 mm, strigillose to glabrous,
acute. Inflorescences axillary and sometimes also terminal, capitate, several flowered, strigillose to glabrous; peduncle compressed, 58 mm; bracts ovate, 12 mm; pedicels to 2 mm.
Flowers subsessile to pedicellate. Calyx with hypanthium portion cupuliform, ca. 1 mm, glabrous; limb 0.81 mm, shallowly
lobed, strigillose to glabrous; lobes broadly triangular, ciliate.
Corolla tubular-funnelform, glabrous outside, villous inside;
tube 78 mm; lobes broadly ovate, ca. 2 mm. Drupes ellipsoid,
78 mm; pyrenes hemispherical, verrucose abaxially. Fl. May,
fr. MayOct.
Forest margins; 3001300 m. Hainan, Yunnan (Xishuangbanna)
[?Vietnam].
A recent analysis of molecular data suggested that this species
may be better placed in Lasianthus, though the authors did not make a
formal nomenclatural transfer (Xiao & Zhu, Bot. Stud. (Taipei) 48:
227232. 2007).

2. Saprosma hainanensis Merrill, Lingnan Sci. J. 9: 44. 1930

[hainanense].
hai nan ran mu shu
Erect shrubs, 12 m tall; branches weakly flattened to terete, glabrous, with epidermis often becoming hardened then
fragmenting. Leaves opposite or sometimes appearing whorled
due to congested nodes with internodes not expanded, sometimes in unequal pairs; petiole 13 mm, glabrous; blade drying

membranous to papery, oblong-ovate to oblong-elliptic, 3.515

1.55.5 cm, adaxially glabrous, abaxially glabrous or hirtellous on principal veins, base rounded to cuneate, apex acute or
shortly acuminate; secondary veins 712 pairs, usually joining
in a submarginal vein, usually with pilosulous or foveolate
domatia; stipules caducous, ovate, 23 mm, glabrous, aristate
with bristle 13 mm. Inflorescences terminal and in uppermost
leaf axils, glomerulate or capitate, few flowered, glabrous;
bracts ovate, usually fused in pairs, 12 mm. Flowers sessile.
Calyx glabrous; hypanthium portion obconic, 12 mm; limb 1
2 mm, lobed for ca. 1/2; lobes 4(6), triangular. Corolla tubular,
glabrous or puberulent outside; tube ca. 3 mm, in throat villous;
lobes 4, elliptic to elliptic-ovate, ca. 2.5 mm. Drupes obovoid,
67 34 mm; pyrenes plano-convex, smooth abaxially. Fl.
Jun, fr. OctNov.
Ravines; 3001700 m. Hainan.
H. S. Lo (in FRPS 71(2): 67. 1999) described the calyx lobes as 5
or 6, but they are consistently 4 on the specimens studied.

3. Saprosma henryi Hutchinson in Sargent, Pl. Wilson. 3: 417.

1916.
yun nan ran mu shu
Shrubs, ca. 3 m tall; branches glabrous. Leaves opposite;
petiole 37 mm, glabrous; blade drying thinly papery, elliptic or
oblong-elliptic, 69 35 cm, both surfaces glabrous, base obtuse, apex shortly acuminate; secondary veins 46 pairs, joining
in a submarginal vein; stipules triangular, 1.52 mm. Inflorescence capitate, few flowered; peduncle 1020 mm, longitudinally ridged. Flowers sessile. Calyx with hypanthium portion
glabrous; lobes triangular, ca. 1.25 mm, subglabrous. Corolla
tubular-funnelform, glabrous outside; tube ca. 10 mm, pilose
inside; lobes ovate-triangular, ca. 2 mm, obtuse. Drupes oblate,
ca. 8 mm; pyrenes verrucose abaxially.
Mountain forests; 13001700 m. Yunnan.

RUBIACEAE

322

4. Saprosma merrillii H. S. Lo, Bot. J. S. China 2: 15. 1993.

qiong dao ran mu shu
Lasianthus hainanensis Merrill, Philipp. J. Sci. 21: 355.
1922, not Saprosma hainanensis Merrill (1930).
Shrubs, 23 m tall; branchlets terete, strigose or hirtellous
to glabrous. Leaves opposite; petiole 1012 mm, hirtellous or
strigillose; blade drying stiffly papery, elliptic-oblong to lanceolate, 1017 3.56 cm, adaxially glabrous, abaxially glabrous
or usually hirtellous along midrib, base cuneate to truncate,
apex acuminate; secondary veins 14 or 15 pairs, joining in a
submarginal vein, without domatia; stipules persistent, narrowly
triangular, 23 mm, strigillose or hirtellous to glabrescent, acute.
Inflorescences axillary along stem and sometimes also terminal,
glomerulate or capitate, sessile, several flowered, strigillose or
hirtellous; bracts apparently reduced. Flowers sessile. Calyx
with hypanthium portion obconic, ca. 1 mm, pilosulous to glabrous; limb ca. 1 mm, deeply lobed; lobes triangular. Corolla
tubular-funnelform, glabrous outside, villous in throat and on
lobes inside; tube 44.5 mm; lobes ovate, ca. 1.5 mm. Drupes
obovate to oblong-obovate, 78(12) mm; pyrenes plano-convex, smooth abaxially. Fl. Apr.
Sparse forests or forest margins; 3001000 m. Hainan.

5. Saprosma ternata (Wallich) J. D. Hooker, Fl. Brit. India 3:

193. 1881 [ternatum].

ran mu shu
Paederia ternata Wallich in Roxburgh, Fl. Ind. 2: 520.
1824; Serissa ternata (Wallich) Kurz.
Erect shrubs 14 m tall, or sometimes small trees 56 m
tall; branches angled, glabrous. Leaves in whorls of 3 or sometimes opposite; petiole 612 mm, glabrous; blade drying thinly
leathery to papery, elliptic, oblong-lanceolate, or oblong-elliptic, 815 36.5 cm, both surfaces glabrous, base acute to obtuse, apex shortly acuminate; secondary veins 710 pairs, not
joining in a submarginal vein, without domatia; stipules caducous, narrowly triangular to lanceolate, 512 mm, with 27 linear teeth or projections. Inflorescences axillary, cymose, glabrous; peduncles 13 per axil, 1040 mm; bracts triangular to
ovate, 15 mm, often aristate, usually deciduous after flowering; pedicels 210 mm. Flowers pedicellate. Calyx glabrous;
hypanthium portion cupuliform, 22.5 mm, glabrous; limb 12
mm, shallowly and sometimes irregularly lobed; lobes triangular. Corolla tubular, puberulent to tomentulose outside; tube 38
mm, pilose in throat; lobes triangular usually with expanded
crisped margins, 34 mm. Drupes ellipsoid or subglobose, 812
68 mm; pyrenes plano-convex, smooth abaxially. Fl. Apr,
Jun, fr. SepNov.
Sparse forests at low to middle elevations, forests in ravines; 400
1000 m. Hainan, Yunnan [NE India, Malaysia].

82. SCHIZOMUSSAENDA H. L. Li, J. Arnold Arbor. 24: 99. 1943.

lie guo jin hua shu
Chen Tao (); Charlotte M. Taylor
Shrubs to small trees, unarmed. Raphides presumably absent. Leaves opposite, apparently without domatia or these infrequent
and rudimentary; stipules persistent at least near stem apex, interpetiolar, generally triangular, 2-lobed. Inflorescences terminal and
sometimes also in uppermost leaf axils and appearing sessile and tripartite, cymose with higher order axes often scorpioid, many
flowered, pedunculate, bracteate with bracts persistent or caducous. Flowers sessile or subsessile, bisexual, distylous. Calyx limb
deeply 5-lobed, on 1 or several flowers of an inflorescence with 1 white, elliptic or ovate calycophyll. Corolla orange-yellow to
orange, salverform with tube slender then abruptly swollen in upper part, inside pubescent at least in throat; lobes 5, notably acuminate, in bud valvate-induplicate with tips pressed together and ascending. Stamens 5, inserted in upper part of corolla tube, included in long-styled form, partially exserted in short-styled form; filaments short; anthers apparently basifixed. Ovary 2-celled,
ovules numerous in each cell on axile peltate placentas; stigmas 2, linear, exserted in long-styled form or included in short-styled
form. Fruit capsular, ellipsoid to obovoid, slightly flattened perpendicular to septum, loculicidally dehiscent through apical half,
woody, with calyx limb persistent, with calycophylls persistent; seeds numerous, small, angled.
One species: SW China, Laos, N Myanmar, N Thailand, N Vietnam.
This species was long included in Mussaenda, but morphological and molecular studies support its separation from that genus. Its morphology
was studied by Puff et al. (Bull. Jard. Bot. Natl. Belg. 62: 3568. 1993). Its lack of raphides has not been described but is presumed based on the
classification of this genus by various authors in Isertieae.

1. Schizomussaenda dehiscens (Craib) H. L. Li, J. Arnold

Arbor. 24: 100. 1943.
lie guo jin hua
Mussaenda dehiscens Craib, Bull. Misc. Inform. Kew
1916: 263. 1916; Emmenopterys rehderi F. P. Metcalf; M.
elongata Hutchinson; M. henryi Hutchinson; Schizophragma
macrosepalum Hu.
Shrubs to small trees, to 8 m tall; branches flattened or
angled to terete, strigose to strigillose becoming glabrescent

with age, often with sparse to rather dense ellipsoid lenticels.

Petiole 0.51.6 cm, strigose to strigillose; leaf blade drying
thinly papery, lanceolate, lanceolate-elliptic, or ovate-lanceolate, 1017 2.56 cm, adaxially sparsely hirsute or strigose at
least along principal veins to glabrescent, abaxially strigose to
strigillose along principal lateral veins, sparsely strigose along
higher order veins, and glabrescent on blade, base cuneate to
rounded, apex acuminate or acute; secondary veins 710 pairs,
infrequently with a few pilosulous domatia; stipules 514 mm,
abaxially strigose to strigillose, 2-lobed for 1/4 to nearly completely. Inflorescences densely hirtellous to strigillose; peduncle

RUBIACEAE

2.59 cm; branched portion 718 717 cm; bracts 315 mm,
acute to acuminate. Calyx puberulent; ovary portion obovoid to
ellipsoid, 1.52 mm; lobes narrowly triangular, 12 mm, acute;
calycophyll puberulent to strigillose at least on veins and margins, blade drying papery, ovate to broadly ovate, 310 36
cm, 5-veined from near base, base cuneate or acuminate, apex
acute or obtuse, with stipe 23 cm. Corolla strigose to strigillose outside; tube 1822 mm, inside densely sulfur-yellow pi-

323

lose in throat and similarly but more sparsely pubescent near

base; lobes broadly ovate, 2.53 mm, acuminate to aristate with
tip or arista 0.51 mm. Anthers ca. 3 mm. Capsule 68 4.55
mm; seeds 0.10.2 mm, minutely foveolate and sulcate. Fl.
MayOct, fr. JulDec.
Forests; 1001000 m. Guangxi, Yunnan [Laos, N Myanmar, N
Thailand, N Vietnam].

83. SCYPHIPHORA C. F. Gaertner, Suppl. Carp. 91. 1806.

ping hua mu shu
Chen Tao (); Charlotte M. Taylor
Shrubs or small trees, unarmed, rather succulent, sometimes with stilt roots; young growth usually mucilaginous or resinous.
Raphides presumably absent. Leaves opposite, decussate, without domatia, notably leathery, with petioles articulate; stipules persistent, united around stem, rounded to truncate, entire, enclosing well-developed colleters. Inflorescence axillary, congested-cymose
with axes often articulate and regularly dichotomous, pedunculate, bracteate with bracts small and fused in pairs, apparently caducous. Flowers sessile or pedicellate, bisexual, monomorphic. Calyx limb cupuliform, subtruncate or (4 or)5-denticulate. Corolla
white or pink, salverform with tube somewhat expanded in throat, pubescent inside; lobes (4 or)5, convolute in bud. Stamens (4 or)5,
inserted in corolla just below throat, partially to fully exserted; filaments short; anthers dorsifixed, bifid at base. Ovary 2-celled,
ovules 2 in each cell, axile and attached in middle of septum with 1 erect and 1 pendulous; stigmas 2, exserted. Fruit drupaceous (or
indehiscent depending on interpretation), ellipsoid-oblong to ellipsoid, often weakly curved, fleshy becoming corky, (6)8-ridged
or -winged, with calyx limb persistent; pyrene 1, 2-celled, with 2 seeds in each cell, ellipsoid and longitudinally ridged or winged;
seeds medium-sized, subcylindrical; testa membranous; endosperm reduced; cotyledons oblong; radicle long.
One species: coastal China, Madagascar, Philippines, Thailand, Vietnam, and Pacific region east to Caroline Islands, New Caledonia, and
Australia.
This unusual species was studied in some detail by Puff and Rohrhofer (Opera Bot. Belg. 6: 143172. 1993), who reported that the flowers have
an ixoroid pollination mechanism, in which the flowers are protandrous and deposit the pollen on the outside of the stigmas and the style for
dispersal. The presence or absence of raphides seems not to have been specifically noted; their absence is presumed here based on the classification by
many authors of this genus in Gardenieae.
W. C. Chen (in FRPS 71(1): 368. 1999) described the funiculi of the ovules as connate, but this condition has not been otherwise reported and
contradicts the work of Puff and Rohrhofer (loc. cit.).

1. Scyphiphora hydrophyllacea C. F. Gaertner, Suppl. Carp.

91. 1806.
ping hua mu
Shrubs or small trees, 14(6) m tall, turning black or dark
brown when dry; branches weakly flattened to angled or terete,
glabrous to puberulent, with nodes sometimes thickened, sometimes with reduced internodes and congested nodes. Petiole
0.51.5 cm, glabrous; leaf blade drying leathery, obovate to
broadly elliptic, 2.57.5 1.54.5 cm, glabrous on both surfaces, adaxially usually shiny, base cuneate to obtuse, apex

rounded; secondary veins not visible or 46 pairs; stipules 1.5

3 mm, margins often sparsely to densely ciliate. Inflorescence
1.53 22.5 cm; peduncle 0.51 cm. Flowers sessile or pedicellate, pedicels or subtending inflorescence axes to 2 mm. Calyx glabrous to puberulent; ovary portion cylindrical-ellipsoid,
34 mm; limb 11.5 mm, truncate to denticulate. Corolla glabrous outside; tube 45 mm; lobes ovate-ligulate, ca. 2 mm, obtuse. Drupe 811 35 mm, glabrous. Fl. JulNov, fr. Aug
Dec.
Foreshore mud at seasides; near sea level. Hainan [Philippines,
Thailand, Vietnam; SE Asia to Pacific islands, Australia, Madagascar].

84. SERISSA Commerson ex Jussieu, Gen. Pl. 209. 1789.

bai ma gu shu
Chen Tao (); Charlotte M. Taylor
Small shrubs, usually much branched, unarmed, fetid when bruised, usually with short shoots. Raphides present. Leaves opposite but often crowded and apparently verticillate, decussate, without domatia; stipules persistent, interpetiolar and often fused to
petioles, generally triangular to truncate, with 18 bristles. Inflorescences terminal on principal stems and/or terminal on axillary
short shoots and apparently axillary, capitate and several flowered or reduced to 1 flower, sessile, bracteate. Flowers sessile or subsessile, bisexual, distylous. Calyx limb 46-lobed essentially to base. Corolla white to pink, funnelform or tubular-funnelform, inside
villous in tube; lobes 46, valvate-induplicate in bud, with margins sometimes crisped. Stamens 46, inserted in upper part of corolla
tube, included or exserted; filaments short to developed, anthers dorsifixed near base. Ovary 2-celled, ovules 1 in each cell, erect,

RUBIACEAE

324

basal; stigma 2-lobed, included or exserted. Fruit drupaceous or tardily capsular, obconic to obovoid, leathery to apparently dry,
tardily dehiscent septicidally then loculicidally across top, with calyx limb persistent and often enlarging and becoming spiny;
pyrenes 2, 1-celled, each with 1 seed, oblong to obovoid, longitudinally densely ridged.
One or two species: China, Japan, Nepal, Vietnam; one or two species (one endemic) in China.
Puff et al. (Rubiaceae of Thailand, 232. 2005) reported that the fruit of Serissa are dehiscent through an apical operculum, releasing two 1seeded pyrenes; the fruit of the Chinese specimens studied appear to split across the top and partly down the sides to release the obovoid striate
pyrenes through the top or disk portion (i.e., the apical section inside the calyx limb), which may correspond to the dehiscence described by Puff et al.
The leaf and flower size and pubescence appear to be widely variable, which probably has fueled the selection that has produced a wide range of
cultivated forms. Serissa is widely cultivated in tropical and warm temperate regions for its foliage, both variegated and solid, and showy flowers,
including frequently as bonsai plants. Normally the cultivated plants in regions outside the native range do not produce fruit. Puff et al. (loc. cit.)
discussed and illustrated some of the numerous cultivated forms, which go under the English name snowrose.
The number of species of Serissa is controversial. W. C. Ko (in FRPS 71(2): 159162. 1999) and Puff et al. (loc. cit.) recognized two species,
others (Govaerts et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/rubiaceae/; accessed on 20 Aug 2007; D. Lorence, Fl. Mesoamericana, in
prep.) recognize only one species. In general, more robust plants have been included in S. serissoides. W. C. Kos treatment is presented here for
reference, with the description of S. japonica augmented with characters from specimens; however, some specimens are not conclusively assignable to
one of these species.

1a. Leaf blade drying stiffly papery to leathery, ovate to oblanceolate, elliptic, elliptic-oblong, or lanceolate,
0.62.2 0.30.6 mm, apex acute to obtuse or acuminate; flowers solitary to several; corolla tube longer
than calyx lobes .............................................................................................................................................................. 1. S. japonica
1b. Leaf blade drying thinly papery, obovate or oblanceolate, 1.54 0.71.3 cm, apex acute or subacute; flowers
solitary or usually several; corolla tube as long as calyx lobes .................................................................................. 2. S. serissoides
1. Serissa japonica (Thunberg) Thunberg, Nov. Gen. Pl. 132.
1798.
liu yue xue
Lycium japonicum Thunberg, Nova Acta Regiae Soc. Sci.
Upsal. 3: 207. 1780; L. foetidum Linnaeus f.; Serissa foetida
(Linnaeus f.) Lamarck.
Small shrubs, 6090 cm tall, presumably evergreen;
branches weakly flattened to terete, puberulent to villosulous or
hirtellous in interpetiolar lines to sometimes uniformly puberulent throughout. Leaves subsessile or with petiole to 2 mm, glabrous or puberulent to villosulous; blade drying leathery to
stiffly papery, ovate to oblanceolate, elliptic, elliptic-oblong, or
lanceolate, 0.62.2 0.30.6 cm, both surfaces glabrous to
hispidulous or villosulous on principal veins to throughout,
often shiny adaxially, base obtuse to acute, apex acute to obtuse
or acuminate; secondary veins 24 pairs; stipules 0.52 mm,
puberulent to villosulous, truncate to triangular, with bristles
0.54 mm. Flowers solitary to several; bracts narrowly triangular to spatulate, 16 mm, glabrous to villosulous, acute, margins entire to ciliate or hispidulous. Calyx glabrous to puberulent or hirtellous; ovary portion obconic, 11.5 mm; limb lobed
essentially to base; lobes narrowly triangular to lanceolate or
linear, 15 mm, entire to ciliolate or hispid. Corolla outside glabrous to puberulent or villosulous; tube 48 mm; lobes narrowly triangular to ovate, 22.5 mm. Drupes 23 mm; pyrenes
22.5 mm. Fl. AprOct, fr. JunNov.
Streamsides or broad-leaved forests on hills; 1001600 m.
Anhui, Fujian, Guangdong, Guangxi, Jiangsu, Jiangxi, Sichuan, Taiwan,
Yunnan, Zhejiang [widely cultivated elsewhere].

The Fl. Taiwan (ed. 2, 4: 324325. 1998) treated one species of

Serissa, which was called S. serissoides but which keys out to S. japonica in the classification of W. C. Ko (in FRPS 71(2): 160. 1999); that
report is accordingly listed here based on this re-identification.

2. Serissa serissoides (Candolle) Druce, Rep. Bot. Soc. Exch.

Club Brit. Isles 1916: 646. 1917.
bai ma gu
Democritea serissoides Candolle, Prodr. 4: 540. 1830;
Leptodermis nervosa Hutchinson; Serissa democritea Baillon,
nom. illeg. superfl.
Small shrubs, to 1 m tall; branches stout, subterete, gray,
pilosulous to glabrescent or puberulent. Leaves usually in clusters, subsessile; blade drying thinly papery, obovate or oblanceolate, 1.54 0.71.3 cm, glabrous except sparsely pubescent
abaxially, base acute, apex acute or subacute; secondary veins 2
or 3 pairs; stipules with lobes subulate, ca. 2 mm, broad at base,
sparsely pubescent. Flowers solitary to usually several; bracts
membranous, elliptic, ca. 6 mm, long acuminate, sparsely ciliolate. Calyx glabrescent; ovary portion obconic, ca. 1 mm; limb
lobed essentially to base; lobes 5, lanceolate-subulate, ca. 4 mm,
sharply acute, ciliate. Corolla outside glabrous; tube ca. 4 mm;
lobes 5, oblong-lanceolate, ca. 2.5 mm. Fruit not seen. Fl. Apr
Jun.
Wastelands, lawns. Anhui, Fujian, Guangdong, Guangxi, Hubei,
Jiangsu, Jiangxi, Taiwan, Zhejiang [Japan].
W. C. Ko (in FRPS 71(2): 162. 1999) reported this species also
from Japan, but the Fl. Japan (3a: 228229. 1993) did not recognize it.

85. SINOADINA Ridsdale, Blumea 24: 351. 1979.

ji zai mu shu
Chen Tao (); Charlotte M. Taylor
Trees, unarmed; buds rounded. Raphides absent. Leaves opposite, decussate, usually with domatia; stipules caducous, interpeti-

RUBIACEAE

325

olar, broadly triangular in outline, deeply bifid. Inflorescences terminal and in axils of uppermost leaves, capitate with 711 globose
heads in a cymose arrangement, many flowered, pedunculate with peduncles occasionally articulate though usually ebracteate in
upper half, bracteate; bracteoles filiform to filiform-clavate. Flowers sessile, bisexual, monomorphic. Calyx limb 5-lobed; lobes
obtuse. Corolla yellow, salverform to slenderly funnelform, pubescent inside; lobes 5, thinly imbricate in bud. Stamens 5, inserted in
upper part of corolla tube, partially exserted; filaments short; anthers basifixed. Ovary 2-celled, ovules 412 in each cell on axile
placentas attached in upper third of septum; stigma obovoid, exserted. Fruiting heads globose. Fruit capsular, obconic, septicidally
dehiscent into 2 valves from base to apex with valves separating along sides or from base to apex and away from persistent septum,
sometimes later splitting loculicidally into 2 more valves, stiffly cartilaginous, with septum persistent or tardily deciduous, with calyx
limb persistent on septum; seeds several, medium-sized, fusiform to spatulate, slightly to rather strongly flattened, winged at both
ends.
One species: China, Japan, Myanmar, Thailand.
Ridsdale (loc. cit.) described the arrangement of the corolla lobes in bud as valvate but subimbricate at the apex; on the specimens studied
these appear to be imbricate with the margins very thinly overlapping, a condition sometimes called subimbricate or thinly imbricate elsewhere in
this treatment (e.g., Timonius). Ridsdale described the seeds as trigonal to tricornute, slightly bilaterally compressed, not winged, but the seeds on
the specimens studied are flattened and shortly winged at the ends (e.g., Fang 8106, MO; Tsui 756, MO).

1. Sinoadina racemosa (Siebold & Zuccarini) Ridsdale,

Blumea 24: 352. 1979.
ji zai mu
Nauclea racemosa Siebold & Zuccarini, Abh. Math.-Phys.
Cl. Knigl. Bayer. Akad. Wiss. 4: 178. 1846; Adina asperula
Handel-Mazzetti; A. mollifolia Hutchinson; A. racemosa (Siebold & Zuccarini) Miquel; Cornus esquirolii H. Lveill; N.
taiwaniana Hayata; N. transversa Hayata.
Semi- to fully deciduous trees, 412 m tall; trunk bark
gray; branches glabrous. Petiole (1)36(8) cm, glabrous or
puberulent; leaf blade drying thinly leathery, ovate, ovate-oblong, or elliptic, (4)915(25) (3)510(18) cm, adaxially
shiny and glabrous or infrequently sparsely hirtellous, abaxially glabrous to pilosulous, base cordate to obtuse, sometimes
slightly inequilateral, apex acute to acuminate; secondary veins
612 pairs, sometimes with foveolate or to pilosulous domatia;
stipules (5)1015 25 mm, puberulent to glabrous, lobes

suborbicular. Inflorescence densely puberulent; peduncles 13

cm; branched portion 510 510 cm, branched to 1 order;
flowering heads 48 mm in diam. across calyces, 1418 mm in
diam. across corollas; bracteoles ca. 1 mm. Calyx villosulous;
ovary portion ellipsoid-obcuneate, 0.71 mm, surrounded at
base by a ring of trichomes 0.51 mm; limb deeply lobed, lobes
spatulate, ca. 0.5 mm, at apex rounded and thickened. Corolla
outside densely woolly puberulent; tube (3)45 mm; lobes
spatulate to deltoid, 0.51 mm, acute to obtuse. Stigma fusiform, 0.30.4 mm, exserted for 46 mm. Fruiting head 1115
mm in diam. Capsules obovoid-cuneate, 57 mm, sparsely hirtellous; seeds 2.53.5 0.51 mm, often bifid at apex. Fl. and
fr. MayDec.
Sunny watersides, forests; 3001000(1500) m. Anhui, Fujian,
Guangdong, Guangxi, Guizhou, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Japan, Myanmar, Thailand].
The synonym Cornus esquirolii was first cited for this species
by Lauener and Ferguson (Notes Roy. Bot. Gard. Edinburgh 32: 103.
1972), based on a type from Guizhou, China.

86. SPERMACOCE Linnaeus, Sp. Pl. 1: 102. 1753.

feng hua cao shu
Chen Tao (); Charlotte M. Taylor
Bigelovia Sprengel (1824), not Smith (1819), nor Sprengel (1820), nor Bigelowia Rafinesque (1817), nor Candolle (1836, nom.
cons.); Borreria G. Meyer (1818, nom. cons.), not Borrera Acharius (1810) [Fungi].
Herbs, annual or perennial, subshrubs, or low shrubs [rarely dioecious], unarmed. Raphides present. Leaves opposite, sometimes borne on very short axillary stems and these plus long-stem leaves appearing whorled or fascicled, without domatia; stipules
persistent, fused to petiole bases or leaves, sheath truncate to broadly rounded or broadly triangular, usually thinly textured, with (1
or)211 bristles to multi-fimbriate, bristles occasionally glandular at apex. Inflorescences terminal and/or axillary, capitate or glomerulate, several to many flowered, sessile, bracteate, often partially enclosed in an expanded stipule sheath; bracts usually filiformlaciniate or stipuliform. Flowers sessile or subsessile, bisexual, monomorphic [or rarely distylous or unisexual]. Calyx limb deeply to
completely 2- or 4[8]-lobed, lobes sometimes unequal in pairs. Corolla white sometimes flushed with blue or pink [to sometimes
pink, red, blue, or violet], salverform to funnelform, inside variously glabrous throughout, pubescent in throat or throughout, or usually with a ring of pubescence at stamen insertion; lobes 4, valvate in bud [and infrequently with abaxial horns or appendages]. Stamens 4, inserted in corolla throat to near base of tube, included or exserted; filaments short or developed; anthers dorsifixed. Ovary
2-celled, ovules 1 in each cell, axile near middle of septum; stigma capitate or 2-lobed with lobes short to linear, included or exserted.
Fruit capsular, ellipsoid to subglobose, septicidally then loculicidally dehiscent with valves remaining joined at base [or sometimes
separating completely or remaining connected at apex or partially schizocarpous with 1 valve loculicidal and 1 valve indehiscent],
papery to thickly textured, with calyx limb persistent; seeds medium-sized, ellipsoid to subglobose, with ventral (i.e., adaxial)
groove, with testa thin, smooth to variously ornamented including minutely pitted to rugose, ruminate, and/or reticulate; endosperm
corneous or fleshy; cotyledons leaflike; hypocotyl terete, basiscopic.

326

RUBIACEAE

About 250300 species: widespread in tropical to warm temperate regions worldwide with several species widely naturalized; seven species
(four introduced) in China.
These species were treated in the genus Borreria by W. C. Ko (in FRPS 71(2): 205210. 1999); B. shandongensis as treated by Ko is here
considered a synonym of Diodia teres. Borreria has traditionally been separated from Spermacoce based on fruit dehiscence, with both of the fruit
valves dehiscent in Borreria vs. one dehiscent and one indehiscent in Spermacoce (vs. both indehiscent in Diodia), but based on pantropical surveys
of this group and molecular data the majority of authors today include Borreria in Spermacoce (Verdcourt, Fl. Trop. E. Africa, Rub. (Pt. 1), 339374.
1976; Deb & Dutta, J. Econ. Taxon. Bot. 5(5): 10371063. 1984; Chaw & Peng, J. Taiwan Mus. 40(1): 7183. 1987; Dessein, Syst. Stud.
Spermacoceae (Ph.D. Diss.), University of Leuven, Belgium, 1403. 2003). The treatment here follows recent neotropical studies as to the separation
of and names used for the adventive neotropical species (Burger & Taylor, Fieldiana, Bot., n.s., 33: 1333. 1993; C. D. Adams, Flora Mesoamericana,
in prep.). In particular, seed coat sculpture or texture is informative at the species level, though it must be observed at high magnification (Dessein, loc.
cit.); Chaw and Sivarajan (Bot. Bull. Acad. Sin., n.s., 30: 1524. 1989) illustrated seed coats of many Chinese species. Pollen is also informative in
this group (e.g., Dessein et al., Australian J. Bot. 18: 367382. 2005) but is incompletely studied.

1a. Corolla relatively small, with tube plus lobes 0.51 mm, with corolla tube shorter than, equal to, or slightly longer
than calyx lobes; calyx lobes 2 or 4; mature fruit 0.61.1 0.31 mm.
2a. Stem angles narrowly winged; leaf blade ovate or elliptic-oblong, 415 mm wide; seed coat apparently covered
by numerous fine horizontal striations or ridges ........................................................................................................... 3. S. exilis
2b. Stems smooth to angled, angles without wings; leaf blade narrowly elliptic-oblong to elliptic, 110 mm wide;
seed coat with coarse rounded pits arranged in longitudinal (i.e., vertical) rows .................................................. 5. S. prostrata
1b. Corollas larger, with tube alone 0.510 mm and longer than calyx lobes; calyx lobes 4; mature fruit
15 13.5 mm.
3a. Fruit 12 11.5 mm; leaves linear-oblong or narrowly elliptic, 2.516 mm wide; corolla tubes 0.51.5 mm.
4a. Leaves linear-oblong, 2.56 mm wide; seeds appearing smooth ........................................................................ 6. S. pusilla
4b. Leaves narrowly elliptic to lanceolate, 416 mm wide; seeds transversely ruminate-rugose with irregular
deep grooves ......................................................................................................................................................... 7. S. remota
3b. Fruit 2.25 1.53.5 mm; leaves elliptic, ovate-oblong, oblong-ellipsoid, obovate, or spatulate, 340 mm wide.
5a. Leaf blade elliptic or ovate-oblong, usually widest near middle, 1275 640 mm; plants often drying
yellowish green; corolla tube 23 mm .................................................................................................................... 1. S. alata
5b. Leaf blade oblong-elliptic, obovate, or spatulate, usually widest above middle, 1030 318 mm; plants
usually drying dull green to grayish; corolla tube 2.510 mm.
6a. Mature seeds 22.5 mm, oblong to elliptic-oblong in outline, brown; corolla tube slender,
6.510 mm ............................................................................................................................................... 2. S. articularis
6b. Mature seeds 2.23 mm, elliptic to elliptic-oblong in outline, black; corolla tube funnelform,
2.54.5 mm ................................................................................................................................................... 4. S. hispida
1. Spermacoce alata Aublet, Hist. Pl. Guiane 1: 55. 1775.
kuo ye feng hua cao
Borreria alata (Aublet) Candolle; B. latifolia (Aublet) K.
Schumann; Spermacoce latifolia Aublet
Herbs, perennial, erect to weak or clambering, sometimes
fleshy, usually drying yellowish green, to 1 m; stems 4-angled,
hispidulous or pilosulous and sometimes also hirsute, angles
rounded to acute or very narrowly winged, wings entire. Leaves
sessile to shortly petiolate; petiole to 4 mm, pilosulous or hirtellous; blade drying papery, elliptic or ovate-oblong, 1275
640 mm, both surfaces sparsely to densely hispidulous to pilosulous, base cuneate to obtuse then long decurrent, apex acute
or obtuse; secondary veins 5 or 6 pairs; stipules hirtellous to
hispidulous, sheath 11.5 mm, with 59 bristles or narrowly triangular lobes 17 mm, ciliate. Inflorescences axillary and infrequently apparently also terminal, 615 mm in diam., few to several flowered, notably hispidulous to pilosulous; bracts filiform,
0.54 mm. Calyx moderately to densely hirtellous or pilosulous; hypanthium portion ellipsoid to obovoid, ca. 0.5 mm;
lobes 4, lanceolate to elliptic or triangular, 12 mm. Corolla
white tinged with blue to pale purple, funnelform, outside pilosulous to hirtellous; tube 23 mm, pubescent in throat; lobes
triangular, 11.5 mm. Capsules ellipsoid to subglobose, 33.5
23 mm, densely hirtellous and often also hirsute on upper
portion, densely puberulent to strigillose on sides, stiffly papery

to cartilaginous, septicidal from apex with valves usually remaining connected at base, then both valves loculicidal through
septum; seeds pale brown or dark brown, ellipsoid, ca. 2 1
mm, obtuse at both ends, shiny or dull, surface with numerous
tiny pits not organized into rows. Fl. and fr. MayNov.
Naturalized in disturbed ground and wastelands; below 100800
m. Fujian, Guangdong, Hainan, Taiwan, Zhejiang [apparently native to
the Neotropics but exact origin unknown; Antilles, Central America,
North America (Mexico, Florida), widespread in tropical South America; naturalized in Africa, S and SE Asia, Australia, Madagascar, and
perhaps North America].
The seeds of this species were illustrated by Chaw and Sivarajan
(Bot. Bull. Acad. Sin., n.s., 30: 20, f. 2527. 1989). This species is
considered an invasive weed in the area of Guangzhou in Guangdong
Province. W. C. Ko (in FRPS 71(2): 207. 1999) described the fruit as
septicidal to base at maturity, septum not caducous, or septum of one
valve caducous, but there seems to be a confusion here and this description is not accurate for this species.
The names Spermacoce alata and S. latifolia (or Borreria alata
and B. latifolia) have been treated as distinct species by many authors
but synonymized by others, variously under each of these names. There
now appears to be only one species here, which takes the name S. alata.
Aublets names were published simultaneously; although the names B.
latifolia and S. latifolia have been more often used, these species were
apparently first synonymized by Hara and Gould (Enum. Fl. Pl. Nepal,
199209. 1979) under the name B. alata.

RUBIACEAE

2. Spermacoce articularis Linnaeus f., Suppl. Pl. 119. 1782.

chang guan cao ye feng hua cao
Borreria articularis (Linnaeus f.) F. N. Williams; Spermacoce flexuosa Loureiro.
Herbs, perennial, or subshrubs, prostrate to weakly ascending, perhaps to 50 cm tall; stems subterete to quadrate, glabrescent on sides, angles sharp to winged with wings to 0.1 mm
wide, ciliolate to ciliate. Leaves sessile or subsessile; blade
drying papery to leathery, oblong-elliptic, obovate, or spatulate,
815(25) 310 mm, both surfaces scaberulous-hispidulous
and sometimes also hirtellous or hirsute, base cuneate to obtuse, margins scaberulous and often revolute, apex obtuse or
rounded; secondary veins 2 or 3 pairs or not visible; stipules
densely puberulent, sheath 12 mm, with 57 bristles 0.52(6)
mm. Inflorescences axillary, 58 mm in diam., with 16 flowers
per axil; bracts linear or infrequently stipuliform, 15 mm. Calyx puberulent to hirtellous or scaberulous; hypanthium portion
ellipsoid, 0.81 mm; lobes 4, linear to narrowly triangular, 1
1.5 mm, ciliolate or ciliate. Corolla perhaps pink to white, very
slenderly funnelform to salverform, glabrous outside; tube
(6.5)910 mm, glabrous in throat; lobes triangular, 12 mm.
Capsules ellipsoid to subglobose, sometimes slightly flattened
perpendicular to septum, 2.22.5 1.52.5 mm, puberulent to
hirtellous, pilosulous, and/or hispidulous, papery to cartilaginous, septicidal from apex with valves usually remaining connected at base, then both valves loculicidal through septum;
seeds brown, oblong to elliptic-oblong in outline, 22.5 mm,
obtuse at both ends, shiny, surface minutely granular or
dimpled. Fl. and fr. MayOct.
On open sandy lands at lower elevations. Fujian, Guangdong
(Nanhai Zhudao), Taiwan (introduced and naturalized) [India, Indonesia, Japan (Ryukyu Islands), Malaysia, Nepal, Pakistan, Philippines,
Sri Lanka, Vietnam; Africa, Australia].
The seeds of this species (as circumscribed here) were illustrated
by Chaw and Sivarajan (Bot. Bull. Acad. Sin., n.s., 30: 19, f. 1718.
1989).
This name is here used differently than in many previous floras in
this region; the commonly collected plants treated by Chaw and Peng (J.
Taiwan Mus. 40(1): 7183. 1987) and W. C. Ko (in FRPS 71(2): 206
207. 1999) as Spermacoce articularis are here treated as S. hispida,
which is the older name. Overall, there is confusion and/or disagreement about the application of these two names as well as the number of
specimens that should be separated among these plants, which have
been studied only based on regional floristic work rather than a systematic review of this species group across its entire range. Deb and
Dutta (J. Econ. Taxon. Bot. 5(5): 10461048. 1984) synonymized S.
hispida and S. articularis and recognized a single species, for which
they described the corolla tube as .31.7 mm ... shorter than the corolla
lobes while illustrating correspondingly a corolla with the tube 46
mm and several times longer than the corolla lobes. Sivarajan and Nair
(Taxon 35: 363369. 1986), followed by Ridsdale (in Dassanayake,
Revis. Handb. Fl. Ceylon 12: 332341. 1998) and Mill (Fl. Bhutan 2(2):
817820. 1999), recognized two species in this group; however, a
number of individual Chinese plants have some characters of each of
these so their distinctions are problematic in our region. Two distinct
groups of plants are separable in our region using the characters listed in
the key to species above, and the protologues of these two names each
correspond to one of these groups; the treatment here generally follows
the annotations of Fosberg (in herb.).

327

3. Spermacoce exilis (L. O. Williams) C. D. Adams, Fieldiana,

Bot., n.s., 33: 316. 1993.
er e feng hua cao
Borreria exilis L. O. Williams, Phytologia 28: 227. 1974,
based on B. gracilis L. O. Williams, Phytologia 26: 487. 1973,
not Scheele (1844), nor Spermacoce gracilis Ruiz & Pavon
(1798); B. repens Candolle; S. decandollei Deb & R. M. Dutta,
nom. illeg. superfl.; S. mauritiana Gideon; S. repens (Candolle)
Fosberg & D. A. Powell (1980), not Willdenow ex Chamisso &
Schlechtendal (1828), nor Sess & Mocio (1893), nor Larraaga (1923).
Herbs, annual to perhaps perennial, slender, creeping to
weakly ascending, to 30 cm tall; stems 4-angled, puberulent to
glabrescent, angles winged, wings 0.10.5 mm wide, ciliate or
ciliolate. Leaves subsessile to shortly petiolate; petiole to
1.5 mm, puberulent to glabrescent; blade drying membranous,
ovate or elliptic-oblong, 0.730 415 mm, sparsely puberulent to glabrous throughout or pilose to hispid along midrib
abaxially, base obtuse to cuneate, apex acute to obtuse; secondary veins 2 or 3 pairs; stipules pilosulous or hirtellous to
glabrescent, sheath 0.51 mm, with 510 bristles 0.52 mm,
often glandular. Inflorescences terminal and in uppermost leaf
axils, 36 mm in diam., several to many flowered; bracts numerous, filiform, 0.51.5 mm. Calyx hirtellous to glabrescent;
hypanthium portion obovoid, ca. 0.3 mm; lobes 2, linearlanceolate to triangular, 0.40.9 mm. Corolla white, rotate to
shortly tubular, 0.50.6 mm, outside glabrous, bearded in throat;
lobes spatulate triangular, as long as tube. Capsules ellipsoid,
weakly to strongly flattened at right angles to septum, 11.1
0.81 mm, glabrescent, membranous and sometimes somewhat hyaline, septicidal from apex then both valves loculicidal through septum or sometimes fragmenting; seeds brownish
yellow, ellipsoid, ca. 0.8 0.4 mm, obtuse at both ends, shiny,
surface apparently with numerous fine horizontal striations or
ridges (at 10; but actually with minute transverse pits, visible
at 40). Fl. and fr. almost year-round.
Naturalized in disturbed humid sites at low elevations. Hainan,
Hong Kong, Taiwan [apparently native to the Neotropics but exact origin unknown; India, Indonesia, Nepal, Sri Lanka, Vietnam; Africa,
Antilles, Australia, Central America, Indian Ocean islands (Mauritius),
North America (Mexico), Pacific islands, N South America].
The seeds of this species were illustrated in detail by Chaw and
Sivarajan (Bot. Bull. Acad. Sin., n.s., 30: 20, f. 2830. 1989, as Spermacoce mauritiana). The taxonomy and circumscription of this adventive
species here follows Burger and Taylor (Fieldiana, Bot., n.s., 33: 316.
1993). The illustration presented by Deb and Dutta (J. Econ. Taxon.
Bot. 5(5): 1045, f. 2. 1984, as S. decandollei) seems to be based on a
mixed collection of S. exilis and S. prostrata according to the circumscription of these species here. Both S. exilis and S. prostrata were
formerly included in a broadly circumscribed, morphologically heterogeneous Spermacoce ocymoides Burm. f. Several authors have concluded that this last name applies to a species found only in SE Asia and
was incorrectly applied to American and adventive plants (Dessein,
Syst. Stud. Spermacoceae (Ph.D. Diss.), University of Leuven, Belgium, 1403. 2003, and references cited there); presumably the species
treated by Bakhuizen f. in the Fl. Java is true S. ocymoides.
The description of Spermacoce exilis here includes observations
from specimens from outside China; this species has probably been introduced to China more than once, thus it seems useful to include the
variation found in adjacent regions that may yet be found in China.

RUBIACEAE

328

4. Spermacoce hispida Linnaeus, Sp. Pl. 1: 102. 1753.

cao ye feng hua cao
Borreria hispida (Linnaeus) K. Schumann.
Herbs, annual or perennial, or subshrubs, prostrate to perhaps weakly ascending, apparently often fleshy, to 50 cm tall;
stems subterete to usually markedly quadrate, glabrous to puberulent or pilosulous on sides, angles cartilaginous to winged,
wings to 0.1 mm wide, ciliolate, hispidulous, or ciliate. Leaves
sessile to shortly petiolate; petiole to 4 mm, hirtellous throughout or ciliolate in lines; blade drying papery to leathery, oblongelliptic, obovate, or spatulate, 1030(40) 515(18) mm,
both surfaces hirtellous to scaberulous, hispidulous, and/or hirsute, base cuneate to obtuse and usually long decurrent, margin
scaberulous or ciliolate and often revolute, apex acute, obtuse,
or rounded; secondary veins 2 or 3(or 4) pairs or not visible;
stipules moderately to densely puberulent, hirtellous, and/or pilosulous often in lines, sheath 13 mm, with 57 bristles 15
mm. Inflorescences axillary, 515 mm in diam., with 16 flowers per axil; bracts linear or infrequently stipuliform, 15 mm.
Calyx puberulent to hirtellous or scaberulous; hypanthium portion ellipsoid, 0.81 mm; lobes 4, linear-lanceolate to narrowly
triangular, 11.5 mm, ciliolate or ciliate. Corolla pink, purple,
or white, funnelform, outside glabrous or hispidulous to pilosulous on upper part; tube 2.54.5 mm, glabrous in throat; lobes
elliptic-oblong, lanceolate, or triangular, 11.8 mm. Capsules
ellipsoid to subglobose, sometimes weakly flattened perpendicular to septum, 2.55 2.53.5 mm, puberulent, hirtellous,
pilosulous, and/or hispidulous, papery to cartilaginous, septicidal from apex with valves usually remaining connected at base
then both valves loculicidal through septum, with calyx lobes
sometimes enlarging, up to 2.2 mm; seeds black, elliptic to
elliptic-oblong in outline, 2.23 mm, obtuse at both ends, shiny
to dull, surface minutely granular or dimpled. Fl. and fr. Mar
Dec.
On open sandy lands at lower elevations; sea level to 100 m. Fujian, Guangdong, Guangxi, Hainan, Taiwan [India, Indonesia, Malaysia,
Philippines, Sri Lanka, Vietnam; Australia].
This name is applied here to most of the plants treated by W. C.
Ko (in FRPS 71(2): 206207. 1999) as Borreria articularis; see the
comments about these two species above, under Spermacoce articularis. The seeds of this species (as circumscribed here) were illustrated
by Chaw and Sivarajan (Bot. Bull. Acad. Sin., n.s., 30: 19, f. 2224.
1989). The leaf measurements included above in parentheses are taken
from Wang & Li W05246 (Taiwan, MO!), which is an exceptionally
robust plant.

5. Spermacoce prostrata Aublet, Hist. Pl. Guiane 1: 58. 1775.

pu fu feng hua cao
Borreria prostrata (Aublet) Miquel.
Herbs, annual or probably perennial, slender, weakly ascending to erect at least at ends of stems, to 65 cm tall; stems
rounded to subquadrate, smooth to sharply angled and/or longitudinally sulcate, scaberulous to glabrescent. Leaves sessile;
blade drying membranous, narrowly elliptic-oblong to elliptic,
1030(45) 17(10) mm, puberulent and/or scaberulous to
glabrescent throughout or sometimes pilose along midrib abaxi-

ally, base obtuse to cuneate, apex acute to obtuse; secondary

veins 25 pairs; stipules puberulent, hirtellous, or glabrescent,
sheath 0.51 mm, with 59 bristles 0.52 mm, often glandular.
Inflorescences terminal and in uppermost leaf axils, several to
many flowered, 34 mm in diam.; bracts numerous, filiform,
0.51 mm. Calyx glabrescent; hypanthium portion obovoid, ca.
0.3 mm; lobes 2 or 4, narrowly triangular, 0.30.7 mm. Corolla
white, rotate to shortly tubular, 0.71 mm, outside glabrous,
pubescent in throat; lobes spatulate-triangular, as long as tube.
Capsules ellipsoid, weakly to strongly flattened at right angles
to septum, 0.60.9 0.30.4 mm, glabrescent, membranous and
sometimes somewhat hyaline, septicidal from apex, then both
valves loculicidal through septum or sometimes fragmenting;
seeds brownish yellow, ellipsoid, ca. 0.5 0.2 mm, obtuse at
both ends, with coarse rounded pits arranged in longitudinal
(i.e., vertical) rows. Fl. and fr. almost year-round.
Naturalized in disturbed wet sites at low elevations. Hainan,
Hong Kong, Taiwan [apparently native to the Neotropics but exact origin unknown; India, Indonesia, Sri Lanka; Antilles, Central America,
Indian Ocean islands (Mauritius), North America (Mexico), Pacific
islands, N South America].
This species has been widely confused with and/or combined with
Spermacoce exilis, and has sometimes been misidentified as S. ocymoides; see the discussion under S. exilis, above.

6. Spermacoce pusilla Wallich in Roxburgh, Fl. Ind. 1: 379.

1820.
feng hua cao
Borreria pusilla (Wallich) Candolle.
Herbs, apparently annual, slender, erect, to 60 cm tall;
stems subterete to 4-angled, glabrous to densely scaberulous at
least along angles or ridges. Leaves subsessile; blade drying papery to leathery, linear-oblong, 1250 2.56 mm, adaxially
densely scaberulous to hispidulous, abaxially glabrous or often
pilosulous, hirtellous, or hispidulous along midrib, base generally straight (i.e., not tapering), margins often revolute at least
when dry, apex acute to acuminate; secondary veins 2 or 3 pairs
or not visible; stipules densely pilosulous or scaberulous often
with unusual clavate trichomes, sheath 12.5 mm, with 57
bristles 26.5 mm. Inflorescences terminal and axillary at most
stem nodes, 512 mm in diam., several to many flowered; bracts
filiform, 14 mm. Calyx glabrescent to densely puberulent or
hirtellous; hypanthium portion turbinate, ca. 0.5 mm; lobes 4,
linear-lanceolate, 11.5 mm. Corolla funnelform, white tinged
red on upper parts, outside glabrous; tube 1.21.5 mm, glabrous
in throat; lobes linear-lanceolate to triangular, 0.81.5 mm,
sometimes pubescent inside near tips. Capsules sometimes
shortly stipitate, oblong or subobovate to ellipsoid, usually
weakly flattened perpendicular to septum, 12 11.5 mm,
glabrescent at base, glabrescent to densely hirtellous near apex,
septicidal from apex with valves often remaining connected at
base, then both valves loculicidal through septum and often partially splitting abaxially; seeds dark brown, narrowly oblong in
outline, 1.32.2 ca. 0.5 mm, obtuse at both ends, shiny,
smooth. Fl. and fr. AugDec.
Grasslands and grassy slopes at lower elevations; 1001500 m.
Anhui, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Jiangxi, Tai-

RUBIACEAE

wan, Yunnan, Zhejiang [Bhutan, India, ?Indonesia, Malaysia, Myanmar,

Nepal, Pakistan, Philippines, Sri Lanka, Thailand, Vietnam; introduced
in tropical Africa].
This species was discussed and well illustrated for Taiwan by
Chaw and Peng (J. Taiwan Mus. 40(2): 5759. 1987); its seeds were
illustrated in detail by Chaw and Sivarajan (Bot. Bull. Acad. Sin., n.s.,
30: 18, f. 1011. 1989).
These plants were treated by W. C. Ko (in FRPS 71(2): 208. 1999)
as Spermacoce stricta Linnaeus f. (Borreria stricta (Linnaeus f.) G.
Meyer), following previous usage by various authors (see discussion in
Chaw & Peng, loc. cit.), but as detailed by Sivarajan and Nair (Taxon
35: 363369. 1986) the identity of the name S. stricta is not at all clear
and very likely actually applies to a species of Hedyotis. W. C. Ko (loc.
cit.) described the seeds of this species as transversely striate/grooved
and with one end mucronate, another end obtuse; however, the seeds
are smooth with both ends obtuse to rounded on all specimens studied,
and as described by Dessein (Syst. Stud. Spermacoceae (Ph.D. Diss.),
University of Leuven, Belgium, 1403. 2003).
Sivarajan and Nair (loc. cit.) separated the Indian plants treated
in Spermacoce stricta into two species, S. pusilla and a newly
described species, S. ramanii Sivarajan & R. V. Nair. They gave the
range of S. ramanii only as India, although they considered several
additional names synonymous, with a consequent tacit expansion of
its range to Thailand, New Guinea, and Java. Dessein (loc. cit.) discussed the separation of these and concluded that there appear to be
two species in India but only one variable species in Africa; he found
the contrasting character states in all possible combinations in Africa
and did not adopt the name S. ramanii for any African plants. The
Chinese plants seen appear to comprise one well-delimited species and
are here all treated as S. pusilla; however, as with the African plants,
several of the features that Sivarajan and Nair used to separate S.
ramanii, which are largely vegetative characters, are found on some
Chinese plants in various combinations with other features. The name S.
ramanii is, therefore, not synonymized nor used here for any Chinese
plants; it has been cited for China by the Kew Rubiaceae checklist
(Govaerts et al., World Checkl. Rubiaceae; http://www.kew.org/wcsp/
rubiaceae/; accessed on 15 Sep 2010) but without documentation of the
report.

7. Spermacoce remota Lamarck, Tabl. Encycl. 1: 273. 1792.

329

guang ye feng hua cao

Borreria assurgens (Ruiz & Pavon) Grisebach; B. remota
(Lamarck) Bacigalupo & E. L. Cabral; Spermacoce assurgens
Ruiz & Pavon.
Herbs, perennial, or subshrubs, ascending to erect, to 65
cm tall; stems subterete to subquadrate, sulcate and/or ridged,
glabrous or ciliolate on angles. Leaves sessile to petiolate; petiole to 3 mm, glabrescent; blade drying papery, narrowly elliptic
to lanceolate, 1045 416 mm, puberulent to glabrescent,
base acute to cuneate, apex acute; secondary veins 2 or 3 pairs;
stipules puberulent or hirtellous to glabrescent, sheath 13 mm,
with 57 bristles 0.52 mm. Inflorescences terminal and in uppermost leaf axils, 512 mm in diam., many flowered; bracts
numerous, filiform, 0.51 mm. Calyx puberulent or hirtellous to
glabrescent; hypanthium portion obovoid, ca. 0.5 mm; lobes 4,
narrowly triangular to linear, 0.81 mm. Corolla white, funnelform, outside glabrous or puberulent on lobes; tube 0.51.5
mm, pubescent in throat; lobes triangular, 11.5 mm. Capsules ellipsoid, weakly to strongly flattened at right angles to
septum, 1.82 11.2 mm, hirtellous or puberulent, papery,
septicidal from apex with valves usually remaining connected
at base, then both valves loculicidal through septum and often
splitting abaxially; seeds brownish yellow, ellipsoid, 1.51.8
0.81 mm, obtuse at both ends, somewhat shiny, transversely
ruminate-rugose with irregular deep grooves. Fl. and fr. Jun
Jan.
Naturalized in disturbed wet sites; below 100300 m. Guangdong,
Taiwan [apparently native to the Neotropics; India, Indonesia, Singapore, Sri Lanka, Thailand, Vietnam; Antilles, Australia, Central
America, Indian Ocean islands (Mauritius), North America (Mexico),
Pacific islands, N South America].
This widely naturalized species was reported from Taiwan by
Chaw and Peng (J. Taiwan Mus. 40(1): 7183. 1987), who noted that it
has been widely misidentified as Borreria laevis, but that name
applies to a distinct Asian species of restricted range. The seeds of this
species were illustrated by Chaw and Sivarajan (Bot. Bull. Acad. Sin.,
n.s., 30: 20, f. 3436. 1989, as Spermacoce assurgens).

87. SPERMADICTYON Roxburgh, Pl. Coromandel 3: 32. 1815.

xiang ye mu shu
Chen Tao (); Charlotte M. Taylor
Hamiltonia Roxburgh.
Shrubs, erect or clambering, unarmed, fetid when bruised. Raphides present. Leaves opposite, decussate, without domatia; stipules persistent, interpetiolar or shortly united around stem, triangular. Inflorescences terminal, cymose, paniculate, or corymbose,
many flowered, pedunculate, bracteate. Flowers subsessile or sessile, bisexual, distylous. Calyx limb deeply 5-lobed. Corolla white,
blue, pink, or violet, slenderly funnelform with tube prolonged, variously glabrous or pubescent inside; lobes 5, valvate in bud. Stamens 5, inserted in corolla throat, included in long-styled form, exserted in short-styled form; filaments short to developed; anthers
apparently basifixed. Ovary 5-celled, ovules 1 in each cell, erect, basal, anatropous; stigma 5-lobed, included in short-styled flowers,
exserted in long-styled flowers. Fruit drupaceous becoming capsular or perhaps schizocarpous, oblong-ellipsoid, dry, with valves or
perhaps mericarps separating septicidally from apex, with calyx limb persistent; pyrenes or perhaps mericarps 5, 1-celled, each with
1 seed, ellipsoid; seeds medium-sized, ellipsoid-oblong or triangular; testa reticulate; embryo straight; radicle basiscopic.
One species: Bangladesh, Bhutan, India, Nepal, Pakistan; cultivated more widely, including in China.
W. C. Ko (in FRPS 71(2): 119. 1999) estimated six species of Spermadictyon, but all other authors seen report only one species. Ko described
the fruit as having a septum disappearing early and the seeds as having a loose aril, but the meaning of these is not entirely clear and does not correspond to morphology described elsewhere.

RUBIACEAE

330

1. Spermadictyon suaveolens Roxburgh, Pl. Coromandel 3:

32. 1815.
xiang hua mu
Hamiltonia suaveolens (Roxburgh) Roxburgh.
Subshrubs, 13 m tall, perhaps rather fleshy; branches
somewhat flattened to quadrangular or subterete, tomentulose
to glabrescent. Petiole 1218 mm, pilosulous or tomentulose to
glabrescent; leaf blade drying papery, elliptic-lanceolate to
elliptic or ovate, 1320 46.5 cm, adaxially glabrous or pilosulous to puberulent to tomentulose, abaxially tomentulose to
villosulous with pubescence denser on principal veins, base
acute to cuneate or rounded, apex acute to obtuse; secondary
veins 1016 pairs; stipules triangular to broadly triangular, 2
4 mm, tomentulose to glabrous, obtuse to subacute. Inflorescences 525 cm, tomentulose to villosulous; peduncles 35 cm;

bracts ovate to triangular or oblanceolate, 14 mm. Calyx

densely pilosulous; ovary portion cylindrical to ellipsoid, ca. 1
mm; limb lobed essentially to base; lobes linear-lanceolate, 1.2
2 mm, acute. Corolla blue or white, outside densely tomentulose to villosulous; tube 812 mm; lobes ovate to triangular,
1.53 mm, acute. Fruit ellipsoid to ovoid, 34 mm, densely
pilosulous.
Cultivated in gardens and perhaps naturalized. Xizang [Bangladesh, Bhutan, India, Nepal, Pakistan; cultivated more widely].
Raizada and Bennet (Indian Forester 108(2): 302303. 1982)
recognized two varieties of this species, Spermadictyon suaveolens var.
suaveolens with pure white flowers and S. suaveolens var. azureum
(Wallich) Bennet & Raizada, based on S. azureum Wallich, with azureblue flowers. Other authors have not separated these color forms taxonomically but regard the color difference as due only to pigmented vs.
albino flowers, a distinction generally not recognized taxonomically any
more and not recognized here.

88. SPIRADICLIS Blume, Bijdr. 975. 18261827.

luo xu cao shu
Chen Tao (); Charlotte M. Taylor
Herbs, annual or perennial, or subshrubs, unarmed. Raphides present. Leaves opposite, sometimes clustered and appearing
whorled, rosulate, or pseudoverticillate, isophyllous to anisophyllous, without domatia; stipules persistent to caducous, interpetiolar,
triangular, entire to 2(5)-lobed. Inflorescences terminal and/or pseudoaxillary, cymose to paniculiform with axes slender and dichasial or often scorpioid, several to many flowered, pedunculate, bracteate or bracts reduced. Flowers sessile to pedicellate, bisexual,
usually if not always distylous. Calyx with ovary portion usually 5-ridged or -winged, limb 5-lobed. Corolla white, pink, purple, or
red, campanulate, funnelform, urceolate, or tubular, inside usually pubescent in throat and with pubescent ring near middle; lobes 5,
in bud valvate or induplicate-valvate, often winged or keeled dorsally, sometimes notably pinnatinerved. Stamens 5, inserted below
middle of corolla tube and included in long-styled flowers, inserted in throat or middle of corolla tube and partially exserted in shortstyled flowers; filaments short to developed; anthers dorsifixed. Ovary 2-celled, ovules numerous in each cell on peltate axile
placentas attached to middle of septum; stigmas 2-lobed, included to shortly exserted in long-styled flowers, included and positioned
near middle of corolla tube in short-styled flowers. Fruit capsular, subglobose, often 5-ridged to -winged, with apical portion prolonged into beak, dehiscing loculicidally and often also simultaneously or subsequently septicidal from top, dividing partially to
completely into 2 or 4 valves with walls usually persistent, papery to stiff, with calyx limb persistent, sometimes elongating; seeds
numerous, small, angled; testa reticulate or alveolate; embryo minute; endosperm fleshy.
At least 40 species: Bhutan, China, India, Indonesia, Myanmar, Vietnam; 35 species (31 endemic, one of unconfirmed occurrence) in China.
Robbrecht (Opera Bot. Belg. 1: 1271. 1988; Opera Bot. Belg. 6: 1200. 1993) accepted earlier conclusions that Spiradiclis is related to Ophiorrhiza; a more recent study based on molecular data suggests that the situation may be more complex and calls into question the separation of these
genera (Rydin et al., Pl. Syst. Evol. 278: 101120. 2009). H. S. Lo (in FRPS 71(1): 86. 1999) described the corolla lobes as valvate in bud, but they
were described as induplicate-valvate by Bakhuizen f. (Fl. Java 2: 289. 1965). This genus does not appear to be well known at all. It has only been
studied regionally, in particular by H. S. Lo et al. (Acta Bot. Austro Sin. 1: 2736. 1983), H. S. Lo (Bull. Bot. Res., Harbin 6(4): 3153. 1986), Deb
and Rout (Candollea 44: 225229. 1989), H. S. Lo (Bull. Bot. Res., Harbin 18: 275283. 1998), and R. J. Wang (Novon 12: 420423. 2002). Ma et al.
(J. Trop. Subtrop. Bot. 13(3): 264270. 2005) studied seed morphology of sixteen Spiradiclis species and found partial correlation with the
infrageneric classification of Lo.
H. S. Lo (loc. cit. 1998: 275276) recognized two subgenera, distinguished as follows:
Spiradiclis subg. Spiradiclis: ellipsoid to linear-oblong capsules that are 24 as long as wide and have straight valves, including eight (or
possibly nine or ten) species in China;
Spiradiclis subg. Sinospiradiclis H. S. Lo: subglobose capsules that are as long as wide and have the valves ultimately twisted, including the
remaining Chinese species.
One subsequently described species, Spiradiclis chuniana R. J. Wang, was not classified to subgenus when it was published because the fruit
were unknown. The key here follows that of H. S. Lo in FRPS (71(1): 8688. 1999), apparently intended as a schematic outline of the genus
classification, with some changes: here the full range of variation that is given in the descriptions has been added to the key leads, and a few species
have been moved to different sections within the key based on Los descriptions. This key does not fully distinguish all the species; however, it cannot
be improved on with the information now available and is here presented to summarize in English the existing information on Chinese Spiradiclis and
to highlight problematic areas of its taxonomy.

RUBIACEAE

331

1a. Capsules narrowly ellipsoid, ellipsoid, oblanceoloid, narrowly oblong, or linear-oblong, 24 as long as wide,
with valves becoming twisted (dehiscence unknown in S. baishaiensis, S. laxiflora, S. longzhouensis) (S. subg.
Spiradiclis).
2a. Capsules linear-oblong, oblanceoloid, or narrowly oblong, 24 as long as wide, glabrous or puberulent.
3a. Stems and leaves glabrous; petioles 34.5 cm; stipule unlobed ........................................................... 23. S. microcarpa
3b. Stems and abaxial leaf veins glabrescent, scaberulous, puberulent, villosulous, or strigillose; petioles
0.21 cm; stipules deeply 2-lobed.
4a. Corolla tube 22.5 mm; corolla lobes apparently perhaps keeled dorsally; fruit 56 mm, with
2 valves or these later partly splitting ...................................................................................................... 4. S. caespitosa
4b. Corolla tube 34 mm; corolla lobes smooth dorsally; fruit 2.54.5 mm with 4 valves.
5a. Capsules 2.54 mm; stipules persistent ..................................................................................... 1. S. arunachalensis
5b. Capsules 44.5 mm; stipules caducous ............................................................................................. 9. S. cylindrica
2b. Capsules narrowly ellipsoid or ellipsoid, ca. 2 as long as wide, glabrous, pilosulous, or villosulous.
6a. Leaves tomentose or pilosulous at least abaxially; capsules glabrous to pilosulous or villous.
7a. Peduncle 59 cm; calyx lobes 34 mm, longer than fruit; capsules glabrous .................................... 2. S. baishaiensis
7b. Peduncle 0.10.5 cm; calyx lobes ca. 1 mm, shorter than fruit; capsules villosulous ........................... 10. S. emeiensis
6b. Leaves glabrous; capsules glabrous.
8a. Stems with 2 lines of dense hispidulous hairs ..................................................................................... 22. S. malipoensis
8b. Stems glabrous or sparsely evenly pubescent.
9a. Leaves 1015 cm, with secondary veins 911 pairs ......................................................................... 16. S. laxiflora
9b. Leaves 1521 cm, with secondary veins 1623 pairs .............................................................. 20. S. longzhouensis
1b. Capsules subglobose, ellipsoid, to ovoid, or subglobose-obconic, as long as wide, valves remaining
straight (fruit unknown in S. chuniana, S. corymbosa, S. ferruginea, S. fusca, S. longipedunculata,
S. napoensis, S. rubescens, S. spathulata, S. xizangensis; fruit dehiscence unknown in S. hainanensis,
S. longibracteata, S. oblanceolata, S. villosa) (S. subg. Sinospiradiclis).
10a. Leaves cordate, cordulate, subtruncate, or broadly obtuse at base.
11a. Leaves 0.61.8 cm; plants creeping.
12a. Calyx lobes 24 mm, in fruit 2 or 3 as long as capsule .................................................... 13. S. guangdongensis
12b. Calyx lobes 1.21.5 mm, in fruit equal to or shorter than capsule .......................................... 14. S. hainanensis
11b. Leaves 1.513 cm, at least some more than 1.8 cm; plants erect or creeping.
13a. Corolla 2021.5 mm; leaves 1.54 cm; plants creeping .......................................................... 33. S. umbelliformis
13b. Corolla 69 mm; leaves 313 cm; plants ascending or acaulescent.
14a. Plants with leaves distributed along developed stems; leaves 36.5 1.63 cm,
broadly obtuse to truncate at base ............................................................................................ 5. S. chuniana
14b. Plants acaulescent or with short stems, with leaves often clustered at base of plant;
leaves 513 25.5 cm, cordate or cordulate at base ................................................................ 7. S. cordata
10b. Leaves acute, cuneate, obtuse, or rounded at base.
15a. Corolla with tube 13.523 mm.
16a. Calyx lobes 11.5 mm, as long as or shorter than corolla tube (i.e., hypanthium portion
together with unlobed basal part of calyx limb).
17a. Stems developed with leaves borne at developed internodes ................................................ 30. S. scabrida
17b. Stems short or hardly developed, with leaves clustered at base .......................................... 31. S. spathulata
16b. Calyx lobes 1.610 mm, longer than corolla tube.
18a. Leaves glabrous on both surfaces; flowers dark red ................................................................ 6. S. coccinea
18b. Leaves pubescent (hirsute, pilose, strigose, hispidulous, hirtellous) on one or both
surfaces and/or ciliate marginally; flowers red, bluish purple, white, purplish red,
purplish white, or pink (flowers unknown in S. xizangensis).
19a. Corolla tube 1922 mm.
20a. Calyx lobes generally equal in size; leaves adaxially densely
hispidulous-strigose; corolla bluish purple, tube 1921 mm ................ 28. S. purpureocaerulea
20b. Calyx lobes unequal in size; leaves adaxially sparsely puberulent
or glabrous; corolla red or purplish red, tube 2122 mm.
21a. Secondary leaf veins 1829 pairs; calyx lobes 310 mm; corolla
tube ca. 22 mm .................................................................................. 18. S. longibracteata
21b. Secondary leaf veins 911 pairs; calyx lobes 3.54.5 mm; corolla
tube ca. 21 mm .......................................................................................... 29. S. rubescens
19b. Corolla tube 1218 mm (only known in bud in S. napoensis).
22a. Calyx lobes unequal in length, one distinctly longer and/or wider than
others; corolla tubes glabrescent or uniformly pubescent over broad
areas inside.

RUBIACEAE

332

23a. Corolla pubescent inside; leaf secondary veins 815 pairs ..................... 11. S. ferruginea
23b. Corolla glabrescent inside; leaf secondary veins ca. 7 pairs .................. 35. S. xizangensis
22b. Calyx lobes equal or subequal in size; corolla tubes with 1 or 2 discrete
white villous rings of pubescence inside.
24a. Leaves rounded, obtuse, or acute at apex, with 1829 pairs of
secondary veins; corolla sparsely purplish red pubescent
outside; stipules broadly ovate .......................................................... 18. S. longibracteata
24b. Leaves acuminate at apex, with 610 pairs of secondary veins;
corolla glabrous or with 5 lines of pubescence outside;
stipules subulate to linear.
25a. Leaves lanceolate, ovate, subelliptic, or subelliptic-oblong,
yellowish brown on lower surface when dry; stipules and
bracteoles 23 mm; corolla with 5 lines of pubescence
outside, lobes narrowly winged on dorsal surface ................................. 15. S. howii
25b. Leaves narrowly ovate to broadly ovate, pale on lower
surface when dry; stipules 28 mm and bracteoles
610 mm; corolla glabrous outside, lobes smooth
on dorsal surface .............................................................................. 25. S. napoensis
15b. Corolla shorter, tube 29 mm (flowers unknown in S. bifida, S. villosa; only buds known in
S. microphylla).
26a. Stems and inflorescences glabrous.
27a. Leaves narrowly elliptic-oblong or oblanceolate, 3.54.5 as long as wide; stipules
long triangular; stems with developed internodes ........................................................... 26. S. oblanceolata
27b. Leaves obovate, broadly obovate, oblong-elliptic, oblong-lanceolate, or subelliptic,
1.53 as long as wide; stipules triangular, triangular-orbicular, suborbicular, or
long triangular; stem internodes developed to reduced.
28a. Stipules suborbicular or triangular-orbicular, cuspidate and 2-lobed; stem
internodes developed ....................................................................................... 19. S. longipedunculata
28b. Stipules triangular to long triangular, at apex acuminate; stem internodes
developed or reduced.
29a. Leaves 1.53.5 cm wide, with 1013 pairs of secondary veins; petioles
0.51.5 cm; corolla tube ca. 9 mm ............................................................................. 12. S. fusca
29b. Leaves 34.5 cm wide, with 79 pairs of secondary veins; petioles
14 cm; corolla tube ca. 5 mm .................................................................... 21. S. luochengensis
26b. Stems and inflorescences pubescent.
30a. Stems short, with leaves crowded at base of plant.
31a. Flowers subsessile; corolla pubescent inside ..................................................................... 17. S. loana
31b. Flowers on pedicels 15 mm; corolla glabrous inside ................................................ 32. S. tomentosa
30b. Stems prolonged, erect to prostrate, with leaves separated by developed internodes.
32a. Calyx and capsules glabrous or subglabrous; leaves 1021 cm, with 1519 pairs
of secondary veins ................................................................................................................ 3. S. bifida
32b. Calyx and capsules pubescent; leaves 125 cm, with 417 pairs of secondary veins.
33a. Capsules villous; calyx lobes ca. 4 mm ................................................................... 34. S. villosa
33b. Capsules puberulent, pilosulous, pilose, or hispidulous; calyx lobes
0.31.5 mm.
34a. Leaves 6.514 cm, with 1117 pairs of secondary veins; corolla tube
7.59 mm ................................................................................................... 8. S. corymbosa
34b. Leaves 15 cm, with 49 pairs of secondary veins; corolla tube
ca. 2 mm (only known in bud in S. microphylla).
35a. Leaves 13 0.54 cm, with 47 pairs of secondary veins;
fruit with 4 valves ......................................................................... 24. S. microphylla
35b. Leaves 35 1.53 cm, with ca. 9 pairs of secondary veins;
fruit valves 2, sometimes tardily splitting into 4 ............................. 27. S. petrophila
1. Spiradiclis arunachalensis Deb & Rout, Candollea 44: 225.
1989.
zang nan luo xu cao
Spiradiclis caespitosa Blume f. subimmersa H. S. Lo.
Herbs, perennial, usually prostrate and rooting on basal

nodes; stems glabrous to densely villosulous. Petiole 0.21 cm;

leaf blade drying papery, oblanceolate to lanceolate-elliptic,
1.57.5 0.31.8 cm, adaxially glabrescent to scaberulous or
strigillose marginally and on principal veins, abaxially strigillose to puberulent along principal veins and glabrescent on
lamina, base cuneate to acute, apex acute; secondary veins 611

RUBIACEAE

pairs; stipules persistent, deeply 2-lobed, lobes narrowly triangular, 410 mm, strigillose to glabrescent, acute. Inflorescences
dichasially branched 1 or 2 times, strigillose or puberulent; peduncles 26 cm; axes scorpioid; bracts linear-lanceolate, 1.53
mm. Flowers subsessile. Calyx glabrous; hypanthium portion
obconic to cylindrical, 1.72.2 mm; limb 0.61 mm, deeply
lobed; lobes triangular to ovate. Corolla white, shortly tubular,
outside puberulent to glabrous; tube 34 mm; lobes ca. 1 mm.
Capsules narrowly oblong to narrowly oblanceoloid, 2.54
1.52 mm, glabrous, valves 4, becoming twisted. Fl. and fr.
Sep.
Moist understories of forests, rice fields at forest margins. Guangxi, Guizhou, Yunnan [India].
See comments about the identity and circumscription of this species under Spiradiclis caespitosa.

2. Spiradiclis baishaiensis X. X. Chen & W. L. Sha, Bull. Bot.

Res., Harbin 8(3): 107. 1988.
bai se luo xu cao
Herbs, perennial, ascending, 2050 cm tall; stems usually
unbranched. Petiole 1.54 cm, yellow pilosulous or -villosulous; leaf blade drying papery, adaxially brown, abaxially strawyellow, ovate-elliptic, obovate, or oblanceolate, 710 36 cm,
adaxially glabrous, abaxially densely yellowish brown tomentose, base cuneate to acute and usually slightly inequilateral,
apex acute or acuminate; secondary veins 1123 pairs; stipules
persistent, ovate-lanceolate, 913 46 mm, obtuse. Inflorescence cymose to paniculiform, glabrous, purplish red; peduncles 59 cm; branched portion 58 cm, dichasial; bracts linearlanceolate, 815 1.52 mm; bracteoles linear, 35 mm; pedicels 34 mm. Calyx glabrous; hypanthium portion ovoid to
ellipsoid, ca. 2.5 mm; limb lobed to base; lobes linear, 34 mm.
Corolla purplish red, becoming brownish yellow when dry, tubular or tubular-funnelform, outside glabrous; tube 813 mm,
inside pubescent near middle or bearded in throat; lobes ovateoblong, ca. 2 mm. Capsules ellipsoid, ca. 3 1.5 mm. Fl. May.
Forests. Guangxi.
In the protologue this species was said to be similar to Spiradiclis
oblanceolata, although this species was classified in S. subg. Spiradiclis
while S. oblanceolata was classified in S. subg. Sinospiradiclis. The
protologue of S. baishaiensis apparently described short-styled flowers
(cf. f. 3, f. 4, and description of style as half as long as corolla and
anthers exserted); however, H. S. Lo (in FRPS 71(1): 106. 1999) reported this to be a description of a long-styled flower. The protologue
described the corollas as pubescent near the middle inside, while H. S.
Lo described the corollas as bearded in the throat.

3. Spiradiclis bifida Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 41: 310. 1872.
da ye luo xu cao
Herbs, to 0.5 m tall, apparently perennial; stems ascending, ?villous. Petiole 13 cm, pubescent; leaf blade drying
thinly papery, elliptic or elliptic-oblong, 1021 36.5 cm,
adaxially sparsely pilose to subglabrous, abaxially pubescent
along veins, base obtuse then decurrent, apex acuminate or subcaudate; secondary veins 1519 pairs; stipules subulate, caudate. Inflorescences paniculate, to 20 cm, ?villous; axes numerous, 12 cm, scorpioid; bracts minute. Flowers shorter than 5
mm. Capsules subglobose, ca. 2 mm in diam., valves 4.

333

Wet places in forests. S and SW Yunnan [NE India].

4. Spiradiclis caespitosa Blume, Bijdr. 975. 18261827.

luo xu cao
Herbs, perennial, usually rooting at basal nodes but ascending in upper parts; stems puberulent to glabrescent. Petiole 0.31 cm; leaf blade drying papery, elliptic to ellipticovate, 16.5 13 cm, puberulent to glabrescent on both surfaces, base cuneate to obtuse, apex obtuse to acute; secondary
veins 37 pairs; stipules deciduous or caducous, deeply 2lobed, lobes narrowly triangular to linear, 45 mm, puberulent
to glabrescent. Inflorescences cymose, puberulent to strigillose, unbranched or dichasially branched 1 time; peduncle 1
4.5 cm; axes scorpioid; bracts filiform. Flowers subsessile. Calyx glabrous to puberulent; hypanthium portion oblanceoloid to
cylindrical, 1.52 mm; limb 0.51 mm, deeply lobed; lobes triangular. Corolla white, shortly tubular, outside glabrous; tube
22.5 mm; lobes 11.5 mm, dorsally keeled. Capsules linearoblong to oblanceoloid, 56 1.51.7 mm, valves 2, becoming
twisted, sometimes later partially splitting.
Moist shady sites, often along streams; near sea level to 1200 m.
Perhaps present in China, no confirmed material seen [Indonesia
(Java)].
This species is here circumscribed more narrowly and somewhat
differently than done by H. S. Lo (in FRPS 71(1): 107110. 1999); it
has not yet been confirmed from China and is included here for comparison, based on the description of Bakhuizen f. (Fl. Java 2: 289. 1965)
who studied plants in the type region. Authors outside China have
separated the Himalayan plants with 4-valved capsules as Spiradiclis
cylindrica (Deb & Rout, Candollea 44: 225229. 1989; Springate &
Wright, Fl. Bhutan 2(2): 772774. 1999) and are followed here; H. S.
Lo et al. (Acta Bot. Austro Sin. 1: 31. 1983) synonymized this name
under S. caespitosa f. cylindrica.
H. S. Lo (Acta Bot. Austro Sin. 1: 3132. 1983) described several
forms of Spiradiclis caespitosa. In FRPS (H. S. Lo, loc. cit. 1999: 110),
f. subimmersa was distinguished by its narrowly lanceolate to lanceolate
leaves 37 0.71.5 cm. The plants included in f. subimmersa were
subsequently treated as a species, S. arunachalensis, by Deb and Rout
(loc. cit.: 225). These plants correspond to S. cylindrica as circumscribed here, rather than to S. caespitosa under which the varietal name
was actually published. Most of the specimens studied and described by
Deb and Rout were not seen by Lo, and similarly Los material was not
seen by them. Spiradiclis arunachalensis is provisionally treated here
pending further study.

5. Spiradiclis chuniana R. J. Wang, Novon 12: 423. 2002.

huan yong luo xu cao
Herbs, annual, to 8 cm tall, mostly unbranched, perhaps
ascending; stems densely villous. Petiole 13 cm, villous; leaf
blade ovate, 36.5 1.63 cm, sparsely villous on both surfaces, base broadly obtuse to truncate and often oblique, apex
rounded to obtuse then mucronate; secondary veins 1012 pairs;
stipules persistent, villous, 25-lobed, lobes linear, 510 mm.
Inflorescences cymose, several flowered, villosulous; peduncle
2.56 cm; branched portion ca. 1.5 cm; axes mixed dichasial
and scorpioid; bracts linear, 1.22 mm; pedicels to 5 mm. Flowers distylous, sessile to pedicellate. Calyx puberulent; hypanthium portion obconic to ellipsoid, ca. 1 mm; lobes narrowly
triangular, ca. 1 mm. Corolla white, funnelform, puberulent outside; tube ca. 7.5 mm, inside with villous ring at middle and

RUBIACEAE

334

sparsely pubescent above; lobes ca. 1.5 mm. Immature capsules

apparently subglobose. Fl. Aug.
Shady and wet places in forests on slopes of limestone hills; ca.
400 m. Guangxi (Nonggang).
This species was not classified to subgenus in the protologue
because the mature fruit were unknown, and predicting the states of
unknown characters is problematic in many Rubiaceae (e.g., Razafimandimbison & Taylor, Novon 10: 7173. 2000).

6. Spiradiclis coccinea H. S. Lo, Bull. Bot. Res., Harbin 6(4):

38. 1986.
hong hua luo xu cao
Herbs, erect, ca. 40 cm tall, presumably perennial; stems
grayish brown when dry, glabrous or subglabrous. Petiole 12
mm; leaf blade drying papery, adaxially grayish brown, abaxially grayish yellow, narrowly elliptic-oblong or elliptic-oblong,
4.59 1.33 cm, glabrous on both surfaces, base cuneate,
apex obtuse to acute; secondary veins 69 pairs; stipules triangular, rapidly narrowed to subulate, long acuminate apex. Inflorescence cymose, ca. 1 cm, with more than 10 flowers; peduncle very short. Flowers distylous. Calyx glabrescent; hypanthium portion obconic, 1.21.5 mm; lobes narrowly lanceolate,
1.72 mm, with gland at each side of base inside. Corolla dark
red, slenderly tubular-salverform, outside glabrous or puberulent; tube 1518 mm, inside with pilose ring above stamens;
lobes broadly ovate to suborbicular, 4.56 mm. Capsules subglobose, 4.55.5 mm in diam., valves 4. Fl. Aug.
On rocks in dense forests. Guangxi (Longzhou).

7. Spiradiclis cordata H. S. Lo & W. L. Sha, Acta Bot. Austro

Sin. 1: 34. 1983.
xin ye luo xu cao
Herbs, fleshy, low, apparently perennial, acaulescent or
stems short, densely hirsute. Leaves often clustered at base of
plant; petiole 17 cm, densely villosulous to hirsute; blade
drying papery, pale to green, elliptic-ovate to elliptic-oblong, 5
13 25.5 cm, adaxially sparsely hispidulous, abaxially hispidulous to villous with pubescence denser along veins, base
cordate to cordulate, apex obtuse to rounded; secondary veins
1519 pairs; stipules deciduous, hispidulous to villous, deeply
2-lobed, lobes linear, 48 mm. Inflorescences cymose to paniculate, puberulent to strigillose; peduncles 616 cm; principal
axes 49, 13 cm, mostly scorpioid; bracts linear or subulate,
ca. 2 mm. Flowers subsessile. Calyx densely puberulent to pilosulous; hypanthium portion obconic, 0.50.8 mm; lobes triangular, ca. 0.8 mm. Corolla white, tubular-funnelform, outside
puberulent to glabrescent; tube ca. 5 mm, villous above middle
inside; lobes subtriangular, ca. 1 mm. Capsules brown, subglobose, ca. 1.8 mm in diam., valves 4, flat. Fl. Aug, fr. AugNov.
Rocks at roadsides; ca. 400 m. Guangxi.

8. Spiradiclis corymbosa H. S. Lo, sp. nov.

mi hua luo xu cao
Type: China. Guangxi: Daxin, Taiping, 4 Jun 1977, D. L.
Chao 264 (holotype, GXMI).
Validating Latin description: that of Spiradiclis corymbosa W. L. Sha e [sic!] X. X. Chen (H. S. Lo, Bull. Bot. Res.,
Harbin 18: 276. 1998).

Herbs, 2430 cm tall, perhaps perennial, unbranched, perhaps ascending; stems densely pubescent. Leaves in slightly
unequal opposite pairs; petiole 0.52.5 cm, pubescent; blade
drying papery, elliptic to long elliptic, 6.514 24.7 cm, both
surfaces pilose along veins, base cuneate, apex acute; secondary
veins 1117 pairs; stipules long triangular, to 23 mm, ciliate,
apex subulate. Inflorescences corymbose, densely many flowered, densely pubescent; peduncles 4.59 cm; axes usually
scorpioid; bracts linear, 318 mm, ciliate; bracteoles linear, ca.
2 mm, ciliate; pedicels to 2 mm. Flowers sessile to pedicellate.
Calyx pubescent; hypanthium portion turbinate, ca. 1.5 mm;
lobes long triangular, ca. 1 mm, ciliate. Corolla red, pubescent
inside and out; tube 7.59 mm; lobes ovate, 1.52 mm. Capsules unknown. Fl. Jun.
Rocks in forests on limestone hills. Guangxi.
This name was previously published by H. S. Lo (loc. cit.) but not
validly so because the type was not indicated in accordance with Art.
37.6 and 37.7 of the Vienna Code. This species as circumscribed here
reportedly (H. S. Lo in FRPS 71(1): 100. 1999) comprises the Chinese
plants previously treated as Spiradiclis leptobotrya (Drake) Pitard var.
longiflora Merrill, in particular as treated under that name by H. S. Lo et
al. (Acta Bot. Austro Sin. 1: 29. 1983).

9. Spiradiclis cylindrica Wallich ex J. D. Hooker, Fl. Brit.

India 3: 76. 1880.
jian ye luo xu cao
Spiradiclis caespitosa Blume f. cylindrica (Wallich ex J.
D. Hooker) H. S. Lo.
Herbs, perennial, often prostrate and rooting on basal
nodes, at apices ascending; stems densely villosulous. Petiole
0.31 cm; leaf blade drying papery, elliptic, narrowly elliptic, or
lanceolate-elliptic, 28 14 cm, adaxially scaberulous to glabrescent, abaxially scaberulous, puberulent, or villosulous at
least along veins, base obtuse to acute, apex acute; secondary
veins 79 pairs; stipules caducous, strigillose to puberulent, 2
10 mm, deeply 2-lobed, lobes narrowly triangular to linear. Inflorescences cymose to paniculate, puberulent; peduncle 28
cm; principal axes dichasially branched 1 or 2 times, higher
order axes scorpioid; bracts linear, 15 mm. Flowers subsessile.
Calyx glabrous to puberulent; hypanthium portion obconic to
oblanceoloid, 1.52 mm; limb ca. 1 mm, lobed nearly to base;
lobes triangular. Corolla white, shortly tubular, outside glabrous; tube ca. 3.5 mm; lobes ca. 1 mm, smooth dorsally. Capsules linear-oblong to oblanceoloid, 44.5 11.7 mm, valves
4, becoming twisted.
Ravines in forests, rice fields at forest margins; 12001500 m.
Guangxi, Guizhou, Xizang, Yunnan [Bhutan, Myanmar, N Vietnam].
See comments regarding the circumscription and recognition of
this species under Spiradiclis caespitosa.

10. Spiradiclis emeiensis H. S. Lo, Acta Bot. Austro Sin. 1: 36.

1983.
e mei luo xu cao
Herbs, decumbent, rather fleshy, presumably perennial;
stems densely pubescent. Petiole 14 cm, densely pubescent;
leaf blade drying thinly papery, grayish green, ovate to elliptic,
49 24 cm, both surfaces sparsely pubescent or often densely

RUBIACEAE

so along abaxial veins, base cuneate then often decurrent, apex

acute to obtuse; secondary veins 912 pairs; stipules persistent,
deltoid, acuminate. Inflorescence cymose to subcapitate, villous; peduncle 0.10.5 cm; branched portion 12 cm, with axes
dichasial; bracteoles subulate, 1.52 mm. Flowers subsessile.
Calyx pilosulous to villous; hypanthium portion ellipsoid-cylindrical, ca. 1.5 mm; limb deeply lobed; lobes triangular-lanceolate, ca. 1 mm. Corolla white, urceolate-tubular, white villous
outside; tube 3.54 mm; lobes lanceolate to elliptic, 22.5 mm.
Capsules narrowly ellipsoid, ca. 6 3 mm, pilosulous to villous, valves 4, becoming slightly twisted. Fl. Jun, fr. Aug.
Dense forests. Sichuan, Yunnan.
This species was described based on fruiting material; later H. S.
Lo (Guihaia 11: 102. 1991) provided a detailed description of the inflorescences and flowers, which apparently represent the short-styled form
of a distylous species.

1a. Infructescence 12 cm, subsessile;

capsules villous ..................................... 10a. var. emeiensis
1b. Infructescence ca. 3.5 cm; peduncles
ca. 0.5 cm; capsules pilosulous ........ 10b. var. yunnanensis
10a. Spiradiclis emeiensis var. emeiensis
() e mei luo xu cao (yuan bian zhong)
Infructescence 12 cm, subsessile. Capsules villous.
Dense forests. Sichuan (Emei Shan).

10b. Spiradiclis emeiensis var. yunnanensis H. S. Lo, Guihaia

11: 102. 1991.
he kou luo xu cao
Infructescence ca. 3.5 cm; peduncles ca. 0.5 cm. Capsule
pilosulous.
Dense forests. Yunnan (Hekou).

11. Spiradiclis ferruginea D. Fang & D. H. Qin, Bull. Bot.

Res., Harbin 13: 334. 1993.
xiu jing luo xu cao
Herbs, prostrate to ascending, 320 cm tall, presumably
perennial; stems densely ferruginous pubescent. Petiole 0.55
cm, densely ferruginous pubescent; leaf blade drying papery,
ovate, ovate-elliptic, or rarely elliptic, 312 1.55 cm and
slightly anisophyllous, both surfaces densely pubescent, base
rounded and sometimes inequilateral, apex acuminate or obtuse; secondary veins 815 pairs; stipules ovate-lanceolate, 8
10 mm, sparsely pubescent, ciliate. Inflorescence cymose, 7
10-flowered, densely pubescent; peduncle 36 cm; bracteoles
lanceolate-linear, 57 mm; pedicels 17 mm. Flowers distylous,
pedicellate. Calyx pubescent; hypanthium portion turbinate, ca.
1 mm; lobes narrowly lanceolate, unequal, one lobe ca. 5 mm,
others ca. 4 mm, ciliate, with a gland in each sinus. Corolla pale
purplish red, tubular-funnelform, outside and inside pubescent;
tube ca. 16 mm; lobes ovate-triangular, ca. 3 mm. Capsules
unknown. Fl. Mar.
On calcareous rocks in forests; ca. 1200 m. Guangxi (Napo).

12. Spiradiclis fusca H. S. Lo, Guihaia 11: 100. 1991.

335

liang guang luo xu cao

Herbs, 3080 cm tall, perhaps perennial, perhaps ascending; stems glabrous, when dry brownish yellow. Petiole
0.51.5 cm; leaf blade drying thinly papery, adaxially grayish
brown, abaxially brownish yellow, oblong-lanceolate to subelliptic, 49 1.53.5 cm, adaxially glabrous, abaxially pilose
on principal veins, base cuneate and somewhat decurrent, apex
acuminate or subacute; secondary veins 1113 pairs; stipules
triangular, 67 mm, long acuminate, usually 2-parted. Inflorescences paniculiform, many flowered, together with peduncles
710 cm, glabrous; bracteoles subulate or linear, 23 mm;
pedicels short. Calyx glabrous; hypanthium portion obconic, ca.
1 mm; lobes sublanceolate, ca. 1.5 mm. Corolla white or pale
purple, funnelform, glabrous outside; tube ca. 9 mm; lobes
subtriangular, ca. 3 mm, apically rostrate-incurved. Capsules
unknown. Fl. Mar.
Limestone hills. Guangdong (Lianxian), Guangxi (Guilin).
The characters given in the protologue, in particular the proportional length of the calyx lobes to the corolla, do not seem to correspond
to the figure of this species presented by H. S. Lo in FRPS (71(1): 102,
t. 22, f. 610. 1999). This species was keyed by Lo in FRPS (loc. cit.:
87) as having pubescent stems and inflorescences, but the description
(loc. cit.: 99100) differed from that and so the species is here included
in a different section of the key.

13. Spiradiclis guangdongensis H. S. Lo, Acta Bot. Yunnan. 9:

299. 1987.
guang dong luo xu cao
Herbs, creeping, perhaps perennial, ascending at apices;
stems sparsely to densely pubescent. Petiole 26 mm, pubescent; leaf blade drying papery, leaden to pale gray, cordiformrounded to broadly ovate, 0.71.5(1.8) 0.51.2 cm, adaxially
sparsely hirsute, abaxially usually glabrous or subglabrous, base
cordulate to broadly obtuse, apex subacute; secondary veins 3
5 pairs; stipules deeply 2-lobed, glabrescent, lobes linear-subulate, 23 mm. Inflorescence 13-flowered, puberulent to pilosulous; peduncles short; bracteoles linear, ca. 1.7 mm; pedicels 12 mm. Flowers distylous, pedicellate. Calyx hispidulous
to glabrescent; hypanthium portion obconic, ca. 1 mm; lobes
oblong-lanceolate, ca. 2 mm, with gland in each sinus. Corolla
white, slenderly funnelform, subglabrous outside; tube ca.
11 mm, in throat densely villous; lobes subovate, ca. 4 mm,
sparsely pinnatinerved. Capsules subglobose-obconic, ca. 1.4
mm, persistent calyx lobes 34 mm, valves 4, straight. Fl. early
spring.
Dense forests, forest margins. Guangdong, Guangxi (Hechi).
As noted by H. S. Lo in the protologue, this species is very similar
to Ophiorrhiza exigua and, in fact, was confused with it in the original
description of that species.

14. Spiradiclis hainanensis H. S. Lo, Acta Bot. Yunnan. 9:

301. 1987.
hai nan luo xu cao
Herbs, creeping, perhaps perennial, ascending at apices;
stems densely villosulous. Petiole 0.10.5 mm, densely villosulous; leaf blade drying thinly papery, adaxially grayish black,

RUBIACEAE

336

abaxially pale gray, cordiform-orbicular to ovate, 0.61.3

0.60.9 cm, both surfaces hispidulous at least on principal veins
to glabrescent, base cordulate to subtruncate, apex obtuse; secondary veins 3 or 4 pairs; stipules deciduous, narrowly triangular, 1.52.5 mm glabrescent. Inflorescences cymose, 2- or
3-flowered, glabrescent, dichasial; peduncle 67 mm. Flowers
subsessile. Calyx densely villosulous (Hainan plants) to glabrescent (Yunnan plants); hypanthium portion subglobose to
oblanceoloid, ca. 1.5 mm; lobes narrowly lanceolate, 1.21.5
mm. Corolla white, funnelform, outside glabrous; tube ca. 8
mm; lobes lanceolate, 11.5 mm. Immature capsules ellipsoidovoid, ca. 2 mm, dehiscence unknown. Fl. Jul, young fr. Oct.
Dense evergreen forests; 21002600 m. Hainan (Sanya), Yunnan.
The description of the flowers and stipules here as well as the extension of the geographic range into Yunnan are taken from two specimens provisionally identified here as Spiradiclis hainanensis, Li Heng
12449 and 12511.

17. Spiradiclis loana R. J. Wang, Novon 12: 420. 2002.

xian rui luo xu cao
Herbs, erect, to 15 cm tall, acaulescent with leaves
crowded at base; stems pubescent. Petiole 14.5 cm, pubescent;
leaf blade oblanceolate, elliptic, or obovate, 612 1.23 cm,
adaxially pilose to hispid, abaxially glabrous and pubescent,
base acute to cuneate, apex acute to obtuse; secondary veins
ca. 10 pairs; stipules persistent, triangular to deeply 2-lobed,
segments 510 mm, pubescent. Inflorescences cymose, many
flowered, pubescent; peduncle 610 cm; bracts linear, ca. 1
mm. Flowers distylous, subsessile. Calyx with hypanthium portion obconic, ca. 1 mm; lobes triangular, ca. 1 mm, acute. Corolla white, tubular-funnelform, puberulent outside; tube 510
mm, inside with villous ring at middle and sparsely strigillose
above; lobes triangular, 11.2 mm, acute. Capsules subglobose,
ca. 2 mm in diam., valves 4. Fl. JulSep, fr. AugOct.
Forests on limestone mountains; 200300 m. Guangxi.

15. Spiradiclis howii H. S. Lo, Bull. Bot. Res., Harbin 6(4): 41.
1986.

18. Spiradiclis longibracteata S. Y. Liu & S. J. Wei, Acta

Phytotax. Sin. 32: 362. 1994.

kuan zhao luo xu cao

chang bao luo xu cao

Herbs, ascending, perennial, or subshrubs; stems subglabrous to densely pilose. Petiole 0.61.5 cm, densely pubescent;
leaf blade drying papery, adaxially grayish green, abaxially yellowish brown, lanceolate, ovate, subelliptic, or subelliptic-oblong, 2.56.5(9) 12(3) cm, adaxially strigose-hirsute, below villosulous along principal veins, base obtuse to sometimes
subrounded, apex acuminate and usually subfalcate; secondary
veins 68(10) pairs; stipules subulate-linear, 23 mm, acuminate. Inflorescence cymose, many flowered, densely pubescent;
peduncle 12.5 cm; bracteoles linear-lanceolate, 22.5 mm;
pedicels 12.5 mm. Flowers distylous, pedicellate. Calyx pilosulous; hypanthium portion subobconic, 0.70.8 mm; limb
deeply lobed; lobes narrowly lanceolate, rigid, 35 mm, equal
or slightly unequal. Corolla white becoming golden yellow
when dry, subtubular, outside with 5 hispidulous or hirsutulous
lines; tube 13.515.5 mm, inside with villous ring near middle
and pubescent above and through throat; lobes triangular-ovate,
ca. 1.5 mm, apically thickened-rostrate, dorsally narrowly
winged. Capsules subglobose, 3.54 mm in diam., pilosulous,
valves 4. Fl. Sep.

Herbs, presumably perennial, decumbent in lower part and

ascending above, sometimes subacaulescent with leaves clustered at base; stems densely villous. Petiole 622 mm; leaf
blade obovate-lanceolate or oblanceolate, 418 16 cm, adaxially sparsely pubescent, abaxially densely pilose along veins,
base acute, apex rounded, obtuse, or acute; secondary veins 18
22(29) pairs; stipules persistent, broadly ovate, 2- or 4-lobed.
Inflorescence cymose, densely many flowered, pubescent; peduncles 715 cm; bracts linear-lanceolate, 1015 mm. Flowers
distylous. Calyx pubescent; hypanthium portion ca. 1.5 mm;
lobes generally equal and 22.5 mm in long-styled form,
unequal and 310 mm in short-styled form. Corolla purplish
red, sparsely purplish red pubescent outside; tube 1618 mm in
long-styled form, ca. 22 mm in short-styled form, inside with
villous ring near middle and another at base; lobes ovate. Capsules globose, 2.53 mm, pilose. Fl. Apr, fr. AugOct.

Rocks in forests; 14001500 m. Yunnan.

Wet places in forests. Guangxi.

The markedly differing sizes and forms of the calyx and corollas
between the long-styled and short-styled flowers are notable.

19. Spiradiclis longipedunculata W. L. Sha & X. X. Chen,

Acta Bot. Austro Sin. 1: 35. 1983.

16. Spiradiclis laxiflora W. L. Sha & X. X. Chen, Acta Bot.

Austro Sin. 1: 35. 1983.

chang geng luo xu cao

shu hua luo xu cao

Herbs, unbranched, ascending, presumably perennial;

stems glabrous. Petiole 36 cm, glabrous; leaf blade drying
papery, adaxially dark green, and abaxially yellowish brown,
obovate or oblong-elliptic, 916 47.5 cm, glabrous on both
surfaces, base cuneate, apex acute; secondary veins 113 pairs;
stipules suborbicular or triangular-orbicular, 815 510 mm,
glabrous, cuspidate and 2-lobed, lobes 48 mm. Inflorescences
cymose-paniculate, ca. 34 cm, many flowered, glabrous; peduncles 1522 cm; bracts linear, ca. 4 mm. Calyx glabrous; hypanthium portion ovoid, ca. 1.5 mm; lobes ca. 0.3 mm. Corolla pale
yellow, tubular, outside glabrous; tube ca. 4 mm, bearded in
throat. Capsules unknown. Fl. Apr.

Herbs, perennial, erect or ascending, 1025 cm tall; stems

glabrous. Petiole 3.55 cm, glabrous; leaf blade drying papery,
adaxially green, abaxially pale, elliptic or obovate-elliptic, 10
15 35 cm, both surfaces glabrous, base acute and decurrent,
apex acute to acuminate; secondary veins 911 pairs; stipules
ovate-triangular, ca. 1 cm, 2-lobed, lobes aristate-acuminate. Inflorescences and flowers not seen. Infructescence paniculate,
ca. 25 cm; peduncle ca. 7 cm; axes dichasially branched 13
times then scorpioid; bracteoles lanceolate or linear-lanceolate,
517 mm. Capsules ellipsoid, ca. 3 1.5 mm, glabrous. Fr. Jun.
Sparse forests on limestone hills. Guangxi (Longzhou).

Guangxi (Daxin).

RUBIACEAE

20. Spiradiclis longzhouensis H. S. Lo, Guihaia 11: 99. 1991.

long zhou luo xu cao
Shrubs or perhaps subshrubs, robust; stems glabrous or
sparsely pubescent. Leaves clustered on upper parts of stems;
petiole 34.5 cm, turning black when dry; blade drying papery,
olive-green, abaxially pale, elliptic or elliptic-oblong, 1521
48.5 cm, glabrous, base usually decurrent, apex abruptly acuminate; secondary veins 1623 pairs; stipules subovate, ca. 10
mm, parallel veined. Inflorescence and flowers not seen. Infructescence ca. 15 cm, lax; bracteoles linear, 35 mm; pedicels 5
10 mm. Capsules ellipsoid, ca. 3.5 1.7 mm, glabrous, persistent calyx lobes linear, 22.5 mm. Fr. Apr.
On rocks in dense forests. Guangxi (Longzhou).

21. Spiradiclis luochengensis H. S. Lo & W. L. Sha, Acta Bot.

Austro Sin. 1: 34. 1983.
gui bei luo xu cao
Herbs, 20 cm or taller, presumably perennial, sometimes
subacaulescent with leaves grouped at base; stems glabrous.
Petiole 14 cm, glabrous, reddish brown when dry; leaf blade
drying papery, broadly obovate, 511 34.5 cm, both surfaces glabrous, base obtuse then decurrent, apex acute, obtuse,
or rounded; secondary veins 79 pairs; stipules long triangular,
glabrous, acuminate. Inflorescences cymose, many branched,
many flowered, glabrous; peduncles slender, 710 cm; pedicels short. Calyx purple, glabrous; hypanthium portion ca. 0.5
mm; lobes triangular-ovate, ca. 1 mm. Corolla white becoming
orange when dry, tubular-funnelform, outside glabrous; tube ca.
5 mm; lobes lanceolate-oblong, 22.5 mm. Capsules ovoid-globose, ca. 3 mm, valves 4. Fl. and fr. AprJun.
Shady places at streamsides. Guangxi.

22. Spiradiclis malipoensis H. S. Lo, Guihaia 11: 98. 1991.

dian nan luo xu cao
Herbs, presumably perennial, 2025 cm tall, decumbent
and rooting at base but ascending in upper parts; stems with 2
lines of dense hispidulous hairs. Petiole 0.51 cm, glabrous;
leaf blade drying papery, olive-green, lanceolate-oblong or
ovate-elliptic, 4.58 1.53.5 cm, glabrous, base cuneate to
obtuse and shortly decurrent, apex obtuse then abruptly acuminate; secondary veins 10 or 11 pairs; stipules lanceolate-linear
or linear, 58 mm, acuminate. Inflorescences and flowers not
seen. Infructescences cymose, 2.53 cm, glabrescent; peduncle ca. 1 cm; pedicels 34 mm. Capsules white, ellipsoid, 5.5
6 mm, glabrous, valves 4, becoming slightly twisted. Fr. Nov
Dec.
Dense forests; ca. 1100 m. Yunnan (Malipo).

23. Spiradiclis microcarpa H. S. Lo, Bull. Bot. Res., Harbin

18: 276. 1998.
xiao guo luo xu cao
Herbs, apparently ascending, presumably perennial, often
turning yellow when dry; stems glabrous. Petiole 34.5 cm, glabrous; leaf blade drying papery, adaxially leaden gray, obovate
or elliptic-obovate, 58 34 cm, both surfaces glabrous, base
obtuse then decurrent, apex obtuse to acute; secondary veins 6

337

or 7 pairs; stipules persistent, triangular to lanceolate, ca. 4 mm,

glabrous, obtuse to acute. Inflorescences and flowers not seen.
Infructescence paniculate, glabrous, branched dichasially to
several orders; peduncle ca. 10 cm; branched portion ca. 4
cm; bracts narrowly triangular, 25 mm. Capsules subsessile,
linear-oblong to oblanceoloid, ca. 2 0.50.6 mm, glabrous
or puberulent, valves 4, becoming twisted. Fr. Jul.
Shady places on limestone hills; ca. 100 m. Guangxi (Daxin).
The name Spiradiclis microcarpa was not validly published by
H. S. Lo (loc. cit.) because what should have been the indication of type
merely cited Guangxi: Daxin, circ. 100 m alt[.], non collecolector [sic!]
s. n. without using the word holotype or type or an equivalent and
without specifying the herbarium in which the type was conserved
(Vienna Code, Art. 37.6 and 37.7). It has not been possible to locate a
specimen from the gathering cited by Lo.

24. Spiradiclis microphylla H. S. Lo, Acta Bot. Austro Sin. 1:

34. 1983.
xiao ye luo xu cao
Herbs, ascending, presumably perennial; stems pubescent.
Petiole 0.41 cm, pubescent; leaf blade drying thinly papery,
brown, ovate, 13 0.51.4 cm, glabrous or pilose on both surfaces, base obtuse then decurrent, apex obtuse to acute; secondary veins 47 pairs; stipules subulate-lanceolate, acuminate or
caudate-acuminate. Inflorescences cymose, several flowered,
with axes scorpioid; peduncles 25 cm; bracts lanceolate or
subulate-lanceolate, 12 mm. Calyx pilose to puberulent; hypanthium portion ca. 0.8 mm; limb deeply lobed; lobes ca. 0.7
mm. Corolla white, shortly tubular, ca. 2.5 mm; lobes ovate-triangular, carinate on dorsal surface. Capsules subglobose, valves
4. Fl. SepOct.
Sparse forests. Jiangxi (Longnan).
This species was keyed by H. S. Lo (in FRPS 71(1): 88. 1999)
based on its glabrous stems and inflorescences, but the description (loc.
cit.: 101) reported it as pubescent. The key here is adjusted accordingly.

25. Spiradiclis napoensis D. Fang & Z. M. Xie, Acta Phytotax.

Sin. 40: 154. 2002.
na po luo xu cao
Herbs, ca. 65 cm tall, presumably perennial, procumbent
at base, ascending above; stems moderately to densely curved
pubescent. Leaves somewhat anisophyllous and clustered near
top of stem; petiole 0.72 cm, densely curved pubescent; blade
drying thinly papery to submembranous, adaxially dark green,
abaxially pale, narrowly ovate to broadly ovate, 47 1.53.5
cm, adaxially pilosulous to glabrescent, abaxially glabrescent or
curved pubescent along principal veins, base rounded to obtuse
then decurrent, apex acuminate; secondary veins 810 pairs;
stipules linear-subulate, 28 mm, glabrous, 2-lobed, lobes ca. 1
mm. Inflorescences cymose, 810-flowered, densely curved pubescent; peduncles 0.30.5 cm; axes 14 mm, puberulent; bracteoles linear, 610 mm, glabrous. Flowers distylous. Calyx puberulent to glabrescent; hypanthium portion ellipsoid, ca. 2
mm; limb deeply lobed; lobes narrowly triangular, ca. 2.5 mm,
with a gland in each sinus. Corolla in bud white tinged with
pink, subtubular, glabrous outside; tube ca. 12 mm, inside villous in throat; lobes subovate, ca. 5 mm, acute. Capsules unknown. Fl. AprMay.

RUBIACEAE

338

Forests in valleys of limestone hills; ca. 1000 m. Guangxi

(Napo).
Details of the long-styled and short-styled flowers are given in the
protologue.

26. Spiradiclis oblanceolata W. L. Sha & X. X. Chen, Acta

Bot. Austro Sin. 1: 35. 1983.
chang ye luo xu cao
Herbs, erect, unbranched, 1325 cm tall, presumably perennial; stems glabrous. Petiole 35 cm, glabrous; leaf blade
drying papery, oblanceolate or narrowly elliptic-oblong, 717
24 cm, glabrous on both surfaces, base cuneate, apex acute;
secondary veins 912 pairs; stipules long triangular, ca. 8 mm,
glabrous, 2-lobed, lobes ca. 3 mm. Inflorescences paniculate,
many flowered, ca. 20 cm, glabrous; peduncles 911 cm; bracts
linear, ca. 3 mm. Calyx with hypanthium portion ovoid, ca. 2
mm. Corolla white, outside glabrous; tube ca. 5 mm, bearded in
throat. Capsules ovoid, ca. 2 mm, glabrous. Fl. and fr. AprJun.
Rocks in forests. Guangxi.

27. Spiradiclis petrophila H. S. Lo, Acta Bot. Austro Sin. 1:

33. 1983.
shi sheng luo xu cao
Herbs, to 30 cm tall, apparently perennial, often procumbent at base, ascending above; stems densely pilosulous to tomentulose. Petiole 0.51.5 cm, densely pilosulous to tomentulose; leaf blade drying papery, ovate-elliptic to elliptic, 35
1.53 cm, densely tomentulose to pilosulous on both surfaces,
base cuneate then decurrent, usually slightly inequilateral, apex
acute; secondary veins ca. 9 pairs; stipules caducous, triangular,
1.52 mm, puberulent to glabrescent, acute and sometimes
erose. Inflorescences paniculate, many flowered, densely pilosulous to tomentulose; peduncles slender, 3.56.5 cm; axes
distally scorpioid, 11.5 cm; bracts narrowly triangular, 0.30.8
mm. Flowers sessile or subsessile, distylous. Calyx densely puberulent to hispidulous; hypanthium portion ellipsoid to obconic, ca. 0.8 mm; lobes triangular, ca. 0.3 mm. Corolla white,
tubular, outside puberulent to glabrescent; tube ca. 2 mm; lobes
triangular, ca. 1 mm, dorsally keeled, apically rostrate-inflexed.
Capsules subglobose to obovoid, ca. 2 mm, somewhat flattened
laterally, valves 2 then tardily splitting into 4. Fl. Sep, fr. Dec.
On limestone rocks and dripping cliffs. Guangdong (Yangchun).

28. Spiradiclis purpureocaerulea H. S. Lo, Bull. Bot. Res.,

Harbin 6(4): 39. 1986.
zi hua luo xu cao
Herbs, presumably perennial; stems densely brown pubescent. Leaves subsessile; blade drying papery, adaxially black,
abaxially pale, ovate, 36.5 1.53 cm, adaxially densely strigose-hispidulous, abaxially densely pubescent, base rounded or
obtuse, apex obtuse; secondary veins 710 pairs; stipules subulate, 23 mm, pubescent, caudate-acuminate. Inflorescence congested-cymose, 1.53 cm, densely pubescent; peduncles short.
Calyx pubescent; hypanthium portion obconic-globose, ca. 2
mm; lobes narrowly lanceolate, 44.5 mm, acuminate. Corolla
bluish purple, slenderly salverform; tube 1921 mm; lobes sub-

ovate, ca. 6 mm, obtuse. Capsules subglobose, 44.5 mm in

diam., pubescent, valves 4. Fl. Aug.
On rocks at roadsides. Guangxi (Longzhou).

29. Spiradiclis rubescens H. S. Lo, Guihaia 11: 101. 1991.

hong ye luo xu cao
Herbs, apparently ascending, low, ca. 6 cm tall, perhaps
annual; stems subglabrous, drying purplish black. Leaf blade
drying papery, purplish red, elliptic-oblong or oblong-ovate, 4
6 22.3 cm, glabrous on both surfaces, base obtuse, margins
ciliate, apex acute to obtuse; secondary veins 911 pairs; stipules triangular to broadly ovate, 45 mm, caudate-acuminate.
Inflorescence cymose, 35-flowered; bracteoles linear, 46 mm,
sparsely ciliate; pedicels short. Calyx glabrous; hypanthium
portion obovoid, ca. 2 mm; lobes narrowly lanceolate, unequal,
larger 44.5 mm, smaller ca. 3.5 mm, acuminate, ciliate, with
gland in each sinus. Corolla red, slenderly salverform-funnelform, outside glabrous, strongly 5-ribbed; tube ca. 21 mm;
lobes ovate, ca. 3.5 mm. Capsules unknown. Fl. Jun.
On rocks in shady places. Guangxi (Tiandong).
This species is noted for its reddened color when dry.

30. Spiradiclis scabrida D. Fang & D. H. Qin, Bull. Bot. Res.,

Harbin 13: 333. 1993.
cao bian luo xu cao
Herbs, perhaps perennial, 4050 cm tall, rooting near base,
apices apparently ascending; stems pubescent to glabrescent.
Leaves often slightly anisophyllous; petiole 0.20.5 cm, glabrous or usually pubescent; blade drying papery, adaxially
olive-green, abaxially yellowish green, ovate, narrowly ovate,
or lanceolate, 29 13.3 cm, glabrous or occasionally
sparsely strigillose or scabridulous at least on principal veins,
base obtuse, apex acuminate; secondary veins 69 pairs; stipules caducous to persistent, subtriangular, 0.71 mm, subglabrous, acute. Inflorescence corymbose-cymose, 324-flowered,
pubescent, puberulent, or glabrescent; peduncle 0.62.7 cm;
bracteoles linear, 25 mm; pedicels to 2 mm. Flowers sessile
to pedicellate. Calyx puberulent; hypanthium portion obovate,
11.5 mm; limb deeply lobed; lobes ovate-lanceolate, 11.5
mm, slightly unequal, with gland in each sinus. Corolla white
sometimes flushed with pink or purple, slenderly salverformfunnelform, glabrous inside and outside; tube 2526 mm;
lobes ovate, ca. 3.5 mm. Capsules subglobose, 34 mm in
diam., glabrescent, valves 4. Fl. Dec, fr. MarApr.
On rocks in forests; 8001200 m. Guangxi (Napo).

31. Spiradiclis spathulata X. X. Chen & C. C. Huang, Guihaia

13: 3. 1993.
chi ye luo xu cao
Herbs, low, erect, perhaps perennial, usually reduced or
acaulescent with leaves clustered at base. Petiole 58 mm; leaf
blade drying papery, adaxially olive-green, abaxially pale, spatulate or obovate-oblanceolate, 813 24.5 cm, adaxially
sparsely pubescent, abaxially densely pubescent at least along
principal veins, base acute to cuneate and usually decurrent,

RUBIACEAE

apex obtuse to rounded; secondary veins 1525 pairs; stipules

lanceolate-linear or linear, 79 mm, densely villosulous. Inflorescences cymose, 10- to many flowered, pubescent; peduncle
712 cm; bracteoles linear-lanceolate, 34 mm; pedicels 23
mm. Calyx pubescent; hypanthium obconic, 1.52 mm; limb
deeply lobed; lobes narrowly lanceolate, 11.3 mm. Corolla
purplish red, slenderly salverform, puberulent outside; tube 15
25 mm, inside with villous ring at middle; lobes subovate, 4
6 mm. Capsules unknown. Fl. May.
On rocks in shady places. Guangxi (Ningming).

32. Spiradiclis tomentosa D. Fang & D. H. Qin, Bull. Bot.

Res., Harbin 13: 330. 1993.
nian mao luo xu cao
Herbs, erect, 323 cm tall, presumably perennial, acaulescent with leaves clustered at base; stems densely gray-viscid
multicellular tomentose. Leaves slightly anisophyllous; petiole
0.54.5 cm, densely tomentose; blade drying thinly papery, oblanceolate, obovate, or rarely elliptic, 3.514 1.55 cm, both
surfaces densely tomentose, base acute, apex cuspidate to
rounded; secondary veins 710 pairs; stipules persistent, triangular, 822 mm, tomentose, narrowed to subulate acuminate
apex. Inflorescences paniculiform, 8- to many flowered, tomentose; peduncles 4.518 cm; bracts linear, 37 mm; pedicels 15
mm. Calyx tomentose; hypanthium portion turbinate, ca. 1 mm;
lobes linear, 1.52 mm. Corolla white, subfunnelform, outside
sparsely pubescent; tube 68 mm, glabrous inside; lobes ovate,
23 mm. Capsules subglobose, ca. 2 mm in diam., valves 4,
persistent calyx lobes 3.54 mm. Fl. Apr, fr. Jun.
Limestone caverns; ca. 500 m. Guangxi (Fengshan).
H. S. Lo (in FRPS 71(1): 99. 1999) described the leaves as cuspidate at apex, but the corresponding figure (p. 103, t. 23, f. 7) showed the
leaves to be apically rounded.

33. Spiradiclis umbelliformis H. S. Lo, Bull. Bot. Res., Harbin

6(4): 36. 1986.
san hua luo xu cao
Herbs, perhaps perennial, creeping or with upper parts perhaps ascending; stems densely reddish brown multicellular villosulous. Petiole 13 cm, reddish brown villosulous; leaf blade
drying papery, adaxially olive-green, abaxially pale, ovate-orbicular, 1.54 13 cm, adaxially glabrescent, abaxially pilosulous to pilose at least along principal veins, base cordulate to
truncate and usually decurrent, apex obtuse to rounded; secondary veins 46 pairs; stipules persistent, pilosulous to glabrescent, deeply 2-parted, lobes linear, to 10 mm. Inflorescences
cymose, umbelliform to subcapitate, 410-flowered, puberulent, pilosulous, or villosulous; peduncles 27 cm; bracts narrowly triangular. Flowers subsessile, distylous. Calyx subgla-

339

brous; hypanthium portion turbinate to subglobose, ca. 1.5 mm;

lobes ovate-triangular, ca. 0.6 mm, usually with gland in sinus.
Corolla white to pale purplish red, funnelform to tubular-funnelform, glabrous outside; tube 1718 mm, with villous ring
above base inside; lobes ovate, ca. 6 mm. Capsules subglobose,
33.5 mm in diam., glabrescent, valves 4, flat. Fl. Apr.
Rock crevices in forests, on limestone. Guangdong (Ruyuan),
Guangxi.

34. Spiradiclis villosa X. X. Chen & W. L. Sha, Bull. Bot.

Res., Harbin 11(3): 17. 1991.
mao luo xu cao
Herbs, 3050 cm tall, perennial, procumbent and rooting
below, perhaps upper parts ascending; stems densely dark
brown villous. Petiole 3.57 cm, densely villous; leaf blade
drying membranous, oblong-elliptic or obovate-elliptic, 1025
38 cm, adaxially dark brown pubescent, abaxially densely
brown villous, base cuneate or acute, apex acute or shortly acuminate; secondary veins 1416 pairs; stipules 2-lobed, densely
dark brown villous, lobes laciniate, 1530 mm. Inflorescences
and flowers not seen. Infructescences cymose-paniculate, many
flowered, densely dark brown villous; peduncles to 27 cm.
Young capsules ovoid, ca. 2 mm, villous, with calyx lobes ca. 4
mm, dehiscence unknown. Young fr. Jul.
On rocks in forests. Guangxi (Longzhou).
H. S. Lo (in FRPS 71(1): 101. 1999) described the stipules as to
30 mm, a relatively large size.

35. Spiradiclis xizangensis H. S. Lo, Bull. Bot. Res., Harbin

6(4): 43. 1986.
xi zang luo xu cao
Herbs, perhaps perennial, procumbent near base, ascending in upper parts; stems densely pubescent, drying purplish brown. Petiole 12 cm, densely pubescent; leaf blade
drying thinly papery, leaden gray with veins dark purple abaxially, ovate, 3.56 1.53.2 cm, adaxially glabrescent except
pubescent along midrib, abaxially pubescent along principal
veins, base obtuse or subrounded, apex acute or slightly obtuse;
secondary veins ca. 7 pairs; stipules sublanceolate, 22.5 mm,
subglabrous, often reflexed. Inflorescence cymose, 46-flowered; peduncle ca. 1 cm; bracteoles linear-lanceolate, ca. 7
mm; pedicels 25 mm. Calyx densely pubescent; hypanthium
portion subellipsoid, ca. 1.8 mm; lobes narrowly lanceolate,
unequal, larger one ca. 2 mm, smaller ones ca. 1.6 mm. Corolla
slenderly salverform-subfunnelform, outside pubescent; tube
ca. 17 mm, glabrescent inside; lobes triangular, ca. 3 mm. Capsules unknown. Fl. May.
Dense forests; 18002100 m. Xizang (Mdog).

89. TARENNA Gaertner, Fruct. Sem. Pl. 1: 139. 1788.

wu kou shu shu
Chen Tao (); Charlotte M. Taylor
Chomelia Linnaeus (1758), not Jacquin (1760, nom. cons.); Cupi Adanson; Webera Schreber.
Shrubs or trees, unarmed, often drying blackened. Raphides absent. Leaves opposite, sometimes with domatia; stipules persis-

340

RUBIACEAE

tent or occasionally tardily deciduous, interpetiolar or united around stem, triangular, often aristate. Inflorescences terminal, sometimes displaced to pseudoaxillary by subsequent growth, cymose to corymbiform, few to many flowered, sessile or pedunculate,
bracteate or bracts reduced. Flowers pedicellate or sessile, bisexual, monomorphic, often fragrant. Calyx limb 5-lobed. Corolla white,
pale green, or yellow, funnelform or salverform, inside glabrous or pubescent in throat; lobes 5, convolute in bud, usually strongly
reflexed at anthesis. Stamens 5, inserted in corolla throat, exserted; filaments short or reduced; anthers dorsifixed. Ovary 2-celled,
ovules 1 to numerous in each cell on axile placentas; stigma fusiform or linear, sulcate or striate, shortly 2-lobed at apex, exserted.
Fruit baccate, leathery or thinly fleshy, globose to ellipsoid, black or perhaps sometimes white, with calyx limb deciduous; seeds
several, medium-sized, plano-convex or concavo-concave, testa membranous, leathery, or crustaceous; endosperm fleshy or corneous; embryo small; cotyledon small, leaflike.
About 370 species: tropical and subtropical Africa, Asia, Madagascar, and Pacific islands; 18 species (12 endemic) in China.
The name Chomelia Linnaeus (1758) was applied to these plants for many years; however, the later homonym Chomelia Jacquin (1760), which
applies to a wholly neotropical genus, is now conserved against the Linnaean name, so the Old World plants formerly known under Chomelia are now
correctly known under Tarenna.
Most of the characters that distinguish species of Tarenna are found in the corollas and calyx limb, thus specimens of Tarenna are often difficult
to identify when they only have fruit from which the calyx limb has fallen. The genus was not well known in China until the work of W. C. Chen
(Acta Phytotax. Sin. 22: 139174. 1984). Tarenna apparently has secondary pollen presentation. W. C. Chen (in FRPS 71(1): 370. 1999) described the
stipules as caducous, but the specimens seen all have persistent or tardily deciduous stipules. The key here closely follows that of W. C. Chen in FRPS
(loc. cit. 1999: 370372), with its emphasis on pubescence characters and number of ovules per locule, for reference. Chen (loc. cit. 1999: 370384)
considered the form (i.e., raised vs. flat vs. impressed) of the leaf midrib adaxially to be consistent within a species, but specimens studied show
variation within species and overlapping among most species.

1a. Flowers with pedicels 1830 mm ..................................................................................................................................... 14. T. sinica

1b. Flowers subsessile or with pedicels to 13 mm.
2a. Calyx lobes subulate, 35 mm .............................................................................................................................. 7. T. lanceolata
2b. Calyx lobes triangular to narrowly triangular, 2 mm or shorter.
3a. Ovary with ovules 1 per cell; fruit with seeds 1 or 2 ....................................................................................... 2. T. attenuata
3b. Ovary with ovules 2 to many per cell; fruit with seeds 1 to many.
4a. Corolla tube longer than corolla lobes.
5a. Corolla villosulous to pilosulous outside; leaf blade densely villosulous to pilosulous ............ 17. T. yunnanensis
5b. Corolla glabrous outside; leaf blade glabrous or sparsely puberulent, strigillose, or
hirtellous abaxially.
6a. Calyx lobes triangular-lanceolate, ca. 1.5 mm, strigillose; leaf blade abaxially glabrous
or sparsely pilosulous or strigillose ........................................................................................... 1. T. acutisepala
6b. Calyx lobes triangular to broadly triangular, 0.51.5 mm, glabrous; leaf blade abaxially
glabrous or sparsely puberulent or strigillose.
7a. Leaves with secondary veins 1214 pairs ........................................................................... 5. T. foonchewii
7b. Leaves with secondary veins 69 pairs.
8a. Ovary with ovules 69 per cell; calyx with hypanthium portion sparsely
puberulent to hispidulous, lobes 0.81.5 mm; leaf blade drying papery or
membranous, with secondary veins 6 or 7 pairs .................................................... 3. T. austrosinensis
8b. Ovary with ovules ca. 17 per cell; calyx with hypanthium portion glabrous,
lobes 0.50.8 mm; leaf blade drying leathery, with secondary veins 79 pairs ...... 12. T. polysperma
4b. Corolla tube as long as or shorter than corolla lobes.
9a. Leaf blade glabrous on both surfaces.
10a. Branches becoming yellowish white or grayish white with age, sometimes resinous
at apex; stipules deciduous usually through fragmentation ................................................. 4. T. depauperata
10b. Branches green to brown or ashy gray, not evidently resinous at apex; stipules
persistent or deciduous through fragmentation.
11a. Leaf secondary veins 310 pairs; petioles 0.82 cm; inflorescences 49 49 cm;
leaf blade 4.515 1.56 cm.
12a. Corolla tube 22.5 mm ........................................................................................... 2. T. attenuata
12b. Corolla tube 57 mm ............................................................................................ 8. T. lancilimba
11b. Leaf secondary veins 713 pairs; petioles 13.5 cm; inflorescences 415 620 cm;
leaf blade 1325 512 cm.
13a. Leaf secondary veins 813 pairs; calyx mealy puberulent; inflorescences
subglabrous or puberulent, ca. 15 20 cm; ovary with 4 ovules per cell ..... 9. T. laticorymbosa
13b. Leaf secondary veins 7 or 8 pairs [or 6 in Japan]; calyx glabrous;
inflorescences glabrous, 410 615 cm; ovary with 2 ovules per cell ............. 18. T. zeylanica
9b. Leaves abaxially strigillose, hirtellous, hispidulous, pilosulous, or villosulous.

RUBIACEAE

341

14a. Leaves pubescent adaxially, at least on midrib.

15a. Corolla glabrous outside; fruit with seeds 16 ............................................................ 13. T. pubinervis
15b. Corolla strigillose or hirtellous outside; fruit with seeds ca. 30.
16a. Corolla tube 57 mm, lobes 1112 mm, much longer than tube; leaf
secondary veins widely spaced, 68 pairs ..................................................................... 10. T. laui
16b. Corolla tube 34 mm, lobes 45 mm, slightly longer than or nearly as
long as tube; leaf secondary veins well spaced to closely set, 512 pairs ......... 11. T. mollissima
14b. Leaves glabrous or glabrescent adaxially.
17a. Corolla tube pubescent outside; leaf blade lanceolate or lanceolate-oblong,
1229 4.510.5 cm, abaxially pubescent along principal veins .................................... 16. T. wangii
17b. Corolla tube glabrous outside; leaf blade narrowly elliptic, obovate, lanceolate,
or oblong-obovate, 616 1.57 cm, abaxially subglabrous to scabrous,
puberulent, or strigillose on blade and veins.
18a. Ovary with ovules 5 or 6 per cell; pedicels rather slender, 510 mm; fruit
ellipsoid; leaf blade abaxially sparsely strigillose ................................................. 6. T. gracilipes
18b. Ovary with ovules 2 per cell; pedicels rather thick, 47 mm; fruit subglobose;
leaf blade scattered subpapillose-pubescent, scabrous, or subglabrous ................... 15. T. tsangii
1. Tarenna acutisepala F. C. How ex W. C. Chen, Acta Phytotax. Sin. 24: 477. 1986.
jian e wu kou shu
Shrubs, 12.5 m tall; branches hispidulous or hirtellous becoming glabrescent, gray to reddish brown. Petiole 522 mm,
hispidulous; leaf blade drying papery or subleathery, ellipticoblong, lanceolate, oblong-ovate, or subovate, 419.5 1.55.6
cm, adaxially glabrous or sometimes sparsely puberulent along
midrib or sparsely strigillose throughout, abaxially strigillose to
hirtellous or sometimes glabrous, base cuneate, obtuse, or acute,
apex acuminate or acute; secondary veins 57 pairs; stipules
persistent, shortly united around stem, triangular, narrowly triangular, or lanceolate, 47 mm, hispidulous to glabrescent,
acute. Inflorescence corymbiform to congested-cymose, 2.53
34 cm, several to many flowered, strigillose, pedunculate;
peduncle 0.51 cm; bracts lanceolate to triangular, 1.52 mm;
pedicels 0.53 mm. Flowers subsessile to pedicellate. Calyx
strigillose; hypanthium portion ellipsoid to ovoid, 11.5 mm;
limb 1.52 mm, deeply lobed; lobes triangular-lanceolate, apex
acute. Corolla pale yellow, glabrous outside; tube ca. 10 mm,
villosulous inside upper part and throat; lobes narrowly elliptic,
ca. 4 mm. Ovules 1620 per cell. Berries subglobose, 57 mm
in diam., strigillose or glabrous; seeds 931. Fl. AprSep, fr.
MayNov.
Forests or thickets at streamsides in valleys or on mountain
slopes; 5001600 m. Fujian, Guangdong, Guangxi, Hubei, Hunan,
Jiangsu, Jiangxi, Sichuan.

2. Tarenna attenuata (J. D. Hooker) Hutchinson in Sargent, Pl.

Wilson. 3: 411. 1916.
jia gui wu kou shu
Webera attenuata J. D. Hooker, Fl. Brit. India 3: 104. 1880;
Ixora attenuata (J. D. Hooker) Kuntze; Tarenna sylvestris Hutchinson.
Shrubs or trees, 18 m tall; branches glabrous, flattened,
becoming ash gray to brown with age. Petiole 0.51.5 cm, glabrous; leaf blade drying papery or thinly leathery, blackish
brown, and somewhat shiny adaxially, oblong-lanceolate, ob-

long-obovate, lanceolate, or obovate, 4.515 1.56 cm, both

surfaces glabrous, base cuneate or acute, margins sometimes
thinly revolute, apex acuminate or abruptly shortly acuminate;
secondary veins 510 pairs, sometimes with pilosulous domatia; stipules persistent, shortly united around stem, narrowly triangular, 49 mm, glabrous, acuminate to cuspidate. Inflorescences congested-cymose, pyramidal to corymbiform, 2.55
46 cm, many flowered, puberulent to glabrescent, pedunculate; peduncle 0.31 cm; bracts narrowly triangular to subulate,
15 mm, acute; pedicels 0.55 mm. Flowers subsessile to pedicellate in dichotomous cymules. Calyx glabrous; hypanthium
portion subglobose to ellipsoid, 1.52 mm; limb 12 mm, lobed
for 1/41/2; lobes triangular, acute. Corolla white or pale yellow, outside glabrous; tube 22.5 mm, villosulous at throat;
lobes narrowly oblong to spatulate, 58 mm, acute to obtuse.
Ovules 1 per cell. Berries subglobose, 57 mm in diam., glabrous; seeds 2. Fl. AprNov, fr. MayJan.
Forests or thickets at streamsides, on hills or mountains, or in
fields; near sea level to 1200 m. Guangdong, Guangxi, Hainan, Yunnan
[Cambodia, India, Thailand, Vietnam].

3. Tarenna austrosinensis Chun & F. C. How ex W. C. Chen,

Acta Phytotax. Sin. 22: 145. 1984.
hua nan wu kou shu
Shrubs, ca. 2 m tall; branches terete to flattened, glabrous,
becoming pale gray. Petiole 515 mm, glabrous; leaf blade
drying papery or membranous, blackish brown, elliptic-oblong
or oblong-lanceolate, 515 24.5 cm, glabrous on both surfaces or puberulent to strigillose abaxially, base cuneate to
acute, apex acuminate; secondary veins 6 or 7 pairs, without
domatia; stipules generally deciduous leaving truncate persistent base, ovate to triangular, 35 mm, glabrous, acuminate. Inflorescences corymbiform to cymose, ca. 3 cm, few flowered,
strigillose; peduncle 45 mm; bracts linear to triangular, 13
mm, acute; pedicels 37 mm. Flowers pedicellate. Calyx with
hypanthium portion cupular to ellipsoid, 11.5 mm, sparsely
puberulent to hispidulous; limb 23 mm, glabrous, lobed for ca.
1/2; lobes triangular, apex acute. Corolla pale green, outside
glabrous; tube ca. 7 mm, villosulous inside and at throat; lobes

RUBIACEAE

342

ovate, 34 mm, apex acute. Ovules 69 per cell. Berries globose, 56 mm in diam.; seeds 614. Fl. AprMay, fr. AugSep.

logue did not describe the number of ovules per ovary locule, which is
not a significant character in Ixora and probably thus was not checked.

Forests on mountains; 8001300 m. Guangdong (Xinyi),

Guangxi (Jinxiu), Hunan (Yizhang).

6. Tarenna gracilipes (Hayata) Ohwi, Repert. Spec. Nov. Regni Veg. 36: 57. 1934.

4. Tarenna depauperata Hutchinson in Sargent, Pl. Wilson. 3:

411. 1916.

bao ye yu xin hua

bai pi wu kou shu

Shrubs or small trees, 16 m tall; branches glabrous,
blackened becoming yellowish white or grayish white with age.
Petiole 418 mm, glabrous; leaf blade drying papery or leathery, dark brown or sometimes yellowish brown abaxially, and
somewhat shiny adaxially, elliptic-obovate, elliptic, or subovate, 415 26.5 cm, both surfaces glabrous, base cuneate or
acute, apex shortly acuminate often abruptly acuminate with
tip often slightly obtuse; secondary veins 511 pairs, without
domatia; stipules deciduous usually through fragmentation,
shortly united around stem, triangular-ovate, 1.53 mm, glabrous, resinous, acute. Inflorescences corymbiform to pyramidal, 2.53 35 cm, few to many flowered, puberulent, hirtellous, or glabrescent, subsessile to pedunculate; peduncle to 1
cm; bracts triangular, 0.31.5 mm; pedicels 03 mm. Flowers
mixed sessile and pedicellate (or perhaps borne on expanded
axes) in dichotomous cymules. Calyx glabrous, puberulent, or
pilosulous; hypanthium portion cupuliform to ellipsoid, ca. 1
mm; limb ca. 1 mm, lobed for 1/23/4; lobes ovate or triangular, ciliolate, acute to rounded. Corolla white, outside glabrous;
tube 34 mm, villous inside and at throat; lobes narrowly oblong to spatulate, ca. 5 mm, acute to obtuse. Ovules 13 per cell.
Berries globose, 68 mm in diam., shiny, glabrescent; seeds 1
or 2. Fl. AprNov, fr. AprJan.
Forests or thickets at streamsides, on hills, or on mountain slopes;
2001700 m. Guangdong, Guangxi, Guizhou, Jiangsu, Yunnan [Vietnam].

5. Tarenna foonchewii (W. C. Ko) Tao Chen, comb. nov.

kuan zhao long chuan hua
Basionym: Ixora foonchewii W. C. Ko, Guihaia 19: 102.
1999.
Small trees, to 3 m tall; branches glabrous. Petiole 1217
mm, glabrous; leaf blade drying papery and olive-brown, elliptic, 1518 59 cm, glabrous, base obtuse, apex cuspidate or
shortly acuminate; secondary veins 12 or 13 pairs; stipules persistent, broadly triangular, 1214 mm, glabrous, long acuminate
to aristate. Inflorescence cymose, ca. 7 5.5 cm, many flowered; peduncle ca. 3 cm, sometimes subtended by or bearing
reduced leaves or leaflike bracts; bracts linear, 34 mm; pedicels ca. 1 mm. Flowers pedicellate. Calyx glabrous; hypanthium cupuliform, ca. 2 mm; limb deeply lobed; lobes triangular, ca. 1 mm. Corolla outside glabrous; tube ca. 13 mm;
lobes narrowly ligulate to narrowly elliptic, ca. 5 3 mm, obtuse. Fruit unknown. Fl. Apr.
Roadsides. Yunnan.
Based on the description of this species as having 5 corolla lobes,
fully exserted anthers, and subcapitate stigmas, as well as its general aspect, this species seems better placed in Tarenna than Ixora. The proto-

Chomelia gracilipes Hayata, Icon. Pl. Formosan. 9: 57.

1920; C. lancifolia Hayata; Tarenna hayataiana Kanehira; T.
lancifolia (Hayata) Kanehira & Sasaki.
Shrubs, to 3 m tall; branches slender, strigillose becoming
glabrescent and brown with age. Petiole 0.51.5 cm, strigillose;
leaf blade drying papery, obovate, lanceolate, or narrowly elliptic, 6.515 24.5 cm, glabrous adaxially, sparsely strigillose
abaxially at least along veins, base cuneate to acute, apex acute
to acuminate; secondary veins 6 or 7 pairs, without domatia;
stipules persistent, shortly united around stem, triangular, 1.53
mm, strigillose to glabrescent, acute to cuspidate. Inflorescences corymbose, 36 310 cm, trichotomous, strigillose,
subsessile to pedunculate; peduncle to 1 cm; bracts narrowly
triangular, 0.22 mm, acute; pedicels slender, 510 mm. Flowers pedicellate. Calyx glabrous to strigillose; hypanthium portion ellipsoid, 11.5 mm; limb 11.5 mm, lobed for 1/41/2;
lobes broadly triangular, acute. Corolla white, glabrous outside;
tube 46 mm, villous at throat; lobes narrowly spatulate-oblong, 910 mm, obtuse. Ovules 5 or 6 per cell. Berries ellipsoid,
58 mm in diam., glabrous. Fl. May, Jul, fr. Dec.
Forests on mountains; 100500 m. Taiwan [Japan].
The specimens studied are from the elevations given; W. C. Chen
(in FRPS 71(1): 378. 1999) gave the elevational range of this species as
up to 2500 m.

7. Tarenna lanceolata Chun & F. C. How ex W. C. Chen, Acta

Phytotax. Sin. 22: 139. 1984.
guang xi wu kou shu
Shrubs, 0.53 m tall; branches densely strigillose becoming glabrescent, often brownish red. Petiole 0.31.8 cm,
densely strigillose; leaf blade drying papery, lanceolate to oblanceolate, 532.5 15 cm, adaxially sparsely scabrous or
strigillose on lamina and densely strigillose on principal veins,
abaxially strigillose to pilosulous with pubescence denser on
principal veins, base acute or cuneate, apex long acuminate;
secondary veins 710 pairs, apparently without domatia; stipules generally persistent becoming brown and hardened, partially fused around stem, lanceolate to triangular or ovate, 34
mm, strigillose or glabrescent on margins, acute to cuspidate.
Inflorescences corymbiform and often nodding, ca. 2 3 cm,
several to many flowered, densely strigillose, sessile or pedunculate; peduncle to 5 mm; bracts linear to narrowly lanceolate,
210 mm; pedicels 13 mm. Flowers pedicellate. Calyx in bud
with hypanthium portion ellipsoid to ovoid, 1.52 mm, densely
pilosulous to puberulent; limb lobed nearly to base, densely
strigillose; lobes linear to narrowly lanceolate, 35 mm, sometimes unequal on an individual flower. Corolla in bud densely
strigillose outside. Berries subglobose, 36 mm in diam., puberulent, strigillose, or glabrescent; seeds 832. Fl. buds Oct,
fr. MayNov.

RUBIACEAE

Forests or thickets in valleys; 7001600 m. Guangxi, Guizhou,

Hunan (Dongkou).
W. C. Chen (in FRPS 71(1): 372. 1999) described the stipules as
610 mm, but the specimens studied (including several paratypes) all
have stipules 34 mm.

8. Tarenna lancilimba W. C. Chen, Acta Phytotax. Sin. 22:

141. 1984.
pi zhen ye wu kou shu
Tarenna attenuata (Voigt) Hutchinson var. puberula Chun
& F. C. How ex W. C. Ko, Fl. Hainan. 3: 584. 1974.
Shrubs or trees, 210(15) m tall; branches slightly compressed, glabrous or mealy pubescent, becoming brown with
age. Petiole 0.82 cm, glabrous; leaf blade drying thinly leathery, dark brown or yellowish abaxially, lanceolate, elliptic, or
obovate-oblong, 515 1.55 cm, both surfaces glabrous, base
cuneate then shortly decurrent, apex shortly acuminate; secondary veins 35 pairs, without domatia; stipules generally persistent, shortly united around stem, triangular, 1.53 mm, glabrous, acute to cuspidate. Inflorescence corymbose, trichotomous, 49 49 cm, many flowered, subglabrous or mealy
pubescent, subsessile; bracts triangular, 0.51.5 mm, ciliate;
pedicels 36 mm. Flowers pedicellate. Calyx glabrous or puberulent; hypanthium portion cupuliform, 11.5 mm; limb 1
1.5 mm, lobed for ca. 1/2; lobes triangular to spatulate, ciliolate.
Corolla white, outside glabrous, sparsely to densely villosulous
at throat and onto lobes; tube 57 mm; lobes ligulate-linear,
56 mm, obtuse. Ovules 2 per cell. Berry globose, 56 mm in
diam., glabrescent; seeds 24. Fl. AprJun, fr. JunJan.
Forests or thickets on hills; 1001000 m. Guangxi (Shangsi), Hainan [Vietnam].

9. Tarenna laticorymbosa Chun & F. C. How ex W. C. Chen,

Acta Phytotax. Sin. 22: 141. 1984.
kuan xu wu kou shu
Shrubs, height not noted; branches glabrous. Petiole 2.5
3.5 cm, glabrous; leaf blade drying membranous or papery and
grayish black, elliptic-oblong or obovate-oblong, 18.525 6
10 cm, glabrous on both surfaces, base cuneate, apex acuminate
or obtuse then mucronate; secondary veins 813 pairs; stipules
ovate-triangular, glabrous. Inflorescence corymbose with axes
spreading, ca. 15 20 cm, subglabrous or puberulent; pedicels
38 mm. Flowers pedicellate. Calyx mealy puberulent; hypanthium portion obconical; limb denticulate. Corolla glabrous outside, pilosulous in throat and onto bases of lobes; tube ca. 6.5
mm; lobes narrowly linear-oblong, ca. 7 mm, rounded. Ovules
4 per cell. Fruit unknown. Fl. May.
Yunnan (Hekou).

10. Tarenna laui Merrill, Lingnan Sci. J. 14: 59. 1935.

ya zhou wu kou shu
Shrubs, 23 m tall; branches densely grayish yellow hispidulous or strigillose becoming glabrescent with age. Petiole
623 mm, densely strigillose; leaf blade drying papery and dark
yellowish brown or blackish brown, oblong-elliptic or oblong-

343

lanceolate, 515 1.56 cm, sparsely hispidulous to strigillose

adaxially, sparsely to densely hispidulous or strigillose abaxially, base cuneate or acute, apex acuminate; secondary veins
68 pairs; stipules generally persistent, shortly united around
stem, broadly triangular, 1.53, densely strigillose, acute. Inflorescences cymose to corymbose and somewhat lax, 610 cm,
densely grayish yellow strigillose, pedunculate; peduncle ca.
1 cm; bracts triangular, 13 mm; pedicels 213 mm. Flowers
pedicellate. Calyx densely strigillose; hypanthium portion cylindrical, ellipsoid, or urceolate, 22.5 mm; limb 12 mm,
lobed for ca. 1/2; lobes ovate-triangular to triangular, apex
acute. Corolla white, outside densely strigillose; tube 57 mm,
villous in throat; lobes narrowly lanceolate to spatulate, 1112
mm, acute to obtuse. Ovules ca. 15 per cell. Berry subglobose, 56 mm in diam., strigillose. Fl. MayJul, fr. JulFeb.
Forests on mountains; ca. 700 m. Hainan (Sanya).

11. Tarenna mollissima (Hooker & Arnott) B. L. Robinson,

Proc. Amer. Acad. Arts 45: 405. 1910.
bai hua ku deng long
Cupia mollissima Hooker & Arnott, Bot. Beechey Voy.
192. 1833; Mussaenda kuliangensis F. P. Metcalf; Stylocoryna
mollissima (Hooker & Arnott) Walpers; Tarenna incana Diels.
Shrubs or small trees, 16 m tall; branches densely gray or
brown pilosulous or tomentulose, becoming glabrescent when
old. Petiole 0.42.5 cm, densely pilosulous or tomentulose; leaf
blade drying papery and blackish brown, lanceolate, oblonglanceolate, or ovate-elliptic, 4.522.5 110 cm, adaxially
moderately to densely hispidulous to pilosulous, abaxially
densely pilosulous, tomentulose, or villosulous, base cuneate,
acute, or obtuse, apex acuminate or long acuminate; secondary
veins 512 pairs, apparently without domatia; stipules generally
persistent, interpetiolar or shortly united around stem, ovate to
triangular, 58 mm, densely strigillose to pilosulous, acute to
cuspidate. Inflorescences corymbose, 48 cm, many flowered,
densely pilosulous to tomentulose, sessile and subtended by a
pair of somewhat reduced leaves; bracts linear, 13 mm; pedicels 36 mm. Flowers pedicellate. Calyx densely tomentulose
to pilosulous; hypanthium portion ellipsoid to subglobose, 1.5
2 mm; limb 11.8 mm, lobed shallowly to deeply; lobes triangular to spatulate, rounded to acute. Corolla white, outside
densely strigillose to hirtellous; tube 34 mm, densely villous at
throat; lobes narrowly oblong, 45 mm, obtuse to rounded.
Ovules ca. 15 per cell. Berry subglobose, 57 mm in diam., pilosulous, villosulous, or sometimes glabrescent; seeds 730. Fl.
MayJul, fr. MayJan.
Forests or thickets at streamsides, on hills, or on mountains; 200
1100 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan,
Jiangxi, Yunnan, Zhejiang [Vietnam].
W. C. Chen (in FRPS 71(1): 377. 1999) described the corollas as
ca. 1.2 cm, but this length has not been seen on the numerous specimens
studied.

12. Tarenna polysperma Chun & F. C. How ex W. C. Chen,

Acta Phytotax. Sin. 22: 144. 1984.
duo zi wu kou shu

RUBIACEAE

344

Shrubs or trees, 1.514 m tall; branches slightly compressed, glabrous, becoming brown when old. Petiole 515
mm, glabrous; leaf blade drying leathery and olive-greenish
brown, oblong-lanceolate or elliptic, 510.5 1.53.8 cm, glabrous on both surfaces, base cuneate, apex shortly acuminate or
abruptly acute; secondary veins 79 pairs, without domatia;
stipules deciduous usually through fragmentation often leaving
a persistent truncate base, interpetiolar, triangular to narrowly
triangular, 34 mm, glabrous, acute. Inflorescences corymbose
and trichotomous, 35 46 cm, 610-flowered, puberulent to
glabrescent, subsessile to pedunculate; peduncle to 1 cm; bracts
filiform, 12 mm, ciliolate; pedicel 3.56.5 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion turbinate, ca. 1
mm; limb 11.5 mm, lobed for ca. 1/2; lobes broadly triangular.
Corolla white, glabrous outside; tube ca. 8 mm, villous at
throat; lobes ovate to elliptic, ca. 4.5 mm, obtuse. Ovules ca. 17
per cell. Berry globose, ca. 7 mm in diam., glabrous; seeds 14
27. Fl. MarApr, fr. MayOct.
Forests on mountains; 9001000 m. Guangdong.
The protologue and W. C. Chen in FRPS (71(1): 382. 1999) described the secondary leaf veins as 79 pairs, but the paratypes seen all
have 5 or 6 pairs, which has complicated herbarium identifications of
this species.

13. Tarenna pubinervis Hutchinson in Sargent, Pl. Wilson. 3:

411. 1916.
dian nan wu kou shu
Shrubs or small trees, 0.56 m tall; branches puberulent to
pilosulous, becoming gray with age. Petiole 525 mm, strigillose to hirtellous; leaf blade drying papery or membranous,
blackish brown, oblong-elliptic, oblong-lanceolate, lanceolate,
or oblanceolate, 622 27.8 cm, strigillose to hirtellous on
both surfaces with pubescence denser on principal veins, base
acute to attenuate, apex caudate-acuminate or acuminate; secondary veins 710 pairs, apparently without domatia; stipules
generally persistent and becoming yellowed and hardened, interpetiolar or shortly united around stem, triangular to ovate, 3
9 mm, pilosulous to strigillose becoming glabrescent, long acuminate to aristate or cuspidate. Inflorescences cymose, ca. 3
3.5 cm, few flowered, densely strigillose, subsessile; bracts
linear, 1.52 mm; pedicels 12.5 mm. Flowers pedicellate. Calyx glabrous or sparsely pubescent; hypanthium portion obconic, 11.5 mm; limb lobed for ca. 1/2; lobes lanceolate, 1
1.75 mm, ciliate. Corolla pale green, glabrous outside; tube 45
mm, villosulous inside; lobes narrowly oblong, 56 mm, obtuse. Ovules ca. 3 per cell. Berry subglobose to ellipsoid, 510
mm in diam., puberulent to glabrescent; seeds 16. Fl. Mar
May, fr. JunJan.
Forests in valleys; 7002700 m. Guangxi, Sichuan, Yunnan.

14. Tarenna sinica W. C. Chen, Acta Phytotax. Sin. 22: 146.

1984.
chang geng wu kou shu
Shrubs, ca. 1.5 m tall; branches hispid. Petiole 0.51.5 cm,
hispid; leaf blade drying papery and blackish brown, narrowly
elliptic or elliptic-lanceolate, 612 24 cm, both surfaces

sparsely strigose except densely hispid-hirsute along principal

lateral veins, base cuneate, apex acuminate; secondary veins 6
8 pairs; stipules persistent becoming hardened and straw-yellow, subtriangular, ca. 2 mm, hispidulous, acute. Inflorescences
corymbiform, ca. 5 cm, few flowered, hispidulous, pedunculate;
peduncle 0.50.7 cm; bracts lanceolate, ca. 2 mm; pedicels 18
30 mm. Flowers pedicellate. Calyx hispidulous; hypanthium
portion ellipsoid, ca. 1.5 mm; limb lobed for ca. 1/2; lobes triangular, ca. 0.7 mm, acute. Corolla white, ca. 12 mm, hispidulous outside; tube densely villosulous inside and at throat; lobes
linear-lanceolate, longer than corolla tube. Ovules 1 per cell.
Berry unknown. Fl. Jun.
Forests on mountain slopes. Guangxi (Jingxi).

15. Tarenna tsangii Merrill, Lingnan Sci. J. 11: 59. 1932.

hai nan wu kou shu
Tarenna tsangii f. elliptica Chun & F. C. How ex W. C.
Ko.
Shrubs or trees, 16 m tall; branches flattened, shortly pilosulous, becoming brown or grayish brown with age. Petiole
515 mm, pilosulous; leaf blade drying papery and blackish
brown, oblong-obovate or lanceolate, 526 1.57 cm, adaxially glabrous, abaxially scabrous and sparsely puberulent to
subglabrous with pubescence denser along principal veins, base
cuneate, apex acuminate or shortly acuminate; secondary veins
47 pairs; stipules triangular, 45 mm, acuminate or apiculate.
Inflorescence corymbose, 47 ca. 6 cm, many flowered, gray
strigillose, pedunculate; pedicels 47 mm. Flowers pedicellate.
Calyx pilosulous; hypanthium portion cylindrical-urceolate, ca.
1.5 mm; limb ca. 1.5 mm, lobed for up to 1/3. Corolla white,
glabrous outside; tube 89 mm, pilosulous inside; lobes narrowly oblong, ca. 10 mm. Ovary with 2 ovules per cell. Berry
subglobose, 57 mm in diam., glabrous; seeds 4. Fl. MayJul,
fr. JulJan.
Forests on hills; 100800 m. Guangdong (Maoming), S Guangxi, Hainan.
Plants from Hainan (Lingshui) with leaf blade elliptic or ovateelliptic and 58 36 cm that fruit in Dec were described as Tarenna
tsangii f. elliptica. These features, however, do not clearly distinguish
the two groups of plants in Hainan, so this name is here synonymized.

16. Tarenna wangii Chun & F. C. How ex W. C. Chen, Acta

Phytotax. Sin. 22: 143. 1984.
chang ye wu kou shu
Shrubs, 25 m tall; branches gray, glabrous. Petiole 13
cm, subglabrous or puberulent; leaf blade drying papery or
membranous and brownish black, lanceolate or lanceolate-oblong, 1229 4.510.5 cm, glabrous on both surfaces or abaxially strigillose along principal veins, base attenuate, apex acuminate or shortly acuminate; secondary veins 911 pairs; stipules ovate-triangular, 34 mm. Inflorescences corymbose, ca. 5
5 cm, strigillose; pedicels 34 mm. Flowers pedicellate. Calyx pilosulous; hypanthium portion obconical; limb subtruncate
or denticulate. Corolla white, villous inside and onto lobes; tube
ca. 5 mm; lobes linear-oblong. Ovules 4 per cell. Berry subglobose, 810 mm in diam.; seeds 2 or 3. Fl. Jun, fr. AugNov.

RUBIACEAE

Forests on hills; 9001000 m. Yunnan.

345

18. Tarenna zeylanica Gaertner, Fruct. Sem. Pl. 1: 139. 1788.

17. Tarenna yunnanensis F. C. How ex W. C. Chen, Acta Phytotax. Sin. 22: 142. 1984.
yun nan wu kou shu
Shrubs or trees, to 3 m tall; branches densely yellowish
brown villosulous to tomentulose. Petiole 0.52.3 cm, densely
villosulous to tomentulose; leaf blade drying papery, elliptic,
obovate, elliptic-oblong, or oblanceolate-oblong, 1132 3.5
12 cm, adaxially subglabrous or sparsely hispidulous with pubescence denser along principal veins, abaxially densely villosulous to pilosulous, base cuneate, obtuse, or acute, apex
abruptly shortly acuminate; secondary veins 812 pairs, apparently without domatia; stipules generally persistent, shortly
united around stem, broadly triangular, 2.54 mm, densely villosulous, acute to cuspidate. Inflorescences cymose to corymbiform, 49 47 cm, many flowered, densely pilosulous, pedunculate; peduncle 13.5 cm; bracts linear to narrowly triangular, 14 mm; pedicels 29 mm. Flowers pedicellate. Calyx
densely pilosulous; hypanthium portion cupulate, ca. 1 mm;
limb 11.5 mm, lobed shallowly or for up to 1/2; lobes triangular. Corolla outside moderately to densely yellowish brown
villosulous to pilosulous; tube 56.5 mm, sparsely pilosulous at
throat; lobes narrowly oblong to spatulate, 44.5 mm, obtuse.
Ovules 35 per cell. Berry subglobose, ca. 5 mm in diam.,
yellowish brown villous. Fl. AprJun, fr. Jun.
Forests or thickets in valleys or at streamsides; 100200 m.
Yunnan.

xi lan yu xin hua

Chomelia kotoensis Hayata; Tarenna kotoensis (Hayata)
Masamune.
Shrubs, evergreen, to 2.5 m tall; branches glabrous, compressed to subquadrangular, green sometimes turning brown.
Petiole 12.5 cm, glabrous; leaf blade drying papery, oblongovate to oblong-obovate, elliptic, or elliptic-oblong, 1322 5
12 cm, glabrous on both surfaces, base cuneate to obtuse, apex
shortly cuspidate or acute; secondary veins 7 or 8 pairs, with pilosulous domatia; stipules generally deciduous through fragmentation often leaving persistent base, shortly united around
stem, triangular, 311 mm, glabrous, acute to acuminate. Inflorescences corymbiform to broadly pyramidal, 410 615 cm,
many flowered, glabrous, subsessile to pedunculate; peduncle
to 1 cm; bracts linear to narrowly triangular, 13 mm, acute;
pedicels 38 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion ellipsoid to cupular, 11.5 mm; limb 11.5 mm,
lobed for ca. 1/2; lobes triangular. Corolla white, glabrous outside; tube ca. 5 mm, villous in throat; lobes narrowly oblong to
spatulate, 712 mm, obtuse. Ovules 2 per cell. Berry globose,
ca. 10 mm in diam., glabrous; seeds 24. Fl. Feb, Apr, fr. Sep,
Dec.
Forests; below 100600 m. Taiwan [Japan, Sri Lanka].
W. C. Chen (in FRPS 71(1): 373. 1999) described the petioles as
up to 5 cm, but no petioles at all similar to this have been seen on specimens studied.

90. TARENNOIDEA Tirvengadum & Sastre, Mauritius Inst. Bull. 8(4): 90. 1979.
ling luo mai shu
Chen Tao (); Charlotte M. Taylor
Trees, unarmed. Raphides absent. Leaves opposite, usually with domatia; stipules caducous, interpetiolar or shortly united
around stem, triangular to ovate. Inflorescences terminal or sometimes pseudoaxillary or appearing leaf-opposed due to anisophylly
with 1 leaf caducous at subtending node, compound-cymose, several to many flowered, pedunculate or sessile and tripartite,
bracteate. Flowers pedicellate, bisexual, monomorphic. Calyx limb campanulate, truncate to 5-dentate. Corolla yellow to white, salverform, inside villosulous in throat; lobes 5, convolute in bud, reflexed at anthesis. Stamens 5, inserted in corolla throat, exserted;
filaments short; anthers dorsifixed. Ovary 2-celled, ovules 1 or usually 26 in each cell on axile placentas; stigma fusiform to cylindrical, exserted. Fruit black, baccate, subglobose, leathery or fleshy, with calyx limb deciduous; seeds 28, medium-sized, ellipsoid
to subglobose.
Two species: S and SE Asia; one species in China.

1. Tarennoidea wallichii (J. D. Hooker) Tirvengadum &

Sastre, Mauritius Inst. Bull. 8(4): 90. 1979.
ling luo mai
Randia wallichii J. D. Hooker, Fl. Brit. India 3: 113. 1880;
Aidia wallichii (J. D. Hooker) T. Yamazaki; Tarenna incerta
Koorders & Valeton; T. pallida (Franchet ex Brandis) Hutchinson; Webera cavaleriei H. Lveill; W. pallida Franchet ex
Brandis.
Trees, 320 m tall; branches rather stout, compressed to
angled, glabrous, with thin epidermis wrinkled and cracked,
coppery brown to reddish purple-brown, deciduous leaving

stems wrinkled to smooth. Petiole 13 cm, glabrous; leaf blade

drying leathery and often paler below, elliptic-oblong, oblanceolate-oblong, or elliptic-lanceolate, 730 2.99 cm, adaxially shiny and glabrous, abaxially puberulent to glabrous, base
cuneate to acute, margins often thinly revolute, apex obtuse to
acuminate with tip often ultimately obtuse; secondary veins
513 pairs, in abaxial axils with foveolate and/or pilosulous
domatia; stipules 410 mm, glabrous, acute to acuminate. Inflorescences 412 813 cm, densely hirtellous or pilosulous to
tomentulose; bracts lanceolate to spatulate, 13 mm, acute to
rounded; pedicels 15 mm. Calyx with ovary portion obconic,
ca. 1 mm, densely puberulent to tomentulose; limb 12.5 mm,
puberulent or tomentulose to glabrescent, with lobes linear to

RUBIACEAE

346

narrowly triangular, 0.50.7 mm, acute. Corolla yellow or white,

outside glabrous [to sericeous], inside pilosulous in throat with
pubescence sometimes extending onto lobes; tube 34 ca.
1.5 mm; lobes spatulate-oblong, 34 mm, obtuse to rounded.
Fruiting pedicels to 10 mm. Berry ovoid to subglobose, 718
mm in diam., glabrous; seeds ca. 5 mm. Fl. MarJun, fr. Jul
Feb.
Forests or thickets at streamsides in valleys or on hills or mountains; 4002200 m. Guangdong, Guangxi, Guizhou, Hainan, Yunnan

[Bangladesh, Bhutan, Cambodia, India, Indonesia, Malaysia, Myanmar,

Nepal, Philippines, Thailand, Vietnam].
Puff et al. (Rubiaceae of Thailand, 70. 2005) noted that the growth
form or architecture of this species is strongly sympodial: approached
a Terminalian branching pattern. The flowers on our specimens appear
to be strongly dichogamous, with the anthers fully developed and apparently dehiscing while the stigmas are still enclosed in the corolla tube;
then after the anthers have released all their pollen the styles apparently
elongate and the stigmas become fully exserted from the corolla.

91. THELIGONUM Linnaeus, Sp. Pl. 2: 993. 1753.

jia fan l shu
Chen Jiarui ( Chen Chia-jui); A. Michele Funston
Cynocrambe Gagnebin.
Herbs, annual or perennial, monoecious. Stems somewhat succulent. Raphides present. Leaves petiolate, somewhat succulent,
lower ones opposite, upper ones apparently alternate due to anisophylly, entire; stipules interpetiolar, membranous, united at base to
petioles. Flowers unisexual, rarely bisexual, axillary in upper nodes, small 13 flowered cymes, bracts reduced. Staminate flowers
sessile; calyx limb reduced; perianth (equivalent to corolla) deeply lobed, lobes 25, valvate, spreading or reflexed at anthesis, 35
parallel nerves; stamens (2)630, sometimes united basally in groups, anthers linear, dorsifixed at anthesis, free slender filaments,
pistillode reduced. Pistillate flowers subsessile; calyx limb reduced; perianth (equivalent to corolla) tubular, oblique, upper part an
elongate constricted mouth, 24 denticulate-lobed; staminodes reduced; ovary 1-celled; style simple, exserted; ovule solitary, basal.
Fruit a nutlike drupe, compressed; seed U-shaped; endosperm fleshy.
Four species: E Asia, Atlantic Ocean islands, Mediterranean region; three species (two endemic) in China.
Traditionally, Theligonaceae have been treated as a monogeneric family of controversial relationship: Rubiales (Cronquist), Myrtales (Engler),
Caryophyllales (Dalla Torre & Harms). However, recent reliable molecular data place it within a highly derived group of Rubiaceae (along with
Galium, Hedyotis, and Rubia; Pl. Syst. Evol. 225: 4375. 2000).

1a. Stamens 57; stem often ca. 10 cm .......................................................................................................................... 1. T. formosanum

1b. Stamens ca. 20; stem 1550 cm.
2a. Annual herbs; leaf blades 25 1.53 cm; stamens more than 20 .................................................................. 3. T. macranthum
2b. Perennial herbs; leaf blades 0.73 0.71.5 cm; stamens 2025 or 1620 ........................................................ 2. T. japonicum
1. Theligonum formosanum (Ohwi) Ohwi & T. S. Liu, Fl.
Taiwan 3: 904. 1977.
tai wan jia fan l
Cynocrambe formosana Ohwi, Acta Phytotax. Geobot. 2:
157. 1933.
Herbs, perennial, small, becoming black when dry. Stem
erect, 1015 cm, branched from base, creeping below, somewhat hairy. Leaf blade broadly ovate, 1.22 11.5 cm,
pilose adaxially, puberulent on veins abaxially, base subcordate, subtruncate, or rounded, decurrent to petiole, apex acute;
petiole 0.51.2 cm; stipules membranous, connate, deltoidovate, 23 mm, ciliate. Flowers white, unisexual. Staminate
flowers: perianth membranous, 3-parted, lobes oblanceolate,
2.53 mm, obtuse, reflexed, 35-veined; stamens 57. Pistillate flowers smaller; perianth 2-lobed, lobes obtuse. Fruit
obliquely obovoid, ca. 4 mm, hairy. Fl. MarJul, fr. JunAug.
Wet places in forests and along mountain trails; ca. 2700 m.
Taiwan (Pingdong).

2. Theligonum japonicum kubo & Makino, Bot. Mag. (Tokyo) 3: [5]. 1889.
ri ben jia fan l
Cynocrambe japonica (kubo & Makino) Makino.

Herbs, perennial, fetid. Stem erect or ascending, 1536 cm,

branched often from base, creeping below, pubescent above.
Leaf blade ovate or elliptic, 0.73 11.5 cm, pubescent on
both surfaces, base rounded or broadly cuneate, apex acute, lateral veins 3 each side; petiole 0.31.5 cm; stipules membranous, connate, ovate or deltoid-ovate, 23 mm, ciliate. Flowers
unisexual. Staminate flowers: perianth membranous, 3-parted,
lobes oblanceolate, 2.53 mm, obtuse, reflexed, 35-veined;
stamens 2025 or 1620, pendulous. Pistillate flowers: minute;
perianth 3- or 4-toothed. Fruit obliquely ovoid, ca. 5 mm, hairy.
Fl. MarJun, fr. JunAug.
Wet places in valleys, by streams; 9001200 m. Anhui, Shaanxi,
Zhejiang [Japan].

3. Theligonum macranthum Franchet, Nouv. Arch. Mus. Hist.

Nat., sr. 2, 10: 71. 1887.
jia fan l
Cynocrambe macrantha (Franchet) Poulsen.
Herbs, annual, fetid. Stem erect, 3050 cm, ferruginous
pubescent. Leaves herbaceous; leaf blade broadly ovate, ovatelanceolate, or suboblong, 25 13 cm, sparsely pubescent on
both surfaces, glabrescent, base subcuneate or subrounded,
somewhat decurrent to petiole, apex acuminate; petiole 0.5

RUBIACEAE

1.8 cm; stipules membranous, connate, ovate or deltoid-ovate,

46 mm, ciliate. Flowers unisexual. Staminate flowers: perianth
membranous, 2-lobed, tube ca. 2 mm, lobes 35 mm, lanceolate, apex obtuse, reflexed, 57-veined; stamens more than 20.

347

Pistillate flowers: smaller; perianth 2-lobed, apex obtuse. Fruit

ovoid, ca. 5 mm. Fl. AprJul, fr. JunAug.
Wet places in forests; 18002400 m. W Hubei, Sichuan, Zhejiang.

92. TIMONIUS Candolle, Prodr. 4: 461. 1830, nom. cons.

hai qian shu shu
Chen Tao (); Charlotte M. Taylor
Trees or shrubs, dioecious [or sometimes polygamo-dioecious], unarmed, terrestrial [or sometimes epiphytic]. Raphides absent.
Leaves opposite or verticillate, usually with domatia, notably leathery [to papery or succulent]; tertiary venation finely lineolate [or
sometimes not visible]; stipules caducous, interpetiolar [or calyptrate], triangular. Inflorescences axillary, cymose with axes dichotomous to scorpioid, few to several flowered with pistillate usually fewer flowered than staminate, pedunculate, bracteate or bracts
reduced; bracteoles sometimes fused in pairs. Flowers sessile, unisexual. Calyx limb cupular, truncate to 4- or 5-lobed. Corolla white
to pale yellow, funnelform or tubular-funnelform with pistillate frequently shorter and broader in diam. than staminate, outside
densely strigose to sericeous, inside glabrous, leathery; lobes 4(10), thinly imbricate in bud (i.e., subimbricate of Darwin 1993,
with margins thinly overlapping; see comment below). Stamens 4(10), inserted in corolla throat, exserted; filaments short; anthers
dorsifixed, at base 2-lobed. Ovary many celled, ovules 1 in each cell, pendulous from apical placentas; disk hirsute; style 412-parted
with segments linear, often unequal; stigmas 412 (i.e., 1 on each style arm), papillose, included or exserted. Fruit red or black,
drupaceous, thinly fleshy, ovoid, subglobose, or oblate (i.e., depressed globose), often 4- or 5-ridged when dry, with calyx limb persistent; pyrenes several to numerous [to 350 or more], 1-celled with 1 seed, narrowly oblong to cylindrical, bony; seeds cylindrical,
straight or curved; testa membranous; endosperm scanty or absent; embryo straight or curved; cotyledons minute, flattened; radicle
elongate, terete, ascending.
About 150180 species: tropical SE Asia and the Pacific islands; one species in China.
Darwin (Allertonia 7(1): 139. 1993) noted that although the corolla lobes have been commonly described as valvate in Timonius, in fact, they
are subimbricate with the margins only thinly overlapping; this arrangement is here called thinly imbricate. A notable characteristic of Timonius
within the Rubiaceae is the multiplication of ovules, or often entire files of ovules, as well as the number of cells in the ovary, from the 2 cells that are
typical of most Rubiaceae, to far past the 59 cells found in Guettarda. Darwin (loc. cit.; Syst. Bot. Monogr. 42: 186. 1994) has monographed part
but far from all of this large genus.

1. Timonius arboreus Elmer, Leafl. Philipp. Bot. 1: 72. 1906

[arborea].
hai qian shu
Trees 512 m tall; branches angled or flattened, rather
stout, glabrous or glabrescent, often markedly sylleptic from
nodes below leaves with markedly elongated lowermost internode, with scars of petioles often enlarged. Leaves opposite or
usually whorls of 3; petiole stout, 15 mm, glabrous; blade
drying stiffly papery, elliptic, elliptic-oblong, or broadly elliptic,
8.417 37.5 cm, adaxially glabrous, abaxially moderately
to sparsely strigillose, base cuneate to acute and often tapered,
apex acuminate; secondary veins 57 pairs, usually with densely
pilosulous domatia; stipules caducous often leaving a ring of
persistent trichomes 0.51 mm, triangular to lanceolate, 57
mm, abaxially moderately strigose, adaxially densely sericeous,
acute to acuminate. Inflorescences strigose to glabrous, stami-

nate 37-flowered, pistillate 1-flowered; peduncles 0.53 cm;

bracts lanceolate to triangular, 0.52 mm, usually fused in pairs.
Flowers subtended at immediate base by 2 to several persistent
bracteoles 0.51 mm, these sometimes shortly fused in pairs.
Calyx sparsely strigillose to glabrous; ovary portion subglobose, 2.53.5 mm in diam.; limb tubular, 12 mm, truncate or
undulate to 5- or 6-denticulate. Corolla outside densely velutinous-strigillose to sericeous; tube 47 mm, glabrous inside;
lobes 68, narrowly triangular, 47 mm, acute. Drupes subglobose to oblate, 810 1014 mm, glabrescent, color not noted;
pyrenes 50 or more, subcylindrical, 67 22.5 mm. Fl. Apr,
Nov, fr. AprJun.
Near sea level to 400 m. Taiwan [Philippines].
The pubescence of the lower leaf surface is nearly colorless and
arranged along the higher order veins; thus, it is often difficult to see,
even with good magnification, and has been overlooked by some
authors.

93. TRAILLIAEDOXA W. W. Smith & Forrest, Notes Roy. Bot. Gard. Edinburgh 10: 74. 1917.
ding qian shu
Chen Tao (); Charlotte M. Taylor
Shrubs, low, erect, unarmed or with spinescent branches; branches at base with a pair of reduced, usually deciduous leaves and a
persistent stipule. Raphides apparently absent. Leaves opposite, sometimes fasciculate on unexpanded axillary stems and appearing
whorled, subsessile, without domatia; stipules deciduous, interpetiolar or fused to petiole bases, 2-lobed, lobes gland-tipped. Inflorescences terminal, pseudoaxillary, or terminal on paired lateral short shoots and appearing axillary, congested-cymose to umbelliform
or fasciculate, few to many flowered, often nodding, sessile to pedunculate; bracts reduced or occasionally leaflike. Flowers pedi-

RUBIACEAE

348

cellate, bisexual, apparently monomorphic. Calyx limb 5-lobed essentially to base. Corolla white, pink, or pale yellow, salverform to
funnelform, glabrous inside; lobes 5, convolute in bud. Stamens 5, inserted in corolla throat, exserted; filaments short; anthers dorsifixed near base. Ovary 2-celled, ovules 1 in each cell, pendulous from apex; style curved; stigma clavate to ellipsoid and 2-lobed for
a third to half its length, exserted. Fruit schizocarpous, dry, oblanceoloid, with calyx limb persistent; mericarps 2, indehiscent, each
with 1 seed, ellipsoid, crustaceous; seeds medium-sized, ellipsoid; testa leathery; endosperm absent; embryo linear-oblanceolate.
One species: China.
The presence or absence of raphides has not been specifically noted anywhere we have seen; their absence is presumed here based on
observations of specimens and the classification by Robbrecht (Opera Bot. Belg. 1: 1271. 1988) of this genus in Antirrhoideae. W. C. Ko (in FRPS
71(2): 1. 1999) described the anthers as partially exserted, but these are fully exserted at anthesis on the specimens studied. On the few specimens
studied (T. T. Y 1348, C. Schneider 2194, Forrest 10713, all A; Boufford et al. 35041, MO), the flowers appear to be protandrous, with the stamens
apparently dehiscing while the stigmas are enclosed in the corolla; on these plants subsequently the style elongates, the stigma is exserted by several
millimeters, and then the two lobes separate and presumably become receptive.

1. Trailliaedoxa gracilis W. W. Smith & Forrest, Notes Roy.

Bot. Gard. Edinburgh 10: 75. 1917.
ding qian
Subshrubs, erect, 2045(60) cm tall, multibranched;
branchlets weakly angled to terete, densely strigillose to pilosulous becoming glabrescent. Leaf blade drying papery to stiffly
papery, obovate, elliptic, or oblanceolate, 0.51 0.30.4 cm,
adaxially glabrous or sparsely pilosulous to strigillose, abaxially
paler and glabrous or villosulous to strigillose or hispidulous at
least along midrib, base acute or very shortly and abruptly truncate, apex rounded or obtuse; secondary veins indistinct; stip-

ules 0.61 mm, pilosulous to glabrescent. Inflorescences 0.51

0.81 cm, 612-flowered, pilosulous to villosulous, sessile or
with peduncle to 5 mm; pedicels 12 mm. Calyx with ovary
portion cylindrical-lanceoloid, ca. 1 mm, densely pilosulous;
lobes narrowly elliptic or oblanceolate to linear, 0.81.5 mm,
glabrescent, acute. Corolla outside sparsely villosulous to glabrescent; tube 1.82.5 mm; lobes elliptic-spatulate, 11.5 mm,
obtuse to rounded. Schizocarps ca. 1.5 1 mm, densely villosulous. Fl. JulAug, fr. Aug.
On rocks or in thickets on mountain slopes in dry and warm
valleys; 14003000 m. Sichuan, Yunnan.

94. UNCARIA Schreber, Gen. Pl. 125. 1789, nom. cons.

gou teng shu
Chen Tao (); Charlotte M. Taylor
Agylophora Necker ex Rafinesque; Ourouparia Aublet.
Woody vines or lianas, climbing by recurved, usually paired spines generally 12 cm. Raphides absent. Leaves opposite, usually with domatia; stipules persistent or caducous, interpetiolar, generally ovate to ligulate in outline, entire to 2-lobed, usually
reflexed. Inflorescences axillary and sometimes also terminal, capitate with heads globose and 1 to several in cymes or fascicles,
pedunculate, bracteate; peduncles usually articulate in middle or upper portion, often with stipuliform bracts at articulation; bracts
enclosing heads sometimes caducous, involucral, stipuliform. Flowers sessile and bracteolate or pedicellate and bracteolate or
ebracteolate, bisexual, monomorphic. Calyx limb 5-lobed. Corolla white to yellow, salverform or funnelform, inside glabrous or
pubescent; lobes 5, imbricate in bud. Stamens 5, inserted in corolla tube near throat, exserted; filaments short; anthers dorsifixed.
Ovary 2-celled, ovules numerous in each cell on axile placentas attached in upper third of septum; stigma globose or clavate,
exserted. Fruiting heads globose, with fruiting pedicels when present often elongating notably. Fruit capsular, fusiform to obovoid,
loculicidally dehiscent into 2 valves that usually remain attached at both ends, thinly to thickly papery or cartilaginous, with calyx
limb persistent; seeds numerous, small to medium-sized, fusiform, flattened, winged with wing deeply bifid.
About 34 species: 29 in tropical Asia through Australia, three in Africa and Madagascar, two in tropical America; 12 species (five endemic) in
China.
Ridsdale (Blumea 24: 4346. 1978) presented an extensive consideration of the morphology, branching, and architecture of Uncaria. The characteristic hooked spines have been interpreted variously as modified plagiotropic shoots (Ridsdale, loc. cit.), peduncles that are modified into spines
(e.g., Steyermark in Lasser, Fl. Venezuela 9: 32. 1974), and short shoots modified into thorns (Robbrecht, Opera Bot. Belg. 1: 1271. 1988). By any
name, these structures function to support the plants as they climb and sometimes bear a terminal inflorescence. Ridsdale (loc. cit.: 69) described the
corolla lobe aestivation as valvate or thinly imbricate at their apices; other authors described them as imbricate. Uncaria was reviewed in detail for
China by How (Sunyatsenia 6: 231262. 1946), who emended the circumscriptions of several species, and then by Hsue and Wu (J. S. China Agric.
Coll. 2(8): 2132. 1981), who recognized ten species. The arrangement of the flowers and fruit, whether sessile or pedicellate, is taxonomically important; however, the pedicels usually elongate shortly before the flowers open and then continue to elongate as the fruit develop, often markedly, so the
pedicel length at anthesis may be difficult to discern from inflorescences in bud.
Uncaria is considered medicinally useful, with uses ranging from general tonics to supposedly curing HIV-AIDS (e.g., K. C. Hsia & X. M. Liu,
Acta Phytotax. Sin. 20: 319320. 1982). Various parts of the plants are apparently used, with the materials generally wild-collected. Uncaria gambir
(W. Hunter) Roxburgh, found from the Malay Peninsula through Borneo, is apparently both cultivated and wild-harvested as the source of gambir or
gambier (Ridsdale, loc. cit.: 82; Mabberley, Mabberleys Pl.-Book, ed. 3, 885886. 2008), a yellowish dry resin chewed together with the betel nut and
sometimes used in tanning. How (loc. cit.) noted that the Chinese drug Kou-Teng is derived from the hardened sterile peduncle with attached
portions of the stem of a species that is probably U. rhynchophylla.

RUBIACEAE

349

1a. Flowers subsessile to pedicellate, fruit pedicellate.

2a. Stipules suborbicular, 1416 mm, entire ........................................................................................................ 12. U. yunnanensis
2b. Stipules ovate, 612 mm, shallowly to deeply 2-lobed.
3a. Leaves drying papery, 28 cm wide; flowers subsessile; fruit 13 mm wide .................................................... 5. U. lanosa
3b. Leaves drying thinly leathery, 612 cm wide; flowers with well-developed pedicels; fruit
45 mm wide .............................................................................................................................................. 6. U. macrophylla
1b. Flowers and fruit sessile to subsessile.
4a. Leaf blade abaxially strigillose, strigose, puberulent, and/or hirsute on lamina (but sometimes with
different pubescence on veins); stems strigillose, tomentulose, strigose, hirtellous, hirsute, or pilosulous;
stipules 2-lobed.
5a. Flowering heads 1825 mm in diam. across calyces; stipule lobes ovate to lanceolate; calyx limb with
tube 24 mm, lobes 23 mm ............................................................................................................................... 1. U. hirsuta
5b. Flowering heads 912 mm in diam. across calyces; stipule lobes narrowly triangular, lanceolate, or
ovate; calyx limb deeply lobed, lobes 0.752 mm.
6a. Corolla lobes ca. 1.25 mm; leaves 2.54 cm wide ............................................................................... 2. U. homomalla
6b. Corolla lobes ca. 2 mm; leaves 35.5 cm wide ........................................................................................ 9. U. scandens
4b. Leaf blade abaxially glabrous to puberulent on lamina; stems glabrous, puberulent, sparsely hirsute, or
sparsely hirtellous; stipules entire or 2-lobed.
7a. Stipules entire or shallowly emarginate, broadly triangular, ovate, or suborbicular ...................................... 11. U. sinensis
7b. Stipules 2-lobed, elliptic-oblong, ligulate, lanceolate, or ovate, with lobes narrowly triangular, linear,
ovate, lanceolate, or triangular-ovate.
8a. Leaf blade drying thinly leathery; calyx lobes 0.10.3 mm.
9a. Leaf blade not glaucous abaxially, with tertiary venation mostly reticulate and not strongly
scalariform on adaxial surface; peduncles simple; corolla lobes externally glabrous; fruit
68 mm .............................................................................................................................................. 3. U. laevigata
9b. Leaf blade often glaucous abaxially, with tertiary venation rather evidently scalariform at
least on adaxial surface; peduncles simple or often branched; corolla lobes externally
sericeous; fruit 1014 mm ......................................................................................................... 10. U. sessilifructus
8b. Leaf blade drying thickly papery to thinly papery; calyx lobes 0.52 mm.
10a. Corollas 79 mm; leaf blade often drying reddish brown or dark red, often glaucous
abaxially; calyx limb ca. 1 mm ............................................................................................... 7. U. rhynchophylla
10b. Corollas 10.514.5 mm; leaf blade drying brown, grayish brown, or green, not glaucous
abaxially; calyx limb 1.54 mm.
11a. Calyx limb ca. 1.5 mm ............................................................................................. 8. U. rhynchophylloides
11b. Calyx limb 24 mm.
12a. Flowering heads 1315 mm across calyces; leaves glabrous ...................................... 4. U. lancifolia
12b. Flowering heads 1820 mm across calyces; leaves glabrous or usually
puberulent at least on veins abaxially ............................................................................... 5. U. lanosa
1. Uncaria hirsuta Haviland, J. Linn. Soc., Bot. 33: 88. 1897.
mao gou teng
Nauclea formosana Matsumura; Ourouparia hirsuta
(Haviland) Yamamoto; Uncaria formosana (Matsumura) Hayata; U. kawakamii Hayata; U. uraiensis Hayata.
Lianas, height unknown. Young stems slender, terete or
weakly 4-angled, sparsely to densely strigillose, strigose, or hirsute. Petiole 36 mm, strigose to strigillose; leaf blade drying
thickly papery to thinly leathery, ovate, lanceolate-oblong, or
elliptic, 812 3.57 cm, adaxially glabrous and smooth to
sparsely puberulent-scabrous on lamina and puberulent on
veins, abaxially sparsely to densely strigose with trichomes
generally produced regularly along and at 90 to tertiary veins,
apex acuminate; secondary veins 711 pairs, usually with hirtellous domatia; stipules generally deciduous, broadly ovate, 710
mm, 2-lobed for at least 2/3, strigillose, puberulent, or glabrescent, lobes ovate to lanceolate, ciliate, acute to acuminate. Inflorescences axillary or frequently paired and terminal on stems
(apparently due to terminal bud not developing further), pilose

to pilosulous; peduncle 2.55 cm, simple; bracts 510 mm;

flowering heads 1825 mm in diam. across calyces, 3040 mm
in diam. across corollas; bracteoles linear or spatulate, ca. 2
mm. Flowers subsessile. Calyx with hypanthium portion obconic, ca. 2 mm, densely pilose to strigose; limb densely strigillose, with tube 24 mm; lobes linear-oblong, 23 mm. Corolla
pale yellow or pale red, salverform, outside sparsely to densely
strigillose; tube 710.5 mm; lobes oblong to elliptic, 22.5 mm,
obtuse to rounded. Fruiting heads 3035 mm in diam. Fruit subsessile, obovoid, 59 mm (not including persistent calyx limb),
densely strigose; seeds ca. 1.5 mm. Fl. and fr. JanDec.
Thickets or forests at streamsides in valleys; below 100500 m.
Fujian, Guangdong, Guangxi, Guizhou, Taiwan.
H. H. Hsue and H. Wu (in FRPS 71(1): 257. 1999) listed Ourouparia setiloba Sakai as a synonym of this species, but that name is
correctly Ourouparia setiloba (Bentham) Sakai, which is a combination based on Uncaria setiloba, here treated as a synonym of U. lanosa
var. appendiculata. Hsue and Wu described the corolla tubes as 710
mm, but these are 810.5 mm on all the specimens studied and in descriptions of other authors.

RUBIACEAE

350

2. Uncaria homomalla Miquel, Fl. Ned. Ind. 2: 343. 1857.

bei yue gou teng
Uncaria tonkinensis Haviland.
Lianas, climbing to 25 m tall. Young stems quadrate,
sparsely to densely, usually ferruginous tomentulose or -hirtellous. Petiole 36 mm, hirtellous to strigose; leaf blade drying
papery, elliptic, lanceolate, elliptic-lanceolate, or ovate-lanceolate, 610 2.54 cm, adaxially rather sparsely but evenly puberulent to strigillose on lamina and densely tomentulose on
costa, abaxially moderately to densely hirsute or strigose with
trichomes produced mostly along veins, base rounded, apex
acuminate or caudate; secondary veins 68 pairs, usually with
well-developed pilosulous domatia; stipules generally deciduous, deeply 2-lobed, lobes narrowly triangular, 45 mm, acute.
Inflorescences axillary and sometimes also terminal, densely
hirtellous to strigose; peduncles simple, 2.53 cm, at articulation with narrowly triangular bracts 23 mm; flowering heads
710 mm in diam. across calyces, 2530 mm in diam. across
corollas; bracteoles apparently absent. Flowers sessile. Calyx
with hypanthium portion obconic, ca. 1.2 mm; limb deeply
lobed; lobes linear to narrowly triangular, 0.751 mm, densely
pilosulous, acute. Corolla yellow [to pale green], outside densely
strigose to strigillose; tube 58.5 mm; lobes ligulate to ellipticoblong, ca. 1.25 mm, obtuse to rounded. Fruiting heads 1220
mm in diam. Fruit sessile, obovoid, ca. 4 2 mm, strigose to
pilosulous; seeds 23 mm. Fl. Apr, May.
Evergreen to seasonal forests; 200600 m. Guangxi, Yunnan
[Bangladesh, Cambodia, India, Indonesia, Laos, Malaysia, Myanmar,
Thailand, Vietnam].
H. H. Hsue and H. Wu (in FRPS 71(1): 258. 1999) described the
leaves as up to 5.5 cm wide, but this has not been seen on specimens
studied nor reported by other authors. Ridsdale (Blumea 24: 95. 1978)
reported this species from Guangdong and Hainan but without vouchers.
The separation here of Uncaria homomalla and U. scandens follows
previous Chinese authors, but the separation of these species (or possibly the application of these names to the Chinese plants) needs further
study. Ridsdale (loc. cit.) separated U. homomalla from U. scandens
based on the size of the flowering heads and leaves; however, Chinese
specimens included previously in U. scandens (e.g., Henry 11868, MO!)
have flowering heads and leaves similar in size to those of U. homomalla.

3. Uncaria laevigata Wallich ex G. Don, Gen. Hist. 3: 470.

1834.
ping hua gou teng
Nauclea laevigata (Wallich ex G. Don) Walpers.
Lianas, height unknown. Young stems slender, quadrangular, glabrous [to puberulent]. Petiole 711 mm, glabrous; leaf
blade drying thinly leathery, elliptic, lanceolate, or elliptic-oblong, 8.512 36 cm, glabrous on both surfaces or puberulent
on veins abaxially, base rounded to cuneate, apex acute or acuminate; secondary veins 47 pairs, usually with hirtellous domatia; stipules caducous, elliptic-oblong to ovate, 46 mm, 1lobed for 1/2 or more, glabrous or sparsely puberulent, lobes
narrowly triangular, acute. Inflorescences axillary and sometimes in terminal groupings, glabrous; peduncles 2.56 cm,

simple; bracts 24 mm; flowering heads 810 mm in diam.

across calyces, 2535 mm in diam. across corollas; bracteoles
linear or subspatulate, 23 mm. Flowers sessile. Calyx with hypanthium portion obconic, 11.5 mm, densely sericeous or strigose; limb densely puberulent to strigillose, with tube 11.5
mm; lobes ligulate, 0.10.3 mm, obtuse. Corolla color unknown,
salverform, glabrous outside; tube 710 mm; lobes oblong, ca.
2 mm, obtuse to rounded. Fruiting head 2530 mm in diam.
Fruit sessile, fusiform, 68 mm, strigose to strigillose; seeds not
seen. Fl. and fr. MayNov.
Forests; 6001300 m. Guangxi, Taiwan, Yunnan [Bangladesh, India, Laos, Myanmar, Thailand, Vietnam].

4. Uncaria lancifolia Hutchinson in Sargent, Pl. Wilson. 3:

407. 1916.
dao gua jin gou
Large lianas, height unknown. Young stems quadrangular,
glabrous. Petiole 35 mm, glabrous; leaf blade drying papery,
oblong-lanceolate or ovate-lanceolate, 912 36 cm, glabrous
on both surfaces, base rounded to truncate or subcordate, apex
acute or acuminate; secondary veins 510 pairs, without domatia; stipules often persistent, ovate, 58 mm, 2-lobed for 1/3
1/2, glabrous, lobes lanceolate, acute. Inflorescences axillary
and terminal, glabrous; peduncles 47 cm, simple; bracts caducous, ca. 5 mm; flowering heads ca. 15 mm in diam. across calyces, ca. 45 mm in diam. across corollas; bracteoles linear or
linear-spatulate, 34.5 mm. Flowers sessile. Calyx with hypanthium portion obconic, ca. 3 mm, densely sericeous to pilose;
limb densely puberulent to strigillose, with tube 12 mm; lobes
spatulate, ca. 2 mm, obtuse to rounded. Corolla greenish white,
salverform, glabrous; tube 912 mm; lobes oblong, 1.52.5
mm, obtuse to rounded. Fruiting head 2535 mm in diam. Fruit
sessile, obovoid, 912 mm, sericeous to strigose; seeds 2.53
mm. Fl. and fr. JunDec.
Subtropical evergreen broad-leaved forests; 15001900 m. Yunnan [Vietnam].
How (Sunyatsenia 6: 252. 1946) first described the flowers of this
species. H. H. Hsue and H. Wu (in FRPS 71(1): 253. 1999) described
the leaves as having domatia; but the type specimen and several additional specimens studied lack these, and How did not mention them.

5. Uncaria lanosa Wallich var. appendiculata (Bentham)

Ridsdale, Blumea 24: 88. 1978.
heng chun gou teng
Uncaria appendiculata Bentham, London J. Bot. 2: 222.
1843; Ourouparia setiloba (Bentham) Sakai; U. lanosa f. setiloba (Bentham) Ridsdale; U. philippinensis Elmer; U. setiloba
Bentham.
Woody vines, height not noted. Young stems quadrangular,
sparsely hirsute to glabrescent. Petiole 410 mm, sparsely hirsute to glabrescent; leaf blade drying papery, ovate, lanceolate,
or lanceolate-oblong, 711 3.58 cm, adaxially shiny and glabrous except sparsely hirsute along veins, abaxially glabrescent
except sparsely hirsute along veins, base rounded, truncate, or
cordulate, apex acute to acuminate; secondary veins 69 pairs,
usually with domatia in axils of secondary and often also ter-

RUBIACEAE

tiary veins; stipules persistent or caducous, ovate, 610 810

mm, 2-lobed for 1/31/2, lobes narrowly triangular, acute to
acuminate. Inflorescences axillary and frequently also terminal,
strigose to hirsute or glabrescent; peduncle 2.54.5 cm; bracts
ca. 10 mm; flowering heads solitary (or 2), 1820 mm in diam.
across calyces, 3037 mm in diam. across corollas; bracteoles
apparently absent. Flowers sessile or subsessile. Calyx densely
strigillose; hypanthium portion fusiform, ca. 2 mm; limb deeply
lobed; lobes linear, ca. 2 mm, obtuse. Corolla color unknown,
salverform, externally sparsely sericeous to glabrescent; tube
ca. 12 mm; lobes oblong, ca. 2.5 mm, obtuse. Fruiting heads
3040 mm in diam.; pedicels 59 mm. Fruit pedicellate, fusiform, 927 13 mm, strigose; seeds 2.53 mm. Fl. Feb, perhaps Oct, fr. Feb.
Forests; ca. 300 m. S Taiwan [Indonesia (Sulawesi), Philippines].
Ridsdale (loc. cit.: 70) keyed Uncaria lanosa based on its stipules
that are subentire, but in our plants as well as the type of U. philippinensis, treated by him as a synonym of U. lanosa (NY, Web!), they
are markedly 2-lobed.
Uncaria lanosa var. lanosa occurs in Australia, Borneo, Indonesia, Malaysia, Myanmar, New Guinea, Pacific islands (Caroline
Islands, Palau, Solomon Islands), Philippines, and Thailand.

6. Uncaria macrophylla Wallich in Roxburgh, Fl. Ind. 2: 132.

1824.
da ye gou teng
Large lianas, height not noted. Young stems weakly to
markedly quadrangular, pilosulous, tomentulose, or glabrescent.
Petiole 325 mm, glabrous to densely tomentulose; leaf blade
drying thinly leathery and yellowish brown adaxially, ovate or
broadly elliptic, 1016 612 cm, adaxially glabrous except
strigillose along veins, abaxially sparsely to densely hirsutulous
with pubescence denser along veins, base rounded, subcordate,
or cordate, apex acute or shortly acuminate; secondary veins 6
9 pairs, usually with pubescent domatia; stipules caducous,
ovate, 612 615 mm, 2-lobed for 1/2 or 2/3, lobes triangular
to ovate. Inflorescences axillary, tomentulose to glabrescent;
peduncle 37 cm; bracts triangular, 68 mm; flowering heads
solitary, 1520 mm in diam. across calyces, 4050 mm in diam.
across corollas; bracteoles absent; pedicels 25 mm. Flowers
pedicellate. Calyx densely strigillose; hypanthium portion obconic, 22.5 mm; limb deeply lobed; lobes linear-oblong, 34
mm, obtuse to rounded. Corolla pale green to white, salverform,
outside strigillose to tomentulose; tube 910 mm; lobes oblong
to ligulate, ca. 2 mm, obtuse to rounded. Fruiting head 810 cm
in diam.; pedicels 618 mm. Fruit pedicellate, fusiform, 1420
45 mm, densely strigillose; seeds 68 mm. Fl. Jul, Sep, Dec,
fr. MarApr, SepNov.
On canopy crowns in secondary forests; 300900 m. Guangdong,
Guangxi, Hainan, Yunnan [Bangladesh, Bhutan, India, Laos, Myanmar,
N Thailand, Vietnam].

7. Uncaria rhynchophylla (Miquel) Miquel ex Haviland, J.

Linn. Soc., Bot. 33: 890. 1897.
gou teng
Nauclea rhynchophylla Miquel, Ann. Mus. Bot. Lugduno-

351

Batavi 3: 108. 1867; Ourouparia rhynchophylla (Miquel) Matsumura; Uncaria rhynchophylla var. koutong Yamazaki.
Lianas, height unknown. Young stems slender, weakly to
markedly quadrangular, glabrous, sometimes glaucous. Petiole
515 mm, glabrous; leaf blade drying papery and often redbrown or dark red, elliptic, lanceolate, or elliptic-oblong, 512
37 cm, glabrous on both surfaces, often glaucous abaxially,
base cuneate, obtuse, or rounded, apex acute to usually acuminate; secondary veins 48 pairs, sometimes with pilosulous
domatia; stipules often deciduous, lanceolate to ovate, 415
mm, 2-lobed for up to 4/5, glabrous, lobes linear to triangularlanceolate or ovate, acute to acuminate. Inflorescences axillary
and terminal, solitary or usually in terminal groups of 711, glabrous; peduncles 1.55 cm, simple; bracts 14 mm; flowering
heads 48 mm in diam. across calyces, 1215 mm in diam.
across corollas; bracteoles linear or linear-spatulate, ca. 2 mm.
Flowers sessile or subsessile. Calyx with hypanthium portion
obconic, ca. 1 mm, densely strigillose to strigose; limb densely
strigillose, ca. 1 mm, lobed for 1/2 or more; lobes triangular to
spatulate, acute to obtuse. Corolla color unknown, salverform,
outside puberulent to glabrous; tube 5.56 mm; lobes ovate,
1.52 mm, rounded to obtuse. Fruiting head 1020 mm in diam.
Fruit sessile or subsessile, obovoid to fusiform, 57 mm, strigillose to strigose; seeds 23 mm. Fl. and fr. MayDec.
Sparse forests or thickets at streamsides in valleys; near sea level
to 1000 m. Fujian, Guangdong, Guangxi, Guizhou, Hubei, Hunan,
Jiangxi, Yunnan, Zhejiang [Japan].
Ridsdale (Blumea 24: 93. 1978) included Uncaria rhynchophylloides as a synonym of U. rhynchophylla; however, these do appear to
be distinct.

8. Uncaria rhynchophylloides F. C. How, Sunyatsenia 6: 257.

1946.
hou gou teng
Lianas, to 13 m tall. Young stems quadrangular, puberulent to glabrescent. Petiole 57 mm, puberulent to glabrous; leaf
blade drying thickly papery and dark brown to grayish brown,
ovate, lanceolate, or elliptic-ovate, 59 2.54.5 cm, both surfaces puberulent to glabrescent, base cuneate to obtuse, rounded,
or cordulate, apex acuminate; secondary veins 57 pairs,
usually with pilosulous domatia; stipules generally persistent,
ovate, deeply 2-lobed, glabrous to puberulent, lobes triangular
to ovate, 34 mm, acute to acuminate. Inflorescences axillary
and often in terminal groups of 35, puberulent to glabrescent;
peduncles 3.56 cm, simple but sometimes with 2 articulations;
bracts 34 mm; flowering heads 911 mm in diam. across
calyces, 2530 mm in diam. across corollas; bracteoles linear or
linear-spatulate, 2.53.5 mm. Flowers sessile. Calyx with hypanthium portion obconic, ca. 1.5 mm, densely sericeous or
strigose; limb deeply lobed, densely strigillose; lobes oblong
to spatulate, ca. 1.5 mm, obtuse to rounded. Corolla color
unknown, salverform, outside puberulent; tube ca. 12 mm;
lobes obovate or oblong-obovate, 22.5 mm, rounded. Fruiting
head 1620 mm in diam. Fruit sessile, obovoid-ellipsoid, 810
33.5 mm, sericeous to strigillose; seeds not seen. Fl. and fr.
MayDec.
Forests, forest margins; 500800 m. Guangdong, Guangxi.

RUBIACEAE

352

This species was illustrated by How (loc. cit.: 257, f. 31). Ridsdale
(Blumea 24: 93. 1978) included Uncaria rhynchophylloides as a synonym of U. rhynchophylla; however, these do appear to be distinct.

9. Uncaria scandens (Smith) Hutchinson in Sargent, Pl. Wilson. 3: 406. 1916.

pan jing gou teng
Nauclea scandens Smith in Rees, Cycl. 24: Nauclea no. 9.
1813; Cephalanthus cavaleriei H. Lveill; Uncaria wangii F.
C. How.
Large lianas, height not noted. Young stems slender,
weakly to markedly quadrangular, densely hirtellous or pilosulous. Petiole 36 mm, hirtellous to pilosulous; leaf blade drying
papery, ovate, ovate-oblong, lanceolate, elliptic, or elliptic-oblong, 1015 35.5 cm, adaxially sparsely to moderately strigillose or scabrous-puberulent on lamina and densely puberulent to strigillose on veins, abaxially moderately to densely pilosulous, hirtellous, and/or strigillose usually with at least some
pubescence spreading, base rounded to truncate, subcordate, or
cordulate, apex acute or usually acuminate; secondary veins 7
10 pairs, usually with pilosulous domatia; stipules deciduous,
ovate, 610 mm, deeply 2-lobed, strigillose, puberulent, or glabrescent, lobes lanceolate, ovate, or narrowly triangular, acute.
Inflorescences axillary and often in terminal groups of 57
heads, densely pilosulous to tomentulose; peduncles 37 cm,
simple; bracts 49 mm; flowering heads 912 mm in diam.
across calyces, 2530 mm in diam. across corollas; bracteoles
linear or linear-spatulate, 13 mm, sparsely pubescent. Flowers
sessile. Calyx with hypanthium portion obconic, 11.5 mm,
densely strigose or pilose; limb 23 mm, lobed for up to 2/3,
densely grayish white strigillose; lobes linear or linear-spatulate, 1.52 mm, obtuse. Corolla pale yellow sometimes flushed
with pink, salverform, outside sparsely to densely hirtellous;
tube 810 mm; lobes obovate to elliptic, ca. 2 mm, rounded.
Fruiting head 2025 mm in diam. Fruit sessile, obovoid to oblanceoloid, 69 mm, hirtellous; seeds orange-yellow, ca. 2 mm.
Fl. Feb, Apr, fr. Jul, Nov.
Sparse forests, broad-leaved forests; 1001500 m. Guangdong,
Guangxi, Hainan, Sichuan, Xizang, Yunnan.
This species was illustrated by How (Sunyatsenia 6: t. 42. 1946,
as Uncaria wangii). See comments about the separation of this species
under U. homomalla.

10. Uncaria sessilifructus Roxburgh, Fl. Ind. 2: 130. 1824.

bai gou teng
Nauclea sessilifructus (Roxburgh) D. Dietrich.
Large lianas, height unknown. Young stems slender, quadrangular, sparsely puberulent or hirtellous to glabrous. Petiole
510 mm, glabrous; leaf blade drying thinly leathery, ovate,
elliptic, or elliptic-oblong, 812 46.5 cm, glabrous on both
surfaces or abaxially puberulent on principal veins, often glaucous abaxially, base cuneate to rounded, apex acute or acuminate; secondary veins 47 pairs, usually with pilosulous domatia; stipules deciduous, ligulate to lanceolate, 710 mm, 2-lobed
for 2/3 or more, glabrous to densely puberulent, lobes narrowly
triangular, acute. Inflorescences axillary or terminal, heads solitary or in groups of 515, strigillose to glabrescent; peduncles

35.5 cm, simple or often branched to 1 order; flowering heads

510 mm in diam. across calyces, 2535 mm in diam. across
corollas; bracteoles linear or subspatulate, 23 mm. Flowers
sessile. Calyx with hypanthium portion obconic, 1.52 mm,
densely strigose to sericeous; limb densely strigillose to pilosulous, with tube 12 mm; lobes oblong to triangular, 0.251 mm,
obtuse to acute. Corolla yellowish white, salverform; tube 610
mm, outside glabrous, pilosulous, or sericeous; lobes oblong,
ca. 2 mm, outside sericeous or hirtellous, rounded to obtuse.
Fruiting head 2535 mm in diam. Fruit sessile, fusiform, 1014
mm, sericeous; seeds not seen. Fl. and fr. MarDec.
Dense forests or thickets in valleys; 3001500 m. Guangxi, Yunnan [Bangladesh, Bhutan, India, Laos, Myanmar, Nepal, Vietnam].
This species was illustrated by Ridsdale (Blumea 24: 91, f. 11.
1978).

11. Uncaria sinensis (Oliver) Haviland, J. Linn. Soc., Bot. 33:

89. 1897.
hua gou teng
Nauclea sinensis Oliver, Hookers Icon. Pl. 20: t. 1956.
1891; Uncaria membranifolia F. C. How.
Lianas, height unknown. Young stems slender, quadrangular, glabrous. Petiole 610 mm, glabrous; leaf blade drying
thinly papery, elliptic to ovate, 914 58.5 cm, both surfaces
sparsely puberulent to glabrous, base obtuse to rounded, apex
acuminate; secondary veins 68 pairs, without domatia; stipules
often persistent, broadly triangular, ovate, or orbicular, 310
mm, glabrous, rounded to truncate or sometimes shallowly
emarginate. Inflorescences axillary, heads solitary or sometimes
in terminal groups of 35, glabrous; peduncles 37 cm, rather
slender; bracts 23 mm; flowering heads 1015 mm in diam.
across calyces, ca. 30 mm in diam. across corollas; bracteoles
linear or subspatulate, 23 mm. Flowers subsessile. Calyx with
hypanthium portion ca. 2 mm, densely strigose to strigillose;
limb deeply lobed, densely strigillose; lobes linear-oblong, 1
1.5 mm, obtuse to rounded. Corolla color unknown, salverform;
tube 78 mm, outside glabrous; lobes ligulate to triangular, ca.
2 mm, outside puberulent, acute to obtuse. Fruiting head 2030
mm in diam. Fruit sessile, ellipsoid, 810 mm, strigillose to strigose; seeds not seen. Fl. and fr. JunOct.
Sparse forests or wet secondary forests at middle elevations;
9001100 m. Gansu, Guangxi, Guizhou, Hubei, Hunan, Shaanxi, Sichuan, Yunnan.
This species was illustrated by How (Sunyatsenia 6: 254, f. 30.
1946, as Uncaria membranifolia).

12. Uncaria yunnanensis K. C. Hsia, Acta Phytotax. Sin. 20:

319. 1982.
yun nan gou teng
Woody vines or lianas, to 1525 m tall. Young branches
weakly quadrangular, ferruginous villous. Petiole glabrous or
ferruginous hirtellous; leaf blade drying leathery, ovate, elliptic,
or elliptic-oblong, 918 58 cm, both surfaces glabrous, base
obtuse, apex shortly acuminate; secondary veins ca. 4 pairs;
stipules caducous, suborbicular, 1416 mm, rounded. Inflorescences axillary and sometimes terminal, with heads solitary,

RUBIACEAE

brown hirtellous; peduncle 3035 mm; flowering heads 1520

mm in diam. in bud. Flowers not seen. Fruiting head ca. 40 mm
in diam.; pedicels 68 mm. Fruit pedicellate, fusiform, ca. 10
mm, brown hirtellous. Fl. Jul, fr. Jan.
Forest margins, thickets. Yunnan (Xishuangbanna).
H. H. Hsue and H. Wu (in FRPS 71(1): 249. 1999) described the

353

petioles as ferruginous pubescent, but the protologue description of

this species says petioli glabri nitiduli, though these petioli are described as structures belonging to pedunculus communis and may
be pedicels rather than petioles.
The possibility cannot be completely excluded based on the information available that this name will be found synonymous with
Uncaria gambir.

95. UROPHYLLUM Wallich in Roxburgh, Fl. Ind. 2: 184. 1824.

jian ye mu shu
Chen Tao (); Charlotte M. Taylor
Trees or shrubs, sometimes dioecious, unarmed. Raphides present. Leaves opposite, often distichous on horizontal branches,
apparently without domatia, with tertiary venation often regularly rectangular-areolate or clathrate and quaternary venation not
visible; stipules persistent or caducous, interpetiolar, narrowly triangular to linear [or sometimes reduced], acute. Inflorescences
axillary, capitate to cymose or often umbelliform, few to several flowered, pedunculate or sessile, bracteate or bracts reduced.
Flowers pedicellate and ebracteate, bisexual and apparently monomorphic or sometimes unisexual. Calyx limb cupular, (4 or)5(7)toothed. Corolla white to yellow, rotate, shortly tubular, or funnelform, inside villous in throat, leathery; lobes (4 or)5(7), valvate in
bud. Stamens (4 or)5(7), inserted in corolla throat, included or exserted; filaments short; anthers apparently dorsifixed. Ovary (4
or)5(7)-celled, ovules numerous in each cell on axile placentas attached at inner corner of cell; style often swollen at base; stigmas
38-lobed, exserted. Fruit white, orange, yellow, or red, baccate, fleshy, ellipsoid to subglobose, with calyx limb persistent; seeds
numerous, small, subglobose; testa crustaceous, areolate; embryo clavate.
About 150 species: widespread in tropical Asia; three species (two endemic) in China.
Although this genus is sometimes said to range into Africa, more recently the African species formerly included in Urophyllum have all been
moved to other genera (Lebrun & Stork, num. Pl. Fleurs Afr. Trop. 1249. 1997). This genus does not seem well known, apart from a consideration
of its delimitation in the Philippines by Bremekamp (J. Arnold Arbor. 21: 3241. 1940). Puff et al. (Rubiaceae of Thailand, 134. 2005) found
Urophyllum to be wholly dioecious in Thailand and noted that the staminate plants sometimes bear what they call sterile pseudo-fruit consisting of
[a] fleshy cuplike calyx and enlarged, empty ovaries [sic]. The reproductive biology of the Chinese species was not noted by H. S. Lo (in FRPS
71(1): 326329. 1999).

1a. Leaves at apex rounded then abruptly caudate with tips narrowly triangular to linear, 1520 mm; corolla
ca. 2 mm ................................................................................................................................................................... 2. U. parviflorum
1b. Leaves at apex acute or tapered to an acuminate apex, with tips narrowly triangular, 315 mm; corolla 34 mm.
2a. Stems flattened; stipules 1019 mm; corolla lobed for ca. 1/2 .............................................................................. 1. U. chinense
2b. Stems subterete; stipules 710 mm; corolla lobed for ca. 1/3 ............................................................................... 3. U. tsaianum
1. Urophyllum chinense Merrill & Chun, Sunyatsenia 2: 19.
1934.
jian ye mu
Shrubs or small trees, 1.54 m tall; branches compressed,
longitudinally sulcate on each interpetiolar side, ascending strigose to -sericeous, sometimes becoming glabrescent. Petiole
715 mm, strigillose to glabrescent; leaf blade drying papery,
olive-green, oblong-elliptic, lanceolate, oblong-lanceolate, or
rarely subovate, 820 2.56.5 cm, adaxially glabrous, abaxially sparsely to moderately strigillose with pubescence denser
on veins, base obtuse to acute, apex acute to acuminate, tip 3
20 mm; secondary veins 79 pairs, tertiary venation subclathrate; stipules persisting on 13 apical nodes, narrowly lanceolate or narrowly oblong, 1019 mm, densely strigose to sericeous, obtuse or acute. Inflorescences cymose, umbelliform, or
corymbiform, several to many flowered, strigose to strigillose;
peduncle 212 mm; bracts lanceolate, 23 mm, acute; pedicels
311 mm. Calyx strigillose or puberulent to glabrescent; hypanthium portion in staminate flowers obconic, ca. 0.5 mm, in pistillate flowers cupuliform, 1.52 mm; limb dentate, ca. 1 mm;
lobes triangular. Corolla white, 34 mm, outside glabrous, vil-

lous in throat, 5-parted for ca. 1/2; lobes ovate to subtriangular.

Berries red or orange-yellow, subglobose, ca. 8 mm in diam. Fl.
JunAug, fr. AugOct.
Thickets on mountains; 400900 m. Guangdong, Guangxi, Yunnan [N Vietnam].

2. Urophyllum parviflorum F. C. How ex H. S. Lo, Bull. Bot.

Res., Harbin 18: 278. 1998.
xiao hua jian ye mu
Shrubs or small trees; branches laterally compressed, often
longitudinally sulcate along interpetiolar sides, glabrous except
sparsely hispidulous on nodes. Petiole 58 mm, hispidulous in
lines; leaf blade drying papery, adaxially greenish gray, abaxially pale brown, elliptic-oblong, oblanceolate, or oblong-lanceolate, 1316 35.5 cm, adaxially glabrous, abaxially hispidulous along principal veins and strigillose to glabrescent on
higher order venation and blade, base acute to rounded, apex
rounded then abruptly caudate, tips narrowly triangular to linear, 1520 mm; secondary veins 710 pairs, tertiary veins generally parallel; stipules caducous or persisting on apical 1 or 2
nodes, narrowly lanceolate to narrowly ligulate, 713 mm,

RUBIACEAE

354

densely sericeous, acute to obtuse. Inflorescences corymbose to

umbelliform, several flowered, strigillose to hispid; peduncle 3
5 mm; bracts lanceolate-linear, ca. 2 mm; pedicels 39 mm. Calyx strigillose; hypanthium portion in pistillate flowers cupulate, ca. 1.5 mm; limb ca. 1 mm, shallowly 4- or 5-lobed; lobes
triangular. Corolla ca. 2 mm, lobed for ca. 1/2; lobes 4 or 5, triangular. Berries ellipsoid to subglobose, ca. 4 5.5 mm. Fl.
May, fr. Aug.
About 700 m. Yunnan (Jinping).

3. Urophyllum tsaianum F. C. How ex H. S. Lo, Bull. Bot.

Res., Harbin 18: 279. 1998.
dian nan jian ye mu
Shrubs or small trees, 26 m tall; branches subterete,

hispidulous. Petiole 58 mm, hispidulous; leaf blade drying

papery, pale brown or greenish gray, narrowly lanceolate, narrowly elliptic-oblong, elliptic-oblong, or lanceolate, slightly
inequilateral, 613.5 24 cm, adaxially glabrous, abaxially
strigillose, base obtuse or acute, apex acuminate; secondary
veins 69 pairs; stipules linear-lanceolate, 710 mm, densely
adpressed sericeous or -villous, acuminate. Inflorescences corymbiform to umbelliform, few to several flowered, hispidulous;
peduncle 510 mm; bracts subulate, ca. 2 mm; pedicels 36
mm, unequal. Calyx hispidulous to glabrous; hypanthium portion cupulate, ca. 2 mm; limb deeply lobed; lobes broadly triangular, ca. 0.6 mm. Corolla yellow or white, tubular, ca. 3 mm,
glabrous outside, lobed for ca. 1/3; lobes triangular. Berries red,
globose, ca. 5 mm in diam. Fl. Jul, fr. JanFeb.
Dense forests on mountains; 10001500 m. S Yunnan.

96. WENDLANDIA Bartling ex Candolle, Prodr. 4: 411. 1830, nom. cons.,

not Willdenow (1799).
shui jin shu shu
Chen Tao (); Charlotte M. Taylor
Shrubs or trees, unarmed; branches sometimes flattened. Raphides absent. Leaves opposite or occasionally in whorls of 3, without or rarely with domatia, with margins rarely denticulate (apparently where scabrous projections form triangular thickenings),
infrequently with venation not visible abaxially; stipules persistent or caducous as a whole or by fragmentation, interpetiolar, triangular to pandurate or leaflike (i.e., generally ovate and narrowed to a stipitate base), entire or rarely bilobed, erect and flat to longitudinally folded and/or spreading to reflexed. Inflorescences terminal, cymose, thyrsoid, or paniculiform, many flowered, sessile to
pedunculate, bracteate. Flowers sessile or pedicellate, bisexual, monomorphic, often fragrant. Calyx limb 5-lobed. Corolla white,
purple, or red, tubular, salverform, or funnelform, glabrous or pubescent inside, with top portion of tube often reflexed at anthesis;
lobes (4 or)5, imbricate in bud, strongly reflexed to revolute at anthesis. Stamens (4 or)5, inserted in corolla tube near throat, partially
to fully exserted; filaments short to developed; anthers dorsifixed, sometimes with connective prolonged in short apical and/or basal
appendages. Ovary 2(or 3)-celled, ovules numerous in each cell on peltate axile placentas; stigma bifid or rarely clavate and shortly
emarginate at apex (Wendlandia pendula), exserted. Fruit capsular, subglobose, loculicidally dehiscent across apical portion into 2
valves with valves later sometimes splitting septicidally, papery to woody, with calyx limb persistent; seeds numerous, small, compressed, sometimes narrowly winged; testa membranous, reticulate-striate; endosperm fleshy.
At least 90 species: mainly in tropical and subtropical Asia and a few in the Pacific region; 31 species (21 endemic) in China.
The flowers seem to open generally all at once on an individual plant and perhaps in the population. They are reported in several species to be
strongly fragrant. W. C. Chen (in FRPS 71(1): 191. 1999) described the fruit as rarely septicidal, but this has not been reported by other authors and
has not been seen on the specimens studied.
Wendlandia has been studied by several authors, notably in a comprehensive monograph by Cowan (Notes Roy. Bot. Gard. Edinburgh 16: 233
313, t. 232235. 1932, with supplemental notes shortly afterward; Notes Roy. Bot. Gard. Edinburgh 18: 183188. 1934) and then in treatments of the
Chinese species by F. C. How (Sunyatsenia 7(12): 3262. 1948) and later W. C. Chen (Acta Phytotax. Sin. 21: 277284, 386403. 1983).
Many Wendlandia species seem to be morphologically variable with much of the variation difficult to partition into distinct clusters, as discussed
by Cowan (loc. cit. 1932). Cowan also evaluated characters used by various authors to distinguish species and concluded that many were variable
and/or incorrectly interpreted, although he used several of these to distinguish infraspecific taxa. With more species known now, there also seems to be
variation within species in some of the characters that Cowan considered reliable to distinguish species. W. C. Chen (loc. cit. 1983) used Cowans
characters to delimit some of his new species, but circumscribed other species to include a relatively wide range of variation in some of the same
features (e.g., W. pingpienensis).
Cowan recognized four series and four subseries, distinguished by stigmas, anther, and stipule morphology as well as habit. These taxa were
accepted by W. C. Chen in FRPS (loc. cit. 1999: 195, 196, 200, 202, 208, 218, 221); however, the classification following Cowans treatment of several Chinese Wendlandia species described by recent authors is problematic because flowers are needed but are unknown for these. The key published
by W. C. Chen (loc. cit. 1999: 192195) closely follows that of Cowan except some leads were numbered incorrectly while others appear to be
missing, and the updated information on morphological variation and geographic distribution was not added to the key so could be problematic to use.
The key to Wendlandia species here follows that of W. C. Chen, to outline for reference the species distinctions in that treatment; however, it has been
augmented with the new morphological and distributional information.
In addition to the species treated here, F. C. How (loc. cit.: 43) reported Wendlandia ternifolia Cowan provisionally from China based on Tsang
21937, Liang 67941, and Zoo 69290. However, W. C. Chen in FRPS (loc. cit. 1999: 192) commented that study of Tsang 21937 showed the calyx to
have stiff pubescence, which is inconsistent with the protologue description of W. ternifolia, while the other two specimens cited have no flowers in

RUBIACEAE

355

adequate condition for identification. Therefore, W. ternifolia was excluded from the Chinese flora; no new or alternative identification was given by
Chen for these specimens.

1a. Most or all flowers with well-developed pedicels 26 mm; stipules triangular to broadly triangular, acute, cuspidate,
or narrowly spatulate.
2a. Shrubs or trees; petioles 0.51.5 cm; corolla lobes 12.5 mm; anthers 0.82 mm, subsessile or with filaments
less than 1 mm.
3a. Corolla with lobes ca. 1 mm, less than half as long as tube; anthers ca. 1 mm .......................................................... 12. W. laxa
3b. Corolla with lobes 1.52.5 mm, half as long as tube to equal to tube; anthers 1.22 mm.
4a. Leaves abaxially with secondary and higher order venation well marked; corolla white to yellow, with
tube 1.54 mm .................................................................................................................................................. 8. W. formosana
4b. Leaves abaxially with secondary and higher order venation hardly visible; corolla purplish red,
with tube 2.33 mm ............................................................................................................................... 16. W. longipedicellata
2b. Low shrubs or sometimes trees up to 12 m tall; petioles 0.10.6 cm; corolla lobes 3.58 mm; anthers
23.5 mm, exserted on filaments 46 mm (W. ser. Montigenae Cowan).
5a. Shrubs or trees 112 m tall; leaves elliptic-oblong, oblong-lanceolate, oblanceolate, or elliptic,
314.5 cm, glabrous on both surfaces; stipules spatulate to lanceolate or leaflike (i.e., ovate and
stipitate); Hainan ............................................................................................................................................... 18. W. merrilliana
5b. Leaves elliptic-lanceolate, lanceolate, narrowly lanceolate, ovate, or suborbicular, 0.83 cm,
pubescent on one or both surfaces; stipules triangular, lanceolate-triangular, or ovate; mainland.
6a. Leaves acute, shortly acuminate, or obtuse at apex, with secondary veins ca. 3 pairs, visible abaxially;
calyx lobes linear-oblong, spatulate, or oblanceolate ...................................................................................... 15. W. longidens
6b. Leaves obtuse at apex, veins not visible abaxially; calyx lobes triangular or lanceolate .............................. 28. W. subalpina
1b. Flowers sessile to shortly pedicellate, with pedicels up to 2 mm; stipules variously shaped, generally triangular,
spatulate, ovate, suborbicular, leaflike, or pandurate.
7a. Stipules triangular, ovate, broadly triangular, or leaflike, at apex acute, cuspidate, ligulate, or aristate, erect
or slightly spreading with age.
8a. Leaves subsessile or shortly petiolate, petioles up to 3 mm.
9a. Leaves opposite or ternate, ovate to ovate-lanceolate, 1.84 cm wide, drying papery, margins not
thickened nor edged with a vein; stigma clavate ............................................................................................... 22. W. pendula
9b. Leaves opposite, narrowly elliptic to narrowly lanceolate, 0.41.2 cm wide, drying thinly leathery,
marginally edged with a thickened vein; stigma deeply bilobed .................................................................... 25. W. salicifolia
8b. Leaves petiolate, petioles 325 mm.
10a. Stipule apex prolonged, ligulate, 57 mm .......................................................................................................... 9. W. grandis
10b. Stipule apex acute, cuspidate, or shortly acicular, 0.52 mm.
11a. Calyx lobes narrowly triangular to linear, 11.2 mm, equal or unequal on an individual flower with
at least one of lobes longer than 1 mm.
12a. Petioles 0.81.2 cm; stipules ca. 5 mm; corolla lobes ca. 1 mm ............................................................. 6. W. cavaleriei
12b. Petioles 0.81.5 cm; stipules 23 mm; corolla lobes 1.52.2 mm ......................................................... 13. W. ligustrina
11b. Calyx lobes lanceolate, triangular, or spatulate, 0.31.2 mm, equal or subequal.
13a. Leaves strigillose to puberulent on both surfaces; corolla tube 8.511 mm ............................................. 1. W. aberrans
13b. Leaves strigillose, puberulent, tomentulose, villous, or glabrescent on both surfaces; corolla tube
1.54 mm.
14a. Calyx lobes 0.30.5 mm ...................................................................................................................... 8. W. formosana
14b. Calyx lobes 0.81 mm.
15a. Inflorescences smaller, 56 36 cm; calyx glabrous or sparsely villosulous; leaves
sparsely pubescent to glabrous abaxially ................................................................................. 4. W. brevipaniculata
15b. Inflorescences larger, 917 622 cm; calyx glabrous to densely pilosulous, hirtellous,
or villosulous; leaves glabrous to strigillose, tomentose, or villous abaxially.
16a. Inflorescences 917 922 cm; corolla outside with tubes and lobes variously glabrous,
strigillose, and/or villosulous (including glabrous on tube and densely villosulous on
lobes in subsp. affinis) .................................................................................................................... 29. W. tinctoria
16b. Inflorescences 912 611 cm; corolla outside glabrous on tube and densely villosulous
or hirtellous on lobes .......................................................................................................................... 31. W. villosa
7b. Stipules triangular, ovate, suborbicular, leaflike, or pandurate, with apex obtuse, rounded, shortly
acuminate, 2-lobed, or ligulate and usually spreading to reflexed.
17a. Anthers linear-lanceolate, 1.32 mm, exserted from corolla on developed filaments; corolla 79.4 mm;
capsule 2.53 mm in diam. (W. ser. Euexsertae Cowan p.p., W. subser. Orbiculares Cowan) ....................... 27. W. speciosa

RUBIACEAE

356

17b. Anthers lanceolate or elliptic, 0.51.7 mm, partially included, sessile or with filaments less than 1 mm;
corolla 213.5 mm; capsule 12.5 mm in diam. (fruit unknown in W. augustinii, W. erythroxylon,
W. myriantha, W. parviflora, W. pubigera) (W. ser. Wendlandia p.p., W. subser. Paniculatae Cowan).
18a. Leaves abaxially densely strigose or sericeous along midrib; inflorescences smaller,
410.5 310 cm; corolla 11.513.5 mm, red or purple; capsule 22.5 mm in diam. ......................... 3. W. bouvardioides
18b. Leaves abaxially glabrous, glabrescent, or sparsely to densely strigillose, puberulent, pilosulous,
tomentose, hispidulous, or hirtellous; inflorescences often larger, 430 425 cm; corolla 26 mm,
white, pale green, pale yellow, or yellowish green; capsule 12 mm in diam. (unknown in
W. augustinii, W. erythroxylon, W. myriantha, W. parviflora, W. pubigera).
19a. Corolla tube 11.5 mm, with lobes equal to, longer than, or sometimes slightly shorter than tube;
leaves 517 28.5 cm.
20a. Leaves glabrescent or sparsely strigillose to pilosulous abaxially; Guangdong, Guangxi ...................... 5. W. brevituba
20b. Leaves densely ferruginous pubescent abaxially; Yunnan .................................................................... 21. W. parviflora
19b. Corolla tube 25 mm, with lobes clearly shorter than tube; leaves 326 1.514 cm.
21a. Leaves drying leathery, abaxially with secondary veins plane and tertiary venation hardly or
not visible.
22a. Calyx densely hirtellous ...................................................................................................................... 14. W. litseifolia
22b. Calyx glabrous to sparsely strigillose .................................................................................................. 20. W. oligantha
21b. Leaves drying papery to leathery, abaxially with secondary veins raised and tertiary venation
easily visible and usually also raised (i.e., most common condition).
23a. Calyx glabrous to sparsely puberulent, with lobes 1.22 mm, entire to ciliate.
24a. Calyx lobes 1.22 mm; corolla tube 23.5 mm; Guangdong, Hainan ................................. 10. W. guangdongensis
24b. Calyx lobes 1.21.5 mm; corolla tube 2.54 mm; Guangxi, Taiwan.
25a. Leaves entire or scabrous to denticulate marginally; stipules entire; Taiwan ........................... 7. W. erythroxylon
25b. Leaves entire, smooth or ciliolate marginally; stipules entire or 2-lobed; Guangxi ................... 19. W. myriantha
23b. Calyx with hypanthium portion and limb glabrous, glabrescent, or pilosulous, puberulent,
tomentose, hirtellous, pilose, strigose, or strigillose, with lobes 0.31 mm.
26a. Leaves abaxially moderately to densely hirtellous, strigillose, pilosulous, pilose, or
strigose with pubescence on lamina spreading and/or partly obscuring its surface.
27a. Leaves scaberulous on lamina adaxially; stipules usually equal to or only slightly
wider than stem; corolla inside with short trichomes confined to lower to upper
part of tube; Yunnan .......................................................................................................................... 26. W. scabra
27b. Leaves sparsely hirtellous, hispidulous or scaberulous adaxially; stipules usually twice
or more as wide as stem; corolla hirsute inside upper part of tube and throat ........................... 30. W. uvariifolia
26b. Leaves abaxially glabrous to sparsely strigillose or hispidulous on lamina and strigillose
to moderately hispidulous or hirtellous on principal veins.
28a. Leaves abaxially glabrous on lamina and sparsely strigillose and/or hirtellous on principal
veins; Taiwan .............................................................................................................................. 17. W. luzoniensis
28b. Leaves abaxially glabrous to hispidulous or strigillose; mainland.
29a. Stipules with apex ligulate, obtuse, often longitudinally folded .................................................... 9. W. grandis
29b. Stipules with apex oblanceolate to suborbicular, folded to usually flat.
30a. Corolla tube 45 mm; Yunnan.
31a. Stipules with apical portion 34 mm wide; leaves with secondary veins 79 pairs ........... 2. W. augustinii
31b. Stipules with apical part broad, ca. 10 mm wide; leaves with secondary veins
713 pairs ..................................................................................................................... 23. W. pingpienensis
30b. Corolla tube 23 mm.
32a. Leaves smaller, 710.5 24 cm, abaxially glabrous on lamina and sparsely
pubescent only on veins; secondary veins 7 or 8 pairs; Yunnan ................................. 11. W. jingdongensis
32b. Leaves larger, 14.518 67 cm, abaxially glabrous to sparsely pubescent on
lamina with pubescence denser along veins; secondary veins 10 or 11 pairs; Guangxi .... 24. W. pubigera
1. Wendlandia aberrans F. C. How, Sunyatsenia 7(12): 44.
1948.
guang xi shui jin shu
Shrubs, 13 m tall; branches terete, yellowish brown,
densely adpressed ferruginous hirsute. Leaves opposite; petiole
310 mm, appressed yellowish brown pubescent; blade drying
papery and reddish yellow, oblong-elliptic or ovate-elliptic, 5

16 25.8 cm, adaxially adpressed puberulent with pubescence

denser along midrib, abaxially sparsely puberulent with pubescence denser along principal veins, base cuneate or acute, apex
shortly caudate-acuminate or acute; secondary veins 612 pairs;
stipules late deciduous to persistent, triangular, 35 mm, yellowish brown pubescent becoming glabrescent, cuspidate. Inflorescences congested-paniculate, narrowly pyramidal in outline, 510 38 cm, appressed ferruginous pubescent, tripartite

RUBIACEAE

and sessile or pedunculate; bracts leaflike or filiform. Flowers

subsessile to shortly pedicellate. Calyx grayish white pilosulous
or hirtellous; hypanthium portion subglobose, 0.81.2 mm;
limb lobed nearly to base; lobes lanceolate to triangular, 0.71.2
mm. Corolla reddish white, slenderly funnelform, glabrous outside; tube 8.511 mm, sparsely white villous at middle inside;
lobes oblong-ovate, 1.52 mm. Anthers linear-oblong, 11.7
mm, subsessile, partially exserted. Stigma 2-lobed, ca. 0.8 mm.
Capsules not seen. Fl. AprDec.
Forests on hill slopes; 9001200 m. Guangxi (Napo).
W. C. Chens key (in FRPS 71(1): 192. 1999) described the leaves
of this species as hispidulous on both surfaces, but this conflicts with
the protologue and Chens species description.

2. Wendlandia augustinii Cowan, Notes Roy. Bot. Gard. Edinburgh 16: 298. 1932.
si mao shui jin shu
Shrubs, 23 m tall; branches terete, densely tomentulose to
hirtellous becoming glabrescent. Leaves opposite; petiole 515
mm, densely tomentulose; blade drying papery, elliptic or elliptic-ovate, 915.5 26 cm, adaxially sparsely hispidulous to
glabrous on lamina and sparsely to densely puberulent on principal veins, abaxially sparsely strigillose to hispidulous, base
cuneate to acute, apex acute to acuminate; secondary veins 79
pairs; stipules generally persistent, pandurate, 57 34 mm,
densely hirtellous, pilosulous, or strigillose, apically spreading,
obtuse to rounded. Inflorescences paniculate, pyramidal in outline, 1112 1114 cm, branched to 2 or 3 orders, densely pilosulous to strigillose, pedunculate; peduncle 0.81.5 cm; bracts
linear to narrowly lanceolate, 11.5 mm. Flowers sessile or subsessile. Calyx densely pilosulous to puberulent; hypanthium
portion ellipsoid to turbinate, 0.81.2 mm; limb deeply lobed;
lobes triangular, ca. 0.3 mm. Corolla white, funnelform, outside
glabrous; tube 45 mm, inside glabrous or puberulent in throat;
lobes ovate, 0.61 mm. Anthers elliptic, subsessile, ca. 0.8 mm,
partially exserted. Stigma bilobed, ca. 0.4 mm. Capsules not
seen.
Forests on mountains; ca. 1300 m. Yunnan (Simao).

3. Wendlandia bouvardioides Hutchinson in Sargent, Pl. Wilson. 3: 393. 1916.

357

narrowly triangular, or lanceolate, 15 mm; pedicels to 1 mm.

Flowers subsessile to pedicellate. Calyx densely strigillose; hypanthium portion turbinate, ca. 1 mm; limb deeply lobed; lobes
lanceolate to narrowly triangular, 0.31.5 mm, unequal on an
individual flower (this entire size range found on one flower).
Corolla red or purple, salverform or tubular, outside glabrous;
tube 1011 mm; lobes oblong to lanceolate, 1.52.5 mm, obtuse
to rounded. Anthers lanceolate, 11.7 mm, partially exserted,
subsessile. Stigma 2-lobed, 0.30.5 mm. Capsules globose, 2
2.5 mm in diam., strigillose. Fl. FebDec, fr. MayDec.
Thickets or forests in valleys, on mountain slopes, or at streamsides; 12001800 m. Yunnan.
This species seems to have been grouped together with Wendlandia species with relatively long, fully exserted anthers by W. C. Chen
(in FRPS 71(1): 192193. 1999), but it actually has relatively short fat
anthers that are not exserted, as noted in the protologue.

4. Wendlandia brevipaniculata W. C. Chen, Acta Phytotax.

Sin. 21: 386. 1983.
chui shu
Trees; branches subterete, yellowish brown velutinous or
mealy pubescent. Leaves opposite; petiole 0.81.2 cm, puberulent; blade drying leathery, elliptic or ovate-elliptic, 4.510.5
2.54.5 cm, sparsely pubescent to glabrescent on both surfaces
with pubescence denser along principal veins, base cuneate to
obtuse, apex abruptly shortly acuminate or acute; secondary
veins 810 pairs; stipules generally persistent, triangular to
ovate, ca. 4 mm, puberulent to strigillose, apex cuspidate, erect.
Inflorescence paniculate, subglobose, 56 36 cm, densely tomentose or strigillose to glabrescent, sessile and tripartite or
pedunculate; peduncle 0.81.8 cm; bracts lanceolate to linear,
15 mm. Flowers subsessile. Calyx glabrous or sparsely villosulous; hypanthium subglobose, ca. 0.8 mm; limb lobed nearly
to base; lobes lanceolate, ca. 0.8 mm. Corolla tubular-salverform, outside glabrous or villosulous on lobes; tube 23 mm,
white hirsute inside; lobes elliptic, ca. 1 mm. Filaments ca. 0.8
mm; anthers elliptic, ca. 0.8 mm, partially exserted. Stigma 2lobed, ca. 0.2 mm. Capsules globose, ca. 2 mm in diam., puberulent, villosulous, or subglabrous. Fl. and fr. Sep.
Forests; 200300 m. Yunnan (Jingdong).

bao ye shui jin shu

5. Wendlandia brevituba Chun & F. C. How ex W. C. Chen,

Acta Phytotax. Sin. 21: 397. 1983.

Shrubs or small trees, 15 m tall; branches terete, densely

strigose to hirsute. Leaves opposite; petiole 0.41.2 cm, strigose
to strigillose; blade drying thinly papery, elliptic, narrowly elliptic, or lanceolate, 520 1.25.5 cm, adaxially strigillose on
principal veins and glabrescent on lamina, abaxially glabrescent
on lamina, strigose to sericeous on principal veins, and strigillose on higher order veins, base acute to attenuate, apex long
acuminate; secondary veins 58 pairs; stipules deciduous, suborbicular, ovate, or leaflike, 35 3.55 mm, strigose to glabrescent, base shortly stipitate, apex spreading to reflexed, obtuse to shortly acuminate. Inflorescences congested-cymose or
thyrsoid, cylindrical in outline, branched to 1 or 2 orders, 4
10.5 310 cm, densely strigose, hirsute, or strigillose, subsessile to pedunculate; peduncle to 1 cm; bracts subulate, linear,

duan tong shui jin shu

Shrubs, 0.53 m tall; branches flattened to terete, densely
hirtellous, pilosulous, or strigillose. Leaves opposite; petiole
0.31.5 cm, densely hirtellous or pilosulous; blade drying papery, elliptic-oblong, elliptic-ovate, or elliptic, 515 26.3
cm, adaxially glabrous or sparsely strigillose on lamina and
densely strigillose on principal veins, abaxially sparsely strigillose to glabrous on lamina and densely strigillose to pilosulous
on principal veins, base cuneate, apex acuminate to shortly acuminate; secondary veins 48 pairs; stipules deciduous, suborbicular, elliptic, or pandurate, 26 1.54 mm, moderately to
densely strigillose, spreading at apex to fully reflexed, apex obtuse to rounded. Inflorescences rather laxly paniculate, pyra-

RUBIACEAE

358

midal in outline, branched to 1 or 2 orders, 47 411 cm,

densely pilosulous to hirtellous, sessile and tripartite or pedunculate; peduncle 0.33.5 cm; bracts linear to triangular, 0.31
mm. Flowers subsessile. Calyx densely strigillose to pilosulous;
hypanthium portion subglobose, ca. 0.8 mm; limb deeply lobed;
lobes lanceolate to triangular, 0.51 mm. Corolla white, tubular
or salverform, sparsely pilosulous or glabrous outside; tube 1
1.5 mm, sparsely pubescent or glabrescent inside; lobes oblongovate or subovate, 11.5 mm, rounded to obtuse. Anthers elliptic, ca. 0.8 mm, subsessile, partially exserted. Stigma bifid, ca. 1
mm. Capsules subglobose, 11.5 mm in diam., pilosulous. Fl.
AprMay, fr. JunDec.
Forests in valleys; 100900 m. Guangdong, Guangxi.

6. Wendlandia cavaleriei H. Lveill, Repert. Spec. Nov. Regni Veg. 10: 434. 1912.
gui zhou shui jin shu
Wendlandia feddei H. Lveill.
Shrubs or small trees, 1.53 m tall; branches brown, subterete, strigillose to glabrescent. Leaves opposite; petiole 0.8
1.2 cm, strigillose; blade drying subleathery, ovate, elliptic, or
oblanceolate-elliptic, 4.513.5 26 cm, adaxially glabrous to
sparsely strigillose on lamina and sparsely pubescent along
midrib, abaxially sparsely hirsute with pubescence denser along
veins, base obtuse to acute, apex acute or acuminate; secondary
veins 710 pairs; stipules generally persistent, broadly triangular, ca. 5 mm, strigillose, apex erect, cuspidate. Inflorescence
paniculate, pyramidal to ovate in outline, 721 615 cm,
densely many flowered, branched to 2 or 3 orders, densely yellowish brown pilosulous, sessile and tripartite or pedunculate;
peduncle 16 cm; bracts triangular, 13 mm. Flowers sessile.
Calyx densely hirtellous to pilosulous; hypanthium portion subglobose, ca. 1 mm; limb deeply lobed; lobes narrowly triangular, 11.2 mm. Corolla white or purple, tubular-funnelform, glabrous to pilosulous; tube ca. 3.5 mm, inside white villosulous;
lobes ovate to elliptic, ca. 1 mm. Anthers elliptic, ca. 0.8 mm,
subsessile, partially exserted. Stigma 2-lobed, ca. 0.7 mm. Capsules globose, ca. 1.5 mm in diam., pilosulous. Fl. MarApr, fr.
Apr.
Forests or thickets on hill slopes; 200700 m. Guangxi (Tianyang), Guizhou.
W. C. Chen (in FRPS 71(1): 203. 1999) described the corollas as
glabrous outside, but Cowan (Notes Roy. Bot. Gard. Edinburgh 16:
263264. 1932) specifically noted that the corollas of Wendlandia cavaleriei vary from glabrous to pubescent in China, as described here.

7. Wendlandia erythroxylon Cowan, Notes Roy. Bot. Gard.

Edinburgh 16: 299. 1932.
hong mu shui jin shu
Trees, ca. 7 m tall; wood firm, red; branches quadrate,
brownish gray, moderately to densely mixed pubescent, hirsute
and pilosulous, sometimes becoming glabrescent. Leaves opposite; petiole 0.51 cm, moderately to densely strigillose to hirtellous; blade drying leathery or papery, elliptic-ovate, obovate,
or ovate, 512 35.5 cm, adaxially sparsely strigillose or hirtellous to glabrescent, abaxially sparsely to moderately hirtel-

lous with pubescence denser along principal veins, base obtuse

to acute, margins scaberulous to denticulate, apex acute to acuminate; secondary veins 6 or 7 pairs; stipules generally persistent, suborbicular to pandurate, 58 45.5 mm, sparsely to
densely strigillose or strigose basally and glabrescent above,
apex usually spreading to reflexed, obtuse to rounded. Inflorescence rather laxly paniculate, 513 312.5 cm, sparsely to
densely hirtellous to hirsute, tripartite and sessile or pedunculate; peduncle 16 cm; bracts ligulate to lanceolate or narrowly
elliptic, 1.25 mm, often glabrescent. Flowers sessile or subsessile. Calyx glabrous or sparsely pilose; hypanthium portion subglobose to ellipsoid, ca. 0.8 mm; limb lobed nearly to base;
lobes ovate-oblong to ligulate, ca. 1.2 mm, ciliate. Corolla tubular to salverform, glabrous outside; tube 2.53 mm, in throat
white hirsute; lobes ovate, 11.2 mm. Anthers elliptic, ca. 0.8
mm, subsessile, partially exserted. Stigma ca. 0.5 mm, 2-lobed.
Capsules unknown. Fl. AprOct.
Forests on hills of low elevations. Taiwan.
The leaf margins of the specimens seen are denticulate, with the
teeth apparently scabrosities rather than dentate mesophyll tissue.

8. Wendlandia formosana Cowan, Notes Roy. Bot. Gard.

Edinburgh 16: 247. 1932.
shui jin jing
Shrubs or trees, 28 m tall; branches flattened to terete,
densely puberulent to glabrescent, reddish brown to gray.
Leaves opposite; petiole 0.72.5 cm, thinly puberulent or glabrous; blade drying papery, elliptic or elliptic-lanceolate, 614
26.5 cm, both surfaces glabrescent or sparsely to densely
puberulent or strigillose at least on principal veins, base acute to
obtuse, apex acute to acuminate; secondary veins 510 pairs;
stipules generally persistent, broadly triangular, 23.5 mm,
densely strigillose to puberulent, apex erect, acute, cuspidate, or
sometimes narrowly spatulate. Inflorescences paniculate, pyramidal in outline, 1220 1020 cm, branched to 2 or 3 orders,
densely puberulent to strigillose, sessile and tripartite or pedunculate; peduncle 24 cm; bracts linear, spatulate, or narrowly
elliptic, 16 mm; pedicels 0.54 mm. Flowers subsessile to
pedicellate. Calyx glabrous; hypanthium portion subglobose,
ca. 0.8 mm; limb lobed nearly to base; lobes triangular, 0.30.5
mm, sometimes ciliolate. Corolla white to yellow, funnelform,
glabrous outside; tube 1.54 mm, villosulous in throat; lobes
spatulate, 1.52.5 mm. Filaments ca. 0.8 mm; anthers linearlanceolate, ca. 2 mm, partially exserted. Stigma 2-lobed, ca. 0.8
mm. Capsules globose, 1.82 22.2 mm, glabrous. Fl. Apr
Jun, fr. MayDec.
Thickets or forests at low elevations or on hills or mountains;
2001600 m. Guangdong, Guangxi, Taiwan, Yunnan [Japan (Ryukyu
Islands), Vietnam].

1a. Leaf secondary veins 710 pairs,

closely set, usually prominent abaxially;
corolla 35 mm, with tube slightly
longer than lobes ................................ 8a. subsp. breviflora
1b. Leaf secondary veins 59 pairs, rather
well spaced, prominulous abaxially;
corolla 56.5 mm, with tube
markedly longer than lobes .............. 8b. subsp. formosana

RUBIACEAE

8a. Wendlandia formosana subsp. breviflora F. C. How, Sunyatsenia 7(12): 38. 1948.
duan hua shui jin jing
Leaf secondary veins 710 pairs, closely set, usually
prominent abaxially. Corolla tube 1.53 mm, lobes 1.52 mm.
Fl. AprJun, fr. MayDec.
Thickets or forests on hills or mountains; 2001600 m. Guangdong, Guangxi, Yunnan [Vietnam].

8b. Wendlandia formosana subsp. formosana

() shui jin jing (yuan ya zhong)
Secondary veins 57 pairs in leaves of flowering branches,
59 pairs in leaves of vegetative branches (Tsang 23991, MO),
rather well spaced, prominulous abaxially. Corolla tube 34
mm; lobes 22.5 mm. Fl. AprJun, fr. JunAug.
Forests on hills at low elevations; 200500 m. Taiwan [Japan
(Ryukyu Islands)].

9. Wendlandia grandis (J. D. Hooker) Cowan, Notes Roy. Bot.

Gard. Edinburgh 16: 261. 1932.
xi zang shui jin shu
Wendlandia tinctoria (Roxburgh) Candolle var. grandis J.
D. Hooker, Fl. Brit. India 3: 38. 1880.
Trees, 34 m tall; branches brown, fissured, strigillose to
glabrescent. Leaves opposite; petiole 0.51.5 cm, strigillose;
blade drying leathery or thickly papery, elliptic, elliptic-oblong,
or obovate-oblong, 517 2.57 cm, adaxially glabrous or
sparsely strigillose, abaxially glabrous or sparsely pubescent on
lamina, sparsely hispidulous along veins, and with pubescence
denser along midrib, base cuneate or acute, apex acute or shortly
acuminate; secondary veins 812 pairs; stipules generally persistent, triangular to ovate, 912 mm, strigillose, apex erect to
spreading, ligulate, 57 mm, often longitudinally folded. Inflorescences paniculate, ovoid, 917 515 cm, densely brown
hirsute; bracts lanceolate, ca. 5 mm. Flowers sessile. Calyx
densely brown hirtellous; hypanthium portion ca. 1 mm; limb
deeply lobed; lobes lanceolate, 0.51 mm. Corolla white, funnelform, glabrous outside; tube ca. 4 mm, white hirsute inside;
lobes triangular, 11.2 mm. Anthers elliptic, ca. 0.8 mm, subsessile, partially exserted. Stigma 2-lobed. Capsules subglobose, ca. 2 mm in diam., pubescent. Fl. and fr. Aug.
Evergreen forests, secondary forests; 7001300 m. Xizang (Mdog) [Bangladesh, Bhutan, India, Myanmar, Nepal].
Springate et al. (Fl. Bhutan 2(2): 754. 1999) described the leaf size
of plants from Bhutan as 1024 511 cm and the corolla tubes there
as 2.55 mm; these measurements might be found in Chinese plants.

10. Wendlandia guangdongensis W. C. Chen, Acta Phytotax.

Sin. 21: 393. 1983.
guang dong shui jin shu
Shrubs or trees, 46 m tall; branches terete, moderately to
densely hirsute. Leaves opposite; petiole 0.31.2 cm, moderately to densely hirtellous; blade drying thickly papery, lanceolate-oblong or ovate-elliptic, 716 38.5 cm, adaxially

359

sparsely strigillose or puberulent to glabrescent on lamina and

moderately to densely strigillose or hirtellous on principal veins,
abaxially sparsely to moderately hirtellous throughout, base
rounded to obtuse, apex obtuse, shortly acuminate, or acute; secondary veins 711 pairs; stipules generally persistent, pandurate, 56 46 mm, glabrescent, with upper part spreading to
reflexed, apex rounded to obtuse. Inflorescences paniculate,
branched to 2 or 3 orders, 1317 1020 cm, densely hirtellous
to hirsute, pedunculate; peduncle 12 cm; bracts narrowly elliptic to oblanceolate, 1.53 mm. Flowers sessile in glomerules.
Calyx glabrous; hypanthium portion subglobose, ca. 0.8 mm;
limb divided nearly to base; lobes lanceolate or narrowly oblong, 1.22 mm, sparsely to densely ciliate. Corolla white or
greenish yellow, tubular to salverform, glabrous outside; tube
23.5 mm, in throat white hirsute; lobes suborbicular, ca. 1 mm.
Anthers elliptic, ca. 0.7 mm, subsessile, partially exserted. Stigma 2-lobed, ca. 0.3 mm. Capsules subglobose, ca. 1.5 mm in
diam., glabrous. Fl. MarApr, fr. May.
Thickets or forests in ravines or on hill slopes; 100800 m.
Guangdong, Hainan.
The ciliate filaments on the margins of the calyx lobes give the
appearance of pilose pubescence on all of the calyx at first glance, but
when teased apart the individual hypanthia on the material seen are
completely glabrous.

11. Wendlandia jingdongensis W. C. Chen, Acta Phytotax.

Sin. 21: 396. 1983.
jing dong shui jin shu
Shrubs, ca. 3 m tall; branches pubescent. Petiole 0.71 cm,
pubescent; leaf blade drying leathery, lanceolate-elliptic, 710.5
24 cm, adaxially glabrous or sparsely puberulent along
veins, abaxially sparsely pubescent along veins and glabrous on
lamina, base cuneate or acute, apex acuminate; secondary veins
7 or 8 pairs; stipules pandurate, as wide as or wider than
branchlets, apex erect, rounded. Inflorescences 915 1015
cm, densely brown hirsute. Flowers sessile. Calyx pubescent;
lobes lanceolate. Corolla pale yellow, glabrous outside; tube
2.53 mm, white hirsute inside; lobes ca. 0.5 mm. Anthers elliptic, ca. 0.6 mm, subsessile, partially exserted. Stigma bifid.
Capsules subglobose, ca. 1.5 mm in diam., densely pubescent.
Fl. and fr. May.
Forests on mountains; ca. 1700 m. Yunnan (Jingdong).

12. Wendlandia laxa S. K. Wu ex W. C. Chen, Acta Phytotax.

Sin. 21: 390. 1983.
shu hua shui jin shu
Small trees, 58 m tall; branches mealy puberulent. Petiole
0.71.5 cm, glabrescent; leaf blade drying papery to thinly
leathery, lanceolate-elliptic to elliptic, 4.513 25 cm, both
surfaces glabrous throughout or puberulent on principal veins,
base acute to obtuse, apex acute to shortly acuminate; secondary veins 6 or 7 pairs; stipules persistent, broadly triangular,
2.44.5 mm, glabrescent, apex erect, cuspidate. Inflorescences
paniculate, pyramidal, 1322 1218 cm, densely puberulent;
pedicels 24.5 mm. Flowers pedicellate. Calyx puberulent to
glabrous; hypanthium portion turbinate; limb lobed nearly to

RUBIACEAE

360

base; lobes ovate. Corolla white, tubular-salverform, outside puberulent to glabrous; tube 3.54 mm, sparsely puberulent inside; lobes subrounded, ca. 1 mm. Anthers elliptic, ca. 0.8 mm,
partially exserted. Stigma 2-lobed. Capsules subglobose, glabrescent. Fl. Nov, fr. Jul, Dec.
Mixed forests; 5001000 m. Yunnan (Jingdong).

13. Wendlandia ligustrina Wallich ex G. Don, Gen. Hist. 3:

518. 1834.
xiao ye shui jin shu
Shrubs, 1.53 m tall; branches flattened to quadrangular,
strigillose to glabrescent. Leaves opposite; petiole 0.81.5 cm,
strigillose to glabrescent; blade drying leathery, dark brown
adaxially, reddish brown abaxially, elliptic or elliptic-ovate, 3
12 1.56 cm, adaxially glabrous, abaxially glabrescent to
sparsely strigillose or puberulent at least along principal veins,
base cuneate and sometimes shortly decurrent, apex acute or
obtuse; secondary veins 68 pairs; stipules generally persistent,
broadly triangular, 23 mm, strigillose to glabrescent, apex
erect, cuspidate. Inflorescences paniculate, pyramidal, 1517
1220 cm, branched to 2 or 3 orders, densely hirtellous to strigillose, sessile and tripartite or pedunculate; peduncle 23 cm;
bracts 1.56 mm; pedicels to 1 mm. Flowers sessile to shortly
pedicellate. Calyx moderately to sparsely strigillose; hypanthium portion ellipsoid, ca. 1 mm; limb lobed nearly to base; lobes
narrowly triangular to linear, 11.5 mm, generally unequal on
an individual flower. Corolla yellow or pale yellowish green,
salverform to tubular-funnelform, glabrous outside; tube 3.55
mm, pilose in throat; lobes narrowly oblong to spatulate, 1.5
2.2 mm. Filaments ca. 1 mm; anthers linear-lanceolate, 1.21.5
mm, partially exserted. Stigma 2-lobed, ca. 1 mm. Capsules subglobose, ca. 2 mm, strigillose. Fl. JunFeb of following year.
Forests in valleys; 15001600 m. Guizhou (Qinglong), Yunnan
[Myanmar].

14. Wendlandia litseifolia F. C. How, Sunyatsenia 7(12): 46.

1948.
mu jiang zi ye shui jin shu
Trees, ca. 10 m tall; branches terete to subquadrangular,
densely brown hirtellous to glabrescent. Leaves opposite; petiole 0.51.2 cm, densely hirtellous to strigillose; blade drying
subleathery, obovate, obovate-elliptic, or rarely elliptic, 6.4
11.5 36.5 cm, adaxially sparsely hispidulous to puberulent
with pubescence denser along midrib, abaxially sparsely to
moderately strigillose or hirtellous with pubescence denser
along veins, base obtuse, apex acute to abruptly and shortly
acuminate; secondary veins 68 pairs, plane to hardly visible
abaxially; stipules generally persistent, pandurate, 35 1.5
2 mm, densely strigillose, apex suberect to spreading, subrounded. Inflorescences paniculate, 710 45 cm, branched to
2 or 3 orders, dark yellow- to reddish brown tomentose to -hirtellous, pedunculate; peduncle 0.50.8 cm; bracts linear to narrowly elliptic, 13 mm. Flowers sessile, densely grouped. Calyx densely hirtellous; hypanthium portion subglobose, ca. 0.8
mm; limb lobed nearly to base; lobes triangular to ovate, ca. 0.5
mm. Corolla pale yellow, tubular-funnelform, glabrous outside;
tube 2.53 mm, sparsely pilose in throat; lobes ovate, ca. 1 mm.

Anthers elliptic, ca. 0.7 mm, subsessile, partially exserted. Stigma 2-lobed, ca. 0.2 mm. Capsules subglobose to ovoid, ca. 2
2 mm, hirtellous. Fl. and fr. Jun.
Forests on mountains or hills; ca. 800 m. Guangxi (Tianlin).
This species was included by How (loc. cit.) and W. C. Chen
(Acta Phytotax. Sin. 21: 391. 1983) in the group of Wendlandia species
with erect, triangular, acute stipules; but examination of an isotype
(MO!) and careful study of the protologue (including the figure) shows
that the stipules are oblanceolate to pandurate, with the apex narrow but
mostly not spreading from the stem on most nodes.

15. Wendlandia longidens (Hance) Hutchinson in Sargent, Pl.

Wilson. 3: 392. 1916.
shui jing ke zi
Hedyotis longidens Hance, J. Bot. 20: 289. 1882; Wendlandia henryi Oliver.
Subshrubs, much branched, 0.21.5 m tall; branches terete, densely strigillose, hirtellous, or hispidulous. Leaves opposite; petiole 0.52 mm, strigillose to hirtellous; blade drying
papery, elliptic-lanceolate, lanceolate, narrowly lanceolate, or
ovate, 0.83 0.31 cm, adaxially densely hispidulous to glabrous, abaxially densely to sparsely hispidulous or strigillose or
glabrous, base acute and usually decurrent, apex acute to shortly
acuminate or rarely obtuse; secondary veins ca. 3 pairs, occasionally with foveolate pilosulous domatia; stipules lanceolate,
triangular, or ovate-triangular, 2.54.5 mm, densely strigillose
to hispidulous, apex erect to spreading, acute to cuspidate. Inflorescence paniculate to cymose, corymbiform to subglobose,
24 24 cm, densely strigillose, hirtellous, or strigose, sessile
and tripartite or pedunculate; peduncle 36 mm; bracts linear to
narrowly triangular, 0.55 mm; pedicels 23 mm. Flowers pedicellate. Calyx densely strigillose to glabrescent; hypanthium
portion turbinate to ellipsoid, 12 mm; limb lobed nearly to
base; lobes linear-oblong to spatulate or oblanceolate, 23 mm.
Corolla white, tubular-salverform, glabrous outside; tube 78
mm; lobes linear-oblong, 78 mm. Filaments 56 mm; anthers
linear-lanceolate, 33.2 mm, exserted. Stigma 2-lobed, ca. 2
mm. Capsule subglobose, 22.5 mm in diam., densely strigillose to glabrescent, with calyx lobes elongating, up to 4 mm. Fl.
MayJul, fr. JulNov.
Thickets at riversides or on mountain slopes; near sea level to
1800 m. Guizhou (Chishui), Hubei (Yichang), Sichuan, Yunnan.

16. Wendlandia longipedicellata F. C. How, Sunyatsenia

7(12): 39. 1948.
chang geng shui jin shu
Shrubs, ca. 1.5 m tall; branchlets ferruginous pubescent.
Leaves opposite; petiole 0.51 cm, glabrous or sparsely pubescent; blade drying leathery, elliptic or lanceolate-elliptic, 48
0.82 cm, adaxially glabrous, abaxially sparsely pubescent
along midrib, base acute and usually decurrent, apex acute; secondary veins not easily visible; stipules generally persistent,
broadly triangular, 1.53 mm, apex erect, acute. Inflorescence
paniculate, lax, pyramidal, 610.5 4.510.5 cm, ferruginous
pilosulous, sessile and tripartite or pedunculate; peduncle 1.8
3.6 cm; pedicels 36 mm. Flowers pedicellate. Calyx sparsely

RUBIACEAE

pilosulous or glabrous; hypanthium portion subglobose, ca.

1.75 mm; limb lobed nearly to base; lobes triangular, ca. 0.75
mm. Corolla purplish red, tubular-salverform, glabrous outside;
tube 2.33 mm, sparsely strigillose inside; lobes narrowly oblong to spatulate, 1.72 mm. Anthers linear-lanceolate, ca. 1.2
mm, subsessile, partially exserted. Stigma 2-lobed, ca. 0.5 mm.
Capsules not seen. Fl. Feb.
Forests on mountains; ca. 1600 m. Yunnan (Luxi).

17. Wendlandia luzoniensis Candolle, Prodr. 4: 412. 1830.

l song shui jin shu
Wendlandia luzoniensis var. membranifolia (Elmer) Cowan; W. membranifolia Elmer.
Shrubs or small trees; branches flattened-quadrate, glabrous to sparsely strigillose. Leaves opposite; petiole 0.81.2
cm, glabrous to sparsely strigillose; blade drying papery, ovateoblong or elliptic, 1020 4.57 cm, glabrous on both surfaces
or sparsely strigillose on principal veins abaxially, base cuneate
to acute, apex acute to shortly acuminate; secondary veins 69
pairs; stipules generally persistent, suborbicular, obovate, oblanceolate, or pandurate, 46 13 mm, glabrous or sparsely
strigillose, apex erect to spreading, obtuse to rounded. Inflorescences paniculate, pyramidal to broadly obconic in outline,
branched to 3 or 4 orders, 1012 1215 cm, densely strigillose to hirtellous, pedunculate; peduncle 0.81 cm; bracts linear
to narrowly lanceolate, 0.52 mm. Flowers sessile or subsessile.
Calyx with hypanthium portion subglobose, 11.5 mm, densely
puberulent to strigillose; limb deeply lobed; lobes triangular to
lanceolate, 0.51 mm, glabrous. Corolla white, tubular or tubular-funnelform, glabrous outside; tube 34 mm, villosulous in
throat; lobes ovate, 0.81.5 mm. Anthers elliptic, ca. 1 mm,
subsessile, partially exserted. Stigma 2-lobed, ca. 0.3 mm. Capsules subglobose, 1.52 mm in diam., moderately to sparsely
strigillose. Fl. and fr. JulAug.
Forests. Taiwan (Taidong) [?India, Philippines, ?Vietnam].
Wendlandia membranifolia was cited by W. C. Chen (in FRPS
71(1): 214. 1999) as a synonym of W. luzoniensis, apparently based on
Cowans 1932 treatment (Notes Roy. Bot. Gard. Edinburgh 16: 294.
1932). However, Cowan subsequently (Notes Roy. Bot. Gard. Edinburgh 18: 187. 1934) separated these plants as W. luzoniensis var. membranifolia; because of the inclusion of the name W. membranifolia, this
other name is here added too.

18. Wendlandia merrilliana Cowan, Notes Roy. Bot. Gard.

Edinburgh 18: 303. 1935.
hai nan shui jin shu
Shrubs or rarely trees, 13(12) m tall; branches terete to
somewhat flattened or quadrangular, sparsely strigillose or pilosulous to usually glabrous. Leaves opposite; petiole 16 mm,
glabrous; blade drying leathery or subleathery, adaxially pale
green to dark gray-green, abaxially rather pale or reddish brown
to dark purple-gray or purple-black, elliptic-oblong, oblong-lanceolate, oblanceolate, or elliptic, 314.5 0.85 cm, glabrous
on both surfaces, base cuneate to rounded, truncate, or cordulate, apex acute to acuminate; secondary veins 59 pairs; stip-

361

ules caducous or persistent, spatulate, lanceolate, or leaflike, 1

10 mm, glabrous, apex erect to spreading, acute. Inflorescence
cymose to paniculate, narrowly pyramidal, 1.510 1.56.6
cm, branched to 2 orders, puberulent to glabrescent, pedunculate; peduncle 0.33 cm; bracts linear, lanceolate, or spatulate, 0.33 mm; pedicels 23.5 mm. Flowers pedicellate. Calyx glabrous; hypanthium portion subglobose to ellipsoid, 1.52
mm; limb deeply lobed; lobes triangular, 0.51 mm. Corolla
white, salverform, glabrous outside; tube 58 mm, pilose in
throat; lobes narrowly oblong, 56 mm. Filaments 44.5 mm;
anthers linear-lanceolate, 33.3 mm, exserted. Stigma 2-lobed,
1.52 mm. Capsules red or purplish black, subglobose to ovoid,
23 2.53.5 mm, glabrous. Fl. and fr. Apr to Jan of following
year.
Open spaces or streamsides and rocks in forests on mountains;
4001400 m. Hainan.
This species was included by Cowan in a group of generally similar species that are all reduced shrubs, but it apparently becomes rather
large compared to the other species; also its stipules are anomalous in
the group, as the other species otherwise have very narrow triangular
stipules.

1a. Branch internodes 0.57 cm; leaves

larger, 4.514.5 1.55 cm; stipules
410 mm, spatulate, lanceolate,
or leaflike .......................................... 18a. var. merrrilliana
1b. Branch internodes 0.42.2 cm; leaves
smaller, 35 0.81.5 cm; stipules
14 mm, spatulate to lanceolate ........... 18b. var. parvifolia
18a. Wendlandia merrilliana var. merrrilliana
() hai nan shui jin shu (yuan bian zhong)
Branch internodes 0.57 cm. Leaves 4.514.5 1.55 cm;
stipules spatulate, lanceolate, or leaflike, 410 mm. Fl. and fr.
Apr to Jan of following year.
Open spaces or streamsides and rocks in forests on mountains;
4001400 m. Hainan.

18b. Wendlandia merrilliana var. parvifolia F. C. How, Sunyatsenia 7(12): 61. 1948.
xi ye hai nan shui jin shu
Branch internodes 0.42.2 cm. Leaves 35 0.81.5 cm;
stipules spatulate to lanceolate, 14 mm. Fr. Oct.
On rocks, infrequent. Hainan (Dongfang).

19. Wendlandia myriantha F. C. How, Sunyatsenia 7(12):

57. 1948.
mi hua shui jin shu
Shrubs, ca. 3 m tall; branches terete to subquadrangular,
brown strigillose. Leaves opposite; petiole 0.71.2 cm, puberulent; blade drying subleathery, elliptic, 712 2.56 cm, adaxially glabrescent or sparsely puberulent along midrib, abaxially
puberulent, base cuneate, margin sparsely ciliolate, apex shortly
acuminate; secondary veins 810 pairs; stipules generally persistent, ovate to pandurate, ca. 7 mm, sparsely puberulent, apex
spreading to reflexed, rounded to 2-lobed. Inflorescences panic-

RUBIACEAE

362

ulate, branched to 24 orders, 1820 1820 cm, puberulent,

sessile and tripartite or pedunculate; peduncle 34 cm; lower
bracts leaflike, upper bracts linear-lanceolate, oblong, oblonglanceolate, or oblanceolate, 26 mm, strigillose. Flowers sessile
in fasciculate groups. Calyx glabrous or sparsely puberulent;
hypanthium portion subglobose, ca. 2 mm; limb lobed nearly to
base; lobes ovate or ovate-triangular, ca. 1.5 mm. Corolla white,
tubular-funnelform, glabrous outside; tube ca. 4 mm, in throat
white hirsute; lobes ovate, ca. 1 mm. Anthers elliptic, ca. 1 mm,
subsessile, partially exserted. Stigma 2-lobed. Capsules not seen.
Fl. Mar.
Forests in ravines; ca. 300 m. Guangxi (Shangsi).

20. Wendlandia oligantha W. C. Chen, Acta Phytotax. Sin. 21:

399. 1983.
long zhou shui jin shu
Shrubs or trees, 310 m tall; branches flattened to quadrangular, grayish brown, glabrous or sparsely or densely puberulent to pilosulous becoming glabrescent. Leaves opposite;
petiole 315 mm, glabrous or mealy puberulent; blade drying
leathery, adaxially dark brown and often shiny, elliptic, ovate,
or ovate-oblong, 310.5 1.55 cm, adaxially glabrous, abaxially glabrous or sparsely to densely strigillose or puberulent
along principal veins, base cuneate or acute, apex acute or
shortly acuminate; secondary veins 57 pairs, hardly visible
abaxially; stipules deciduous, spatulate to pandurate, 1.52.5
mm, pilosulous or puberulent, apex spreading to reflexed,
rounded to obtuse. Inflorescences laxly paniculate, branched to
2 or 3 orders, 720 418 cm, moderately to densely pilosulous, sessile and tripartite or pedunculate; peduncle 0.51.5 cm;
bracts linear, narrowly oblanceolate, or triangular, 15 mm.
Flowers sessile to subsessile. Calyx glabrous to sparsely strigillose; hypanthium portion subglobose, ca. 0.5 mm; limb lobed
nearly to base; lobes triangular to lanceolate, 0.30.5 mm. Corolla white, salverform to funnelform, outside glabrous; tube ca.
2.5 mm, rather abruptly narrowed at base, sparsely villous in
throat; lobes triangular, ca. 1 mm. Anthers elliptic, ca. 0.7 mm,
subsessile, partially exserted. Stigma 2-lobed, ca. 0.2 mm. Capsules subglobose, ca. 1.5 mm in diam., glabrous. Fl. JulAug,
fr. AugDec.
Rocks in forests or thickets in valleys; 3001000 m. Guangxi.

21. Wendlandia parviflora W. C. Chen, Acta Phytotax. Sin.

21: 394. 1983.
xiao hua shui jin shu
Small trees, 45 m tall; branches ferruginous pubescent.
Leaves opposite; petiole 0.81.5 cm, densely ferruginous pubescent; blade drying leathery, elliptic-oblong or ovate-elliptic,
8.517 4.58.5 cm, adaxially sparsely strigose except densely
ferruginous pubescent along veins, abaxially densely ferruginous pubescent, base cuneate, acute, or sometimes obtuse, apex
acuminate; secondary veins 813 pairs; stipules pandurate, 4
5 mm wide, pilosulous, apex spreading to reflexed. Inflorescences paniculate, ca. 19 15 cm, ferruginous pubescent; bracts
ovate, ca. 1 mm. Flowers sessile. Calyx glabrous or pilose; hypanthium portion ca. 0.8 mm; limb deeply lobed; lobes semiorbicular, ca. 0.5 mm. Corolla pale green, outside glabrous; tube

11.5 mm, white hirsute in throat; lobes ca. 1 mm. Anthers

elliptic, ca. 0.5 mm, subsessile, partially exserted. Stigma 2lobed. Capsules not seen. Fl. Dec.
Forests on flat land. Yunnan (Mengla).

22. Wendlandia pendula (Wallich) Candolle, Prodr. 4: 412.

1830.
chui zhi shui jin shu
Rondeletia pendula Wallich in Roxburgh, Fl. Ind. 2: 140.
1824.
Shrubs, often straggly, 13 m tall; branches pendulous, terete, hispidulous or puberulent to glabrescent. Leaves opposite
or ternate, subsessile; blade drying papery, ovate-lanceolate
or ovate, 3.510 1.84 cm, adaxially glabrous to sparsely
hispidulous, abaxially sparsely to moderately scaberulous or
hispidulous with pubescence denser on principal veins, base
rounded to cuneate, apex acute to weakly acuminate; secondary
veins 47 pairs; stipules caducous to persistent, triangular, 1.5
2 mm, glabrescent, apex erect, cuspidate. Inflorescences paniculate, pyramidal in outline, 1015 49 cm, branched to 2 or 3
orders, densely hispidulous to puberulent, sessile and tripartite
or pedunculate; peduncle 24.5 cm; bracts triangular to ovate,
0.51 mm; pedicels 0.51.2 mm. Flowers shortly pedicellate.
Calyx puberulent to glabrous; hypanthium ellipsoid to subglobose, 11.5 mm; limb lobed nearly to base; lobes lanceolate or
triangular, 0.50.8 mm. Corolla red, tubular-funnelform, outside glabrous; tube 45 mm, inside pubescent in upper part;
lobes oblong to elliptic, ca. 2 mm. Anthers linear-oblong, 1.3
1.5 mm, partially exserted. Stigmas clavate, ca. 1.3 mm. Capsules subglobose, 1.52 mm in diam., glabrous or pilose. Fl.
and fr. DecFeb of following year.
Forests or thickets in ravines; 6001300 m. Yunnan [Bhutan, India, Myanmar, Nepal].
W. C. Chen (in FRPS 71(1): 221. 1999) described the stipules as
caducous, but these are persistent on all the specimens studied.

23. Wendlandia pingpienensis F. C. How, Sunyatsenia 7(12):

51. 1948.
ping bian shui jin shu
Shrubs or trees, 315 m tall. Petiole 13 cm, sparsely
strigillose; leaf blade drying papery, elliptic-oblong or oblanceolate-oblong, 7.523 48 cm, adaxially glabrous or sometimes sparsely hispidulous along midrib, abaxially glabrous or
sparsely hispidulous along principal veins, base cuneate or
attenuate, apex acute or shortly caudate; secondary veins 713
pairs; stipules ca. 2 as wide as branchlets, apex reflexed, subrounded. Inflorescences laxly paniculate, to 30 24 cm, ferruginous strigillose. Flowers sessile. Calyx pilosulous to strigillose; hypanthium portion subglobose, 11.5 mm; limb deeply
lobed; lobes triangular. Corolla white, funnelform, glabrous outside; tube 45 mm, white hirsute inside upper part; lobes triangular, ca. 1 mm. Anthers elliptic, ca. 1 mm, subsessile, partially
exserted. Stigma 2-lobed. Capsules subglobose, 1.52 mm in
diam., pubescent or subglabrous. Fl. AprOct, fr. JunNov.
Forests or thickets in valleys; 2001500 m. Yunnan.

RUBIACEAE

24. Wendlandia pubigera W. C. Chen, Acta Phytotax. Sin. 21:

395. 1983.
da ye mu lian hong
Shrubs, to 2 m tall; branches pubescent. Leaves opposite
or ternate; petiole 0.81.8 cm, pubescent; blade drying thinly
leathery, obovate-oblong or elliptic, 14.518 67 cm, adaxially glabrous or sparsely puberulent along principal veins, abaxially sparsely pubescent with pubescence denser along principal
veins, base cuneate, apex shortly acuminate; secondary veins 10
or 11 pairs; stipules generally persistent, pandurate, 7.59
2.53 mm, nearly as wide as to slightly wider than branches,
apex spreading, rounded. Inflorescence paniculate, pyramidal in
outline, ca. 12 1215 cm, branched to 2 or 3 orders, yellowish
brown pubescent, sessile and tripartite or pedunculate; peduncle
1.83 cm. Flowers sessile. Calyx pubescent; hypanthium portion ca. 1.8 mm; limb lobed nearly to base; lobes ovate to deltoid, ca. 1 mm. Corolla white, tubular-salverform, outside glabrous; tube 23 mm, white villous inside upper part; lobes
rounded to spatulate, ca. 1 mm. Anthers elliptic, ca. 0.7 mm,
subsessile, partially exserted. Stigma 2-lobed. Capsules not
seen. Fl. Mar.
Dense forests. Guangxi (Shangsi).
Some of the characters in the description here are augmented with
details from the protologue figure.

25. Wendlandia salicifolia Franchet ex Drake, J. Bot. (Morot)

9: 208. 1895.
liu ye shui jin shu
Ligustrum thea H. Lveill & Dunn.
Shrubs, to ca. 1 m tall; branches flattened to terete,
hirtellous or strigillose to glabrescent. Leaves opposite; petiole
0.53 mm; blade drying thinly leathery, narrowly lanceolate to
narrowly elliptic, 26.5 0.41.2 cm, glabrous on both surfaces
or sparsely pubescent along midrib abaxially, base acute, margin usually edged with thickened vein, apex acute to weakly
acuminate; secondary veins 57 pairs; stipules generally persistent, triangular, 1.54 mm, densely strigillose to hirtellous, apex
erect, acute to aristate. Inflorescences paniculate, pyramidal in
outline, 25 13 cm, branched to 2 or 3 orders, densely hirtellous to strigillose, sessile and tripartite or pedunculate; peduncle 0.40.7 cm; bracts linear, 15 mm; pedicels 0.31 mm.
Flowers subsessile to pedicellate. Calyx moderately to sparsely
strigillose; hypanthium portion ellipsoid to subglobose, ca. 0.8
mm; limb lobed nearly to base; lobes triangular, 0.30.5 mm.
Corolla pale reddish white, funnelform, glabrous outside; tube
25 mm, apparently pubescent inside upper part; lobes linearoblong, 22.5 mm. Filaments ca. 1 mm; anthers linear-lanceolate, ca. 1.4 mm, partially exserted. Stigma 2-lobed, 0.81 mm.
Capsules subglobose to ovoid, ca. 1.5 2 mm, moderately to
sparsely strigillose. Fl. Nov, fr. Jan of following year.

363

Shrubs or trees, 112 m tall; branches terete to quadrangular, densely brown hirtellous to -tomentose. Leaves opposite;
petiole 0.52.7 cm, densely brown tomentulose to -hirtellous;
blade drying papery or leathery, elliptic-obovate, elliptic, or
ovate, 6.518 2.89 cm, adaxially sparsely to moderately
scaberulous on lamina and densely strigillose to tomentulose on
principal veins, abaxially sparsely to densely strigillose, pilosulous, hirtellous, strigose, or pilose, base obtuse, acute, or
rounded, apex acute or acuminate; secondary veins 610 pairs;
stipules generally persistent, spatulate to pandurate, 56 mm,
strigillose or tomentulose to glabrescent, apex spreading, obtuse. Inflorescences paniculate, pyramidal, 2030 2025 cm,
branched to 2 or 3 orders, erect and congested (var. scabra, var.
pilifera) or pendulous and lax (var. dependens), densely hirtellous to tomentose, pedunculate; peduncle ca. 4 cm; bracts linear, 12.5 mm. Flowers sessile or subsessile. Calyx densely hirtellous; hypanthium portion turbinate to ellipsoid, ca. 0.5 mm;
limb deeply lobed; lobes triangular, 0.50.8 mm. Corolla white,
tubular-funnelform, glabrous outside or pubescent on lobes
(var. pilifera); tubes 2.53 mm, inside glabrous or sparsely pubescent in lower part (var. scabra, var. dependens) or white
villous or hispid in upper part (var. pilifera); lobes oblong, 1
1.25 mm. Anthers elliptic, 0.751 mm, subsessile, partially exserted. Stigma 2-lobed, ca. 0.7 mm. Capsules subglobose, ca. 2
mm in diam., hirsute. Fl. MarMay, fr. MayJul.
Forests or thickets on mountains; 1001800 m. Guangxi, Guizhou, Yunnan [Bangladesh, India, Myanmar, Nepal, Thailand, Vietnam].
The application of this name and the report of this species seem to
be based on Cowans incorporation of Kurzs report into his treatment
and may deserve re-evaluation.

1a. Leaves densely strigillose or scaberulous

adaxially, densely tomentulose abaxially
.................................................................... 26c. var. scabra
1b. Leaves glabrescent adaxially, sparsely
strigose or pilose abaxially.
2a. Leaves sparsely strigose abaxially;
inflorescences lax, pendulous;
corolla tubes inside glabrous or
pubescent near middle ................ 26a. var. dependens
2b. Leaves pilose abaxially;
inflorescences compact, erect;
corolla tubes inside white villous
or hispid in upper part ...................... 26b. var. pilifera
26a. Wendlandia scabra var. dependens Cowan, Notes Roy.
Bot. Gard. Edinburgh 16: 292. 1932.
xuan hua shui jin shu
Leaves glabrescent adaxially, sparsely strigose abaxially.
Inflorescence lax, pendulous. Corolla tube inside glabrous or
pubescent near middle; lobes glabrous outside. Fl. MarApr.

Streamsides in ravines and forests on hill slopes; 100200 m.

Guangxi (Donglan), Guizhou (Guanling), Yunnan [Laos, Vietnam].

Forests or thickets on mountains; 5001800 m. W Yunnan.

26. Wendlandia scabra Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat.

Hist. 41: 310. 1872.

26b. Wendlandia scabra var. pilifera F. C. How ex W. C.

Chen, Acta Phytotax. Sin. 21: 395. 1983.

cu ye shui jin shu

mao cu ye shui jin shu

RUBIACEAE

364

Leaves glabrescent adaxially, pilose abaxially. Inflorescence compact, erect. Corolla tube white villous in upper part
inside; lobes pubescent outside. Fl. Apr.
Thickets on mountains. Guangxi (Shangsi).

26c. Wendlandia scabra var. scabra

() cu ye shui jin shu (yuan bian zhong)
Wendlandia paniculata (Roxburgh) Candolle subsp. scabra (Kurz) Cowan; W. zooi F. C. How.
Leaves strigillose and/or scaberulous adaxially, tomentulose abaxially. Inflorescence erect, compact. Corolla tube inside
sparsely pubescent in lower part or glabrous; lobes glabrous
outside. Fl. AprMay, fr. MayJul.
Forests or thickets on mountains; 1001600 m. Guangxi, Guizhou, Yunnan [Bangladesh, India, Myanmar, Nepal, Thailand, Vietnam].

27. Wendlandia speciosa Cowan, Notes Roy. Bot. Gard. Edinburgh 16: 254. 1932.
mei li shui jin shu
Wendlandia speciosa var. forrestii Cowan.
Shrubs or trees, 112 m tall; branches flattened, strigillose
to glabrescent. Leaves opposite; petiole 0.53 cm, strigillose to
glabrescent; blade drying papery or subleathery, ovate, obovate,
ovate-lanceolate, or elliptic, 619 2.511 cm, on both sides
sparsely to moderately strigillose or hirtellous to glabrescent
with pubescence often denser on principal veins, base acute or
cuneate, apex acute or acuminate; secondary veins 512 pairs;
stipules generally persistent, spatulate to pandurate, 47 mm,
strigillose to glabrescent, apex spreading, rounded. Inflorescence paniculate, pyramidal, 2030 2030 cm, branched to 2
or 3 orders, densely strigillose, pedunculate; peduncle ca. 4.5
cm; bracts linear, spatulate, or lanceolate, 13 mm, acute. Flowers subsessile. Calyx strigillose; hypanthium portion turbinate,
ca. 1 mm; limb lobed nearly to base; lobes lanceolate to triangular, 11.5 mm. Corolla white to cream, yellowish white, salverform or tubular, glabrous outside; tube 57 mm, white villous inside; lobes narrowly oblong to oblanceolate, 22.5 mm,
obtuse to rounded. Anthers linear-lanceolate, exserted, 1.32
mm, at base 2-lobed; filaments ca. 1.5 mm. Stigma 2-lobed, 1
1.2 mm. Capsule subglobose, 2.53 mm in diam., pubescent.
Fl. and fr. MarNov.
Forests in ravines, forest margins; 15002800 m. Xizang (Mdog), Yunnan [Bhutan, India].

28. Wendlandia subalpina W. W. Smith, Notes Roy. Bot.

Gard. Edinburgh 9: 142. 1916.
gao shan shui jin shu
Shrubs, tufted to prostrate, 0.31 m tall; branches subterete, puberulent. Leaves opposite, subsessile; petiole to ca. 1
mm, glabrescent; blade drying leathery, ovate or suborbicular,
0.41 0.30.6 mm, glabrescent on both surfaces, base obtuse
to cuneate, apex obtuse, subacute, or shortly acuminate; secondary veins indistinct; stipules deciduous, triangular, ca. 1 mm,
glabrescent, apex erect, apparently acute. Inflorescences paniculate to cymose, 612-flowered, 0.61 0.61 cm, branched to

2 orders, puberulent, pedunculate; peduncle ca. 0.2 cm; bracts

ca. 1 mm; pedicels 23 mm. Flowers pedicellate. Calyx puberulent; hypanthium portion ellipsoid, ca. 1.5 mm; limb deeply
lobed; lobes triangular or lanceolate, 1.52 mm. Corolla white
or pale yellow, tubular, outside glabrous; tube ca. 2.5 mm;
lobes linear-oblong, ca. 3.5 mm. Filaments ca. 3 mm; anthers
linear-lanceolate, ca. 2 mm, exserted. Stigma 2-lobed, ca. 1.2
mm. Capsules subglobose, 22.5 mm in diam., pubescent. Fl.
MayJul, fr. SepOct.
Open spaces or thickets on mountain slopes; 18003100 m.
Yunnan.

29. Wendlandia tinctoria (Roxburgh) Candolle, Prodr. 4: 411.

1830.
ran se shui jin shu
Rondeletia tinctoria Roxburgh, Fl. Ind. 2: 134. 1824.
Shrubs or trees, to 6 m tall; branches somewhat flattened
to terete or quadrangular, densely puberulent, velutinous, or hirtellous usually becoming glabrescent. Leaves opposite; petiole
0.52 cm, densely puberulent or strigillose to glabrous; blade
drying papery to leathery, oblong-lanceolate, elliptic-ovate, or
obovate, 5.520 2.510 cm, adaxially sparsely strigillose at
least on principal veins to glabrous throughout, abaxially
sparsely to moderately strigillose or tomentose to glabrescent,
base acute to obtuse, apex acute to acuminate; secondary veins
1012 pairs, occasionally with pubescent and/or foveolate
domatia; stipules generally persistent, triangular to ovate, 35.5
mm, densely strigillose or puberulent to glabrescent, apex cuspidate, erect or slightly spreading with age. Inflorescences paniculate, pyramidal in outline, 917 922 cm, branched to 24
orders, densely strigillose, pilosulous, velutinous, tomentose,
hirtellous, or villosulous, sessile and tripartite or pedunculate;
peduncle 0.84 cm; bracts linear to narrowly elliptic, 15 mm;
pedicels to 0.8 mm. Flowers sessile to shortly pedicellate. Calyx
densely hirtellous or pilosulous to glabrous; hypanthium portion
subglobose to ellipsoid or turbinate, 0.81 mm; limb lobed
nearly to base; lobes spatulate, triangular, or lanceolate, 0.81
mm. Corolla white, tubular-funnelform, outside glabrous and/or
variously densely villosulous or strigillose; tube 34 mm, pilose
in throat; lobes elliptic to ovate, 0.81 mm. Anthers oblong, ca.
0.5 mm, subsessile, partially exserted. Stigma 2-lobed, 0.50.8
mm. Capsules ovoid, ca. 1.5 22.5 mm, hirtellous to glabrous. Fl. and fr. JanDec.
Dry sparse forests, dense forests, or thickets in valleys, in ravines,
or on mountain slopes; 2002800 m. Guangxi, Guizhou, Yunnan [Bangladesh, Bhutan, India, Myanmar, Nepal, Thailand, Vietnam].
Cowan recognized seven infraspecific taxa of this species, with
five found in China; two additional infraspecific taxa from China were
described by F. C. How. In Cowans circumscription, Wendlandia tinctoria subsp. tinctoria was widespread in India and also found in Bhutan,
Myanmar, Nepal, and Thailand but not known from China.

1a. Inflorescences glabrous or puberulent;

calyx glabrous or sparsely pubescent.
2a. Corolla lobes strigose outside
................................................. 29d. subsp. floribunda
2b. Corolla lobes glabrous outside
................................................... 29g. subsp. orientalis

RUBIACEAE

1b. Inflorescences densely pilosulous or

tomentose.
3a. Calyx with hypanthium portion
glabrous, glabrescent, or puberulent,
lobes glabrous, glabrescent, or
pilosulous.
4a. Corolla lobes pilosulous
outside ................................ 29b. subsp. barbata
4b. Corolla lobes glabrous
outside ............................ 29f. subsp. intermedia
3b. Calyx hirtellous or pilosulous.
5a. Corolla lobes villosulous
outside ................................... 29a. subsp. affinis
5b. Corolla lobes glabrous outside.
6a. Calyx densely
hirtellous ................. 29c. subsp. callitricha
6b. Calyx pilosulous or
sparsely hirtellous ...... 29e. subsp. handelii
29a. Wendlandia tinctoria subsp. affinis F. C. How ex W. C.
Chen, Acta Phytotax. Sin. 21: 389. 1983.
mao guan shui jin shu
Inflorescence densely tomentose. Calyx hirtellous or pilosulous. Corolla lobes villosulous outside. Fl. and fr. FebJun.
Forests or thickets in valleys or on mountain slopes; 7001400
m. Guangxi (Longzhou), Yunnan.
In the protologue, this taxon is treated as subsp. in the text (both
key and description) but its figure is labeled var.

29b. Wendlandia tinctoria subsp. barbata Cowan, Notes Roy.

Bot. Gard. Edinburgh 16: 268. 1932.
cu mao shui jin shu

365

Inflorescence glabrous or puberulent. Calyx glabrous or

sparsely puberulent. Corolla lobes strigillose outside. Fl. Jul.
Dry sparse forests on mountain slopes; ca. 1100 m. Yunnan
(Menglian) [Myanmar, Thailand].

29e. Wendlandia tinctoria subsp. handelii Cowan, Notes Roy.

Bot. Gard. Edinburgh 16: 267. 1932.
ma li shui jin shu
Inflorescences densely pilosulous. Calyx pilosulous or
sparsely hirtellous. Corolla lobes glabrous outside. Fl. and fr.
MarDec.
Forests or thickets on mountain slopes or in ravines; 2001900
m. Guangxi (Wuming), Guizhou (Wangmo), Yunnan.

29f. Wendlandia tinctoria subsp. intermedia (F. C. How) W.

C. Chen, Acta Phytotax. Sin. 21: 390. 1983.
hong pi shui jin shu
Wendlandia tinctoria var. intermedia F. C. How, Sunyatsenia 7(12): 43. 1948.
Inflorescence densely tomentose. Calyx with hypanthium
portion glabrous or puberulent, lobes pilosulous. Corolla lobes
glabrous outside. Fl. and fr. MarMay.
Forests or thickets in valleys; 14001600 m. Yunnan.

29g. Wendlandia tinctoria subsp. orientalis Cowan, Notes

Roy. Bot. Gard. Edinburgh 16: 268. 1932.
dong fang shui jin shu
Inflorescence glabrous or puberulent. Calyx glabrous or
rarely sparsely puberulent. Corolla lobes glabrous outside. Fl.
MarMay, fr. AprOct.

Inflorescence densely tomentose. Calyx with hypanthium

portion glabrous or glabrescent, lobes glabrous or pilosulous.
Corolla lobes pilosulous outside. Fl. and fr. FebNov.

Thickets or forests on mountains; 2002100 m. Guangxi, Yunnan

[India, Myanmar, Thailand].

Thickets or forests in valleys; 10001800 m. Guangxi, Yunnan

[Vietnam].

shui jin shu

29c. Wendlandia tinctoria subsp. callitricha (Cowan) W. C.

Chen, Acta Phytotax. Sin. 21: 389. 1983.

Shrubs or trees, 215 m tall; branches terete to subquadrangular, tomentulose, pilosulous, or hirsute, often with pubescence of two or more types. Leaves opposite; petiole 0.53.5
cm, densely ferruginous hirtellous or -pilosulous; blade drying
papery, broadly elliptic, elliptic-oblong, ovate, or oblong-lanceolate, 726 414 cm, adaxially sparsely hirtellous, hispidulous, or scaberulous on lamina and moderately to densely tomentulose to strigillose along principal veins, abaxially moderately to densely hirtellous to pilosulous, base acute to obtuse,
apex shortly to abruptly acuminate; secondary veins 812 pairs;
stipules persistent or sometimes deciduous, obovate to pandurate, 512 512 mm, hispid, apex spreading to reflexed,
rounded. Inflorescences paniculate, pyramidal in outline, 1020
1020 cm, branched to 24 orders, densely tomentulose,
pilosulous, and/or hirtellous, sessile and tripartite or pedunculate; peduncle 14 cm; bracts linear-lanceolate to spatulate, 14
mm. Flowers sessile or subsessile. Calyx densely grayish pilose, -strigose, -hirtellous, -tomentose, or subglabrous (subsp.
pilosa); hypanthium obconic, ca. 0.8 mm; limb lobed nearly to

hou mao shui jin shu

Wendlandia tinctoria var. callitricha Cowan, Notes Roy.
Bot. Gard. Edinburgh 16: 265. 1932.
Inflorescence densely tomentose. Calyx densely hirtellous.
Corolla lobes glabrous outside. Fl. and fr. year-round.
Forests or thickets on mountain slopes or in ravines; 4002800 m.
Guangxi, Yunnan [Myanmar].

29d. Wendlandia tinctoria subsp. floribunda (Craib) Cowan

in Craib, Fl. Siam. 2: 23. 1932.
duo hua shui jin shu
Wendlandia glabrata Candolle var. floribunda Craib, Bull.
Misc. Inform. Kew 1911: 386. 1911; W. floribunda (Craib)
Craib.

30. Wendlandia uvariifolia Hance, J. Bot. 8: 73. 1870.

RUBIACEAE

366

base; lobes triangular to lanceolate, ca. 1 mm. Corolla white,

tubular-funnelform, glabrous outside; tube 2.53 mm, inside upper part and throat white hirsute; lobes oblong to elliptic, ca. 1
mm. Anthers elliptic, ca. 0.8 mm, subsessile, partially exserted.
Stigma 2-lobed, ca. 0.3 mm. Capsules subglobose, 12 mm in
diam., densely strigillose to strigose. Fl. JanMay, fr. AprOct.
Thickets, forests, or forest margins on mountains or hill slopes, in
ravines, or at streamsides; below 1001200 m. Guangdong, Guangxi,
Guizhou, Hainan, Taiwan, Yunnan [Vietnam].
Cowan discussed in detail the wide morphological variability in
this species, in both vegetative and flower characters.

1a. Calyx pilose or subglabrous ................... 30b. subsp. pilosa

1b. Calyx densely grayish white hirsute,
-strigose, -tomentulose, or -strigillose.
2a. Leaves rather narrow, ellipticoblong or oblong-lanceolate,
pilosulous abaxially .................. 30a. subsp. chinensis
2b. Leaves wider, broadly elliptic
or ovate, densely grayish brown
pilosulous abaxially ................ 30c. subsp. uvariifolia
30a. Wendlandia uvariifolia subsp. chinensis (Merrill) Cowan,
Notes Roy. Bot. Gard. Edinburgh 16: 288. 1932.
zhong hua shui jin shu
Wendlandia chinensis Merrill, Philipp. J. Sci. 15: 257.
1919.
Leaves rather narrow, elliptic-oblong or oblong-lanceolate,
pilosulous abaxially. Calyx densely grayish white hirsute,
-strigose, -tomentulose, or -strigillose. Fl. MarApr, fr. AprJul.
Forests or thickets in ravines or on hill slopes; below 100600
m. Guangdong, Guangxi, Hainan.

30b. Wendlandia uvariifolia subsp. pilosa W. C. Chen, Acta

Phytotax. Sin. 21: 393. 1983.
shu mao shui jin shu
Leaves densely pilosulous abaxially. Calyx pilose or subglabrous. Fl. Nov.

30c. Wendlandia uvariifolia subsp. uvariifolia

() shui jin shu (yuan ya zhong)
Wendlandia dunniana H. Lveill; W. rotundifolia HandelMazzetti; W. uvariifolia subsp. dunniana (H. Lveill) Cowan;
W. uvariifolia subsp. rotundifolia (Handel-Mazzetti) Cowan;
W. uvariifolia subsp. rufula Cowan; W. uvariifolia subsp. yunnanensis Cowan.
Leaves broadly elliptic or ovate, densely grayish brown
pilosulous abaxially. Calyx densely grayish white hirsute, -strigose, -tomentulose, or -strigillose. Fl. JanMay, fr. AprOct.
Thickets, forests, or forest margins on mountains or at streamsides; below 1001200 m. Guangdong, Guangxi, Guizhou, Hainan, Taiwan, Yunnan [Vietnam].

31. Wendlandia villosa W. C. Chen, Acta Phytotax. Sin. 21:

388. 1983.
mao ye shui jin shu
Shrubs or trees; branches densely villous becoming glabrescent. Leaves opposite; petiole 810 mm, villous becoming
glabrescent; blade drying leathery, elliptic or elliptic-oblong,
4.513 2.56 cm, adaxially sparsely strigose or glabrous,
abaxially densely yellowish brown villous, base cuneate, apex
broad then shortly acuminate; secondary veins 79 pairs; stipules generally persistent, triangular to ovate, ca. 6 mm, villous,
apex erect, cuspidate. Inflorescence paniculate, pyramidal in
outline, 912 611 cm, branched to 2 or 3 orders, densely
yellowish brown villosulous to -hirtellous, sessile and tripartite
or pedunculate; peduncle ca. 3 cm; bracts lanceolate, ca. 1.5
mm. Flowers sessile or subsessile. Calyx densely hirtellous or
villosulous; hypanthium portion ellipsoid, ca. 0.8 mm; limb
lobed nearly to base; lobes lanceolate, 0.81 mm, ciliate. Corolla tubular-salverform; tube 34 mm, outside and inside glabrous; lobes elliptic to ovate, ca. 1 mm, densely villosulous or
hirtellous outside. Anthers elliptic, ca. 0.7 mm, subsessile, partially exserted. Stigma 2-lobed, ca. 0.2 mm. Capsules not seen.
Fl. Apr.
Forests. Yunnan.

Forests; ca. 900 m. Yunnan (Mengla).

97. XANTHOPHYTUM Reinwardt ex Blume, Bijdr. 989. 18261827.

yan huang shu shu
Chen Tao (); Charlotte M. Taylor
Paedicalyx Pierre ex Pitard; Xanthophytopsis Pitard.
Small trees, shrubs, or subshrubs, sometimes unbranched (i.e., monocaulous), unarmed, often fleshy; young growth usually
densely sericeous to lanate, hirsute, or tomentose with trichomes drying golden yellow to ferruginous. Raphides present. Leaves
opposite, isophyllous [or sometimes markedly anisophyllous], without domatia; stipules persistent or deciduous, interpetiolar,
generally triangular or somewhat leaflike, sometimes markedly parallel-veined or -fibrous, entire or 2-lobed. Inflorescences axillary,
cymose to paniculate or subcapitate, several to many flowered, sessile to pedunculate, bracteate or bracts reduced. Flowers sessile to
pedicellate, bisexual, distylous or monomorphic, at least sometimes fragrant. Calyx limb shallowly to deeply 5-lobed; lobes sometimes unequal on an individual flower. Corolla white, yellow, or purple, tubular to funnelform, inside with pubescent ring in upper
part of tube; lobes 5, valvate in bud. Stamens 5, exserted or included, inserted near middle to base of corolla tube; filaments reduced
to developed; anthers apparently dorsifixed. Ovary 2-celled, ovules numerous in each cell on peltate placentas attached at middle of
septum; stigmas clavate to 2-lobed, included or exserted. Infructescences often with peduncle, axes, and pedicels elongating notably.
Fruit indehiscent, schizocarpous, or capsular, ovoid to subglobose, dry, with calyx limb persistent or deciduous, sometimes splitting

RUBIACEAE

367

septicidally into 2 indehiscent mericarps or loculicidally dehiscent valves, these each 1-celled, ellipsoid to plano-convex, each with
numerous seeds; seeds small, angled, smooth, often brown.
About 30 species: China, Indonesia (including Borneo, with most of the species), Laos, Malaysia, New Guinea, Pacific islands (Fiji), Philippines, Vietnam; four species in China.
Xanthophytum was reviewed for China by Chun and F. C. How (Sunyatsenia 4: 1015. 1939, as Paedicalyx) and then by H. S. Lo (Bull. Bot.
Res., Harbin 6(4): 2133. 1986), who formally synonymized Xanthophytopsis and Paedicalyx in this region based on previous comments by Bakhuizen but making the necessary combinations. Axelius (Blumea 34: 425497. 1990) provided the only comprehensive review of Xanthophytum. She
recognized four types of trichomes characteristic of the genus (loc. cit.: 427, f. 1); also notable in Xanthophytum are the fruit with numerous tiny seeds
apparently enclosed in drupaceous locules or mericarps, and the stamen filaments that are often fused to the corolla only at the base and appear free or
nearly free. Several authors have described the fruit as drupaceous and splitting, but the fruit morphology apparently corresponds better to schizocarpous in the terminology used in this treatment. Axelius observed also that distyly in Xanthophytum was apparently not noted by Chinese authors,
resulting in somewhat confused descriptions of floral morphology.

1a. Inflorescences subcapitate to congested-cymose, subsessile to shortly pedunculate, with peduncles to 0.5 cm
in flower, sometimes to 1 cm at fruiting; leaves with secondary veins 916 pairs ............................................ 3. X. kwangtungense
1b. Inflorescences subcapitate to laxly cymose or paniculate, pedunculate with peduncles 0.511 cm; leaves
with secondary veins 930 pairs.
2a. Fruit septicidally dehiscent; inflorescences laxly cymose to paniculate; peduncles 411 cm; leaves with
secondary veins 915 pairs ..................................................................................................................................... 2. X. balansae
2b. Fruit indehiscent; inflorescences subcapitate to laxly cymose or paniculate; peduncles 0.56 cm; leaves
with secondary veins 1430 pairs.
3a. Inflorescences subcapitate; corolla tube 22.2 mm ..................................................................................... 1. X. attopevense
3b. Inflorescences laxly cymose to paniculate; corolla tube 1.52 mm ............................................................ 4. X. polyanthum
1. Xanthophytum attopevense (Pierre ex Pitard) H. S. Lo,
Bull. Bot. Res., Harbin 6(4): 32. 1986.

Xanthophytopsis balansae Pitard in Lecomte, Fl. IndoChine 3: 90. 1922.

qiong dao yan huang shu

Shrubs, ca. 1 m tall; stems subterete, densely sericeous.

Petiole 515 mm, puberulent to sericeous; leaf blade drying
membranous or thinly papery, green above, pale brown below,
elliptic or elliptic-oblong to lanceolate, 917.5 2.55 cm,
adaxially glabrous or sparsely strigose, abaxially densely sericeous, base cuneate to acute and often decurrent, apex acute; secondary veins 915 pairs; stipules persistent, narrowly ovate, ca.
10 3 mm, acuminate. Inflorescence laxly cymose to paniculate, pilose to strigillose; peduncles flexuous, 411 cm; branched
portion broadly pyramidal, 27 210 cm; bracts elliptic, up to
2.5 9 cm, acute; bracteoles reduced; pedicels 12 mm. Flowers unknown. Fruit septicidally dehiscent, subglobose to ovoid,
ca. 2 mm in diam., densely strigillose to strigose, with persistent
calyx lobes obovate or oblanceolate, 13 ca. 1 mm, ciliate;
seeds yellow. Fl. and fr. JunOct.

Paedicalyx attopevensis Pierre ex Pitard in Lecomte, Fl.

Indo-Chine 3: 88. 1922.
Shrubs or herbs, to 1 m tall; branches flattened to subterete, densely sericeous to pilose. Petiole 0.22 cm, densely
sericeous; leaf blade drying membranous or thinly papery, narrowly elliptic or oblanceolate-oblong, 1020 3.56 cm, adaxially moderately to sparsely villous to hirsute, abaxially rather
densely hirtellous to hirsute on veins and densely sericeous on
lamina, base acute and often decurrent, apex acuminate or
shortly acuminate; secondary veins 1730 pairs; stipules persistent, ovate or lanceolate, 1217 48 mm, sericeous or pilose
to glabrescent, longitudinally veined, caudate or acuminate. Inflorescences subcapitate, densely sericeous; peduncles 0.51.5
cm; heads subglobose, 0.51 cm in diam.; bracts triangular,
1.53 mm; bracteoles reduced; pedicels 01 mm. Calyx densely
villous or strigose; ovary portion obconic, ca. 1.5 mm; lobes
obovate, 12.5 mm, sometimes unequal, glabrescent. Corolla
white, outside pilosulous at least on lobes; tube 22.2 mm;
lobes oblong-lanceolate, ca. 1 mm. Fruit indehiscent, subglobose to didymous, ca. 2 2.5 mm, hirsute to sericeous. Fl. Jan
Mar, fr. MayAug.
Dense forests. Hainan [Laos, Vietnam].
This species is circumscribed here following Axelius; many of the
specimens that were included in this species by Chinese authors are here
separated in Xanthophytum polyanthum.

2. Xanthophytum balansae (Pitard) H. S. Lo, Bull. Bot. Res.,

Harbin 6(4): 31. 1986.
chang geng yan huang shu

Streamsides in dense forests. Guangxi (Shiwan Dashan) [N Vietnam].

3. Xanthophytum kwangtungense (Chun & F. C. How) H. S.

Lo, Bull. Bot. Res., Harbin 6(4): 32. 1986.
yan huang shu
Xanthophytopsis kwangtungensis Chun & F. C. How, Sunyatsenia 4: 14. 1939.
Shrubs, 0.51 m tall; branches flattened to subterete,
densely sericeous to tomentose-pilose. Petiole 0.53 cm,
densely sericeous to pilose; leaf blade drying papery, elliptic,
elliptic-oblong, or ovate, 520 2.57 cm, adaxially glabrous
or sparsely pilose at least along veins, abaxially densely sericeous or sericeous-villous, base cuneate to acute and usually long
decurrent, apex acuminate; secondary veins 916 pairs; stipules
drying membranous, persistent, ovate to triangular, 915 57

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RUBIACEAE

mm, sericeous or pilose to glabrescent, parallel-veined, acute to

acuminate or bilobed for up to 1/2, margins often ciliolate.
Inflorescences subcapitate to congested-cymose, many flowered, densely strigose to strigillose; peduncles 0.10.5 cm;
flowering portion subglobose, ca. 1 cm in diam.; bracteoles
linear-lanceolate, ca. 2 mm; pedicels 03 mm. Calyx densely
strigillose to sericeous; ovary portion ellipsoid, 1.11.2 mm;
limb lobed essentially to base, glabrous internally; lobes subspatulate to ovate or oblanceolate, 1.22.5 mm, obtuse to
rounded, often ciliate. Corolla pale yellow, campanulate-funnelform, outside pilosulous at least on lobes; tube 2.22.3 mm;
lobes ovate-triangular to spatulate, 1.21.3 mm, obtuse to acute.
Infructescences often borne at lower leaf nodes or below leaves,
with peduncles up to 1 cm, pedicels up to 2 mm. Fruit capsular,
subglobose to ovoid, ca. 2 mm in diam., strigillose to sericeous.
Fl. May, fr. JulOct.
Wet places in forests. SE Guangxi, S Yunnan (Hekou) [Vietnam].
The calyx lobe measurements given here are based on Chinese
specimens studied and on H. S. Lo (in FRPS 71(1): 24. 1999); Axelius
(Blumea 34: 467469. 1990) reported that these range up to 4.1 mm
throughout the range of the species (i.e., in Vietnam). Several specimens
of this species, in particular W. T. Tsang 23975 and 24532, were distributed as Xanthophytum chinense Merrill, but that name has not
been published.

4. Xanthophytum polyanthum Pitard in Lecomte, Fl. IndoChine 3: 91. 1922.

duo hua yan huang shu

Low shrubs, ca. 0.5 m, little branched; branches weakly
angled or subterete, sometimes rather stout, densely sericeous.
Petiole 0.85 cm, densely sericeous; leaf blade drying papery,
lanceolate to elliptic, elliptic-oblong, or oblanceolate, 930
3.58 cm, adaxially sparsely or moderately hirtellous to villous
to subglabrous, abaxially densely pilose to sericeous, base cuneate to acute and usually decurrent, apex acute to acuminate;
secondary veins 1425 pairs; stipules persistent, lanceolate to
ovate, 1.82.3 0.60.9 cm, sericeous to glabrescent, acuminate or lobed for up to 1/3. Inflorescences laxly paniculate or
cymose, densely villous to hirtellous; peduncles 26 cm;
branched portion pyramidal, 1.55 25 cm; bracts triangular,
25 mm; bracteoles narrowly triangular, 12 mm; pedicels 03
mm. Calyx densely villosulous; ovary portion subglobose to
ellipsoid, ca. 0.5 mm; limb lobed nearly to base, glabrous inside;
lobes spatulate to ovate or obovate, 0.62.1 mm, obtuse to
rounded. Corolla tubular to funnelform, glabrous or with a few
stout hairs outside; tube 1.52 mm; lobes 0.51.5 mm, acute.
Fruit indehiscent or perhaps tardily splitting septicidally, ellipsoid to subglobose or didymous, 1.53 mm, villosulous to hirtellous. Fl. Feb, fr. Apr.
Wet forests; ca. 1400 m. Hainan (Jianfeng Ling) [N Vietnam].
The specimens described here were included by Chinese authors
in Xanthophytum attopevense but are here separated following Axelius
(Blumea 34: 470472. 1990).