Vol. XI

Edited by K.
Kubitzki
Volume XI
Flowering Plants. Eudicots
Malpighiales
THE FAMILIES
AND GENERA
OF VASCULAR PLANTS
Edited by K. Kubitzki
For further volumes see list at the end of the book and:
http://www.springer.com/series/1306
.
The Families
and Genera
of Vascular Plants
Edited by K. Kubitzki
XI
Volume Editor:
Flowering Plants Eudicots
Malpighiales

K. Kubitzki
With 74 Figures
Editor
Professor Dr. Klaus Kubitzki
Universität Hamburg
Biozentrum Klein-Flottbek und Botanischer Garten
22609 Hamburg
Germany
ISBN 978-3-642-39416-4 ISBN 978-3-642-39417-1 (eBook)

DOI 10.1007/978-3-642-39417-1
# Springer Heidelberg New York Dordrecht London
Library of Congress Control Number: 2013954369
# Springer-Verlag Berlin Heidelberg 2014

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Preface
It is with great satisfaction that, with this volume, the major part of the families of the
order Malpighiales is now published within this book series. Malpighiales are a
relatively new ordinal concept because the member families have few evident common
characters and, therefore, could be recognised as a monophyletic group only after the
application of DNA sequence studies. Submission of manuscripts for the families
belonging to this order extended over a very long time, and I regret the long delay in
publication for those authors who complied early. Now, all but three families of the
order (Achariaceae, Malpighiaceae and Salicaceae) are treated in this series and a
concise overview of all families is given in the introduction to this volume, including
those (the Clusiaceae Alliance and Passifloraceae Alliance) already published in Vol. IX
of this series.
My deep thanks go to all authors of this volume for their interesting and scholarly
contributions, and to all those who have freely shared additional information and/or
have commented on earlier drafts. Dr. C.C. Davis is warmly thanked for permission to
prepare a simplified version of his consensus tree of malpighialean families for the
introduction of this volume. I am also most grateful to all copyright holders who so
kindly gave permission to reproduce illustrations published under their responsibility,
including the Director and Board of Trustees, Royal Botanic Gardens, Kew, and the
editors of Blumea (Leiden, The Netherlands) and Austrobaileya (Brisbane, Queens-
land). The artist Bobby Angell, New York, deserves my continued appreciation for her
generous authorization to use illustrations published under her authorship.
The volume has greatly profited from the critical eyes of the copy editor,
Dr. Monique Delafontaine, whose dedicated work deserves my warmest thanks.
I would also like to gratefully acknowledge the pleasant collaboration with Dr. Andrea
Schlitzberger from the staff of Springer Verlag, and with SPi Technologies India PvT
Limited.
Hamburg Klaus Kubitzki

13 October 2013
v
.
List of Contributors
Amaral, Maria do Carmo E. do Depto de Biologı́a Vegetal, Instituto de Biologı́a,

Universidade Estadual de Campinas, Cx. P. 6109, Campinas
SP 13083-970, Brazil, mceamaral2009@gmail.com
Ballard, Jr., Harvey E. Ohio University, Environmental and Plant Biology,
Athens, OH 45701-2979, USA, ballardh@ohio.edu
Bayer, Clemens Palmengarten der Stadt Frankfurt, Siesmayerstr. 61,
60323 Frankfurt/M., Germany, clemens.bayer@stadt-
frankfurt.de
Bittrich, Volker Rua Mário de Nucci, 500, Campinas SP 13083-290,
Brazil, folcar2007@gmail.com
Dickison, William C.{ Formerly at University of North Carolina, Chapel Hill,
NC, USA, folcar2007@gmail.com
Dressler, Stefan Forschungsinstitut Senckenberg, Senckenberganlage
25, 60325 Frankfurt/M., Germany, stefan.
dressler@senckenberg.de
Kubitzki, Klaus Biozentrum Klein-Flottbek, Ohnhorststr. 18, 22609
Hamburg, Germany, klaus.kubitzki@gmail.com
Levin, Geoffrey A. Illinois Natural History Survey, University of Illinois,
1816 South Oak Street, Champaign, IL 61820, USA,
glevin@inhs.illinois.edu
Paula-Souza, Juliana de Universidade de São Paulo, Instituto de Biosciências,
Rua de Matão 277, Ed. Sobre as Ondas, São Paulo
SP 05508-090, Brazil, jupsouza@gmail.com
Prance, Sir Ghillean T. Royal Botanic Gardens, Kew, Richmond, Surrey
TW9 3AS, UK, siriain01@yahoo.co.uk
Repplinger, Miriam Johannes Gutenberg-Universität, Institut für Spezielle
Botanik, Mainz 55099, Germany, siriain01@yahoo.co.uk
Schwarzbach, Andrea Dept. of Biological Sciences, University of Texas, 80 Fort
Brown St., Brownsville, TX 78520, USA, andrea.
schwarzbach@utb.edu
Wahlert, Gregory University of Utah, 257 South 1400 East, Salt Lake City,
UT 84112, USA, rinorea@gmail.com
Webster, Grady L.{ Formerly at Section of Plant Biology, University of
California, Davis, CA, USA, rinorea@gmail.com
vii
.
Contents
Introduction to Malpighiales K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Balanopaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
Caryocaraceae G.T. PRANCE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Centroplacaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
Chrysobalanaceae G.T. PRANCE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 19
Ctenolophonaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 29
Dichapetalaceae G.T. PRANCE . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 33
Elatinaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 39
Erythroxylaceae V. BITTRICH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 43
Euphorbiaceae G.L. WEBSTER { . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 51
Euphroniaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 217
Goupiaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 219
Humiriaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 223
Irvingiaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 229
Ixonanthaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 233
Linaceae S. DRESSLER, M. REPPLINGER, and C. BAYER . . . . . 237
Lophopyxidaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
Medusagynaceae W.C. DICKISON {, and K. KUBITZKI . . . . . . . . . . . . . 249
Ochnaceae M.C.E. AMARAL and V. BITTRICH . . . . . . . . . . . . . . . 253
Pandaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 269
Putranjivaceae G. LEVIN . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 273
Quiinaceae K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 277
Rhizophoraceae A.E. SCHWARZBACH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 283
Trigoniaceae V. BITTRICH . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 297
Violaceae H.E. BALLARD, J. DE PAULA-SOUZA,
and G.A. WAHLERT . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 303
ix
x Contents
Addition to Peridiscaceae C. BAYER and S. DRESSLER . . . . . . . . . . . . . . . . . . . . . . 323

General References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 325
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 327
Introduction to Malpighiales
K. K U B I T Z K I
Conspectus of Families including those treated in numerous, inner stamens staminodial; ovary locules
previous volumes and those not received for and stylodia 4–20; ovule 1 per locule. 1/27, neotropical
Caryocaraceae
inclusion into this volume
– Stamen filaments not vesiculate; leaves rarely 3-folio-
1. Plants with unclear differentiation of stems, roots, and late but then not long-petiolate 6
leaves, or even lacking these completely 2 6. Plants with the combination of epitropous ovules,
– Green land plants with stems, roots, and leaves nucellar beaks and obturators; fruit structure basi-
3 cally a septicidal/loculicidal capsule/schizocarp, also
2. Echlorophyllous endophytic stem or root parasites on splitting from the persistent columella; mesocarp
Tetrastigma [Vitac.]; flowers often very large; stamens often splitting from endocarp. Euphorbiaceae s. l.
12–40 adnate to a central column; ovary inferior; [as circumscribed in this volume] 7
loculi irregular; ovules very many/carpel; embryo – Not this combination of embryological traits and fruit
undifferentiated. 3/20, SE Asia, W Malesia structure [although epitropus ovules are common and
For the time being treated as Rafflesiaceae but per- nucellar beaks have a scattered occurrence in Mal-
haps to be included into Euphorbiaceae pighiales] 9
[see Tribe Rafflesieae in Vol. II: 560–561] 7. Ovule 1/locule; laticifers + or 0; stems sometimes
– Green cataract-dwellers with unclear differentiation of succulent; vessel elements mostly with simple per-
stems, roots and leaves; fertile pollen and fertilisable forations; leaves alternate, less often opposite or ver-
embryo sacs developed underwater. 49/280, worldwide ticillate; stipules + or 0, with axillary colleters,
in tropical and warm-temperate regions sometimes transformed into glands or spines; petals
Podostemaceae [see Vol. IX: 304–344] 1–8; stamens 1–many; pollen grains usually 3-colpo-
3. Monoecious lianes, climbing by leaf tendrils; vessel rate; inner integument vascularized. 213/c. 5,900.
elements with simple perforations; leaves alternate, ser- Worldwide
rulate to crenate; stipules knob-like; flowers 5-merous, Euphorbiaceae [s. str., "uniovulate euphorbs"]
disk of 5 connate circumgynoecial glands; ovary 5(4)- – Ovules 2/locule; laticifers 0; stems not succulent ["bio-
locular, each locule with 2 apical axile ovules sur- vulate euphorbs"] 8
mounted by small obturator-like appendages; stylodia 8. Vessel elements usually with scalariform perforation
5 (4), minute, subulate, stigmatic; fruit 5-winged, plates; leaves simple and entire (trifoliolate and den-
1-seeded, indehiscent. 1/1, from Malay Peninsula to tate); petals + or 0; stamens usually 3–8; pollen grains
Melanesia Lophopyxidaceae 3–7-colporate (-porate), lacking spines; caruncle
– No lianes climbing by leaf tendrils 4 mostly 0. 59/1,750, pantropical, most in Malesia,
4. Plants producing glucosinolates, woody; dioecious; some temperate
vessel elements with scalariform perforation plates; Euphorbiaceae subfam. Phyllanthoideae
leaves alternate, entire, stipulate; sepals 3–6, petals 0; [alternatively: Phyllanthaceae]
stamens 2–50; disk intrastaminal or 0; ovary 1–3(–6)- – Vessel elements usually with simple perforations;
locular, stylodia as many as locules; ovules 2 per loc- leaves simple or palmately compound; petals mostly
ule embedded in massive obturator; placentation 0; stamens 2–30; pollen grains 3–7-zono-brevicolpor-
axile; fruit drupaceous; seeds 1 per locule (or fruit, oidate or -porate or 10–40-pantoporate, echinate to
by abortion); contain glucosinolates, cucurbitacines. verrucose; caruncle mostly +. 26/95, pantropical, pref-
2/225, pantropical and subtropical, mostly in Africa erably S Hemisphere
and Malesia Putranjivaceae Euphorbiaceae subfam. Oldfieldioideae
– Glucosinolates 0 5 [alternatively: Picrodendraceae]
5. Stamen filaments vesiculate towards apex; leaves 9. Carpels sometimes producing false septa; flowers
long-petiolate, trifoliolate, serrate to entire, stipulate, mostly diplostemonous; lamina vernation usually
alternate to opposite; plants woody; vessel elements involute 10
with simple (scalariform) perforations; stamens – False septa 0, except for some Salicaceae 11
K. Kubitzki (ed.), Flowering Plants. Eudicots, The Families and Genera of Vascular Plants 11, 1
DOI 10.1007/978-3-642-39417-1_1, # Springer-Verlag Berlin Heidelberg 2014
2 K. Kubitzki
10. Woody; vessel elements with simple perforations; sta- axile placentas and a solitary ovule in each locule;
men filaments and style folded in bud; nectary disk stylodia mostly distinct, rarely connate into a common
usually conspicuous; ovary 5(2)-locular; ovules 2 (1) style; woody, often lianescent; secondary growth often
per locule; style simple; seeds winged or arillate. 4/23, anomalous; hairs T-, V- or Y-shaped; leaves usually
pantropical Ixonanthaceae opposite, entire, with paired stipules. x ¼ 6 and 9. 68/
– Herbaceous and woody; vessels with simple or scalar- 1,250, worldwide tropical/subtropical, most in America
iform perforations; stamen filaments not folded in Malpighiaceae
bud; nectary weakly developed; ovary 2–8-locular; sty- [not yet in this volume but see the website of W.R.
lodia distinct, as many as carpels; ovules 2/locule; Anderson et al., Malpighiaceae, http://herbarium.lsa.
seeds sometimes arillate. 13/255, widely distributed umich/edu/malpigh/index.html]
in tropical and mostly northern temperate regions – Calyx lobes without glands producing fat oil 17
Linaceae 17. Plants produce a brownish resin which is also excreted
11. Flowers obliquely monosymmetric; fertile stamens as granulous exudate on the stems; annual and peren-
abaxial, connate, staminodes adaxial; plants nial herbs or subshrubs; leaves opposite, entire or
woody; vessel elements mostly with simple perfora- serrate, with scarious stipules; ovary 2–5-locular;
tions; leaves alternate, entire 12 ovules many per locule; placentation axile or basal;
– Flowers not obliquely monosymmetric or only stylodia 3–5; capsule septicidal; endosperm scant or 0.
slightly so (Malpighiaceae) 15 x ¼ 6. 2/35, nearly worldwide Elatinaceae
12. Flowers with distinctive floral cup lined by a nectary – Plants lacking superficial granulous exudate 18
13 18. Leaves opposite or alternate, entire, estipulate, with
– Flowers lacking distinctive floral cup 14 colleters; schizogenous canals or cavities often + and
13. Corolla of 3 petals; androecium with a large anterior containing clear, black or coloured exudate; vessel
pointed staminode flanked by 2 stamen pairs and elements with simple perforations 19
2 posterior groups of 2–3 tooth-like staminodes; – Leaves usually stipulate, plants usually lacking schi-
ovary 3-locular; ovules 2 per carpel, syntropous, col- zogenous canals or cavities containing exudates 22
lateral, axile; style simple; capsule septicidal. 1/3, 19. Woody or herbaceous; leaves opposite (alternate);
northern South America Euphroniaceae sepals and petals 3–5; stamens distinct or fasciculate;
– Corolla of 5 (0) petals; androecium of (2–)8–20 to very ovary 3–5-locular; ovules 1–many per locule; stylodia
numerous stamens, abaxially best developed; ovary 1- distinct, at least distally, or style simple; aril 0; tri-
locular with 2 ovules or 2-locular with 1 ovule in each chomes, if multicellular, stellate. 9/540, worldwide
locule; style gynobasic; drupe 1-seeded. 18/531, Hypericaceae [see Vol. IX: 194–201]
pantropical, mostly neotropical Chrysobalanaceae – Strictly woody; leaves opposite or alternate 20
14. Inflorescence sometimes adnate to petiole or midvein 20. Anthers often with complex or simple apical glands;
of leafblade; stamens 5 (4), up to 3 of them sometimes leaves alternate to opposite, entire, often with resin
lacking anthers; pollen grains 3-colporate; disk lobate canals; stamens not obviously fasciculate; ovary 1–15-
or a ring; ovary 2–5-locular; ovules 2 per locule; sty- locular; ovules 1–many per locule; style simple, usu-
lodia distinct or more often connate into a single style ally long; seeds exarillate, small to large, when large,
provided with distal style branches; fruit a drupe; consisting almost entirely of the huge cotyledons. 13/
plants contain the poisonous fluoracetic acid. 3/170, 460, pantropical Calophyllaceae
pantropical, also subtropical regions of India and [see Vol. IX under Clusiaceae subfam. Kielmeyeroi-
South Africa Dichapetalaceae deae, pp. 48–66]
– Inflorescence not adnate to petiole or midvein; fertile – Anthers lacking apical glands 21
stamens 4–8 in abaxial part of the flower, staminodes 21. Leaves alternate, supervolute, serrulate-setose; ovary
0 or up to 6 in adaxial part; pollen grains 3–5-porate; 3–5-locular; stylodia distinct or connate into a simple
nectary glands [staminodial?] at base of standard + or or distally branched common style; capsule with per-
0; ovary 3(4)-locular; ovules 1–numerous per locule; sistent column. 3/35, SE Asia and Neotropics
style simple; capsule septicidal. 5/28, Neotropics, Bonnetiaceae [see Vol. IX: 36–39]
Madagascar, Malesia Trigoniaceae
15. Pistillate flowers with the naked ovary surrounded by – Leaves opposite, entire, often with resin canals, verna-
spirally arranged cupular bracts; staminate flowers in tion often flat; stamens sometimes fasciculate; plants
catkins, the perianth reduced to one or several bracts mostly dioecious; ovary 1–5-locular; stylodia widely
or vestigial perianth lobes; ovary 2–3-locular; ovules separate or connate into a common short style; seeds
2 per locule, axile, basal; stylodia repeatedly bifurcate; arillate or not; hypocotyl much enlarged, cotyledons
plants woody; dioecious; vessel elements with scalari- 1/10 or less length of embryo. 14/595, pantropical
form perforation plates; leaves alternate, toothed; sti- Clusiaceae (s. str.)
pules minute. 1/9, Queensland to Fiji Balanopaceae [see Vol. IX under Clusiaceae subfam. Clusioideae
– The naked ovary not surrounded by spirally arranged and Symphonioideae, pp. 48–66]
cupular bracts 16 22. Placentation axile 23
16. All or 4 of the 5 calyx lobes with paired abaxial glands – Placentation parietal 33
producing fat oil [Neotropics] or sometimes nectar 23. Leaves opposite 24
[Palaeotropics]; ovary multilocular (inferior), with – Leaves alternate 27
Introduction to Malpighiales 3
24. Style simple; vessel elements with simple and scalari- staminal tube; ovary 2-locular; ovule 1(2) per carpel;
form perforation plates 25 stylodia distinct or style simple; fruit a 1-seeded drupe
– Common style 0; stylodia as many as carpels; vessel or 2–3-seeded capsule; seeds rarely arillate. 4/240,
elements mostly with simple perforations 26 pantropical, especially neotropical
25. Leaves often serrate; plants sometimes with Erythroxylaceae
subepidermal laticiferous idioblasts; vessel elements – Flowers not heterostylous; petals eligulate; woody; ves-
with scalariform and/or simple perforations; hairs sel elements with scalariform perforation plates
simple; petals usually with a terminal arista and fili- [Bhesa]; leaves alternate, stipulate, colleters + or 0;
form appendages on two lobes, enwrapping 1–5 flowers bisexual or unisexual, 5-merous, haplostemo-
stamens; nectary disk intrastaminal, entire or lobed; nous; disk +; ovary 2- or 3-locular; ovules 2 per locule,
ovary often inferior, 2–20-locular with the locules collateral, obturator 0; stylodia widely divergent; capsule
often incompletely separated at anthesis; ovules 2–6 loculicidal; seed 1 per locule, almost completely envel-
per locule; style simple. 14/145, pantropical oped by an exostomal fleshy, sheet-like, coloured aril;
Rhizophoraceae seed coat exotegmic, thick-walled, ribbon-shaped; endo-
– Leaves opposite, entire; vessel elements with scalari- sperm copious; embryo small. 2/7, West Africa, Indo-
form perforation plates; hairs tufted, stellate and sim- Malesia Centrolepidaceae
ple; nectary disk extrastaminal; petals unappendaged; 31. Stylodia on outer shoulders of the 5 carpels; woody;
ovary 2-locular; ovules 2 per locule, placentation api- vessel elements with scalariform perforation plates;
cal; style simple, apically bifurcate; fruit a capsule; inflorescence umbellate; petals induplicate-valvate,
seed solitary, persistent on columella, with hairy- slender; nectary annular; stamens 5, with very short
papillose arillode. 1/2(3?), W Africa, Malesia filaments; ovary 5-locular; placentation basal-axile;
Ctenolophonaceae ovules few per locule; fruit a few-seeded berry; seeds
26. Leaves opposite, simple, toothed, stipules 0; phloem not arillate. 1/3, C and N South America Goupiaceae
stratified; cristarque cells 0; pollen 3-porate; ovary – Pistil with a single style 32
16–25-carpellate, each carpel with a short stylodium; 32. Woody, contains balsamic juice; stamen filaments not
ovules 2 per locule; fruit capsular, with persistent folded in bud; vessels elements with scalariform per-
columella. 1/1, endemic to Mahé, Seychelles Islands foration plates; leaves alternate, involute; stamens
Medusagynaceae 10–30+; stamen filaments at base connate into a
– Leaves opposite/whorled, simple, sometimes lobed or tube; anthers apiculate or linguiform, thecae with
compound, stipulate; cristarque cells and lysigenous separated superposed loculi; nectary disk intrastam-
mucilage canals +; tertiary veins parallel or plumose inal; ovary 4–7-locular; ovule 1 (2) per locule; style
reticulate and very closely spaced; ovary 2–14-locular simple; fruit a drupe; endocarp with "resin"-filled cav-
(gynoecium apocarpous), ovules 2 per locule; fruit ities, 1(2)-seeded. 8/50, Central and South America to
baccate (folliculate). 4/51, neotropical Quiinaceae S Brazil, one sp. in W Africa Humiriaceae
27. Nectary disk 0 28 – Wood; balsamic juice 0; stamen filaments plicately
– Nectary disk + 29 folded in bud; vessel elements with simple perfora-
28. Flowers unisexual, actinomorphic; stamens 5–15; tions; leaves alternate, vernation involute; stipules
plants dioecious; woody, vessel elements with scalari- very large, intrapetiolar, encircling terminal bud;
form (simple) perforation plates; leaves alternate, ver- stamens 10 (9), anther loculi parallel; nectary disk
nation involute; nectary 0; ovary 2–5-locular; ovule obvious; ovary 5–4(2)-locular; ovule 1 per locule; pla-
strictly 1/locule, pendulous; stylodia 2–5, short, or 0; cental obturator +; style simple, short. 3/10, Old
stigmas spreading; fruit a drupe. 3/15, Old World World tropics Irvingiaceae
tropics Pandaceae 33. Flowers usually strongly monosymmetric; sepals and
– Flowers bisexual, actinomorphic to zygomorphic, petals 5; stamens 5 (3), distinct or filaments connate
often polystemonous; woody; vessel elements mostly into a tube, anthers commonly bearing a dorsal con-
with simple perforations; cristarque cells +; leaves nective appendage; ovary 1-locular, placentation pari-
alternate, simple, entire to serrate, stipulate; anthers etal; ovules 1–many; style simple; fruit capsular,
dehiscing by slits or 1 or 2 pores; nectary 0; ovary usually 3-valvate; woody (herbaceous, lianescent);
1–15-locular, either entire and provided with a simple vessel elements with simple and scalariform perfora-
apical style, or divided into 5 or more uniovulate lobes tion plates; leaves alternate (opposite), with salicoid
and the style gynobasic; ovules 1–many per locule; teeth, stipulate. 22 [+ 9 undescr.]/830, worldwide
fruit capsular or drupaceous, or separating into sev- Violaceae
eral drupelets. 27/495, pantropical, mostly in Brazil – Flowers polysymmetric; connective appendages 0
Ochnaceae 34
29. Ovary 2–3-locular 30 34. (Andro)gynophore often +; calyx and corolla forming
– Ovary 4–5(–7)-locular 31 a floral tube 35
30. Flowers often heterostylous; petals sometimes with – Gynophore and floral tube 0 [the latter + in some
fringed bilobed ligule; woody; vessel elements with Flacourtiaceae] 37
simple perforations; cristarque cells +; leaves alter- 35. (Andro)gynophore 0; petals contorted; floral tube
nate, entire, involute, colleters +, stipules mostly campanulate or funnel-shaped, rarely with weakly
intrapetiolar; nectary glands sometimes on outside of developed corona; nectary near base of tube; ovules
4 K. Kubitzki
1–numerous per placenta (1 basal ovule); seeds aril- herbaria of that time were limited in size and
late, pitted; woody and herbaceous; vessel elements permitted an easier overview than our vast pres-
with simple and sclariform perforations; hairs some-
times stellate. 10/205, Africa, America
ent collections, recognised hardly any relation-
Turneraceae [see Vol. IX: 458–466] ship among at least some of the Malpighialean
– [Andro]gynophore usually present; petals cochlear; families. It remains enigmatic how Hutchinson,
corona often present and strikingly coloured 36 who in his "Families of Flowering Plants" used the
36. Herbaceous to subwoody; vessel elements usually with Malpighialean concept, in the third edition of
simple perforations; stamens 5; pollen grains tricolpo- this work (Hutchinson 1973) included in it 14
rate; ovules numerous on parietal placentae; stylodia
inserted beneath the top of ovary; seeds exarillate;
families of which only five did not belong there
calyx persistent in fruit; tendrils 0. 1/24, Andean according to the modern ordinal concept. There
region of N Chile and S Peru were no obvious morphological characters unit-
Malesherbiaceae [see Vol. IX: 247–249] ing these families, and Hutchinson’s finding
– Woody, vessel elements with simple or scalariform must have been either fortuitous or perhaps due
perforation plates; branch tendrils often present; to his vast experience or instinct, but little atten-
stamens 4, 5, or many; pollen grains 3–12-colporate
or -foraminate; ovules numerous; stylodia inserted on
tion was paid to it until the advent of molecular
top of ovary; seeds arillate; contains often cyanogenic systematics.
compounds. 17/705, pantropical In the first broad-based molecular systemat-
Passifloraceae [see Vol. IX: 270–281] ics analysis of the seed plants by Chase et al.
37. Woody; vessel elements with simple or simple and (1993), a clade within their rosids was disclosed
scalariform perforation plates; leaves supervolute-
curved or involute, alternate, toothed or serrate,
which comprised representatives of 11 families
often with the distinctive "salicoid teeth"; flowers previously never found in association. This
3–6-merous; sepals (0)3–15; petals often 0; nectarifer- clade was confirmed and expanded by further
ous disk or nectar glands intra-, inter- or extrastam- work, and the Angiosperm Phylogeny Group
inal; stamens (1 in Lacistemateae) 2–many; (APG 1998) adopted the ordinal name Malpigh-
placentation parietal (axile); fruit a capsule (berry,
drupe); seed coat commonly not vascularized; plants
iales for it. Molecular analyses specifically
contain benzoylated glacosides but lack cyanogenic devoted to this group (Davis et al. 2005; Wurdack
glycosides (except Banara). 55/1010, pantropical and and Davis 2009; Korotkova et al. 2009) employed
temperate [not received] Salicaceae an increasing number of gene regions, eventually
– Woody (herbaceous and climbing); vessel element of all three genomes, and underpinned its mono-
with simple, scalariform or simple and scalariform phyly; they also confirmed or provided evidence
perforations; leaves alternate, entire, rarely toothed or
serrate but then leaf teeth not salicoid; petiole often
for family relationships and revealed suprafami-
geniculate; sepals 2–5; petals 4–15 (Acharieae: 3–4 and lial clades around the Violaceae/Salicaceae, the
connate), either whitish and papery or with a gland Ochnaceae and the Chrysobalanaceae, but failed
inside at base; nectary 0; stamens 5–many; anthers to resolve deeper relationships among the 16 mal-
sometimes [Chiangiodendron, Kiggelaria] poricidal; pighialean subclades previously identified by
seed coat thick; testa commonly strongly vascularized;
plants contain cyclopentenoid cyanogenic glucosides
Wurdack and Davis (2009). More recently, how-
and/or cyclopentenyl fatty acids [gynocardin]. 32/145, ever, the use of phylogenomic approaches and the
pantropical [not received] Achariaceae inclusion of 82 plastid genes led Xi et al. (2012) to
the elaboration of a well-resolved, taxon-rich
molecular tree for the Malpighiales, in which 12
The Ordinal Concept of Malpighiales. The term additional clades and further increased resolution
Malpighiales had already been created but hardly among its deeper nodes were obtained. This new
used by nineteenth century botanists, at a time phylogenetic hypothesis also proved helpful for
when family and ordinal concepts most gener- an understanding of the detailed morphological
ously were proposed for nearly every characteris- analyses of the Chrysobalanaceae, Rhizophora-
tic family or even genus. In the 19th and ceae, Ochnaceae and their presumed relatives
practically the whole 20th century, the allocation by Matthews and Endress (2008, 2011) and
of the families presently comprised in the Mal- Matthews et al. (2012).
pighiales was scattered over many different Thus, the concept of Malpighiales is the direct
orders. Even the perspicacious botanists of the outcome of molecular systematics, which may
19th century, who had the advantage that the explain its dearth of easily recognisable
distinctive marks. It is only recently that P.F. tion following the concept of the author of the
Stevens (2001 onwards) has succeeded in elabor- family. The sister group relationship of the bio-
ating a list of possible apomorphies for the whole vulate euphorbs with the linoid clade needs atten-
order and its constituent families and family tion because their closeness had been considered
groups, but the order as a whole is characterised by several earlier authors, and a relationship
by nothing more than "leaf margin toothed (teeth between Linaceae and Ixonanthaceae, already
with a single vein running into a congested recognised by Hallier (1923), is now strongly
deciduous apex); stigma dry; exotegmen supported in the analysis of Xi et al. (2012). Pre-
fibrous". viously, there had been much confusion as to the
According to classificatory concepts, Mal- relationships of the Ixonanthaceae, and both
pighiales now comprise 37–42 families, 716 genera morphological and molecular workers had often
and 15,935 species (Stevens 2001 onwards), and associated them with the (equally problematic)
within the eudicots are resolved as a member of Irvingiaceae. The Rafflesiaceae (which are not in
the Fabidae (Rosidae) where they occupy a sister Xi’s consensus tree) are holoparasites living
position to the Oxalidales, both these orders being embedded inside their host plants (species of
sister to the Celastrales (Wang et al. 2009; Moore Tetrastigma), and cladistically have been found
et al. 2010). nested in the Euphorbiaceae between the Peroi-
deae and the rest of the uniovulates (Davis et al.
Phylogeny. Fig. 1 gives the best resolved consen- 2007). If one should not wish to incorporate the
sus tree for malpighialean families available at the three rafflesiaceous genera into the euphorbs, the
time of writing (Jan. 2013), which is redrawn reinstatement of the Peroideae at family rank
from Xi et al. (2012) and will be used to explain would allow the intercalation of the Rafflesiaceae
the phylogenetic interrelationships among the between them. More recently, however, Xi et al.
families. In it, the previously known subclades (2012, in the SI) have placed the Rafflesiaceae as
of Malpighiales, which hitherto were unresolved sister to the Euphorbiaceae + Peroideae, which
in relation to one another, are represented in warns against making premature rearrange-
three well-supported major clades. The first of ments; the loss of plastids and the consequences
these is composed of two subclades; one contains of horizontal gene transfer in the Rafflesiaceae
the euphorbioids and linoids, the other the make the issue very complicated.
Humiriaceae, the parietal clade, and the salicoids. The sister clade to euphorbioids and linoids
The euphorbioid-linoid clade is remarkable comprises the Humiriaceae with the parietal clade
because it brings (albeit with weak support) the and the salicoids. Support for the placement of the
uniovulate euphorbs close to the biovulate ones Humiriaceae is low; relationships previously sug-
(the phyllanthoids), which traditionally have gested for them were quite diverse but never con-
been included in a single family, Euphorbiaceae vincing, and their closest relatives may still
(s. l.). Based on early results of molecular system- remain unknown. In the parietal clade, the Achar-
atics, the euphorbs have been split into three iaceae, reshaped by Chase et al. (2002), are sister
families, these being the uniovulate Euphorbia- to the Violaceae + Goupiaceae, the latter two
ceae, and the biovulate Phyllanthaceae and Picro- being strongly divergent in life form and gynoe-
dendraceae; the latter share many characters and cium structure, but still sharing other floral traits.
more recently have been resolved as sister clades. Further members of this clade are the closely
Although the three-family classification for the related Malesherbiaceae, Turneraceae and Passi-
traditional euphorbs is now generally accepted, floraceae, and ultimately the Salicaceae, again
the common possession of peculiar capsular reshaped by Chase et al. (2002). I cannot see any
fruits (p. 58) and a remarkable combination of cogent reason for maintaining Lacistemataceae,
embryological characters (p. 55) shared by the Samydaceae and Scyphostegiaceae separately
uniovulate and biovulate euphorbs is a momen- from the Salicaceae into which, in a future treat-
tous hint that must not be overlooked; it calls for ment for this series, they would be better
further inquiry into the deeper nodes of the integrated.
Malpighialean phylogeny. In this volume, The second principal clade comprises three
euphorbs are treated in the broad circumscrip- subclades in a trichotomy strongly supported
6 K. Kubitzki
uniovul. Euphorbiac.
biovul. Euphorbiac.
Linaceae
Ixonanthaceae
Salicaceae
Passifloraceae
Turneraceae
Malesherbiaceae
Violaceae
Goupiaceae
Achariaceae
Humiriaceae
Hypericaceae
Podostemaceae
Calophyllaceae
Clusiaceae
Bonnetiaceae
Ochnaceae
Quiinaceae
Medusagynaceae
Rhizophoraceae
Erythroxylaceae
Ctenolophonaceae
Pandaceae
Irvingiaceae
Chrysobalanaceae
Euphroniaceae
Dichapetalaceae
Trigoniaceae
Balanopaceae
Malpighiaceae
Elatinaceae
Centroplacaceae
Caryocaraceae
Putranjivaceae
Lophopyxidaceae
Fig. 1. Maximum likelihood bootstrap consensus tree of matrices; redrawn and simplified from Xi et al. (2012)
the Malpighiales, based on analyses of 82 plastid genes with the kind permission from the authors. Branches
from 58 species added to the existing DNA sequence with low support hatched.
and characterised by the possession of cristarque interesting association between Hypericaceae

cells, with Medusagyne as the only exception and Podostemaceae. Both clusioids and ochnoids
(elsewhere in Malpighiales, cristarque cells are usually lack nectar production and are polyste-
known from Hugonioideae/Linaceae and Irvin- monous; they have brought forth reproductive
giaceae). The first subclade is composed of the specialisations like several kinds of floral secre-
clusioids and the ochnoids, both with a consoli- tions or simply pollen as reward for pollinators,
dated family classification and containing the culminating in buzz pollination. Another
subclade, the strongly supported rhizophorids, plete disappearance of conifers from tropical
comprises the Erythroxylaceae, Rhizophoraceae biomes. They also suggested that the rise of the
and Ctenolophonaceae, which initially had been Malpighiales might have been paralleled by other
recognised as a clade by Wurdack and Davis tropical clades. Recent broader insights into the
(2009; see also the morphological work by Mat- age and diversification of the angiosperms have
thews and Endress 2011). The third subclade, the revealed that the Malpighiales were but one part
pandoids, contains the until recently unplaced of the vast rosid clade comprising most extant
Pandaceae, which for lack of a better alternative representatives of tropical and temperate tree
sometimes had been included in the Euphorbia- species (70,000 species and 140 families), the
ceae. They are here in a weakly supported clade lineages of which underwent a rapid diversifica-
with the Irvingiaceae, the latter to date a family of tion within a short period of time in the Creta-
highly controversial placement. This new alliance ceous. The timing of the inferred radiation of the
is backed by several morphological characters rosids (108 to 91 Mya) and their main branches
listed by Stevens (2001 onwards; see also family Fabidae and Malvidae (107–83 Mya) supports a
treatment of Irvingiaceae). rapid rise of angiosperm-dominated forests in
The third principal clade is composed of four the Cretaceous (Wang et al. 2009; see also
subclades which, except for the position of the Bell et al. 2010; Moore et al. 2010).
Centroplacaceae, had already been recognised by With the invasion of modern trees, the habi-
Wurdack and Davis (2009). The five families tats of those times underwent profound abiotic
composing the chrysobalanoids appear in the and biotic changes. New interactions developed
well-consolidated topology that has already been among plants and their substrates, the soil, as
used as the point of reference for the morpholog- well as their pollinators, dispersers, predators,
ical work by Matthews and Endress (2008). In the symbiotic partners, etc., thereby triggering an
malpighioids, Elatinaceae and Malpighiaceae are enormous codiversification. The increase of
strongly supported as a clade, whereas their sister shade reduced the evapotranspiration of the sub-
position with the Centroplacaceae is only weakly canopy including the soil and vegetation, whereas
supported. A clade formed by the isolated the canopy of the angiospermous forest supplied
Putranjivaceae and Lophopyxidaceae and the increased transpiration. This became particularly
isolated Caryocaraceae belongs also to the third important for the forest belt of the ever-wet equa-
major clade. torial west wind zone, i.e. the tropical rain forest,
the water balance of which is maintained largely
Malpighiales in Time and Space. Davis et al. by its own transpiration and, to a smaller degree,
(2005) used molecular divergence time estimates, by the humidity brought by the western winds
calibrated on reliable fossils, to determine the age from the oceans (Salati and Vose 1984). In addi-
of the Malpighiales clade. They resolved a much tion, biogenous aerosol particles in the atmo-
higher age than was expected at that time, i.e. well sphere yield condensation nuclei for clouds and
before the Cretaceous/Tertiary boundary, 112–94 precipitation, and sustain the hydrological cycle
million years ago (Mya) in the Albian and Cen- (P€oschl et al. 2010). Thus, there is an interdepen-
omanian. The extant representatives of this clade dence between vegetation and climate: the equato-
are mostly species of shrubs and small trees of the rial humid west wind zone cannot exist without the
understorey of tropical rain forests, though some humid equatorial forest and vice versa (Kubitzki
of them are tall rain forest trees (Goupia and the and Krutzsch 1996; Boyce and Lee 2010).
Irvingiaceae), and various families have exten- Very little is known about the place of origin
sions into seasonally dry habitats; the Ochnaceae of the Malpighiales and its component lineages.
can hardly be seen as a forest family. Davis et al. More than half of its families have a pantropical
(2005) extrapolated that the early Malpighiales distribution, but in most cases it is unknown
were shade-tolerant invaders into the under- whether this is the result of vicariance or dis-
storey of the Cretaceous conifer-dominated persal. Lineages that have propagules capable of
forests, where they led to the decline of the flotation (Clusiaceae, Chrysobalanaceae and per-
light-demanding cycadophytes, pteridophytes haps others) appear prone to long-distance dis-
and pteridosperms and eventually to the com- persal and, in view of the very many documented
8 K. Kubitzki
intercontinental dispersal events in other groups, vol.) will demonstrate the value of an ample fossil
it is likely that this may have occurred in the record for an understanding of extant distribu-
Malpighiales as well. The only pertinent case tion. By the end of the Cretaceous, Ctenolophon
studies known to me that demonstrate intercon- appears in West Africa, where it survives to the
tinental dispersal in this plant group are those by present with a single species, soon followed by
Li et al. (2009) on Bridelia and by Bardon et al. several species in southern Arabia. In the Palaeo-
(2013) on the Chrysobalanaceae. Marked differ- cene the genus enters into northern South Amer-
ences in diversification rates among the different ica where its record ends by the Eocene. In the
Malpighiales clades have been found by Xi et al. Palaeocene the genus starts also drifting north-
(2012), disclosing significant rate decelerations in wards on the Indian Plate and there undergoes its
eight clades, among them Balanopaceae, Ctenolo- greatest diversification (up to 9 pollen types) but
phonaceae, Goupiaceae and Lophopyxidaceae, disappears from India in the Neogene, after hav-
and rate accelerations in Malpighiaceae, Passi- ing spread to SE Asia and Malesia where it has
floraceae and three others. Elevated diversifica- survived to the present day in the Malay penin-
tion rates may indeed be related to specialised sula, Sumatra, Borneo, the Philippines, and New
plant–pollinator interactions, as suggested by Xi Guinea. About 50 years ago my unforgettable
et al. (2012), whereas each lineage in decline will mentor, Prof. C.G.G.J. van Steenis, alerted me to
need its own specific explanation. Lophopyxis, for look for living Ctenolophon in the swamp forests
instance, because of its highly specialised shoot of northern South America and, though I did not
morphology and anatomy, may hardly have a see it there, I have pleasure in passing his sugges-
chance for further diversification. tion onto another generation because it still
To finish this section, the story of Ctenolo- makes sense!
phon (for documentation see family account, this For references, see under General References.
Balanopaceae
Balanopaceae Benth. in Benth. & Hook., Gen. Pl. 3: 341 (1880), nom. cons. (‘Balanopseae’).
K. K U B I T Z K I
Dioecious small to tall evergreen trees; older bark and Fiji), and bound to forest and scrub commu-
with white, circular, prominent lenticels; hairs nities; on New Caledonia growing on both ser-
simple, unicellular. Leaves basically alternate, pentine and gneiss.
dimorphic, shoots with minute scale leaves proxi-
mally and foliage leaves distally, the latter some- VEGETATIVE MORPHOLOGY AND ANATOMY. According
times subverticillate and restricted to the shoot tip; to the study by Carlquist (1980), the leaves are
foliage leaves petiolate, with prominently to dimorphic: prior to the fully formed leaves, scale
obscurely toothed margins; stipules a pair of min- leaves appear on each shoot. In some species
ute teeth at the base of the petiole. Inflorescences (confined to New Caledonia), the foliage leaves
usually subtended by scale leaves, less commonly form subverticillate clusters at the tips of
by foliage leaves, in staminate plants a catkin; the shoots only. The leaf margins are toothed to
staminate flowers short-pedicellate and axillary to various degrees. The teeth are several cell layers
scales on the catkin axis, subtended by one or thick, and the cells are filled with a dark content
several bracts or vestigial perianth lobes, these which is thought to be tannins. Both the scale
often fewer than stamens; stamens (1–)3–6(12), leaves and the foliage leaves are each provided
filaments short, anthers dehiscing laterally; a ves- with a pair of lateral, dark-coloured stipules.
tigial pistil sometimes present; pistillate plants Hairs are non-glandular, unicellular and early
with solitary subsessile or short-pedicellate flowers vanishing from the leaves, but persistent in the
in the axils of scale leaves, with numerous densely involucral bracts, which subtend the solitary
crowded, spirally arranged deltoid “cupular” female flower and fruit and form the “cupule”.
bracts subtending the naked ovary, or a terminal Balanops has trilacunar nodes. For leaf anat-
flower surrounded by further lateral flowers also omy, see Carlquist (1980); a notable anatomical
subtended by cupular bracts; staminodes 0; gynoe- feature is the scattered occurrence of cristarque
cium syncarpous, 2(3)-carpellate; ovary 2(3)-locular, cells in the leaves. The involucral bracts contain
the locules imperfectly septate at anthesis but fully brachysclereids. The wood has no growth rings.
septate in fruit; ovules 2 per carpel, collateral, The vessel elements are relatively long and pos-
borne axile-basally, bitegmic and intermediate sess scalariform or reticulate-scalariform perfo-
between apotropous and epitropous; stylodia ration plates. The wood parenchyma is diffuse,
connate at the base and once or twice bifurcate and the rays are multiseriate and uniseriate, with
to form ribbon-like lobes stigmatic on the adaxial a predominance of erect cells.
surface. Fruit a yellow, orange or brown drupe
with rather thin fleshy mesocarp, subtended at REPRODUCTIVE STRUCTURES. In contrast to earlier
the base by a cup of numerous imbricate, con- reports to the contrary, Sutter and Endress (2003)
crescent bracts; pyrenes 2–3; endosperm fleshy have shown that the ovary of Balanops is
during fruit maturation, but crushed at maturity, completely septate, that the ovules are weakly
and forming only a thin layer around the embryo; crassinucellate, bitegmic with a multilayered
embryo large, green, straight. n ¼ 20 (21). inner integument (in contrast to all previous
A single genus comprising nine spp., indications), and intermediate between apotropus
distributed across “old lands” in the S Pacific and epitropous, and that an obturator is not
(N Queensland, New Caledonia, New Hebrides present.
10 K. Kubitzki
A F F I N I T I E S . The affinities of Balanops were

dubious to early workers such as Engler (1897)
and have remained so for a long time. After dis-
cussing all taxa that had been suggested as candi-
dates for a relationship to Balanops, Carlquist
(1980) focused on Hamamelidaceae, Myricaceae,
Pittosporales sensu Thorne, and particularly
Daphniphyllaceae. Takhtajan (1997) combined
Balanopaceae and Daphniphyllaceae in mono-
typic orders in a superorder Daphniphyllanae,
but in sequence analyses of the rbcL gene (Litt
and Chase 1999; Savolainen, Fay et al. 2000; Chase
et al. 2002) Balanopaceae appeared within the
Malpighiales in a clade in which they are sister
to Dichapetalaceae/Trigoniaceae plus Chrysoba-
lanaceae/Euphroniaceae. In contrast, Sutter and
Endress (2003) and Matthews and Endress (2008)
argued that features of Balanops such as
the unisexual flowers, the lack of a perianth, the
repeatedly bifurcate stylodia and the weakly cras-
sinucellar ovules would fit much better with
Fig. 2. Balanopaceae. Balanops sparsiflora. A Branchlet a euphorbialean than the chrysobalanoid rela-
with pistillate flowers. B Pistillate flowers enclosed by tionship. However, the multi-gene analyses of
cupular bracts. C Pistillate flower. D Same, transversally
sectioned. (Takhtajan 1980) Wurdack and Davis (2009), Soltis et al. (2011)
and Xi et al. (2012) provide strong support for
the earlier molecular findings.
EMBRYOLOGY. Unknown. Only one genus:
POLLEN MORPHOLOGY. The pollen is 3–5-colpate, Balanops Baill. Fig. 2

with colpi that show no sign of an endoaperture.
The exine sculpturing consists of small spinules. Balanops Baill., Adansonia 10: 117 (1871); Carlquist, Aller-
The exine structure is tectate-granular to colu- tonia 2: 191–246 (1980), rev.
mellate. The tectum is traversed by microperfora- Trilocularia Schlechter (1906).
tions (Erdtman 1952; Zavada and Dilcher 1986). Description as for family. Nine species, for distri-
bution see above.
POLLINATION. Probably by wind.
F R U I T A N D S E E D . The pericarp consists mostly Selected Bibliography

of thin-walled parenchyma cells, but below the
epidermis two or three layers of thick-walled bra-
Carlquist, S. 1980. Anatomy and systematics of Balanopaceae.
chysclereids have been observed (Guillaumin Allertonia 2(3): 191–246.
1925; Carlquist 1980). Dispersal is probably by Carlquist, S. 1989. Balanopaceae. In: Flora of Australia, vol.
birds (Carlquist 1980). 3: 93–95. Canberra: Government Publishing Service.
Chase, M.W. et al. 2002. See General References.
Engler, A. 1897. Balanopsidaceae. In: Engler & Prantl, Nat.
P H Y T O C H E M I S T R Y . Carlquist (1980) and Sutter Pflanzenfam., Nachtr. I zu T. III, 1. Leipzig: W.
and Endress (2003) mention idioblasts contain- Engelmann, pp. 114–116.
ing dark-coloured substances, supposedly tan- Erdtman, G. 1952. See General References.
Guillaumin, A. 1925. Recherches sur l’anatomie et la clas-
nins, but it is unknown of which type these are. sification des Balanopsidacées. Rev. Gén. Bot. 37:
The occurrence of trivial triterpenes gives no clue 433–449.
for affinities (Hegnauer 1964). Hegnauer, R. 1964. See General References.
Balanopaceae 11
Hjelmquist, H. 1948. Studies on the floral morphology Soltis, D.E. et al. 2011. See General References.
and phylogeny of the Amentiferae. Bot. Not. Suppl. Sutter, D.M., Endress, P.K. 2003. Female flower and
2: 1–171. cupule structure in Balanopaceae, an enigmatic
Litt, A., Chase, M.W. 1999. The systematic position of rosid family. Ann. Bot. 92: 459–469.
Euphronia, with comments on the position of Bala- Takhtajan, A.L. 1980. See General References.
nops: an analysis based on rbcL sequence data. Syst. Takhtajan, A.L. 1997. See General References.
Bot. 23: 401–409. Wurdack, K.J., Davis, C.C. 2009. See General References.
Matthews, M.L., Endress, P.K. 2008. See General References. Xi, Z. et al. 2012. See General References.
Nemirovich-Dachenko, E.N. 1991. Balanopaceae. In: Zavada, M.S., Dilcher, D.L. 1986. Comparative pollen
Takhtajan, A. (ed.) Anatomia seminum comparativa, morphology and its relationships to phylogeny in
vol. 3. Leningrad: Nauka, pp. 115–116. the Hamamelidae. Ann. Missouri Bot. Gard. 73:
Savolainen, V., Fay, M.F. et al. 2000. See General References. 348–381.
Soltis, D.E. et al. 2000. See General References.
Caryocaraceae
Caryocaraceae Voigt (1845), nom. cons.
G.T. P R A N C E
Trees or shrubs. Leaves trifoliolate, opposite or structure is given in Prance and Silva (1973). Bro-
alternate, the margins of the leaflets serrate, dentate chidodromous and camptodromous nervation are
or crenate or rarely entire, often with stipels at base the usual patterns, although Anthodiscus trifolia-
of leaflets; stipules 2–4, usually caducous, or 0. tus is hyphodromous. Branched sclerenchymatous
Inflorescences of terminal racemes; pedicels articu- idioblasts are present in the leaf mesophyll and
lated. Flowers large, hermaphrodite, actinomorphic; petioles of both genera. The palisade cells fre-
sepals 5(6), imbricate; petals 5(6), imbricate, cadu- quently contain crystals. Stomata are confined to
cous, distinct or rarely slightly connate at base or the abaxial leaf surface, and are usually anomocy-
connate at apex to form a calyptra in Anthodiscus; tic and occasionally either anisocytic or paracytic.
stamens numerous, 55–750; filaments usually con- The wood of Anthodiscus can easily be distin-
nate in a ring at the base, long and slender and guished from that of Caryocar by the greater
usually with some much shorter sterile interior abundance (15/sq. mm) of vessels (3/sq. mm
ones which are often recurved, apical portion with in Caryocar), the quantity of gum deposits in
numerous vesicles, the sterile filaments often with the ray cells, and the fibers without septa.
spirally arranged vesicles along entire length, or the The vessels of Caryocar vary from 74–577 mm in
filaments with a row of sterile staminodes at base of tangential diameter and are solitary or in radial
interior; anthers basifixed or attached at middle, multiples of 2–5 cells. The vessels of Anthodiscus
bilocular; the stamens frequently caducous as a have 50–100 mm mean tangential diameter and
unit together with the petals after pollination; are solitary or in multiples of 2–6 cells. Wood of
ovary compound, superior, 4(–6)-carpellate in Car- Caryocar is described in Barghoorn and Renteria
yocar and 8–20-carpellate in Anthodiscus, with as (1967), Loureiro and Silva (1968), Mello (1970)
many stylodia as carpels (a common style being and Araújo and Mattos Filho (1973).
absent), each with a distal punctiform stigma; the
carpels unilocular each with a single ovule; the F L O R A L S T R U C T U R E . In Anthodiscus the five
ovules basal, erect, anatropous or atropous, biteg- petals fall as a unit, whereas in Caryocar petals
mic or unitegmic. Fruit a drupe, with 1–4 seeds and stamens fall as a unit. The floral structure was
developing in Caryocar or 8–20 in Anthodiscus; studied by Dickison (1990) who found that Caryo-
mesocarp indehiscent, usually fatty or fleshy; endo- car has bitegmic ovules and simple stylodia, and
carp hard and woody, muricate, tuberculate or spi- Anthodiscus has unitegmic ovules and compound
nulose on outer surface, eventually splitting into stylodia (formed by complete coalescence of the
1-seeded pyrenes or mericarps. Seeds often reni- upper ventral halves of adjacent carpels). There is
form, endosperm thin or lacking, the embryo with also more fusion of major traces in Anthodiscus.
a straight, arcuate or spirally twisted radicle, a fleshy The gynoecia are richly vascularised by a complex
hypocotyle, and two small cotyledons. network of wall veins but dorsal or median carpel-
Two genera and 27 spp., in the American lary bundles are absent. Small prismatic crystals
tropics from Costa Rica to southern Brazil but and large deposits of resin-like material are widely
not native to the West Indies. distributed throughout all floral parts. The fila-
ments have unique vesicles which are enlarged
A N A T O M Y A N D M O R P H O L O G Y . A summary of epidermal cells with numerous membrane-covered
anatomical features with an emphasis on leaf wall perforations. Dickison suggests that these cells
14 G.T. Prance
function in the rapid release and uptake of water tissue rich in cytoplasm. It remains unknown
during filament erection and subsequent anther whether it is a nectary or an oil gland.
dehiscence.
The inner staminodes and the inner side of K A R Y O L O G Y . The chromosome number of three
the floral tube of Caryocar are nectariferous, species of Caryocar is 2n ¼ 46 (Ehrendorfer et al.
whereas a nectary has not been found in Antho- 1984).
discus (Vogel 1968; Dickison 1990; Matthews and
Endress 2011). On the outside of the sepals of P O L L I N A T I O N . The genus Caryocar is bat-polli-
Caryocar, Matthews and Endress (2011) found a nated (Vogel 1968), and C. brasiliense, which is
glandular structure reminiscent of an oil gland, in pollinated mainly by glossophagine bats, is self
which the cuticle is separated from the epidermis compatible but sets more fruits when crossed
and the latter is differentiated into small-celled (Barradas 1972; Gribel and Hay 1993). Sphingids
may also act as pollinators of Caryocar (Gribel
and Hay 1993), and Anthodiscus is probably
insect-pollinated (Prance and da Silva 1973).
F R U I T A N D S E E D . The fruits are drupes with 1–4

seeds in Caryocar splitting into 1-seeded cocci
and 8–20 seeds in Anthodiscus. In Caryocar the
mesocarp is soft and fleshy and the endocarp is
protected by a layer of spines in most species.
D I S P E R S A L . Little is known about dispersal of the
seeds but agoutis have been reported as dispersers
of several species of Caryocar. Gribel (1986) found
that rheas eat and defecate the stone and seed of
C. brasiliense, and were the main agent of dispersal
before they became so rare.
P H Y T O C H E M I S T R Y . Since the oil of the fruit is
comestible, there are many papers about the fatty
acid content of Caryocar oils, for C. villosum Georgi
(1929) and Hilditch and Rigg (1935), for C. brasi-
liense Ferreira and Motidome (1962) and Handro
and Barradas (1971), and for C. coriaceum Sales
(1973), Lima et al. (1981) and Alencar et al. (1983).
Caryocar oils are rich in palmitic and oleic acids,
and there is a difference between the mesocarp and
kernel oils but little difference between species.
The leaves of Caryocar brasiliense contain
various triterpenes, ellagic acid (Oliveira et al.
1968) and large amounts of condensed tannins
(K. Kubitzki, pers. comm.).
A F F I N I T I E S . Caryocaraceae have been placed in
the Theales in most traditional systems of classi-
fication and are there still in Takhtajan (2009).
Molecular studies have placed the family firmly in
Fig. 3. Caryocaraceae. A–C Caryocar harlingii. A Branch the Malpighiales, but no close relatives could be
with inflorescence. B Flower bud. C Section of fruit; note revealed in the vast sequence analyses from many
spiny endocarp. D–F Anthodiscus peruanus. D Flowering
branchlet. E Petals forming a single caducous unit. F gene regions of Wurdack and Davis (2009) and
Ovary and stylodia. (Orig.) Soltis et al. (2011). In a 3-gene analysis Soltis et al.
Caryocaraceae 15
(2007) recovered a clade, albeit with low support, KEY TO THE GENERA
in which Caryocaraceae + [Linaceae + Irvingia-
ceae] were sister to Erythroxylaceae + Rhizo- 1. Leaves opposite, calyx deeply lobed; ovary and fruit 4
phoraceae. From their careful comparative study (–6)-locular; stylodia 4; radicle of embryo straight
of the floral structure of these families, however, 1. Caryocar
Matthews and Endress (2011) concluded that a – Leaves alternate; calyx truncately dentate and reduced;
clade of Linaceae, Irvingiaceae and Caryocara- ovary and fruit 8–20-locular, stylodia 8–15; radicle of
ceae is not well supported by floral morphology, embryo spirally twisted 2. Anthodiscus
and that Caryocaraceae are morphologically most
divergent among these families. This has been
confirmed by the analysis of Xi et al. (2012) in 1. Caryocar L. Fig. 3
which Caryocaraceae are somewhat lost in a mod- Caryocar L., Mant. plantarum 2: 247 (1771); Prance &
erately supported tetratomy where they stand Silva, Fl. Neotrop. Monogr. 12: 1–75 (1973), rev.
alongside the chrysobalanoids, malpighioids and Large trees or rarely shrubs or suffrutices.
putranjivoids. Leaves opposite, often with 2–4 stipels at base of
leaflets; stipules absent or present and soon cadu-
D I S T R I B U T I O N A N D H A B I T A T S . Both genera cous. Flowers large; calyx distinctly 5(–6)-lobed;
range from Costa Rica to eastern Brazil. Antho- petals fused at base together with the base of the
discus occurs in the Guianas and western Amazo- filaments and often caducous with the filaments;
nia but is absent from central and eastern stamens 55–750, the inner often staminodial and
Amazonia. Most species are found in lowland basally with glandular nectar-secreting tissue;
rainforest and flooded riverside habitats, but spe- ovary 4(–6)-locular; ovules bitegmic. Fruit 4–6-
cies of Caryocar occur in other habitats. C. brasi- locular, a drupe with 1–4 loculi developing, and
liense is common throughout the cerrados of dehiscing into 1-seeded cocci; endocarp tubercu-
central Brazil and C. cuneatum in the arid caa- late or spinous on exterior. Seeds reniform or
tinga of the Brazilian northeast. C. montanum subreniform, the embryo with a straight to arcu-
occurs at about 1,000 m in the Guayana Highland ate radicle.
and C. gracile is endemic to white sand areas of Eighteen spp., from Costa Rica, Colombia,
north-western Amazonia. Peru, Ecuador, Venezuela, the Guianas to central
and Atlantic coastal Brazil.
E C O N O M I C I M P O R T A N C E . The uses of Caryocar
are summarised in Prance (1990). The wood of
most species is durable and finishes well, is resis- 2. Anthodiscus G.F.W. Meyer
tant to insect attack and is much used in boat
Anthodiscus G.F.W. Meyer, Prim. Fl. Esseq. 193–195
building. The fruit of most species have an edible (1818); Prance & Silva, Fl. Neotrop. Monogr. 12: 1–75
mesocarp and a kernel that is used like a nut or (1973).
for the extraction of a comestible oil rich in oleic
acid. The fruit of C. glabrum is also used as a fish Trees or shrubs. Leaves alternate; stipels 0;
poison (Prance 1973; Kawanishi et al. 1986), the stipules 0. Flowers medium sized; calyx cupuli-
mesocarp of C. brasiliense to prepare a liqueur, form with very small, reduced, indistinct dentate
and there are many other local indigenous uses. lobes; petals circumscissile at base and fused at
apex to form a caducous calyptra; stamens
C O N S E R V A T I O N . Caryocar costaricense has been 100–280; ovary 8–12(–15)-locular; ovules uniteg-
seriously overexploited for its wood and is listed mic. Fruit an 8–15-locular drupe with most loculi
in Appendix 2 of the CITES Treaty, and therefore developing. Seeds small, laterally compressed.
requires permits for trade. Nine spp. in Guyana, Venezuela, Colombia,
Peru and western Amazonia, with one sp. in
16 G.T. Prance
Costa Rica and Chocó, Colombia and one in Handro, W., Barradas, M.M. 1971. Sôbre os óleos do fruto
Atlantic coastal Brazil. e da semente do pique – Caryocar brasiliense Camb.
(Caryocaraceae), pp. 110–113. In: III Simpósio sôbre
o Cerrado. Ed E. Bl€ ucher, Univ. São Paulo.
Hilditch, T.P., Rigg, J.G. 1935. The component glycerides
of piqui-a fats. J. Soc. Chem. Indust. 54: 109.
Selected Bibliography Kawanishi, K., Raffauf, R.F., Schultes, R.E. 1986. The Car-
yocaraceae as a source of fish poisons in the north-
west Amazon. Bot. Mus. Leafl. 30: 247–253.
Alencar, J.W., Alves, P.B., Craveiro, A.A. 1983. Pyrolysis Lima, M.T., Maia, G.A., Guedes, B.L., Oria, H.F. 1981.
of tropical vegetable oils. J. Agric. Food Chem. 31: Composição de acidos graxos da fração lipidica do
1368–1270. piqui (Caryocar coriaceum Wittm.). Ciencia Agron.
Araújo, P.A. de M., Mattos Filho, A. de 1973. Estrutura das 12: 93–96.
madeiras de Caryocaraceae. Arch. Bot. Rio de Janeiro Loureiro, A.A., Silva, M.F. 1968. Catálogo das madeiras do
19: 5–47. Amazônia 1: 154–164. Belém.
Barghoorn, A.W., Renteria, R.M. 1967. Estudio anatomico Matthews, M.L., Endress, P.K. 2011. See General References.
y fisico-mecanico del cagui (Caryocar costaricense Mello, E.C. 1970. Estudo anatômico das madeiras do gên-
Donn. Sm.). Bol. Inst. Forest. Lat.-Amer. Merida 24: ero Caryocar Linn. Brasil Florestal 1: 54–62.
35–57. Oliveira, M.M. de, Gilbert, B., Mors, W.B. 1968. Triter-
Barradas, M.M. 1972. Informação frutificação e dispersão penes in Caryocar brasiliense. An. Acad. Brasil. Ciên-
do piqui Caryocar brasiliense Camb. (Caryocara- cias. 40: 451–452.
ceae). Ciência e Cultura 24: 1063–1068. Prance, G.T. 1973. Ethnobotanical notes from Brazil.
Dickison, W.C. 1990. A study of the floral morphology Econ. Bot. 26: 221–237.
and anatomy of the Caryocaraceae. Bull. Torrey Bot. Prance, G.T. 1990. The genus Caryocar L. (Caryocara-
Club 117: 123–137. ceae): an underexploited tropical resource. Adv.
Ehrendorfer, F., Morawetz, W., Dawe, J. 1984. The neo- Econ. Bot. 8: 177–188.
tropical angiosperm families Brunelliaceae and Car- Prance, G.T., Silva, M.F. da 1973. Monograph of Caryocar-
yocaraceae: first karyosystematical data and aceae. Flora Neotropica 12: 1–75. New York: Hafner.
affinities. Pl. Syst. Evol. 145: 183–191. Sales, F.J.M. 1973. O oleo no fruto de piquizeiro, Caryocar
Ferreira, P.C., Motidome, M. 1962: Estudo quimico do coriaceum Wittm. Turrialba 23: 108–109.
óleo de piquı́. An. Fac. Farm. Odont. Univ. São Soltis, D.E., Gitzendanner, M.A., Soltis, P.S. 2007. A 567-
Paulo 19: 25–30. taxon data set for angiosperms: the challenges posed
Georgi, C.D.V. 1929. Piqui-a fruit oils. Malayan Agric. J. by Bayesian analyses of large data sets. Int. J. Pl. Sci.
17: 166–170. 168: 137–157.
Gribel, R. 1986. Ecologia da polinização e da dispersão de Soltis, D.E. et al. 2011. See General References.
Caryocar brasiliense Camb. (Caryocaraceae) na Takhtajan, A. 2009. See General References.
região do Distrito Federal. M. Sc. thesis, Univ. de Vogel, S. 1968, 1969. Chiropterophilie in der neotro-
Brası́lia, xii + 109 pp. pischen flora. Flora 157: 562–602; 158: 195–202,
Gribel, R., Hay, J.D. 1993. Pollination ecology of Caryocar 289–323.
brasiliense (Caryocaraceae) in Central Brazil cerrado Wurdack, K.J., Davis, C.C. 2009. See General References.
vegetation. J. Trop. Ecol. 9: 199–211. Xi, Z. et al. 2012. See General References.
Centroplacaceae
Centroplacaceae Doweld & Reveal (2005).
K. K U B I T Z K I
Trees. Leaves alternate, stipulate, colleters + or 0. and Davis (2009) believe that the aril arises at the
Inflorescences racemose; pedicels articulated. exostome, but Leandri (1957: 207) thought it to be
Flowers bisexual or unisexual and then plants an outgrowth of the "fruit axis".
dioecious, 5-merous, haplostemonous, stamens The seeds of both genera agree in the small
antesepalous; gynoecium syncarpous, ovary 2- size of the embryo, the copious endosperm, and
or 3-locular; stylodia widely divergent, with the 1-layered exotegmen of ribbon-like cells (see
slightly expanded stigmas; ovules 2 per locule, Corner 1976, figs. 80 and 82; Stuppy 1996: 202 and
collateral. Capsule loculicidal. Seed 1 per locule, t. 85; Tokuoka and Tobe 2001). Such a seed coat is
almost completely enveloped by an exostomal also found in genera of the Phyllanthoideae.
fleshy, sheet-like, coloured aril; seed coat exoteg-
mic, thick-walled, ribbon-shaped; endosperm R E L A T I O N S H I P S . This family concept is the out-
copious; embryo small. come of recent molecular studies. Bhesa, for long
A family comprising two genera and seven considered a strange element within Celastraceae
spp., West Africa, Indo-Malesia. in terms of gross morphology and anatomy, has
been recognised as malpighialean by Zhang and
M O R P H O L O G Y A N D A N A T O M Y . Bhesa has vessel Simmons (2006). Centroplacus, after having been
elements with scalariform perforation plates and assigned to Flacourtiaceae and Celastraceae, was
paratracheal parenchyma +; stomata are anisocytic referred by Hutchinson (1912) to Euphorbiaceae,
(Centroplacus) and laterocytic (Bhesa); the ovules where its affinities remained contentious. The
are epitropous (Centroplacus) and apotropous absence of an obturator and the minute embryo,
(Bhesa). together with the presence of an aril may exclude
Centroplacus from the biovulate Euphorbiaceae.
P O L L E N M O R P H O L O G Y . The pollen is tricolpo- The multi-gene analysis of Wurdack and Davis
rate, that of Centroplacus tectate and microperforate (2009) provided strong support for a clade con-
and very similar to that of (uniovulate!) Microdes- sisting of Centroplacus and Bhesa, which is weakly
mis (see also the discussion by Punt 1962: 89), supported as sister to a clade of Malpighiaceae +
whereas that of Bhesa is finely striate (Wurdack Elatinaceae (Xi et al. 2012). These four families
and Davis 2009). share axile placentation, crassinucellar ovules
without endothelium, and sepals persistent in
F R U I T A N D S E E D . The loculicidal fruits contain fruit (Xi et al. 2012).
a single seed per locule (one ovule aborts),
invested with a coloured, sheet-like, fleshy aril
that envelops nearly the entire seed. It is orange KEY TO THE GENERA
or red in Bhesa and red in Centroplacus; this may
be related to bird dispersal. Stuppy (1996) cites a
1. Flowers unisexual, plants dioecious; leaves denticulate;
collector’s note according to which the valves of pistillate flowers apetalous; ovary 3-merous
the mature fruit of Centroplacus fall away 1. Centroplacus
completely and leave behind the brown seed – Flowers bisexual; leaves entire; pistillate flowers petali-
wrapped into a fleshy bright orange aril. Wurdack ferous; ovary 2(1)-merous 2. Bhesa
18 K. Kubitzki
1. Centroplacus Pierre to racemose. Flowers bisexual, 5-merous; calyx

Centroplacus Pierre, Bull. Mens. Soc. Linn. Paris II, 14: 144 contorted; anthers extrorse or introrse; pollen
(1899); Hutch., Fl. Trop. Afr. 6(1): 629 (1912); Webster, spheroidal, 3-colporate, finely striate; disk lobed
Ann. Missouri Bot. Gard. 81: 54 (1994); Radcl.-Sm., Gen. or not; ovary 2(1)-locular, ovules basal, erect, apo-
Euphorb. 78, fig. 8 (2001). tropous. Fruit capsular, fusiform or deeply 2-lobed,
Dioecious trees; indumentum simple. Leaves loculicidally dehiscent. Seeds with coloured, sheet-
2-ranked, denticulate, eglandular; stipules minute, like aril; exotegmic cells fibrous.
persistent. Inflorescences axillary, paniculate. Flow- Six spp., Indo-Malesia.
ers unisexual, pedicellate; staminate flowers: sepals
and petals 5, imbricate; disk extrastaminal, cupular,
its lobes antepetalous; stamens 5, anthers basifixed, Selected Bibliography
introrse, dehiscence oblique-apical; pollen oblate-
spheroidal, 3-colporate, tectate-microperforate; Corner, E.J.H. 1976. See General References.
Forman, L.L. 1966. The reinstatement of Galearia Zoll. & Mor.
pistillode columnar; pistillate flowers apetalous; and Microdesmis Hook.f. in the Pandanaceae, with
disk annular; ovary 3-locular; ovules subapical, appendices by C. R. Metcalfe and N. Parameswaran.
epitropous; stylodia spreading. Fruits septicidal Kew Bull. 20: 309–321.
and loculicidal capsules dehiscing from the base; Leandri, J. 1957. Notes systématiques sur les Euphorbiacées-
Phyllanthées de Madagascar. Mém. Inst. Sci. Madagascar,
columella 0. Seeds 1 per locule, carunculate, with Sér. B, Biol. Vég. 8: 205–261.
black and shining testa, exotegmic cells flattened, Nowicke, J.W. 1984. A palynological study of the Pandaceae.
elongate at right angles to mesotegmic fibres; coty- Pollen Spores 26: 31–42.
ledons twice as long as radicle. Pierre, P.J.L. 1894. Flore forestière de la Cochinchine.
Vol. 3. Paris: Octave Doin.
A single sp., C. glaucinus Pierre, rain forests Punt, W. 1962. Pollen morphology of the Euphorbiaceae
of West Africa (Cameroon to Gabon). with special reference to taxonomy. Wentia 7: 1–116.
Stuppy, W. 1996. Systematische Morphologie und Anatomie
der Samen der biovulaten Euphorbiaceen. Dissert.,
2. Bhesa Buch.-Ham. ex Arn. Univ. Kaiserslautern.
Bhesa Buch.-Ham. ex Arn., Edinb. New Philos. Soc. J. 16: Tokuoka, T., Tobe, H. 2001. Ovules and seeds in subfamily
315 (1834); Simmons, this book series vol. VI: 41 (2004). Phyllanthoideae (Euphorbiaceae): structure and syste-
matic implications. J. Plant Res. 114: 75–92.
Buttressed trees, glabrous. Leaves spiral, con- Wurdack, K.J., Davis, C.C. 2009. See General References.
duplicate, entire; petioles pulvinate apically; sti- Xi, Z. et al. (2012). See General References.
Zhang, L.-B., Simmons, M.P. 2006. Phylogeny and delimi-
pules with colleters, almost encircling the stem, tation of the Celastrales inferred from nuclear and
caducous. Inflorescence axillary, thyrsopaniculate plastid genes. Syst. Bot. 31: 122–137.
Chrysobalanaceae
Chrysobalanaceae R. Br. (1818), nom. cons.
G.T. P R A N C E
Trees, shrubs or suffrutices. Leaves alternate, understorey shrubs, especially in the genus
simple, margins entire, pinnately nerved; stipules Hirtella. At least six species are geoxylic suffru-
small and caducous to large and persistent, some- tices, one in Florida, two in the Cerrado region of
times adnate to petiole. Inflorescence racemose, Brazil and three (Parinari capensis and two spe-
paniculate or cymose. Flowers bracteate and usu- cies of Magnistipula) in southern Africa. Many
ally with 2 prophylls, actinomorphic to zygomor- species are tall trees of the rainforest canopy and
phic, bisexual or rarely polygamous or unisexual, a few, especially in Parinari, are emergents. A few
markedly perigynous; receptacle short to elon- species of Dactyladenia and Hirtella have scan-
gate-cylindrical, sometimes gibbous at base; disk dent branches, but there are no lianas. Those
always present, forming a lining to receptacle species which have been studied exhibit the archi-
or an annular or shortly tubular structure at its tectural model of Troll. Leaves are simple, alter-
mouth; sepals 5, imbricate; petals 5, imbricate, nate and usually medium sized, but very large in a
often unequal, rarely unguiculate, sometimes few species (up to 50 cm long in Licania gentryi).
absent; stamens 2–100 (300) inserted on margin There are frequently glands on the undersurface
of disk or basally adnate to it, forming a complete or on the petioles. Stipules are always present but
circle or unilateral in zygomorphic flowers; fila- sometimes very early caducous. They range from
ments free, connate at base or ligulately connate, minute to quite large in some species of Licania,
included to far exserted; anthers dorsifixed, lon- Parinari and Magnistipula. They are either axil-
gitudinally dehiscent; ovary superior, inserted at lary or, in many species of Licania, inserted on
base, middle or mouth of the receptacle, either the lower portion of the petiole. Seven species of
unilocular with 2 ovules, or bilocular with 1 ovule Hirtella section Myrmecophila have myrmecoph-
in each loculus; ovules erect, epitropous with ilous inflations at the junction of the leaf lamina
micropyle directed towards base; style filiform with the petiole. These are inhabited by ants of
arising from receptacle at base of ovary; stigma the genera Allomerus, Azteca or Solenopsis. Mag-
distinctly or indistinctly 3-lobed. Fruit a dry or nistipula bimarsupiata also has two small infla-
fleshy drupe; endocarp thin and bony to thick tions at the base of the lamina, and some
and woody, often with a special mechanism for internodes are swollen and perforated. It also
seedling escape, often densely hairy within. Seed has the coarse hispid tomentum characteristic of
erect, almost exalbuminous; cotyledous plano- ant plants. The trunks of tree species may be
convex, fleshy, rarely ruminate; germination either cylindric or buttressed. The bark of most
cryptocotylar or phanerocotylar. species chips into small fragments with a sharp
A pantropical family with eighteen genera and metallic ring when hit with a machete, due to the
423 of the 531 spp. in the Neotropics, 56 in Africa presence of abundant silica grains.
and Madagascar, and 43 in Asia and the Pacific.
Extending beyond the tropics with one species F L O W E R S T R U C T U R E . The flowers are actino-
each into the southern U.S.A. and southern Brazil. morphic with the ovary inserted at the base of
the receptacle in Chrysobalanus, Licania and
V E G E T A T I V E M O R P H O L O G Y . The majority of Parastemon (Fig. 4), and zygomorphic with the
species of Chrysobalanaceae are leptocaul trees ovary inserted at the mouth or middle of the
but there are also many savannah shrubs and receptacle in the remaining genera (Fig. 5). The
20 G.T. Prance
Fig. 4. Chrysobalanaceae. Licania granvillei. A Flowering

branchlet. B Flower; note lack of corolla. C Medial section
of flower showing gynobasic style. D Entire and sectioned
fruit with attachment scar at top. (Reproduced with kind
permission of the artist Bobbi Angell)
calyx is imbricate in bud, 5-lobed, entire; stipitate

glands occur in some species of Dactyladenia and
Hirtella. There are usually 5 free, imbricate petals,
except in Licania subgenus Licania and section
Leptobalanus which are apetalous. In a few spe-
cies distinctly unguiculate petals occur, especially Fig. 5. Chrysobalanaceae. Parinari montana. A Flower-
ing branchlet. B Abaxial leaf surface with part of woolly
in Kostermanthus where the 2 posterior petals are pubescence removed to show venation. C Stipulate node
larger and enclose the stamens in bud. Stamens with glandular petiole and base of panicle with prophyll.
range from 2 in Parastemon to over 300 in some D Flower. E Same, medial section, with perched gynoe-
species of Couepia. The filaments are usually dis- cium, cup-shaped hypanthium and petal. F Fruit with
attached scar. G Seed with corrugated endocarp. (Repro-
tinct, but are connate into a unilateral, strap-like duced with kind permission of the artist Bobbi Angell)
ligule in Acioa, Dactyladenia and Kostermanthus;
they may be connate for up to half their length in E M B R Y O L O G Y . The embryology was studied by
a few species of Licania and in Chrysobalanus. Tobe and Raven (1984). The tetrasporangiate
The stamens may be inserted around a complete anthers have five-layered walls including a glan-
circle or unilateral, and the filaments either dular tapetum with cells two-nucleate. Cytokinesis
exserted or included. The anthers are dorsifixed in the microspore mother cells is simultaneous.
and dehisce laterally. A disk is always present, The nucellus is very small and the nucellar tissue
forming a lining to the receptacle or an annular soon disintegrates, except for the megaspore or
structure at its mouth. The ovary is superior and embryo sac; antipodal cells are absent. The ovule
inserted at the base, middle or mouth of the is bitegmic; the inner integument (ii) and the
receptacle, either unilocular with two ovules or outer integument (oi) are initiated dermally; the
bilocular with one tenuinucellate ovule in each ii is up to five or eight cells thick and the oi more
loculus. The ovules are erect and epitropous, with than five cells thick; the inner epidermis of the ii
the micropyle directed towards the base. The develops into the endothelium, which directly
style is filiform and always arises from the base borders the embryo sac and accumulates starch
of the ovary (gynobasic); it usually equals or grains; the micropyle is formed by both integu-
exceeds the stamens in length. ments. Endosperm formation is of the nuclear
Chrysobalanaceae 21
type and the seed is exalbuminous. The young range of dispersal agents. The fruit may be dry
seed coat is composed of both testa and tegmen. or fleshy, the endocarp thick or thin, fibrous or
bony, often with a special mechanism for the
P O L L E N M O R P H O L O G Y . The pollen of Chryso- seedling to escape, two basal obturators in Par-
balanaceae is very uniform. The pollen is tricol- inari and lateral plates in Maranthes, Grangeria
porate but in some species four furrows may and Parastemon. There are longitudinal lines of
occur. There are no special features, excepting dehiscence in Chrysobalanus and Hirtella. The
equatorial constrictions. The grains are usually seed is erect, exalbuminous, with planoconvex
distinctly triangular in shape in polar view, fleshy cotyledons, and ruminate cotyledons in
except when four-furrowed; they are elliptical to Atuna. Chrysobalanus icaco and several species
circular in equatorial view and are oblate-sphe- of Atuna are dispersed by ocean currents. Several
roidal, prolate-spheroidal or subprolate in shape. riverine species of Licania drop their fruit into
The polar length x 100 divided by the equatorial rivers at flood time, and are eaten and presum-
length ¼ 85–150, but the size is quite variable ably dispersed by fish. Various species including
from one genus to another; the polar area is Licania elaeosperma and Acioa guianensis are
usually small, sometimes medium, but never water-dispersed by rivers. Species of Couepia,
large. The exine is medium to rather thick with Licania and Parinari are frequently eaten by
very little patterning on the walls, usually sca- bats. Licania splendens is dispersed by the fruit
brous to verrucose. See Prance and White pigeon, Ducula aenea. Various species of Pari-
(1988), and also Barth and Silva (1963), nari are known to be dispersed by bats, elephants,
Demchenko (1973), Patel et al. (1983). baboons and other primates, a scatter-hoarding
squirrel, fruit pigeons, rheas, emus, agoutis and
K A R Y O L O G Y . The chromosomes have been fish. Agoutis have been observed transporting the
counted for nine species: 2n ¼ 20 in Maranthes fruits of two species of Couepia. Most species of
and Parinari and 2n ¼ 22 in Chrysobalanus, Hirtella, with their small fleshy fruits, are bird-
Dactyladenia and Licania (Mangenot and dispersed. Maranthes corymbosa is dispersed by
Mangenot 1962; de Souza 1979). fruit pigeons and hornbills.
P O L L I N A T I O N . Chrysobalanaceae have a wide P H Y T O C H E M I S T R Y . The chemistry of Chrysoba-

range of pollinators. The small-flowered genera lanaceae has been little studied, except for fatty
Chrysobalanus, Parinari, Exellodendron, Licania, acids in the seeds, including some quite unusual
etc. are mainly pollinated by a wide variety of ones: a-elaeostearic acid (Z-octadeca-9c, 11t, 13t-
small bees. Most species of Couepia are night- trienoic), a-licanic acid (4-keto-a-elaeostearic),
flowering and are pollinated by hawk-moths, with a-parinaric acid (Z-octadeca-9c, 11t, 13t, 15c tet-
the exception of the two flagelliflorous species C. raenoic) and 4-keto-a-parinaric acid. For details,
longipendula and C. dolichopoda which are polli- see Table 2 in Prance and White (1988). As a
nated by bats (Vogel 1968–1969). Maranthes poly- result, there are several useful oils in Chrysobala-
andra is also bat-pollinated (Lack 1978). Hirtella is naceae (see section on Economic Importance
mainly pollinated by butterflies, except for H. below). Phenolics have been studied in a few
rugosa from the mountains of Puerto Rico. The species and the following compounds have been
bright red petals of this species do not open fully reported: myricetin, delphinidin, quercetin, cya-
but form a tube, and it is hummingbird-pollinated. nidin, kaempferol, quercitin 3-O-glycoside, nar-
Hummingbirds have also been observed visiting ingenin 7-O-clycoside, quercetin aglycone (Gibbs
the flowers of several other species of Chrysobala- 1974; Coradin et al. 1985). All species analysed
naceae, especially Couepia, but are probably only contained kaempferol and quercetin, and only
secondary pollinators. Souza (1979) showed Chry- some contained myricetin.
sobalanus icaco to be self-compatible.
S Y S T E M A T I C S . Prance and White (1988) divided
F R U I T A N D S E E D . The fruits of Chrysobalana- the family into four tribes. More recent molecular
ceae are always drupes and they are quite work (Yakandawala et al. 2010; C.A. Sothers, in
uniform, yet have become adapted to a wide prep.) does not support this tribal division,
22 G.T. Prance
nor those of other earlier workers. Molecular data Several species of Parinari have edible fruits or
show the family as presented here to be a seeds, and charred kernels of a species of Parinari
well-defined monophyletic group, but they did have been found at archaeological sites in Malawi.
not support obvious tribal subdivisions. The In Ambuina a dish called koku koku is prepared
tribes of Prance and White were found to be from the mashed seeds of Atuna excelsa mixed
paraphyletic and are not recognised here. with fish, ginger, onions, etc. A cooking oil is
extracted locally in Amazonia from Acioa edulis
A F F I N I T I E S . Most early classifications regarded and Couepia longipendula. The oil of Licania rigida
the Chrysobalanaceae as a subfamily of the Rosa- and Afrolicania elaeosperma was formerly used in
ceae. It is clearly different in terms of many fea- paints and varnishes as a substitute for tung oil,
tures such as the gynobasic style, erect ovules and and L. arborea produces a flammable oil which was
wood anatomy. Most later authors who treated used for lighting by local people in Mexico. The
Chrysobalanaceae as a family distinct from Rosa- wood of Chrysobalanaceae is extremely hard to
ceae left it in Rosales. Dahlgren and Thorne work due to the presence of silica. It has many
(1984) questioned this placement without offer- local uses, especially for marine and river pilings
ing an alternative, but they suggested it might be because of its resistance to marine borers. Some
useful to look at the Myrtales. Tobe and Raven species are used locally for fuel and charcoal.
(1984) showed that the embryology does not Throughout Amazonia the bark of various species
resemble that of Myrtales, and suggested Theales. of Licania and Couepia is burnt and the ash mixed
Many other placements have been postulated on with clay to strengthen pottery because of the silica
morphological grounds, including Linaceae, granules. In the Solomon Islands the seeds of
Polygalaceae, Limnanthaceae, Dichapetalaceae, Atuna excelsa, known as putty nut, is used for
Trigoniaceae, Geraniaceae, Tropaeolaceae, Sapin- caulking boats. The oil from the seed of the same
daceae, Rhizophoraceae, Vochysiaceae and Pro- species is used as a hair dressing in the Caroline
teaceae. The Chrysobalanaceae do not even Islands and as a massage oil in Samoa and Tonga.
remotely resemble most of these families. How- In Brazil, Couepia subcordata and Licania tomen-
ever, molecular data suggested that Chrysobala- tosa commonly serve as shade trees.
naceae are a member of Rosidae (rosid group 1)
nearest to Trigoniaceae (Chase et al. 1993) in a C O N S E R V A T I O N . Many species of Chrysobala-
clade that later became the broadly construed naceae are of extremely restricted distribution
order Malpighiales (APG 1998). Further molecu- and grow in highly threatened areas such as the
lar multigene analyses placed Chrysobalanaceae Atlantic coastal forest of Brazil or the lowland
into a well-supported clade including Chrysoba- forest of Western Ecuador, and are therefore
lanaceae + Euphroniaceae and Trigoniaceae + potentially threatened with extinction. The
Dichapetalaceae, with Balanopaceae basal to this genus Grangeria of two species in Madagascar
clade (e.g. Davis and Chase 2004; Wurdack and and the Mascarenes must be one of the most
Davis 2009), a grouping that (without Balanops) endangered, and the states of the monotypic
had been recognised by Hallier (1921) as Chryso- genera Bafodeya and Neocarya from West Africa
balanaceae (s. l.). The close relationship among is critical.
these four families is also supported by important
traits of their floral structure that have been
KEY TO THE GENERA
revealed in the comparative study of Matthews
and Endress (2008). 1. Ovary inserted at or near base of receptacle 2
– Ovary inserted laterally at or near mouth of
E C O N O M I C I M P O R T A N C E . The fruits of many receptacle 5
2. Endocarp with distinct longitudinal ridges
species are edible and are frequently used by local corresponding with lines of dehiscence
peoples. Those of Chrysobalanus icaco are tinned 5. Chrysobalanus
or bottled in Venezuela and Colombia under the – Endocarp without longitudinal ridges, indehiscent or
name icacos. Various species of Couepia, especially with two lateral plates 3
C. bracteosa and C. rufa, are sold in local markets. 3. Flowers unisexual (Africa) 2. Afrolicania
Chrysobalanaceae 23
– Flowers bisexual (South America and Malesia) 4 GENERA OF CHRYSOBALANACEAE

4. Endocarp without lateral plates which allow seedling
to escape 13. Licania 1. Acioa Aubl.
– Endocarp dehiscing on germination by means of a
Acioa Aubl., Hist. Pl. Guiane 2: 698, t. 280 (1775).
pair of large lateral plates 16. Parastemon
5. Endocarp with two small basal plugs (obturators); Trees or shrubs. Leaves with 1 or 2 pairs of con-
lower leaf surface usually with stomatal crypts 6 spicuous glands at base of lamina and several
– Endocarp without two basal plugs but sometimes ger- smaller discoid glands, glabrous on lower surface.
minating by means of two large lateral plates; leaf
lower surface rarely with stomatal crypts 7
Inflorescence a rather lax thyrse with flattened
6. Receptacle turbinate-campanulate; fertile stamens
axes; bracts and prophylls not enclosing flower
6–8 17. Parinari buds, eglandular; receptacle obconic and slightly
– Receptacle saccate; fertile stamens 12–17 curved or cyathiform, glabrous within; petals 5,
18. Neocarya exceeding sepals; stamens 10–20, ligulately con-
7. Stamens far exserted beyond calyx lobes 8 nate or free, unilateral, glabrous; ovary unilocu-
– Stamens not or barely exceeding calyx lobes 14 lar, inserted at mouth of receptacle. Fruit without
8. Stamens united into a ligule 9 any special mechanism to allow seedling to
– Stamens free to base or nearly so, not ligulate 11 escape; endocarp thick, hard, fibrous. Germina-
9. Sepals very unequal in size, 3 large, 2 very small, the tion phanerocotylar, first two eophylls opposite,
outer sepals with 1 or 2 large discoid glands otherwise alternate.
1. Acioa Four spp. in Venezuela, the Guianas and
– Sepals subequal, without discoid glands on surface Amazonian Brazil.
10
10. Two posterior petals unguiculate, enclosing stamens
in bud 12. Kostermanthus 2. Afrolicania Mildbr.
– Posterior petals not unguiculate 7. Dactyladenia
11. Ovary bilocular 12 Afrolicania Mildbr., Notizbl. Bot. Gart. Berlin-Dahlem 7:
483 (1921).
– Ovary unilocular 13
12. Fruit dehiscing by two lateral plates; cotyledons not Small to medium-sized tree. Leaves usually gla-
ruminate; exocarp smooth 15. Maranthes brous, with two glands on petiole. Inflorescence
– Fruit with no lateral plates; cotyledons ruminate; exo- many-flowered terminal and subterminal panicle
carp warted crustaceous 3. Atuna
13. Stamens 3–9; fruit dehiscing by longitudinal lines; of racemes; bracts small, membranous, eglandu-
endocarp thin, bony 10. Hirtella lar. Flowers polygamous andro-dioecious; recep-
– Stamens 15–300; fruit indehiscent; endocarp thick, tacle flattened-turbinate, pubescent within; petals
fibrous 6. Couepia absent; stamens ca. 20, around complete circle,
14. Ovary unilocular 15 included, free; ovary unilocular, inserted at base
– Ovary bilocular 16 of receptacle, absent in male flowers but repre-
15. Receptacle symmetrical; fruit dehiscing by two lateral sented by a slight swelling. Fruit a dry drupe,
plates 9. Grangeria
epicarp warted, without a mechanism for seedling
– Receptacle gibbous; fruit indehiscent
14. Magnistipula escape. Germination cryptocotylar, eophylls
16. Leaf undersurface with stomatal cavities; fruit exocarp alternate.
sparsely lenticellate; receptacle much swollen to one One sp., A. elaeosperma Mildbr., widespread
side 4. Bafodeya in W and W Central Africa from Sierra Leone to
– Leaf undersurface glabrous or arachnoid pubescent; Gabon.
receptacle symmetrical or only slightly swollen to one
side 17
17. Endocarp opening by lines of weakness which allow 3. Atuna Rafin.
seedling to escape 11. Hunga
Atuna Rafin., Sylva Tellur.: 153 (1838).
– Endocarp without lines of weakness 18
18. Receptacle markedly asymmetrical; endocarp thick, Trees. Leaves with a pair of glands on the midrib
hard and bony, not ridged at base 14. Magnistipula
at or near base of lower surface, glabrous on
– Receptacle only slightly swollen to one side; fruit with
hard, bony endocarp thin, with a ridge on one side at
lower surface. Inflorescence a raceme or a
the base 8. Exellodendron sparsely branched contracted panicle; bracts and
24 G.T. Prance
prophylls not enclosing flower buds in groups, Three spp., one widespread in tropical Africa,
eglandular; receptacle obconic, pubescent America and the Caribbean, one endemic to the
throughout within; petals 5, shorter than sepals; West Indies, and one in submontane forests of
stamens 10–25; filaments distinct, far exceeding Venezuela.
sepals, glabrous, unilateral, tiny denticulate sta-
minodes opposite; ovary bilocular, inserted at
mouth of receptacle. Fruit without any mecha- 6. Couepia Aubl.
nism for seedling to escape; endocarp hard and Couepia Aubl., Hist. Pl. Guiane 1: 519, t. 207 (1775);
thick, cotyledons strongly ruminate. Germination Prance, Fl. Neotrop. 9: 202 (1972).
cryptocotylar, eophylls alternate. Trees or shrubs. Leaves often with 1 or 2 pairs of
Eight spp. from southern India eastwards to glands at base of lamina, sometimes with several
Samoa, and most abundant in the Malay Penin- small marginal glands especially near apex, gla-
sula, Borneo and Indonesia. brous or arachnoid pubescent beneath or rarely
with stomatal crypts. Inflorescence most often a
4. Bafodeya Prance congested thryse or raceme, rarely flowers soli-
tary or densely crowded in a long-pedunculate
Bafodeya Prance in F. White, Bull. Jard. Bot. Nat. Belg. 46:
271 (1976). compound corymb; bracts and prophylls usually
not enclosing buds in small group, eglandular;
Small trees. Leaves with stomatal crypts filled with receptacle turbinate to narrowly cylindric, usually
densely matted hairs on lower surface, eglandu- glabrous inside except at throat, hairy throughout
lar. Inflorescence a terminal cymose panicle; in a few species; petals 5, more or less equalling
bracts and prophylls not enclosing flower buds, sepals; stamens 15–100 or rarely more; filaments
eglandular; receptacle obliquely campanulate, far exceeding sepals, free, glabrous, usually form-
markedly zygomorphic, much swollen to one ing a complete circle less frequently unilateral.
side, pubescent within; petals 5, equalling sepals; Ovary unilocular, inserted at mouth of receptacle.
stamens ca. 7; filaments more or less equalling Fruit without any mechanism to allow seedling to
sepals, unilateral with 4–10 staminodes opposite; escape; endocarp hard, thick, granular. Germina-
ovary bilocular, inserted at mouth of receptacle. tion cryptocotylar, eophylls alternate.
Fruit without special mechanism for seedling to Seventy one spp., all neotropical, ranging
escape; endocarp hard, thin, smooth. from Mexico to S Brazil, but most abundant in
A single sp., B. benna (Scott Elliot) Prance, in the Guianas and Amazonia.
W Africa in Sierra Leone and adjacent Guinea.
7. Dactyladenia Welw.
5. Chrysobalanus L.
Dactyladenia Welw., Apont., Ann. Cons. Ultram. 1: 572
Chrysobalanus L., Gen. pl.: 365 (1737); Sp. pl.: 513 (1753). (1859); Prance & White, Philos. Trans. Roy. Soc. London
B, 320: 133–145 (1988), rev.
Shrubs to large trees. Leaves glabrous, usually
with two or more glands on lower surface. Inflo- Small trees or shrubs occasionally scandent.
rescence a few-flowered short raceme of cymules Leaves glabrous or with stiff appressed hairs,
or cymose or a subsessile fascicle; bracts and rarely with lanate arachnoid pubescence beneath.
prophylls not enclosing flower buds, eglandular; Inflorescence a raceme or a panicle of racemes,
receptacle cupuliform, pubescent within; petals 5, rarely a subcapitate spike; bracts and prophylls
longer than sepals; stamens 12–26, filaments not enclosing flower buds, often with stalked or
twice as long as sepals, slightly united in groups sessile glands; receptacle elongate cylindrical to
at base, around complete circle, hairy; ovary uni- obconic-tubular, glabrous within except at throat;
locular, inserted at base of receptacle. Fruit lon- sepals more or less equal; petals 5, equalling
gitudinally ridged, with lines of fracture that sepals; stamens 10–75, far exceeding calyx lobes,
allow the seedling to escape, endocarp thin and filaments united into a ligule for most of length,
bony. Germination cryptocotylar, eophylls alter- unilateral with short staminodes opposite; ovary
nate. 2n ¼ 22. unilocular, inserted at mouth of receptacle. Fruit
Chrysobalanaceae 25
with no plates or lines of dehiscence, endocarp cophilous swellings at base. Inflorescence raceme,
thin and hard. Germination cryptocotylar, thyrse, corymb or panicle; bracts and prophylls
eophylls opposite. 2n ¼ 22. with sessile or stipitate glands or eglandular not
Thirty spp. in tropical Africa, mostly in the enclosing flower buds in groups; receptacle sub-
Guineo-Congolian region. campanulate to narrowly cylindrical, glabrous
within except at throat; sepals usually almost
equal; petals 5, not exceeding sepals; stamens
8. Exellodendron Prance
3–9, filaments far exserted, free, glabrous, usually
Exellodendron Prance, Fl. Neotrop. 9: 195, t. 31–32 (1972). unilateral with short staminodes opposite; ovary
Trees or shrubs. Leaves with a pair of small glands unilocular, inserted at mouth of receptacle. Fruit
or ill-defined glandular areas on upper surface at fleshy, usually with longitudinal lines of dehis-
junction with petiole, glabrous or lanate pubescence, endocarp thin, bony. Germination crypto-
cent beneath. Inflorescence a simple or branched cotylar cataphylls ca. 5, minute, eophylls
raceme of small, congested cymes; bracts and alternate.
prophylls not enclosing flower buds, eglandular; 107 spp. in tropical America from Mexico to S
receptacle subcampanulate, slightly swollen on Brazil, and one sp. in E Africa and Madagascar.
one side, pubescent within; petals 5, equalling
sepals; stamens ca. 7; filaments equalling sepals, 11. Hunga Pancher ex Prance
free, glabrous; ovary bilocular, inserted at mouth
Hunga Pancher ex Prance, Brittonia 31: 79 (1979).
of receptacle tube. Fruit without any mechanism
for seedling to escape; endocarp thin, smooth, Shrubs or small trees. Leaves with a pair of often
bony. obscure marginal glands towards base, lower sur-
Five spp. in Amazonia, the Guianas, central face glabrous or lanate pubescent. Inflorescence a
and eastern Brazil. few-flowered terminal or axillary raceme of cym-
ules; bracts and prophylls not enclosing flower
buds, eglandular; receptacle subcampanulate,
9. Grangeria Commerson ex Juss.
slightly asymmetric, pubescent within; petals 5,
Grangeria Commerson ex Juss., Gen. pl.: 340 (1789). not exceeding sepals; stamens 5–9, filaments
Trees or shrubs. Leaves glabrous. Inflorescence a shorter than calyx lobes, free, unilateral, gla-
simple, or rarely branched axillary or terminal brous, ovary bilocular, inserted midway up recep-
raceme, bracts and prophylls not enclosing flower tacle. Fruit bilocular, with 4–6 longitudinal lines
buds, often with a single apical gland; receptacle of weakness which allow seedling to escape; endo-
obliquely turbinate, slightly asymmetric, gla- carp thin, hard, bony.
brous within; petals 5, slightly shorter than Eleven spp., three of which occur in Papua
sepals; stamens 7–8 or 15; filaments slightly New Guinea and eight in New Caledonia and the
exceeding sepals, glabrous, distinct, inserted Loyalty Islands.
around complete circle or unilateral; ovary uni-
locular, inserted laterally at mouth of receptacle. 12. Kostermanthus Prance
Fruit with two large lateral plates which break
Kostermanthus Prance, Brittonia 31: 91 (1979).
away on germination and allow endocarp to
escape; endocarp thin and bony. Large trees. Leaves glabrous and papillate
Two spp., one in Mauritius and Réunion, the beneath. Inflorescence an unbranched or little-
other in Madagascar. branched terminal or axillary raceme bearing
congested cymules proximally and solitary flow-
ers distally; bracts and prophylls eglandular, not
10. Hirtella L.
enclosing flower buds; receptacle broadly obco-
Hirtella L., Sp. pl.: 34 (1753); Prance, Fl. Neotrop. 9: 259 nic, asymmetric, hairy within, sepals 5, markedly
(1972).
unequal; petals 5, exceeding calyx lobes, mark-
Trees or shrubs. Leaves glabrous or with strigose edly unequal, 2 posterior ones larger, unguicu-
hairs or hirsute, occasionally with 2 large myrme- late, enclosing stamens in bud; stamens 30–75, far
26 G.T. Prance
exceeding calyx lobes, filaments united into a within; sepals 5, unequal; petals 5, exceeding
ligule for ¾ length, unilateral with opposite stami- sepals; stamens 7–9 scarcely exceeding sepals,
nodes; ovary unilocular, inserted at mouth of recep- united towards base, unilateral with staminodes
tacle. Fruit with no lines or plates of dehiscence; opposite; ovary usually unilocular, bilocular in 1
endocarp thick, hard. Cotyledons slightly ruminate. sp., inserted at mouth of receptacle. Fruit fleshy,
Three spp. in Malesia from the Malay Penin- with no plates or lines of dehiscence; endocarp
sula to Sulawesi. usually thin and fibrous, rarely thick and woody.
Germination cryptocotylar, cataphylls absent,
eophylls opposite or in fours.
13. Licania Aubl. Fig. 4
Twelve spp. in Africa and Madagascar.
Licania Aubl., Hist. Pl. Guiane 1: 119, t. 45 (1775); Prance,
Fl. Neotrop. 9: 21 (1972).
15. Maranthes Blume
Large to small trees, shrubs or rarely suffruticose.
Leaves glabrous, lanate, pulverulent or with sto- Maranthes Blume, Bijdr. Fl. Nederl. Ind.: 89 (1825);
Prance & White, Philos. Trans. Roy. Soc. London B, 320:
matal crypts on lower surface. Inflorescence most 121–129 (1988), rev.
frequently a panicle of racemes, less often a sim-
ple raceme, a spike, a glomerule or a branched Trees. Leaves with a pair of glands at the junction
panicle of shortly stalked cymules; bracts and of the lamina and petiole, glabrous or arachnoid
prophylls not enclosing flower buds except in pubescent on lower surface. Inflorescence usually
L. licaniiflora, eglandular; receptacle variable in a many-flowered corymbose panicle, rarely a lax
shape, usually campanulate, cupuliform or urce- few-flowered thryse or a raceme of few-flowered
olate, rarely turbinate or patelliform, always monochasial cymes; bracts and prophylls not
pubescent within; petals either 5, equalling sepals, enclosing flower buds, eglandular; receptacle
or absent; stamens 3–40, filaments included to far obconical, slightly to strongly curved, nearly
exceeding sepals, usually glabrous, free in most always solid and almost completely filled by nec-
species, rarely united for half length in groups, tariferous tissue, glabrous or glabrous on one side
usually glabrous, rarely hairy; ovary unilocular, and hairy on the other within; petals 5, more or
inserted at or near base of receptacle. Fruit with less equalling sepals; stamens 20–60; filaments
no plates or lines of dehiscence, endocarp thick, free, glabrous, inserted around complete circle,
hard and woody or thin and fibrous. Germination far exceeding sepals in length, usually in a tangled
cryptocotylar, eophylls alternate. 2n ¼ 22 in two mass; ovary bilocular, inserted at mouth of recep-
species counted. tacle. Fruit with two lateral plates which break
218 spp., all but 4 of which are American away on germination to allow seedlings to escape;
largely in lowland tropical South America but endocarp very hard, thick, with a rough fibrous
extend north to Mexico, Florida and (L. mixauxii) exterior. Germination phanerocotylar, first
the Gulf states of the U.S.A., and two in the 2 eophylls opposite, the others opposite or alter-
Malesian region. nate. 2n ¼ 20.
Twelve spp., 10 of which occur in tropical
Africa, one each in tropical America (Nicaragua
14. Magnistipula Engl. to Panama) and Malesia.
Magnistipula Engl., Bot. Jahrb. 36: 226 (1905); Prance &
White, Philos. Trans. Roy. Soc. London B, 320: 152–162
(1988). 16. Parastemon A.DC.
Parastemon A.DC., Ann. Sci. Nat. Bot. II, 18: 208 (1842).
Trees, shrubs or suffrutices. Leaves glabrous or
with few strigose hairs beneath. Inflorescence a Trees or shrubs. Leaves glabrous, with two small
simple or branched raceme of cymules or rarely a discoid glands at base of lamina. Inflorescence
raceme or sessile glomerule; bracts and prophylls and axillary or rarely terminal simple or little-
not enclosing flower buds, eglandular or with 1 or branched raceme; bracts and prophylls not
2 pairs of sessile glands or with shortly stipitate enclosing flower buds, eglandular; receptacle
glands; receptacle usually curved and gibbous at patelliform or shallowly cupuliform, pubescent
base, rarely turbinate or campanulate, hairy within; petals 5, more or less equalling sepals;
Chrysobalanaceae 27
stamens 5 around circle or 2 with 3 staminodes reflexed hairs at throat, glabrous towards base
opposite; filaments glabrous, free, much shorter within; stamens 12–17; filaments slightly exceed-
than sepals; ovary unilocular, inserted at base of ing sepals, unilateral with ca. 6 toothed stami-
receptacle. Fruit with 2 large lateral plates which nodes opposite; ovary bilocular, inserted at
break away on germination to allow seedlings to mouth of staminal tube. Fruit with 2 basal obtura-
escape; endocarp thin, hard, bony. Germination tors which allow seedling to escape; endocarp
cryptocotylar, first eophylls opposite. thick, hard, with a rough fibrous surface. Germi-
Three spp. from the Nicobar Islands and the nation cryptocotylar.
Malay Peninsula to New Guinea. A single species, N. macrophylla (Sabine)
Prance, in W Africa from Senegal to Liberia.
17. Parinari Aubl. Fig. 5
Parinari Aubl., Hist. Pl. Guiane 1: 514, t. 204–206 (1775);
Prance, Fl. Neotrop. 9: 178 (1972); Prance & White, Philos. Selected Bibliography
Trans. Roy. Soc. London B, 320: 110–112 (1988), Afr. spp.
APG (The Angiosperm Phylogeny Group) 1998. See Gen-
Small to large trees or more rarely shrubs or eral References.
suffruticose. Leaves with two discoid glands on Barth, O.M., Silva, A.F. 1963. Catálogo sistemático dos
upper surface of petiole and often with small polens das plantas arbóreas do Brasil meridional.
marginal glands along entire length of lamina, IV. Cunoniaceae, Rosaceae e Connaraceae. Mem.
Inst. Oswaldo Cruz 61: 411–427.
usually with stomatal crypts filled with densely Chase, M.W. et al. 1993. See General References.
matted hairs on lower surface or rarely in two Coradin, L., Giannasi, D.E., Prance, G.T. 1985. Chemosys-
Malesian species glabrous. Inflorescence a many- tematic studies of the Chrysobalanaceae. I. Flavo-
flowered complex cyme or cymose panicle; bracts noids in Parinari. Brittonia 37: 169–178.
Dahlgren, R., Thorne, R.F. 1984. The order Myrtales:
and prophylls enclosing small groups of flower circumscription, variation and relationships. Ann.
buds, eglandular; receptacle subcampanulate, Missouri Bot. Gard. 71: 633–699.
slightly swollen to one side, pubescent within; Davis, C.C., Chase, M.W. 2004. See General References.
Demchenko, N.I. 1973. The pollen morphology of the
petals 5, equalling or shorter than sepals; stamens family Chrysobalanaceae. In: Pollen and spore mor-
6–10; filaments shorter than sepals, free, gla- phology of recent plants (Proc. 3rd Int. Palynol.
brous, unilateral with ca. 6 minute subulate sta- Conf. Acad. Sci. USSR.), pp. 60–73. (In Russian).
minodes opposite; ovary bilocular, inserted at Gibbs, R.D. 1974. Chemotaxonomy of flowering plants.
Vols 1–4. Montreal, London: McGill-Queens Univer-
mouth of receptacle. Fruit with 2 basal obturators sity Press.
which allow seedling to escape; endocarp thick, Hallier, H. 1921. See General References.
hard, with a rough fibrous surface. Lack, A. 1978. The ecology of the flowers of the savanna
Thirty-nine species, pantropical with 19 in tree Maranthes polyandra and their visitors, with
particular reference to bats. J. Ecol. 66: 287–295.
tropical America, 6 in tropical Africa and 15 in Mangenot, S., Mangenot, G. 1962. Enquête sur les nom-
tropical Asia and the Pacific region extending to bres chromosomiques dans une collection d’espèces
Fiji, Tonga and Samoa. tropicales. Rev. Cytol. Biol. Vég. 25: 411–447.
Matthews, M.L., Endress, P.K. 2008. See General References.
Patel, V.C., Skvarla, J.J., Raven, P.H. 1983. Pollen ultra-
18. Neocarya Prance structure of Chrysobalanaceae. Vidya 26: 1–10.
Prance, G.T. 1972. Flora Neotropica, monograph no. 9
Neocarya Prance in F. White, Bull. Jard. Bot. Nat. Belg. 46: (Chrysobalanaceae). 409 pp. New York: Hafner.
308 (1976). Prance, G.T. 1979. Chrysobalanaceae. Flora of Ecuador
(eds. G. Harling & B. Sparre), vol. 10, pp. 1–23.
Shrubs or small trees. Leaves with several small, Prance, G.T. 1979. New genera and species of Chrysoba-
sessile glands near base of lamina, with stomatal lanaceae from Malesia and Oceania. Brittonia 31:
crypts filled with densely matted hairs on lower 79–95.
Prance, G.T. 1982. Chrysobalanaceae. Flora de Venezuela,
surface. Inflorescence a terminal raceme of vol. 4(2), pp. 325–482. Carácas: Ediciones Fundación
almost sessile cymules, sometimes unbranched Educación Ambiental.
but more frequently with one or more short to Prance, G.T. 1983. Chrysobalanacées. Flore da la Nouvelle-
elongate branches at base; bracts and prophylls Calédonie et dépendances, vol. 12, pp. 105–123. Paris:
Muséum national d’Histoire naturelle
enclosing one or more flower buds, eglandular; Prance, G.T.1988. Flora Neotropica, monograph no. 9 (sup-
receptacle asymmetric, saccate, gibbous, with plement). New York: The New York Botanical Garden.
28 G.T. Prance
Prance, G.T. 1989. Chrysobalanaceae. Flora Malesiana Ser. Tobe, H., Raven, P.H. 1984. An embryological contribu-
1.10: 635–678. tion to systematics of the Chrysobalanaceae. I. Tribe
Prance, G.T., Sothers, C.A. 2003. Chrysobalanaceae 1: Chrysobalaneae. Bot. Mag. Tokyo 97: 397–411.
Chrysobalanus to Parinari. Species Plantarum: Vogel, S. 1968–1969. Chiropterophilie in der neotro-
Flora of the World Part 9: 1–319. pischen Flora. Flora (Abt. B) 157: 562–602 (1968);
Prance, G.T., Sothers, C.A. 2003. Chrysobalanaceae 2: 158: 289–323 (1969).
Acioa to Magnistipula. Species Plantarum: Flora of White, F. 1976. The taxonomy, ecology and chorology of
the World Part 10: 1–268. African Chrysobalanaceae (excluding Acioa). Bull.
Prance, G.T., White, F. 1988. The genera of Chrysobala- Jard. Bot. Nat. Belg. 46: 265–350.
naceae: a study in practical and theoretical taxonomy White, F. 1976. Chrysobalanaceae. Distributiones plan-
and its relevance to evolutionary biology. Philos. tarum africanarum, vol. 10, maps 281–334.
Trans. Roy. Soc. London B, 320: 1–148. Wurdack, K.J., Davis, C.C. 2009. See General References.
Souza, S. de 1979. Contribution à l’étude biologique et Yakandawala, D., Morton, C., Prance, G.T. 2010. Phyloge-
écologique de quelques Chrysobalanacées des genres netic relationships of the Chrysobalanaceae inferred
Chrysobalanus, Parinari et Maranthes au Bénin. from chloroplast, nuclear and morphological data.
Thesis, Université de Bordeaux III. Ann. Missouri Bot. Gard. 97: 259–281.
Ctenolophonaceae
Ctenolophonaceae Exell & Mendonça (1951).
K. K U B I T Z K I
Buttressed trees with simple and stellately tufted hairs cells (crystalliferous cells with a unilaterally thick-
on young shoots, stipules, and the outside of sepals ened and lignified cell wall), and crystals and
and petals. Leaves opposite, petiolate, entire, simple, druses are of a more common occurrence (van
pinnately veined; stipules interpetiolar, caducous. Welzen and Baas 1984).
Inflorescences thyrso-paniculate, axillary or terminal; In the secondary xylem, the vessels are exclu-
bracts present, prophylls 0. Flower buds elongate- sively solitary; perforation plates are predominantly
ellipsoid; flowers hypogynous, bisexual, 5-merous, scalariform with numerous thick bars; axial paren-
diplostemonous, regular, hypogynous; sepals basally chyma is paratracheal-abaxial; wood rays are uniseri-
shortly connate, quincuncially imbricate, subequal, ate, of upright cells, and multiseriate heterogeneous,
indurate, swollen and persistent in fruit; petals con-
torted, caducous, spoon-shaped at the base and often
shortly clawed; disk well developed, extrastaminal,
cup-like; stamens 10, inserted on inner side of the
disk and basally connate into an androecial tube,
dorsally surrounded by a 10-lobed corona-like tube;
antepetalous stamens shorter than antesepalous
ones; anthers dorso-versatile, 2-celled, introrse; con-
nective protruding, acute-triangular; gynoecium syn-
carpous, 2-carpellate; ovary 2-celled; ovules 2 per cell,
axile, collateral, pendant, anatropous, epitropous,
bitegmic; style simple, apically cleft with 2 capitate
stigmas; ovary tube joining near its base the androe-
cial tube to form a short androgynophore. Fruit a
1-celled capsule, the woody pericarp eventually
splitting lengthwise into 2 valves. Seed solitary, per-
sisting after falling of the pericarp and pendulous
from the top of a filiform columella; aril pectinate-
fimbriate, surrounding the lower half of the seed;
endosperm copious; embryo straight.
Monogeneric, probably with 2 extant spp.,
C. englerianus Mildbr. in West Africa, C. parvifolius
Oliv. from Malay Peninsula to New Guinea.
V E G E T A T I V E A N A T O M Y . Ctenolophon differs
from its presumed allies in possessing tufted
hairs, stellate in appearance, which are found on
young vegetative and mature floral parts, and
Fig. 6. Ctenolophonaceae. Ctenolophon parviflorus. A
anomocytic to anisocytic stomata. The simple Flowering branchlet. B Flower. C Same, longitudinal sec-
arc of collateral vascular tissue in the midrib and tion. D Stamens. E Fruiting twig. F Fruit. G Same, longi-
petiole, the scattered occurrence of cristarque tudinal section. H Seed. (Pungga 1996; drawn by J. Pao)
30 K. Kubitzki
up to 3 cells wide; fibre tracheids have conspicuously parvifolius or “C. philippinensis” has been found
bordered pits; solitary crystals occur in chambered to exhibit a sculpture similar to that of the engler-
parenchyma and ray cells (Heimsch and Tschabold ianus type (Thanikaimoni et al. 1984; van der Ham
1972). The heartwood of C. parvifolius is reddish to 1989).
brown, very hard, and heavier than water. Saad (1962) pointed to pollen morphological
similarities between Ctenolophon and the Malpighia-
F L O W E R S T R U C T U R E . The flowers of Ctenolo- ceae which, however, appeared superficial to van der
phon are tubular, as the petals bend outwards only Ham (in van Hooren and Nooteboom 1988).
apically. In bud (and later?) the overlapping por-
tions of the petals are postgenitally connected by the P O L L I N A T I O N . The bright petals and well-
indumentum of their dorsal surfaces and margins. developed nectary disk indicate that Ctenolophon
In C. parvifolius the petals are white to yellow, is entomophilous.
orange or bright red (van Hooren and Nooteboom
1988). The stamens are basally connate into an F R U I T A N D S E E D . The fruit of C. parvifolius is
androecial tube and are surrounded by a tubular yellow to pink and later brown, the seed brown or
disk, which distally extends into ten lobes that alter- glossy black, and the aril white to orange or brilliant
nate with the stamens (the corona of Matthews and vermilion. Seeds are exotegmic; the embryo is
Endress 2011). The disk surrounds the ovary up to folded (Boesewinkel and Bouman 2000).
three quarters of its height. As the petals bend out-
wards only apically, the floral tube can retain large A F F I N I T I E S . Ctenolophon had been referred to
amounts of nectar. The tissue of the disk is formed various families including Oleaceae, Icacinaceae
by thin-walled, typical adenoid cells. Close to the and Celastraceae, until Pierre (1893) suggested a
margin of both sides of the disk, numerous open relationship with Linaceae. Winkler (1931)
anomocytic stomata are diffusely distributed, which included in it Ctenolophon as a subfamily,
may function in the release of the nectar (Link 1992). together with Ixonanthoideae and Humirioideae.
Due to partial disintegration of the thin septa, the These subfamilies have since been elevated to the
ovary becomes partly unilocular. The ovules have a rank of family; among them, Ctenolophon has
distinct endothelium (Matthews and Endress 2011). been considered close to Linaceae s.str., with
which it shares the extrastaminal disk. Link
P O L L E N M O R P H O L O G Y . Pollen morphology of (1992) stressed the similarities between Humiria-
Ctenolophon has been studied by, among others, ceae and Ctenolophon, particularly the perianth
Thanikaimoni et al. (1984) and van der Ham structure, the broadened and markedly elongate
(1989). The pollen is typically zonocolporate, the anther connectives, and the histology of the disk,
number of apertures ranging from 3–9. Two main including the presence of the marginally
pollen types can be distinguished, which by and distributed nectarial stomata. In this context it
large correspond to the two extant species. The appeared less important to him whether the disk
parvifolius type has 3–8 apertures and is sub- is intrastaminal (Humiriaceae) or extrastaminal
spheroidal, whereas the englerianus type has 6–9 (Ctenolophon), or the filament bases are adnate to
apertures and exhibits distinct ridges on the the disk (Ctenolophon) or distinct (Humiriaceae).
mesocolpia that extend to the polar area, where Matthews and Endress (2011) identified impor-
they fuse into a ring and make the pollen grains tant structural traits uniting Ctenolophon, Rhizo-
characteristically angular and barrel-shaped. phoraceae and Erythroxylaceae. These groups
Equatorial size ranges from 31 to 66 mm; the have also been retrieved as a clade [Ctenolopho-
thickness of the exine ranges from 3–6 mm, and naceae [Rhizophoraceae + Erythroxylaceae]] by
its stratification is obscure under the light micro- Wurdack and Davis (2009) and Xi et al. (2012).
scope and difficult to determine. The infratectal
layer appears labyrinthic and sometimes exhibits D I S T R I B U T I O N A N D H A B I T A T S . Ctenolophon
granular-columellate structures. The tectum is englerianus is known from West Africa, where it
perforate to foveolate in the parvifolius type, and grows in riverine forests. C. parvifolius is
psilate or finely perforate in the englerianus type. distributed in Malesia from the Malay Peninsula
Ctenolophon from the Philippines classified as C. to New Guinea, and grows in primary forest but
Ctenolophonaceae 31
also swamp and heath forest on a wide variety of but not reported from Java, the Lesser Sunda
soils over peat, sandstone and ultrabasic rocks, Islands, Celebes and the Moluccas. The two species
commonly in lowland and hill regions and in are very similar, the only differences being the
Borneo up to 1,650 m. simple cymose panicle and larger flowers (stamens
up to 15 mm long) in C. englerianus vs. compound
P A L A E O B O T A N Y . The fossil record of Ctenolo- cymose panicles and smaller flowers (stamens up
phon is extensive and starts in the Maastrichtian to 10 mm long) in C. parvifolius, apart from the
of Central Africa with pollen of the englerianus differences in pollen morphology.
type, which through a nearly uninterrupted
record can be linked with the Recent African
distribution of C. englerianus. This pollen type
is recorded also from the Palaeocene and Eocene Selected Bibliography
of northern South America and the Eocene and
Miocene of India but disappeared from these Boesewinkel, F.D., Bouman, F. 2000. Ctenolophonaceae. In:
Takhtajan, A. (ed.) Anatomia seminum comparativa,
regions in the course of the Tertiary (Krutzsch vol. 6, pp. 13–14. St. Petersburg: Nauka. (In Russian)
1989). The parvifolius type first appeared in the Erdtman, G. 1955. Pollen grains of cf. Ctenolophon from
Palaeocene of Africa, but continued there only to Tertiary deposits in India. Bot. Not. 108: 143–145.
the early Eocene. In India (Kutch) it appeared in Heimsch, C., Tschabold, E.E. 1972. Xylem studies in the
Linaceae. Bot. Gaz. 133: 242–253.
the early Palaeocene, from where it spread in the Kar, R.K. 1985. The fossil flora of Kutch. IV. Tertiary
course of the Tertiary all over India and, in the palynostratigraphy. The Palaeobotanist 34: 1–279.
Miocene with up to 9 pollen types, attained its Krutzsch, W. 1989. Paleogeography and historical phyto-
greatest diversity (Kar 1985). Erdtman (1955) geography (paleochorology) in the Neophyticum. Pl.
Syst. Evol. 162: 5–61.
first described and characterised the fossil pollen Link, D.A. 1992. Floral nectaries of the Geraniales and
from this region. In the course of the Eocene, the their systematic implication. IV. Ctenolophonaceae
parvifolius type became established in Southeast Badre. Flora 187: 103–107.
Matthews, M.L., Endress, P.K. 2011. See General References.
Asia/Malesia, again being linked with the distri- Pierre, J.B.L. 1893. Flore forestière de la Cochinchine, IV.
butional area of its extant name-giving species. Fasc. 18, t. 281. Paris: Doin.
Thanikaimoni et al. (1984) and van der Ham Pungga, R.S. 1996. Ctenolophonaceae. In: Soepadmo, E.,
(1989) have discovered that populations of Wong, K.M., Saw, L.G. Tree flora of Sabah and Sara-
wak, vol. 2: 151–153. Kuala Lumpur: Forest Res. Inst.
C. parvifolius from the Philippines exhibit pollen Malaysia.
that resembles the pollen of the englerianus type, Saad, S.I. 1962. Pollen morphology of Ctenolophon. Bot.
giving rise to the suggestion (W. Krutzsch, pers. Notis. 115: 49–57.
comm., Nov. 2003) that the ridges of the engler- Thanikaimoni, G., Caratini, C., Venkatachala, B.S., Rama-
nujam, C.G.K., Kar, R.K. (eds.) 1984. Selected Tertiary
ianus type pollen are plesiomorphic and have angiosperm pollens from India and their relationship
been lost on the pollen grains of C. parvifolius, with African Tertiary pollens. Pondichéry: Institut
which is supported by rudimentary ridges that Français de Pondichéry.
have been observed by W. Krutzsch. van Hooren, A.M.N., Nooteboom, H.P. 1984. Linaceae and
Ctenolophonaceae especially of Malesia, with notes
on their demarcation and the relationships with Ixo-
Only one genus: nanthaceae. Blumea 29: 547–563.
van Hooren, A.M.N., Nooteboom, H.P. 1988. Ctenolopho-
Ctenolophon Oliver Fig. 6 naceae, in: Flora Malesiana I, 10: 629–634.
van der Ham, R.W.J.M. 1989. New observations on the
Ctenolophon Oliver, Trans. Linn. Soc. 28: 516 (1873); pollen of Ctenolophon Oliver (Ctenolophonaceae),
Bullock, Kew Bull. 14: 41 (1960); Badré, Fl. Gabon 21: with remarks on the evolutionary history of the
genus. Rev. Palaeobot. Palynol. 59: 153–160.
43–44 (1973); van Hooren & Nooteboom in Fl. Males. I,
van Welzen, P.C., Baas, P. 1984. A leaf anatomical contri-
10: 629–634 (1988); R.S. Pungga, Tree flora of Sabah and
bution to the classification of the Linaceae complex.
Sarawak 2: 151–153 (1996). Blumea 29: 453–479.
Winkler, H. 1931. Linaceae. In: Engler & Prantl, Nat.
Two spp., C. englerianus Mildbr. from Angola, Pflanzenfam., ed. 2, 19a. Leipzig: W. Engelmann.
Zaire, Nigeria and Gabon, and C. parvifolius Wurdack, K.J., Davis, C.C. 2009. See General References.
Oliver from the Malay Penisula to New Guinea, Xi, Z. et al. 2012. See General References.
Dichapetalaceae
Dichapetalaceae Baill. (1886), nom. cons.
Chailletiaceae R. Br. (1818).
G.T. P R A N C E
Trees, shrubs or lianas. Leaves alternate, simple, Three genera, 170 species, tropical Asia,
entire, pinnately nerved; stipules often early Africa and America extending to subtropics in
caducous, entire, lobed, partite or fimbriate. South Africa and India.
Inflorescences axillary, sometimes on leafless
axillary or terminal shoots, cymose, distinctly V E G E T A T I V E M O R P H O L O G Y . Most species of
branched to subcapitate or fasciculate, the ped- Dichapetalum are lianas, a few are small trees or
uncle free or adnate to petiole or more rarely shrubs. Some species may be either shrubs or lianas
the midrib; bracts and prophylls usually small. or even small trees. Species of Tapura and Stephano-
Flowers small, actinomorphic or zygomorphic, podium are generally small to medium-sized trees or
hermaphrodite or unisexual; pedicels usually shrubs but T. africana can be a tree of 20 m and 100
articulated; sepals 5(4), imbricate, equal to very cm d.b.h. Two African species of Dichapetalum are
unequal, free or slightly united or rarely forming suffruticose. Tree and shrub forms of variable species
a tube; petals 5(4), either free and almost equal tend to occur in shady habitats and lianas in more
or connate into a tube with the lobes equal to open, well-lit spaces. Most liana species begin as
very unequal, the lobes usually bifid at apex and shrubs and then become lianescent. The lianas are
most frequently bicucullate or inflexed, often not generally twining and do not have tendrils, but
clawed at base; stamens (4) 5, up to 3 of them climb by the production of short, hook-like plagio-
sometimes lacking anthers, antesepalous, dis- tropic branches. Two models of architecture have
tinct to base of receptacle or adnate to corolla been reported, Roux’s Model in two African species
tube, generally with filaments, rarely with of Dichapetalum and Cook’s Model in Tapura guia-
anthers sessile; anthers bilocular, introrse, lon- nensis. In the latter species with the inflorescences
gitudinally dehiscent; 1–5 variously shaped inserted at the distal end of the petiole, a branch-like
equal or unequal hypogynous glands ("stami- character occurs on the leaf which is itself part of a
nodes") or disk lobes alternating with stamens, leaf-like branch. The stipules in many species are
distinct or connate into a disk; ovary superior, small and early caducous, in Dichapetalum they
2–4(5)-locular, with 2 ovules in each loculus, may be fimbriate, pinnatisect or palmately divided
ovules epitropous, anatropous, bitegmic, tenui- into filiform segments. The lower leaf surface is often
nucellate, pendulous from top of each loculus, glandular, but the glands are not arranged in any
raphe ventral; style usually simple with 2–4(5) regular pattern, and less frequently glands also
lobes, or more rarely 2–4(5) distinct stylodia; occur on the upper surface. In young leaves the
stigma punctate. Fruit a dry or fleshy drupe, glands are nectariferous.
1–3(4)-seeded; exocarp most frequently
appressed pubescent, sometimes dehiscent; V E G E T A T I V E A N A T O M Y . As is common in many
mesocarp thin to thick; endocarp hard or parch- lianas, the wood is often deeply divided by intruding
ment-like, indehiscent, glabrous or pubescent phloem which runs from the exterior in bands
within. Seeds pendulous, with little or no endo- towards the centre. The stem is often clearly five
sperm; embryo large, erect, with plano-convex lobed because of the intruding phloem, but in
cotyledons. Germination hypogeal, first leaves some species there are many more strands. The
opposite or alternate. wood has vessels solitary or in small groups, with
34 G.T. Prance
simple or scalariform perforation. Parenchyma is show no trace of the floral bundle, but in a few cases
predominantly paratracheal around the vessels, floral bundles have been noted in sterile petioles.
with some short irregular wings. Rays are typically The petiolar inflorescence appears to have arisen
5–6 cells wide, heterogeneous, sometimes uniseriate. by a gradual fusion of the peduncle and the petiole,
The leaves are dorsiventral, glabrous or with rather than by true epiphylly.
unicellular hairs. The hairs may be long and arach- The sieve tubes contain non-dispersive protein
noid or short, stiff and bristle-like and often barbed bodies.
or flexible. The leaf indumentum is quite variable.
The petiole of species with an axillary inflorescence I N F L O R E S C E N C E A N D F L O W E R S T R U C T U R E . The
has the characteristic horseshoe-shaped bundles flowers are rather small, seldom exceeding 5 mm in
which are less compact towards the base and are size. Flowers are generally bisexual but unisexual
often flanked by small lateral bundles. In species flowers occur in some species of Stephanopodium
with an epiphyllous inflorescence, the inflore- and some Asiatic Dichapetalum. In Africa some
scence usually emerges above the midpoint, but species with apparently bisexual flowers are func-
the floral and leaf bundles are separate from near tionally unisexual. The inflorescence is always axil-
to the base of the petiole. The sterile petioles usually lary and cymose; it is a fascicle arising from the
petiole in Stephanopodium and most species of
Tapura. It is branched and either inserted in the
axil or on the petiole in most species of Dichapeta-
lum. The petals are distinct almost to their base in
Dichapetalum, but connate into a tube in Tapura
and Stephanopodium. In the latter genus the anthers
are sessile on the tube. The petal lobes are usually
cucullate, except in Stephanopodium. Breteler (1973)
has suggested that the petals are of staminal origin,
and that the lobes are homologous with anthers and
the lower united part homologous with filaments.
This seems quite possible, and the bicucullate lobes
strengthens this theory. For convenience in descrip-
tions, however, all authors have termed the tube and
lobes petals. At the base of the petals of most
species, there are five hypogynous epipetalous struc-
tures variously termed glands, disk or staminodes.
These staminodes are often bilobed at their apex,
one of the reasons for considering them as stami-
nodes rather than disks. It is interesting that within
the family a stepwise progression from distinct sty-
lodia through a style with distal style branches to an
unbranched style is documented. See also the
detailed studies on floral structure of the family
and its relatives by Matthews and Endress (2008).
P O L L E N M O R P H O L O G Y . A eurypalynous family
in which Punt (1975) distinguished 29 pollen
types. The pollen is 3-colporate or in a few species
Fig. 7. Dichapetalaceae. A–D Dichapetalum spruceanum. inaperturate, and a small number of 4-colporate
A Leaf and inflorescence. B Petal. C Flower, vertical section. grains occur in some specimens. The grains are
D Flower. E–H Stephanopodium aptotum. E Leaf and inflore- spheroidal, suboblate or oblate. The longest axis
scence. F Flower. G Flower, vertical section. H Corolla and
stamens. I–L Tapura amazonica. I Flower. J Leaf and inflore- is usually under 20 mm, rarely up to 35 mm. The
scence. K Flower, vertical section. L Corolla and stamens. ornamentation is quite varied from reticulate or
(Orig.) microreticulate to pertectate.
Dichapetalaceae 35
K A R Y O L O G Y . A study of 16 African species of brilliantly coloured pulp which may attract birds
Dichapetalum (Gadella 1969, 1970, 1972) found all (Breteler 1973, 1981). Some neotropical species
species to have the same basic number 2n ¼ 24. are eaten by bats.
There is one report of 2n ¼ 20 in a species where
Gadella found 2n ¼ 24. Arends and van der Laan P H Y T O C H E M I S T R Y . Some African species of
(1986) gave a summary of all known chromosome Dichapetalum are extremely poisonous due to
numbers in Dichapetalaceae, and reported an octo- the presence of fluoroacetic acid (FCH2COOH)
ploid of 2n ¼ 96 in D. crassifolium var. crassifolium and fluoro fatty acids. The leaves of D. toxicarium
and a tetraploid of 2n ¼ 48 in Tapura africana. All contain monofluoroacetate and fluoride ions
50 other counts in 21 species were 2n ¼ 24. The (Ward et al. 1964; Vickery and Vickery 1972).
chromosomes are 1–2 mm in length and have pri- Species with fluoride compounds are extremely
mary constrictions in the median region. poisonous to cattle and are a problem in some
African pastures. The fruits of D. rudatsii and
P O L L I N A T I O N . Very little is known about polli- D. toxicarium are used to kill mice and rats.
nation. The flowers of Tapura guianensis are sweet
scented and visited by small bees (Trigona and A F F I N I T I E S . Many different relationships have
Melipona). A sweet scent has also been noticed in been suggested for the family including Geraniales,
several African species of Dichapetalum. Rosales, Thymelaeales, Celastrales and Euphor-
biales. Although often placed near to the Euphor-
F R U I T A N D S E E D . The fruit is a 1–3(4)-seeded biaceae, it does not seem to fit well there. A
fleshy or dry drupe sometimes with pyrenes free combined morphological/molecular analysis
from each other, but more often one-seeded. The (Nandi et al. 1998) placed Dichapetalaceae in close
pyrenes are always 1-seeded, since only one ovule proximity with Trigoniaceae and Chrysobalanaceae,
from each loculus develops; they are always free and several subsequent molecular studies provided
from each other within the fruit pulp. In most very strong support for a subclade of the Malpigh-
species the exocarp is pubescent with a short iales in which Dichapetalaceae + Trigoniaceae are in
appressed velutinous pubescence, but in some a sister position and together are sister to Chryso-
African Dichapetalum long barbed, and irritating balanaceae + Euphroniaceae (Davis and Chase 2004;
Mucuna type hairs occur. The endocarp is quite Wurdack and Davis 2009; Xi et al. 2012). Matthews
distinct from the exocarp, which may be either and Endress (2008) compared the floral structure of
leathery or bony. An aril has been observed in these families and provided support for the close
two species of Dichapetalum but the seeds are relationship of Dichapetalaceae and Trigoniaceae.
normally without arils. The mature seed coat
consists only of a tanniniferous exotesta with D I S T R I B U T I O N A N D H A B I T A T S . In the Neotropics
thickened cell walls, while the mesophyll and Dichapetalum and Tapura range from Mexico to C
inner epidermis of the testa and the tegmen dur- Brazil and Stephanopodium from Costa Rica to
ing seed development are compressed and largely northern and western South America and Atlantic
resorbed (Boesewinkel and Bouman 1980). The coastal Brazil but not to Amazonia. In Africa the
embryo is large and surrounded by a thin endo- centre of distribution is in C Africa in Cameroon,
sperm; the cotyledons are plano-convex. Germi- Gabon, Zaire, etc. but ranging from West to East
nation of all species examined is hypogeal, the tropical Africa, and two species of Dichapetalum
cotyledons remaining within the endocarp and extend south into subtropical South Africa. There
the taproot and epicotyl emerging from a slit at are seven species of Dichapetalum in Madagascar, of
the suture of the endocarp. The first leaves are not which six are endemic (Breteler 1986). The Asiatic
markedly different from later leaves but may be species of Dichapetalum are found in SE Asia, the
either opposite or alternate. Little is known about Philippines, Malaysia, Australia, New Guinea and
seed dispersal. Several species have fruits with Melanesia, with one species in Fiji.
edible pulp and are presumably dispersed by ani- The family is found predominantly in forests.
mals. Dichapetalum integripetalum is eaten by Only two African species of Dichapetalum are
chimpanzees. Species with barbed hairs on the found in semi-arid conditions. Several neotropical
exocarp open, exposing the pyrenes covered by species of Dichapetalum and Stephanopodium are
36 G.T. Prance
found at altitudes of 2,000 m. One species of Tapura polygamo-dioecious; petals united into a long
is common in the savannas of central Brazil. obconical or cylindrical tube, with 5 equal, broadly
ovate lobes which are shorter than the tube,
E C O N O M I C I M P O R T A N C E . The family is of little lobes entire at apex or slightly bicucullate; stamens
economic importance, the only uses being recorded 4–5, anthers sessile, adnate to interior of corolla
are in a few folk medicines and as a poison. tube.
Thirteen spp. in Costa Rica, N W South America
C O N S E R V A T I O N . Many species are of very and Atlantic coastal Brazil.
restricted distribution especially in the Neotropics,
Africa and Madagascar. The most vulnerable species
3. Tapura Aubl.
must be those of the Atlantic coastal forests of Brazil,
especially in Stephanopodium, and the Madagascan Tapura Aubl., Pl. Guiane 1: 126, t. 48 (1775); for revisions,
see Bibliography.
species of Dichapetalum.
Trees or shrubs. Inflorescence usually a small sessile
KEY TO THE GENERA or shortly pedunculate glomerule adnate to petiole,
rarely, in Africa, an axillary glomerule borne on a
1. Petals unequal, 1–2 distinctly larger, usually only 2–3
long peduncle free from petiole. Flowers usually
stamens fertile, rarely 5 3. Tapura
hermaphrodite, rarely unisexual; petals connate at
– Petals equal, stamens all fertile 2
2. Petals distinctly bilobed, distinct to base or connate base to form a distinct tube or free almost to base,
into a tube, anthers usually filamentose with 1–2 large broad lobes with bicucullate divided
1. Dichapetalum apex, the other 2–4 smaller, linear-lanceolate,
– Petals not bilobed, slightly bifid in S. estellense, always entire; stamens 2–3 or 5 fertile, adnate to corolla
united into a tube; anthers sessile on tube tube or to base of corolla in species with distinct
2. Stephanopodium
petals, with distinct filaments.
Twenty neotropical spp. and 8 in Africa.
GENERA OF DICHAPETALACEAE
Acknowledgement I am grateful to F.J. Breteler for reviewing
1. Dichapetalum Thouars this contribution.
Dichapetalum Thouars, Gen. Nov. Madag.: 23 (1806);
Prance, Kew Bull. 52: 213–219 (1997), key to Neotrop.
spp.; Breteler (1969–1982), rev. Afric. spp. Selected Bibliography
Lianas, shrubs or rarely small trees. Inflore-
scence axillary, adnate to petiole or in axil, usually Arends, J.C., van der Laan, F.M. 1986. Cytology. In: Breteler F.J.
a pedunculate cyme, rarely a sessile glomerule. Flow- 1986, op. cit.
Barth, F. 1896. Anatomie comparée de la tige et de la feuille
ers usually hermaphrodite, rarely unisexual; petals des Trigoniacées et des Chailletiacées (Dichapétalées).
equal, distinct to base or shortly connate, entire or Bull. Herb. Boissier 4: 497–520.
bilobed at apex; stamens 5, equal, distinct to base or Boesewinkel, F.D., Bouman, E. 1980. Development of
connate at extreme base only, with distinct fila- ovule and seed coat of Dichapetalum mombuttense
Engl. with notes on other species. Acta Bot. Neerl. 29:
ments. 2n ¼ 24 or 96. 103–115.
About 135 spp., 90 in Africa, 7 Madagascan, Breteler, F.J. 1969. The African Dichapetalaceae 1. Acta
20 neotropical and 16 in SE Asia, Malesia and the Bot. Neerl. 18: 375.
Pacific. Breteler, F.J. 1973. The African Dichapetalaceae (III). A taxo-
nomical revision. Species a–b. Meded. Landbouwhoge-
school Wageningen 73-13.
Breteler, F.J. 1978. The African Dichapetalaceae IV. A taxo-
2. Stephanopodium Poepp. & Endl. nomical revision. Species c–f. Meded. Landbouwhoge-
school Wageningen 78-10.
Stephanopodium Poepp. & Endl., Nov. Gen. et Sp. 3: 40, t.
Breteler, F.J. 1979. The African Dichapetalaceae V. A taxo-
246 (1842); Prance, Kew Bull. 50: 295–305 (1995), rev. nomical revision. Species g–l. Meded. Landbouwhoge-
Small to medium-sized trees or shrubs. Inflore- school Wageningen 79-16.
Breteler, F.J. 1981. The African Dichapetalaceae VII. A taxo-
scence a small sessile or shortly pedunculate glome- nomical revision. Species m–q. Meded. Landbouwhoge-
rule, adnate to petiole. Flowers hermaphrodite or school Wageningen 81-10.
Dichapetalaceae 37
Breteler, F.J. 1982. The African Dichapetalaceae VIII. A taxo- Leenhouts, P.W. 1956. Some notes on the genus
nomical revision. Species r–z. Meded. Landbouwhoge- Dichapetalum in Asia, Australia and Melanesia. Rein-
school Wageningen 82-8. wardtia 4: 75–87.
Breteler, F.J. 1986. The African Dichapetalaceae IX. A taxo- Leenhouts, P.W. 1957. Dichapetalaceae. In: Fl. Malesiana
nomical revision. Agric. Univ. Wageningen Papers 86-3. II, 5: 305–316.
Breteler, F.J. 1991. Dichapetalaceae. In: Flore du Gabon Matthews, M.L., Endress, P.K. 2008. See General Refer-
32: 1–221. ences.
Davis, C.C., Chase, M.W. 2004. See General References. Nandi, O.I., Chase, M.W., Endress, P.K. 1998. A combined
Descoings, B. 1960. Révision de Dichapetalum de Madagascar. cladistic analysis of angiosperms using rbcL and
Mém. Inst. Sci. Madag. B9: 63–120. non-molecular data sets. Ann. Missouri Bot. Gard.
De Wildeman, E. 1919. Notes sur les espèces Africaines du 85: 137–212.
genre Dichapetalum Thon. Rev. Zool. Afr. 4(2), Prance, G.T. 1972. Dichapetalaceae. Flora Neotropica
Suppl. Bot.: 1–75. Monograph 10: 1–84.
Engler, A. 1896. Dichapetalaceae. In: Engler & Prantl, Nat. Prance, G.T. 1995. A synopsis of the genus Stephanopodium
Pflanzenfam. III, 4: 345–351. (Dichapetalaceae). Kew Bull. 50: 295–305.
Engler A., Krause, K. 1931. Dichapetalaceae. In: Engler & Punt, W. 1975. Pollen morphology of the Dichapetalaceae
Prantl, Nat. Pflanzenfam. 2nd ed. 19c: 1–11. with special reference to evolutionary trends and
Gadella, Th.W.J. 1969. Chromosome numbers of some mutual relationships of pollen types. Rev. Palaeobot.
angiospermae collected in Cameroun and the Ivory Palynol. 19: 1–97.
Coast. Proc. Kon. Nederl. Akad. Wet. C, 72: 306–310. Vickery, B., Vickery, M.L. 1972. Fluoride metabolism in
Gadella, Th.W.J. 1970. Chromosome numbers of some Dichapetalum toxicarium. Phytochemistry 11:
Angiospermae collected in Cameroun and the Ivory 1905–1909.
Coast II. Acta Bot. Neerl. 19: 431–435. Vickery, B., Vickery, M.L., Ashu J.T. 1973. Analysis of
Gadella, Th.W.J. 1972. Cytological notes on some flowering plants for fluoracetic acids. Phytochemistry 12:
plants collected in Africa. Bull. Jard. Bot. Nat. Belg. 42: 145–147.
393–402. Ward, P.F.N., Hall, R.J., Peters, R.A. 1964. Fluoro fatty
Hauman, L. 1958. Dichapetalaceae. In: Flore du Congo acids in the seeds of Dichapetalum toxicarium.
Belge et du Ruanda-Urundi 7: 287–348. Nature 201: 611–612.
Keay, R.W.J. 1958. Dichapetalaceae. In: Flora of West Wurdack, K.J., Davis, C.C. 2009. See General References.
Tropical Africa ed. 2. 1(2): 433–439. Xi, Z. et al. 2012. See General References.
Elatinaceae
Elatinaceae Dumort., Anal. Fam. Pl. 44, 49 (1829), ‘Elatinideae’, nom. cons.
K. K U B I T Z K I
Perennial or annual herbs of aquatic or moist anomocytic (Tucker 1986). The wood of Bergia
terrestrial habitats, rarely (Bergia suffruticosa) suffruticosa has been studied by Carlquist (1984).
suffrutescent; nodes unilacunar 1-trace (Elatine); He observed vessel elements with simple perfora-
indumentum of unicellular and multicellular mul- tion plates, vasicentric tracheids and fibriform
tiseriate capitate trichomes, or plants glabrous; a vessel elements, predominance of uniseriate
resinous secretion widely distributed in the tis- rays, vertical orientation of scalariform vessel-
sues and deposited in granular form on the sur- ray pitting, absence of intraxylary phloem, pres-
face of the stems. Leaves opposite and decussate, ence of brownish deposits in the parenchyma, and
rarely (Elatine alsinastrum) verticillate, entire or occurrence of druses and solitary crystals. Sto-
coarsely serrate; stipules minute, scarious, distinct mata have 4–8 irregularly shaped subsidiary
or interpetiolar. Inflorescences axillary, dichasial, cells. In contrast to most other families of Mal-
or flowers solitary. Flowers hermaphroditic, pighiales, the sieve tube plastids of Elatinaceae
small, actinomorphic, hypogynous, usually incon- lack starch inclusions (Behnke 1991).
spicuous, occasionally cleistogamous; sepals 2–5
(6), distinct or connate basally; petals 2–5, dis- E M B R Y O L O G Y . The tapetum is secretory and
tinct, membranaceous, imbricate, persistent; micosporogenesis simultaneous. Pollen grains
stamens (2)3–6(–10), in 1 or 2 whorls, the outer are 2-celled (Bergia) or 3-celled (Elatine) when
whorl antesepalous; anthers broadly ovoid, dorsi- shed. The ovules are pendulous apical and epi-
fixed, introrse, dehiscing by longitudinal slits; tropous, bitegmic and crassinucellate, and the
ovary superior, (depressed-)ovoid, (2)3–5-locular, micropyle is formed by both integuments. The
the partitions not reaching the top of the ovary in embryo sac is of the Polygonum type; endosperm
some Bergia, stylodia 3–5; stigmas capitate, papil- development is Nuclear. Embryogeny is of the
late; placentation axile or basal; ovules numerous, Solanad type (Kajale 1939; Raghavan and Srini-
anatropous, bitegmic, weakly crassinucellar. Fruit vasan 1940; Dathan and Singh 1971).
a thin-walled septicidal capsule. Seeds ellipsoid to
oblong, 0.5–1.5 mm long, with finely reticulate or P O L L E N M O R P H O L O G Y . Pollen is tricolp(oroid)
smooth surfaces; endosperm reduced to 1–5 cell ate with an equatorially constricted colpus but no
layers (Bergia) or 0 (Elatine); embryo straight or clearly differentiated endoaperture and finely
less often curved, filling nearly the entire seed; reticulate (Erdtman 1952; Melikian and Dildarian
germination epigaeal. x ¼ 6, 9. 1977).
Two genera and about 35 spp., nearly cosmo-
politan. K A R Y O L O G Y . Most Bergia are based on x ¼ 6,
most Elatine on x ¼ 9, up to high polyploid levels
VEGETATIVE MORPHOLOGY AND ANATOMY. (8 x or perhaps higher), but in both genera x ¼ 10
Bergia capensis has dimorphic roots (D’Al- is also found (Raghavan and Srinivasan 1940;
meida 1941): those anchoring the plants in the IPCN, version Feb. 2004).
mud are sparsely ramified but are almost
completely covered by roots hairs, whereas the REPRODUCTIVE BIOLOGY AND POLLINATION.
plumose water roots lack root hairs but frequently Most Elatine species are adapted to reproduce
have chloroplasts in the cortex. Stomata are very quickly on the mud of periodically dry
40 K. Kubitzki
habitats. Salisbury (1967) documented the enor- walled inner epidermis with tannin (Netolitzky
mous reproductive capacity of the annual E. hex- 1926; Dathan and Singh 1971).
andra and showed that germination is independent
from day length but depends on exposure to light P H Y T O C H E M I S T R Y . Bergia accumulates resin-
of sufficient intensity and duration. ous substances in parenchymatous cells of the
Chasmogamous flowers of Elatine have small axis and leaf, and these substances are also
nectaries but insect visitors are unknown. In Swe- excreted on the stem surface. From Elatine, pro-
den, self-pollination occurs frequently; the fila- cyanidin, delphinidin and ellagic acid have been
ments elongate and bring the anthers in contact reported, while the search for other compounds
with the stigmas. The same mechanism leads to remained unsuccessful (Hegnauer 1969, 1989).
pollination in cleistogamous underwater flowers
known in several species, whereby the pollen A F F I N I T I E S . Previously, Elatinaceae had been
tubes germinate in situ and grow through the considered to be relatives of either the Caryophyl-
anther wall into the stigma. In the stamens of laceae or the Clusiaceae and Hypericaceae; the
cleistogamous flowers, the endothecium may be latter view was still defended by Takhtajan
lacking and the number of microsporangia is (2009). Features common to them include oppo-
reduced (Frisendahl 1927). site leaves, distinct stylodia with capitate stigmas,
capsular fruits and exotegmic seeds, but Clusia-
F R U I T A N D S E E D . The seed coat consists of two ceae differ from Elatinaceae in the lack of stipules.
thin-walled layers (probably testal), the outer tan- In the analysis of Savolainen, Fay et al. (2000),
niniferous, the inner with annular thickenings Elatine was linked with Malpighiaceae, which was
(this layer lacking in Bergia), and a thick-walled, strongly corroborated by further studies (Davis
stellate-undulate layer lignified or not but not and Chase 2004; Wurdack and Davis 2009). Traits
fibrous and probably exotegmic, plus the tegmic such as opposite or whorled stipulate leaves, uni-
mesophyll which is early crushed, and thin- cellular and glandular hairs, the production of
resin (Elatinaceae) or latex (some Malpighia-
ceae), the persistent calyx and the base number
x ¼ 6 may link both families. The problematic
Centroplacaceae have recently been included,
although with weak support, in the “malpighioid”
clade [Centroplacaceae [Malpighiaceae + Elatina-
ceae]], in turn forming a well-supported clade
together with strange companions such as the
chrysobalanoids, putranjivoids and Caryocara-
ceae (Xi et al. 2012).
D I S T R I B U T I O N A N D H A B I T A T S . Bergia is found
throughout the warmer regions of the world, with
a few species in temperate zones. Elatine is
almost cosmopolitan, the majority of species
occurring in temperate regions. Both genera pre-
fer moist disturbed habitats often on sand bars
along rivers, on shores of lakes and ponds, or in
mud flats. Several species are weeds in rice fields,
those of Bergia in the Old World, and some of
Elatine in California, Java and Japan. About half
of the Bergia species are aquatic and semiaquatic,
while the remainder grow on moist soil. Most
Fig. 8. Elatinaceae. A–C Elatine alsinastrum. A Aquatic
plant. B Terrestrial plant. C Seeds. D–F Elatine hydropi-
Elatine thrive in shallow water that seasonally
per. D Terrestrial plant. E Fruit vertically sectioned. F dries out. They are also very common in periodi-
Seed. (Takhtajan 1981, redrawn from Seubert 1845) cally drained fishponds.
Elatinaceae 41
KEY TO THE GENERA Selected Bibliography

1. Plants glabrous; flowers 2–4-merous; sepals obtuse,
without visible midrib; capsules globose or depressed- Behnke, H.-D. 1991. See General References.
globose 2. Elatine Carlquist, S. 1984. Wood and stem anatomy of Bergia
– Plants glandular pubescent throughout; flowers suffruticosa: relationships of Elatinaceae and broader
significance of vascular tracheids and fibriform ves-
5-merous; sepals acute, with a conspicuous, thickened
sel elements. Ann. Missouri Bot. Gard. 71: 232–242.
midrib; capsules ovoid 1. Bergia Corner, E.J.H. 1976. See General References.
D’Almeida, J.F.R. 1941. A contribution to the study of the
biology of Indian marsh and aquatic plants. Part II. J.
1. Bergia L. Bombay Nat. Hist. Soc. 43: 92–96.
Dathan, A.S.R., Singh, D. 1971. Embryology and seed
Bergia L., Mant. Pl. 2: 152 (1771); Leach, J. Adelaide Bot. development in Bergia L. J. Indian Bot. Soc. 50:
Gard. 11: 75–100 (1989), Austral. spp. 362–370.
Davis, G.L. 1966. See General References.
Annual or perennial, herbaceous or suffrutescent; Davis, C.C., Chase, M.W. 2004. Elatinaceae are sister to
roots much branched from a conspicuous tap- Malpighiaceae; Peridiscaceae belong to Saxifragales.
root; stems herbaceous but woody and thickened Am. J. Bot. 91: 262–273.
at base. Leaves (glandular-)pubescent on both Erdtman, G. 1952. See General References.
€
Frisendahl, A. 1927. Uber die Entwicklung chasmo- und
surfaces or glabrous; flowers in dichasia or soli- kleistogamer Bl€ uten bei der Gattung Elatine. Acta
tary; sepals 5, distinct, mucronate, with a thick- Horti Gothob. 3: 99–142.
ened midvein and scarious margins; petals 5; Hegnauer, R. 1969, 1989. See General References.
stamens 5 or 10; ovary 5(6)-locular. Seeds oblong, Kajale, L.B. 1939. A contribution to the life history of
Bergia ammanioides Roxb. J. Indian Bot. Soc. 18:
slightly curved. x ¼ 6. 157–167.
About 24 species, most of them native to Old Melikian, A.P., Dildarian, B.I. 1977. Comparative anato-
World tropics including Australia; B. capensis L. mical and palynological study of representatives of
Elatinaceae family. Biol. Zhurn. Armenii 30(11):
the most widespread species, ranging from south- 44–49. (in Russian with Armenian summary)
ern Africa to India and Indonesia. Metcalfe, C.R., Chalk, L. 1950. See General References.
M€uller, F. 1877. Untersuchungen u €ber die Struktur einiger
Arten von Elatine. Flora 60: 481–496, 519–526.
Netolitzky, F. 1926. See General References.
2. Elatine L. Niedenzu, F. 1925. Elatinaceae. In: Engler & Prantl, Nat.
Elatine L., Sp. pl. 1: 367 (1753). Pflanzenfam., 2nd edn, 21: 270–276.
Raghavan, T.S., Srinivasan, V.K. 1940. A contribution to
Small aquatic or emergent annuals or short-lived the life history of Bergia capensis Linn. J. Indian Bot.
perennials; stems soft, chlorophyllous, with 5–10 Soc. 19: 283–291.
Salisbury, E.J. 1967. On the reproduction and biology of
air chambers visible in cross section; flowers sol- Elatine hexandra (Lapierre) DC. (Elatinaceae); a typ-
itary, (2)3(4)-merous; sepals inconspicuous; ical species of exposed mud. Kew Bull. 21: 139–149.
petals as long as sepals, in some spp. with floral Savolainen, V., Fay, M.F. et al. 2000. See General Refer-
nectaries; anthers 4-sporangiate (2–3-sporangiate ences.
Seubert, M. 1845. Elatinarum monographia. Nov. Acta
in cleistogamous flowers); ovaries broadly ovoid, Acad. Caes. Leop.-Carol. Nov. Curios. 21(1): 34–60.
(2)3(4)-locular; placentation basal (axile). Cap- Takhtajan, A. 1981. See General References.
sules delicate, the seeds visible within; seeds Takhtajan, A. 2009. See General References.
Tucker, G.C. 1986. The genera of Elatinaceae in the south-
straight or slightly curved. x ¼ 9. eastern United States. J. Arnold Arb. 67: 471–483.
About 10 species, in all continents, most of Wurdack, K.J., Davis, C.C. 2009. See General References.
them in North America and Eurasia. Xi, Z. et al. 2012. See General References.
Erythroxylaceae
Erythroxylaceae Kunth in H.B.K., Nov. Gen. Sp. 5, ed. 4: 175, ed. f: 153 (1822), nom. cons.
Nectaropetalaceae Exell & Mendonça (1951).
V. B I T T R I C H
Glabrous trees or shrubs, evergreen or deciduous, In Erythroxylum they are often longitudinally
sap sometimes milky or coloured; alkaloids com- striate or ribbed. Unbranched finger- or awl-
mon. Leaves alternate (opposite), simple, entire, shaped colleters are present in the axils of the
pinnately veined, with intrapetiolar (interpetio- stipules of all genera (Stevens 2001 onwards;
lar), often caducous stipules, colleters present. Thiebaut and Hoffmann 2005), but easily over-
Flowers small, generally bisexual, more rarely looked. Leaf texture and duration is often corre-
unisexual, distylous [Erythroxylum], axillary, sol- lated with the habitats of the species (Rury 1981).
itary or in fascicles, rarely in pedunculate sub- Many evergreen species are sclerophyllous. In
umbelliform cymes. Calyx (4)5-lobed, persistent, Erythroxylum persistent cataphylls, which
lobes valvate or imbricate in bud; petals (4)5, correspond to the stipules of reduced leaf
distinct, imbricate in bud, often shortly clawed, laminas, are common on lateral twigs
early caducous, mostly with a ligular bilobed mainly near their base (Weberling et al. 1980).
appendage near the base; stamens 10(–12), biseri- The aggregation of these cataphylls is often
ate, filaments connate at the base into a cup or termed "ramenta".
tube, generally persistent; anthers basifixed, tet- Leaf anatomy was studied by Rury (1981) and
rasporangiate, longitudinally dehiscent; ovary van Welzen and Baas (1984). Hairs are absent. A
superior, 2–3(–4)-carpellate, all or only one of papillose abaxial epidermis occurs in several Ery-
the locules fertile, bearing 1(2) pendulous crassi- throxylum spp. Mucilage cells in the leaf epider-
nucellate ovules; stylodia distinct or a single style. mis are reported from Erythroxylum spp. and
Fruit a monospermous drupe (a 2- or 3-seeded Aneulophus, and crystalliferous epidermis cells
capsule). Seeds with starchy endosperm (absent), containing a solitary crystal of calciumoxalate
embryo straight, aril absent (present). were found in some Erythroxylum (Solereder
Four genera and about 240 spp., pantropical/ 1908). The stomatal type is paracytic or paralle-
subtropical. The large genus Erythroxylum is locytic. The midrib shows mostly a simple arc of
pantropical, the other genera are restricted to collateral vascular tissue, very rarely more com-
Africa south of the Sahara. plex vascular systems with pith bundles. Cristar-
que cells, which are either restricted to the bundle
VEGETATIVE MORPHOLOGY AND ANATOMY. Growth sheath or occur additionally in the ground tissue
forms include prostrate shrubs, rosette shrubs, of petiole and midrib, are reported by van Welzen
small and tall trees; they are generally correlated and Baas (1984). Mesophyll sclereids are com-
with the habitats (Rury 1985); the buds are mon, but typically absent in deciduous and in
perulate (Stevens 2001 onwards). The leaves are many mesophytic evergreen leaves (Rury 1981).
alternate, opposite in Aneulophus, entire, and Data about stem anatomy were reported by
involute in bud in Erythroxylum. This vernation Heimsch (1942), Metcalfe and Chalk (1950), and
type often results in the presence of two parallel Rury (1985). Cork develops superficially: epider-
lines on the mature leaf surface. The stipules are mal in Aneulophus, subepidermal in Erythroxy-
mostly intrapetiolar (interpetiolar in Aneulophus) lum; secretory cells with gum-like contents are
and usually more or less connate; they include present in the cortex; nodes are unilacunar; cor-
the terminal bud and are persistent or caducous. tical bundles are usually present in young stems;
44 V. Bittrich
compressed inflorescence axis. Pedicels are

usually present and have two or rarely four (Aneu-
lophus) tiny bracteoles at the base; above the brac-
teoles the pedicel is articulated (Eichler 1878).
The flowers are basically 5-merous, actino-
morphic, bisexual or rarely unisexual. The sepals
are united below, forming a mostly campanulate
and persistent calyx; the lobes are imbricate or
valvate in bud. The petals are caducous, alternat-
ing with the calyx lobes, imbricate in bud, and
mostly shortly clawed; a nectariferous entire or
bilobed appendage is inserted at the apex of the
Fig. 9. Erythroxylaceae. Aneulophus africanus. A Flower- claw, more rarely it is absent. Narayana (1960)
ing branchlet. B Flower with basal bracts. C Androecium reports the presence of obdiplostemony for
and gynoecium. D Gynoecium. E Same, vertically sec- two Erythroxylum but its absence for another spe-
tioned. F Ovary, transversal section. G Fruit. H Seed. cies. There are 10(–12) stamens, subequal or
(Badré 1973)
unequal (then forming two sets) in length, and
persistent; the filaments are basally united into
vessel perforations are simple; in most cases ever- a sometimes glandular cup or tube of various
green species have longer narrower vessel ele- length and, according to Eichler (1878), a disk
ments than deciduous species (Rury 1981); the participates in their formation; the anthers are
pits to the parenchyma are generally simple. basifixed, tetrasporangiate, two-locular, and
Vasicentric tracheids are present. Fibrous ele- dehisce by longitudinal slits. The ovary is supe-
ments consist of fibre tracheids or libriform rior, 2–3-carpellate, 2–3-loculate, with one or two
fibres. The axial parenchyma is predominantly axile pendulous, epitropous ovules per locule, or it
apotracheal-diffuse and paratracheal, rarely with has only one uniovulate fertile locule (Erythroxy-
some apotracheal banded parenchyma; rays are lum); the stylodia are free or basally or completely
uni- to triseriate, rarely pluriseriate (up to 5 cells connate; the stigmas are capitate, depressed-capi-
wide), heterocellular or nearly homocellular. Uni- tate or clavate, rarely acute, free or rarely connate
and biseriate rays show silica grains mainly in and then bilobed (Pinacopodium).
species of two neotropical sections of Erythroxy-
lum; prisms of calcium oxalate occur in the E M B R Y O L O G Y . Both integuments of Erythroxy-
parenchyma strands of certain Erythroxylum lum coca are of dermal derivation (Boesewinkel
spp. and in ray cells of Nectaropetalum and Pina- and Geenen 1980). Embryo sac development is of
copodium (Rury 1981; Chalk 1983). According to the Polygonum type, the antipodals degenerate
Rury (1985) xylem features, life form and leaf early. The nucellus is partly resorbed by the
structural features are strongly interrelated, embryo sac. The inner layer of the inner integu-
reflecting habitat conditions. Wood anatomical ment is differentiated into a pigmented endothe-
specialization within the family may be of xero- lium (Tunmann and Jenzer 1910). Endosperm
morphic as well as of mesomorphic nature. Sieve- formation is nuclear with later centripetal wall for-
element plastids of Erythroxylum spp. contain mation. The tapetum is of the secretory type (Nar-
about ten rectangular protein crystals and no ayana 1960). The mature pollen grains are 3-celled.
starch (type P5c, Behnke 1988).
P O L L E N M O R P H O L O G Y . An overview is given by
I N F L O R E S C E N C E A N D F L O W E R S T R U C T U R E . Sin- Oltmann (1968). Pollen grains are suboblate to
gle flowers or few- to many-flowered sessile to prolate and generally tricolporate; in Aneulophus
subsessile fascicles are borne in the axils of leaves the colpi are strongly reduced. Pollen size was
or cataphylls, rarely distinctly pedunculate inflor- observed to be significantly different in longer
escences are formed axillary or on axillary short and shorter stamens of the same flower of Ery-
shoots (Pinacopodium). According to Schulz throxlum coca, and pollen from thrum flowers are
(1907) the fascicles are cymes with a strongly larger than pollen from pin flowers (Ganders 1979).
Erythroxylaceae 45
R EPRODUCTIVE S YSTEMS AND POLLINATION. Since

Darwin (1877) it is known that Erythroxylum
is often heterostylous. Many species of this
genus and of Nectaropetalum have dimorphic
flowers. There are also reports of three or four
different flower morphs (see Payens 1958), but
according to Ganders (1979) this is probably a
misinterpretation due to continuous variation in
the relative lengths of the two sets of stamens in
the flowers. As common in heterostylous plants,
pin morphs often produce more but smaller pol-
len grains than the drum morphs (Ganders 1979;
Pailler et al. 1998). In E. coca, Ganders (1979)
found a strong self-incompatibility system linked
with floral dimorphism. In the self-compatible
pin morph of E. novogranatense he observed
illegitimate cross-compatibility, whereas Pailler
et al. (1998) reported slight illegitimate cross-
compatibility in the self-incompatible pin morph
of E. laurifolium, but without any effect on popu-
lation structure. Burck (1895, cited in Payens
1958) observed abundant fruiting in introduced
plants of E. novogranatense in Java, where only
the partial self-compatible pin morphs occur.
He also discussed the possibility that in some
species the heterostyly might have led to dioecy
(see Barrett and Richards 1990). In these
cases, the flowers of the pin morph possess abor-
Fig. 10. Erythroxylaceae. Erythroxylum macrophyllum. tive stamens, the flowers of the drum morph only
A Flowering branch with cataphylls and stipules. B Cata- a rudimentary pistil (Schulz 1907). A dioecious
phyll. C Floral bud with floral bract. D Flower. E Same,
with petals and two sepals removed. F Adaxial, lateral and species was also reported by Bawa and Opler
medial section of petal. G Androecium and pistil. (1975) from Costa Rica. As an exception to
H Medial section of androecium and pistil, with cross "Baker’s law", Pailler et al. (1998) found distyly
section of ovary (right) and detail of anther. I Pistil. in three species of Erythroxylum endemic to
J Part of infructescence. K Fruit and seed. (Reproduced
with the kind permission of the artist Bobbi Angell) the Mascarene islands and a heteromorphic
incompatibility system in the one species,
The tectum is finely reticulate or punctate. The E. laurifolium, studied in detail. In E. undulatum
colpi are of different length and in some species the flowers are monomorphic (pin morph) and
of Erythroxylum united at the pole. The pori are produce apomictic seeds from sporadically devel-
isodiametric or not; in a few species of Erythrox- oping aposporous embryo sacs; this species is
ylum they are united in a ring in the equatorial zone probably a sterile triploid of hybrid origin
(zonorate). In various Erythroxylum opercula are (Berry et al. 1991).
developed. According to Oltmann (1968) pollen of Erythroxylum flowers are reported to be fra-
the Old World representatives is more heteroge- grant and nectar-producing. The petal appen-
neous and shows the basal types within the family. dages may restrict the access of flower visitors
to the nectar. Various insects, including Hyme-
K A R Y O L O G Y . Most counts show that Erythrox- noptera, Diptera and Lepidoptera, with bees or
ylum has a chromosome base number of x ¼ 12. wasps as the most important pollinators, were
The counts of Berry et al. (1991) in the agamos- observed as flower visitor in Erythroxylum (Schulz
permic E. undulatum Plowman of n ¼ ca. 18 1907; Domı́nguez et al. 1997; Barros 1998).
suggest that this species is a triploid of hybrid Flower visitors of E. havanense were found to
origin. not distinguish between the two floral morphs,
46 V. Bittrich
but fruit set of thrum plants was significantly nolic compounds include flavonoids, mainly var-
higher than that of pin plants (Domı́nguez et al. ious flavonols (cf. Johnson et al. 1998) as well as
1997). This difference is probably due to the caffeic acid and derivatives and methyl salicylate,
much higher male sterility of the thrum flowers, which were found in the essential oil of coca
leading to a partially gynodioecious system with leaves. A mixture of essential oils and resins in
pin flowers behaving as bisexual flowers. Species the wood of Erythroxylum contains mono-, di-
of Erythroxylum with quite similar flower mor- and sesquiterpenoid metabolites. Triterpenoid
phology may occur sympatrically and flower syn- compounds were found in the waxes of leaves
chronously for months. Such species were and fruits. Saponins were reported from a num-
observed to partly share the same pollinators ber of species.
(Barros 1998).
S U B D I V I S I O N A N D A F F I N I T I E S . Aneulophus
F R U I T , S E E D A N D D I S P E R S A L . The fruits of Ery- deviates remarkably in several characters from
throxylum and Nectaropetalum are drupes, the other genera. In the cladogram of Schwarz-
monospermous in Erythroxylum and probably bach and Ricklefs (2000), based on DNA
bispermous in Nectaropetalum. Erythroxylum sequence data, it forms the basal branch of the
spp. have mostly red drupes with a well-devel- family, and Nectaropetalum and Pinacopodium
oped endocarp. Aneulophus has a 3–4-locular appear as sister taxa to Erythroxylum. The mono-
septicidal capsule, with one seed per locule. The phyly of the vast genus Erythroxylum would need
mature fruit of Pinacopodium is unknown. further confirmation.
The ovule (Erythroxylum) is anatropous, Formerly, the family was generally consid-
bitegmic and crassinucellate with an elongated ered as closely related to the Linaceae s.str. from
nucellus (Corner 1976; Boesewinkel and Geenen which it differs by traits such as the presence of
1980). The micropyle is formed by the inner colleters, mostly appendaged petals, the number
integument only. The ventral raphe shows simple of (functional) stamens, mostly drupaceous
or multiple bundles. The tegmen is multiplicative. fruits, and possibly the chromosome base num-
In Erythroxylum the exotegmen with fibre trac- ber. Hallier (1923) stressed similarities with
heids forms the mechanical layer of the seedcoat; Hugoniaceae (Linaceae s.l.), which also have
the other layers are crushed during seed develop- drupacous fruits. A comparison of leaf anatomi-
ment. The embryo is straight, an endosperm is cal characters revealed various similarities with
present or absent. The seeds of Aneulophus afri- Linaceae, Hugoniaceae, Ixonanthaceae and
canus are arillate (Badré 1973). Humiriaceae (van Welzen and Baas 1984). Boe-
The drupaceous, reddish, sometimes sweet sewinkel and Geenen (1980) listed seed charac-
fruits of most Erythroxylaceae suggest ornithoch- ters common to Erythroxylum and Linum, but it
ory. Birds were observed already by Martius to remains questionable which of these are syna-
feed on Erythroxylum fruits (Schulz 1907). pomorphic.
rbcL sequences analyzed by Setoguchi et al.
P H Y T O C H E M I S T R Y . As summarized by (1999), Savolainen et al. (2000) and Schwarzbach
Hegnauer (1981, 1989), the present knowledge is and Ricklefs (2000) as well as the multi-gene
restricted to species of Erythroxylum. Condensed analyses of Wurdack and Davis (2009), Soltis
tannins are common, ellagitannins were very et al. (2011) and Xi et al. (2012) all resolved a
rarely reported. Alkaloids are common, especially sister group relationship of Erythroxylaceae and
tropane and pseudotropane alkaloids, but it is Rhizophoraceae and finally a clade comprising
only in E. coca and E. novogranatense that con- [Ctenolophonaceae [Erythroxylaceae + Rhizo-
siderable amounts of cocaine and cinnamylco- phoraceae]]. This is also supported by the shared
cain were found. In contrast to tropine occurrence of hygroline and pyrrolodine alka-
alkaloids, hygroline ecgonine alkaloids seem to loids, sieve tube plastids with protein crystals
be restricted to Erythroxylaceae and Rhizophor- and involute leaf blades in Erythroxylaceae and
aceae, ecgonine alkaloids possibly exclusive to Rhizophoraceae. However, the merging of these
Erythroxylaceae, or are very rare in angiosperms. two families in a broadened family concept, as
Various esters of tropanols were reported. Phe- suggested by APG II (2003), could cause more
Erythroxylaceae 47
confusion than it would help to clarify and is not monies (Davis 1997). For this reason, plans for
followed here. the eradication of coca plants were strongly
opposed by ethnobotanists like the late Timothy
D I S T R I B U T I O N A N D H A B I T A T S . Erythroxylum Plowman. In the last century, coca extracts some-
has a pantropical distribution with centres in times mixed with red wine (“Vin Tonique
the Neotropics and on Madagascar. The other Mariani”), but also in form of candies, cigarettes,
genera are restricted to tropical and subtropical soft drinks, etc., became extremely popular as
Africa south of the Sahara; one species of Nectar- stimulants and for various medicinal purposes.
opetalum was reported from Madagascar. After 1906 cocaine had to be removed from Coca-
Nectaropetalum occurs in forest fringes of Cola and similar soft drinks, but the residue of
low altitude and in dry forests. Erythroxylum the coca leaf extracts is still used for flavouring
species are found in various habitats, such as (Davis 1997). In Brazil “catuaba”, a Tupi name for
the understorey of tropical wet lowland and mon- certain Erythroxlum spp. but also used for a few
tane forests, evergreen and deciduous forests, species of other families, is renowned as a tonic
savannah woodland, sandy or rocky shores and and aphrodisiac in form of a tea or alcoholic
hills (Malaysia), and restingas and caatingas (NE tincture prepared from the bark. Several species
Brazil). Deciduous species of Erythroxylum are of Erythroxylum and Nectaropetalum have hard
usually found in dry or seasonally dry habitats, and durable wood used for various purposes.
whereas evergreen ones occur in a broad range of Some local uses of Old World Erythroxylaceae
habitats (Rury 1981). Aneulophus africanus are reported by Hegnauer (1981).
occurs in swampy forests and Pinacopodium
spp. in moist forests of West Africa. KEY TO THE GENERA
P A L A E O B O T A N Y . Fossils from the Eocene of 1. Leaves opposite 1. Aneulophus
Argentina are considered to belong to Erythrox- – Leaves alternate 2
ylaceae. 2. Flowers in pedunculate subumbelliform inflores-
cences; style bearing a bilobed stigma
E C O N O M I C I M P O R T A N C E . Erythroxylum coca 2. Pinacopodium
and E. novogranatense are the source of cocaine, – Flowers solitary or in sessile or subsessile fasci-
a dangerous narcotic drug. Besides its economic cles; stylodia distinct or connate into a single
importance for the coca planters in NW South style with distinct stigmas 3
America, it has become notorious for its social, 3. Stylodia (2)3, distinct or only partly connate;
economic and political impact on Central and petal appendages large (usually longer than the
South America and in the First World. Cocaine sepals) 3. Erythroxylum
has lost its importance as a local anaesthetic to – Style 1, stigmas 2; petal appendages small and
synthetic products. Chewing of coca leaves often hidden by the sepals or absent
with some unslaked lime or ashes of plants as an 4. Nectaropetalum
alkaline catalyst has been a common custom in
north-western South America for several thou- GENERA OF ERYTHROXYLACEAE
sand years; throughout the northwest Amazon
region, powdered coca leaves mixed with ash are 1. Aneulophus Benth. Fig. 9
slowly dissolved in the mouth (e.g. Rury and
Aneulophus Benth. in Benth. & Hook.f., Gen. pl. 1:
Plowman 1983; Davis 1997). Chewing of the
244 (1862).
leaves confers endurance to hunger and exhaus-
tion. Coca leaves not only contain cocaine, but Shrubs with opposite leaves; stipules interpetiolar.
are very rich in vitamins and minerals. Thus, they Flowers numerous (up to 50) in congested inflor-
are sometimes considered not as a drug but a escences; pedicel with 4 minute bracteoles at base;
food with mildly stimulating effects essential for petals much longer than sepals, white, shortly
the survival of the native Andean people. These unguiculate, unappendaged, thickened along the
revere coca as a sacred plant, which formerly midrib; stamens very shortly united at base;
often played an important role in religious cere- stylodia 3(4) united below, ovary 3(4)-locular
48 V. Bittrich
with two ovules in each locule. Capsule septicidal Shrubs or tree. Leaves alternate, with pointed tips
with one arillate seed in each locule. or acuminate, stipules mostly completely united.
One or two spp., tropical West Africa. Flowers axillary, single or in small groups; sepals
valvate, petals white, clawed or not, with or more
rarely without small appendages near the base;
2. Pinacopodium Exell & Mendonça
filaments united below into a shallow cup, some-
Pinacopodium Exell & Mendonça, Bol. Soc. Brot. times bearing 5 glands outside; the single style
II, 25: 105 (1951). with 2 distinct stigmas; ovary 2-locular, with one
Morelodendron Cavaco & Normand (1951). ovule in each locule.
Eight spp., South Africa to Kenya, Zaire,
Trees with alternate leaves. Flowers in axillary
Madagascar.
pedunculate, subumbelliform cymes; sepals val-
vate; petal appendages indistinct; stamens basally
united into a short tube; style 1, very short, stigma
capitate, bilobed; ovary two-locular with one Selected Bibliography
ovule in each locule. Fruit unknown.
Two spp. in tropical moist forests of West APG II. 2003. The Angiosperm Phylogeny Group. See
Africa (Gabon and Mayombé). General References.
Badré, F. 1973. Erythroxylaceae. In: Aubreville, A. et al.
(eds.) Flore du Gabon, fasc. 21: 49–54. Paris:
3. Erythroxylum P. Browne Fig. 10 Muséum National d’Histoire Naturelle.
Barrett, S.C.H., Richards, J.H. 1990. Heterostyly in tropi-
Erythroxylum P. Browne, Civ. Nat. Hist. Jamaica: cal plants. Mem. New York Bot. Gard. 55: 35–61.
Barros, M.A.G. 1998. Sistemas reprodutivos e polinização
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18–161 (1907). throxylaceae) do Brasil. Rev. Brasil. Bot. 21: 159–166.
Bawa, K.S., Opler, P.A. 1975. Dioecism in tropical forest
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compressed at apex, often with persistent cata- Behnke, H.-D. 1988. Sieve-element plastids and system-
phylls. Leaves alternate, leaf ptyxis involute; sti- atic relationships of Rhizophoraceae, Anisophylla-
pules united to the apex or bifid, mostly ceae, and allied groups. Ann. Missouri Bot. Gard.
75: 1387–1409.
bicarinate and striate, and 2–3-setulose. Flowers Behnke, H.-D. 1991. Distribution and evolution of forms
in axillary fascicles subtended by bracts, often and types of sieve-element plastids in the dicotyle-
dimorphic, sometimes unisexual; sepals valvate dons. Aliso 13: 167–182.
Berry, P.E., Tobe, H., Gómez, J.A. 1991. Agamospermy
or imbricate; petals with a usually bifid append- and the loss of distyly in Erythroxylum undulatum
age on the inner face; filaments connate below; (Erythroxylaceae) from northern Venezuela. Amer.
stylodia distinct or connate up to the middle; J. Bot. 78: 595–600.
ovary (2)3-locular, usually only one cell fertile; Boesewinkel, F.D., Geenen, J. 1980. Development of ovule
and seed-coat of Erythroxylum coca Lamk. Acta Bot.
drupe reddish, monospermous. Neerl. 29: 231–241.
Ca. 230 spp., tropical, especially Neotropics Chalk, L. 1983. Axial parenchyma. In: Metcalfe, C.R.,
and Madagascar. E. coca Lam. and E. novograna- Chalk, L. (eds.) Anatomy of the dicotyledons, ed. 2.
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introduction. Oxford: Clarendon Press, pp. 15–20.
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subdivision of the genus in sections by Schulz Darwin, C. 1877. The different forms of flowers on plants
(1907) is often artificial (Plowman, cited in Rury of the same species. London: Murray.
1985). Davis, W. 1997. One river – science, adventure and hallu-
cinogenics in the Amazon basin. London: Simon &
Schuster.
Domı́nguez, C.A., Ávila-Sakar, G., Vázquez-Santana, S.,
4. Nectaropetalum Engl. Márquez-Guzmán, J. 1997. Morph-biased male ste-
Nectaropetalum Engl., Bot. Jahrb. Syst. 32: 109 rility in the tropical distylous shrub Erythroxylum
havanense (Erythroxylaceae). Amer. J. Bot. 84:
(1902). 626–632.
Peglera Bolus (1907). Eichler, A.W. 1878. Bl€ uthendiagramme, 2. Teil. Leipzig:
Wilhelm Engelmann, pp. 342–343.
Erythroxylaceae 49
Ganders, F.R. 1979. Heterostyly in Erythroxylum coca Rury, P.M., Plowman, T. 1983 (1984). Morphological
(Erythroxylaceae). Bot. J. Linn. Soc. 78: 11–20. studies of archeological and recent coca leaves (Ery-
Hallier, H. 1923. Beitr€age zur Kenntnis der Linaceae (DC. throxylum spp.). Bot. Mus. Leafl. Harvard Univ. 29:
1819) Dumort. Beih. bot. Centralbl. 39: 1–178. 297–341.
Hegnauer, R. 1981. Chemotaxonomy of Erythroxylaceae Savolainen, V. et al. 2000. See General References.
(including some Ethnobotanical notes on Old World Schulz, O.E. 1907. Erythroxylaceae. In: Pflanzenreich, IV.
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Heimsch, C. 1942. Comparative anatomy of the secondary nities of Rhizophoraceae and Anisophylleaceae, and
xylem in the "Gruinales" and "Terebinthales", of intergeneric relationships within Rhizophoraceae,
Wettstein, with reference to taxonomic grouping. based on chloroplast DNA, nuclear ribosomal DNA,
Lilloa 8: 83–198. and morphology. Amer. J. Bot. 87: 547–564.
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chem. Syst. Ecol. 26: 743–759. Solereder, H. 1908. Systematische Anatomie der Dicotyle-
Metcalfe, C.R., Chalk, L. 1950. See General References. donen. Erg€anzungsband. Stuttgart: Ferdinand Enke,
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Oltmann, O. 1968. Die Pollenmorphologie der Erythrox- Stevens, P.F. 2001 onwards. See General References.
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Euphorbiaceae
Euphorbiaceae Juss. (1789), nom. cons.
Phyllanthaceae Martinov (1833).
Peraceae Klotzsch (1859).
Picrodendraceae Small (1917), nom. cons.
G.L. W E B S T E R { 1
Monoecious or dioecious trees, shrubs, or herbs, deciduous in fruit; petals + or 0, greenish to

sometimes succulent or scandent; stems with or white, yellow, pink, or red; disk + or 0, extrastam-
without laticifers; indumentum simple, malphigh- inal or less commonly interstaminal, entire to
iaceous, stellate, or lepidote, sometimes glandular toothed, lobed, or dissected; stamens (1–)3–50
or 0. Leaves alternate, spiral or distichous, less (400), filaments distinct or connate; anthers
commonly opposite or whorled; leaf blades simple mostly 2-locular, dehiscing longitudinally or later-
to palmately lobed or compound, pinnately to ally, extrorse or less commonly introrse; gynoe-
palmately nerved, sometimes with basal, laminar, cium syncarpous, ovary (1)2–5(20)-locular;
or marginal glands; stipules persistent or decidu- placentation axile (basal); ovules 1 or 2 per locule,
ous, sometimes reduced or 0. Inflorescences axil- anatropous or hemitropous, epitropous, usually
lary or terminal, sometimes cauliflorous, thyrsoid, inserted beneath an obturator. Fruits mostly schi-
paniculate dichasial, glomerulate or synanthial, or zocarpic, splitting from the columella, commonly
flowers solitary; bracts sometimes colored, glandu- explosively dehiscent, mesocarp often separating
lar, or hypertrophied. Flowers unisexual, usually from endocarp, sometimes baccate or drupaceous.
actinomorphic; perianth segments distinct or Seeds often carunculate; seed coat mostly exoteg-
connate, imbricate to valvate, rarely obsolete or mic; endosperm copious, often fleshy, or 0, some-
suppressed; sepals (–)3–6(10), entire or lobed, times oily; embryo straight, curved or plicate.
sometimes glandular, the pistillate persistent or
1
By the end of 2004 Dr. Grady L. Webster sent me a far advanced version of his treatment of the Euphorbiaceae for this
book series and, in June 2005 a few months before his death (Oct. 27, 2005), I received an additional document from
him dealing with a review of suprageneric and generic taxa in the family. That year, I could not immediately take care
of these documents because I was busy with other volumes of this series. In 2008, through the courtesy of Drs. K.J.
Wurdack and P.E. Berry, I received what I thought to be further updated documents left by Grady at the time of his
death in his laboratory; contrary to my expectation, however, these did not differ from those supplied earlier by Grady
himself. Although the author had made perusal of many taxonomic revisions and publications on pollen morphology
and seed anatomy, it was evident that the manuscript needed substantial revision in the light of more recent
publications in molecular systematics accrued before and after Grady’s death. Regrettably, Dr. Wurdack, the born
heir of Grady, did not wish to revise the manuscript so that this task fell onto me, in an effort to not loose Grady’s work
spanning several years. The paper thus contains many changes introduced by me without being able to secure the
author’s formal approval, but these were essential for this paper to reflect the high standard that distinguished
Dr. Webster’s scientific production. Whereas the molecular-cladistic papers on the euphorbs in the widest sense,
mainly those by Wurdack and Berry and their co-authors, were of greatest value for me in this task, I must emphasise
that the main objective of this series is to give baseline information, whereas further elaboration of a cladistic
classification of the family would have been beyond my desire and capacity.
My thanks go to Drs. K.J. Wurdack and P.E. Berry for showing me the manuscripts left by the late Dr. Webster.
Prof. W.S. Armbruster is warmly thanked for going through parts of the manuscript lying within his interest and for
updating the section on reproductive biology. I am also most grateful to Dr. R. Secco for his comments on Amazonian
euphorbs, and to Dr. V.W. Steinmann for advice on subtribe Euphorbiinae.
K. Kubitzki
52 G.L. Webster
The family is here construed to include 299 succulent Euphorbias to the umbracular leaves of
genera with 8,000 species; these are grouped into Macaranga gigantea with blades exceeding 50 cm
47 tribes in seven subfamilies, the biovulate Phyl- in diameter. Leaves are usually stipulate, although
lanthoideae and Oldfieldioideae, and the uniovulate stipules have been lost in various taxa, notably in
Peroideae, Cheilosoideae, Acalyphoideae, Crotonoi- Croton and Euphorbia (Uhlarz 1978). Leaf blades
deae, and Euphorbioideae (see below under CIRCUM- are prevailingly simple or lobed, but palmate/pin-
SCRIPTION AND SUBDIVISION OF THE FAMILY). Although the nately dissected laminae occur in Oldfieldioideae
family is nominally cosmopolitan (except for the (Piranhea), and Crotonoideae (Hevea, Joannesia,
polar regions), it is poorly represented in cool tem- Leeuwenbergia, and species of Manihot). There is
perate zones and best developed in subtropical and some variation in ptyxis (leaf blade folding in bud):
tropical regions. the study of Cullen (1978), although based on a
limited sample, indicates that most of the sub-
V E G E T A T I V E M O R P H O L O G Y . Euphorbiaceae are families have conduplicate ptyxis, but Euphorbio-
highly variable in vegetative structure, with a great ideae are mainly supervolute. Leaves are persistent
range of life forms, including trees, shrubs, and in tropical rainforest taxa, but usually deciduous in
perennial and annual herbs. Sympodial growth genera adapted to monsoonal or arid climates.
related to inflorescence production is common, Leaf blade venation is mostly brochidodromous;
especially in subfamilies Crotonoideae and Euphor- margins may be entire or toothed with "theoid" or
bioideae (Hallé 1971). Scandent life forms are "violoid" teeth (Hickey and Wolfe 1975). An atlas of
rare in Euphorbioideae (tribe Stomatocalyceae), leaf venation has been published by Klucking (1997,
unknown in Oldfieldioideae and Crotonoideae 2003).
(except Omphalea), but common in Acalyphoideae The widespread adaptation of Euphorbiaceae to
tribe Plukenetieae. In subfam. Phyllanthoideae the arid habitats has led to characteristic morphological
branching pattern tends to be monopodial and and physiological traits. The succulent life form is
inflorescences axillary, and there is a striking ten- well represented in Euphorbiaceae, especially in
dency toward monopodial plagiotropic branching subfamilies Crotonoideae and Euphorbioideae.
as in Flueggea and Phyllanthus. A further speciali- Succulent species, mostly caudiciform or geophytic,
zation of this pattern yields the deciduous, flori- also occur in African and American Jatropha.
ferous, phyllomorphic branchlets found in several The immense diversity of the genus Euphorbia,
genera of the Phyllanthinae, including Breynia, Glo- which comprises over 2,000 recognized species, is
chidion, Phyllanthus, and Sauropus. This highly paralleled by enormous structural, ecological and
specialized growth from, first documented by Din- physiological variation; it is the only land plant
gler (1885) and designated as "phyllanthoid branch- genus in which the three major photosynthetic
ing" by Webster (1956), includes some unique pathways are expressed. In a phylogenetic analysis
patterns such as the relegation of photosynthetic Horn et al. (2012) identify a woody, non-succulent
leaves and flowers to specialized deciduous branch- growth form with C3 photosynthetic pathway as
lets in Phyllanthus and related genera (Dingler ancestral. Cactiform succulents with CAM photo-
1885; Webster 1956; Roux 1968; Bancilhon 1971; synthesis amount to nearly 650 species and have
Rossignol and Rossignol 1985). Subfam. Oldfield- arisen in two clades within Euphorbia; most of them
ioideae resembles the Phyllanthoideae in having are concentrated in eastern and southern Africa.
relatively unspecialized monopodial branching pat- An independent evolution of herbs has taken place
terns. The most divergent life form in the Euphor- in the Esula Clade and Chamaesyce Clade of
biaceae is found in a single neotropical section Euphorbia, in the latter accompanied by the acqui-
(Salviniopsis) of Phyllanthus. Mature plants float sition of all three photosynthetic pathways.
unrooted on the surface of white-water streams
and lakes in the Amazonian region. V E G E T A T I V E A N A T O M Y . Wood anatomy in the
Phyllotaxis in Euphorbiaceae is predominantly Euphorbiaceae was reviewed by Pax (1884) in his
alternate (spiral or distichous), but opposite or first study of the family, and treated in greater detail
whorled leaves are common in the Oldfieldioideae by students of Radlkofer (Rittershausen 1892;
and Euphorbioideae. Leaves in the Euphorbiaceae Froembling 1896; Rothdauscher 1896; Herbert
vary in size from scale leaves less than 1 mm long in 1897), and by Solereder (1908). Metcalfe and Chalk
Euphorbiaceae 53
(1950) synthesized much of these data in their clas- phoideae. Rudall (1994) suggests that sclereids
sic work on the anatomy of the dicots. From the (in at least some instances) may be homologous
studies of Mennega (1987) on Phyllanthoideae and with laticifers.
Hayden (1994) on Oldfieldioideae, it appears that The indumentum of leaves and stems is highly
the apparently least specialized wood structure is diverse in the Euphorbiaceae (Metcalfe and Chalk
present in Phyllanthoideae tribes Wielandieae and 1950), and has often been used as a diagnostic
Antidesmateae, evidenced by thick-walled, non-sep- character. In the biovulate subfamilies it consists
tate fibers, abundant axial parenchyma, and vessel usually of simple unicellular or uniseriate tri-
elements with scalariform perforations. In other chomes; bifurcate trichomes are known from Aus-
tribes of Phyllanthoideae, there are trends toward trobuxus. Both glandular and branched hairs are
wood with septate, thin to moderately thick-walled common in the uniovulate subfamilies. In Croton,
fibers, scanty parenchyma, and porous vessel ele- there is great diversity in the size, color, and num-
ment perforations. In Oldfieldioideae, fibers are ber of rays of branched trichomes (Webster 1993).
non-septate, the rays narrow and heterocellular, Dibrachiate (malpighiaceous) trichomes are
axial parenchyma is abundant and diffuse or in characteristic of various genera in the Acalyphoi-
wide bands, and vessel perforations are simple deae and Crotonoideae. Particularly notable are the
with exception of Podocalyx (and Paradrypetes), stinging hairs found in Acalyphoideae (Plukene-
which have multiple perforation plates in very long tieae) and Crotonoideae (Cnidoscolus). The urticant
vessel elements. With some exceptions (Acalypha, hairs have clearly evolved independently in the two
Clutia, Pogonophora), the basic pattern of wood subfamilies and are basically different in morpho-
structure is similar in Acalyphoideae, Crotonoideae logy: Cnidoscolus stinging hairs are of the Urtica
and Euphorbieae. Scalariforn vessel perforation type—hollow, with thin walls and deciduous tip
plates are lacking; intervascular and vessel/ray (Thurston and Lersten 1969); those of the Plukene-
pitting is medium to very large; axial parenchyma tieae represent a unique type, with a central crystal-
is apotracheal diffuse or banded; rays are numerous, liferous cells forming a kind of hypodermic syringe
narrow, heterocellular, and often vertically fused; (Thurston 1976).
fibers are non-septate, often wide and thin-walled Detailed analyses of foliar vention patterns
(Mennega 1987). and leaf structure in Phyllanthoideae and Old-
Laticifers appear to be completely absent in fieldioideae have been presented by Levin
Phyllanthoideae and Oldfieldioideae but are (1986a, b) and Hayden (1994). Accordingly, sto-
well-developed in subfamilies Crotonoideae and mata are paracytic, brachyparacytic, or anisocytic
Euphorbioideae (Rudall 1987). Most laticifers are in Phyllanthoideae, and brachyparacytic
unicellular (non-articulated) and produce whitish (derived) in Oldfieldioideae, with exception of
latex, but colored (yellow or red) latex occurs in the anomocytic Podocalyx (and paracytic Para-
drypetes). The occurrence of tanniniferous cells
some Crotonoideae. In the two basal tribes of
in the adaxial leaf epidermis has been confirmed
Crotonoideae (Micrandreae and Manihoteae),
for supertribe Antidesmodae. An atlas of leaf
the laticifers are mostly multicellular (arti- venation in all the subfamilies has been published
culated), with mainly whitish latex. According to by Klucking (1997, 2003).
Rudall (1994), the multicellular laticifers of Hevea Foliar glands are rare in the Phyllanthaceae,
and Manihot may have evolved from the non- but common in the uniovulate subfamilies. Lam-
articulated type, and the articulated laticifers inar (embedded) glands appear to be usually
reported in Jatropha (Dehgan and Craig 1978) derived independently of those at the leaf base
may have evolved independently. The latex in or apex of petiole (Bernhard 1966). Inflorescence
tribe Euphorbieae is unusual in containing dis- bracts in the Crotonoideae and especially the
tinctively shaped (rod- or dumbbell-like) starch Euphorbioideae often have glands that appear
grains; the distribution of these starch grain types homologous to foliar glands, although morpho-
is considered to be of taxonomic significance logically divergent. The glands of the cyathium of
(Mahlberg 1975). Latex of Euphorbieae is often Euphorbia are anatomically similar to and appear
toxic, due to triterpenes and diterpenes. Foliar to be homologous with those of the thyrses of the
sclereids have been detected in various Acaly- Hippomaneae. Although the function of foliar
54 G.L. Webster
glands may be to attract ants that can deter her- Uhlarz (1982) and Hoppe (1985) studied the
bivores, bracteal glands of genera such as Mabea morphogenesis of several species of Euphorbia
appear to be a major source of nectar to attract and interpreted their findings within the classical
pollinators (Steiner 1981, 1983). morphological framework. Prenner and Rudall
(2007) and Prenner et al. (2008), who included in
I N F L O R E S C E N C E S T R U C T U R E . The basic inflore- their study a broader sampling of genera of the
scence pattern in Euphorbiaceae is cymose, as in Euphorbieae such as Anthostema, Dichostemma,
many angiosperm families. The prevailingly cymose Neoguillauminia, Calycopeplus, and Euphorbia,
nature of the Euphorbiaceous inflorescence is most suggested several scenarios for a derivation of the
explicit in the dichasial inflorescences of genera cyathium; their findings imply that the nectari-
such as Cnidoscolus (Bawa et al. 1982) and Jatropha ferous appendages may not all be of stipulate deri-
(Dehgan and Webster 1979), where a terminal pis- vation, as previously believed.
tillate flower is subtended by lateral staminate axes.
These inflorescences are usually protogynous; the F L O R A L S T R U C T U R E . Since the classic essays on
spatial and temporal separation of the staminate floral structure by Baillon (1858) and Eichler
and pistillate flowers have the effect of limiting the (1875), there have been few comprehensive
degree of geitonogamy in pollination. In surveys besides that of Michaelis (1924). Herma-
many genera, the dichasia are reduced to axillary phroditic flowers are rare and sporadic (e.g., in
glomerules, and separation of the sexes is com- Jatropha, and less commonly in Croton and Phyl-
monly achieved by dioecy or by spatial separation. lanthus), and are not consistently present in any
A common modification is a spicate arrangement of single species. The basic floral pattern is penta-
cymules (the thyrse). Paniculate and thyrsopanicu- merous, as found in the Phyllanthoideae tribe
late inflorescences, arising from elaboration of Wielandieae (e.g., in Heywoodia, Wielandia).
dichasia or aggregation of thyrses, appear to repre- The presumably plesiomorphic and apomorphic
sent an end-point in elaboration. Primitively, each states of some of the salient floral characters have
flower in an inflorescence is bracteate. Bracts may been tabulated by Webster (1994a). In virtually
be deciduous or persistent, and are sometimes dis- all genera, the flowers are hypogynous. Whereas
tinctively colored, especially in aggregations of more than half of the genera of the Phyllanthoi-
flowers into pseudanthia. deae have well-developed petals in at least one of
Pseudanthia have arisen in all of the subfamilies the sexes, nearly all members of the Oldfieldioi-
except for the Cheilosoideae and Crotonoideae. deae are apetalous. The sepals in some apetalous
In Uapaca (Phyllanthoideae), the pedunculate genera may be distinctly petaloid (e.g., in Cnidos-
pseudanthia with 5–10 imbricate colored bracts colus, Manihot, and Phyllanthus). Aestivation of
resemble single flowers (the pistillate flowers are the floral parts, especially the sepals, has been
solitary). In Peroideae, the flowers of Pera form used as a diagnostic character; sepals are imbri-
involucrate capitula enclosed in involucres of 1–2 cate in most of the Pyllanthoideae, whereas in the
smaller outer and two larger inner valvate bracts. majority of genera of Acalyphoideae the stami-
Most species of Dalechampia (Acalyphoideae) have nate sepals, at least, are valvate. In pistillate flow-
spectacular bisexual, bilaterally symmetrical pseu- ers petals are present in some genera of all the
danthia in which the paired bracts may vary greatly subfamilies except the Euphorbioideae; they are
in size, color, and lobing; the pistillate dichasium of usually inconspicuous but can be distinctly
three flowers is ventral to a staminate dichasium of showy in some Crotonoideae (e.g., Aleurites,
mostly 4–21 flowers associated with a resiniferous Jatropha). Except for the Euphorbioideae, a floral
gland derived from modified bractlets (Webster disk is generally present although sometimes
and Armbruster 1991). Finally, in the Euphorbioi- reduced in pistillate flowers. In the majority of
deae an entire tribe, the Euphorbieae, is character- genera, the androecium includes 2–20 stamens,
ized by the pseudanthial cyathium, which closely but the stamen number is reduced to 1 in Dalem-
resembles a bisexual flower and traditionally is bertia and the Euphorbieae, and in some Acaly-
interpreted as an inflorescence with a single termi- phoideae and Crotonoideae it has proliferated to
nal pistillate flower surrounded by clusters of 50–100 stamens or more. In many instances, the
highly reduced staminate flowers. Hoppe and filaments of the stamens are united into a column,
Euphorbiaceae 55
and sometimes (e.g., in Phyllanthus and Ompha- Punt (1962), using light microscopy, produced a
lea) the anthers are connate as well. This may survey of pollen in the family and established
also apply to Androstachys and Stachyandra many palynological "types" that were more or less
(Oldfieldioideae), the stamens of which have well correlated with phylogenetic position. K€ohler
been interpreted as spirally inserted; for details, (1965), also using LM, proposed a number of taxo-
see under the generic treatment (p. 93). M€ uller nomic realignments, notably a new subfamily, the
(1866) used anther dehiscence (introrse or Oldfieldioideae. Detailed palynological surveys
extrorse) as a diagnostic character, and it is still using SEM and TEM have been made for the Old-
a useful indicator of affinity in many cases. Some- fieldioideae (Levin and Simpson 1994), the Croto-
times, there is a rudimentary gynoecium (pistil- noideae (Nowicke 1994), the Peroideae, the
lode) in the center of the staminate flower. The Cheilosoideae, and the Acalyphoideae (Nowicke
perianth of pistillate flowers is often meristically et al. 1998, 1999, Takahashi et al. 2000; Nowicke
the same as in staminate flowers, but the sepals and Takahashi 2002). Although no complete atlas
often become modified during fruit development of pollen in Phyllanthoideae and Euphorbioideae
to enlargement (accrescence) in some cases, but has been published, there have been a number of
in other cases the sepals are deciduous in fruit. studies of infrasubfamilial scope. A detailed survey
The Euphorbiaceous gynoecium is generally of pollen morphology in the tribe Plukenetieae
described as hypogynous and 3-carpellate. How- has been provided by Gillespie (1994). The genus
ever, carpel number may vary from 2 (sporadi- Phyllanthus and its satellites display the most
cally) to 5 or 10, and occasionally the gynoecium extensive variation of exine sculpturing as well as
may be unilocular or 20-locular (Hura). Stylodia aperture structure and number, which has led to
vary greatly in configuration, but most commonly elevated numbers of colpi (often diploporate) and
are bifid, and often are connate at least at the even to pantoporate grains (Meewis and Punt 1983;
base; in various taxa they are unbranched (e.g., Punt 1987; Webster and Carpenter 2002; Sagun and
Alchornea, Hippomaneae) or may be repeatedly van der Ham 2003).
bifurcate (e.g., in Croton). Placentation is axile; Pollen features have been used as diagnostic
Berg (1975) is of the opinion that the Euphorbia- characters to distinguish the seven subfamilies
ceae show a variation of the axile structure that is recognized here. The putatively plesiomorphic
linked to the explosive capsular fruit. Each locule states are found in the Phyllanthoideae (tribes
contains one or two ovules, which are anatropous Wielandieae, Phyllantheae): prolate 3-colporate
and epitropous (i.e., with raphe adaxial) and cras- grains with reticulate (semitectate) exine; trans-
sinucellate; the nucellus is extended and may mission electron microscopy has revealed a per-
protrude as a nucellar beak far out of the micro- forate or discontinuous ektexine with distinct
pyle. There is usually an obturator projecting columellae and a continuous homogeneous end-
from the placenta over the micropylar region of exine (Levin and Simpson 1994). In subfam. Old-
the ovule. Epitropous ovules are common fieldioideae the grains are mostly oblate-globose
among the Malpighiales, and obturators are not and the apertures are contracted, grading from
restricted to the Euphorbiaceae, but nucellar brevicolpate to porate and from zonoporate to
beaks appear to be very rare elsewhere (only in pantoporate; basal taxa have at least four aper-
Malpighiaceae, Polygonaceae). However, the tures, but in the pantoporate condition their
combination of epitropous ovules, obturators and number can reach 40; the exine is modified to
extended nucellar beaks has not been reported for radially elongate, tapering spines that are conti-
any other angiosperm family, and therefore pro- nuous with the tectum. The Peroideae are quite
vides a strong morphological argument for the diversified palynologically and it is not easy to
monophyly of the Euphorbiaceae s.l. (Sutter and recognize a common pattern, whereas the Chei-
Endress 1995; Merino Sutter et al. 2006). losoideae are characterized by their echinate tec-
tum. In Acalyphoideae the pollen is invariably
P O L L E N M O R P H O L O G Y . The taxonomic signifi- tricolporate but exhibits a great diversity in struc-
cance of pollen morphology in the Euphorbiaceae, ture and sculpture. Although in Crotonoideae
first demonstrated by Erdtman (1952), has stimu- colporate, colpate and porate pollen is present,
lated investigations by a number of later workers. most members of the group have inaperturate
56 G.L. Webster
grains with an exine sculpture of triangular that recorded for some woody Hawaiian species
(rounded) structures in continuous arrays— of Chamaesyce by Koutnik (1987). The absence of
now widely known as "Croton pattern". Euphor- self-incompatibility is also well documented in
bioideae have 3-colporate pollen; among them, Mercurilis annua. Both hermaphroditic and
tribe Euphorbieae appears to have strange con- androdioecious populations are reported, and
spicuous intine bands bordering the colpi both exhibit mixed mating (i.e., a mix of self-
(Suárez-Cervera et al. 2001). fertilization and outcrossing; Korbecka et al.
2011). Self-compatibility is also the rule in Dale-
K A R Y O L O G Y . It is apparent from the excellent champia (e.g., Sazima et al. 1985).
survey of Hans (1973), and later additions by It seems likely that primitive Euphorbiaceae
Urbatsch et al. (1975) and others, that the were dioecious with binucleate pollen (Webster
Euphorbiaceae are quite diversified in chromo- and Rupert 1973); it is possible that the near-
some number. There is no single base number absence of self-incompatibility in the family is a
from which the chromosomal variation can be consequence of the evolution of monoecious taxa
derived, although Hans (1973) thought x ¼ 11 from dioecious ancestors. Only 19 of the 300
to be the base number both in the Acalyphoideae genera of Euphorbiaceae include both mono-
and Crotonoideae. However, as 9 and 10 are ecious and dioecious species, and the number of
nearly equally frequent base numbers, the ques- monoecious (131) and dioecious (142) genera is
tion remains whether these numbers have an nearly equal (Webster 1994a). The Oldfield-
aneuploid connection and in which direction ioideae are mostly dioecious and the Euphorbio-
this took place. In Euphorbia s. l., where very ideae are mostly monoecious, but both types are
many counts are available, x ¼ 11 is rare. It well represented in the other subfamilies. As pre-
seems that here descending aneuploidy has led dicted by Bawa (1980) and others, taxa with
to secondary base numbers 10, 9, 8, 7, 6, which ornithochorous dispersal are mainly dioecious,
were the starting points for effective polyploid and myrmecochorous taxa are somewhat more
speciation. The Synadenium clade (x ¼ 18) and likely to be monoecious. Taxa with binucleate
the Monadenium clade (x ¼ 19, 18, 17, 16, and 12) pollen grains may be monoecious or dioecious,
apparently represent a mostly decreasing aneu- but it appears that trinucleate grains are pro-
ploid series from an ancestral base number of x ¼ duced only in monoecious species.
9 (Jones and Smith 1969). A similar descending Prevention of autogamy in the self-compatible
aneuploid series can be found elsewhere in monoecious taxa is generally facilitated by spatial
Euphorbia, beginning with a base number of x separation of the sexes, often in separate inflore-
¼ 10 (Hans 1973). The few chromosome numbers scences (e.g., in Acalypha), and/or by dichogamy
known from Hippomaneae are based on 11, 9, (usually protogyny). In the inflorescences of a num-
and 7. According to the survey of Hans (1973) ber of genera (notably Cnidoscolus, Jatropha, and
polyploidy is more frequent in the family than in Euphorbia), the pistillate flowers terminate the
other woody plant families. Bischofia javanica (2n main axis or lower dichotomies, and open before
¼ 198, 28-ploid if based on x ¼ 7), systematically the staminate flowers (Bawa et al. 1982). In Euphor-
and geographically isolated, seems to represent bia, which has bisexual pseudanthia, monoecious
the final stage of development in a polyploid species are self-compatible, but in some species
complex, i.e., a "relictual polyploid". there is functional andromonoecy, with some
early-produced cyathia lacking functional pistillate
R E P R O D U C T I V E S Y S T E M S . In Euphorbiaceae, flowers (Narbona et al. 2002). The staminate cyathia
with open-pollinated strictly unisexual flowers, are proximal in the inflorescence and open earlier
self-incompatibility is rare, and earlier reports than the cyathia with pistillate flowers, so there is
of it appear to be incorrect. Self-incompatibility effective protandry, in contrast to the usual proto-
has been shown to be absent or incomplete in gyny and proximal position of pistillate flowers. As
Chamaesyce (herbaceous species, Ehrenfeld noted by Narbona et al. (2002), andromonoecy
1976), Hevea (Bouharmont 1962), and Manihot appears to be more frequent in perennial species,
(Jennings 1963; George and Shifriss 1967). A which are known to have a higher percentage of
documented instance of self-incompatibility is outcrossing. In Bridelia tomentosa, a rarely
Euphorbiaceae 57
documented (but possibly frequent, see Sapinda- tion by a variety of pollinator species seeking
ceae in Vol. X of this book series) temporal mating nectar, particularly those belonging to the insect
system has been analyzed (Luo et al. 2007), in which orders Hymenoptera and Diptera. These plants
each individual produces subsequently staminate, include Croton (Crotoneae), Jatropha (Jatro-
then pistillate, and then staminate flowers, with the pheae), and Tragia (Plukenetieae; see review in
resting periods between the flowering phases pre- Webster 1994). Specialized pollination (by few
cluding selfing, and asynchrony among individuals pollinator species or functional groups) is seen
ensuring mating partners (duodichogamy). in some species of Macaranga (Acalypheae; by
Although polyploidy is common in large thrips, Moog et al. 2002), Breynia, Glochidion,
genera such as Croton, Euphorbia, and Phyll- and some Phyllanthus (Phyllantheae; by moths,
anthus, published records of interspecific hybrid- Kawakita 2010). These four cases represent the
ization are rare (Webster 1967). Polyploidy is recent discovery of the brood-reward systems in
associated with apomixis in some species of the Euphorbiaceae. Here the pollinators lay eggs
Euphorbia (Cesca 1961). on or near the flowers and the larvae feed on
floral tissues, much as do fig wasps on figs and
P O L L I N A T I O N . Consistent with their floral and yucca moths on yuccas. Moderately specialized
inflorescence diversity, Euphorbiaceae show a lepidopteran pollination has been reported in
great diversity of pollination systems and polli- Cnidoscolus (Manihoteae; by sphingid moths,
nator species. Pollination occurs abiotically via butterflies, and even hummingbirds, depending
wind in Acalypha, some Macaranga and Mallotus on species). Moderately specialized pollination
species, Mercurialis (Daumann 1972; Korbecka by mammals has been reported in Central Amer-
et al. 2011), Ricinus (Alex 1957), and probably ican Mabea occidentalis (Hippomaneae; by bats,
many other Acalypheae. Wind pollination has marsupials, and the woolly opossum, Steiner
also been reported in Bernardia (Bernardieae; 1983) and in Central American Hura (Hureae;
Bullock 1994), Bertya (Ricinocarpeae; Fatemi by bats, Steiner 1982). These last two cases are
and Gross 2009), "Celaenodendron" (¼ Piranhea, somewhat intermediate between individual-
Oldfieldioideae; Bullock 1994), some Croton spe- flower and pseudanthial functioning in that
cies (Crotoneae; Bullock 1994) and possibly in mammal pollinators respond to clusters of stami-
two or more species of Phyllanthus (Subba- nate flowers, although only one or a few pistillate
Reddi and Reddi 1984). Anemophilous taxa flowers are involved; the less important insect
characteristically have a reduced or lacking peri- pollinators of Mabea presumably respond to
anth and nectary disk combined with a multipli- individual flowers (Steiner 1983). Incidentally,
cation of stamens, catkin-like inflorescences, and the primarily bat-pollinated Mabea fistulifera in
smaller pollen grains than their zoophilous rela- disturbed semi-deciduous forest and woodland of
tives; Mallotus may be transitional in this respect southeastern Brazil has become the meeting point
(Lock and Hall 1982). The extraordinarily elabo- of many forest animals, including more than 30
rate androecium of Ricinus, with branching fila- bird species many of which are frugiverous, two
ments and hundreds of anthers, doubtlessly ceboid primates, four frugivorous bat species, the
reflects adaptation to anemophilous pollination, common opossum, and the coati (Nasua nasua),
and is not a primitive character, as claimed by the majority of which probably can act as polli-
Venkata Rao and Ramalakshmi (1968). nators (Torres de Assumpção 1981; Vieira and Car-
Species with biotic pollination can be divided valho-Okano 1996; Olmos and Boulhosa 2000). The
into two groups: those in which unisexual flowers unspecialized reproductive system of this tree allows
are visited individually and those in which the its continued existence in biotically impoverished
flowers operate pseudanthially, i.e., the unisexual habitats, and at the same time maintains a signifi-
flowers are united into generally bisexual blos- cant part of the pollinator and frugivore community
soms and pollinators visit several flowers (usually during a period when fruits are scarce or not avail-
of both sexes) simultaneously (with simultaneous able.
or sequential receptivity). Most euphorbs in the Euphorbs with fully pseudanthial inflores-
former category (visitation of individual flowers) cences have either "normal" unisexual flowers
probably have ecologically generalized pollina- (Dalechampia, Pera) or highly reduced unisexual
58 G.L. Webster
flowers united in a cyathium (Euphorbia and The seed coat in Euphorbiaceaous seeds,
relatives in Euphorbieae). In the latter case the investigated by Netolitzky (1926), Wunderlich
staminate flowers are reduced to a single stamen (1968), Corner (1976), Stuppy (1996), and
and the pistillate flowers to a single pistil, with Tokuoka and Tobe (1995, 1998, 2001, 2002,
perianth absent in both cases. In both situations, 2003), usually consists of two primary layers:
pollinators visit the clusters of flowers as if they the testa, derived from the outer integument of
were a single flower—hence, application of the the ovule, and the tegmen, derived from the outer
term pseudanthium (false flower) or blossom. layer(s) of the inner integument. The testa usually
Pollination of euphorbs with pseudanthial inflor- has few cell layers, except in seeds with a sarco-
escences is often highly generalized. Pollination testa, which may have 10 cell layers or more
by several to many types of insects occurs in (Stuppy 1996). In indehiscent fruits, however,
some species of Malagasy Dalechampia (Pluke- the testa may be thin or even rudimentary. Cor-
netieae; Armbruster and Baldwin 1998), in South ner (1976) recognized that the variable structure
American Pera (Pereae; Freitas et al. 2011), and of the sclerified cells derived from the outer layer
many or possibly most species of Euphorbia. of the inner integument (exotegmen) furnished
Other pseudanthiate euphorbs exhibit highly important clues to relationships between genera.
specialized pollination. Most neotropical Dale- In biovulate euphorbs, the exotegmen is often
champia are pollinated by females of one or a fibrous, but the studies of Stuppy (1996) and
few species of resin-collecting bees (Armbruster Tokuoka and Tobe (2001) have revealed such a
1984, 1988). The rewards are oxygenated triter- great variation in its structure that these authors
pene resins secreted by modified bractlets borne even considered the exclusion of the aberrant
by the staminate subinflorescence (Armbruster genera Oldfieldia, Picrodendron, Hymenocardia
et al. 1997). Specialized pollination by fragrance- and Didymocistus from the family. Among the
collecting, male euglossine bees has evolved uniovulate subfamilies, Peroideae (except for the
independently from resin-reward ancestors at problematic Pogonophora) have a peculiar seed
least three times (Armbruster 1993; Armbruster coat consisting of only two persistent cell layers,
et al. 2009). The fragrance rewards are secreted by the well-developed exotesta of radially elongated
the stigmatic surface in two of these clades, and by a cells and the more weakly developed tracheoidal
modified "resin gland" in one clade (Armbruster exotegmen. All remaining uniovulate Euphorbia-
1993). Specialized pollination by hummingbirds ceae have the palisadal exotegmen composed of
is seen in the Pedilanthus clade of Euphorbia; radially elongated cells. Carunculate seeds are
Cacho et al. 2010). lacking in the Phyllanthoideae but occur, though
not consistently, in all other subfamilies. The
F R U I T A N D S E E D . The typical Euphorbiaceous caruncle can function as an elaiosome and, after
fruit is a 3-locular capsule (schizocarp) that the ballistic or barochorous ejection of the seeds,
dehisces, often explosively, into three 1- or 2- attracts ants that are implicated in a further dis-
seeded units (cocci or mericarps). The seeds are persal event (Berg 1975). Bianchini and Pacini
released when the cocci separate septicidally and (1996) and Lisci et al. (1996) point out that the
sometimes loculicidally as well; in some genera caruncle may also have functions other than
the cocci remain more or less closed. The axis of mymecochory, such as facilitation or inhibition
the fruit, including the three placentae, persists as of germination.
a columella, which may be massive or appen- In Euphorbiaceous seeds, starch as a storage
daged (e.g., in Stillingia), but is sometimes deci- product is replaced by protein and fat oil.
duous. However, in a sizeable number of genera
scattered through all the subfamilies, the fruit D I S P E R S A L . The typical Euphorbiaceous fruit
remains indehiscent and the carpellar walls (upon drying) dehisces explosively into three 1-
become fleshy, sometimes with a sclerified endo- or 2-seeded merocarps (cocci). This kind of auto-
carp. One-seeded fruits are almost always inde- chorous dispersal can be quite effective in genera
hiscent. Somewhat counter-intuitively, Stuppy such as Hevea or Hura, whose seeds may be
(1996) recorded in biovulate genera only 19 dispersed as far as 43 m (Swain and Beer 1976).
(28%) with 2 seeds per locule in fruit. However, in many genera the seed may produce
appendages or a sarcotesta, and the primary
Euphorbiaceae 59
autochorous dispersal is followed or superseded P H Y T O C H E M I S T R Y . Euphorbiaceae are one of

by animal dispersal. Seeds with fleshy integumen- the chemically most diverse angiosperm families
tal elaiosomes (caruncles) are dispersed by ants. (Hegnauer 1966, 1989; Seigler 1994). Many taxa
In Australia, myrmecochory is especially promi- produce toxic or medicinally important com-
nent, perhaps reflecting the widespread open and pounds (e.g., ricinin in the seed coat and castor
xeromorphic vegetation (Berg 1975). As reported oil in the endosperm of Ricinus seeds); in the
by Clifford and Monteith (1989), seed dispersal of tropics, they have acquired a wide use for many
Australian Petalostigma pubescens implies three purposes such as for arrow poisons (Sapium) or
distinct phases, beginning with the ingestion and as fish poisons (Phyllanthus). A number of genera
transport of the drupes by migrating emus. (Cnidoscolus, Dalechampia, Tragia) are impli-
Voided endocarps, freed from the fleshy epicarp, cated in dermatitis. The sap of several Euphor-
upon exposure to the sun dehisce explosively and bioideae (Euphorbia subg. Euphorbia, Hippomane,
project the seeds up to 2.5 m from the point of Ophthalmoblapton, Synadenium, Excoecaria agal-
origin, and the seeds can be carried off by ants locha ¼ "blind-your-eye") is dangerously toxic.
feeding on the conspicuous elaiosomes. In Cro- Among the more significant secondary meta-
ton, where myrmecochory is a very widespread bolites produced by Euphorbiaceae are tannins,
mode of dispersal, the typical caruncles have been alkaloids, cyanogenic glycosides, diterpenes,
lost in sect. Cuneati with riverine species such as glucosinolated lipids, and triterpenes. Both
C. cuneatus and C. yavitensis showing an arillate condensed and hydrolysable tannins are present
structure possibly adapted to attract fish (Riina in the family, mostly in woody species. The ellagi-
et al. 2010). Dispersal of diaspores by fish during tannin geraniin has been found in all subfamilies,
their upstream-directed spawning migrations and in the uniovulate ones it is often accompa-
is a frequent phenomenon in the periodically nied by the similar mallotusinic acid. Alkaloids
overflooded river plains of Amazonia, where are particularly diverse in Phyllanthoideae, Cro-
large fish such as tambaqui (Colossoma macro- tonoideae (especially Aleurites, Croton, Jatropha)
pomum) and jatuarana (Brycon sp.) show strong and in some Acalyphoideae (Alchornea, Mallotus,
preferences for the fruits of Hevea spruceana and Ricinus), but there is no structural type of alka-
H. brasiliensis (Goulding 1980) but have also been loids characteristic for the family. Cyanogenic
found to disperse the propagules of Piranhea glycosides are widespread in Phyllanthoideae
trifoliata and Mabea sp. (Andrachne, Bridelia, Flueggea, Phyllanthus)
Ornithochory seems to be confined to the and Crotonoideae (Aleurites, Croton, Elateio-
tropics, especially in Africa and Asia, but is less spermum, Hevea, Manihot), but have also been
common in the Oldfieldioideae and Euphorbio- reported from Acalyphoideae (Acalypha, Mercur-
ideae (Webster 1994a). Seeds with fleshy coats ialis, Ricinus) and several Euphorbioideae. A con-
(sarcotesta), found in all five subfamilies, are siderable variety of diterpenes occur in
generally dispersed by birds. In Baccaurea, for Euphorbiaceae. Cyclic diterpenes are especially
example, which has non-explosive fruits, some prominent in Crotonoideae (Bertya, Croton,
species have orange arils and are dispersed by Jatropha). Tetracyclic diterpenes isolated from
birds, whereas other species have berries with species of Croton have been implicated as co-
sweet pulp and are dispersed by rats, fruit bats, carcinogens. Triterpenes are common in many
or squirrels (Ridley 1930: 427). The fruits of the Euphorbiaceae, especially the Euphorbioideae.
shade-intolerant riverine tree Trewia nudiflora In Euphorbia, investigations of triterpene compo-
(¼ Mallotus) are large, hard, and dull-colored sition have produced suggestive evidence for de-
upon ripening, unattractive to the usual seed dis- limiting subgenera and sections (Ponsinet and
persers such as monkeys, bats, and birds, but Ourisson 1965). Terpene-rich latex in many
regularly eaten by the greater one-horned Asian Euphorbiaceae is also accompanied by rubber
rhinoceros (Rh. unicormis). Seeds deposited into (polyisoprenes). Although Hevea is the best
the animals’ dung on open grassland develop into known source of rubber, it is also produced by
robust, fast-growing seedlings, providing an other Crotonoideae such as Cnidoscolus, Mani-
interesting example of megafaunal dispersal hot, and Micrandra, as well as Chamaesyce and
(Dinerstein and Wemmer 1988). Euphorbia (Rizk 1987).
60 G.L. Webster
P A L E O B O T A N Y . The Euphorbiaceae, despite DISTRIBUTION AND H A B I T A T S . Although

their contemporary prominence and broad distri- Euphorbiaceae are a pantropical family, they also
bution, do not have an impressive fossil record. extend into subtropical and warm temperate
Fossil woods, such as the much-cited Paraphyl- regions, and some herbaceous species of Euphorbia
lanthoxylon from the Cretaceous of North America occur in subpolar and alpine areas. Generic diver-
(Bailey 1924; Cahoon 1972), cannot be convin- sity for Phyllanthoideae is greatest in Africa, for
cingly verified as Euphorbiaceous. M€adel (1962), Oldfieldioideae in Australasia, and for Acalypho-
in a study of Cretaceous fossils from South Africa, ideae and Euphorbioideae in Asia; diversity is
plausibly suggested that Paraphyllanthoxylon approximately equal in American and Asiatic Cro-
capense is similar to Margaritaria discoidea. tonoideae (Webster 1994a). In total generic diver-
M€adel’s Securinegoxylon biseriatum can perhaps sity, American leads with 111 genera, followed by
be compared with Heywoodia lucens, which grows Asia with 107, Africa with 102, and Australasia with
in South Africa today. 77. Madagascar, with 51 genera (17 endemic), is
There is greater certainty in the relationships the most distinctive subregion.
of the Eocene fossils of fruits from America and Among the Phyllanthoideae there are
Europe that have been referred to the Hippo- genera—mostly smaller ones—that are restricted
maneae by Mazer and Tiffney (1982) and Dilcher to Africa/Madagascar, such as Wielandia, Lachno-
and Manchester (1988). Both Wetherellia and stylis/Securinega, and Uapaca. Some large genera
Palaeowetherellia, in fruit morphology and in such as the Meineckia, Flueggea, and Margaritaria
the depositional facies in which they occur, can have a pantropical distribution that includes the
be associated with the living genus Hippomane; African/Malagasy region. There are remarkable
Dilcher and Manchester consider Crepetocarpon disjunctions between closely related genera, such
to be unquestionably close to Hippomane. as between the African Heywoodia and the neo-
Fossil flowers of Euphorbiaceae are little tropical Chascotheca + Astrocasia. The closest
known, but the Eocene Hippomaneoidea of relatives of the strictly African/Malagasy Wielan-
Crepet and Daglian (1982) is well-preserved, and dia are the east Asian Dicoelia + Chorisandrachne,
the pollen suggests a relationship to Senefeldera, a and African Hymenocardia is close to neotropical
neotropical genus considerably less specialized Didymocistus. This highly disjunct distribution of
than Hippomane. the tribes and genera is most likely the result of
Dispersed fossil pollen can be hazardous to long-distance dispersal, as has been suggested for
identify but, according to Muller (1984), genera Bridelia, which very likely dispersed from tropical
resembling the Oldfieldioideae, Acalyphoideae, Africa to Asia and Australia once or twice from 10
and Crotonoideae are known from the early Ter- million years ago onward (Li et al. 2009).
tiary. A notable record of Euphorbiaceous pollen In the Oldfieldioideae, which are best devel-
from the Eocene in France (Gruas-Cavagnetto oped in Australasia, much more compact distri-
and K€ ohler 1992) includes a wide range of taxa: bution patterns of the (sub)tribes can be
Actephila, Antidesma, Bischofia, Leptopus, and recognized. The whole group is highly disjunct,
Phyllanthus (Phyllanthoideae); Austrobuxus and with the tribally isolated neotropical Podocalyx
Dissiliaria (Oldfieldioideae); Alchornea and Rici- and tribe Picrodendreae disjunct between the
nus (Acalyphoideae). In Australia, Oldfield- New World and Africa/Madagascar. The genera
ioideae appear in the Paleocene and, in the early of Mischodontinae, with exception of the Ceylo-
Eocene, in greater diversity (Martin 1974, 1978). nese Mischanthus, all are Madagascan. Tribe Cale-
Because the pollen of most Phyllanthoideae is tieae is a group of 14 genera centered in Australia
morphologically unspecialized, it may be difficult with a paleontological record dating back there
to follow the record of fossil pollen of the Euphor- into the Paleocene (Martin 1974, 1978).
biaceae back into the Cretaceous. Taking the Peroideae are utterly disjunct, and Cheiloso-
available evidence into account, however, it ideae restricted to the Oriental region. Acalypho-
seems reasonable to surmise that the Phyllantho- ideae are widely distributed in the neo- and
ideae and Oldfieldioideae, at least, had differen- paleotropical region including Australia, and
tiated by the upper Cretaceous and that the only few and smaller tribes are restricted to a
radiation of the subfamilies had occurred by the single continent, such as neotropical Adelieae,
Paleocene. Asian Erismantheae, and Australian Ampereeae;
Euphorbiaceae 61
the large tribes Acalypheae and Plukenetieae are Ericoid life forms are common in Australia,
pantropical. Crotonoideae are well represented in especially in Bertya, Beyeria, Micrantheum, and
the New World, with Crotoneae, Micrandreae, Pseudanthus. Ultramafic (serpentinic) outcrops,
etc., or in the Old World with inclusion of the especially in Cuba and New Caledonia, have
Australasian region (Codieae, Ricinocarpeae), housed a spectacular radiation of sclerophyll spe-
but have a weak representation in Africa. In cies of Phyllanthus. Sand dunes in subtropical
Euphorbiodeae the four genera of Stomatocaly- arid regions support psammophile species of
ceae, two of them monotypes, present a highly Chamaesyce, Chrozophora, and Croton.
disjunct picture, with one genus in the New
World, two in Africa, and one in Australasia. A CIRCUMSCRIPTION AND SUBDIVISION OF THE
similar trend is evident in the equally relictual F A M I L Y . The circumscription of the Euphorbia-
basal subtribes of the Euphorbieae, the Antho- ceae had remained surprisingly stable over a long
stematinae and Neoguillauminiinae. Subtribe period of time (Webster 1987), whereas its infra-
Euphorbiinae, i.e., Euphorbia s. l., is sub- familial classification remained contentious
cosmopolitan, with the Rhizanthium clade (reviewed in Webster 1987, 1994a). Delimitation
including mostly southern African xerophytes, of infrafamilial taxa was long obfuscated by
the Esula clade with predominantly northern M€ uller’s unnatural binary division of the family
hemisphere herbaceous species, the Euphorbia into Platylobeae and Stenolobeae (M€ uller 1866),
clade representing the greatest diversity within which—despite its rejection by Bentham
the genus formed mainly by African/Madagascan (1880)—was adopted by Pax (1890) and Pax and
xerophytes, and the mainly New World Chamae- Hoffmann (1931). Bentham (1880) and later
syce clade. Hutchinson (1969) recognized only the tribal
Despite their diversity, Euphorbiaceae are level as the major infrafamilial rank, whereas
predominantly shrubs or short-lived trees Pax (1884, 1890) recognized two main subfami-
adapted to secondary succession in dry to mesic lies, Phyllanthoideae and Crotonoideae, for the
seasonal habitats. In savannas, tropical deciduous biovulate and uniovulate taxa, a critical distinc-
forests, and deserts, genera such as Chamaesyce, tion not grasped by M€ uller and Bentham but
Cnidoscolus, Croton, Euphorbia, Hymenocardia, accepted since then by most systematists. Never-
and Jatropha are well represented. However, theless, the heterogeneity within these two major
there are also a significant number of genera in infrafamilial groups prompted Webster (1975,
lowland tropical rainforest: in America, Alchor- 1994b) to propose a system of five subfamilies:
nea, Amanoa, Hevea, Hieronyma, Micrandra, and Phyllanthoideae, Oldfieldioideae, Acalyphoideae,
Richeria; in Africa, Maesobotrya, Spondianthus, Crotonoideae, and Euphorbioideae. This was
and other Phyllanthoideae; in Asia and Austral- generally adopted, notwithstanding the claims
asia, Antidesma, Aporosa, Baccaurea, Glochidion, by various authors to elevate superficially ano-
Macaranga, and Mallotus. Adaptation to diverse malous genera such as Hymenocardia, Uapaca,
habitats can be seen even within single genera. Bischofia, etc. to family rank—after closer inspec-
For example, while most Dalechampia are twin- tion, these were found to be untenable. In
ing vines or lianas adapted to secondary environ- the system of Webster (1994b), the number of
ments with very high light levels, one clade of 3–4 tribes was increased from 14 to 49, a number
species, sect. Cremophyllum, is a small mono- again slightly increased by Radcliffe-Smith
podial shrub largely restricted to the dark under- (2001).
storey of primary rainforests. In tropical regions, Corner’s (1976) studies in seed coat structure
some Euphorbiaceae have adopted specialist of the dicotyledons led him to question the homo-
roles. Along rivers, there are a number of rheo- geneity of the family, an opinion that received
phytes adapted to surviving strong currents and support from upcoming DNA sequence studies.
submergence, including species of Phyllanthus Molecular systematics, beginning with the work of
and Homonoia. Rooted aquatics are found in Wurdack (2002), led to a radical dismemberment
Caperonia and Phyllanthus; the only floating of the family into three families, the Phyllantha-
aquatic in the family is the Amazonian Phyl- ceae and Picrodendraceae (¼ Oldfieldioideae),
lanthus fluitans. both biovulate, and the uniovulate
62 G.L. Webster
Euphorbiaceae, apart from the hardly disputed is highly significant; the formation of cocci, sepa-
expulsion of some extraneous genera relegated to rating from a central column, is a characteristic of
the Putranjivaceae, Centroplacaceae, and Panda- the family and has been the basis of the earlier
ceae. More recently, and not unexpectedly, ordinal name, Tricoccae. The cocci and their xero-
increased molecular resolution of the deeper chastic sclerenchymatic endocarp tissue instru-
nodes of the Malpighiales tree has resolved Phyl- mental in the explosive dehiscence are the basic
lanthaceae and Picrodendraceae (Oldfieldioideae) elements of the explosive seed release characteristic
as strongly supported sisters (Tokuoka and Tobe for the family.
2006; Wurdack and Davis 2009; Korotkova et al. These structural traits support the close
2009; Soltis et al. 2011); this means that, for those relationship between the uni- and biovulate
who want to maintain different families for bio- euphorbs, and have led us to retain the broad
vulate and uniovulate euphorbs, there is no longer family concept of the family. Further molecular
any compelling necessity for distinguishing inquiry into the deeper nodes of the phylogeny
between Phyllanthaceae s. str. and Picrodendra- of the Malpighiales beyond the interesting findings
ceae. of Xi et al. (2012) is impatiently expected for a
Broad-based molecular phylogenetic analyses clarification of this issue.
of representatives of the traditional uniovulate
subfamilies Acalyphoideae, Crotonoideae, and E C O N O M I C I M P O R T A N C E . As a consequence of
Euphorbioideae have recovered the uniovulate their chemical diversity, the Euphorbiaceae
euphorbs as monophyletic (Wurdack et al. 2005; include a considerable number of economically
Tokuoka 2007; Xi et al. 2012), and have brought important species (Schultes 1987). The currently
to light novel groupings that largely correlate most valuable commodity is probably the starchy
with the distribution of traditional characters tubers of cassava (Manihot), widely cultivated in
such as latex and laticifers, pollen morphology, America and Africa. The Pará rubber tree
and ovular and seed coat characters. Two (Hevea), the most important Euphorbiaceous
strongly supported basal clades have been pro- crop in the 19th century, is still the major source
posed as additional subfamilies (Peroideae and of non-synthetic commercial rubber. Seed oils of
Cheilosoideae), whereas the rest of the family is tung (Aleurites) and castor bean (Ricinus) con-
divided into seven major lineages. Among them, tinue to be widely used, but in South America
the core Acalyphoids and the Euphorbioideae are there is now considerable interest in the seed oil
monophyletic groups, whereas the Crotonoideae of inchi (Caryodendron), which is considered
are resolved as several distinct and partly not superior to the oil of the African oil palm, Elais
well-supported clades. (Schultes 1987). Waxes from neotropical species
Strong molecular evidence supports the mono- of Sapium and Euphorbia (especially E. antisyphi-
phyly both of the biovulate and uniovulate Euphor- litica) are still a minor source of commercial
biaceae, whereas molecular proof for the waxes. Use of the latex of Euphorbia tirucalli
monophyly of the Euphorbiaceae s. l. is still lacking. plants as "gasoline trees", once vigorously advo-
The most recent analysis (Xi et al. 2012) places cated by Calvin (1987), appeared not competitive
the biovulate and uniovulate euphorbs, the latter with sugar cane.
together with Linaceae and Ixonanthaceae, in a As pointed out by Schultes (1987), many
weakly supported clade. There are, however, Euphorbiaceae have been cultivated in tropical
important similarities in embryological, gynoecial areas for thousands of years. Schultes observed
and fruit structures between Phyllanthaceae s. l. Amerinidians in the Amazon basin exploiting
and Euphorbiaceae s.str. First, the combination of Euphorbiaceae in a variety of ways, including
nucellar beaks, epitropous ovules, and obturators food, medicines, and piscicides. For example,
within the angiosperms is restricted to Euphorbia- the reddish latex of some neotropical species of
ceae s. l. (Sutter and Endress 1995; Merino Sutter Croton ("Sangre de Drago") is widely used in
et al. 2006; see above under FLORAL STRUCTURE). South America as a salve applied to cuts and
Second, the fruit morphology of the Euphorbiaceae sores. The alkaloids in Sangre de Drago latex
s. l., which has attracted the interest of botanists appear to be non-toxic to humans, but other
since the time of Baillon (1858; see also Berg 1975), species of Croton produce phorbol esters that
Euphorbiaceae 63
have been implicated in human cancer (King- itudinally, and that their petioles are not
horn 1979). Herbaceous species of Phyllanthus pulvinate;
have long been used in folk medicine in the – rarely occurring character states are given in paren-
tropics to treat kidney and urinary problems theses ( ), whereas square brackets [ ] are used in
and jaundice (Webster 2002), and pharmaceuti- taxonomic descriptions for comments and expla-
cal research has revealed many useful applica- nations.
tions of the antiviral compounds of Ph. amarus, Users of this treatment who want to make
which are of potential value in treating the hep- determinations of genera of Euphorbiaceae may
atitis B virus. use this multichoice key:
Because of their generally small inconspicuous Hoffmann, P., Kirkup, D., Galster, A.M.,
flowers, the Euphorbiaceae include relatively few Challen, G., Radcliffe-Smith, A. 2005. Key to the
ornamental plants, but species of a few genera are genera of Euphorbiaceae sensu lato. Available at
commonly cultivated in tropical gardens or green- www.kew.org/herbarium/keys/euphorbs
houses for their colorful inflorescences or foliage:
Acalypha, Codiaeum, Jatropha, Euphorbia. Succu- CONSPECTUS OF THE SUBFAMILIES OF EUPHOR-
lent taxa of Euphorbia subgenus Euphorbia—e.g., BIACEAE
sects. Monadenium, Pedilanthus, Synadenium— 1. Locules of ovary each with 2 ovules; milky latex, intraxy-
are popular in tropical and subtropical gardens. lary phloem, and stinging hairs 0; indumentum simple
Dalechampia aristolochiifolia Kunth is a recently (lepidote or dendritic); embedded foliar glands rare;
marketed horticultural vine increasingly cultivated pollen grains binucleate 2
in warm temperate regions for its colorful bracts – Locules of ovary each with 1 ovule; latex + or 0; indu-
(generally advertised on the internet incorrectly as mentum various; pollen grains binucleate or trinucle-
ate 3
D. dioscoreifolia). 2. Leaves alternate (opposite), stipulate, simple and
Some Euphorbiaceae have a distinctly nega- unlobed [except in Bischofia]; petals + or 0; pollen
tive economic value; the family includes many grains tricolporate to -porate, sexine lacking
taxa adapted for secondary succession, and spines [except in Amanoa and Croizatia]; seeds ecar-
hence there are a number of aggressive weeds. In unculate
I. Subfam. Phyllanthoideae (p. 64)
temperate regions, the most damaging invaders of
– Leaves alternate, opposite, or whorled, stipulate or esti-
cultivated fields, especially in North America, are pulate, simple to trifoliolate; petals 0; pollen grains
species of Euphorbia section Esula, especially colporoidate to porate, sexine spiny; seeds carunculate
E. esula and related species. Some weedy taxa— or ecarunculate
species of Euphorbia, Mercurialis, Phyllanthus— II. Subfam. Oldfieldioideae (p. 90)
3. Ovules with thin integuments [inner integument < 6
are particularly obnoxious because of their toxic-
cells thick]; seeds arillate; seed coat with tracheoidal
ity to livestock (Burrows and Tyrl 2001). Recently exotegmen
exotic woody Euphorbiaceae, including species III. Subfam. Peroideae (p. 102)
of Glochidion, have invaded the southern United – Ovules usually [Adenoclineae excepted] with thick
States. The most aggressive species is the Chinese integuments [inner integument > 8 cells thick]; seeds
tallow tree, Triadica sebifera (Sapium sebiferum), arillate or not; seed coat with palisadal exotegmen 4
4. Pollen exine echinate; flowers apetalous; seeds exaril-
which has invaded forests and even grasslands late IV. Subfam. Cheilosoideae (p. 105)
from Florida to Texas and has become established – Pollen exine not echinate; flowers petaliferous or apet-
in California. alous; seeds arillate or exarillate 5
5. Milky latex 0; laticifers, if present, non-articulate; pol-
SUBFAMILIES, TRIBES AND GENERA OF len grains binucleate
EUPHORBIACEAE V. Subfam. Acalyphoideae (p. 106)
In all generic descriptions given below, – Latex + (scanty or 0), milky or colored; laticifers articulate
or non-articulate; pollen grains binucleate or trinucleate 6
– for the sake of briefness it is generally 6. Latex reddish or yellowish to milky; laticifers articu-
assumed—if not stated otherwise—that the late or non-articulate; leaves unlobed to lobed or
compound; indumentum simple, stellate, dendritic,
plants have alternate simple leaves with entire or lepidote; bracts usually not biglandular at base;
margins, distinct stipules, lack foliar glands sepals imbricate to valvate, usually completely cover-
and domatia, have anthers dehiscing long- ing anthers in bud; petals mostly + [at least in
64 G.L. Webster
pistillate flowers]; pollen grains tricolporate or more and molecular evidence accrued since 1990 con-
often porate or inaperturate, with hexagonal sexinous firms Bischofia as a member of Phyllanthoideae.
processes
VI. Subfam. Crotonoideae (p.156)
On the other hand, Putranjivaceae have now been
– Latex whitish (yellow), often caustic or toxic; laticifers excluded definitely from the Euphorbiaceae on
non-articulate; leaves usually unlobed; indumentum the basis of chemical and molecular studies (e.g.,
simple or lepidote (dendritic in some Hippomaneae); Chase et al. 1993; Wurdack 2002), corroborated
bracts often biglandular at base; sepals imbricate to by Tokuoka and Tobe (1999) in the demonstra-
reduced or 0, anthers mostly not covered in bud; petals tion of an impressive number of embryological
0; pollen grains tricolporate, exine mostly perforate-
reticulate
differences between the reinstated Putranjivaceae
VII. Subfam. Euphorbioideae (p. 190) and Euphorbiaceae and the findings by Xi et al.
(2012), which place Putranjiva in a clade with
Lophopyxis.
I. SUBFAM. PHYLLANTHOIDEAE Beilschm. (1833). In the classification of Phyllanthoideae,
which largely follows the one given by Hoffmann
Phyllanthaceae Martinov (1820).
and Wurdack (2006), the group is subdivided
Monoecious or dioecious trees, shrubs, or herbs; into 10 tribes and 12 subtribes, which correspond
vessel elements with simple to scalariform per- with strongly supported clades of the multigene
forations; indumentum mostly simple. Leaves analysis of Kathriarachchi et al. (2005). In spite of
alternate (opposite), spiral to distichous, usually this, morphologically they are often utterly dispa-
stipulate, simple and entire, trifoliolate and den- rate and, in extreme cases, determination of spe-
tate in Bischofia, without embedded glands, sto- cies and genera is said to depend on molecular
mata mostly paracytic or anisocytic; petiole analyses.
eglandular. Inflorescences axillary (terminal), of
racemoid or spikelike thyrses or reduced to glo-
KEY TO THE TRIBES OF SUBFAM. PHYLLANTHOI-
merules or flowers solitary; bracts eglandular.
DEAE
Sepals mostly 4–6, imbricate (valvate), usually
basally connate; petals and disk + or 0; 1. Leaves simple; inflorescence mainly of glomerules or
stamens (2)3–8(60), filaments distinct or thyrses; flowers with or without petals; stylodia entire
to multifid; fruits capsular, baccate, or drupaceous 2
connate; anthers dehiscing horizontally to verti-
– Leaves palmately 3–5-foliolate; inflorescence panicu-
cally; pollen grains mostly 3–4(7)-colporate late or racemose; flowers lacking petals and disk; sty-
(porate), exine mostly semitectate (echinate); lodia entire; fruits drupaceous
male gametophyte binucleate; pistillode + or 0; 10. Bischofieae
ovary (1)2–5(20)-locular, ovules 2 in each loc- 2. Fruits winged or lobed; petals and disk 0; ovary locules
ule, anatropous or hemitropous, nucellar beak 2; pollen zonoporate
sometimes prominent; stylodia mostly distinct, 5. Antidesmateae-Hymenocardiinae
usually bifid, sometimes unlobed. Fruit mostly – Fruits not winged; inflorescences mostly not ament-
like [except in Antidesmeae]; stylodia entire to bifid
capsular, less commonly baccate or drupaceous; (multifid); floral disk + or 0; pollen rarely zonoporate
columella often persistent. Seeds ecarunculate 3
[except for Celianella]; endosperm + or 0; coty- 3. Stems with resinous exudate; inflorescences paniculate
ledons plane, incrassate, or folded. x ¼ 12 or 13 or capitulate 4
(8, 10 or 11). – Stems lacking resin; inflorescences not paniculate or
A diverse subfamily comprising about capitate 5
59 genera, sometimes recognized as a separate 4. Petals +; inflorescence terminal, paniculate; pollen not
spinulose; fruit capsular 8. Spondiantheae
family Phyllanthaceae or Antidesmataceae. Airy
– Petals 0; inflorescence axillary, capitate; pollen spinu-
Shaw (1965) split off from it four families—Bis- lose; fruit drupaceous 9. Uapaceae
chofiaceae, Hymenocardiaceae, Stilaginaceae, 5. Leaves with tanniniferous cells in upper epidermis; flow-
and Uapacaceae; Meeuse (1990) added one more ers in axillary or terminal spike-like or raceme-like
segregate, Putranjivaceae. Except for Bischofia, inflorescences, with discernible axes 6
none of Airy Shaw’s proposed segregates is suffi- – Leaves without tanniniferous cells in upper epidermis;
ciently aberrant to make its membership in the flowers in axillary glomerules, usually without discern-
ible axes [except in Flueggea gracilis and some Phyl-
Phyllanthoideae questionable, and all structural lanthus] 7
Euphorbiaceae 65
6. Thecae usually separate and connective enlarged, was first suggested by Wurdack et al. (2004) and
anther locules pendulous; leaves not glandular later confirmed by, among others, Vorontsova
5. Antidesmateae et al. (2007). The clade comprises genera that
– Thecae not separate and connective not enlarged formerly belonged to three different tribes, and
(enlarged in some Aporosa), anther locules not pendu-
lous; leaves sometimes glandular 6. Scepeae the group (and even some of its genera!) cannot
7. Indumentum 0; pistillate sepals persistent; inflorescence be recognized on the basis of morphological char-
sometimes [Celianella] pedunculate 7. Jablonskieae acters alone. x ¼ 11, 12.
– Indumentum + or 0; pistillate sepals usually not per-
sistent; inflorescences not pedunculate 8
8. Foliar glands rarely +; sepals distinct or nearly KEY TO THE GENERA OF PORANTHEREAE (from
completely connate; petals 0 [+ in Heterosavia]; disk Voronts. & Hoffm. 2008).
+ or 0; pistillode + or 0; ovules hemitropous or rarely
1. Anther dehiscence poricidal; flowers appearing to be in
[Heterosavia] anatropous; seeds exarillate or rarely
terminal umbels or racemes (in fact in dense aggrega-
with colored sarcotesta 4. Phyllantheae
tions of axillary flowers); fruit deeply 6-lobed; no sec-
– Foliar glands 0; sepals distinct; petals + or 0; disk +; ondary leaf venation visible with the naked eye.
pistillode +; ovules anatropous [hemitropous in Australia and New Zealand 4. Poranthera
Andrachne]; seeds exarillate 9
– Anther dehiscence longitudinal; flowers clearly axil-
9. Petals + in staminate flowers, 0 in Chascotheca, usually lary, staminate flowers sometimes on short densely
equaling the sepals 3. Wielandieae bracteate inflorescences; fruit 3(5)-lobed or globose;
– Petals + or 0 in staminate flowers 10 secondary leaf venation visible with the naked eye,
10. Sepals 4–6(7), imbricate (valvate in Bridelia, Clei- usually prominent 2
stanthus); stamens 4–10(19); fruit dehiscent or inde- 2. Petals in staminate flowers 0 or, if present, less than half
hiscent 2. Bridelieae the sepal length, if petals longer, then of unequal size or
– Sepals (3)5–6, imbricate; stamens 3–6; anthers rarely filiform [5(6) perianth segments clearly visible]; petals
poricidal [Poranthera]; fruit explosively dehiscent in pistillate flowers, if present, very similar to those in
1. Poranthereae staminate ones; disk annular, entire or shallowly undu-
late; stipules usually deciduous 3
– Petals in staminate flowers 5(6), nearly as long
as sepals, all equal in size, white [10(12) perianth seg-
IA. SUPERTRIBE PHYLLANTHODAE G.L. Webster, ments clearly visible]; petals in pistillate flowers, if
supertrib. nov.1 present, less than half the sepal length, often hidden
by the disk; staminate disk glands distinct or fused at
Subfam. Phyllanthoideae Kostel. (1836). the base, bilobed, or almost 10-crenate (Notoleptopus)
or irregularly crenate (Pseudophyllanthus); pistillate
disk glands distinct or disk annular, more or less cre-
1. TRIBE PORANTHEREAE Gr€
uning (1913), nate; stipules persistent 4
recircumscribed by Hoffmann et al., Kew Bull. 61: 41 3. Seeds smooth; endosperm scanty or 0; disk convex,
(2006), Vorontsova et al., Amer. J. Bot. 94: 2026–2040 thick and fleshy, becoming thinner toward edge; fila-
(2007), mol. syst.; Vorontsova & Hoffmann, Kew Bull. 63: ments distinct or connate at base only; petals in stami-
41–59 (2008), classif. nate flowers 0 or, if present, unequal or filiform;
Staminate disk in distinct parts or extrastam- stipules deciduous 2. Actephila
inal-annular, pistillate disk simple; stamens 3–6; – Seeds rugose to fenestrate; endosperm copious; disk
filaments distinct or variously connate; anthers concave, slightly cupular, usually thin and chartaceous;
longitudinally dehiscing or poricidal; thecae not filaments connate for 1/4 to 9/10 of their length, rarely
separate and connective not enlarged; pistillode (former Zimmermannia and Zimmermanniopsis) dis-
tinct; petals in staminate flowers 0 or rarely (subg.
usually +; ovary 3-locular; stylodia distinct or Petaliferae) 5, equal, less than half of the sepal length;
completely connate; fruits explosively dehiscent; stipules deciduous or persistent 7. Meineckia
seeds exarillate. 4. Plant including branches, leaves, sepals, ovaries and
Eight genera, (sub)tropical regions, world- fruit densely covered with hairs about 1 mm long,
wide. The circumscription of this group of genera visible to the naked eye; seed intensely black, rugose
6. Notoleptopus
– Plant glabrous, with sparse covering of simple hairs, or
rough with gland-tipped hairs not visible to the naked
eye; seed whitish to dark brown, smooth to rugose 5
1
Supertribe Phyllanthodae G.L. Webster, supertrib. nov., 5. Procumbent herbs to subshrubs up to 0.5 m tall (erect
floribus cymulis axillaribus, cellulis epidermalibus foliorum in Andrachne fruticulosa); ovule adnation hemitro-
non tanniniferis, embryonibus plerumque non chlorophyl- pous; seed rough, with star-shaped sculpture visible at
losis, fructibus saepe capsularibus. Typus: Phyllanthus L. high magnification when dry 8. Andrachne
66 G.L. Webster
– Erect shrubs to trees 0.2–6 m tall (Leptopus clarkei 2. Actephila Blume

procumbent); ovules anatropous; seed smooth, rugose
or pitted, without star-shaped sculpture 6 Actephila Blume, Fl. Ned. Ind.: 581 (1825); Forster, Aus-
6. Staminate disk irregularly crenate to almost lacerate; trobaileya 7: 57–98 (2005), Austral. spp.; Vorontsova &
pistillate disk annular, finely crenulate; filaments usu- Petra Hoffm., Kew Bull. 63: 46–47 (2008).
ally connate for about one half of their length, rarely Monoecious trees to subshrubs; indumentum
distinct. Seed undulate to shallowly rugose; endosperm
invaginate independently of seed coat sculpture simple or 0. Leaves subverticillate or subopposite,
5. Pseudophyllanthus entire or rarely toothed; stipules deciduous. Inflo-
– Staminate disk regularly divided into 5 rounded rescences axillary, fasciculate, or flowers soli-
bilobed segments; pistillate disk regularly divided into tary; pedicels usually elongated, not articulate.
5 emarginate segments; filaments distinct; seeds Staminated flowers: sepals 4–6; petals 4–6(0),
smooth, rugose or pitted; endosperm entire in cross
section or with undulations following seed coat curva-
shorter than sepals; disk annular, entire or shal-
ture 7 lowly undulate; stamens (3–)5(6), filaments dis-
7. Stylodia bifid for half their length or less; stigmas acute tinct or connate basally with pistillode; anthers
to dilated; seed smooth 3. Phyllanthopsis introrse; pollen prolate to subprolate, exine reticu-
– Stylodia bifid to base or almost so; stigmas capitate to late-striate; pistillode apically 3(5)-fid. Pistillate
dilated; seed smooth, rugose or pitted flowers: pedicels longer than staminate ones; sepals
1. Leptopus
and petals as in staminate; disk annular; ovary 3-
locular, ovules 2 per locule, anatropous; stylodia
1. Leptopus Decne.
short, distinct or nearly so, entire to bifid to base at
Leptopus Decne., Voy. Inde 4 (Bot.): 155 (1836); Voronts. apex. Fruit capsular, loculicidally 3-valved or
& Petra Hoffm., Kew Bull. 63: 46 (2008) and 64: 627–644 separating into 2-valved cocci; columella apically
(2009), rev.
Archileptopus P. T. Li (1991). 3-pronged, basally dilated. Seeds 2(1) per locule,
sometimes ecarunculate, ovoid, carinate; testa
Monoecious (dioecious) herbs or shrubs; indu- smooth and dry; endosperm scanty or 0; embryo
mentum simple or 0. Leaves entire; stipules per- folded, cotyledons thick, fleshy, sometimes plicate.
sistent. Inflorescences axillary, fasciculate but About 35 spp., Asia, Australia and Melanesia,
mostly flowers solitary; staminate flowers some- in rainforest understorey.
times on short bracteate inflorescences. Staminate
flowers with pedicels articulate above base; sepals 3. Phyllanthopsis (Scheele) Voronts. & Petra
5(6), petals 5(6), shorter than or as long as sepals; Hoffm.
disk of 5(6) contiguous, bilobed segments; Phyllanthopsis (Scheele) Voronts. & Petra Hoffm., Kew
stamens 5, filaments distinct; anthers longitudi- Bull. 63: 47 (2008).
nally dehiscent; pollen prolate to subprolate, exine Phyllanthus subg. Phyllanthopsis Scheele (1853).
tectate to microperforate; pistillode of 3 distinct Andrachne sect. Phyllantopsis (Scheele) M€ull. Arg. (1866);
segments or 3-lobed. Pistillate flowers with pedi- Petra Hoffm., Bot. Jahrb. 116: 321–331 (1994).
cels apically dilated; perianth as in staminate flow- Monoecious or dioecious shrubs; indumentum
ers but petals often minute; disk annular, regularly simple or 0. Leaves entire; stipules persistent.
divided into 5 emarginate segments; ovary 3–4(5)- Inflorescences axillary, fasciculate or flowers soli-
locular; ovules anatropous; stylodia distinct, bifid tary. Staminate flowers pedicellate; sepals 5; petals
to base or nearly so; stigmas apically dilated to 5, slightly shorter than sepals; disk annular, divided
capitate. Fruit 3(5)-lobate to subglobose; colu- into 5 bilobed segments; stamens 5; filaments dis-
mella 4–5 times longer than its narrowest width. tinct; anthers longitudinally dehiscent; pistillode of
Seeds 2 per locule, ecarunculate; endosperm copi- 3 distinct segments. Pistillate flowers pedicellate,
ous, in cross section entire to deeply invaginate; sepals 5; petals 5, much shorter than sepals; disk
embryo curved to almost straight; cotyledons as annular, divided into 5 emarginate segments; ovary
long as radicle or longer, wider than radicle. 2n ¼ 3-locular; ovules anatropous; stylodia distinct,
22, 24. bifid for up to half their length; stigmas acute to
Nine spp., deciduous forests, Caucasus to apically dilated. Fruit rounded to 3-lobed; colu-
Malesia. mella 6 times longer than its narrowest width.
Seeds 2 per locule, ecarunculate, smooth; endo-
sperm copious, entire in cross section; embryo
Euphorbiaceae 67
curved; cotyledons broad, thin, longer and wider

than radicles.
Two spp., Mexico, USA, in dry scrub.
4. Poranthera Rudge Fig. 11
Poranthera Rudge, Trans. Linn. Soc. London 10: 302
(1811); Halford & Henderson, Austrobaileya 7: l–27
(2005); Voronts. & Petra Hoffm., Kew Bull. 63: 50 (2008).
Oreoporanthera (Gr€uning) Hutch. (1969).
Monoecious (dioecious) annual or perennial

herbs, sometimes suffruticose; indumentum 0.
Leaves alternate (opposite), sessile or petiolate,
entire, 2–20 times longer than wide; stipules pale,
scarious. Inflorescences axillary but appearing to
be in terminal umbels or racemes, fasciculate or
flowers solitary. Staminate flowers: sepals (3)5,
not accrescent; petals (3)5 or 0, smaller than
sepals; disk of (3)5 distinct globular entire glands,
stamens (3)5; filaments distinct; anthers dehiscing
by terminal pores; pollen prolate(spheroidal),
exine tectate-perforate to reticulate; pistillode 3-
Fig. 11. Euphorbiaceae-Phyllanthoideae. Poranthera
fid. Pistillate flowers: sepals and petals as in stami- obovata. A Flowering branchlet. B Staminate flower. C
nate; disk annular and divided into (3)5 erose to Pistillate flower. D Branchlet with stipulate leaf. E Fruit
undular segments; ovary 3-locular, 6-angled; sty- seen from above. F Same from side. (Halford & Hender-
lodia bifid to base; stigmas rounded to capitate. son 2005, drawn by W. Smith)
Fruit deeply 6-lobed; columella 8 times longer
than its narrowest width, persistent. Seeds 2 per
locule, ecarunculate, smooth, reticulate, rugose or pistillode 3-lobed. Pistillate flowers: sepals and
fenestrate; endosperm copious; embryo curved or petals as in staminate; disk annular, finely crenu-
straight; cotyledons shorter than radicle. late; ovary 3-locular; stylodia distinct, bifid to base;
Fifteen spp., Australia, Tasmania, and New stigmas capitate. Fruit 3-lobed; columella 2.5–5
Zealand, in temperate grassland and woodland. times longer than its narrowest width. Seeds ecar-
This is the morphologically most aberrant genus unculate; endosperm copious, deeply invaginate
of the tribe. independently of outer seed coat sculpture; embryo
almost straight; cotyledons longer and wider than
5. Pseudophyllanthus (M€
ull. Arg.) Voronts. & radicle.
Petra Hoffm. One sp., P. ovalis (E. Mey. ex Sond.) Voronts.
& Petra Hoffm., southern Africa, deciduous for-
Pseudophyllanthus (M€ ull. Arg.) Voronts. & Petra Hoffm., ests.
Kew Bull. 63: 50 (2008).
Andrachne sect. Pseudophyllanthus M€ ull. Arg. (1866),
Petra Hoffm., Bot. Jahrb. 116: 321–331 (1994). 6. Notoleptopus Voronts. & Petra Hoffm.
Dioecious (monoecious) pubescent shrubs to small Notoleptopus Voronts. & Petra Hoffm., Kew Bull. 63: 50
(2008).
trees. Leaves entire; stipules membranaceous.
Inflorescences axillary, fasciculate or flowers soli- Monoecious, densely pubescent shrubs. Leaves
tary or in twos, staminate flowers on short densely entire; stipules persistent. Inflorescences axillary,
bracteate inflorescences leaving pedicel stumps fasciculate or flowers solitary; pedicels articulate.
("podia"); pedicels articulate. Staminate flowers: Staminate flowers: sepals 5; petals 5; disk annular,
sepals 5; petals 5; disk annular, irregularly crenate regularly divided into 5 bilobed segments; stamens
to almost lacerate; stamens 5; filaments distinct to 5; filaments distinct; anthers longitudinally dehis-
connate for about half their length; pollen subpro- cent; pollen prolate, exine tectate-perforate to
late, exine tectate-perforate to micro-reticulate; microreticulate; pistillode of 3 distinct segments.
68 G.L. Webster
Pistillate flowers: sepals 5; petals 5, much shorter of Vorontsova et al. (2007) were found to be basal
than sepals; disk annular, regularly and deeply to the otherwise apetalous species of M. subg.
divided into 5 emarginate or erose segments; Meineckia; morphologically, they are also very
ovary 3-locular, densely pilose; stylodia distinct, close to Pseudophyllanthus (see Hoffmann, Adan-
bifid to base; stigmas rounded to apically dilated. sonia III, 22: 123–133. 2000).
Fruit 3-lobed; columella 6 times longer than its
narrowest width. Seeds ecarunculate, rounded tri- 8. Andrachne L.
quetrous, rugose; endosperm copious, deeply
Andrachne L., Sp. Pl.: 1014 (1753); Petra Hoffm., Bot. Jahrb.
invaginate; cotyledons broad, thin, as long as radi- 116: 321, figs. 1, 2 (1994), Adansonia III, 22: 123 (2000).
cle, wider than radicle.
One sp., N. decaisnei (Benth.)Voronts. & P. Monoecious annual to perennial herbs or sub-
Hoffm., from Java to New Guinea, Australia, in shrubs; indumentum simple, glandular, or 0.
open forest and savannah. Although habitually Leaves with persistent stipules. Inflorescences axil-
very similar to Leptopus (see Vorontseva et al. lary, fasciculate, or flowers solitary, the staminate
2007), molecular evidence placed this taxon in glomerules, the pistillate solitary; pedicels artic-
close to the Southern Hemisphere Pseudophyl- ulate. Staminate flowers: sepals 5(6), distinct; petals
lanthus and Poranthera. 5, slightly shorter than sepals in staminate flowers;
disk in staminate flowers of 5 contiguous bilobed
7. Meineckia Baill. segments or distinct globular glands; stamens 5;
filaments distinct to connate up to half of their
Meineckia Baill., Étude Gén. Euphorb.: 587 (1858); Web-
ster, Acta Bot. Neerl. 14: 323–365 (1965), rev.; Vorontsova length; pollen (spheroidal-)prolate, exine striate
& Petra Hoffm., Kew Bull. 63: 53–55 (2008). to micro-reticulate; pistillode of 3 distinct segments
Zimmermannia Pax (1910). or 3-lobed. Pistillate flowers: sepals 5(6), distinct;
Zimmermanniopsis Radcl.-Sm. (1990). petals much shorter than sepals to minute or 0;
Monoecious or dioecious shrubs or subshrubs; disk either divided into 5 emarginate segments or
indumentum simple or 0. Leaves distichous, 5 distinct globular glands; ovary 3-locular; ovules
epeltate or rarely (M. peltata) peltate; stipules hemitropous; stylodia distinct, bifid, stigmas
persistent or deciduous. Inflorescences axillary, capitate. Fruit rounded to 3-lobed; columella 5–6
fasciculate or flowers solitary; staminate flowers times longer than its narrowest width. Seeds ecar-
sometimes on short densely bracteate inflores- unculate, rounded-triquetrous, rough, with star-
cences, on which the pedicels leave "podia"; shaped sculpture visible at high magnification
pedicels articulate. Staminate flowers: sepals 5 (see illustration in Stuppy 1996: 283); endosperm
(6); petals 0 (5 small ones in subg. Petaliferae); copious, entire in cross section; embryo curved;
disk annular, entire or shallowly undulate; cotyledons broader than and about equaling radi-
stamens 5, filaments distinct to connate up to cle in length. 2n ¼ 24.
9/10 of their length; pollen prolate to speroidal, Twenty-two spp., in semi-deserts of South Asia,
exine reticulate to verrucate; pistillode subentire northern Africa, southern Europe and the Americas;
to 3-lobed. Pistillate flowers: sepals 5, imbricate, two subgenera, A. subg. Andrachne, 16 spp., Old
persistent in fruit; petals 0; disk annular; ovary World, and A. subg. Phyllanthidea (Didr.) Voronts.
3-locular, glabrous or densely pilose; stylodia & Petra Hoffm., 6 spp., also in the New World.
distinct or connate at base, entire to bifid to
base, stigmas apically dilated to capitate. Fruit 2. TRIBE BRIDELIEAE M€ull. Arg. (1864),
3-lobed; columella slender, persistent. Seeds 1 or recircumscribed by Hoffmann et al., Kew Bull. 61:
2 per locule, ecarunculate; endosperm copious, 37–53 (2006).
deeply invaginate; cotyledons much broader Armature 0 or rarely +; leaves entire or slightly
than radicle. crenate; stipules distinct or rarely intrapetiolar;
Thirty spp., mostly in deciduous forests of pedicels sometimes articulated; sepals 4–6 (7),
Meso-America and South America, Africa, Mada- imbricate (valvate); petals (3)4–6 or 0; staminate
gascar and South Asia. Two of them represent M. disk extrastaminal-annular or in distinct parts or
subg. Petaliferae Voronts. & Petra Hoffm.; these 0; pistillate disk simple or double; stamens 4–10
have minute petals and in the molecular analysis (19), distinct or variously fused; thecae not
Euphorbiaceae 69
separate and connective not enlarged; ovary Dioecious shrubs or trees. Leaves deciduous; sti-
locules 2–4; stylodia distinct or completely con- pules deciduous. Inflorescences axillary glomerules,
nate; fruit explosively or tardily dehiscent or the staminate dense glomerules, the pistillate few-
indehiscent; seeds exarillate. to 1-flowered; bracts obscure; pedicels articulated.
Twelve genera in five subtribes. Staminate flowers subsessile; sepals 3–5, imbricate;
disk annular; petals 5, imbricate, shorter than
2a. SUBTRIBE AMANOINAE Pax & K. Hoffm. (1922). sepals, or 0; stamens 5, filaments distinct; anthers
basifixed, introrse; pistillode 3-fid. Pistillate flowers:
A monotypic subtribe. sepals 5, distinct, imbricate; petals 5 or 0; ovary 3-
locular, sometimes pubescent; stylodia distinct,
9. Amanoa Aubl. (1775). bifid, branches slender. Fruit capsular; columella
Amanoa Aubl., Fl. Gui. Fr.: 256 (1775); Hayden, Brittonia 3-angled distally, persistent. Seeds mostly 1 per
42: 260–290 (1990), Neotrop. spp. locule, smooth; endosperm copious, embryo
Monoecious (dioecious) glabrous trees or shrubs. straight; cotyledons much longer than radicle.
Leaves entire; stipules persistent, distinct or intra- Two or three spp., neotropical. Other species
petiolar. Inflorescences terminal or axillary, spici- formerly included in this genus have been trans-
form-thyrsoid; pedicels not articulated. Staminate ferred to Heterosavia (Tribe Phyllantheae) or
flowers: sepals 5, imbricate, persistent; petals 5, Wielandia (Tribe Wielandieae) (Hoffmann and
scale-like; disk extrastaminal-annular; stamens 5, McPherson 2007; Hoffmann 2008).
distinct; anthers introrse; pollen oblate, 3-colpo-
rate, intectate or tectum widely perforate; pistil- 11. Gonatogyne M€
ull. Arg.
lode columnar. Pistillate flowers: sepals 5, Gonatogyne M€ ull. Arg., Fl. Bras. 11(2): 13 (1873).
deciduous in fruit; petals much shorter than sepals Savia sect. Gonatogyne (M€ ull. Arg.) Pax & K. Hoffm.
or sometimes 0; disk patelliform; staminodes 0; (1922).
ovary subglobose, 3-locular; ovules anatropous; Dioecious evergreen trees; indumentum simple.
stylodia connate into a massive column, tips Leaves entire; stipules deciduous. Inflorescences
dilated, stigmatiform. Fruit 3-angled, tardily axillary bracteate glomerules; sepals 5, imbricate,
dehiscent; columella massive, dilated toward the pistillate deciduous in fruit; petals 5, imbri-
base and at apex. Seeds usually 1 per locule; endo- cate, as long as sepals; staminate disk fleshy,
sperm scanty or 0; cotyledons fleshy. corrugated; stamens 5, filaments connate into a
Sixteen spp., 13 in tropical South America, column; anthers stipitate, erect, introrse; pollen
mostly northern, and 3 in west and west-central grains 3-colporate, reticulate; pistillode atop col-
tropical Africa. This is the only genus of umn; pistillate flowers pedicellate; staminodes 0;
Bridelieae that occurs in both the New World disk multiglandular; ovary 3(4)-locular, seri-
and the Old. ceous; stylodia twice bifid. Fruit capsular, persi-
stent; seeds smooth; endosperm copious;
2b. SUBTRIBE SAVIINAE M€
ull. Arg. (1865), as cotyledons flat, greenish.
’Savieae’. One sp., G. brasiliensis (Baill.) M€ ull. Arg.,
Croizatieae G.L. Webster (1994). Brazil (São Paulo).
Dioecious; pedicels sometimes articulated; sepals
imbricate, persistent; staminate disk extrastaminal- 12. Croizatia Steyerm.
annular; pistillate disk simple; staminodes rarely +; Croizatia Steyerm., Fieldiana 28: 308 (1952); Webster
stylodia 2-fid or 4-fid; fruit dehiscent. et al., Syst. Bot. 12: 1–8, figs. 1–8 (1987); Dorr, Sida 18:
Five genera, neotropical. 831–836 (1999).
Pseudosagotia Secco (1985).
10. Savia Willd. Dioecious trees or shrubs. Leaves alternate to

Savia Willd., Sp. Pl. 4: 771 (1805); P. Hoffmann & McPher-
subopposite; stipules deciduous or persistent.
son, Ann. Miss. Bot. Gard. 94: 519–55 (2007); P. Hoff- Inflorescences axillary, glomerular; bracts incon-
mann, Brittonia 60: 136–166 (2008), rev. spicuous. Staminate flowers pedicellate; sepals 5,
Kleinodendron L.B. Sm. & Downs (1964). distinct, imbricate; petals 5, very small; disk
70 G.L. Webster
annular, glabrous; stamens 5, filaments distinct; Dioecious trees. Stipules deciduous. Inflore-
anthers introrse; pollen grains spheroidal, 3-bre- scences axillary, glomerular, sessile or subses-
vicolporate, sexine tectate-perforate and spinose; sile. Staminate flowers pedicellate; sepals 5,
pistillode 3–4-lobed. Pistillate flowers pedicellate; imbricate; petals 5, equaling or exceeding sepals;
sepals 5, distinct, persistent in fruit; petals 5, disk obscure; stamens 5 or [T. amanoifolia Huft]
much shorter than sepals; disk annular, glabrous; 14–19, filaments distinct; pollen grains 3-colpo-
ovary 3-locular, pubescent; ovules hemitropous; rate, exine striate- reticulate; pistillode colum-
stylodia distinct, slender, twice 2-fid. Fruits cap- nar; pistillate flowers pedicellate; sepals 3 and
sular, 3-lobed, cocci reticulate; columella distally "petals" 3, persistent in fruit, or sepals 5 and
expanded in papery wings. Seeds 1 or 2 per locule, petals 0; disk annular (?); ovary 3-locular, seri-
ecarunculate; testa smooth; endosperm 0; embryo ceous; stylodia bifid. Fruit capsular; columella
large, cotyledons thin, contorted. scarcely winged. Seeds apparently 1 per coccus,
Four spp., Panama to Venezuela and Ecuador angled in cross section, testa smooth; endo-
and possibly Peru. Because of its spinose pollen, sperm scant; embryo large; cotyledons thin,
Croizatia was placed in the Oldfieldioideae (Web- contorted.
ster 1994; Radcliffe-Smith 2001), but there is now Three spp., Panama and South America, still
strong molecular support for its inclusion in this unsatisfactorily known.
subtribe.
2c. SUBTRIBE KEAYODENDRINAE P. Hoffm. (2006).
13. Discocarpus Klotzsch
A monotypic subtribe, distinguished by the
Discocarpus Klotzsch, Archiv. Naturg. 7: 201 (1841); Hay- combination of apetalous flowers, indehiscent,
den & Hayden, Ann. Missouri Bot. Gard. 83: 153–167
(1996).
1-locular and 1-seeded fruits.
Dioecious trees or shrubs. Stipules deciduous. 15. Keayodendron Leandri

Flowers in axillary glomerules, the staminate
Keayodendron Leandri, Bull. Soc. Bot. France 105: 517
compounded. Staminate flowers sessile; sepals (1959); Breteler, Bull. Jard. Bot. Nat. Belg. 62: 187–190 (1993).
5, basally connate; petals 5, often reduced; disk
cupular, lobed or toothed, stamens (4)5; fila- Dioecious trees; indumentum simple. Stipules
ments basally connate, exserted from perianth; deciduous. Flowers in axillary glomerules. Pedi-
anthers longitudinally introrsely dehiscent; pol- cels nor articulated. Staminate flowers subsessile
len grains oblate, 4- or 5-colporate, exine or pedicellate; sepals 5–7, imbricate; petals 0; disk
finely reticulate; pistillode divided into 2 or 3 extrastaminal-annular; stamens 5–6; filaments
narrow segments; pistillate flowers subsessile or connate; anthers introrse, dehiscing longitudi-
pedicellate; sepals and petals as in staminate; nally; pistillode dilated and subpeltate. Pistillate
disk slightly lobed, glabrous; ovary 3-locular, flowers subsessile or pedicellate; perianth similar
densely hirtellous; ovules anatropous; stylodia to staminate, sepals 5 or 6, persistent in fruit; disk
dilated, bifid or irregularly incised. Fruits capsu- double, minutely pubescent; ovary 2-locular; sty-
lar, dehiscent into 1- or 2-seeded mericarps; lodia 2-fid. Fruit drupaceous, 1-locular. Seed 1
columella massive, 3-winged distally. Seeds subper fruit, linear-compressed, deeply invaginate
globose, testa smooth and shiny; endosperm ventrally; endosperm scanty, embryo curved,
scanty or 0; embryo large, cotyledons thin, con- cotyledons much longer than radicle.
torted. One sp., K. bridelioides Leandri, semi-decidu-
Four spp., northern South America: Amazon ous forests of west tropical Africa (Ivory Coast to
and Orinoco basins, Guayana region, E Brazil. Cameroun).
14. Tacarcuna Huft 2d. SUBTRIBE PSEUDOLACHNOSTYLIDINAE Pax &

K. Hoffm. (1922).
Tacarcuna Huft, Ann. Missouri Bot. Gard. 76: 1080
(1989); Webster, Ann. Missouri Bot. Gard. 81: 130 Armature rarely +; leaves entire, rarely slightly
(1994); Wurdack in Wurdack et al., Amer. J. Bot. 91: crenate; pedicels sometimes articulated;
1892 (2004). sepals imbricate or valvate; staminate disk
Euphorbiaceae 71
extrastaminal-annular; pistillate disk usually dou- pollen grains subglobose, 3-colporate, exine striate;
ble; stamens 4–7; fruit dehiscent or indehiscent. pistillode at apex of column, 3-lobed. Pistillate flow-
Four Old World genera. ers sessile or pedicellate; sepals mostly 5, valvate,
connate basally, usually persistent in fruit; petals 5,
16. Bridelia Willdenow (1806). scale-like, adnate to sepalar cup; disk annular or
Bridelia Willd., Sp. Pl. 4(2): (1806; as Briedelia); Webster,
cupular; ovary (2)3(4)-locular; ovules anatropous,
Ann. Missouri Bot. Gard. 81: 39 (1994); Dressler, Blumea rarely hemianatropous; stylodia 1–4-lobate or -fid.
41: 263–331 (1996); Radcl.-Sm., Gen. Euphorb.: 19 (2001). Fruit capsular, septicidal or tardily dehiscent to
nearly indehiscent [sect. Chartacei Jabl.], with sub-
Monoecious (dioecious) trees or shrubs, branches
tending podium; columella usually persistent.
sometimes thorny; indumentum simple. Leaves
Seeds 1 or 2 per coccus; testa smooth and shiny;
entire or subentire; stipules persistent or decidu-
endosperm copious; embryo straight; cotyledons
ous. Flowers in axillary glomerules, flowering
longer and broader than radicle. 2n ¼22.
branches sometimes appearing racemose due to
About 140 spp. distributed from Africa and
reduction of leaves; bracts persistent. Staminate
Madagascar to Malesia, Australia, and Polynesia.
flowers pedicellate or subsessile; sepals 5, valvate;
The genus is highly variable in various characters
petals 5, erect or inflexed, smaller than sepals; disk
including foliar venation, pollen morphology and
annular or somewhat cupular, glabrous or pubes-
seed anatomy (Stuppy 1996; Tokuoka and Tobe
cent; stamens 5, filaments basally connate; pollen
2001), is polyphyletic and needs profound taxo-
grains slightly oblate, 3-colporate, exine reticulate
nomic readjustment.
with striate pattern; pistillode 2–4-lobed or -
divided. Pistillate flowers sessile or subsessile;
sepals 5, basally connate into a hypanthium, val- 18. Pentabrachion M€
ull. Arg.
vate, persistent in fruit; petals as in staminate; disk Pentabrachion M€ ull. Arg., Flora 47: 532 (1864); Pax & K.
duplex, outer part annular and adnate to hypan- Hoffm., Pflanzenr. 147: XV: 188, fig. 15 (1922); Radcl.-Sm.,
thium, inner part inflexed as cupule surrounding Gen. Euphorb.: 15 (2001).
the ovary; ovary 1- or 2-locular; stylodia distinct or Monoecious or dioecious subglabrous shrubs or
basally connate, bifid to subentire. Fruits pseudo- trees. Leaves entire; stipules persistent. Inflore-
baccate with fleshy exocarp but sometimes dehis- scences axillary glomerules. Staminate flowers
cent or drupaceous, 1- or 2-locular; columella not pedicellate; sepals (4)5(6), imbricate; petals 4–6,
persistent. Seeds usually 1 per locule, plano-con- smaller than sepals; disk annular, unlobed;
vex, adaxially grooved [in 2-locular fruits] or ven- stamens 5(7), filaments basally connate; pollen
trally invaginate [in 1-locular fruits]; endosperm grains oblate sphaeroidal, 3-colporate, colpi c. 2/3
copious, excavated adaxially; embryo chlorophyl- length of grain, exine coarsely heterobrochate;
lous, curved, cotyledons broad, not fleshy. 2n ¼ 26. pistillode 3-fid, segments laminar. Pistillate flow-
About 50 spp. extending from Africa and ers long-pedicellate; sepals 5, imbricate; disk
Madagascar to Australia and the Pacific islands. annular; ovary glabrous; stylodia twice bifid.
Fruits capsular, 3-lobed. Seeds smooth, endo-
17. Cleistanthus Hook. f. ex Planch. (1848). sperm copious, embryo with cotyledons longer
than radicle.
Cleistanthus Hook. f. ex Planch. in Hook. Ic. Pl. 8: t. 779 One sp., P. reticulatum M€ ull. Arg., western
(1848); Kathriarachchi et al., Molec. Phylogenet. Evol. 36:
112–134 (2005); Li et al., Syst. Bot. 34: 529 (2009).
equatorial Africa.
Monoecious (dioecious) shrubs or trees; indumen- 19. Pseudolachnostylis Pax

tum simple. Leaves entire; stipules persistent or
deciduous. Flowers in axillary, rarely terminal glo- Pseudolachnostylis Pax, Bot. Jahrb. 28: 19 (1899); Radcl.-
Sm., Fl. Trop. E. Afr., Euphorb. 1: 80, fig. 10 (1987), Gen.
merules or axillary thyrses; bracts deciduous. Sta- Euphorb.: 28 (2001).
minate flowers usually sessile; sepals (4)5(6),
valvate; petals (4)5(6), scale-like and shorter than Dioecious shrubs or trees; buds perulate; indu-
sepals; disk annular or cupulate, entire or lobed; mentum simple. Leaves entire; stipules decidu-
stamens (4)5(6); filaments connate at base; ous. Inflorescences axillary or arising below the
72 G.L. Webster
leaves, staminate flowers in few-flowered pedun- two genera and of S. congesta M€ ull. Arg.
culate or subsessile cymes, pistillate flowers (sub) (¼Jablonskia) by Webster (1965–1984), resulting
solitary. Staminate flowers sessile; sepals 5(6), in a core Malagasy group characterized by dis-
imbricate; petals 0; disk annular-lobed; stamens tinctive spiny pollen and smooth seeds (Wurdack
(4)5–7; filaments adnate to pistillode; pollen et al. 2004). Lachnostylis has a strongly supported
grains oblate, 3-colporate, coarsely reticulate; relationship with Securinega capuronii Leandri,
pistillode 3-fid. Pistillate flowers: pedicels 2-brac- despite the lack of morphological resemblance
teolate; sepals 5(6), imbricate, deciduous in fruit; (Wurdack et al. 2004).
petals 0; disk dentate or lobed; ovary 3-locular;
ovules hemitropous; stylodia bifid. Fruit a tardily 21. Lachnostylis Turcz. (1846).
dehiscent woody capsule with fleshy exocarp;
columella not persistent. Seeds 1 per locule, ellip- Lachnostylis Turcz., Bull. Soc. Imp. Naturalistes Moscou
19: 503 (1846); Dyer, Gen. S. Afr. Pl. 1: 309 (1975); Radcl.-
soid, hilum submedian; endosperm copious; Sm., Gen. Euphorb.: 12 (2001).
cotyledons much longer than radicle. 2n ¼ 26.
Apparently only one variable sp., P. maprounei- Dioecious shrubs; indumentum simple. Leaves
folia Pax, distributed from Zaire and Tanzania to small; stipules deciduous. Inflorescences axillary
South Africa. glomerules; bracts scarious; female flowers often
solitary. Staminate flowers pedicellate; perianth
2e. SUBTRIBE SECURINEGINAE M€
ull. Arg. (1866). abaxially sericeous; sepals 5, distinct, imbricate,
persistent; petals 5, equaling the sepals; disk
Two Afro-Madagascan genera without evident extrastaminal, annular, massive, unlobed, hirtel-
similarities, but strongly supported in a subclade lous; stamens 5, filaments connate below, villous;
that is sister to the remaining four subtribes. anthers dehiscing longitudinally; pollen grains
subprolate, 3-colporate, exine reticulate; pistil-
20. Securinega Comm. ex Juss. (1789). lode 3–4-fid, villous, adnate to staminal column.
Securinega Comm. ex Juss., Gen. Pl.: 388 (1789), nom. Pistillate flowers pedicellate; perianth similar to
cons.; Leandri, Fl. Madag. 111(1): 107–116 (1958); the staminate; disk massive, unlobed, villous;
Radcl.-Sm., Gen. Euphorb.: 31 (2001); Schatz, Generic ovary 2- or 3-locular, tomentose, ovules hemitro-
Tree Fl. Madagascar: 164, fig. 171 (2001). pous; stylodia bifid. Fruits capsular, valves
Dioecious shrubs. Leaves entire; stipules decidu- tomentose; each coccus with 1(2) seeds. Seeds
ous. Flowers in axillary glomerules, the pistillate subglobose, testa smooth, endosperm scanty;
flowers solitary or few; pedicels not articulated. cotyledons plicate, longer than the radicle.
Staminate flowers pedicellate; sepals 5–7, imbri- One variable or perhaps 2 spp., T. capensis
cate, deciduous; petals 0; disk in 5 distinct seg- Turcz., South Africa.
ments; stamens (4)5(10), filaments distinct,
exserted from calyx; pollen grains subglobose, 3- 3. TRIBE WIELANDIEAE Baill. ex Hurus. (1954),
colporate, colpi long, exine spiny or finely reticu- recircumscribed by Hoffmann et al., Kew Bull. 61:
late; pistillode mostly 2–3-fid. Pistillate flowers 37–53 (2006).
pedicellate; sepals 5, imbricate, persistent in fruit;
Monoecious or dioecious; leaf base sometimes
petals 0; disk annular; ovary 3(4)-locular; stylodia 3
peltate; petioles sometimes pulvinate; sepals 4–6
(4), sessile, recurved, 2-fid. Fruit capsular; colu-
(8), imbricate, distinct; petals 4–6, sometimes
mella deciduous. Seeds ovoid, testa dark, smooth;
reduced or 0; disk + or rarely 0; staminate disk
hilum submedial, narrowly elliptic; endosperm
annular or in distinct parts, pistillate disk simple;
copious; embryo straight, cotyledons longer and
stamens (3)4–6(12); filaments distinct or vari-
broader than the radicle.
ously connate; ovary 3–6-locular; fruits explo-
Five spp., Madagascar, Réunion and Mauri-
sively dehiscent.
tius. This genus had been circumscribed to
Six genera, New and Old World, except Australia.
include Flueggea by M€ uller (1866), in which he
was followed by Pax and Hoffmann (1931) who
3a. SUBTRIBE ASTROCASIINAE G.L. Webster (1992).
added Meineckia. This broad circumscription
was reduced by the removal of the latter Three genera, two New World, one African.
Euphorbiaceae 73
22. Heywoodia Sim tillode 3-fid, adnate to staminal column. Pistillate

Heywoodia Sim, For. Fl. Cape Col.: 326, t. 140/1 (1907);
flower long-pedicellate, sepals and disk as in stami-
Hutchinson, Bull. Misc. Inf. Kew 1922: 115 (1922); G. nate, persistent, petals 0; ovary 3-locular, ovules
L. Webster, Ann. Missouri Bot. Gard. 81: 36 (1994); anatropous; stylodia short, 2-fid. Fruit capsular,
Radcl.-Sm, Fl. Zamb. 9(4): 6, t. 1 (1996), Gen. Euphorb.: venose; columella slender, persistent. Seeds 1 per
6 (2001). locule; testa dry, smooth; reticulate; chalaza
Dioecious glabrous trees. Leaves of seedlings and appearing dorsal, hilum in depression beneath ter-
sucker-shoots peltate; stipules persistent. Inflor- minal beak; endosperm copious; embryo deflexed,
escences axillary bracteate glomerules, staminate cotyledons about equaling radicle.
flowers many, pistillate 3 or 4. Staminate flowers One sp., Ch. neopeltandra (Griseb.) Urb.,
sessile or nearly so; perianth poorly differen- Greater Antilles (Cuba and Hispaniola).
tiated, sepals 3(4), distinct, imbricate, unequal;
petals (4)5, distinct, longer than sepals, imbricate; 24. Astrocasia B. L. Rob. & Millsp.
disk extrastaminal, lobed, projecting between fila-
Astrocasia B. L. Rob. & Millsp., Bot. Jahrb. 36, Beibl. 80: 19
ments; stamens 8–12, in 2 whorls, filaments (1905); G.L. Webster, Syst. Bot. 17: 311–323 (1992); Jimé-
distinct, anthers dorsifixed, introrse, muticous, nez R. & Gordillo, Acta Bot. Mex. 55: 1, fig. 1 (2001).
dehiscing longitudinally; pollen 3-colporate-
spheroidal; pistillode minute, trifid. Pistillate Dioecious or subdioecious evergreen or deciduous
flowers pedicellate; sepals and petals as in stami- trees or shrubs; indumentum 0. Leaves sometimes
nate; staminodes 6–8, filiform; ovary 4–5-locular; peltate or stipellate; stipules ribbed, deciduous.
stylodia 4 or 5, stigmatiform, bilobed. Fruit cap- Inflorescences axillary glomerules, pistillate flow-
sular, 4–5-locular, separating into 4–5 2-valved ers sometimes solitary. Staminate flowers pedicel-
cocci; columella not persistent. Seeds 1(2) per late; sepals 5, usually unequal; petals 5, longer than
locule, smooth (minutely striate); endosperm sepals, prominently veined; stamens 3–5, fila-
papyraceous, embryo small, cotyledons chloro- ments connate into a column; anthers extrorse in
phyllous, inflexed, much longer than the radicle. bud, dehiscing horizontally or deflexed; pollen
One sp., H. lucens Sim, eastern and southern grains subglobose, 3-colporate, colpi elongated,
Africa, disjunct along mountain ranges from Tan- exine reticulate; pistillode discoid, sessile or stipi-
zania to South Africa (Transkei and Natal). Levin tate atop the staminal column. Pistillate flowers
(1986b) suggested a close relationship between long-pedicellate [1.5 cm or more in fruit]; sepals
Heywoodia and Astrocasia, which is confirmed and petals similar to staminate, deciduous; disk
by the molecular evidence. Webster (1994) con- annular to cupular; ovary 3(4)-locular; stylodia
sidered Heywoodia as an ancient relict genus distinct, 2-fid. Fruit capsular, thin-walled; cocci
because he interpreted traits such as dioecy, the veiny; columella slender, persistent. Seeds 1 or
poorly differentiated perianth, the variability of 2 per locule; testa dry, smooth or roughened,
stamen and carpel number, and the small embryo raphe conspicuous; endosperm copious; cotyle-
as plesiomorphic. dons much longer and broader than radicle.
Six spp., Cuba and from Mexico south to
23. Chascotheca Urb. Bolivia and eastern Brazil.
Chascotheca Urb., Symb. Ant. 5: 14 (1904); Alain, Fl. Cuba

3: 44, fig. 8 (1953); Radcl.-Sm., Gen. Euphorb.: 26, fig. 3 3b. SUBTRIBE WIELANDIINAE Pax & K. Hoffm. (1922).
(2001).
Chaenotheca Urb. (1902; nom. illeg.). Three genera, western Indian Ocean, East Africa
and SE Asia.
Dioecious shrubs; indumentum simple. Leaves nar-
rowly peltate at base; stipules persistent. Staminate 25. Dicoelia Benth.
flowers in axillary glomerules, the pistillate solitary
Dicoelia Benth., Hook. Icon. Pl. 13, 70: t. 1289 (1879); Pax &
and axillary. Staminate flower subsessile; sepals 5,
K. Hoffmann, Pflanzenr. 147 XV: 15, fig. 3 (1922); Airy
imbricate, somewhat unequal; petals 0; disk annu- Shaw, Kew Bull. 36: 285 (1981).
lar; stamens 5, filaments connate at base; anthers
dorsifixed, dehiscing longitudinally; pollen grains Monoecious trees or shrubs; indumentum sim-
prolate, 3-colporate, exine coarsely reticulate; pis- ple. Stipules deciduous. Inflorescences axillary,
74 G.L. Webster
thyrsoid, pedunculate; glomerules often bisex- Morphologically, Chorisandrachne and Dicoelia

ual, staminate flowers several, pistillate 1/node. have little in common, and the seed coat of Chor-
Staminate flowers pedicellate; sepals 5, valvate, isandrachne shows some similarity with that of
shorter than petals; petals 5, valvate, convex, Chascotheca (Stuppy 1996).
apically thickened, with 2 paired cavities; disk
0; stamens 5, filaments distinct, basally adnate to 27. Wielandia Baill.
pistillode, exserted from perianth; anthers
Wielandia Baill., Étude Gén. Euphorb.: 568 (1858); P.
introrse, thecae fitted into paired cavities in the
Hoffmann, Adansonia III, 20: 333–340 (1998); P. Hoff-
petals in bud; pollen large, prolate, the exine mann & McPherson, Ann. Missouri Bot. Gard. 94:
with nearly continuous tectum; pistillode mas- 519–553 (2007), rev.
sive, apically 3–5-fid. Pistillate flowers pedicel- Savia Willd. sect. Wielandia (Baill.) M€ull. Arg. (1863).
late; sepals 5, valvate, shorter than petals; disk 0; Petalodiscus Baill. (1858).
ovary 3-locular, pubescent; ovules anatropous; Savia sect. Petalodiscus Baill. (1858).
Blotia Leandri (1957).
stylodia unlobed, elongated, erect. Fruits
spheroidal, capsular, thin-walled. Seeds with Monoecious trees or shrubs; indumentum simple
smooth testa; endosperm copious; cotyledons and scanty or 0. Petioles channeled or terete;
orbicular. stipules sometimes peltate, persistent or decidu-
One sp., D. beccariana Benth., Malaya, Sumatra, ous. Inflorescences axillary or cauline thyrses, or
Borneo. Kathriarachchi et al. (2005) provided flowers solitary; staminate and pistillate flowers
molecular support for the relationship of Dicoelia in the same or separate fascicles. Staminate flow-
to Chorisandrachne, and gave a detailed discussion ers pedicellate; pedicels inarticulate; sepals (4)5
of its peculiar, strongly autapomorphic appearance (6); petals (4)5(6), half as long to 3 x as long as
that, up to very recently, has camouflaged its true sepals; disk extrastaminal, glabrous (pubescent),
relationship. of antesepalous lobes, crenate, annular, or lacer-
ated, sometimes of different structure in stami-
26. Chorisandrachne Airy Shaw nate and pistillate flowers; stamens 5(6),
filaments distinct or partly connate; anthers
Chorisandrachne Airy Shaw, Kew Bull. 23: 40 (1969), 26:
323 (1972), Hooker’s Icon. Pl. 38: t. 3707 (1974). introrse; pistillode 3(5)-fid. Pistillate flowers ped-
icellate, perianth and disk as in staminate, ovary 3
Apparently dioecious trees or shrubs; indumen- (5)-locular; stylodia 3(5), distinct, 2-fid to
tum simple. Leaves at the base asymmetric, base, stigmas acute to slightly capitate. Fruits
shortly petiolate; stipules subulate, deciduous. solitary 3(5)-lobed schizocarps with irregular
Inflorescences axillary, flowers pedicellate, the dehiscence; columellae 3(5)-angled, base and
staminate ones in upper axils of branches soli- apex thickened; disk and perianth usually persis-
tary or in twos, the pistillate solitary in lower tent in fruiting stage. Seeds 1 or 2 per locule,
axils. Staminate flowers: sepals 5, broadly usually with distinct perichalazal annulus;
obovate, surmounted by the obtuse, unguiculate endosperm very sparse; embryo either with hemi-
petals; disk large, flat, subentire; stamens 5, fila- spherical to hemi-ovoid cotyledons and short
ments at the base connate, distally distinct, radicle, or with thin and folded cotyledons and
divaricate; anthers small, subglobose; pistillode prominent radicle.
narrowly cylindric. Pistillate flowers [incom- Thirteen spp., Madagascar (10 endemic), Sey-
pletely known]: sepals 5; petals 5, exceeding the chelles, Comoro Islands, and SE Kenya, most in
sepals, persistent; disk broadly cupulate, sinu- humid forest.
ous-pentagonous. Fruits capsular, subtended by
persistent disk and calyx; columella persistent. 4. TRIBE PHYLLANTHEAE Dumort. (1829).
Seeds 2 per locule, orbicular, plano-convex,
much flattened. Monoecious or dioecious trees, shrubs, or herbs;
A single sp., Ch. diplosperma Airy Shaw, south- leaves simple, alternate, entire, petioles not pul-
west Thailand. Both Webster (1994) and Radcliffe- vinate; inflorescences axillary or rarely on
Smith (2001) included Chorisandrachne in Lepto- branches; pedicels not articulated; sepals 4–6(7);
pus, but the molecular data (Kathriarachchi et al. petals 0 or very rarely +; disk + or 0; stamens
2005: 129) placed it as sister to Dicoelia beccariana. 2–15; anthers usually with thecae not separate
Euphorbiaceae 75
and connective not enlarged; ovary 2–6(15)- 4a. SUBTRIBE FLUEGGEINAE M€

ull. Arg. (1865).
locular; ovules hemitropous or rarely (Hetero-
savia) anatropous; fruits dehiscent or indehis- Dioecious; branches persistent; flowers in axillary
cent; seeds exarillate. glomerules or thyrses; sepals 4–7, imbricate, the
A tribe of worldwide distribution comprising pistillate persistent in fruit; petals 0; disk annu-
10 genera. lar; stamens 4–7, filaments distinct, anthers
extrorse; pollen grains prolate, 3-colporate, ora
circular, exine finely reticulate; pistillode +; ovary
KEY TO THE GENERA OF PHYLLANTHEAE 2–6-locular, ovules hemitropous; stylodia bifid;
fruits capsular or baccate; seeds ventrally invagi-
1. Phyllotaxy spiral or distichous on lateral axes but
deciduous branchlets not developed; pistillode mostly nated, testa dry or fleshy; endosperm copious,
+ in staminate flowers; seeds ventrally invaginated. 4a. embryo, curved.
Flueggeinae 2 This subtribe of four genera appears closely
– Phyllotaxy spiral on penultimate axes, distichous on related to the Phyllanthinae, and earlier (Webster
deciduous branchlets; pistillode 0; seeds with or without 1994) was more broadly circumscribed to include
ventral invaginations. 4b. Phyllanthinae 7
that group.
2. Petals +; petioles 2-winged to channeled adaxially
31. Heterosavia
– Petals 0; petioles terete 3 28. Plagiocladus Brunel ex Petra Hoffm. Fig. 12
3. Monoecious; stamens 15–21, inserted on disk; pistillate
calyx accrescent 30. Lingelsheimia Plagiocladus Brunel ex Petra Hoffm., Kew Bull. 61: 45
(2006).
– Dioecious or monoecious; stamens 2–7, not exserted
on disk; pistillate calyx not accrescent 4 Phyllanthus diandrus Pax (1904); Breteler Bull. Mus. Hist.
4. Monoecious; terminal branches distinctly flattened; nat. Paris IV, 12 B Adansonia: 293–295, fig. 1 (1991).
stamens 2 28. Plagiocladus Phyllanthus sect. Diandri Pax & K. Hoffm. in Engler &
– Dioecious; terminal branches not distinctly flattened Drude, Veget. der Erde 9 (Pflw. Afr. III, 2): 29 (1921).
(sometime slightly so in Margaritaria); stamens more Monoecious, glabrous shrub; terminal branches
than 2 5
5. Capsule with 1 seed per locule; staminate flowers sub-
distinctly flattened. Leaves shortly petiolate, entire;
sessile; flowers in racemoid paniculate thyrses; seed stipules unequally subcordate, deciduous. Inflor-
exotegmen vittate 32. Richeriella escences axillary; staminate flowers axillary glo-
– Capsule usually with 2 seeds per locule; staminate merules; pistillate long-pedicellate, solitary.
flowers pedicellate; flowers in axillary glomerules; Staminate flowers: sepals 6; disk flat, annular;
seed exotegmen palisadal 6 stamens 2, filaments distinct; anthers dehiscing
6. Pistillode + in staminate flowers; seed testa 2-layered,
exotegmen 3- or 4-layered, not woody; leaves spiral longitudinally. Pistillate flower sepals and disk as
33. Flueggea in staminate; ovary glabrous, 3-locular; stylodia
– Pistillode 0; seed testa 6–11-layered, distinctly fleshy; deeply 2-fid. Fruit a 3-carpellate schizocarp;
exotegmen 6–14-layered, + bony; leaves distichous endocarp hard; dehiscence explosive loculicidal,
29. Margaritaria septicidal and septifrague. Seeds deeply invagi-
7. Floral disk usually +, not completely adnate to calyx; nated (?).
pollen grains 3–4(6)-colporate/monoporate; seeds with
dry testa, not ventrally invaginated 34. Phyllanthus One sp., P. diandrus (Pax) Brunel ex Petra
– Floral disk 0 or completely adnate to calyx; pollen Hoffm., western C Africa, most common in
grains 3–6(16)-colporate/monoporate or diporate; Gabon. Plagiocladus is strongly supported as sister
seeds mostly not ventrally invaginated, with dry or to Margaritaria, but is easily distinguished from it
fleshy testa 8 by the difference in sepal and stamen number and
8. Stylodia unlobed; anthers apiculate; pollen grains 3–6-
the distinctly flattened terminal branches.
colporate-monoporate; ovary 3–8-locular; seed coat
usually fleshy 35. Glochidion
– Stylodia bifid or emarginate; pollen grains up to 16- 29. Margaritaria L. f.
colporate/diploporate; anthers not apiculate; ovary 3-
locular 9 Margaritaria L. f., Suppl. Pl.: 66 (1781); Webster, Ann.
9. Seed coat dry; staminate calyx discoid, not turbinate; Missouri Bot. Gard. 54: 217, fig. 2 (1968), J. Arnold Arb.
pollen grains 6–16-colporate 36. Sauropus 60: 403–444, figs. 11–13 (1979), rev.
– Seed coat fleshy; staminate calyx turbinate-truncate; pol- Dioecious (monoecious) trees or shrubs; indu-
len grains 4–12-colporate 37. Breynia mentum simple. Leaves evergreen or deciduous,
76 G.L. Webster
straight or curved, cotyledons longer than the

radicle. 2n ¼ 26.
Fourteen spp., extending through most tropi-
cal regions from America and the Caribbean
through Africa and Madagascar to tropical Asia
and N Australia.
30. Lingelsheimia Pax

Lingelsheimia Pax, Bot. Jahrb. Syst. 43: 317 (1909);
Mildbr., Wiss. Erg. Deut. Zentr.-Afr. Exped., Bot. 2: t. 54,
55 (1912); Pax & K. Hoffmann, Pflanzenr. 147. XV: 279
(1922); Léonard, Bull. Soc. Roy. Bot. Belg. 84: 49 (1951),
Bull. Jard. Bot. État. 32: 513 (1962); Radcl.-Sm., Gen.
Euphorb.: 49 (2001).
Danguyodrypetes Leandri (1939).
Aerisilvaea Radcl.-Sm. (1990).
Monoecious shrubs; indumentum 0. Stipules
persistent. Inflorescences axillary glomerules. Sta-
minate flowers pedicellate; sepals (4–)6–7, imbri-
cate; petals 0; disk glandular-lobed and intrusive
between filaments; stamens 15–35, filaments dis-
tinct; anthers basifixed; pollen grains somewhat
prolate, 3-colporate, ora circular, exine reticulate;
pistillode rudimentary or 0. Pistillate flowers pedi-
cellate; sepals 6, imbricate, persistent and accres-
cent in fruit; petals 0; disk massive, angled; ovary
3-locular, ovules anatropous [?]; stylodia distinct,
2-fid or -partite. Fruits capsular, dehiscing into
three 2-valved cocci or 6 valves. Seeds 1 per locule,
ventrally invaginated, testa smooth, dry.
Seven spp., tropical Africa (Zaire, Tanzania),
Fig. 12. Euphorbiaceae-Phyllanthoideae. Plagiocladus dia- and Madagascar. This genus was placed with Dry-
ndrus. A Branchlet with glomerules of staminate and petes and later with genera like Meineckia, but in
single pistillate flowers. B Leaf axil with stipule. C Pistillate
flower. D Staminate flower. E Same, only disk and stamens.
the molecular analyses of Kathriarachchi et al.
(Breteler 1990; drawn by W. Wessel-Brand) (2005, 2006) it has been recovered among the
basal genera of Phyllantheae. This placement is
also compatible with pollen morphology (K€ohler
distichous; stipules deciduous or somewhat per- 1965), leaf morphology (Levin 1986b), and seed
sistent. Inflorescences axillary glomerules, the pis- coat structure (Tokuoka and Tobe 2001).
tillate ones sometimes of solitary flowers.
Staminate flowers pedicellate; sepals 4, distinct,
biseriate; petals 0; disk annular, sometimes 31. Heterosavia (Urb.) Petra Hoffm.
slightly lobed; stamens 4, filaments usually dis- Heterosavia (Urb.) Petra Hoffm., Brittonia 60: 152 (rev.:
tinct; anthers extrorse; pollen subglobose, 3-col- 136–166) (2008).
Savia sect. Heterosavia Urb. (1902).
porate, semitectate; pistillode 0. Pistillate flowers
pedicellate; sepals 4, persistent in fruit; disk annu- Dioecious trees or shrubs; indumentum simple.
lar; ovary (2)3–4(6)-locular, ovules hemitro- Petioles 2-winged to channeled adaxially; stipules
pous; stylodia distinct or basally connate, 2-fid, persistent. Inflorescences axillary glomerules; sta-
often dilated. Fruit capsular, irregularly dehis- minate flowers 10–25 per inflorescence, subsessile;
cent, endocarp papery. Seeds 2(1) per locule, ven- pistillate flowers solitary, rarely in twos or threes.
trally invaginated; testa fleshy, bluish or greenish; Pedicels not articulated; staminate flowers: sepals
tegmen bony; endosperm copious; embryo and petals (4)5(6)-merous, sepals imbricate, petals
Euphorbiaceae 77
shorter than sepals; disk extrastaminal, annular, They differ significantly in inflorescence structure
crenate to entire, narrow, thick; stamens (4)5(6); and seed anatomy (Stuppy 1996; Tokuoka and
filaments distinct or connate for up to half of their Tobe 2001).
length; anthers introrse; pollen 3(long)-colporate,
tectate-perforate vermiculate; pistillode 3-fid to 33. Flueggea Willd.
base or nearly so, the branches erect; pistillate
flowers sepals, petals and disk as in staminate Flueggea Willd., Sp. Pl. 4: 637 (Fl€ uggea), 757 (Fluggea)
(1806); Webster, Allertonia 3: 259–312 (1984), rev.; Hay-
flowers but petals as long as or hardly shorter den, Brittonia 39: 268, fig. 1 (1987).
than sepals; staminodes sometimes +; ovary 3(4)
locular; stylodia 3(4), 2-fid to about half of their Dioecious (monoecious) shrubs or trees; branches
length; stigmas acute to obtuse. Fruits explosive sometimes spiny; indumentum mostly 0. Stipules
schizocarps, solitary, rarely in twos, subglobose, 3 persistent or deciduous. Inflorescences axillary
(4)-lobed, glabrous to pilose, dehiscence septici- glomerules. Staminate flowers pedicellate; sepals
dal, loculicidal and septifragous; columella 1.5–2 4–7, imbricate; petals 0; disk dissected, or seg-
times as long as wide; perianth persistent in fruit. ments confluent; stamens 4–7, filaments dis-
Seeds 2 per locule, smooth; endosperm copious; tinct; anthers extrorse, dehiscing longitudinally;
cotyledons straight, 3–4 times as long as radicle; pollen grains tricolporate, colpi short; pistillode
radicle 3–4 times as long as wide. 2–3-fid, rarely rudimentary or 0. Pistillate flowers
Four spp., Caribbean, from Florida to Swan pedicellate; sepals 4–7, imbricate, persistent in
Islands (Honduras). Recognized as distinct from fruit; petals 0; disk annular or lobed; ovary (2)3
Savia mainly on molecular evidence; distinguishing (4)-locular; ovules hemitropous; stylodia distinct,
morphological characters are subtle, and species 2-fid, sometimes dilated. Fruits capsular or bac-
identification may be problematic unless pistillate cate; columella persistent. Seeds usually 2 per loc-
material is available. ule, ventrally invaginated; testa smooth to
verruculose; endosperm copious; embryo straight
32. Richeriella Pax & K. Hoffm. to curved; cotyledons longer than radicle. 2n ¼ 26.
About fifteen spp. from the West Indies
Richeriella Pax & K. Hoffm., Pflanzenreich 147, XV: 30 through South America, tropical Africa and
(1922); Airy Shaw, Hook. Icon. Pl. 38: t. 3703 (1974),
Kew Bull. Add. Ser. 4: 190 (1975).
southeast Asia to the Pacific islands.
Dioecious trees; indumentum 0. Stipules decidu- 4b. SUBTRIBE PHYLLANTHINAE Pax (1890).
ous. Inflorescences axillary spiciform thyrses.
Staminate flowers sessile or subsessile; sepals 5, Monoecious or dioecious trees, shrubs, or herbs;
imbricate; petals 0; disk-segments 5; stamens 5, branching mostly "phyllanthoid" with main axis
filaments distinct, long-exserted; anthers dorsi- distally bearing scale-like leaves that subtend
fixed, extrorse; pollen grains subprolate, 3-colpo- leafy flowering deciduous branchlets; indumen-
rate, coarsely reticulate; pistillode 2–3-fid. tum simple or 0; leaves spiral on main axes,
Pistillate flowers subsessile in flower but pedi- distichous on deciduous branchlets; stipules usu-
cellate in fruit; sepals 5, imbricate; petals 0; disk ally persistent; flowers in axillary glomerules on
annular; ovary 3-locular; ovules anatropous?; sty- branchlets, sometimes cauliflorous; staminate
lodia distinct, 2-fid. Fruits capsular. Seeds mostly sepals (4)5(6), imbricate, entire or dentate; petals
1 per locule, ventrally invaginated, testa dry, 0; disk annular or segmented; stamens (1)2–4
smooth; endosperm scanty; embryo curved, coty- (10), filaments free or connate; anthers
ledons thin, longer than the radicle. extrorse, dehiscing horizontally to longitudinally;
One sp., R. gracilis (Merr.) Pax & K. Hoffm. [¼ pollen grains prolate to oblate, 3–6-colpate,
Flueggea gracilis (Merr.) Petra Hoffm.], S and SE porate, or panporate, exine reticulate; pistillode
Asia from India to Borneo. Richeriella and Flueg- 0; pistillate flowers pedicellate; sepals mostly 4–6;
gea are similar in leaf anatomy (Levin 1986a) and disk annular or segmented; ovary 3-locular,
pollen morphology (various studies), and the ovules anatropous to hemitropous; stylodia bifid
molecular results (Kathriarachchi et al. 2005: or entire; fruits mostly capsular (baccate or dru-
127) confirm the close relationship between them paceous); columella persistent; seeds usually 2/
but fail to prove that the one is nested in the other. locule, smooth to striate or ribbed.
78 G.L. Webster
This subtribe, with 5 genera and nearly 1,000

species, is the most species-rich taxon in the
Phyllanthoideae, due to the large size of Phyl-
lanthus. Most of the species, except in some sec-
tions of Phyllanthus, are characterized by the
unique phyllanthoid branching pattern (Webster
1956). The Phyllanthinae appear to have evolved
from the Flueggeinae, as suggested by the close
relationship between some species of Flueggea
and Phyllanthus subg. Isocladus.
34. Phyllanthus L. Fig. 13

Phyllanthus L., Sp. Pl. 981 (1753); M€ ull. Arg. in DC.,
Prodr. 15(2): 274 (1866), Fl. Bras. 11(2):23, t. 4–10
(1873); Hutch., Fl. Trop. Afr. 6(1): 692 (1912); Fawc. &
Rend., Fl. Jam. 4: 251, fig. 85 (1920); Pax & K. Hoffm., Nat.
Pflanzenfam. ed. 2, 19c: 60, figs. 28–30 (1931); Alain, Fl.
Cuba 3: 44, fig. 9 (1953); Leandri, Fl. Madag. 111(1): 30, t.
7–14 (1958); G.L. Webster, J. Arnold Arb. 37: 91–122,
217–256, 340–357 (1956); 38: 51–79, 170–198, 295–373
(1957); 49–100, 111–212, t. XIII–XXXII (1958), J. Arnold
Arb. 48: 332 (1967), Jablonski, Mem. N. Y. Bot. Gard. 17:
85, figs. 17–22 (1967); Webster, Ann. Missouri Bot.
Gard.54: 220; fig. 3 (1968); Bancilhon, Boissiera 18: 1–81
(1971); Radcl.-Sm., Fl. E. Trop. Afr., Euphorb. 1: 9, figs.
2–5 (1987); Santiago, Bradea 5(2): 44 (1988); Howard, Fl.
Less. Ant. 5: 70, fig. 34 (1989); Schmid, Fl. Nouv. Caléd. 17:
31, figs. 7–68 (1991); J.R. Wheeler, Fl. Kimberley Reg.: 619,
fig. 190 (1992); Webster, Ann. Missouri Bot. Gard. 81: 44
(1994); Friedmann, Fl. Seychelles, Dicot.: 362, figs.
100–103 (1994); Murillo & Franco, Euf. Reg. Araracuara
129, figs. 35–38 (1995); Hunter & Bruhl, Fl. Victoria 3: 74
(2000); Radcl.-Sm., Gen. Euphorb.: 38 (2001); Schatz,
Generic Tree Fl. Madag.: 63 (2001); Webster, Contr.
Univ. Michigan Herb. 23: 376, fig. 2 (2001), Novon 12:
290, figs. 1, 2 (2002); Li Bingtao & M.G. Gilbert, Fl. China
11: 180–190 (2008).
Niruri Adans. (1763).
Cicca L. (1767).
Kirganelia Juss. (1789).
Cathetus Lour. (1790).
Emblica Gaertn. (1790).
Nymphanthus Lour. (1790).
Epistylium Sw. (1800).
Eriococcus Hassk. (1843).
Macraea Wight (1852).
Fig. 13. Euphorbiaceae-Phyllanthoideae. Phyllanthus zor-
Hemicicca Baill. (1858). nioides. A Flowering branch system. B Staminate flower. C
Reverchonia A. Gray (1880). Stamens with glands. D Pistillate flower. E Distal (left) and
Diasperus L. ex Kuntze (1891). lower leaf. (Radcliffe-Smith 1996; drawn by J.M. Fothergill)
Phyllanthodendron Hemsl. (1898).
Monoecious or dioecious trees, shrubs, or herbs;
branching phyllanthoid or unspecialized; indu- floriferous deciduous branchlets; lamina pin-
mentum simple, unicellular or multicellular. nately veined, often inaequilateral at base; petioles
Leaves on penultimate axes alternate, spiral, mostly much shorter than blade; stipules persis-
those on ultimate axes spiral or distichous on tent or deciduous. Flowers in axillary glomerules
Euphorbiaceae 79
on persistent or deciduous branchlets, sometimes Monoecious (dioecious) trees or shrubs; branch-

cauliflorous. Staminate flowers pedicellate; sepals ing phyllanthoid. Leaves alternate, those on penul-
(4) 5–6, imbricate, entire or dentate; petals 0; disk timate axes spiral and reduced to cataphylls; leaves
usually dissected, sometimes annular; stamens (1) on deciduous branchlets distichous, often inaequi-
2–4(10), filaments distinct or connate, anthers lateral at base, entire; stipules persistent. Flow-
dehiscing horizontally to longitudinally, muticous ers in axillary glomerules, bracts persistent.
or apiculate; pollen grains prolate to oblate, 3–4 Staminate flowers pedicellate; sepals 5–9, imbicate;
(6)-colporate, rarely diploporate, porate, or pan- petals 0; disk 0; stamens 3–8, filaments connate;
porate, exine patterns various; pistillode 0. Pistil- anthers apiculate, extrorse, dehiscing longitudi-
late flowers pedicellate; sepals (4)5–6, imbricate, nally; pollen grains slightly prolate, 3–6-colpo-
usually entire, persistent in fruit; petals 0; disk rate-monoporate, reticulate; pistillode 0. Pistillate
annular or dissected; ovary 3-locular, ovules hemi- flowers pedicellate; sepals usually 6, imbricate,
tropous, inner integument thin, 3–5 cell layers; persistent in fruit; petals and disk 0; ovary 3–15-
stylodia bifid or entire, sometimes dilated. Fruits locular, ovules hemitropous; stylodia unlobed,
capsular, baccate, or drupaceous; columella usu- rarely emarginate or bifid, often connivent or con-
ally persistent in dehiscent fruits. Seeds usually nate. Fruit capsular; columella persistent. Seeds
2 per locule, trigonous, mostly not ventrally inva- usually 2 per locule, ventrally invaginated, testa
ginated; testa smooth, striate, ribbed, foveolate, or fleshy; endosperm copius; embryo curved, cotyle-
verruculose; endosperm copious; embryo straight dons much longer and broader than the radicle.
or curved, cotyledons broader than and equal to or 2n ¼ 26, 52.
longer than the radicle. 2n ¼ 16, 24, 26, 28, 52. Over 300 spp., Asia and Australasia; the spp.
Over 800 spp., pantropical, and also entering described from Madagascar appear of dubious
warm temperate regions. Molecular systematic affinity.
studies (Kathriarachchi et al. 2005, 2006) found
three of the eight subgenera of Phyllanthus to be 36. Sauropus Blume
polyphyletic, and the genus in its traditional cir-
cumscription to be paraphyletic. Sauropus, Brey- Sauropus Blume, Bijdr. Fl. Ned. Ind.: 595 (1826); Airy
Shaw, Hook. Icon. Pl. 38: t. 3708, 3709 (1974); Webster,
nia and Glochidion are deeply embedded within Ann. Missouri Bot. Gard. 81: 46 (1994); J.T. Hunter &
Phyllanthus s. str., and ultimately may be included Bruhl, Austrobaileya 4: 661 (1997); Philcox, Fl. Ceylon
into an expanded generic concept that would com- 13: 99 (1999); Radcl.-Sm., Gen. Euphorb.: 46 (2001); van
prise over 1,250 species. At present, however, the Welzen, Blumea 48: 319–391 (2003), Males. & Thai spp.;
Pruesepan et al., Ann. Bot. 102: 1007–1018 (2008), mol.
nomenclatural changes conforming to the molec-
syst.
ular results hardly have been initiated, and the Aalius Rumph. ex Lam. (1793).
non-monophyletic subtaxa of Phyllanthus s. str. Ceratogynum Wight (1852).
and the genera deeply embedded in it will require Diplomorpha Griff. (1854) not Meissn. (1841).
careful taxonomic revision of the different ele- Synostemon F. Muell. (1858).
ments before being amalgamated in the expanded Breyniopsis Beille (1925).
Heterocalymnantha Domin (1927).
generic concept, as has already been initiated for
Sauropus by the work of van Welzen (2003), Sagun Monoecious shrubs, subshrubs, or herbs; branch-
and van der Ham (2003), and Pruesapan et al. ing phyllanthoid or phyllotaxy entirely spiral;
(2008). At present, merging Phyllanthus with its leaves on deciduous branchlets alternate, disti-
embedded genera would displace the lack of a chous; lamina pinnately veined, margins entire;
workable taxonomic structure for the expanded stipules persistent. Flowers in axillary glomerules.
genus only to the infrageneric level. Staminate flowers pedicellate; sepals 6, connate,
the calyx sometimes flattened and discoid; petals
35. Glochidion J.R. & G. Forst. and disk 0 but sometime inflexed sepal tips mim-
icking a disk; stamens 3, filaments connate into a
Glochidion J.R. & G. Forst., Char. Gen. Pl.: 57 (1775), nom. column; anthers sessile, extrorse, dehiscing lon-
cons.; A.C. Sm., Fl. Vit. Nova 2: 467 (1981); McPherson &
Tirel, Fl. Nouv. Caléd. 17: 18, t. 4 (1991); Webster, Ann. gitudinally; pollen grains oblate, 6–16-colpo-
Missouri Bot. Gard. 81: 46 (1994); Florence, Fl. Polynésie rate-diploporate, reticulate; pistillode 0. Pistillate
Française 1: 66, figs. 9–16 (1997). flowers pedicellate; sepals 6, connate, persistent
80 G.L. Webster
and sometimes accrescent in fruit; petals and disk Molecular data suggest a reclassification as indi-
0; ovary 3-locular, ovules hemitropous; stylodia cated under the previous genus.
bifid. Fruit capsular but pericarp often fleshy;
columella persistent. Seeds 2 in each locule, tri- IB. SUPERTRIBE ANTIDESMODAE G.L. Webster,
gonous, hilum linear, within a large ventral invag- supertrib. nov.1
ination, testa dry; endosperm copious; embryo
straight or curved, cotyledons broader and longer Subfam. Antidesmatoideae Hurus. (1954).
than radicle. 2n ¼ 24. Antidesmatinae Pax (1890), "Antidesminae".
Traditionally comprising about 80 spp. in Dioecious or rarely monoecious; leaves simple
tropical Asia, Malesia, and Australia except for (compound: Bischofia), entire (subentire or
the widespread Sauropus bacciformis. The toothed); leaf epidermis usually with tanninifer-
molecular data of Pruesapan et al. (2008) suggest ous cells; inflorescences usually thyrsoid with
the splitting of Sauropus into two clades, the distinct axes; flowers usually apetalous; ovary
mainly southeast Asian Sauropus s. str., which locules 1–5; ovules anatropous; fruits indehiscent
should be united with Breynia under the latter or tardily dehiscent (explosively dehiscent). Tro-
name, and the mainly Australian Sauropus, for pics and subtropics.
which the former name Synostemon should be This supertribe corresponds to the tannini-
reinstated. ferous clade of Wurdack et al. (2004); in molecu-
lar analyses (Kathriarachchi et al. 2005; Samuel
37. Breynia J.R. & G. Forster et al. 2005), it is placed sister to Phyllanthodae.
Breynia J.R. & G. Forst., Char. Gen. Pl.: 73 (1775; nom.
cons.); Airy Shaw, Kew Bull. Add. Ser. 4: 61 (1975), 8: 38 5. TRIBE ANTIDESMATEAE Benth. (1873).
(1980); McPherson & Tirel, Fl. Nouv.-Caléd. 17: 14, fig. 3,
6–10 (1991); Webster, Ann. Missouri Bot. Gard. 81: 46 Dioecious (monoecious); exudate 0; indumentum
(1994); Philcox, Fl. Ceylon 11: 238 (1997); Radcl.-Sm.,
Gen. Euphorb.: 46 (2001). simple (lepidote); petioles sometimes pulvinate;
Foersteria Scop. (1777). foliar glands and domatia 0 or +; sepals (3)4–6
Melanthesa Blume (1826). (8), imbricate to nearly completely connate;
Melanthesopsis M€ ull. Arg. (1863). petals 0 or [Thecacoris] +, small; disk + or 0;
Monoecious shrubs; branching phyllanthoid; stamens 2–8(13); thecae usually separate and
indumentum 0. Leaves on penultimate axes spi- connective enlarged; pistillode + (0); ovary
ral, reduced to scales, those on deciduous branch- locules (1)2–4(5); ovules anatropous; stylodia
lets alternate, distichous; lamina entire; stipules + or 0; fruits indehiscent or dehiscent; seeds
persistent. Flowers in axillary glomerules, the exarillate, usually albuminous.
pistillate solitary. Staminate flowers pedicellate; Eight genera, pantropical, in five apparently
sepals 6, connate into a turbinate cup, sepal tips isolated mono- or digeneric subclades (Kathriar-
inflexed; petals and disk 0; stamens 3, filaments achchi et al. 2005; Samuel et al. 2005).
connate; anthers basifixed, muticous, dehiscing
longitudinally; pollen grains 4–12-colporate- KEY TO THE GENERA OF ANTIDESMATEAE
diploporate; pistillode 0. Pistillate flowers pedi- 1. Ovary 2-locular above, appearing 4-locular below;
cellate; sepals 6, nearly distinct to connate, imbri- fruits 4-gonous; pollen with atria 42. Martretia
cate, persistent and sometimes accrescent in fruit; – Ovary locules not as above; fruits commonly 1–3-
petals and disk 0; ovary 3-locular, rounded to gonous; pollen grains exatriate 2
turbinate, ovules hemitropous; stylodia distinct 2. Fruit winged; floral disk 0; tanninferous cells in leaf
or basally connate, emarginate to bifid. Fruits epidermis 0 3
somewhat fleshy, capsular; columella usually not
persistent. Seeds 2 in each locule, trigonous, ven-
trally invaginated, testa fleshy; endosperm copi- 1
Supertribe Antidesmodae G.L. Webster, supertrib. nov.,
ous; embryo curved, cotyledons equaling or floribus inflorescentiis paniculatis spicatisve, cellulis epi-
longer than radicle. 2n ¼ 26, 52. dermalibus foliorum saepe tanniniferis, embryonibus pler-
umque chlorophyllosis, radicula quam cotyledonibus
Traditionally comprising about 35 spp., many breviore, fructibus plerumque baccatis vel drupaceis.
difficult to distinguish, east Asia and Australasia. Type: Antidesma L.
Euphorbiaceae 81
– Fruit not winged; floral disk + or 0; tanninferous cells 39. Thecacoris A. Juss. Fig. 14
in leaf epidermis + 4
3. Fruit capsular; inflorescence paniculate; stylodia Thecacoris A. Juss., Euphorb. Tent.: 12 (1824); Léonard,
lacerate 40. Didymocistus Bull. Jard. Bot. Nat. Belg. 64: 13–52 (1995).
– Fruit samaroid; inflorescence racemoid; stylodia papil- Cyathogyne M€ ull. Arg., Flora 47: 536 (1864).
lose 41. Hymenocardia
4. Indumentum lepidote; fruit indehiscent
Dioecious or rarely monoecious trees, shrubs,
44. Hieronyma subshrubs or perennial herbs; indumentum sim-
– Indumentum 0 or of simple or stellate hairs 5 ple. Leaves entire or repand; stipules deciduous
5. Ovary 1-locular; fruit indehiscent 38. Antidesma or persistent. Inflorescences axillary, sometimes
– Ovary 2–5-locular; fruit dehiscent 6 fasciculate, racemoid or spiciform; bracts persis-
6. Disk 0; pistillode 0; ovary 4–5-locular tent, 1-flowered. Staminate flowers sessile or ped-
45. Leptonema icellate; sepals 5(6), imbricate; petals 5, small or 0;
– Disk +; pistillode +; ovary 3–4-locular 7 disk segments 5, distinct; stamens 5, filaments
7. Dioecious; ovary 3-locular; stylodia 2-fid; endosperm
copious 39. Thecacoris
distinct; anthers pendent; anther thecae discrete,
– Monoecious; ovary 4-locular; stylodia entire; seeds parallel and pendulous at first, later divaricate
exalbuminous; embryo massive 43. Apodiscus and erect; connective enlarged; pollen grains per-
prolate, 3-colporate, pores lalongate, exine finely
reticulate; pistillode massive, dilated. Pistillate
5a. SuBTRIBE ANTIDESMATINAE M€
ull. Arg. (1865).
flowers pedicellate, pedicel geniculate in fruit;
Two genera, Old World. sepals 5, imbricate, persistent in fruit; petals vari-
able in number, sometimes 0; disk annular; ovary
38. Antidesma Burm. ex L. 3(4)-locular; stylodia distinct or connate at base,
bifid. Fruits capsular, septicidal; columella persis-
Antidesma Burm. ex L., Sp. Pl.: 1027 (1753); Airy Shaw,
Kew Bull. Add. Ser. 4: 207 (1975), 8: 208 (1980); Léonard,
tent. Seeds usually 1 per locule; testa dry and
Fl. Afr. Centr. Euph. (2): 16–40 (1995); Chakrabarty & shiny; endosperm copious; embryo greenish,
Gangopadkyay, J. Econ. Tax. Bot. 24: 1 (2000). cotyledons flat, much longer and broader than
Stilago L. (1767). radicle.
Dioecious trees and shrubs; indumentum simple. About 25 spp., four in Madagascar and the
Leaves sometimes domatiiferous; marginal rest in tropical Africa from Sierra Leone to
glands rarely + (A. vaccinioides); stipules mostly Angola and Tanzania. In the concept of Léonard,
entire. Inflorescences axillary or terminal, some- Cyathogyne includes five herbaceous species of
times fasciculate. Flowers in catkin-like, some- Africa and Madagascar. Webster (1994) and Rad-
times branched thyrses; bracts 1-flowered. cliffe-Smith (2001) followed Leandri (in Fl.
Staminate flowers sessile to pedicellate; calyx cup- Madag., 1958) in reducing Cyathogyne to a sec-
ular or 3–5(8)-lobed, sepals imbricate; petals 0; tion of Thecacoris, with Th. usambarensis bridg-
disk segments distinct or connate; stamens ing the gap between the two genera. Léonard’s
mostly (2)3–5(6); filaments distinct; anthers 2- case (1995) for maintaining Cyathogyne distinct
lobate, thecae distinct, divergent; connective may find support in the pollen and seed coat
enlarged; pollen grains perprolate, 3-colporate, structure (K€ohler 1965; Stuppy 1996).
pores conspicuously lalongate, exine tectate-
punctate, scabrate; pistillode small or 0. Pistillate 5b. SUBTRIBE HYMENOCARDIINAE P. Hoffm. (2006).
flowers pedicellate; calyx usually cupular-lobed, Two genera, one neotropical, the other African.
persistent in fruit; disk annular or cupular; ovary
1(2)-locular; stylodia usually 2, bifid. Fruit drupa- 40. Didymocistus Kuhlm.
ceous, asymmetric and flattened, endocarp retic-
ulate or foveolate. Seeds 1(2) per fruit; endosperm Didymocistus Kuhlm., An. Prim. Reun. Sud-Amer. Bot. 3:
82 (1940); Radcl.-Sm., Gen. Euphorb.: 76, fig. 7 (2001).
fleshy, not copious; embryo flat, cotyledons much
longer and broader than radicle. 2n ¼ 26. Dioecious trees or shrubs; indumentum simple.
At least 150 spp., mostly from India to Mal- Leaves with sessile abaxial glands; stipules decid-
esia, southern Japan, Australia, and the Pacific uous. Inflorescences terminal, paniculate; bracts
islands, only 10 in Africa/Madagascar. deciduous. Staminate flowers subsessile; sepals 5,
82 G.L. Webster
Peru; seed dispersal probably hydrochorous

(Wurdack et al. 2004).
41. Hymenocardia Wall. ex Lindl. Fig. 15

Hymenocardia Wall. ex Lindl., Nat. Syst. ed. 2, 441 (1836);
J.G. Adam, Mém. Mus. natl. Hist. Nat., N.S. Bot. 20: 481, t.
174 (1971); Léonard & Mosango, Fl. Afr. Centr., Hymeno-
cardiaceae: 2 (1985); Radcl.-Sm., Fl. Zamb. 9: (4): 113, fig.
18 (1996).
Dioecious trees or shrubs; indumentum simple.
Leaves abaxially with domatia and laminar
glands; stipules deciduous. Staminate inflores-
cences axillary, spicate or catkin-like; pistillate
flowers solitary or in axillary or terminal, race-
moid inflorescences. Staminate flowers subses-
sile; sepals (4)5(8), slightly imbricate, connate
into a dentate or lobed cup; petals 0; disk 0;
stamens usually 5, filaments distinct; anthers
introrse in bud, anther sacs discrete, dehiscing
longitudinally; pollen grains suboblate, 3-porate
or brevicolporate, pores marginate, exine sca-
brate, tectum rugulose; pistillode cylindric or 2-
lobed. Pistillate flowers pedicellate; sepals 5
(4–9), distinct, deciduous; petals 0; disk 0;
ovary 2-locular, flattened perpendicular to sep-
tum; stylodia 2, unlobed. Fruits samaroid,
dehiscing into two 1-seeded winged indehiscent
mericarps; columella persistent. Seeds com-
pressed, testa striate, shiny; endosperm copious;
embryo flattened, cotyledons longer and broader
than radicle. 2n ¼ 26, 28.
Six or seven spp., all African except one in
Fig. 14. Euphorbiaceae-Phyllanthoideae. Thecacoris tricho- southeast Asia. A very distinctive genus that has
gyne. A Branch tip with pistillate flowers. B Infructes- been accepted as a separate tribe or even family
cence. C Staminate flower. D Staminate disk glands and by Airy Shaw (1965), Léonard and Mosango
petals. E Pistillate flower. F Fruit. (Radcliffe-Smith 1996;
drawn by J.M. Fothergill) (1985), and Meeuse (1990). The ament-like inflor-
escences and unique samaroid fruits make it one
imbricate, distinct; petals 0; disk 0; stamens 5, of the more aberrant genera of Phyllanthoideae.
filaments distinct, exserted, anthers introrse, dor- However, the pollen evidence weighted so heavily
sifixed; pollen grains globose, 3-colporate, exine by Léonard must be seen in connection with the
scabrate, tectum rugulose; pistillode 2–3-lobed; complete anemophily found in the genus. In con-
pistillate flowers sessile or subsessile; calyx 5- trast to the pollen data, the wood anatomy of
lobed, petals 0; disk 0; ovary 2-locular, stylodia Hymenocardia, according to Mennega (1987), is
unlobed, plumose. Fruits capsular, valves thin, compatible with genera of Antidesmeae, and the
inflated, deciduous; columella persistent. Seeds foliar venation studies of Levin (1986a, b) show
hemitropous, endosperm copious; embryo no incompatibility with Phyllanthoideae. The
straight, cotyledons much longer and broader placement of Didymocistus with Hymenocardia
than radicle. appears strongly supported by the similarities of
One sp., D. chrysadenius Kuhlm., in flood- pollen structure demonstrated by Simpson and
plain forests of Amazonian Brazil, Colombia and Levin (1994), the possession of multicellular
Euphorbiaceae 83
42. Martretia Beille

Martretia Beille, C. R. Séances Acad. Sci. Paris 145: 1294
(1907), Bull. Soc. Bot. France 55 (Mém. 8b): 64 (1908);
Léonard, Bull. Jard. Bot. Nat. Belg. 59: 319 (1989); Radcl.-
Sm., Gen. Euphorb.: 80 (2001).
Dioecious; indumentum 0 on stems and leaves.
Tanniniferous epidermal cells 0. Stipules entire,
deciduous. Inflorescences axillary, spiciform to
racemoid, sometimes compound. Staminate flow-
ers pedicellate; sepals 4 or 5, distinct; petals 0;
disk 0; stamens 4–8, filaments distinct; anthers
latrorse, dehiscing longitudinally, connective
glandular at the apex and pubescent; pollen sub-
spheroidal, 3-colporate, angulaperturate, colpi
narrow, unbordered, exine semitectate, vermicu-
late-rugulose; pistillode 2-fid, pubescent apically.
Pistillate flowers pedicellate; sepals 4–6, not per-
sistent in fruit; petals 0; disk of 5 or 6 minute
segments; ovary 2-locular, appearing 4-locular
below due to intercalary partitions, glabrous or
puberulent; stylodia 2, basally connate, unlobed,
subulate, elongated. Fruits capsular, 4-gonous,
dehiscing into four 1-seeded segments. Seeds
albuminous, ellipsoid; testa smooth, fleshy; coty-
ledons suborbicular.
One sp., M. quadricornis Beille, from W and C
Africa (Sierra Leone to C.A.R., Congo and Gabon),
in swamp and riverine forests. Unique in Phyl-
lanthoideae for its pollen apertures and false-par-
titioned fruit. Beille (1908) suggested a placement
in Antidesmeae, and it was placed in Antidesminae
by Pax and K. Hoffmann (1922, 1931).
Fig. 15. Euphorbiaceae-Phyllanthoideae. Hymenocardia
ulmoides. A Branchlet with staminate flowers. B Stami- 43. Apodiscus Hutch.
nate flower. C Stamen, rear view, showing gland. D
Branchlet with pistillate flowers. E Pistillate flower. F Apodiscus Hutch., Bull. Soc. Bot. France 58, Mém. 8: 205
Fruiting branch. G Fruit. (Radcliffe-Smith 1996; drawn (1912), and in Hook. Ic. Pl. 31: t. 3032 (1915); Keay, Fl. W.
by Pat Halliday). Trop. Afr. ed. 2, 1: 373 (1958); Radcl.-Sm., Gen. Euphorb.:
63 (2001).
secretory leaf glands in the absence of tanninifer- Monoecious trees; indumentum simple, confined
ous leaf epidermal cells, the tanniniferous endo- to the inflorescence. Leaves deciduous; stipules
tegmic seed coat (Tokuoka and Tobe 2001), and deciduous. Inflorescences axillary, spiciform, fas-
the molecular data of Wurdack et al. (2004) and ciculate; bracts uniflorous; pistillate flowers 1 or
Kathriarachchi et al. (2005). 2 at base of inflorescence. Staminate flowers ses-
sile; sepals 5, imbricate; petals 0; disk segments 5,
5c. SUBTRIBE MARTRETIEAE P. Hoffm. (2006). pubescent; stamens 5, filaments distinct; anthers
introrse, thecae subglobose, longitudinally dehis-
Two genera held together by strong molecular cent, connective slightly produced; pollen prolate,
support; they had been of dubious affinity and tricolporate, colpus transversalis elongate; pistil-
have little in common morphologically. lode depressed-globose, pubescent. Pistillate flow-
ers pedicellate; sepals 5, imbricate; petals 0; disk 0;
84 G.L. Webster
ovary (3)4(5)-locular; stylodia unlobed, recurved. Dioecious (monoecious) shrubs; indumentum

Fruits capsular, septicidally dehiscent into 3 cocci. simple. Leaves circular to heart-shaped; stipules
Seeds exalbuminous; embryo massive, cotyledons persistent. Inflorescences axillary, staminate
much longer and broader than radicle. racemoid or corymbiform, pistillate brachyblas-
One imperfectly known sp., A. chevalieri tic; bracts subulate; flowers 1 per bract. Staminate
Hutch., from W Africa (Guinea, Liberia and flowers pedicellate; sepals 5, connate at base,
Sierra Leone); it appears aberrant in the Antides- imbricate; petals 0; disk 0; stamens 5, filaments
mateae because of its monoecy and exalbuminous distinct, capillary, exserted; anther connective
seeds. globose, glandular; thecae separate; pollen sacs
pendulous; pollen grains 3-colporate, pores
5d. SUBTRIBE HIERONYMINAE M€
ull. Arg. (1865). transversely elongate; exine very finely reticulate;
pistillode 0. Pistillate flowers pedicellate; sepals 5,
Monotypic. persistent in fruit; petals 0; disk 0; ovary 4–5-
locular; stylodia bifid. Fruits capsular, splitting
44. Hieronyma Allemão into 4–5 2-valved cocci; columella persistent.
Hieronyma Allemão, Pl. Nov. Bras.: 22 (1848); Pax & K. Seeds 2 per locule, testa verruculose.
Hoffm., Pflanzenreich 147, XV: 31, fig. 8 (1922); Jablonski, Two spp. endemic to Madagascar, still poorly
Mem. N. Y. Bot. Gard. 17: 122 (1967); Webster, Ann. Mis- known.
souri Bot. Gard. 75: 1094 (1988) and ibid. 81: 52 (1994);
Franco, Bot. Jahrb. Syst. 111: 297, figs. 1–16 (1990).
Hyeronima Allemão (orth. var.) 6. TRIBE SCEPEAE Horan. (1847).
Dioecious trees or shrubs; indumentum lepidote Aporuseae (Lindl. ex Miq.) Airy Shaw (1974).
or rarely simple. Stipules usually small, cochleate,
often soon deciduous. Inflorescences axillary, Dioecious; exudate 0; leaves often with laminar
compound racemoid to subspicate thyrses; glands; inflorescences axillary or terminal, some-
bracts inconspicuous. Staminate flowers subses- times cauliflorous; sepals 4–5(7), distinct; petals
sile; calyx 4- or 5-lobed; petals 0; disk pulvinular 0; disk lobed to dissected or 0; pollen grains 3-
or cupular; stamens 3–6, filaments distinct; colporate, reticulate or tectate; pistillode +; ovary
anthers introrse; thecae divergent; anther sacs 2–5-locular; stylodia bifid or unlobed; fruit cap-
pendent, dehiscing longitudinally; pollen perpro- sular or baccate; seed coat dry or fleshy.
late, 3-colporate, pores elliptic, exine tectate-per- This subtribe of eight genera, all paleotropical
forate or microreticulate; pistillode columnar. except for Richeria, was subsumed within the
Pistillate flowers pedicellate; calyx 4- or 5-lobed, Antidesminae by Pax (1924) and Pax and K.
persistent in fruit; petals 0; disk cupular; ovary Hoffmann (1922, 1931). Airy Shaw (1974) was
2(3)-locular, glabrous or lepidote; inner integu- the first to formally recognize it, as tribe Aporu-
ment thin [2–4 layers]; stylodia very short, stig- seae, which was accepted by Webster (1975) and
matoid. Fruits drupaceous, 1–2-locular by more recently by Thin (1995).
suppression; exocarp fleshy, endocarp hard and
bony or fibrous. Seeds usually 1 per fruit by KEY TO THE GENERA OF SCEPEAE
abortion; endosperm fleshy; embryo green, coty- 1. Pistillate disk dissected; stylodia 2-lobate or 2-fid
ledons much longer and broader than radicle. 46. Protomegabaria
Approximately 15(40?) spp., from S Mexico – Pistillate disk cupular or 0; stylodia 2-fid or unlobed
and Cuba south to Bolivia and S Brazil. 2
2. Staminate disk +; pistillate disk + or 0 3
5e. SUBTRIBE LEPTONEMATINAE M€uLL. ARG. (1865). – Staminate and pistillate disk 0 6
3. Pistillate disk +; sepals persistent in fruit 4
Monotypic. – Pistillate disk 0; sepals deciduous 5
4. Capsule loculicidal; staminate flowers mostly 1 per
bract 51. Maesobotrya
45. Leptonema A. Juss. – Capsule septicidal; staminate flowers several per bract
Leptonema A. Juss., Euphorb. Tent.: 19 (1824); Leandri, 47. Richeria
Fl. Madag. 111 (1): 12, fig. IV, 1–6 (1958); Schatz, Generic 5. Staminate sepals distinct, shorter than stamens; stami-
Tree Fl. Madag.: 157 (2001). nodes usually 0; stipules narrow 52. Baccaurea
Euphorbiaceae 85
– Staminate sepals basally connate, longer than stamens; and exserted from calyx; anthers introrse, versa-
staminodes + in pistillate flower; stipules broad tile; pollen grains prolate, 3-colporate, semitec-
53. Nothobaccaurea tate-reticulate; pistillode cylindric, hirtellous.
6. Leaves spiral; stamens 5 or 6; pistillode massive, peltate
50. Ashtonia Pistillate flowers pedicellate; calyx 3–5-lobed,
– Leaves spiral or distichous; stamens mostly 2 [unknown sepals imbricate; petals 0; disk cupular; ovary
in Distichirrhops]; pistillode small or 0 2–3-locular, sericeous; stylodia 2- or 3-fid. Fruit
7 capsular, tardily dehiscent septicidally; columella
7. Leaves spiral; pistillate flowers subtended by 1 bract; flattened, winged, persistent. Seeds 1 per locule,
pistillate disk + but obscure; pedicels not articulated
ovoid; testa fleshy, exotegmen vittate; endo-
48. Aporosa
– Leaves distichous; pistillate flowers subtended by 3 sperm copious; cotyledons flat, much longer
bracts; pistillate disk 0; pedicels articulated than radicle, both cotyledons and radicle chloro-
49. Distichirrhops phyllous.
Two or three spp., distributed from the Lesser
46. Protomegabaria Hutch. Antilles and Panama to Bolivia and Brazil.
Protomegabaria Hutch., Hook. Icon. Pl. 30: t. 2929 (1911);
Léonard, Bull. Jard. Bot. Nat. Belg. 64: 53–63 (1995), Fl. 48. Aporosa Blume
Afr. Cent., Euph. 2: 79, t. 13 (1995). Aporosa Blume, Bijdr.: 514 (1826; orth. cons.); Airy Shaw,
Dioecious, semipachycaul trees; trunk buttressed Hook. Icon. Pl. 38: pl. 3701 (1974), Kew Bull. Add. Ser. 4:
30 (1975), ibid. 8: 28 (1980); Schot, Blumea 40: 449 (1995).
and stilt-rooted; indumentum simple or 0. Stipules Scepa Lindl. (1836).
fugacious. Inflorescences racemose to subspicate,
axillary, sometimes cauliflorous, solitary or clus- Dioecious trees and shrubs [flowers bisexual in
tered; pedicels articulate; staminate flowers in four spp.]; indumentum simple. Leaves often
clusters partly enclosed by cupular central bractlet; with laminar glands, petioles long, often genicu-
pistillate flowers 1 per bract. Staminate flowers: late and distally glandular; stipules mostly persis-
sepals 5(4), imbricate; petals 0; disk segments tent, sometimes foliaceous or falcate.
usually 5, fleshy; stamens usually 5, distinct; Inflorescences usually axillary, often fasciculate,
anthers introrse; pollen grains prolate, 3-colporate, the staminate spiciform and catkin-like, the pis-
pores laterally elongated, exine finely reticulate; tillate racemoid. Staminate flowers sessile or sub-
pistillode columnar, small. Pistillate flowers: sepals sessile; sepals 3–6, imbricate; petals 0; disk 0;
5, imbricate, persistent in fruit; petals 0; disk annu- stamens 2(5), filaments distinct; anthers sub-
lar; ovary 3(4)-locular; stylodia 2-lobate or 2-fid. globose, basifixed; pollen grains subprolate, 3-
Fruit capsular, somewhat lignified; columella per- colporate, pores lalongate, exine semitectate-
sistent, dilated at base and clavate distally. Seeds reticulate; pistillode minute or 0. Pistillate flowers
ellipsoid, testa smooth and shiny; endosperm pedicellate; sepals 4 or 5, imbricate, persistent in
copious; embryo straight. 2n ¼ 26. fruit; petals 0; disk small; ovary 2(4)-locular;
Three African spp. (Guinea to Gabon and stylodia usually bipartite, often papillose or
Congo), in lowland tropical rainforest. laciniate. Fruit capsular; pericarp leathery,
dehiscing into valves or irregularly; columella
47. Richeria Vahl persistent. Seeds 1 or 2 per fruit, with colored
sarcotesta; endosperm copious; embryo chloro-
Richeria Vahl, Eclog. Amer. 1: 30, t. 4 (1797); Secco &
Webster, Bol. Mus. Para. Emilio Goeldi, N.S., Bot. 6:
phyllous, straight, cotyledons much longer and
141–158 (1990), rev. broader than radicle. 2n ¼ 26, 52.
About 75 spp., distributed from India and Sri
Dioecious trees or shrubs; indumentum simple Lanka to the Philippines and New Guinea.
or 0. Leaves entire or crenulate, sometimes glan-
dular near the base; stipules deciduous. Inflores- 49. Distichirrhops Haegens
cences racemoid or spicate, axillary; staminate
flowers in dense glomerules; pistillate flowers Distichirrhops (’Distichirhops’) Haegens, Blumea Suppl.
12: 193, figs. 3.28, 3.29 (2000).
1 per bract. Staminate flowers sessile; calyx
3–5-lobed, tips of lobes imbricate; petals 0; disk- Dioecious trees; indumentum simple. Leaves dis-
segments 3–5; stamens 3–6, filaments distinct tichous, entire, with marginal glands; petioles
86 G.L. Webster
apically pulvinate; stipules deciduous. Inflores- small, 1–3, or more often 0; disk intrastaminal,
cences axillary or ramiflorous, thyrsoid [only pis- segments confluent, usually pubescent;
tillate plants known], unbranched, 10–15- stamens 4–6(7), filaments distinct, usually
flowered. Pistillate flowers in triads, each sub- exserted from calyx, anthers introrse, connective
tended by 3 bracts, with articulated pedicels; not enlarged; pollen grains subprolate, 3-colpo-
sepals 4 or 5, deciduous to persistent; petals, rate, pores elliptic, exine semitectate, finely
staminodes and disk 0; ovary 2–4-locular; ovules reticulate; pistillode cylindric or clavate, pubes-
2 per locule; stylodia unlobed to 2-fid. Fruits 0–4- cent. Pistillate flowers: sepals and petals as in
seeded, baccate or capsular and tardily dehiscent, staminate; disk cupular, sometimes lobed, rim
often glandular, 0–4-seeded. Seeds albuminous, usually ciliate; ovary 2–3(4)-locular, glabrous or
with fleshy testa. slightly pubescent; stylodia bifid. Fruits capsular,
Three incompletely known spp., Borneo and dehiscing loculicidally; pericarp thin; columella
New Guinea. According to Haegens, the genus not persistent. Seeds 1 or 2 per locule; testa fleshy
seems to be related to Aporosa. and often blue; endosperm copious; embryo chlo-
rophyllous, cotyledons flat, much longer than
50. Ashtonia Airy Shaw radicle. 2n ¼ 26.
Some 20 spp., mostly in the West Africa/
Ashtonia Airy Shaw, Kew Bull. 21: 357 (1968), Hook. Icon.
Pl. 38: t. 3702 (1974), Kew Bull. Add. Ser. 4: 42 (1975).
Congo lowland rainforest area, one in Uganda
and another in Zambia.
Dioecious trees; indumentum 0. Leaves bigland-
ular at base; stipules deciduous. Inflorescences 52. Baccaurea Lour.
spiciform [staminate] and racemoid [pistillate],
Baccaurea Lour., Fl. Cochinch.: 661 (1790); Haegens, Blu-
solitary; staminate flowers in glomerules, pistil- mea Suppl. 12: 80–129, figs. 3.1–3.27 (2000).
late solitary. Staminate flowers subsessile; sepals
3 or 4, imbricate; petals 0; disk obscure or 0; Dioecious trees; indumentum simple or stellate.
stamens 5 or 6, filaments distinct, shorter than Leaves entire to undulate or obscurely crenate,
sepals, anthers latrorse, dehiscing longitudinally; often with laminar and marginal glands; petiole
pollen subglobose, 3-colporate, colpi margins apically and rarely basally pulvinate; stipules
broad, exine coarsely reticulate; pistillode mas- deciduous or subpersistent. Inflorescences axil-
sive. Pistillate flowers pedicellate; sepals 3 or 4, lary or ramiflorous, often fasciculate, racemoid to
deciduous; petals 0; disk 0; ovary 3 or 4(5)-locu- paniculate; staminate flowers mostly in bracteate
lar; stylodia 2-lobed, stigmatiform. Fruits capsu- triads, pistillate 1 per bract. Flowers pedicellate;
lar with fleshy pericarp; columella persistent. staminate flowers sepals (3)4 or 5(8), connate,
Seeds with thin fleshy testa; endosperm layer imbricate; petals 0; disk segments mostly small or
thin, exotegmen vittate; embryo massive, cotyle- obsolete, sometimes confluent; stamens 3–10,
dons flat, chlorophyllous. filaments distinct; anthers basifixed to dorsifixed,
Two spp. from SE Asia: Thailand to Malaya introrse; pollen grains subprolate, 3-colporate,
and Borneo. pores lalongate, exine semitectate-reticulate; pis-
tillode massive, sometimes peltate. Pistillate flow-
51. Maesobotrya Benth. ers sepals 4–6(9), deciduous or persistent;
petals 0; disk 0; staminodes very rarely +; ovary
Maesobotrya Benth., Hook. Icon. Pl. 13, 75: t. 1296 (1879); 2–4-locular; ovules 2 per locule; stylodia unlobed
Léonard, Fl. Afr. Centr., Euph. 2: 46–79, t. 7–12 (1995).
to bifid, often minutely dentate or laciniate. Fruits
Dioecious trees or shrubs; indumentum simple. baccate or capsular, dehiscent or indehiscent;
Leaves entire or dentate; stipules subulate (foli- persistent columella lacking. Seeds 0–2 per locule
aceous), deciduous or persistent. Inflorescences [or capsule]; testa fleshy; endosperm copious;
terminal or axillary, sometimes ramiflorous; sta- embryo straight, cotyledons much longer and
minate flowers in racemoid thyrses, sometimes broader than radicle. 2n ¼ 26.
fasciculate or paniculate; pistillate flowers soli- About 45 spp., mainly in southeast Asia and
tary in axils of bracts on spiciform thyrses; ped- New Guinea, with one sp. in India and two in the
icels shortly articulated. Staminate flowers: sepals Pacific, mostly in rainforest and freshwater
4–6, imbricate, distinct or basally connate; petals swamp forest. Studies of Levin (1986a, b) and
Euphorbiaceae 87
Haegens (2000) indicated that Baccaurea is prob- Monoecious glabrous shrubs or trees. Leaves dis-
ably most closely related to African Maesobotrya, tichous, shortly petiolate, with minute pigment
which was confirmed by the molecular data streaks and basal laminar glands; tanniniferous
(Kathriarachchi et al. 2005). cells 0; stipules deciduous. Inflorescences axil-
lary, glomerules bracteate. Flowers sessile or sub-
53. Nothobaccaurea Haegens sessile; staminate flowers sepals 5, imbricate;
Nothobaccaurea Haegens, Blumea Suppl. 12: 198, figs. petals 0; disk segments 5; stamens 5, filaments
3.30, 3.31 (2000). distinct; anthers introrse, versatile, dehiscing lon-
gitudinally; pollen grains prolate, 3-colporate,
Dioecious trees or shrubs; indumentum simple. pores lalongate, exine tectate-perforate; pistillode
Leaves alternate or opposite, often with marginal much shorter than calyx. Pistillate sepals persis-
and laminar glands; petioles apically pulvinate; tent in fruit, petals 0; disk patelliform; ovary 3-
stipules deciduous. Inflorescences axillary or locular; stylodia distinct, bifid, erect. Fruit a bac-
ramiflorous, solitary or fasciculate, racemoid or cate thin-walled capsule, dehiscing irregularly;
spiciform; pedicels articulate or not. Staminate columella subpersistent. Seeds 2 per locule; testa
flowers sepals 3–7, fused at base; petals and disk thin and fleshy; endosperm copious; cotyledons
0; stamens 5–7, longer than sepals, filaments dis- broader than and about equaling radicle.
tinct; anthers dehiscing longitudinally or with One sp., J. congesta (Benth. ex M€ ull. Arg.)
apical slit; pistillode cylindrical, glabrous. Pistil- G.L. Webster, rainforests of Amazonian South
late flowers: sepals 5 or 6, slightly imbricate, per- America.
sistent in fruit; petals and disk 0; staminodes +;
ovary 2-locular; stylodia bifid. Fruits baccate or 55. Celianella Jabl.
tardily dehiscent; pericarp fleshy and glandular.
Seed coat fleshy; embryo with cotyledons much Celianella Jabl., Mem. New York Bot. Garden 12(3): 176,
fig. 28 (1965); Radcl.-Sm., Gen. Euph.: 68 (2001).
longer and broader than radicle.
Two spp., Melanesia: Solomons and Fiji. Dioecious shrubs; indumentum 0. Leaves subses-
Appears quite similar to Baccaurea and the Afri- sile, semisucculent, minutely puncticulate abaxi-
can Maesobotrya. ally; stipules deciduous, leaving conspicuous scars.
Inflorescences pedunculate, axillary or subtermi-
7. TRIBE JABLONSKIEAE Petra Hoffm. (2006). nal, staminate racemoid, pistillate 3–1-flowered
brachyblasts; bracts uniflorous, entire, deciduous.
Monoecious or dioecious; indumentum and exudate Staminate flowers pedicellate; sepals 5, imbricate;
0; inflorescences racemoid or glomerular; sepals 5, petals 0; stamens 5, filaments distinct; anthers
imbricate, petals 0; disk +; stamens 5, distinct. introrse, pendulous; pollen grains prolate, 3-col-
Two neotropical genera, which differ in several porate, exine finely tectate-perforate; disk central,
traits such as the reproductive system and the 5-lobate; pistillode 0. Pistillate flowers pedicellate;
structure of the anthers, fruits, and seeds (but see sepals 5, imbricate, entire, strongly veined, persis-
Tokuoka and Tobe 2002), but share similarities in tent and accrescent in fruit; petals 0; disk annular;
pollen (Webster 1984) and wood structure (Men- ovary 3-locular; ovules 2 per locule, pendulous
nega 1987), and appear in a strongly supported beneath an obturator; stylodia connate in lower
sister position (Kathriarachchi et al. 2005). half, 2-fid. Fruits capsular, septicidal; columella
persistent. Seeds fusiform, testa reticulate, micro-
KEY TO THE GENERA OF JABLONSKIEAE pyle with minute caruncle; endosperm copious;
1. Monoecious; leaves distichous; flowers in axillary glo- embryo straight, cotyledons flat, much longer
merules; fruit dehiscing irregularly 54. Jablonskia and broader than radicle.
– Dioecious; leaves spiral; staminate flowers in peduncu- One sp., C. montana Jabl., on sandstone
late pseudo-racemes, pistillate ones in few-flowered tepuis of southern Venezuela above 1,000 m.
brachyblasts; fruit septicidal 55. Celianella
54. Jablonskia G.L. Webster 8. TRIBE SPONDIANTHEAE G.L. Webster (1975).

Jablonskia G.L. Webster, Syst. Bot. 9: 232 (1984); Radcl.- Dioecious; exudate reddish; inflorescences
Sm., Gen. Euph.: 62 (2001). mostly terminal, paniculate, bracts small; flowers,
88 G.L. Webster
at least the staminate, petaliferous, subsessile; 57. Uapaca Baill. Fig. 16

staminate disk dissected; stamens 5, filaments Uapaca Baill., Étude Gén. Euph.: 595 (1858); Benth.,
distinct; pistillode +; pistillate disk lobed; stylodia Hook. Icon. Pl. 13: t. 1287 (1879); Pax & K. Hoffm.,
distinct, bilobed; fruit loculicidally dehiscent; Pflanzenr. 147, XV: 298–311 (1922); Leandri, Fl. Madag.
seed testa dry; endosperm scanty; cotyledons 111(1): 163 (1958); Radcl.-Sm., Fl. Zamb. 9(4): 93 (1996).
much broader than the radicle. Dioecious pachycaul trees or shrubs with reddish
A single, monotypic African genus. resinous exudate; trunks often stilt-rooted; indu-
mentum simple or microlepidote. Leaves petio-
56. Spondianthus Engler late or subsessile, entire; stipules deciduous or 0.
Spondianthus Engler, Bot. Jahrb. 36: 215 (1905); Hutch- Inflorescences axillary, pedunculate, globose-
inson, Hook. Icon. Pl. 30: t. 2986 (1911); Léonard & capitular, solitary or clustered, bracts 5–12, con-
Nkounkou, Bull. Jard. Bot. Nat. Belg. 59: 133–149 (1989). spicuous, becoming reflexed, each with 5–10 sta-
Dioecious trees; stems with reddish exudate; minate or solitary pistillate flowers. Staminate
indumentum simple or 0. Petioles with paired flowers sessile; calyx small, truncate to 4–6-
minute glands; stipules deciduous. Inflorescences lobate, lobes imbricate; petals and disk 0; stamens
terminal or subterminal, paniculate; flowers brac- 4–6, filaments distinct; anthers introrse, erect;
teate, staminate in glomerules, pistillate solitary. pollen grains oblate spheroidal, 3-colporate,
Staminate flowers subsessile; sepals (4)5, imbri- angulaperturate, colpi narrow, pore large and
cate; petals (4)5, smaller than sepals; disk seg- rectangular, exine semitectate-reticulate; pistil-
ments 5; stamens 5, filaments distinct; anthers lode clavate to pileiform, sometimes lobate. Pis-
introrse, connective glandular; pollen subprolate, tillate flowers sessile, perianth similar to
3-colporate, pores dumbbell-shaped, exine tec- staminate; disk 0; ovary mostly (2)3(5)-locular;
tate-perforate; pistillode massive, apically flat- stylodia distinct, dilated, recurving, distally lacin-
tened. Pistillate flowers pedicellate; sepals and iate. Fruit drupaceous with mostly 3 dorsally
petals as in the staminate, or petals 0; disk cupu- carinate pyrenes, tardily loculicidal into 2 valves
lar, fleshy; ovary 3(5)-locular; ovules 2 per locule; each; columella persistent. Seeds usually 1 per
stylodia 3, distinct, shortly 2-lobate, reflexed; pyrene, compressed; endosperm copious; embryo
stigmas papillose. Fruit capsular, dehiscing locu- straight, cotyledons green, flat or slightly plicate,
licidally; columella persistent. Seeds usually 1 per longer and broader than the radicle. 2n ¼ 26.
locule, compressed, testa reddish; endosperm About 50 spp. in Madagascar and Africa;
scanty; embryo straight, cotyledons broad, flat. growing from humid evergreen to semi-decidu-
A single sp., S. preussii Engl., widespread in ous and sclerophyllous forest and from sea level
tropical Africa: Guinea to Angola, Uganda and up to over 2,000 m elevation. The largest number
Tanzania. Due to the presence of fluoroacetic acid of species are known from the Congo basin and
(Hegnauer 1989; Neuwinger 2000), Spondianthus is are in need of reevaluation, 12 are endemic to
strongly toxic, and shares with Uapaca the unique Madagascar. The involucrate heads of disk-less
presence of resinous exudate in the Phyllanthoi- flowers have always set Uapaca apart from other
deae. The molecular data place these genera in a Euphorbiaceae, so that Airy Shaw (1965) and
sister position but with low support. The available similarly Meeuse (1990) removed it to the sepa-
seed anatomical data are inconclusive. rate family Uapacaceae. However, the basic floral
and fruit structure is clearly euphorbiaceous.
9. TRIBE UAPACEAE Hutch. (1969).
Uapacaceae (M€
ull. Arg.) Airy Shaw (1965). 10. TRIBE BISCHOFIEAE (M€
ull. Arg.) Hurus. (1954).
Dioecious; exudate reddish; inflorescences axil- Bischofiaceae (M€
ull. Arg.) Airy Shaw (1965).
lary, capitular, conspicuously bracteate; flowers Dioecious; exudate 0; indumentum simple,
apetalous, disk 0; sepals 5, connate; stamens 4–6; sparse. Leaves palmately 3(5)-foliolate; stipules
pistillode large; ovary mostly 3-locular; stylodia caducous. Inflorescences axillary, paniculate;
laciniate; fruits drupaceous. flowers apetalous. Sepals 5, imbricate, margins
A monotypic Afro-Malagasian tribe. induplicate; disk 0; stamens 5; pollen grains
Euphorbiaceae 89
flowers pedicellate; sepals 5, imbricate, cucullate-

concave and enclosing anthers; petals and disk 0;
stamens 5, filaments distinct, much shorter than
sepals; anthers extrorse, muticous; pollen grains
subglobose, 3-colporate, nearly syncolpate, exine
tectate-perforate; pistillode peltate. Pistillate flow-
ers pedicellate; sepals 5, imbricate, deciduous;
petals and disk 0; staminodes minute or 0; ovary
3(4)-locular; stylodia 3(4), partially connate, subu-
late, unlobed; stigmas terete, smooth. Fruits oblate,
baccate, epicarp thin, mesocarp fleshy, endocarp
corneo-pergamaceous; locules 1–2-seeded; colu-
mella not persistent. Seeds smooth; endosperm
copious; embryo green, curved, cotyledons much
broader and longer than radicle. 2n ¼ 196.
Two spp., B. javanica Blume being wide-
spread from India to eastern Asia, Melanesia,
and Polynesia (to Samoa and Niue); B. polycarpa
(Lévl.) Airy Shaw is endemic to China.
The distinctive characters of Bischofia, espe-
cially the trifoliolate leaves, the marsupiform
domatia, and the sepals partly enclosing the
anthers led Airy Shaw (1965) to move it to the
segregate family Bischofiaceae, but the evidence
provided by Bhatnagar and Kapil (1974), Levin
(1986a), Mennega (1987), and Tokuoka and Tobe
(2001) indicates that Bischofia belongs to Phyl-
Fig. 16. Euphorbiaceae-Phyllanthoideae. Uapaca heude- lanthoideae without showing a clear affinity with
lotii. A Flowering branch. B Staminate inflorescence. C, D any other genus. In the molecular analyses it
Staminate flowers. E Pistillate flower. F Pistil. (Pax &
Hoffmann 1922) appears always in basal position within the Tan-
niniferous Clade.
Incertae sedis:
subglobose, 3-colporate; pistillode +; ovary 3(4)-
locular; stylodia simple, entire; fruits baccate.
59. Chonocentrum Pierre ex Pax & K. Hoffmann
A monotypic paleotropical tribe.
Chonocentrum Pierre ex Pax & K. Hoffmann, Pflanzenr.
58. Bischofia Blume 147, XV: 205 (1922); Webster, Ann. Missouri Bot. Gard.
81: 37 (1994); Hayden & Hayden, Ann. Missouri Bot.
Bischofia Blume, Bijdr.: 1168 (1826); Hook., Icon. Pl. 9: t. Gard. 83: 165 (1996); Radcl.-Sm., Gen. Euphorb.: 12, fig.
844 (1852); Pax & K. Hoffm., Pflanzenr. 147, XV: 312, fig. 2 (2001).
26 (1922); Li Bingtao & Gilbert, Fl. China 11: 217 (2008).
Dioecious (monoecious) trees; vessel element per- Leaves alternate, entire; stipules persistent. Flow-
forations simple and scalariform; wood fibers sep- ers in axillary glomerules. Staminate flowers sub-
tate; indumentum simple, very sparse. Leaves sessile; sepals 4–5, connate, apically pubescent;
palmately 3(5)-foliolate; leaflets petiolulate, cre- petals 0; disk cupular; stamens 4–6, filaments
nate-serrate, abaxially with marsupiform domatia distinct; anthers erect, basifixed, longer than fila-
and with paired glands on the petiole and the ments, introrse, pubescent, connective not
petiolule of the terminal leaflet [interpreted by enlarged; pollen grains prolate, 3-colporate,
Wurdack et al. 2004 as stipels]; stipules deciduous. exine apparently tectate; pistillode large, cyathi-
Inflorescences axillary, paniculate, bracts 1-flow- form, 4- or 5-lobed, glabrous. Pistillate flowers
ered, deciduous; pedicels not articulated. Staminate and fruits unknown.
90 G.L. Webster
One sp., Ch. cyanophorum (M€ ull. Arg.) Pax & tectate, mostly with conspicuous spines; male
K. Hoffm., Amazonian Brazil, only once collected gametophyte binucleate; pistillode + or 0. Pistillate
(Spruce 3781, K, P). Originally described by sepals (3)4–8(13), distinct, imbricate; disk annu-
M€ uller (1873) as Drypetes cyatophora, it was lar to lobed, dissected or 0; ovary 2–4(5)-locular;
regarded to be related to Discocarpus by Pax ovules 2 per locule; stylodia entire, often dilated (2-
and Hoffmann (1922). S.M. and W.J. Hayden fid). Fruit capsular (drupaceous). Seeds 1 or 2 per
(1996) have shown that it is not closely related locule, often carunculate, testa usually smooth and
to Discocarpus and instead suggested placing it in shiny; endosperm usually copious; cotyledons
the Antidesmeae, but there it would be anoma- plane or plicate, usually much longer and broader
lous in its cupular staminate disk and non-versa- than the radicle. x ¼ 12 or 13.
tile anthers. A predominantly Southern Hemisphere
group of three tribes with 26 genera and c. 95
species. Relationships within Oldfieldioideae
II. SUBFAM. OLDFIELDIOIDEAE Eg. K€ohler & have been greatly clarified by the anatomical
G.L. Webster (1967)1. and morphological studies of Hayden (1994) and
Hyaenanchoideae Baillon ex Hassk. (1859) (’Hyaenach-
Levin and Simpson (1994), by the analysis of seed
neae’ [sic!]). structure by Stuppy (1996), and by the molecular
Picrodendraceae Small (1917), nom. cons. analyses of Wurdack (2002) and Wurdack et al.
(2004). The latter work suggests the maintenance
Monoecious or dioecious trees, shrubs or sub-
of three tribes, of which the monotypic Podocaly-
shrubs; vessel elements (excepting Podocalyceae)
ceae are basal to Picrodendreae and Caletieae, the
mainly with simple perforations; wood fibers not
latter being sister to each other. In Picrodendreae,
septate except in Parodiodendron; indumentum
Tetracoccus is first-branching, followed by a sub-
simple [trichomes unicellular or uniseriate]. Leaves
clade in which Picrodendron is followed by Paro-
alternate, opposite or whorled; leaf blades simple
diodendron and then by Piranhea. Hyaenanche
or palmately compound, venation mostly brochi-
(formerly included in Caletieae) appears sister to
dodromous, margins entire or dentate; stomata
Oldfieldia and both are sister to Miscodontinae,
mostly brachyparacytic; stipules mostly deciduous
with which they agree in the whorled leaves and
or 0. Inflorescences axillary, glomerulate, race-
also in seed and general anatomy, but differ from
moid, capitulate, or paniculate; bracts eglandular.
them in the lack of stipules and among themselves
Flowers mostly apetalous; staminate sepals (3)4–8
in floral traits. Therefore, for these two genera
(12), imbricate, usually distinct; disk mostly
Subtribe Paveusinae is upheld. In Caletieae, from
intrastaminal or 0; stamens (2)3–30, filaments dis-
which Securinega and Hyaenanche have been
tinct or connate; anthers mostly extorse; pollen
removed, Petalostigma occupies a basal position,
spheroidal to oblate, 3–7-zono-brevicolporoidate
whereas the morphologically clearly recognizable
or -porate or 10–40-pantoporate, microperforate-
Dissilariinae and Pseudanthinae are also sup-
ported by the molecular analyses.
1
Reveal in his Website (since 1995) indicates that the sub-
family name Hyaenanchoideae Baill. & Hassk. should have KEY TO THE TRIBES OF SUBFAM. OLDFIELDIOI-
priority over Oldfieldioideae Eg. K€ ohler & G.L. Webster. DEAE
Because the latter name has been in use in very many
publications during the last four and a half decades, it is
1. Pollen grains with 4 apertures; vessel elements at least
exasperating to have to replace it with an obscure,
in part with scalariform perforation plates; staminate
completely unfamiliar and originally misspelled name that
disk extrastaminal or 0; leaves simple
obviously never has been used since its proposal by Hass-
1. Podocalyceae
karl (1859) by taxonomists working on the Euphorbiaceae. – Pollen grains [except in Tetracoccus] usually with more
For the sake of better communication, we are continuing to than 4 apertures, rarely inaperturate; vessel elements
use the younger, well-introduced name. This is also so with simple perforation plates; staminal disk intras-
because there is so much still unearthed but nomenclatu- taminal or 0; leaves simple, unifoliolatae, or palmately
rally relevant botanical literature especially from the 19th compound 2
century—e.g., in seed lists and annual reports from botanic 2. Leaf blades compound [if simple, then lamina stipellate
gardens or colleges and grammar schools—that there is no or stipules adnate to petiole]; pollen grains zonoporate
end in sight for discoveries of even older botanical names [except Androstachys]; dioecious trees or shrubs
and/or authorships. 2. Picrodendreae
Euphorbiaceae 91
– Leaf blades undivided [pseudo-trifoliolate in 60a. Paradrypetes Kuhlm.

Micrantheum]; pollen grains zonoporate or pantopo-
rate; monoecious or dioecious trees, shrubs, or sub- Formerly included in Oldfieldioideae but, on the
shrubs 3. Caletieae basis of molecular findings by Wurdack (2002)
and Wurdack and Davis (2009), this genus has
1. TRIBE PODOCALYCEAE G.L. Webster (1994). been shifted (close) to the Rhizophoraceae.
After the exclusion of Tetracoccus (to Picroden- Although there is a general resemblance in habit
dreae) and Paradrypetes (to Rhizophoraceae), the between Paradrypetes and genera such as Cassi-
tribe is monotypic. It is remarkable for its vessel pourea, none of the Rhizophoraceae has echinate
elements, 4-aperturtate pollen and ecarunculate pollen, such strongly pronounced unisexual and
seeds; in the molecular analysis of Wurdack apetalous flowers, thin, distinct and imbricate
(2002), Podocalyx is sister to all other Oldfieldioi- sepals, and sessile stigmas (see Levin 1992; Mat-
deae. thews and Endress 2011). Therefore, doubts on
the implications of the molecular data persist,
and a clarification of the issue would be highly
60. Podocalyx Klotzsch desirable.
Podocalyx Klotzsch, Arch. Naturgesch. 7: 202 (1841);
Radcl.-Sm., Gen. Euphorb.: 84 (2001).
2. TRIBE PICRODENDREAE (Small) G.L. Webster
Richeria sect. Podocalyx (Klotzsch) M€ull. Arg. (1866). (1975).
Dioecious trees; indumentum simple; vessel ele- Picrodendraceae Small (1917).
ments with simple, scalariform, and reticulate Monoecious or dioecious; leaves alternate or oppo-
perforations. Leaves simple; petioles thickened at site, simple, unifoliolate or 3–9-parted; stipules
both ends, stomata anomocytic; stipules appear- fused to petiole or 0; flowers in axillary cymes,
ing obsolete [precociously deciduous?]. Inflores- racemes, or glomerules; sepals 4–8 or obsolete;
cences axillary or pseudo-terminal, staminate disk intrastaminal or 0; stamens 4–50, distinct;
spiciform with dense pubescent capitular glomer- pollen [sub]oblate, 4–7-zoniporate, rarely [Picro-
ules, pistillate racemoid, often fasciculate. Flowers dendron] 5–8-brevicolporate or [Androstachys]
apetalous; staminate flowers pedicellate, sepals 5–7-pantoporate, echinate; ovary 2- or 3-locular;
usually 4, scarcely imbricate, basally connate; fruit capsular or drupaceous; seeds carunculate or
stamens usually 4, filaments distinct, far exserted ecarunculate; endosperm usually copious.
above the calyx; anthers extrorse; pollen oblate, 4- This heterogeneous tribe includes 9 genera in
zoniporate, echinate; pistillode represented by 4 subtribes: 2 neotropical and 2 paleotropical.
massive lobed intrastaminal disk. Pistillate flow- The West Indian genus Picrodendron has
ers pedicellate, sepals 4, entire, distinct, persistent been placed by some authors in a separate family
but not accrescent in fruit; disk cupular, 5-lobed; Picrodendraceae because of its unusual foliage,
ovary 3-locular, ovules anatropous; stylodia stig- flowers, and fruits, but the pollen indicates its
matoid, dilated. Fruiting pedicels massive, lenti- membership in the Oldfieldioideae. Androsta-
cellate; fruits capsular, loculicidally and chys, placed in family Androstachydaceae by
septicidally dehiscent; columella persistent. Airy Shaw (1965) because of its highly specialized
Seeds 1/locule, hypostase basal, hilum subtermi- flowers, also has pollen typical of the Oldfieldioi-
nal; testa smooth, blackish, exotegmen cells elon- deae. The tribal position of Tetracoccus would
gated; endosperm copious; embryo straight, flat, merit further inquiry; see the discussion by
cotyledons much longer and broader than radicle. Stuppy (1996: 172).
A single sp., P. loranthoides Klotzsch, wide-
spread and common in Amazonian riparian and KEY TO THE SUBTRIBES OF PICRODENDREAE
inundated forests of Colombia, Venezuela, and
1. Pollen grains with 4 apertures, interspinal tectum ver-
Brazil. Treated by M€ uller (1873) as a section of rucate ; pistillate sepals persistent in fruit; ovary 2–5-
Richeria (Phyllanthoideae-Antidesmateae), Podo- locular; seeds carunculate 2a. Tetracoccinae
calyx has spinose pollen grains typical of Oldfiel- – Pollen grains [at least in part] with 5 apertures or more;
dioideae, but is an isolated genus in the subfamily. pistillate sepals persistent or deciduous; ovary 2–3-loc-
ular; seeds carunculate or ecarunculate 2
Doubtful genus (for its formal treatment, see 2. Stipules adnate to petiole and persistent, or intrapetio-
under Rhizophoraceae): lar-connate and deciduous; pollen grains with
92 G.L. Webster
interspinal tectum psilate; seeds ecarunculate petiole or intrapetiolar-connate; pollen 5–8-

2b. Mischodontinae zonoporate [Androstachys pantoporate]; stylodia
– Stipules distinct, deciduous or 0; interspinal tectum connate.
baculate or verrucate; seeds carunculate or ecaruncu-
late 3
This subtribe of 5 genera is African/Madagas-
3. Leaves alternate; staminate sepals distinct; capsule sep- can in distribution, except for Mischodon in Ceylon.
ticidal or indehiscent 2e. Picrodendrinae
– Leaves alternate or more often opposite or whorled; KEY TO THE GENERA OF MISCHODONTINAE
staminate sepals connate; capsule loculicidal 4 1. Staminate "flowers" of 30–50 solitary stamens allegedly
4. Leaves digitately foliolate; stamen filaments as long as spirally arranged on an elongated column; stylodia con-
or longer than anthers 2c. Paivaeusiinae nate into a common style with distal style branches 2
– Leaves simple, whorled [at least in part]; filaments – Staminate flowers compact, with 6–25 stamens and the
shorter than anthers 2d. Hyaenanchinae filaments in 1 or 2 whorls; stylodia distinct or basally
connate 3
2a. SUBTRIBE TETRACOCCINAE G. Levin (1994). 2. Stipules discrete, adnate to petiole; leaves 3–7-foliolate,
leaflets pinnately veined; pollen grains zonoporate
Dioecious; inflorescences racemoid or paniculate; 66. Stachyandra
stamens 5–10, distinct; pollen grains 4-zoniporo- – Stipules connate, free from petiole; leaf blade simple,
rate; ovary 2–5-locular; fruit capsular. palmately veined; pollen grains pantoporate
A monogeneric North American subtribe. 65. Androstachys
3. Leaves alternate, 1–3-foliolate; staminate disk 0; pistil-
lode + 62. Aristogeitonia
61. Tetracoccus Engelm. ex Parry – Leaves opposite or whorled, simple; staminate disk
and/or pistillode 0 4
Tetracoccus Engelm. ex Parry, W. Amer. Sci. 1: 13 (1885); 4. Staminate disk 0, pistillode +; pistillate sepals decidu-
Croizat, Bull. Torrey Bot. Club 69: 456 (1942); Dressler, ous 63. Mischodon
Rhodora 56: 49 (1954), rev. – Staminate disk +; pistillode 0; pistillate sepals accres-
Halliophytum I.M. Johnston (1923). cent in fruit 64. Voatamalo
Dioecious shrubs; indumentum simple. Leaves
62. Aristogeitonia Prain
alternate, opposite or whorled, 1-veined or pin-
nately veined, entire or dentate; stipules 0. Stami- Aristogeitonia Prain, Kew Bull. Misc. Inf. 1908: 338 (1908);
nate inflorescences axillary, racemoid or Hook. Icon. Pl. 30: t. 2926 (1911); Radcl.-Sm., Fl. E. Trop.
Afr. Euphorb. 1: 118 (1987).
paniculate, sometimes fasciculate; pistillate flow- Paragelonium Leandri (1939).
ers axillary, solitary or clustered. Staminate flow-
ers pedicellate; petals 0; sepals 4–10, filaments Dioecious or monoecious trees or shrubs; indu-
distinct, exserted; anthers extrorse, dehiscing mentum simple. Leaves long-petiolate, 1–3-folio-
longitudinally; pollen grains spheroidal, zono-4- late, leaflets entire; stipules subulate, adnate to
porate, echinate; disk [pistillode?] intrastaminal, petiole. Inflorescences axillary, ramiflorous, in
lobed. Pistillate flowers pedicellate; petals 0; glomerules. Staminate flowers pedicellate; sepals 6,
sepals 5–13, imbricate, persistent in fruit; disk distinct, biseriate, imbricate; petals and disk 0;
lobed; ovary (2)3–4(5)-locular, ovules anatropous; stamens 11–15, filaments distinct; anthers extrorse,
stylodia distinct, unlobed. Fruits capsular; colu- dehiscing longitudinally; pollen grains oblate sphe-
mella persistent. Seeds carunculate; testa smooth, roidal, 5–7-zono-brevicolporate, exine echinate,
shiny, blackish, exotegmen tracheoidal; endo- microperforate; pistillode lobed. Pistillate flowers
sperm copious; embryo straight, green; cotyledons pedicellate, sepals 6, distinct, biseriate, imbricate,
flat, much longer and broader than radicle. deciduous; petals 0; disk annular, somewhat lobed;
Five spp., North America, in deserts of Cali- ovary 3-locular, slightly pubescent; stylodia short,
fornia, Arizona, and northern Mexico. flat, stigmatiform. Fruits capsular, septicidal;
columella not persistent. Seeds 1 or 2 per locule;
2b. SUBTRIBE MISCHODONTINAE M€
ull. Arg. (1865). testa shiny, 3–5 cell layers thick, ecarunculate;
Androstachydaceae Airy Shaw (1965). endosperm copious; embryo straight, cotyledons
much longer and broader than radicle.
Monoecious or dioecious; leaves simple or Seven spp., disjunct in Angola, Kenya/Tanza-
3–7-foliolate; stipules persistent and adnate to nia, one of them in Madagascar.
Euphorbiaceae 93
63. Mischodon Thwaites 65. Androstachys Prain Fig. 17

Mischodon Thwaites, Hook. J. Bot. Kew Gard. Misc. 6: 299 Androstachys Prain, Kew Bull. Misc. Inf. 1908: 438 (1908);
(1854); Raju, J. Econ. Tax. Bot. 5: 165 (1984); Philcox, Rev. Airy Shaw, Adansonia II, 10: 519 (1970); J.-F. Leroy, C.R.
Handb. Fl. Ceylon 11: 267 (1997). Acad. Sci. D 283: 147 (1976); Radcl.-Sm., Fl. Zambesiaca 9
(4): 120, t. 20 (1996).
Dioecious trees; indumentum simple. Leaves
whorled, simple, entire; stipules minute, adnate Dioecious trees; indumentum simple. Leaves
to petiole, deciduous [sometimes obsolete]. Inflor- opposite, simple or 3–7-foliolate; stipules intra-
escences axillary, paniculate [pistillate reduced petiolar, connate, sheathing, deciduous. Inflor-
to glomerules]. Staminate flowers pedicellate; escences axillary; the staminate ones in triad of
sepals 5–8, distinct, imbricate; petals 0; disk 0; elongate spikes; pistillate flowers solitary. Sta-
stamens usually 6(5–10), filaments distinct, minate flowers subtended by 3–5 distinct, nar-
exserted; anthers extrorse, dehiscing longitudi- row sepals; petals and disk 0; stamens 30–50,
nally; pollen grains spheroidal, 5–7-brachycolpo- allegedly spirally but probably not so inserted
rate, echinate; pistillode 3-lobed. Pistillate flowers on an elongate "receptacle" probably formed by
subsessile but pedicels elongating and thickened connation of the filaments; anthers extrorse,
in fruit; sepals 6, distinct, imbricate, deciduous in elongated, connective setose, dehisicing longitu-
fruit; petals 0; disk annular; ovary 3- or 4-locular, dinally; anther connective hispidulous; pollen
glabrous; stylodia stigmatiform. Fruits capsular, grains spheroidal, pantoporate; exine echinate;
thin-walled; columella persistent. Seeds with pollen grains spheroidal, 5–7-pantoporate, echi-
smooth shiny testa; exotegmen 1-layered; endo- nate; pistillode 0. Pistillate flowers pedicellate;
sperm copious; embryo straight, cotyledons sepals 5 or 6, lanceolate, distinct, imbricate,
much longer and broader than radicle. n ¼ 24. deciduous or persistent in fruit; petals 0; disk
A single sp., M. zeylanicus Thwaites, 0; ovary 3(5)-locular; stylodia connate into a
restricted to Ceylon and southern India. long columnar style; stylar branches unlobed.
Fruits capsular, depressed-globose; endocarp
64. Voatamalo Capuron ex Bosser thin and crustaceous; columella persistent.
Seeds 2 per locule, carunculate or not; testa
Voatamalo Capuron ex Bosser, Adansonia II, 15: 333
(1976); Radcl.-Sm., Gen. Euphorb.: 108 (2001). smooth and shiny, 4 or 5 cell layers thick, exo-
tegmen uniseriate; endosperm copious; embryo
Dioecious trees; indumentum 0 or very sparse. green, cotyledons much longer and broader
Leaves opposite, simple; stipules intrapetiolar, than radicle.
connate, deciduous. Inflorescences axillary, One sp., A. johnsonii Prain, Madagascar and
cymose. Staminate flowers pedicellate; sepals 6 southeast Africa.
(7), biseriate, distinct, imbricate; petals 0; disk In Androstachys and Stachyandra, the male
irregularly lobed, partly intrastaminal; stamens flowers are usually interpreted as possessing
9–14, filaments distinct, exserted; anthers numerous stamens spirally inserted on an elon-
extrorse; dehiscing longitudinally; pollen grains gated column (Radcliffe-Smith 2001), but from
spheroidal, 5- or 6-zonoporate; sexine echinate, the morphological point of view this is unlikely
microperforate; pistillode 0. Pistillate flowers in the extreme, because flowers with numerous
pedicellate; sepals 6, biseriate, distinct, imbricate, spirally arranged stamens on elongate floral axes
persistent in fruit; petals 0; disk annular; ovary are unknown in the higher eudicots. It is likely
3–5-locular; stylodia connate in lower half, that the androecia of these probably anemophi-
unlobed, distally dilated. Fruits capsular; colu- lous plants simply have strongly developed
mella persistent. Seeds carunculate. androecia but are not catkin-like pseudanthia,
Two spp., endemic to Madagascar, said by as probably erroneously assumed by Airy Shaw
Bosser to be related to Austrobuxus; however, he (1965) and Leroy (1976). In all other characters,
also compared Voatamalo with Androstachys. including the characteristic stipules, fruits and
The distinctive intrapetiolar stipules strongly seeds, the two genera agree perfectly with the
support a relationship with Androstachys and rest of the subtribe, and there is no need for an
Stachyandra. elevated taxonomic status.
94 G.L. Webster
Four spp., endemic to Madagascar. Closely

related to Androstachys and combined with it by
Schatz (2001) on the basis of the close resem-
blance in floral morphology.
2c. SUBTRIBE PAIVAEUSINAE Pax & K. Hoffm. (1922).

Dioecious; leaves alternate, opposite, or in whorls,
long-petiolate, digitately 3–8-foliate, entire; sti-
pules 0; exotegmen 1 or 4–5 cell layers thick.
A single African genus.
67. Oldfieldia Benth. & Hook. f. Fig. 18

Oldfieldia Benth. & Hook. f., Hook. J. Bot. Kew Gard.
Misc. 2: 184, t. 6 (1850); Pax & K. Hoffm., Pflanzenr. IV,
147: 297 (1922); Léonard, Bull. Jard. Bot. État 26: 338
(1956); Radcl.-Sm., Fl.Trop. E. Afr., Euphorb. 1: 114, t.
21 (1987), Fl. Zambesiaca 9(4): 117, t. 19 (1996).
Paivaeusa Welw. ex Benth. (1867).
Leaves alternate, opposite or in whorls of 3, long-
petiolate, digitately 3–8-foliolate; leaflets entire;
stipules 0. Inflorescences axillary, staminate
cymose, densely congested, pistillate 1–3-flow-
ered. Staminate flowers pedicellate or subsessile;
sepals 5–8, basally connate, imbricate; petals 0;
disk intrastaminal; stamens 4–12, filaments dis-
tinct, exserted, inserted between lobes of disk;
anthers extrorse, dehiscing longitudinally; pollen
spheroidal, 5–8-zoniporate or brachycolporate,
echinate; pistillode small or 0. Pistillate flowers
pedicellate in fruit; sepals 5–8, imbricate, persis-
tent in fruit; petals 0; disk annular; ovary 2- or
Fig. 17. Euphorbiaceae-Oldfieldioideae. Androstachys 3-locular; stylodia unlobed, apically dilated.
johnsonii. A Distal portion of fruiting branch. B Staminate Fruits capsular, tardily loculicidally dehiscent;
inflorescence. C Staminate flower. D Pistillate inflores-
cence. E Pistillate flower. F Columella. G Fruit. (Rad-
columella persistent. Seeds 1 or 2/locule, carun-
cliffe-Smith 1996; drawn by J.M. Fothergill) culate, testa fleshy; exotegmen 4 or 5 cell layers
thick; endosperm copious; embryo green, cotyle-
dons flat, much longer and broader than radicle.
66. Stachyandra Leroy ex Radcl.-Sm. Four spp., tropical Africa. Stuppy (1996) has
Stachyandra Leroy ex Radcl.-Sm., Kew Bull. 45: 562 (1990). questioned the position of Oldfieldia in the Old-
fieldioideae—and indeed in the Euphorbiaceae—
Dioecious trees; indumentum simple. Leaves because of its aberrant fruits and seeds. However,
opposite, 3–7-foliolate; leaflets pinnately veined, the spinose pollen as well as the anatomical and
entire; stipules intrapetiolar, laterally connate, molecular data do not provide any reason to
sheathing, caducous. Inflorescences and staminate remove Oldfieldia from the Euphorbiaceae.
floral structures as in Androstachys; pollen grains
spheroidal, 4–7-zonoporate; exine echinate. Pistil- 2d. SUBTRIBE HYAENANCHINAE Baill. ex M€
ull. Arg.
late flowers pedicellate; sepals 6, verticillate, per- (1865).
sistent in fruit; disk 0; ovary 3-locular; stylodia
connate into a columnar style; tips unlobed. Fruits Dioecious; disk 0; pollen grains 6- or 7-zonopo-
capsular; apiculate; columella persistent. Seeds rate; stylodia elongated, dilated; seeds carunculate.
2 per locule, carunculate, testa smooth. A single genus endemic to the Cape region.
Euphorbiaceae 95
anthers introrse to extrorse, dehiscing longitudi-

nally; pollen oblate, 6–8-zoniporate, echinate;
pistillode 0. Pistillate flowers pedicellate; sepals
3–8, distinct, deciduous in fruit; petals 0; ovary
3–4-locular, sericeous; stylodia elongated,
dilated. Fruit capsular; columella persistent.
Seeds carunculate, caruncule subterminal, hypos-
tase subbasal; testa black, shiny, exotegmen 1 cell
layer thick; endosperm copious, embryo green,
somewhat bent, cotyledons much longer and
broader than radicle.
A single sp., H. globosa (Gaertn.) Lamb. &
Vahl, Cape region of South Africa. The molecular
data (Wurdack et al. 2004) show Hyaenanche out
of place among the Old World genera of Caletieae,
where it had been accommodated, and suggest a
place close to Oldfieldia, from which it differs in,
among other things, its entire, whorled leaves.
2e. SUBTRIBE PICRODENDRINAE (Small) G.L. Webster

(1994).
Dioecious; leaves simple [unifoliolate] or digi-
tately 3-foliolate; pollen 5–8-porate or brevicol-
porate, echinate; fruit capsular or drupaceous.
This subtribe of three neotropical genera
represents the Picrodendraceae of Small (s. str.).
KEY TO THE GENERA OF PICRODENDRINAE

1. Leaves simple [unifoliolate], subsessile; pistillate disk
6-lobed 70. Parodiodendron
– Leaves mostly 3–5-foliolate, long-petiolate 2
Fig. 18. Euphorbiaceae-Oldfieldioideae. Oldfieldia dacty- 2. Fruits capsular; ovary 3-locular; staminate flowers ped-
lophylla. A Fruiting branch. B Staminate inflorescence. icellate, in spiciform thyrses; disk intrastaminal
C Staminate flowers. D Fruit, partly cut away to show 71. Piranhea
attachment of seed. (Radcliffe-Smith 1987; drawn by
Christine Grey-Wilson)
– Fruits indehiscent; ovary 2-locular; staminate flowers
sessile or subsessile, in catkins; disk 0
69. Picrodendron
68. Hyaenanche Lamb.
69. Picrodendron Planch.
Hyaenanche Lamb., Descr. Cinchona: 52, t. 10 (1797);
M€ull. Arg. in DC., Prodr. 15(2): 479 (1866); Connell, Fl., Picrodendron Planch., Hook. London J. Bot. 5: 579 (1846;
Pl. S. Afr. 21: t. 837 (1941); Dyer, Gen. S. Afr. Pl. 1: 311 nom. cons.); Fawcett & Rendle, J. Bot. 55: 268 (1917);
(1975); Radcl.-Sm., Gen. Euphorb.: 89 (2001). Correll, Fl. Bahama Arch. 410, t. 165 (1982); Hayden et al.,
Toxicodendrum Thunb. (1796); non Toxicodendron P. J. Arnold Arb. 65: 109, t. 1, 2 (1984); M.I. Hakki, Bot. Jahrb.
Miller (1754). 107: 379 (1985); Radcl.-Sm., Gen. Euphorb.: 103 (2001).
Dioecious shrubs; indumentum simple [inflores- Dioecious (monoecious) trees; indumentum sim-
cences only]. Leaves opposite or in whorls of 4, ple. Leaves long petiolate, 3-foliolate, leaflets
simple, entire; stipules 0. Inflorescences axillary, entire; stipules rudimentary, deciduous. Staminate
in glomerules or panicles. Staminate flowers ped- inflorescences axillary, spicate, catkin-like, flowers
icellate; sepals 4–8(12), basally connate; petals 1 per bract; pistillate flowers solitary. Staminate
0; disk 0; stamens 8–30, filaments distinct, shorter flowers sessile or subsessile; perianth apparently
than anthers, inserted on a convex receptacle; 0; stamens subtended by (1–)3(7) bracts; disk 0;
96 G.L. Webster
stamens (3–)10–15(55), filaments inserted on petioles (Hayden 1994: 183) suggest that the
convex receptacle, shorter than anthers; anthers leaves are unifoliolate.
slightly extrorse, basifixed, apically puberulent,
dehiscing longitudinally; pollen oblate, 5–8-zoni- 71. Piranhea Baill.
brevicolporate, echinate; pistillode 0. Pistillate Piranhea Baill., Adansonia 6: 235 (1866); Jablonski, Mem.
flowers long-pedicellate; sepals [bracts?] 4 or 5, New York Bot. Gard. 17: 121 (1967); Radcl.-Sm. & Ratter,
valvate, mostly persistent in fruit; petals 0; disk 0; Kew Bull. 51: 543 (1996), rev.
ovary 2-locular, ovules anatropous; stylodia con- Celaenodendron Standl. (1927).
nate c. halfway, distal branches lanceolate, revo- Dioecious trees or shrubs; indumentum simple.
lute. Fruits drupaceous; exocarp with vesicles. Leaves 3-foliolate, long-petiolate; leaflets entire;
Seeds usually 1 per fruit, labyrinthic, with invagi- stipules deciduous. Inflorescences axillary, stami-
nations of the tegmen into the endosperm that nate flowers in spiciform thyrses, pistillate flowers
coats the plications of the cotyledons. solitary or in racemoid thyrses. Staminate flowers
One sp., P. baccatum (L.) Krug & Urb., pedicellate; sepals 4–6, distinct, imbricate; petals
endemic to the Greater Antilles, Bahamas, Cayman 0; disk intrastaminal, lobed; stamens (3–)6–15,
Islands, and Swan Islands. Stuppy (1996) provided filaments distinct; anthers introrse, dehiscing lon-
the first correct analysis of the labyrinthic seed of gitudinally; pollen grains spheroidal, 6-zonopo-
Picrodendron and pleaded for its exclusion from rate, echinate; pistillode 0. Pistillate flowers
Euphorbiaceae, but the anatomical data summar- subsessile to long-pedicellate; sepals 4–6, biseri-
ized by Hayden et al. (1984), the typically Old- ate, imbricate, persistent or deciduous in fruit;
fieldioid pollen grains, and the molecular data petals 0; disk divided into subulate lobes; ovary
(Wurdack 2002) argue strongly for a position of 3-locular; stylodia unlobed, recurved. Fruits cap-
Picrodendron in Oldfieldioideae. sular, 3-angled; columella persistent. Seeds 1 per
locule, ecarunculate, testa smooth; endosperm
70. Parodiodendron Hunz. copious; embryo straight, cotyledons flat, much
Parodiodendron Hunz., Kurtziana 5: 331, t. 1–3 (1969); longer and broader than radicle.
Radcl.-Sm., Gen. Euphorb.: 102 (2001). Four neotropical spp., 3 from Brazil, 1 from
Dioecious trees; wood with septate fibers; indu- western Mexico.
mentum simple. Leaves unifoliolate, subsessile,
entire; stipules deciduous; apex of petiole with a 3. TRIBE CALETIEAE M€
ull. Arg. (1865).
pair of minute stipels. Inflorescences axillary, sta-
Dioecious or monoecious; staminate sepals 4–8
minate flowers in glomerules, the pistillate soli-
(13); stamens 4–30; pollen grains spheroidal,
tary. Staminate flowers pedicellate; sepals (5)6
zonoporate or pantoporate, exine echinate; pistil-
(9), distinct, imbricate; petals 0; disk intrastam-
late sepals 3–8(13); seeds carunculate or ecar-
inal; stamens 13–19, filaments distinct, inserted in
unculate; endosperm usually copious.
glandular disk, exserted beyond calyx; anthers
This is the largest tribe of Oldfieldioideae,
extrorse, dehiscing longitudinally; pollen grains
with 14 genera in 4 subtribes. It is entirely Old
spheroidal, 6–8-zonoporate, echinate, tectum
World in distribution, confined to Australia and
smooth; pistillode 0. Pistillate flowers pedicellate;
neighboring Melanesia.
sepals 6, biseriate, imbricate, foliose, deciduous;
petals 0; disk annular, 6-lobed; ovary 3-locular;
stylodia basally connate, short, unlobed, KEY TO THE SUBTRIBES OF CALETIEAE
recurved. Fruits capsular, spheroidal; columella 1. Pollen grains pantoporate; capsule dry; cotyledons as
persistent. Seeds 1 or 2/locule; hilum subterminal; broad as or broader than radicle 3b. Pseudanthinae
testa smooth, 5 or 6 cell layers thick, exotegmen – Pollen grains zonoaperturate; capsule dry or fleshy;
1-layered; endosperm copious; embryo straight, cotyledons broader than radicle; trees or shrubs 2
cotyledons flat, much longer and broader than 2. Leaves opposite; stamens distinct; anthers not apicu-
late; pistillate sepals persistent in fruit; capsule dry
radicle. 3c. Dissilariinae
One sp., P. marginivillosum (Speg.) Hunz., – Leaves alternate; stamens connate; anthers apiculate;
Bolivia and northern Argentina. The minute pistillate sepals deciduous; capsule fleshy
acropetiolar stipels and the often disarticulated 3a. Petalostigmatinae
Euphorbiaceae 97
3a. SUBTRIBE PETALOSTIGMATINAE Pax & K. Hoffm. – Stamens (7–)10–50, variously connate; staminate flow-
(1922). ers without central disk 78. Stachystemon
3. Stipules foliaceous, appearing as if 2 further leaves per
Dioecious; stamens 18–40 or more, filaments con- node 76. Micrantheum
nate; pollen oblate, 5- or 6- zonoporate, brevie- – Stipules not foliose 4
chinate; stylodia petaloid-dilated; fruits capsular; 4. Leaves caudate-acuminate; staminate sepals 4; stamens
10–12 73. Kairothamnus
seeds carunculate.
– Leaves not caudate-acuminate; staminate sepals 4–7;
A monogeneric Australasian subtribe. stamens 4–6 5
5. Ovules 1/locule; pistillode columnar; pistillate sepals
72. Petalostigma F. Muell. deciduous in fruit 75. Scagea
– Ovules 2/locule; pistillode replaced by intrastaminal
Petalostigma F. Muell., Hook. J. Bot. Kew Gard. Misc. 9: 16
disk; pistillate sepals persistent 74. Neoroepera
(1857); Airy Shaw, Kew Bull. 35: 661 (1980); Forster & van
Welzen, Blumea 44: 104–107 (1999).
73. Kairothamnus Airy Shaw
Kairothamnus Airy Shaw, Kew Bull. 34: 596 (1980), Kew
Leaves simple, entire; stipules deciduous. Inflor- Bull. Add. Ser. 8: 121 (1980); Radcl.-Sm., Gen. Euphorb.:
escences axillary, glomerular. Staminate flowers 97 (2001).
pedicellate; sepals 4, distinct, imbricate; petals 0;
disk 0; stamens 18–85, filaments basally connate Dioecious trees or shrubs; indumentum simple.
into a column; anthers extrorse, dehiscing Leaves alternate; stipules deciduous. Inflores-
longitudinally; pollen grains globose, 5- or 6- cences axillary, thyrsoid. Staminate flowers pedi-
porate, breviechinate; exine psilate-striate; pistil- cellate; sepals 4, distinct, imbricate; petals 0; disk
lode usually 0. Pistillate flowers pedicellate, sepals annular; stamens 10–12, filaments distinct,
4–6, imbricate, deciduous; disk 0; ovary (3)4-loc- inserted in convex receptacle; anthers extrorse,
ular, ovules anatropous; stylodia entire, dilated, dehiscing longitudinally; pollen grains 10–12-
petaloid. Fruits capsular, exocarp fleshy; colu- pantoporate, echinate, tectum granular; pistillode
mella slender, persistent. Seeds somewhat 0. Pistillate flowers pedicellate; sepals 6 , biseri-
compressed, carunculate; testa smooth, mostly ate, distinct, dorsally carinate, deciduous; petals
4 cell layers thick; endosperm copious; embryo 0; disk 0; ovary 3-locular, strigose; stylodia stig-
straight, cotyledons flat, much longer and matoid, ovate. Fruit capsular, stigmas persistent
broader than radicle. at apex. Seeds ecarunculate, testa dark and
Five spp., Australia, one of these reaching smooth, with 2- or 3-cell layers; endosperm copi-
eastern New Guinea (Papua). The pollen of Peta- ous; embryo straight, cotyledons flat, much lon-
lostigma was found to be close to that of Hyae- ger and broader than radicle.
nanche (K€ ohler 1965), while Stuppy (1996) One sp., K. phyllanthoides (Airy Shaw) Airy
regards the seed structure as suggesting an affin- Shaw, eastern New Guinea.
ity with subtribe Pseudanthinae. In the analyses
of Levin and Simpson (1994), Petalostigma 74. Neoroepera M€
ull. Arg. & F. Muell.
appears as the sister group to subtribe Pseu- Neoroepera M€ ull. Arg. & F. Muell. in DC., Prodr. 15(2):
danthinae and in the analyses of Wurdack et al. 488 (1866); Airy Shaw, Kew Bull. 35: 658, t. 5, A1–4 (1980);
(2004) as sister to the rest of tribe Caletieae. Henderson, Austrobaileya 3: 618, figs. 1–3 (1992); 81: 59
(1994).
3b. SUBTRIBE PSEUDANTHINAE M€
ull. Arg. (1865).
Monoecious arborescent shrubs; indumentum
Monoecious (dioecious) trees, shrubs, or herbs; simple. Leaves short-petiolate; stipules minute
stamens 3–20; pollen pantoporate. x ¼ 12. or obsolete. Inflorescences of axillary glomerules,
An Australasian subtribe of six genera. the pistillate flowers often solitary. Staminate
flowers pedicellate; sepals (4–)6(8), biseriate,
KEY TO THE GENERA OF PSEUDANTHINAE distinct, imbricate; petals 0; stamens mostly 5 or
6, filaments distinct; anthers extrorse, dehiscing
1. Fruit 1-locular and 1-seeded by abortion 2
longitudinally; pollen grains spheroidal, 16–25-
– Fruit (2)3-locular with all locules fertile 3
2. Stamens 3–6, distinct; staminate flowers with central pantoporate, sexine echinate, tectum granular or
disk 77. Pseudanthus psilate; pistillode replaced by central
98 G.L. Webster
intrastaminal disk. Pistillate flowers pedicellate, extrorse, dehiscing longitudinally; pollen grains
sepals usually 6, biseriate, distinct, persistent in prolate-spheroidal, panto[30–40]-porate, pores
fruit; petals 0; disk annular, lobed; ovary 3-locu- prominently marginate, exine echinate, tectum
lar; ovules 2/locule; stylodia proximally connate, granular; pistillode glandular, lobed. Pistillate flow-
spreading. Fruits capsular. Seeds carunculate, ers pedicellate or subsessile; sepals 4–6, distinct,
testa smooth and shiny or minutely pitted, 2 cell imbricate, persistent in fruit; petals 0; disk + or 0;
layers thick; endosperm copious; embryo ovary 2- or 3-locular; ovules anatropous, 2/locule;
straight, cotyledons much broader and longer stylodia connate at the base, unlobed. Fruits cap-
than radicle. sular; columella persistent. Seeds 1/locule, carun-
Two spp., Australia (Queensland). culate, testa smooth, 2 cell layers thick, exotegmen
uniseriate; endosperm copious; embryo green,
75. Scagea McPherson straight, cylindric, cotyledons longer than radicle.
Scagea McPherson, Bull. Mus. Nat. Hist. Nat. Paris, IV B
Three spp., endemic to Australia. The leaves
Adansonia 7: 247 (1985); McPherson & Tirel, Fl. Nouv.- interpreted by Gr€ uning and Jeanes as ternate
Caléd. 14(1): 90, t. 18 (1987); Radcl.-Sm., Gen. Euphorb.: clusters seem more plausibly interpreted as spi-
97 (2001). rally arranged leaves with foliose stipules.
Monoecious trees or shrubs; indumentum simple.
Leaves short-petiolate; stipules deciduous. Inflor- 77. Pseudanthus Sieber ex Spreng. Fig. 19
escences axillary, racemoid, unisexual or bisex- Pseudanthus Sieber ex Spreng., Syst. Veg. 4(2): 22, 25
ual. Staminate flowers pedicellate; sepals (5)6 (1827); Halford & Henderson, Austrobaileya 6: 497–532
(7), distinct, imbricate; petals 0; disk 0; stamens (2003), rev.
4–6, filaments distinct, shorter than anthers; Pseudanthus Sieber ex Spreng. sect. Pseudanthus
anthers extrorse, dehiscing longitudinally; pollen Monoecious ericoid shrubs; branchlets longitudi-
grains spheroidal, 16–20-pantoporate, echinate, nally ridged by decurrent margins of stipules along
tectum rugulose, granular; pistillode usually pres- internodes; indumentum simple, sparse or 0.
ent. Pistillate flowers pedicellate; sepals 6, dis- Leaves alternate or opposite, margins entire and
tinct, imbricate, deciduous; petals 0; disk thickened; petioles short; stipules persistent. Flow-
segments 3; ovary 3-locular, pubescent; ovules ers in upper leaf axils solitary or 2 or 3; distal
anatropus, 1/locule; stylodia short, erect, thick. branchlet internodes often contracted to produce
Fruit capsular; columella persistent, distally terminal flower clusters. Staminate flowers pedicel-
enlarged. Seeds carunculate, testa smooth, 3 or 4 late; sepals (5)6, distinct; petals 0; disk 0; stamens
cell layers thick, exotegmen uniseriate; endo- (3–)6, filaments distinct; anthers dehiscing longitu-
sperm copious; embryo green, cotyledons flat, dinally; pollen grains spheroidal, panto(6–)10–14
much longer and broader than radicle. (25)-porate, brevi-echinate, exine granular; disk
Two spp. endemic to New Caledonia, origi- generally 3-lobed, fleshy. Pistillate flowers sessile or
nally described in the Crotonoideae because of subsessile; sepals (4–)6, distinct, persistent in fruit;
the (secondarily) uniovulate carpels. However, petals 0; disk 0; ovary (2)3-locular; ovules (1)2/
the pollen is typical for Oldfieldioideae. locule, anatropous; stylodia connate at the base or
to about halfway into a style, the stigmatic branches
76. Micrantheum Desf. undivided. Fruits capsular, unilocular by suppres-
Micrantheum Desf., Mém. Mus. Hist. Nat. Paris 4: 253, t. sion, 1-seeded; splitting at maturity into 3 bivalved
14 (1818); Gr€uning, Pflanzenr. 147, XV: 21 (1913); Jeanes, segments. Seed solitary, carunculate, testa smooth,
Fl. Victoria 4: 70 (1999); Radcl.-Sm., Gen. Euphorb.: 98 2 cell layers thick, exotegmen uniseriate; endo-
(2001). sperm copious; embryo cylindric, cotyledons a lit-
Monoecious ericoid shrubs or subshrubs; indu- tle broader than radicle.
mentum simple. Leaves pseudo-3-foliate [the Nine spp., eastern Australia.
stipules as large as the lamina], 1-veined. Inflores-
cences of axillary glomerules, pistillate flowers 78. Stachystemon Planch.
often solitary. Staminate flowers pedicellate; sepals Stachystemon Planch., Hooker’s Lond. J. Bot. 4: 471, t. 15
4 or 6, biseriate, distinct, imbricate; petals 0; disk 0; (1845); Halford & Henderson, Austrobaileya 6: 497–532
stamens 3–6(9), filaments distinct; anthers (2003), rev.
Euphorbiaceae 99
Nine spp., endemic to southwestern Western

Australia.
3c. SUBTRIBE DISSILARIINAE Pax & K. Hoffm. (1922).

Monoecious or dioecious; stamens 8–50, fila-
ments distinct; pollen grains 5–7-zonoporate,
echinate.
Seven Australasian genera with c. 25 species.
The subtribe seems clearly monophyletic, but
generic boundaries are subject to revision.
KEY TO THE GENERA OF DISSILARIINAE

1. Dioecious 2
– Monoecious 5
2. Staminate flowers without pistillodes; seeds semiellip-
tic in outline, laterally compressed 80. Dissilaria
– Staminate flowers with pistillodes; seeds globose or
ovoid, not laterally compressed 3
3. Stylodia linear in outline; fruit subglobose, strongly
tricoccous with remnants of stylodia widely separated
84. Choriceras
– Stylodia cordate-ovate in outline; fruit globose, rem-
nants of stylodia in close proximity 4
Fig. 19. Euphorbiaceae-Oldfieldioideae. Pseudanthus 4. Ovary 3–4-locular; seeds with arilloid caruncle
ligulatus subsp. ligulatus. A Branchlet with flowers and 79. Austrobuxus
fruit. B Staminate flower from side. C Staminate flower – Ovary 2-locular; seeds ecarunculate 81. Canaca
with sepals removed. D Stamen. E Pistillate flower with 5. Stipules very large, conspicuous, > 8 mm long; pistil-
sepals and stamens removed showing central disk. F Fruit late flowers with 3 sepals; receptacle of staminate flow-
with persistent sepals. G Carunculate seed. (Halford & ers glabrous 83. Sankowskya
Henderson 2003; drawn by W. Smith)
– Stipules small, inconspicuous, > 8 mm long, or 0;
pistillate flowers with 2+2 or 2+3 sepals; receptacle of
Monoecious shrubs; branchlets ridged by decur- staminate flower hairy 6
6. All flowers with glandular disk; stylodia linear in out-
rent stipular margins; indumentum sparse or 0. line; stamens > 45; pollen spiny 82. Whyanbeelia
Leaves alternate or opposite, thickened along – All flowers without glandular disk, stylodia cordate-
margins; stipules persistent. Flowers few or soli- ovate in outline; stamens < 45; pollen smooth
tary in upper leaf axils; distal branchlet inter- 85. Longetia
nodes often contracted to produce terminal
flower clusters. Staminate flowers pedicellate or 79. Austrobuxus Miq.
rarely sessile; sepals (3)4–6(10); receptacle Austrobuxus Miq., Fl. Ned. Ind. Suppl.: 444 (1861); Airy
hemispherical to elongated; stamens 7 to numer- Shaw, Kew Bull. 29: 303 (1974); A.C. Smith, Fl. Vitiensis
ous; filaments variously connate or distinct, Nova 2: 495 (1981); McPherson & Tirel, Fl. Nouv.-Caléd.
mostly bifid distally; anthers of 2 separate contig- 14(1): 193, figs. 40–43 (1987); Forster, Austrobaileya 4:
uous cells, each transverse on the apex of the 619–626 (1997).
Bureavia Baill. (1873).
filament, dehiscing longitudinally; disk 0. Pistil- Choriophyllum Benth. (1879).
late flowers sessile or shortly pedicellate, sepals 4
or 6, persistent; disk 0; ovary 2(3)-locular; ovules Dioecious (monoecious) trees or shrubs; indu-
2/locule; stylodia 2(3), at the base shortly connate mentum simple. Leaves opposite, mostly entire;
or distinct, entire. Fruit capsular, unilocular by stipules 0. Inflorescences axillary, cymose, some-
suppression, 1-seeded, splitting at maturity into 2 times paniculate. Staminate flowers pedicellate;
(3) bivalved segments. Seeds solitary, caruncu- sepals 4–6, distinct, imbricate; petals 0; disk 0 or
late, endosperm copious; cotyledons several represented by convex receptacle; stamens 8–27,
times broader than radicle. filaments distinct, inserted in the raised
100 G.L. Webster
receptacle; anthers extrorse, dehiscing longitudi- cellate; sepals 4, imbricate, persistent in fruit;
nally; pollen grains 6–7-zonoporate, echinate, petals 0; disk annular; ovary 2-locular; stylodia
exine between spines rugulose; pistillode 0. Pistil- very short, stigmatoid. Fruits capsular, endocarp
late flowers subsessile or pedicellate; sepals 4–6, thin; columella persistent or deciduous. Seeds sub-
distinct, imbricate, persistent in fruit; petals 0; orbicular, compressed tangentially, rugose, pale,
disk cupular or 0; ovary 3(4)-locular; stylodia ecarunculate; endosperm copious; cotyledons flat,
very short, dilated, stigmatiform. Fruits capsular; much longer and broader than the radicle.
columella persistent. Seeds smooth, caruncle Seven spp., all endemic to New Caledonia.
laciniate; testa smooth, 3 or 4 cell layers thick, This genus was combined with Austrobuxus by
exotegmen uniseriate; endosperm copious; McPherson and Tirel (1987); however, Airy Shaw
embryo green, straight or curved, cotyledons (1971), who first reduced Canaca to synonymy,
much longer and broader than radicle. said that it "almost merits generic recognition"
A genus of c. 20 species, from Malaysia and because of the very distinctive seeds. The distinc-
Australia to New Caledonia and Fiji. tive indumentum and 2-locular thin-walled cap-
sule with deciduous columella also support the
80. Dissiliaria F. Muell. ex Baill. generic distinctiveness of Canaca.
Dissiliaria F. Muell. ex Baill., Adansonia I, 7: 366 (1867);
Forster, Austrobaileya 5: 9–27 (1997), rev.; Radcl.-Sm., 82. Whyanbeelia Airy Shaw & Hyland
Gen. Euphorb.: 91, fig. 10 (2001).
Whyanbeelia Airy Shaw & Hyland, Kew Bull. 31: 375
Monoecious or dioecious trees or shrubs; indu- (1976) and ibid. 35: 691 (1980); Hyland & Whiffin,
mentum simple. Leaves opposite, entire or Austral. Trop. Rain For. Trees 2: 152 (1993).
crenulate; stipules interpetiolar, deciduous. Dioecious (?) trees; indumentum simple. Leaves
Inflorescences axillary, glomerular. Staminate opposite, short-petiolate; stipules obsolete. Inflor-
flowers pedicellate; sepals 3+3, imbricate; petals escences axillary, cymosely paniculate. Staminate
0; extrastaminal disk 0; stamens 8–26, filaments flowers pedicellate; sepals 6, biseriate, imbricate;
distinct, shorter than sepals, inserted on slightly petals 0; disk 0; stamens 50–55, inserted on the
convex receptacle; anthers dehiscing longitudi- pubescent central receptacle; filaments distinct,
nally; pollen grains 5–7-zonoporate, pores bor- exserted; anthers subspheroidal; pollen grains
dered; exine echinate, tectum baculate; pistillode oblate, 5–6-zonoporate, echinate, interspinal tec-
0. Pistillate flowers pedicellate; sepals 3+3, persis- tum verrucate; pistillode 0. Pistillate flowers pedi-
tent in fruit; petals 0; disk annular; ovary 2 or cellate; sepals 6; petals 0; disk dissected into
3-locular; stylodia unlobed, shortly connate at subulate segments, glabrous; ovary 3-locular,
the base. Fruits capsular, dehiscing septicidally pubescent; stylodia unlobed, recurved. Fruit cap-
into 2 or 3 bivalved cocci. Seeds laterally com- sular. Seeds carunculate, testa 4 or 5 cell layers
pressed, carunculate; endosperm copious; cotyle- thick, exotegmen uniseriate; cotyledons green.
dons broad, flat. A single sp., W. terra-reginae Airy Shaw &
Six spp., Australia (Queensland). B. Hyland from Australia (Queensland), with
the habit of Dissiliaria. Although the genus was
81. Canaca Guillaumin first described as monoecious, Airy Shaw (1971)
characterized it as dioecious.
Canaca Guillaumin, Arch. Bot. Caen 1: 74 (1927); Airy
Shaw, Kew Bull. 25: 508 (1971); Radcl.-Sm., Gen.
Euphorb.: 91 (2001). 83. Sankowskya P.I. Forster Fig. 20
Dioecious trees or shrubs; indumentum of abaxial Sankowskya P.I. Forst., Austrobaileya 4: 329, fig. 1 (1995);
leaf blades appressed, crystalloid, dark, partly
malpighiaceous. Leaves opposite; stipules 0. Monoecious trees; indumentum simple. Leaves
Inflorescences axillary, dichasial. Staminate flow- opposite, petiolate; stipules interpetiolar, decidu-
ers pedicellate, sepals 4, imbricate; petals 0; disk 0; ous. Inflorescences axillary, cymose, bracteate. Sta-
stamens (8–)15–26, filaments distinct, inserted on minate flowers pedicellate; sepals 4, biseriate,
convex receptacle; anthers dehiscing longitudi- imbricate; petals 0; disk 0; stamens 12–15, filaments
nally; pistillode 0. Pistillate flowers distinctly pedi- distinct, inserted on convex receptacle; anthers
Euphorbiaceae 101
cymose, unisexual or bisexual. Staminate flowers

pedicellate; sepals 4–6, biseriate, imbricate; petals
0; disk 0; stamens 4–6, distinct, inserted on the
pubescent central receptacle; anthers extrorse,
dehiscing longitudinally; pollen grains oblate, 6-
zonoporate, exine pilate-wrinkled, scabrate; pis-
tillode small, pubescent. Pistillate flowers pedicel-
late; sepals 6, biseriate, imbricate; petals 0; disk of
3 segments; ovary 3(4)-locular, sericeous; stylodia
distinct, undivided, recurved. Fruits capsular,
apex trifid due to persistent stylodia bases; colu-
mella persistent [?], slender. Seeds 1 or 2 per
locule, ecarunculate; testa smooth, 2 or 3 cell
layers thick, tegmen uniseriate; endosperm copi-
ous; embryo straight, cotyledons flat, much lon-
ger and broader than radicle.
One or two spp., tropical Australia and New
Guinea. Forster and van Welzen (1999) reduced
C. australiana Benth. to synonymy. Pollen mor-
phology indicates that the genus is closely related
to Longetia.
85. Longetia Baill.

Longetia Baill., Adansonia I, 6: 352 (1866); Pax &
K. Hoffmann, Pflanzenr. 147, XV: 289 (1922); McPherson
Fig. 20. Euphorbiaceae-Oldfieldioideae. Sankowskya
& Tirel, Fl. Nouv.-Caléd. 14(1): 188 (1987).
stipularis. A Fruiting branchlet. B Base of leaf pair
with interfoliar stipule. C Staminate inflorescence. Monoecious shrubs; indumentum simple, very
D Staminate flower. E Pistillate inflorescence. F Fruit.
G Carunculate seed, adaxial view. (P.I. Forster 1995; sparse. Leaves opposite, glabrous, simple, entire;
drawn by W. Smith) stipules 0. Inflorescences terminal or axillary,
cymose-paniculate, unisexual or bisexual. Stami-
extrorse, longitudinally dehiscent; pistillode 0. Pis- nate flowers pedicellate; sepals 6, imbricate; petals
tillate flowers pedicellate; sepals 3, imbricate; petals 0; disk 0; stamens 9–17, filaments distinct; anthers
0; disk 0; ovary 3-locular; stylodia distinct, extrorse, dehiscing longitudinally; pollen grains
unlobed, papillose, recurving. Fruit capsular; endo- zono(5–)6–7-brevicolporate, apertures bordered;
carp thin. Seeds carunculate, testa smooth. exine rugulose, microspinulose; pistillode present.
A single sp., S. stipularis P.I. Forst. from Aus- Pistillate flowers pedicellate; sepals 6, biseriate,
tralia, NE Queensland. Forster (1995) suggests imbricate; petals 0; disk dissected; ovary 3-locular;
that Sankowskya may be closest to Longetia, ovules anatropous; stylodia short, stigmatoid,
with which it shares smooth pollen. stigmas emarginate. Fruit capsular, columella per-
sistent. Seeds 1/locule, tangentially compressed,
carunculate; testa smooth; endosperm copious;
84. Choriceras Baill. embryo green, straight, cotyledons flat, much lon-
Choriceras Baill., Adansonia I, 11: 119 (1873); Airy ger and broader than radicle.
Shaw, Kew Bull. 35: 604 (1961), Muelleria 4: 220 (1980); A single sp., L. buxoides Baillon, endemic to
Hyland & Whiffin, Austral. Rain For. Trees: 310 (1993); New Caledonia. Pax & Hoffmann (1922) had
Forster & van Welzen, Blumea: 44: 99–101 (1999).
included several species now placed in Austro-
Monoecious trees; indumentum simple. Leaves buxus within Longetia, but the pollen is so differ-
opposite, subentire or crenulate, short-petiolate; ent in the two genera that they cannot be very
stipules deciduous. Inflorescences axillary, closely related.
102 G.L. Webster
III. SUBFAM. PEROIDEAE Baill. & Hassk. (1859), 3. Flowers in axillary glomerules, not involucrate; stamens
taken up by K. Wurdack & P. Hoffmann in Amer. 5–15; stylodia bifid, elongated; capsule echinate; leaves
stipulate 2. Chaetocarpeae
J. Bot. 92: 1413 (2005).
– Flowers enclosed in involucres of bibracteolate bracts;
stamens 2–6; stylodia very short, stigmatoid; capsule
Dioecious (monoecious) trees, shrubs, or herbs; not echinate; leaves exstipulate 3. Pereae
latex 0; indumentum simple, malpighiaceous,
stellate, or lepidote. Leaves alternate (opposite), 1. TRIBE CLUTIEAE (M€
ull. Arg.) Pax (1890).
simple, entire; stipules + or 0. Inflorescences axil-
lary, mostly strongly condensed, rarely [Pera] A monogeneric tribe, containing only the African
surrounded by involucral bracts; sepals 2–6 [in genus Clutia.
Pera 0 in pistillate flowers and sometimes rudi-
mentary in staminate flowers]; petals + or 0; disk 86. Clutia L.
+, 0 in Pera; stamens 2–20; pollen prolate to Clutia L., Sp. Pl.: 1042 (1753); Léonard, Fl. Congo Rwa.-
oblate spheroidal, 3(4)-colporate, mostly tectate- Bur. 8(1): 93, t. 7 (1962); Radcl.-Sm., Fl. E. Trop. Afr.,
perforate; pistillode + [0 in Pera but reduced Euphorb. 1: 331, fig. 63 (1987), Kew Bull. 47: 111, figs. 1–3
pistillate flowers surrounding staminate flowers (1992), Fl. Zambesiaca 9(4): 123, t. 21, 22 (1996).
in some spp.]; ovary 3(4)-locular; stylodia bifid to Dioecious (monoecious) shrubs or perennial
bipartite; ovule 1 per locule. Fruit dehiscent herbs; indumentum simple or 0. Leaves alternate,
(indehiscent); septa membranous, fragile, with- entire, often pellucid-punctate; stipules small or
out visible vascularization. Seeds black, shiny, obsolete. Flowers in axillary glomerules, the pis-
smooth, carunculate; seed coat with tracheoidal tillate often solitary. Staminate flowers pedicel-
exotegmen [Pogonophora excepted]; endosperm late; sepals and petals 5, distinct, imbricate; disk
usually copious; cotyledons longer and wider segments 5 or more, lobed or glandular; disk of
than radicle. numerous glands in 1–3 series at the base of the
Four genera with about 125 spp., pantropical. perianth and staminal column; stamens 5, fila-
Peroideae are characterized by fruits with ments connate into a column; anthers introrse;
membranous, fragile septa without visible vascu- muticous; pollen grains prolate, 3-colporate,
larization, and very peculiar seed coats which in colpi inoperculate, exine tectate-perforate/reticu-
Clutia, Chaetocarpus and Pera have a large tanni- late; pistillode inserted on top of staminal col-
niferous exotesta and a tracheoidal exotegmen, umn. Pistillate flowers pedicellate; sepals 5,
whereas Pogonophora, like the cheilosoids, imbricate, entire, persistent in fruit; petals 5, usu-
acalyphoids, crotonoids, and euphorbioids, has ally persistent in fruit; disk segments 5; ovary 3
a palisadal exotegmen (Tokuoka and Tobe (4)-locular, glabrous or pubescent; ovules anatro-
2003). Therefore, the inclusion of Pogonophora pous, inner and outer integuments thin, mostly 3
in the Peroideae remained uncertain (Tokuoka or 4 cell layers; stylodia distinct, bifid. Fruits
2007), but its inclusion in a strongly supported capsular; columella persistent, with deciduous
clade with Pera and Clutia (Xi et al. 2012) now wings in distal half. Seeds carunculate, testa dry,
leaves little doubt of this. mostly smooth, exotegmen tracheoidal; endo-
sperm copious, cotyledons broader and some-
KEY TO THE TRIBES OF SUBFAM. PEROIDEAE what longer than radicle.
1. Staminate flowers with 5 imbricate sepals and petals; About 75 African spp., most in South Africa,
stamens 5; pollen exine tectate-perforate; indumentum two in Arabia.
simple or malpighiaceous; seeds carunculate; dioecious
2 2. TRIBE CHAETOCARPEAE (M€
ull. Arg.) G.L. Webster
– Staminate flowers not with 5 imbricate sepals and (1975).
petals (or if so, cotyledons not broader than radicle);
stamen number variable; pollen grains various 3 Dioecious; indumentum simple or 0; leaves entire;
2. Petals adaxially barbate; filaments distinct; disk seg- flowers in axillary bracteate glomerules; sepals 4 or
ments not glandular-lobed; seeds ecarunculate; leaves
not pellucid-punctate 4. Pogonophoreae 5, imbricate; petals + or 0; stamens 5–15; fruits
– Petals not adaxially barbate; filaments connate; disk capsular; seeds carunculate, exotegmen tracheoidal.
segments glandular-lobed; seeds carunculate; leaves Two genera; their close relationship is con-
usually pellucid-punctate 1. Clutieae firmed by Tokuoka (2007).
Euphorbiaceae 103
KEY TO THE GENERA OF CHAETOCARPEAE

1. Petals +; anthers extrorse, subsessile on staminal
column; ovary smooth; endosperm 0
87. Trigonopleura
– Petals 0; anthers introrse or latrorse, filaments well
developed; ovary tuberculate or echinate; endosperm +
88. Chaetocarpus
87. Trigonopleura Hook. f.

Trigonopleura Hook. f., Fl. Brit. Ind. 5: 399 (1887), Icon.
Pl. 18: t. 1753 (1888); Pax & K. Hoffm., Pflanzenr. 147, III:
95, fig. 30 (1911); van Welzen et al., Blumea 40: 363–374,
fig. 2 (1995).
Dioecious trees; indumentum simple, scale-like
on young growth. Leaves simple, petiolate [petiole
adaxially sulcate], entire; stipules deciduous.
Flowers in axillary glomerules, the staminate
multiflowered, the pistillate 1–3-flowered; bracts
minute. Staminate flowers pedicellate, articulate
at or above the middle; sepals 5, coriaceous,
imbricate, distinct; petals (4) 5, distinct, valvate,
pubescent; disk segments 5, erect; stamens (5–)
8–15, biseriate, filaments connate into a column;
5 anthers of lower whorl subsessile on column,
3 upper anthers equaling filaments; anthers
extrorse; apiculate; pollen grains subprolate,
3-colporate, colpi inoperculate, exine tectate-
punctate, tectum psilate; pistillode adnate to
staminal column, apically trifid. Pistillate flowers
pedicellate, pedicels mid-articulate; sepals 5,
imbricate, entire, deciduous in fruit; disk-seg-
Fig. 21. Euphorbiaceae-Peroideae. Chaetocarpus rabar-
ments 5; ovary 3-locular, tomentose; stylodia dis- aba. A Flowering branchlet. B Staminate flower. C
tinct, erect, bipartite, adaxially papillose. Fruits (Pseudo?)hermaphrodite flower. D Stylodia. E Fruiting
capsular, subglobose, tomentose, carinate-angled, branchlet. F Young fruit. G Two-valved coccus. H Colu-
reticulate-venose; columella persistent, trans- mella with seeds. (Capuron 1972; drawn by D. Godot de
Mauroy)
lucent-winged. Seeds carunculate, the fleshy
caruncle partially covering testa; exotegmen tra-
cheoidal, mesotesta vascularized; hilum lobed; Ceylon 11: 177 (1997); Radcl.-Sm., Fl. Zambesiaca 9(4):
endosperm 0. 135, t. 23 (1996).
Three spp., Malay Peninsula and Sumatra to
the Philippines and Borneo. According to Now- Dioecious trees or shrubs; indumentum simple.
icke et al. (1998), the pollen of Trigonopleura is Leaves distichous; stipules persistent, sometimes
different from that of Chaetocarpus. Although the foliose (deciduous). Flowers in axillary glomer-
seeds of Trigonopleura are exalbuminous, the ules. Staminate flowers pedicellate, pedicels mid-
genus is resolved as sister to Chaetocarpus articulate; sepals 3–5, imbricate, distinct or con-
(Tokuoka 2007). nate; petals 0; disk segments 5; stamens 5–15,
filaments proximally connate; anthers introrse or
88. Chaetocarpus Thwaites Fig. 21 latrorse, basifixed; pollen grains subprolate, 3-col-
porate, exine tectate-perforate, tectum microru-
Chaetocarpus Thwaites, Hook. J. Bot. Kew Gard. Misc. 6: gulose; pistillode distinct from stamens, trifid,
300, t. 10a (1854); Léonard, Fl. Congo Rwa.-Bur. 8(1): 127,
fig. 8 (1962); Capuron, Adansonia II, 12: 209–211, t. villose. Pistillate flowers pedicellate; sepals 3–8,
2 (1972); Alves, An. Jard. Bot. Madrid 51: 302 (1994); imbricate, distinct, deciduous in fruit; petals 0;
van Welzen, Rheedea 4: 93, fig. 1 (1994); Philcox, Fl. disk cupular, lobed; ovary 3(4)-locular, hirsute;
104 G.L. Webster
ovules anatropous, inner and outer integuments

thin, 3–6 cell layers; stylodia bifid, tips laciniate.
Fruits capsular, muricate to echinate; columella
persistent. Seeds ovoid, compressed, carunculate;
testa dry, smooth and shiny; exotegmen tracheoi-
dal; endosperm copious; cotyledons longer and
broader than radicle.
Fivteen spp. scattered from the West Indies
and South America to Africa, Madagascar, India,
and Malesia. The seeds of Chaetocarpus, with
shiny blackish testa, are strikingly similar to
those of Pera, and suggest a close relationship
between the two genera.
3. TRIBE PEREAE (Klotzsch & Garcke) Pax &

K. Hoffm. (1919).
Monogeneric; represented only by the New
World genus Pera.
89. Pera Mutis Fig. 22

Pera Mutis, Kongl. Vetensk. Acad. Nya Handl. 5: 299
(1784); Pax & K. Hoffm., Pflanzenr. 147, XIII: 2, figs. 1,
2 (1919); Correll, Fl. Bahama Arch.: 834, fig. 347 (1982); L.
Gillespie & Armbruster, Smiths. Contr. Bot. 86: 8, figs. 2, 3
(1997); Bigio & Secco, Rodriguésia 63: 163–207 (2012),
spp. of Braz. Amazonia.
Dioecious (monoecious) trees or shrubs; indu-
mentum lepidote or stellate (simple). Leaves Fig. 22. Euphorbiaceae-Peroideae. Pera anisotricha.
A Flowering branchlet. B Staminate inflorescence with
alternate (opposite), simple, entire, exstipulate. involucre. C Three staminate flowers. D Four pistillate
Inflorescences pseudanthial, axillary, fascicled; flowers subtended by the opened involucre. E Two of
each pseudanthium usually unisexual, of 3 or 4 these flowers with basal staminodes. F Part of infructes-
flowers subtended by a laterally dehiscing com- cence with two septifragal capsules. (Bigio & Secco 2012;
drawn by B. Alvarez)
pound involucrate bract [of 2 connate bracts];
petals and disk 0. Staminate flowers sessile, sur-
rounded by "pistillodes" [reduced pistillate flow- 4. TRIBE POGONOPHOREAE (M€
ers]; calyx irregularly lobed; stamens (2)3–4(8), (1975).
filaments distinct or connate; anthers introrse to A single genus with a disjunct African/South
extrorse; pollen grains suboblate to subprolate, 3- American distribution.
colporate, exine intectate or tectate, reticulate to
microrugulate. Pistillate flowers sessile or subses- 90. Pogonophora Miers ex Benth.
sile; perianth and disk 0; ovary 3-locular; ovules
anatropous, inner and outer integuments thin, Pogonophora Miers ex Benth., Hook., J. Bot. Kew
Gard. Misc. 6: 372 (1854); Letouzey, Adansonia II, 9:
3–5 cell layers; stylodia very short and connate 275, fig. 1 (1969); Secco, Revisāo dos gêneros Anomalo-
into a thick peltate stigmatic structure. Fruits calyx Ducke, Dodecastigma Ducke, Pausandra Radlk.,
capsular, valves woody; columella slender, usu- Pogonophora Miers ex Bent. e Sagotia Baill.: 88, figs. 26,
ally not persistent. Seeds carunculate, testa dry, 27 (1990).
smooth, dark, shiny, exotegmen tracheoidal. Dioecious trees or shrubs; indumentum simple,
About 30–35 neotropical spp. In its floral alpighiaceous in inflorescence. Leaves alternate;
morphology the genus is very distinct from all stipules small or obsolete. Inflorescenes of axillary
other Euphorbiaceae, but the seed morphology thyrses or spikes; bracts persistent, eglandular.
(Tokuoka and Tobe 2003) suggests a relationship Staminate flowers sessile; sepals 5, distinct, scari-
with Chaetocarpus. ous, imbricate, unequal, outer bract-like; petals 5,
Euphorbiaceae 105
distinct, imbricate, longer than sepals, barbate – Ovary 2-locular; stamens 5–8 (9), anthers muticous;
adaxially; stamens 5, filaments distinct; anthers leaves with laminar glands (except N. kingii); bracts
introrse, basifixed; pollen grains subprolate, 3- glandular 92. Neoscortechinia
colporate, exine tectate-perforate, tectum psilate;
disk intrastaminal, urceolate, enclosing the 2–3- 91. Cheilosa Blume
fid pistillode. Pistillate flowers pedicellate; sepals Cheilosa Blume, Bijdr.: 613 (1826); M€ ull. Arg. in DC.,
and petals similar to staminate, the calyx persis- Prodr. 15(2): 1123 (1866); J. J. Sm., Meded. Dept. Landb.
tent in fruit; disk cupular; ovary 3-locular; inner Ned.-Ind. 10: 604 (1910); Pax & K. Hoffmann, Pflanzenr.
and outer integuments thin, 5 or 6 cell layers; 147, XV: 12, fig. 3 (1912); Whitmore, Tree Fl. Malaya 2: 77
(1973); van Welzen et al., Blumea 38: 162, fig. 1 (1993).
stylodia basally connate, bifid, branches fimbri-
ate-papillate. Fruits capsular, columella slender, Dioecious trees; indumentum simple or fascicled.
persistent. Seeds carunculate; testa smooth or Leaves dentate to crenate, teeth with abaxial dis-
slightly rugulose; exotegmen palisadal. coid glands; petiole apically pulvinate; stipules
Three spp., widely disjunct, P. schomburgki- deciduous or subpersistent. Inflorescences
ana Miers ex Benth. neotropical, the recently axillary or terminal, the staminate thyrsoid-
described P. africana Letouzey and P. letouzeyi paniculate, the pistillate thyrsoid-racemoid;
Feuillet from West Africa (Gabon). bracts eglandular, persistent, sometimes foliose.
Staminate flowers pedicellate; sepals 5 (6), dis-
IV. SUBFAM. CHEILOSOIDEAE (M€
ull. Arg.) tinct, imbricate; petals 0; disk annular, 5-
K. Wurdack & P. Hoffm. (2005). lobed, pubescent; stamens 9 or 10, biseriate,
filaments distinct; anthers basifixed, introrse,
Dioecious trees; latex 0; indumentum simple and/ apiculate; pollen grains suboblate, 3-colporate,
or stellate. Leaves alternate, simple, entire or colpi inoperculate, tectum echinate and rugulose;
toothed. Inflorescences pseudo-terminal or axil- pistillode mainly 3-lobed, hirsutulous. Pistillate
lary, thyrsopaniculate. Flowers apetalous; stami- flowers pedicellate; sepals (4) 5 (6), distinct
nate flowers: sepals 4–6, distinct, imbricate; disk or nearly so, imbricate, entire, persistent in
with 4 or 5 segments; stamens 5–10, distinct; pollen fruit; petals 0; disk annular, pubescent; ovary
grains suboblate, 3-colporate, colpi inoperculate, 3-locular, pubescent; ovules anatropous, inner
exine echinate; pistillode pubescent; pistillate flow- and outer integuments moderately thick, the
ers: sepals 5 or 6, distinct, imbricate, persistent in outer vascularized; stylodia basally connate,
fruit; disk annular or 0; ovary 2–3-locular; ovules unlobed, recurving. Fruits capsular, rugulose;
with thick integuments, the outer vascularized; mesocarp woody; columella cylindrical, persis-
stylodia bifid. Fruits capsular; columella persistent. tent. Seeds 1–3 per fruit, ovoid, ecarunculate,
Seeds with sarcotesta; mesotesta vascularized; exo- exotesta fleshy.
tegmen palisadal; endosperm copious. A single sp., Ch. montana Blume, widespread
Only one tribe: in western Malesia: Malay Peninsula and Sumatra
to Borneo and the Philippines.
1. TRIBE CHEILOSEAE (M€
ull. Arg.) Airy Shaw &
G.L. Webster (1975). 92. Neoscortechinia Pax
Two closely related genera of southeast Asia Neoscortechinia Pax, Nat€url. Pflanzenfam. Nachtr. 1: 213
and Malesia. Cheiloseae are remarkable for their (1897); Pax & K. Hoffm., Pflanzenr. 147, XIV: 52 (1919);
Whitmore, Tree Fl. Malaya 2: 119 (1973); van Welzen,
echinate exine sculpture; the seed structure with Blumea 39: 301–320, fig. 3 (1994).
the fleshy exotesta and vascularized mesotesta
(Tokuoka and Tobe 2003) is similar to that of Dioecious trees; indumentum of simple and
Trigonopleura. fasciculate hairs. Leaves usually with 2 raised
Two genera, 7 spp., SE Asia to Melanesia. adaxial basal glands, subentire to glandular-
dentate, petioles pulvinate; stipules decidu-
KEY TO THE GENERA OF CHEILOSEAE ous. Inflorescences axillary or pseudo-terminal,
1. Ovary 3-locular; stamens 8–10, anthers apiculate; thyrsoid, staminate compounded; bracts
leaves without laminar glands; bracts eglandular entire, sometimes glandular; flowers subsessile.
91. Cheilosa Staminate sepals 4–5(6), imbricate; petals 0;
106 G.L. Webster
disk dissected; stamens (4)5–9, filaments dis- isolated Erismantheae, which may occupy a sister
tinct; anthers basifixed, introrse to latrorse, position to the rest of the subfamily.
muticous; pollen grains subglobose, 3-colpo-
rate, colpi inoperculate, exine echinate, tectum KEY TO THE TRIBES OF SUBFAM. ACALYPHOIDEAE
microrugulose; pistillode 2–3-fid. Pistillate 1. Leaves opposite; pollen exine reticulate; seeds
flowers pedicellate, sepals 4 or 5, imbricate, ecarunculate 1. Erismantheae
persistent or deciduous in fruit; petals and – Leaves alternate; pollen exine rugulose or micro-
disk 0; ovary 2-locular, hirsute; ovules anatro- punctate; seeds carunculate or not 2
pous, inner and outer integuments thick, the 2. Cotyledons scarcely broader than radicle; monoecious
outer vascularized; stylodia stigmatoid. Fruits (dioecious) subshrubs with ericoid foliage; seeds
carunculate 9. Ampereeae
capsular, oblong, ribbed; columella persistent
– Cotyledons distinctly broader than radicle; foli-
or deciduous. Seeds 1 (2) per fruit, exotesta age generally not ericoid; seeds carunculate or
fleshy, endotesta smooth. ecarunculate 3
Six spp., mainly western Malesian, extending 3. Petals +, at least in staminate flowers 4
to Burma in the West and Borneo, New Guinea – Petals 0 6
and the Solomons in the East. 4. Monoecious (dioecious); indumentum malpighiac-
eous, stellate, or lepidote [simple in Speranskiinae
and Philyra]; anther connective not enlarged, sacs
V. SUBFAM. ACALYPHOIDEAE Beilschm. (1833). not pendulous; pollen sexine often distinctly hetero-
Subfam. Ricinoideae Baillon ex Hassk. (1859). brochate 11. Chrozophoreae
Subfam. Dysopsidoideae Baillon ex Hassk. (1859). – Dioecious; indumentum simple; anther connective
Trees, shrubs, or herbs; milky latex 0 [colored enlarged, sacs pendulous; pollen sexine coarsely reticu-
late, not heterobrochate 5
exudate in some genera]; indumentum simple,
5. Stylodia bifid; pistillate petals usually present; inflor-
malpighiaceous, or lepidote. Leaves alternate escences paniculate, racemose, or spiciform; pollen
(opposite); lamina simple and pinnately veined, spheroidal; leaves stipulate 3. Agrostistachydeae
or palmately veined or lobed , often with foliar or – Stylodia multifid; pistillate petals 0; inflorescences
petiolar glands. Inflorescences axillary or termi- capitellate; pollen prolate; leaves exstipulate
nal, racemose, spicate, or paniculate, or reduced 4. Sphyranthereae
to axillary glomerules or solitary flowers; bracts 6. Stylodia connate, unlobed; plants often scandent;
sometimes glandular. Staminate flower: sepals stinging hairs sometimes present; pollen tectum
microverrucate 14. Plukenetieae
imbricate or valvate [sometimes connate in bud – Stylodia distinct or basally connate; plants rarely
and rupturing at anthesis]; petals and disk + or 0; scandent; stinging hairs 0; pollen tectum psilate to
stamens 2–100+, filaments distinct or connate; microverrucate 7
pollen grains binucleate, mostly 3- or 4-colporate 7. Pollen tectum perforate, psilate to scabrate 8
(3-colpate in some Plukenetieae), exine mostly – Pollen tectum usually rugulose, microverrucate 11
semitectate-reticulate (echinate); pistillode + or 8. Staminate disk 0; indumentum stellate 10. Epiprineae
0. Pistillate flower: sepals mostly 3–6 (2–12), – Staminate disk + [except in Adenophaedra]; indu-
imbricate or open at anthesis; petals and disk + mentum simple 9
or 0; ovary mostly 2–4-locular, ovules solitary in 9. Pistillate sepals deciduous; staminate disk massive,
each locule, anatropous, inner integument non- intrastaminal; stamens 4–15; pistillode + or 0
2. Caryodendreae
vascularized; stylodia entire to multifid. Fruit
– Pistillate sepals usually persistent; staminate disk not
capsular (baccate or drupaceous). Seeds caruncu- massive and extrastaminal; stamens (2–)8–100 or
late or ecarunculate; seed coat with palisadal exo- more; pistillode usually 0 10
tegmen; endosperm usually copious; cotyledons 10. Pollen colpi distinctly marginate; indumentum simple
longer and broader than radicle [except in or stellate; anthers sometimes 4-locellate
Ampereeae]. x mostly ¼ 9, 10, 11. 13. Bernardieae
The Acalyphoideae, with 99 genera in 14 tribes – Pollen colpi emarginate; indumentum simple; anthers
2-locellate 8. Pycnocomeae
and a total of over 3,000 species, are the largest and 11. Pollen grains with operculate colpi; seeds mostly
most complex of the seven subfamilies. They are ecarunculate, testa not fleshy; stipules deciduous or
resolved as monophyletic with exception of the obsolete 12
Euphorbiaceae 107
– Pollen grains with inoperculate colpi; seeds carun- – Pistillode deeply trifid into slender branches; staminate
culate or ecarunculate; testa dry or fleshy; stipules inflorescence not closely bracteate; staminate pedicel <
deciduous or persistent 13 1.5 cm long; stamens 4–11 2
12. Leaves without laminar glands; stylodia subentire to 2. Stipules persistent, foliaceous and cordate at the base;
multifid; inflorescences axillary; pollen sexine finely staminate pedicels > 2 mm long; cymules unisexual;
perforate-tectate 12. Adelieae pistillate flowers petaliferous 94. Moultonianthus
– Leaves usually with embedded laminar glands; – Stipules deciduous, triangular, not cordate; staminate
stylodia mostly entire, if divided, then inflorescences pedicels < 1 mm long; cymules bisexual; pistillate
terminal; pollen exine rugulose to striate flowers apetalous 95. Syndyophyllum
5. Alchorneeae
13. Seeds carunculate; inflorescences terminal; indu-
mentum stellate or 0 6. Ricineae 93. Erismanthus Wall. ex M€
ull. Arg.
– Seeds mostly ecarunculate; inflorescences axillary Erismanthus Wall. ex M€ ull. Arg. in DC., Prodr. 15(2): 1138
or terminal; indumentum simple or stellate (1866); Pax & K. Hoffm., Pflanzenr. 147. III: 33, fig. 9
7. Acalypheae (1911); Airy Shaw, Kew Bull. 36: 294 (1981); van Welzen,
Blumea 40: 379, fig. 2 (1995).
1. TRIBE ERISMANTHEAE G.L. Webster (1975). Monoecious trees or shrubs, branching sympo-
Monoecious trees or shrubs; indumentum sim- dially; indumentum simple, often inconspicuous.
ple; leaves opposite, entire or obscurely dentate; Leaves opposite, minutely glandular-serrulate,
stipules interpetiolar; inflorescences axillary, petiole not pulvinate; stipules interpetiolar, sub-
racemoid or spicate; staminate sepals 4 or 5, persistent or deciduous. Inflorescences unisexual
imbricate (valvate); petals 5 or 0, disk minute or bisexual, staminate flowers in bracteate catkins
or 0; stamens 5–15, filaments distinct; pollen grains or capitula, pistillate flowers solitary. Staminate
3-colporate, angulaperturate, colpi emarginate, flowers pedicellate; sepals (4) 5, imbricate; petals
inoperculate, sexine tectate-reticulate; pistillode 5, shorter than sepals; disk 0; stamens 12–15,
+; pistillate sepals 5 or 6, imbricate; petals + or 0; filaments distinct; anthers biseriate, basifixed,
disk 0; stylodia distinct or connate, bifid; fruit dehiscing latrorsely; pollen grains subprolate,
capsular; seeds ecarunculate, testa smooth and dry. 3-colporate, exine tectate-punctate/reticulate,
Three tropical Asian genera with a total of 5 reticulum slightly vermiculate; pistillode long-
spp. Nowicke et al. (1998) note the anomalous exserted, slender, trifid at apex. Pistillate flowers
nature of the pollen grains of Syndyophyllum, pedicellate; sepals 5, imbricate, subentire,
which are operculate and resemble the pollen of foliose, persistent and reflexed in fruit; petals
Alchornea, but in the rbcL analysis of Wurdack and disk 0; ovary 3-locular, hirsute; stylodia
et al. (2005), Moultonianthus and Syndyophyllum proximally connate, distally bifid, slender, papil-
form a monophyletic group. Later (Wurdack and lose. Fruit oblate, capsular; columella persistent,
Davis 2009), Moultonianthus has been resolved in apically dilated. Seeds 1–3 per fruit, subglobose,
a position sister to the rest of subfamily Acaly- ecarunculate, testa smooth.
phoideae s.str., thus confirming the placement of Two spp., southeast Asia: Vietnam to Suma-
Erismantheae by Webster (1994). tra and Borneo.
The apetalous flowers of Erismanthus (and
possibly of Syndyophyllum as well) and their 94. Moultonianthus Merr. Fig. 23
elongate stylodia and catkin-like staminate Moultonianthus Merr., Phil. J. Sci. Bot. 11: 70 (1916); Pax
inflorescences are indicative of wind pollination. & K. Hoffm., Pflanzenr. 147, XIV: 41 (1919); van Steenis,
Bull. Bot. Gard. Buitenz. III, 17: 404 (1948); van Welzen,
The opposite leaves are thought to be due to Blumea 40: 384, fig. 3 (1995).
strong shortening of each second internode
beginning in the seedling stage (see Radcliffe- Monoecious trees or shrubs; indumentum simple
Smith 2001: 121). but axes early glabrescent. Leaves opposite,
shallowly crenate; stipules interpetiolar, foliose,
KEY TO THE GENERA OF ERISMANTHEAE cordate, persistent. Inflorescences axillary, uni-
sexual, racemiform; bracts small, eglandular. Sta-
1. Pistillode elongated, clavate; staminate inflorescence
covered with closely imbricate bracts; staminate minate pedicel articulate; sepals 5, imbricate,
pedicel > 1.5 cm long; stamens 12–15; pistillate flowers biseriate; petals 5 (7), longer than sepals, white,
apetalous 93. Erismanthus entire; disk rudimentary, 5-lobed; stamens 8–11,
108 G.L. Webster
One sp., M. borneensis Merr., Sumatra and

Borneo.
95. Syndyophyllum Lauterb. & K. Schum.

Syndyophyllum Lauterb. & K. Schum., Fl. Schutzgeb.
S€
udsee: 403 (1901); Pax & K. Hoffm., Pflanzenr. IV. 147:
104 (1911); Airy Shaw, Hook. Ic. Pl. 38(1): t. 3722 (1974);
van Welzen, Blumea 40: 388, fig. 4 (1995).
Monoecious trees; indumentum simple, glabres-
cent. Leaves opposite, distichous, simple,
minutely puncticulate, sometimes with abaxial
domatia, obscurely crenulate; stipules interpetio-
lar, deciduous. Inflorescences axillary, bisexual or
staminate thyrses, pistillate flowers solitary;
bracts minute. Staminate flowers subsessile;
sepals 5, imbricate, basally connate; petals shorter
than sepals or 0; disk 0; stamens 4–10, filaments
distinct, in 1 or 2 series, much longer than
anthers; anthers basifixed, apiculate, introrse;
pollen grains subspheroidal, 3-colporate, colpi
operculate, pistillode deeply trifid, shorter than
filaments. Pistillate flowers subsessile; sepals 4 or
5, imbricate, basally connate, entire, persistent
and reflexed in fruit; petals and disk 0; ovary 3-
locular, hirsute; stylodia connate halfway, tips
apically bifid and adaxially papillose. Fruits cap-
sular; columella persistent, trigonous and api-
cally dilated. Seeds subglobose, ecarunculate,
testa smooth.
Two spp. from Sumatra, Borneo, and New
Guinea.
Fig. 23. Euphorbiaceae-Acalyphoideae. Moultonianthus
leembruggianus. A Branchlet with separate staminate
and pistillate flowers. B Staminate flower. C Staminate
2. TRIBE CARYODENDREAE G.L. Webster (1975).
flower with disk lobes, androecium and pistillode.
D Pistillate flower with perianth lobes removed, showing Dioecious; indumentum simple; leaves with basal
disk lobes and pistil. E Outline of young fruit. F Fruit laminar glands; inflorescences terminal or axil-
column after dehiscence. G Seed. (van Welzen 1995; lary, spiciform; staminate sepals 3–5, valvate;
drawn by J. van Os) petals 0; disk massive, pubescent; stamens 4–15;
pollen grains oblate, 3-colporate, colpi not mar-
ginate, inoperculate; pistillode + or 0; pistillate
biseriate, filaments distinct; anthers basifixed, flowers sessile, sepals 4–6; disk pubescent; ovary
latrorse to introrse, apiculate; pollen grains sub- mostly 3-locular; stylodia distinct, unlobed; fruit
prolate, 3-colporate; pistillode deeply trifid, capsular; seeds ecarunculate, testa dry or fleshy.
branches slender. Pistillate pedicel articulate; As treated here, Caryodendreae comprise
sepals 5, imbricate, entire or glandular-denticu- three genera, two from the New World, and one
late, persistent in fruit; petals similar to stami- African. The group will need a critical revision: in
nate; disk 5-lobed; ovary 3-locular, tomentose; the molecular analysis (Wurdack et al. 2005), Car-
ovules anatropous, inner and outer integuments yodendron is resolved in a clade with two bernar-
very thick, the outer vascularized; stylodia basally dioid genera (Bernardia and Adenophaedra),
connate, bifid, branches dilated and papillate. which is sister to the Plukenetieae, whereas Alchor-
Fruit capsular, spheroidal; columella winged. neopsis and, farther away, Discoglypremna appear
Seeds 1–3 per fruit, subglobose, ecarunculate; at the base of the "Alchorneoid" grade together
testa smooth. with agrostistachyoid genera and Mareyopsis.
Euphorbiaceae 109
KEY TO THE GENERA OF CARYODENDREAE Dioecious trees; indumentum simple. Leaves often
1. Leaves pinnately veined, stipulate; pistillode massive; crowded near apices, pinnately veined and tripli-
seed coat dry 96. Caryodendron nerved, subentire, with abaxial embedded glands
– Leaves triplinerved, exstipulate; pistillode slender or 0; and biglandular at junction with petiole; stipules
seed coat fleshy 2 deciduous. Inflorescences terminal, paniculate,
2. Staminate disk dissected; anthers apiculate; pistillode 0; bracts minute, eglandular; flowers subsessile, the
seeds trigonous, testa foveolate 97. Discoglypremna staminate several per axil, the pistillate solitary.
– Staminate disk entire, receptacular; anthers muticous; Staminate flowers pedicellate, calyx closed in
pistillode +; seeds lenticular, testa reticulate
98. Alchorneopsis bud, sepals 3–4(5), valvate; disk pubescent, dis-
sected, segments extra- and intrastaminal; petals
96. Caryodendron Karsten 0; stamens (5–)8–12(15), filaments distinct;
anthers basifixed, introrse, locules pendulous,
Caryodendron Karsten, Fl. Colombiae 1: 91, t. 45 (1860); connective apiculate; pollen grains subprolate to
M€ull. Arg., Fl. Brasil. 11(2): 706 (1874); Pax & K. Hoffm.,
Pflanzenr. 147, VII: 263 (1914); Ducke, Trop. Woods 76: prolate spheroidal, 3-colporate, colpi narrow,
18 (1943); Webster, Ann. Missouri Bot. Gard. 54: 287 inoperculate and emarginate, endoaperture
(1968); Huft, Ann. Missouri Bot. Gard. 76: 1077 (1989); circular, sexine tectate; pistillode 0. Pistillate
Radcl.-Sm., Gen. Euphorb.: 153, fig. 15 (2001). flowers pedicellate, articulate near base; sepals
Dioecious trees, glabrous except for inflores- mostly 5, valvate, acute, persistent in fruit; petals
cences; indumentum simple. Leaves biglandular 0; disk dissected into 6–8 distally pubescent seg-
adaxially near base and with small dispersed ments; ovary 3(4)-locular, glabrous or sericeous;
glands; stipules lanceolate, entire, deciduous. inner and outer ovular integuments thick, the
Inflorescences mostly terminal or subterminal, outer vascularized; stylodia distinct, unlobed,
spiciform, the staminate sometimes compound; adaxially fimbriate. Fruit 3-lobed, capsular but
bracts eglandular, scarious, rounded to truncate exocarp somewhat fleshy; cocci reticulate; colu-
with ciliate margins, multiflorous. Staminate mella slender, persistent. Seeds: exotesta fleshy,
flowers pedicellate; calyx closed in bud, sepals 3 reddish, endotesta blackish, foveolate, mesotesta
or 4, valvate; petals 0; disk massive, pulviniform, vascularized. n ¼ 11.
instrastaminal, pubescent; stamens 4–7, filaments A single sp., D. caloneura (Pax) Prain, rain-
distinct; anthers introrse, locules pendulous, forests of West Africa (including São Tomé) to
connective apiculate; pollen grains suboblate, 3- Uganda.
colporate, colpi narrow and emarginate, inoper-
culate; endoaperture ovate to lalongate, sexine 98. Alchorneopsis M€
ull. Arg.
tectate-perforate; pistillode 0. Pistillate flowers Alchorneopsis M€ ull. Arg., Linnaea 34: 156 (1865), and in
with stout pedicels articulated at base; sepals 5 DC., Prodr. 15(2): 764 (1866); Pax & K. Hoffm., Pflanzenr.
or 6, imbricate, persistent and accrescent in fruit; 147: 267 (1914); Liogier, Descr. Fl. Puerto Rico 2: 352, fig.
petals 0; disk annular, sometimes angled; ovary 3- 59–4 (1988); Burger & Huft, Fieldiana Bot. n.s. 36: 62
(1995); Radcl.-Sm., Gen. Euphorb.: 156, fig. 16 (2001).
locular, glabrous, ovules with inner integument
moderately thick, outer integument thin; stylodia Dioecious trees; indumentum scanty, simple.
basally connate, unlobed. Fruits thick-walled, Leaves triplinerved, biglandular abaxially at
(2)3(4)-locular, loculicidally and septicidally base, entire or remotely crenulate; stipules 0.
dehiscent; columella persistent. Seeds ovoid; Inflorescences axillary, the staminate often
testa smooth, dry, thin-walled, hilum elliptic. fascicled, spiciform [flowers subsessile or shortly
Three or four closely related neotropical pedicellate]; bracts minute, eglandular; staminate
spp., from Costa Rica to Amazonian Brazil. flowers in glomerules, pistillate solitary. Stami-
Caryodendron orinocense H. Karst., the "inchi", nate flowers pedicellate, calyx closed in bud;
is cultivated for the seed oil in Colombia. lobes 3 or 4, valvate; petals 0; disk massive,
hirsutulous; stamens (5)6(8), filaments distinct;
97. Discoglypremna Prain anthers introrse, locules unequal; pollen grains
subspheroidal, 3-colporate, colpi inoperculate,
Discoglypremna Prain, Kew Bull. 1911: 317 (1911); Radcl.-
Sm., Fl. E. Trop. Afr. Euphorb. 1: 222, fig. 44 (1987); emarginate, endoaperture lalongate; sexine tectate,
Léonard, Fl. Afr. Centr. Euphorb. 3: 11, t. 1 (1996). granulate; pistillode 3-lobed, strigose-hirtellous.
110 G.L. Webster
Pistillate flowers pedicellate, articulate; sepals 4 (1912); Gagnep., Fl. Indochine 5: 465, t. 57 (7–11), 58 (1–2)
or 5, slightly imbricate, acute, persistent in fruit; (1926); Airy Shaw, Kew Bull. Add. Ser. 4: 25 (1975); Sevilla
& van Welzen, Blumea 46: 76–89, figs. 1–3 (2001).
petals 0; disk annular, pulviniform, hirsutulous;
ovary 3-locular; pubescent; inner and outer Dioecious (monoecious) trees or shrubs;
ovular integuments thin; stylodia unlobed or branches resinous; indumentum simple. Leaves
emarginate. Fruit capsular; columella persistent, alternate, lamina decurrent on the petiole, entire
broadly 3-winged. Seeds flattened, ecarunculate; or dentate; stipules deciduous. Inflorescences
exotesta fleshy, endotesta reticulate-striate. axillary, racemoid or spicate; bracts glumaceous,
A single sp., A. floribunda (Benth.) M€ ull. Arg., indurate, sometimes glandular, persistent, each
ranging from the Greater Antilles to Costa Rica subtending 1–10 staminate flowers or solitary pis-
and Brazil. tillate flowers. Staminate flowers subsessile or
short-pedicellate; calyx closed in bud, dehiscent
3. TRIBE AGROSTISTACHYDEAE (M€
ull. Arg.) into 2–5 valvate segments; petals (0)5–8, imbri-
G.L. Webster (1975). cate; disk segments (4)5(7); stamens (8)10
(13), biseriate, filaments distinct or basally
Dioecious (monoecious) trees or shrubs; indu- connate; anthers basifixed, introrse, minutely
mentum simple or 0; inflorescences racemoid or apiculate; pollen grains subspheroidal, angula-
spicate; the calyx closed in bud, splitting into 2–5 perturate, 3-colporate, colpi inoperculate with
valvate lobes; petals 3–8, imbricate in bud [except narrow to broad margo, endoaperture lalongate,
Cyttaranthus]; disk receptacular or extrastaminal sexine tectate, intrareticulate; pistillode 2–3-fid or
and dissected; stamens 10–120; anthers introrse, entire (0). Pistillate flowers articulate-pedicellate;
connective enlarged or apiculate, thecae often sepals (4) 5, imbricate, entire or denticulate, per-
pendulous; ovary 3-locular, often tomentose; sty- sistent in fruit; petals 5, longer than sepals; disk
lodia distinct, bifid, sometimes lacerate; fruit cap- annular to 5-lobed, sometimes with staminodes;
sular; seeds carunculate, not fleshy. ovary (2)3-locular, glabrous or pubescent; ovules
Four genera, two African and two Asian. pachychalazal, inner integument thick, outer
Nowicke et al. (1999) stated that Cyttaranthus integument thin; stylodia distinct or basally con-
and Chondrostylis differ in pollen structure from nate, bifid, papillate. Fruits capsular, thin-valved;
Agrostistachys and Pseudagrostistachys, implying columella persistent, narrowly 3-winged. Seeds
that the Agrostistachydeae may not be monophy- subglobose, ecarunculate; testa smooth; hilum
letic. This is confirmed by the molecular data of deltoid-reniform, large.
Wurdack et al. (2005). Ten spp., from India and Sri Lanka to New
Guinea, five in Malesia.
KEY TO THE GENERA OF AGROSTISTACHYDEAE
1. Petals +; anther connective narrow; flowers in spici- 100. Pseudagrostistachys Pax & K. Hoffm.
form or racemoid thyrses; anthers extrorse; pistillate
disk annular 2 Pseudagrostistachys Pax & K. Hoffm., Pflanzenr. 147, VI:
96, fig. 18 (1912); Léonard, Fl. Congo 8(1): 183, fig. 15
– Petals 0; anther connective flattened, broader than (1962); Airy Shaw, Kew Bull. 36: 248, t. 1 A-G (1981);
high; flowers in thyrses or panicles; pistillate disk 5-
Radcl.-Sm., Fl. E. Trop. Afr. Euphorb. 1: 166, fig. 30
lobed 102. Chondrostylis
(1987), Fl. Zambesiaca 9(4): 136 (1996).
2. Dioecious; staminate petals imbricate in bud; petals
present in pistillate flower 3 Dioecious shrubs or trees; indumentum simple,
– Monoecious; staminate petals not imbricate in bud; sparse except in inflorescences. Leaves subentire
pistillate flowers apetalous; leaves with laminar glands
or somewhat dentate, with dispersed laminar
101. Cyttaranthus
3. Staminate disk dissected; stamens 8–10; stipules distinct, glands and 2–4 glands near junction with petiole;
leaves lacking laminar glands 99. Agrostistachys stipules connate, sheathing, deciduous, leaving a
– Staminate disk receptacular; stamens 20–55; stipules circular scar. Inflorescences axillary or ramiflor-
connate, leaving an annular scar; leaves with laminar ous, sessile to short-pedunculate, flowers usually
glands 100. Pseudagrostistachys 1 per bract; bracts glumaceous, ciliate, persistent,
eglandular. Staminate flowers pedicellate, articu-
99. Agrostistachys Dalzell late near base; calyx closed in bud, splitting into
Agrostistachys Dalzell, Hook. J. Bot. Kew Gard. Misc. 2: 41 2–5 valvate lobes; petals 5–8, distinct, imbricate,
(1850); Pax & Hoffm., Pflanzenr. 147, VI: 98, figs. 19, 20 longer than sepals; disk receptacular, pubescent;
Euphorbiaceae 111
stamens 20–55, filaments distinct, erect in bud; Monoecious shrubs; indumentum simple to
anthers extrorse, connective thickened and nearly 0. Leaves subsessile, serrate, usually
glandular, thecae pendulous; pollen grains sub- minutely punctate with dispersed embedded
prolate, 3-colporate, colpi inoperculate, emargin- glands and biglandular at base; stipules decidu-
ate, endoaperture obscure; sexine tectate- ous. Inflorescences axillary (ramiflorous), panic-
reticulate; pistillode small or 0. Pistillate flowers ulate, unisexual; bracts glumaceous, entire,
pedicellate; sepals mostly 4 or 5, scarcely imbri- eglandular. Staminate flowers short-pedicellate;
cate, entire, persistent in fruit; petals 4 or 5; disk calyx splitting into 3 or 4 valvate segments; petals
thick, annular, pitted; ovary 3-locular, tomentose; 0; disk of discrete interstaminal segments;
stylodia distinct, bipartite, branches papillose. stamens 50 +; anthers basifixed, introrse, connec-
Fruit capsular, 3-lobed, valves crustaceous; colu- tive enlarged, thecae divergent; pollen grains
mella persistent. Seeds subglobose, ecarunculate, spheroidal, angulaperturate, 3-colporate, colpi
hilum elliptic; testa smooth, shiny. inoperculate, emarginate, endoaperture lalon-
Two spp., West Africa, one of these reaching gate, sexine tectate-perforate and microverrucate;
Uganda and Zambia. pistillode 0. Pistillate flowers subsessile; sepals 5,
imbricate, connate into a cup; petals 0; disk annu-
101. Cyttaranthus Léonard lar, tomentose; ovary 3-locular, pubescent; stylo-
Cyttaranthus Léonard, Bull. Jard. Bot. Brux. 25: 286 dia distinct, bifid, branches adaxially fimbriate.
(1955); Fl. Congo Rwa.-Bur. 8(1): 180, t. 12 (1962). Fruit capsular, valves thin; columella persistent,
slender, trigonous. Seeds spheroidal, ecaruncu-
Monoecious shrubs; indumentum simple. Leaves late, testa smooth.
subentire to crenate or dentate, trinerved at base Two spp., distributed from Thailand to
and pinnately veined, with scattered discoid Sumatra and Borneo.
glands abaxially, stipellate at junction with peti-
ole; stipules minute, deciduous. Inflorescences 4. TRIBE SPHYRANTHEREAE Radcl.-Sm. (2001).
axillary, usually unisexual, spiciform; bracts con-
cave, glumaceous, persistent. Staminate flower A monotypic tribe containing only the genus
pedicellate, articulate at base; calyx splitting into Sphyranthera.
2 or 3 valvate lobes; petals 3(6–8), not imbricate,
103. Sphyranthera Hook. f.
shorter than calyx lobes; disk segments glabrous,
intrastaminal, on the convex receptacle; stamens Sphyranthera Hook. f., Hook. Ic. Pl. 18: t. 1702 (1887);
25–40, filaments distinct; anthers extrorse, con- Chakrabarty & Vasudeva Rao, J. Econ. Tax. Bot. 5: 959
(1984); 6: 429 (1985); Radcl.-Sm., Gen. Euphorb.: 135,
nective apiculate, thecae pendulous; pollen grains fig. 12 (2001).
prolate spheroidal, 3-colporate, colpi inopercu-
late, scarcely marginate; sexine irregularly tec- Dioecious shrubs; indumentum simple, scanty.
tate-perforate; pistillode 0. Pistillate flowers Leaves glandular; stipules 0. Inflorescences axil-
pedicellate, pedicel distally dilated and articulate lary or extraaxillary, sometimes fasciculate,
at base; sepals 3, connate into a cup, persistent in pedunculate and subumbellately capitulate, the
fruit; petals 0; disk swollen, appearing as a gyno- staminate many-flowered capitula, the pistillate
phore; ovary 3-locular, pubescent; stylodia 1–4-flowered; bracts crowded at apex of pedun-
slightly connate basally, deeply bifid. Fruit capsu- cle. Staminate flowers pedicellate, basally articu-
lar, invaginated between cocci; columella persis- lated; sepals (3)4(5), valvate; petals 4, entire to
tent. Seeds oblong to subglobose, ecarunculate, bifid; disk segments 4, apically bilobed; stamens
testa smooth. (8–)12–20, filaments distinct; connectives
A single sp., C. congolensis Léonard, rainfor- enlarged and minutely bifid, anthers subglobose;
ests and semi-deciduous forests in the Congo. pollen prolate, 3-colporate, sexine reticulate; pis-
tillode 0. Pistillate flowers pedicellate; sepals 3 or
102. Chondrostylis Boerl. 4 (5), valvate or open in bud, entire, deciduous
in fruit; petals 0; disk segments 3 or 4; ovary
Chondrostylis Boerl., Ic. Bogor. 1: t. 23 (1897); Pax & 3-locular, pubescent; stylodia distinct, bifid,
K. Hoffm., Pflanzenr. 147, VII: 15 (1914); Airy Shaw,
Kew Bull. 14: 358 (1960), Kew Bull. Add. Ser. 4: 69
somewhat lacerate. Fruits capsular; columella
(1975), Kew Bull. 36: 276, fig. 3C (1981); Sevilla & van persistent, emarginate. Seeds globose, ecaruncu-
Welzen, Blumea 46: 89–93, fig. 4 (2001). late, testa smooth.
112 G.L. Webster
Two spp., endemic to the Andaman/Nicobar – Stamens 2 or 3; stylodia stigmatiform; seeds minutely
archipelago. The curious malleiform anthers sug- carunculate; leaf blade not stipellate; inflorescences
gest a possible relationship with Chondrostylis. axillary or ramiflorous 108. Bocquillonia
5. TRIBE ALCHORNEEAE (Hurus.) Hutch. (1969). 5a. SUBTRIBE ALCHORNEINAE Hurus. (1954).
Dioecious (monoecious) trees or shrubs; wood Indumentum simple or stellate; staminate inflor-
rays with lysigenous canals; indumentum simple escences axillary; stamens 2–10; pollen exine with
or stellate; leaves entire or dentate, sometimes complex infratectum of columellae; pistillate
stipellate or with laminar glands; stipules mostly sepals eglandular; stylodia mostly unlobed, often
deciduous or obsolete; inflorescences terminal or dilated or stigmatiform; ovary 2–3(4)-locular.
axillary; staminate flowers subsessile; calyx closed Six genera with c. 70 spp. in both the Neo-
in bud, splitting into 2–5 valvate segments; petals tropics and Paleotropics. Generic limits within
0; disk + or 0; stamens (2–)4–60; pollen grains 3- the Alchorneae are still controversial; Orfilea,
colporate, colpi operculate; pistillode rudimen- Coelebogyne, and Aparisthmium possibly might
tary or 0; pistillate flowers subsessile; sepals 3–8, be included within Alchornea s. lat.
imbricate; petals 0; disk mostly rudimentary or 0;
ovary 2–3(4)-locular; stylodia entire to multifid; 104. Orfilea Baill.
fruits capsular; columella persistent; seeds carun- Orfilea Baill., Étude Gén. Euphorb.: 452 (1858); Pax & K.
culate or ecarunculate, testa smooth or tubercu- Hoffmann, Pflanzenr. 147. VII: 253 (1914, under Lautem-
late, not fleshy. bergia); Coode, Fl. Mascar. 160: 55, fig. t. 11 (1982, under
Lautembergia); Radcl.-Sm., Gen. Euphorb.: 192 (2001);
The Alchorneae include 10 genera grouped Schatz, Generic Tree Fl. Madag.: 162, fig. 166 (2001).
into 3 subtribes; the monogeneric subtribe Mar-
eyopsinae is added provisionally because Mar- Dioecious (monoecious) trees or shrubs; indu-
eyopsis is resolved in the Alchorneoid clade of mentum simple. Leaves entire or crenulate,
Wurdack et al. (2005). biglandular at base or eglandular; stipules subu-
late, subpersistent. Inflorescences terminal and
KEY TO THE GENERA OF ALCHORNEEAE axillary, spiciform or paniculate; pistillate bracts
trifid. Staminate flowers subsessile; calyx closed
1. Stylodia bifid; indumentum stellate; stamens 15–60
109. Conceveiba
in bud, splitting into 2–4 valvate segments;
– Stylodia unlobed (or if bifid then indumentum simple); petals and disk 0; stamens (4)5–10, filaments
stamens 2–12; seeds ecarunculate or caruncle, minute 2 basally connate; anthers 2-celled; pollen grains
2. Indumentum stellate; ovary 2-locular; stylodia unlobed, prolate spheroidal, angulaperturate, 3-colporate,
elongated, slender; pollen exine coarsely rugulose operculate, emarginate; endoapertures lalongate;
105. Alchornea tectum microrugulose, microverrucate; pistillode
– Indumentum simple; ovary 3-locular; pollen exine 0. Pistillate flowers sessile or subsessile; sepals
finely tectate-punctate 3
3. Dioecious or monoecious; stamens 7–12 4 5 (6), entire, persistent in fruit; disk of 5 segments
– Monoecious; stamens up to 8 5 or 0; ovary 3-locular, glabrous; stylodia connate
4. Dioecious; staminate disk of 8 segments in 4 pairs; basally, bifid. Fruits capsular; columella persis-
stamens 8; fruit indehiscent 111. Mareyopsis tent. Seeds rounded, ecarunculate, testa smooth.
– Monoecious; staminate disk 0; stamens 7–12; fruit Four spp. from Madagascar and Mauritius.
dehiscent 110. Aubletiana
5. Stylodia distinctly bifid; leaf blades pinnately veined, 105. Alchornea Sw. Fig. 24
not stipellate; stamens 8 104. Orfilea
– Stylodia entire to emarginate; stamens 2–8; leaf blades Alchornea Sw., Prodr.: 98 (1788), Fl. Ind. Occ. 2: 1153, t. 24
pinnately to palmately veined 6 (1800); Thin, Tâp Chi Sinh Hoc 6 (3): 26 (1984); Burger &
6. Stamens usually 8, distinct; stigmas smooth; pistillate Huft, Fieldiana Bot. II. 36: 59 (1995); Radcl.-Sm., Fl. Zam-
sepals glandular at base; leaf blades spinose-dentate besiaca 9 (4): 151, t. 31 (1996), Gen. Euphorb.: 192 (2001);
106. Coelebogyne Secco, Fl. Neotrop. 93: 55–134 (2004); van Welzen &
– Stamens 2–4, filaments basally connate; leaf blades not Bulalacao, Syst. Bot. 32: 803–818 (2007), Males. spp.
spinose-dentate 7 Bossera Leandri (1962).
7. Stamens 4; stylodia elongated, apically dilated;
seeds ecarunculate; leaf blade stipellate; inflorescences Dioecious (monoecious) trees and shrubs;
terminal 107. Aparisthmium indumentum simple or stellate. Leaves entire to
Euphorbiaceae 113
sepals usually (3)4(6), imbricate, entire or den-

ticulate, persistent in fruit; petals 0; disk obso-
lete; ovary 2(5)-locular, usually pubescent
(cristate); ovules with outer integument vascular-
ized; stylodia nearly distinct, unlobed (emargin-
ate). Fruits capsular, smooth or tuberculate;
columella persistent. Seeds subglobose or elliptic,
ecarunculate (with rudimentary caruncle), testa
smooth or tuberculate; mesotesta vascularized.
n ¼ 9, 18.
About 42 spp., distributed in the Neotropics
(23 spp.), Africa and Madagascar, and in Asia and
Malesia. The American taxa differ from the Old
World taxa in a number of characters such as
indumentum and carpel number, and the palyno-
logical studies of Takahashi et al. (2000) indicate
that the pollen grains of neotropical species have
a prominently vermiculate-rugose exine, in dis-
tinct contrast with the punctate-spinulose tectum
of the paleotropical species. Bossera was distin-
guished by the elevated number of stamens (10)
and the cristately ornamented ovary, which is
also present in A. alnifolia.
106. Coelebogyne Js. Sm.

Coelebogyne Js. Sm., Proc. Linn. Soc. London 1: 41 (1839);
Baill., Étude Gén. Euphorb.: 416 (1858); Pax & K. Hoffm.,
Pflanzenr. 147, VII: 255, fig. 38 (1914); Radcl.-Sm., Gen.
Euphorb.: 194 (2001).
Dioecious shrubs; indumentum simple. Leaves
dentate, with basal and scattered laminar glands;
stipules persistent. Inflorescences axillary, spici-
Fig. 24. Euphorbiaceae-Acalyphoideae. Alchornea cordi-
folia. A Branchlet. B, C Base of upper resp. lower leaf
form; bracts eglandular. Staminate flowers sub-
surface with extrafloral nectaries. D Part of staminate sessile; calyx closed in bud, splitting into usually
inflorescence, flowers in bud stage. E Staminate flower. 4 valvate segments; petals and disk 0; stamens
F Tip of pistillate inflorescence, with anthetic flowers. 8, filaments distinct; anthers muticous; pollen
G Pistillate flower. H Fruit. I Seed. (Radcliffe-Smith
1987; drawn by Judy Dunkley) grains spheroidal, 3-colporate, colpi operculate,
sexine tectate-perforate, microverrucate; pistil-
dentate, with laminar glands abaxially, sometimes lode 0. Pistillate flowers subsessile; sepals 5
stipellate; stipules persistent to deciduous, some- or 6, imbricate, entire, basally glandular; petals
times rudimentary. Inflorescences axillary, spici- and disk 0; ovary (2)3-locular; glabrous; stylodia
form, often compound; bracts eglandular. stigmatiform. Fruits capsular; columella persis-
Staminate flowers subsessile, in glomerules; tent. Seeds ecarunculate; testa smooth.
calyx closed in bud, splitting into 2–5 valvate seg- One or two spp., Australia (Queensland, NSW),
ments; petals 0; disk receptacular, confluent with very similar to the paleotropical spp. of Alchornea
base of filaments; stamens (3–)6–8(10), filaments (Cladodes) and often merged with that genus.
basally slightly connate; anthers introrse or
extrorse, muticous; pollen grains spheroidal, 107. Aparisthmium Endl.
angulaperturate, 3-colporate, colpi operculate,
Aparisthmium Endl., Gen. Pl.: 1112 (1840); Webster, Fl.
endoapertures small and obscure, sexine tectate- Venez. Guayana 5: 99, fig. 97 (1999); Radcl.-Sm., Gen.
rugulose, microverrucate; pistillode 0 or rudi- Euphorb.: 194 (2000); Secco, Fl. Neotrop. 93: 134–143,
mentary. Pistillate flowers sessile or pedicellate; fig. 56 (2005).
114 G.L. Webster
Dioecious trees or shrubs; indumentum simple. glandular; ovary 2–3-locular, inner integuments
Leaves glandular-crenate, with basal laminar equaling or thicker than outer integuments; sty-
glands, stipellate at junction with petiole; stipules lodia bifid; seeds carunculate.
deciduous. Inflorescences terminal; staminate Two genera, one neotropical, the other African.
spicate-paniculate, pistillate racemose; bracts
biglandular. Staminate flowers glomerulate, sub- 109. Conceveiba Aubl.
sessile; calyx closed in bud, splitting into 3(4)
Conceveiba Aubl., Hist. Pl. Gui.: 923, t. 353 (1775); M€ ull.
valvate segments; petals and disk 0; stamens 3–5 Arg. in DC., Prodr. 15(2): 895 (1866), Fl. Brasil. 11(2): 370,
(10), filaments basally connate; anthers latrorse, t. 55 (1874); Pax & K. Hoffm., Pflanzenr. 147, VII: 214, fig.
muticous; pollen grains subspheroidal, 3-colpo- 32 (1914); Macbride, Field Mus. Nat. Hist., Bot. 13 (3A, 1):
rate, colpi operculate; sexine finely tectate-perfo- 152 (1951); Jablonski, Mem. N. Y. Bot. Gard. 17: 131
rate, microverrucate; pistillode 0. Pistillate flowers (1967); Secco, Fl. Neotrop. 93: 143–177 (2005).
Gavarretia Baill. (1860).
pedicellate, with 2 basal glands; sepals 4, valvate, Conceveibastrum (M€ ull. Arg.) Pax & K. Hoffm. (1914).
entire, persistent in fruit; petals and disk 0; ovary Veconcibea (M€ ull.Arg.) Pax & K. Hoffm. (1914).
3-locular, ovules anatropous, smooth, pubescent;
inner integuments thick, outer integuments thin- Dioecious (monoecious) trees or shrubs; indu-
ner; stylodia thick, dilated, shallowly bifid at tip, mentum stellate and sometimes simple as well.
papillose. Fruits capsular; columella persistent. Leaves glandular-crenate or entire; stipules per-
Seeds ellipsoid, ecarunculate, testa smooth. sistent or deciduous, or 0. Inflorescences terminal
A single sp., A. cordatum (A. Juss.) Baill., or axillary, the staminate paniculate or racemose,
widespread in tropical South America. the pistillate racemose or spicate; bracts often
biglandular. Staminate flowers subsessile or
108. Bocquillonia Baill. pedicellate; calyx closed in bud, splitting into
3 or 4 valvate segments or lobes; petals and disk
Bocquillonia Baill., Adansonia I, 2: 225 (1862); Airy Shaw, 0; outer staminodes sometimes +; stamens 7–60,
Kew Bull. 29: 321 (1974); McPherson & Tirel, Fl. Nouv.
Calédonie 14(1): 114–143, t. 23–29 (1987). filaments distinct or slightly connate at base; up
to 35 inner stamens sometimes infertile; anthers
Dioecious (monoecious) trees or shrubs; indu- extrorse or introrse, connective sometimes
mentum simple, but usually very sparse or 0. enlarged; pollen grains oblate to prolate spheroi-
Leaves dentate, with dispersed laminar glands; dal, 3-colporate, colpi operculate, endoapertures
stipules persistent but often minute. Inflores- lalongate, sexine finely tectate-perforate, micro-
cences axillary, spiciform, sometimes reduced to verrucate; pistillode 0 (+). Pistillate flowers pedi-
glomerules; bracts persistent. Staminate flow- cellate; sepals 3–8, distinct or connate, glandular
ers subsessile; calyx closed in bud, splitting into at base, deciduous in fruit; petals and disk 0;
2 or 3 valvate segments; petals and disk 0; ovary 2–3-locular, pubescent; ovules anatropous,
stamens 2–4, filaments basally connate; anthers inner and outer integuments moderately thick;
latrorse or extrorse, muticous; pollen grains sub- stylodia usually completely distinct and distally
oblate to prolate spheroidal, 3-colporate, colpi to deeply bipartite. Fruits capsular, smooth to
operculate, sexine finely tectate-perforate, micro- rugose; columella not persistent. Seeds oblong,
verrucate; pistillode vestigial or 0. Pistillate flow- carunculate, hilum triangular, testa smooth.
ers subsessile; sepals 4 or 5, imbricate, entire, About 12 spp., tropical Central and South
persistent in fruit but inconspicuous; petals America.
and disk 0; ovary 3-locular, pubescent; ovules
anatropous, inner integuments moderately 110. Aubletiana J. Murillo-A. Fig. 25
thick, outer thin; stylodia unlobed, ovate,
Aubletiana J. Murillo-A., Rev. Colomb. Cienc. 24: 360
dilated. Fruits capsular; columella persistent. (2000).
Seeds oblong, ecarunculate (?), testa smooth. Conceveiba Aubl. quoad Thomas, Ann. Missouri Bot.
Fourteen spp., all endemic to New Caledonia. Gard. 77: 856, fig. 1 (1990) et Breteler & Mennega, Bull.
Jard. Bot. Belg. 63: 209–217, t. 1 (1994).
5b. SUBTRIBE CONCEVEIBINAE G.L. Webster (1975).
Monoecious trees; indumentum simple. Leaves
Leaf blades not stipellate; indumentum stellate; dentate with often gland-tipped teeth; the blade
floral disk 0; stamens 15–60; pistillate sepals often with two glands at the junction of the petiole;
Euphorbiaceae 115
splitting into 3 1-seeded cocci, leaving no column.

Seeds ellipsoid, with a triangular hilum, ecarun-
culate, exarillate.
Two West African spp., Cameroun, Gabon.
5c. SUBTRIBE MAREYOPSINAE G.L. Webster (2004).

Dioecious; indumentum simple; leaves alternate,
lamina puncticulate; inflorescences axillary; flow-
ers subsessile; staminate disk segments and
stamens 8; fruits indehiscent, seeds ecarunculate.
A monogeneric African tribe.
111. Mareyopsis Pax & K. Hoffm.

Mareyopsis Pax & K. Hoffm., Pflanzenr. 147, XIV: 13
(1919); Léonard, Bull. Jard. Bot. Nat. Belg. 65: 15, fig. 3
(1996), Fl. Afr. Centr. Euphorb. 3: 16, t. 2 (1996); Breteler,
Bull. Jard. Bot. Belg. 66: 133, figs. 1, 3 (1997); Radcl.-Sm.,
Gen. Euphorb.: 223 (2001).
Leaves translucent-punctate with scattered
embedded glands, glandular-dentate; stipules
deciduous. Inflorescences axillary, often rami-
florous, generally fasciculate, spiciform; bracts
entire, persistent. Staminate flowers subsessile;
calyx closed in bud, splitting into 3–5 valvate
segments; petals 0; disk-segments 8, in 4 pairs,
extrastaminal; stamens 8, biseriate, filaments dis-
tinct, flattened and ligulate; anthers muticous or
apiculate, connective somewhat enlarged, thecae
pendulous; pollen grains 3-angled, 3-colporate,
Fig. 25. Euphorbiaceae-Acalyphoideae. Aubletiana lep-
colpi granulose, neither marginate nor opercu-
tostachys. A Branchlet. B Same, with staminate inflores- late; sexine tectate-perforate and rugulose,
cence. C Leaf margin with gland seen from above. D Same, microverrucate; pistillode prominent, 2- or 3-
from beneath. E Base of leaf blade with glands. F Detail of lobed. Pistillate flowers subsessile; sepals 3–4(5),
inflorescence. G Staminate flower. H Androecium and
perianth of staminate flower. I Pistillate flower. J Fruit.
basally connate, open in bud; petals 0; disk 3- or
(Breteler 1994; drawn by H. de Vries) 4-lobed; ovary 2–3(4)-locular, pubescent; stylodia
nearly distinct, unlobed, papillate. Fruits indehis-
cent, exocarp coriaceous. Seeds ecarunculate,
testa smooth.
stipules lateral. Staminate inflorescences axillary Two spp. of tropical West Africa. Mareyopsis
panicles with contracted 3–4-flowered cymes; pis- was incorrectly treated by Webster (1994) as a
tillate inflorescences terminal spikes or flowers synonym of Mareya, but Léonard (1996) conclu-
solitary. Staminate flowers: sepals 4 or 5, for sively demonstrated that the two genera are not
lower half connate, valvate; petals and disk 0; closely related.
stamens 7–12, sometimes with a few staminodes;
pollen tricolporate-operculate [Murillo fide 6. TRIBE RICINEAE Bartl. (1830).
Punt]; pistillode +, glabrous or pubescent. Pistil-
late flowers solitary, sepals sometimes basally Monoecious or dioecious trees or shrubs; indu-
glandular; petals and disk 0; ovary 3-carpellate; mentum stellate or 0; leaves simple to deeply
ovule 1 per locule; stylodia at the base connate lobed, dentate; inflorescences terminal or oppo-
into a style provided with 3 undivided or divided site leaves; spiciform or paniculate; staminate
style branches. Fruit papillose, 3-lobed, dehiscent, calyx closed in bud, splitting into valvate
116 G.L. Webster
segments; petals and disk 0; stamens 30–1,000, persistent. Seeds oblong, carunculate, testa pale,
filaments distinct or connate; anthers biloculate; rugulose.
pollen grains spheroidal, 3–5-colpate, colpi inop- Two variable spp., Australia.
erculate, emarginate; pistillate sepals 3–5, imbri-
cate; petals and disk 0; ovary 3-locular; stylodia 113. Ricinus L.
bifid; fruits capsular; seeds carunculate. Ricinus L., Sp. Pl.: 1007 (1753), Gen. Pl. ed. 5: 437 (1754);
The genera comprised in this tribe earlier had M€ ull. Arg. in DC., Prodr. 15(2): 1016 (1866); Pax &
been included in two different subtribes of the K. Hoffm., Pflanzenr. 147, XI: 119 (1919); Webster,
Acalypheae (Webster 1994), from which they dif- J. Arnold Arb. 48: 379, fig. 4 (1967); Purseglove, Trop.
fer in many respects such as the carunculate seeds Crops, Dicot. 1: 180, fig. 27 (1968); van Welzen, Blumea
43: 151 (1998).
and terminal inflorescences. The exine structure,
which is very similar in both genera (Takahashi Monoecious shrub or tree, herbaceous in temper-
et al. 2000), led Webster to unite them in a tribe. ate regions; indumentum simple or 0. Leaves pal-
The molecular data of Wurdack et al. (2005) place mately lobed, peltate, denticulate to serrulate;
both genera in the alcalyphoid clade A4, but Rici- petiole elongated, glandular basally and at junc-
nus as sister to Speranskia, and Adriana as sister tion with blade; stipules connate into a deciduous
to Monotaxis, though both with low support. sheath, leaving a circumaxial scar. Inflorescences
Available data are insufficient for improving this terminal, sometimes pseudo-axillary, paniculate;
unsatisfactory placement. staminate cymules proximal, distal cymules
bisexual or pistillate; bracts papery, glandular at
KEY TO THE GENERA OF RICINEAE base. Staminate flowers pedicellate, articulate in
1. Dioecious; filaments distinct; pistillate sepals persistent the middle; calyx closed in bud, splitting into 3–5
in fruit; stipules distinct, glandular 112. Adriana valvate segments; petals and disk 0; stamens very
– Monoecious; filaments connate into fascicles; pistillate numerous, to 1,000, filaments partially connate
sepals deciduous; stipules connate, deciduous and irregularly branched; anthers introrse,
113. Ricinus
locules distinct and subglobose; pollen grains
subspheroidal, angulaperturate, 3-colporate,
112. Adriana Gaud.
colpi not operculate, sexine finely tectate-perfo-
Adriana Gaud., Ann. Sci. Nat. Paris 5: 223 (1825); Pax, rate/reticulate and microverrucate; pistillode 0.
Pflanzenr. 147, II: 17 (1910); Airy Shaw, Kew Bull. 35: 589 Pistillate flowers pedicellate; sepals 3–5, entire,
(1980); C.L. Gross & M.A. Whalen, Austral. Syst. Bot. 9:
757, figs. 1–4 (1996); Radcl.-Sm., Gen. Euphorb.: 202, deciduous; petals and disk 0; ovary 3-locular,
fig. 25 (2001). prominently echinate, glabrous; ovules pachycha-
lazal, with inner integuments thick, outer integu-
Dioecious shrubs; indumentum stellate or 0. ments thin; stylodia basally connate, bifid,
Leaves alternate or opposite, petiolate or sessile, conspicuously papillate. Fruits capsular; colu-
pinnately or palmately veined, coarsely dentate; mella persistent, distally winged. Seeds ellipsoid-
stipules persistent, glandular. Inflorescences compressed, carunculate, testa smooth. 2n ¼ 20.
terminal or opposite the leaves, spiciform, the The single sp., R. communis L., is probably
pistillate shorter and more condensed than the native originally to east Africa but early dispersed
staminate; bracts glandular. Staminate flowers by man to India. It is highly variable due to its
subsessile; calyx splitting into 4–5 valvate seg- cultivation for the oil from the seeds ("castor
ments; petals and disk 0; stamens numerous, fila- beans"). The combination of a thick inner integ-
ments distinct; anthers extrorse, linear, much ument, a pachychalazal ovule, and an arillate seed
longer than filaments, connective conspicuously makes Ricinus unique among Acalyphoids
apiculate; pollen grains spheroidal, 3–5-colpo- (Tokuoka and Tobe 2003).
rate, colpi inoperculate, sexine finely tectate-per-
forate and microverrucate; pistillode 0. Pistillate 7. TRIBE ACALYPHEAE Dumort. (1829).
flowers pedicellate; sepals 6–8, imbricate, persis-
tent in fruit; petals and disk 0; ovary 3-locular, Monoecious or dioecious trees, shrubs, or herbs;
stellate-tomentose, sometimes muricate; ovules indumentum simple or stellate; leaves simple or
with outer integuments vascularized, equaling lobed, pinnately or palmately veined, glandular
inner integuments in thickness; stylodia bipar- or eglandular; inflorescences terminal or axillary,
tite, coarsely papillate. Fruits capsular; columella spiciform or racemose to paniculate, unisexual or
Euphorbiaceae 117
bisexual; staminate calyx closed in bud, splitting 8. Stylodia bifid; inflorescences axillary, racemoid or
into 2–5 valvate segments; disk intrastaminal, spiciform; staminate flowers pedicellate
7a. Cleidiinae
interstaminal, or 0; stamens 4–many; anthers
– Stylodia unlobed; inflorescences terminal, dichasial;
sometimes apiculate, anther sacs sometimes pen- staminate flowers subsessile
dulous; pollen grains spheroidal, 3–4-colpo- 7e. Avellanitinae
rate, colpi inoperculate; sexine tectate-perforate
or rugulose, usually microverrucate; pistillode 7a. SUBTRIBE CLEIDIINAE G.L. Webster (1975).
usually 0; pistillate sepals mostly 3–6, distinct;
petals 0; disk cupular or 0; ovary 2–4-locular; Monoecious or dioecious; indumentum simple;
stylodia unlobed to multifid or lacerate; fruits leaves with laminar glands, stipulate; inflores-
capsular or drupaceous; seeds carunculate or cences axillary, spiciform or racemose; staminate
ecarunculate; testa dry or fleshy. flowers without petals or disk, sepals 3 or 4,
This is the largest tribe in the Acalyphoideae, stamens 25–80, anthers 2(4)-celled, usually apic-
with 9 subtribes including a total of 28 genera, ulate; pollen grains 3-colporate, colpi not opercu-
and over 1,000 species. With the removal of sub- late, sexine tectate-perforate or rugulose-
tribes Adrianinae and Ricininae, tribe Acalypheae vermiculate, microverrucate; pistillode 0; pistil-
is somewhat more homogeneous but still remark- late sepals 3–6, imbricate; ovary (2)3-locular; sty-
ably diverse, and further revision of its bound- lodia bifid, branches elongate; fruit capsular;
aries will be necessary. seeds ecarunculate, testa not fleshy.
Three genera, two restricted to southeast Asia
and Australia, Cleidion widespread. The very
KEY TO THE SUBTRIBES OF ACALYPHEAE
graceful, elongate stylodia of Cleidion and Wetria
1. Filaments connate, distally ramified; stylodia unlobed and their distantly placed pistillate flowers (see
7i. Lasiococcinae our Figs 26 and 27 and Fig. 27 in Radcliffe-Smith
– Filaments distinct or basally connate, not ramified; 2001) are strongly indicative of anemophily.
stylodia bifid or unlobed 2
2. Indumentum stellate and abaxial leaf surfaces with
minute beadlike glands [0.1 mm in diam.]; stylodia KEY TO THE GENERA OF SUBTRIBE CLEIDIINAE
unlobed, often plumose or laciniate; seed testa often 1. Staminate flowers subsessile, 1 per bract; stamens 15–25;
fleshy 7h. Rottlerinae pollen exine tectate-perforate 115. Sampantaea
– Indumentum mostly simple; abaxial leaf surfaces lack- – Staminate flowers pedicellate, usually 2 or more per
ing minute beadlike glands; stylodia lobed or unlobed; bract; stamens 25–80; pollen exine tectate-rugulose 2
seed testa dry or fleshy 3 2. Pistillate sepals valvate; anthers muticous, 2-celled,
3. Pollen exine finely reticulate; syles unlobed; seeds dehiscing longitudinally; stamens 25–30 114. Wetria
carunculate; mostly herbaceous 4
– Pistillate sepals imbricate; anthers apiculate, 4-celled,
– Pollen exine tectate-perforate to rugulose; stylodia dehiscing cruciately; stamens mostly 30–80
lobed or unlobed; seeds carunculate or ecarunculate; 116. Cleidion
mostly woody 5
4. Staminate sepals discrete; anthers muticous; pollen
grains distinctly colporate; cotyledons broader than 114. Wetria Baill. Fig. 26
radicle 7c. Mercurialinae Wetria Baill., Étude Gén. Euphorb.: 409 (1858); J. J. Smith,
– Staminate sepals connate; anthers apiculate; pollen Med. Dept. Landb. 10: 470 (1910); Pax & Hoffm., Pflan-
grains with rudimentary colpi; cotyledons scarcely zenr. 147,VII: 219 (1914); Forster, Austrobaileya 4:
broader than radicle 7d. Dysopsidinae 139–143, fig. 1 (1994); Webster, Ann. Missouri Bot.
5. Anthers apicifixed, thecae divergent from filament; Gard. 81: 86 (1994); van Welzen, Blumea 43: 156, fig. 5
stylodia unlobed, sometimes plumose or laciniate 6 (1998); Radcl.-Sm., Gen. Euphorb.: 209, fig. 27 (2001).
– Anthers basifixed or dorsifixed, thecae adnate to
connactive, not divergent from filament; stylodia bifid Dioecious trees; indumentum simple, scanty.
to multifid 7 Leaves subentire, with several glands near base;
6. Anthers not vermiform; pollen grains colporate; stami- stipules deciduous. Inflorescences axillary, slen-
nate disk present 7b. Claoxylinae
der, the staminate spiciform, nodes with solitary
– Anthers vermiform; pollen grains pseudo-porate; sta-
minate disk 0 7f. Acalyphinae
flowers or glomerules, the pistillate racemoid,
7. Staminate disk of interstaminal segments, pistillate with solitary flowers; bracts entire, acute, persis-
disk 0; seed testa fleshy 7g. Blumeodendrinae tent. Staminate flowers pedicellate; calyx closed
– Staminate and pistillate disk 0; seed testa dry 8 in bud, splitting into 3 or 4 valvate segments;
118 G.L. Webster
Lobreau-Callen (1996) and van Welzen (1998). In

the molecular analysis of Wurdack et al. (2005),
Wetria is resolved in a clade with the African/
Malagasy genera Argomuellera and Pycnocoma
(in the combined analysis) and in a clade with
the Thai genus Sampantaea (see note under fol-
lowing genus).
115. Sampantaea Airy Shaw

Sampantaea Airy Shaw, Kew Bull. 26: 328 (1972), Hook.
Ic. Pl. 38: t. 3717 (1974); Radcl.-Sm., Gen. Euphorb.:
212 (2001).
Dioecious trees; indumentum simple. Leaves sub-
sessile or short-petiolate, entire, with 1–3 small
glands near base; stipules deciduous. Inflores-
cences axillary, solitary or fascicled, spiciform,
flexuous; bracts persistent, eglandular. Staminate
flowers sessile; calyx closed in bud, splitting into
2 (3) valvate segments; petals and disk 0; stamens
15–25, filaments nearly suppressed; anthers sub-
sessile, oblong, minutely apiculate; pollen grains
subprolate, 3-colpate or rarely 3-colporate,
endoaperture minute; sexine tectate-perforate
and microverrucate; pistillode 0. Pistillate flowers
subsessile; sepals 5, imbricate, entire; petals and
disk 0; ovary 3-loculare, pubescent; stylodia
basally connate, erect, thin, elongate. Fruits not
seen.
Fig. 26. Euphorbiaceae-Acalyphoideae. Wetria austra- A single sp., S. amentiflora (Airy Shaw)
liensis. A Branchlet with flowers and fruits. B, C Pistillate Airy Shaw, known from Thailand and Cambodia.
flower in side and face view. D, E Staminate flower in side
and face view. F Stamen. G, H Fruit in side and face view. In spite of the pollen morphological diff-
I Capsule with seed. J, K Seed in ventral and lateral view. erence between Sampantaea and Cleidion/Wetria
L Seedling. (P.I. Forster 1994; drawn by W. Smith) (Fernández-González and Lobreau-Callen 1996),
the genus is strongly resolved as sister to Wetria
petals and disk 0; receptacle pubescent; stamens in the trnL-F analysis of Wurdack et al. (2005).
25–30, filaments distinct; anthers latrorse, muti-
cous or apiculate, 2-celled; pollen grains prolate 116. Cleidion Blume Fig. 27
spheroidal, (3)4-colporate, colpi emarginate, sex- Cleidion Blume, Bijdr. Fl. Ned. Ind.: 612 (1826); Leandri,
ine tectate-vermiculate, microverrucate; pistil- Adansonia II, 12: 193 (1972); A.C. Smith, Fl. Vitiensis Nova
lode 0. Pistillate flowers pedicellate; sepals 5, 2: 514, fig. 136 (1981); McPherson & Tirel, Fl. Nouv.-Caled.
imbricate, persistent in fruit; petals and disk 0; 14: 143–169, t. 30–34 (1987); Webster, Ann. Missouri Bot.
Gard. 81: 86 (1994); Philcox, Fl. Ceylon 9: 166 (1997); Kulju
ovary 3-locular, pubescent; ovules anatropous, & van Welzen, Blumea 50: 197–219, figs. 1–6 (2005); Qiu
inner integument thick, outer integument thin; Huaxing & Gilbert, Fl. China 11: 244 (2008).
stylodia bifid, branches thin, elongate. Fruits cap- ? Polyandra Leal, Arch. Jard. Bot. Rio de Janeiro 11: 63, fig.
sular, 3-lobed; columella persistent, angled and 1 (1961); Webster & Huft, Ann. Missouri Bot. Gard. 75:
apically dilated. Seeds spheroidal, testa smooth. 1087–1144 (1988); Secco, Fl. Neotrop. 93: 13, 178 (2004).
Two spp., one in southeast Asia and Malesia, Monoecious or dioecious trees or shrubs; indu-
the other in Queensland and New Guinea. In mentum simple or 0. Leaves often dentate,
Webster (1994) Wetria was placed in sub- abaxially with laminar glands; stipules persistent
tribe Cleidiinae, a disposition supported by the or deciduous. Inflorescences axillary, usually uni-
more recent studies of Fernández-González and sexual; staminate spiciform, sometimes compound,
Euphorbiaceae 119
ument thin; stylodia basally connate, erect,

deeply bifid, branches elongate, thin. Fruits cap-
sular; columella persistent. Seeds subglobose,
testa smooth.
About 25 spp., 5 neotropical [except the West
Indies], 2 in west Africa and Madagascar, the
remainder from India and China to islands in
the southwest Pacific, 12 endemic in New Caledo-
nia. In the molecular analysis (Wurdack et al.
2005), Cleidion is resolved as basal to Blumeoden-
dron, Macaranga and Mallotus, but lacks the vas-
cular bundles in the outer integuments of those
genera.
Polyandra, which is only known from a single
staminate specimen collected in central Amazo-
nia, possesses 4-locellate stamens, which led
Webster and Huft (1988) to propose a relation-
ship to Cleidion. Murillo-A. (2000) synonymised
Cleidion prealtum Croizat with Polyandra and
transferred it into Conceveiba. The identity of
Polyandra with C. prealtum is still uncertain
because the type of the latter species has only
solitary pistillate flowers, which would exclude it
from Conceveiba (Secco 2004). Although both
Polyandra and the type collection of C. prealtum
come from the same region of Central Amazonia
near Borba, a clarification of the status of Poly-
andra and confirmation of its place in subtribe
Cleidiinae would require pistillate material of
Polyandra.
Fig. 27. Euphorbiaceae-Acalyphoideae. Cleidion javani-
cum. A Branch of staminate plant. B Domatium on lower 7b. SUBTRIBE CLAOXYLINAE Hurus. (1954).
leaf surface. C Staminate bud. D Staminate flower.
E Anthers with cruciate cells. F Pistillate flower. G Fruit. Monoecious or dioecious trees, shrubs, or herbs;
H Seed. (Kulju and van Welzen 2005; drawn by J. van Os) indumentum simple; inflorescences axillary, race-
mose or paniculate; staminate sepals 2–5; disk
flowers in glomerules, pistillate racemose, some- annular and extrastaminal or segmented and inter-
times reduced to 1 or 2 flowers; bracts eglandular, staminal; stamens 5–40(200); filaments distinct;
inconspicuous. Staminate flowers subsessile or anthers extrorse, thecae discrete, erect from
pedicellate; calyx closed in bud, splitting into enlarged connective; pollen grains 3-colporate,
3 or 4 valvate segments; petals and disk 0; recep- colpi inoperculate, emarginate; pistillate sepals
tacle convex, stamens 20–100, filaments distinct, 2–4, open or imbricate; disk annular or dissected;
straight in bud; anthers with enlarged connec- ovary mostly 2- or 3-locular; stylodia unlobed,
tive, apiculate, 4-celled, on both sides 2 cells lacerate; fruits capsular, columella deciduous;
above each other, dehiscing introrsely and cruci- seeds ecarunculate, testa usually fleshy.
ately; pollen grains oblate to prolate spheroidal, Subtribe Claoxylinae is a paleotropical group
3-colporate, colpi inoperculate, sexine tectate- of 6 genera, best represented in Africa. Amyrea,
punctate, tectum rugulate or microverrucate; pis- included here in Webster (1994), is now placed in
tillode 0. Pistillate flowers long-pedicellate; sepals the Pycnocomeae. The systematic position of Mar-
3–5, imbricate, entire, persistent in fruit; petals eya is still controversial. Baillon (1860) originally
and disk 0; ovary (2)3-locular, pubescent; ovules noted similarities to Claoxylon and Pycnocoma,
anatropous, inner integument thick, outer integ- while M€uller (1866) saw an affinity with Alchornea.
120 G.L. Webster
Pax and Hoffman (1919) inserted Mareya in their sile or pedicellate; calyx closed in bud, splitting
Bernardiiformes [¼ Bernardieae], but it clearly into (2) 3 or 4 valvate segments; petals 0; disk of
does not belong in the same taxon as Bernardia. numerous interstaminal segments; stamens (9)
Webster (1994) assigned Mareya to Acalypheae 10–40, filaments distinct; anther connective bifur-
subtribe Claoxylinae, but Radcliffe-Smith (2001) cate, locules pendent; pollen grains subspheroi-
created a new subtribe Mareyinae to include both dal, angulaperturate, 3-colp(or)ate, inoperculate,
Mareya and Mareyopsis. Pollen evidence (Fernán- emarginate, endoapertures very narrow; sexine
dez-González and Lobreau-Callen 1996) indicates tectate-perforate, microverrucate; pistillode usu-
that Mareya, but not Mareyopsis, has pollen char- ally 0. Pistillate flowers subsessile or pedicellate;
acteristic of the Claoxylinae. In the molecular anal- sepals 3–5, imbricate or open in aestivation,
ysis of Wurdack et al. (2005), Mareyopsis goes with entire, persistent in fruit; petals 0; disk annular-
Alchorneopsis and Alchornea, and the Old World lobed; ovary 3-locular, pubescent; ovules with
Mareya is in a clade with New and Old World inner integument thick, outer integument thin;
Acalypha. Morphological similarities between the stylodia distinct, unlobed, plumose or lacini-
two genera include traits such as the laciniate ate. Fruits capsular; columella persistent. Seeds
stylodia, the pendulous anther locules, and seed subglobose, testa smooth, not fleshy.
and pollen morphology. It is remarkable that in Three spp., tropical West and Central Africa.
the molecular analysis Mercurialis is basal to the
Claoxylinae clade. 118. Claoxylon A. Juss.
Claoxylon A. Juss., Euphorb. Tent.: 43, t. 14 (1824); A.C.
KEY TO THE GENERA OF SUBTRIBE CLAOXYLINAE Sm., Fl. Vit. Nova 2: 516 (1981); Coode, Fl. Mascar. 160:
58, t. 12, 13 (1982); N. Rani & N.P. Balakr., Rheedea 5: 113,
1. Leaf blades stipellate; seed coat not fleshy 117. Mareya figs. 1–8 (1995); Florence, Fl. Polynésie Française 1: 54,
– Leaf blades not stipellate or, if so, then seed coat figs. 6–8 (1997); Forster, Austrobaileya 7: 451–472 (2007),
fleshy 2 Austral. spp.; Qiu Huaxing & Gilbert, Fl. China 11:
2. Indumentum stellate 119. Lobanilia 245–246 (2008).
– Indumentum simple 3 Claoxylopsis Leandri (1938); Radcl.-Sm., Kew Bull. 43:
3. Stipules usually persistent; buds perulate 625–647 (1988).
122. Erythrococca
– Stipules usually deciduous; buds not perulate 4 Dioecious (monoecious) trees and shrubs; indu-
4. Racemes interrupted; capsules crustaceous; leaf blades mentum simple or 0. Leaves entire or dentate,
stipellate 121. Micrococca often purplish when young, abaxially minutely
– Racemes uniformly floriferous; capsules coriaceous; punticulate and eglandular; stipules minute,
leaf blades not stipellate 5 deciduous. Inflorescences axillary, racemoid; sta-
5. Staminate disk of interstaminal segments; stamens minate bracts with 1–several flowers, pistillate
10–200 118. Claoxylon
– Staminate disk urceolate, extrastaminal; stamens 6–12
bracts with 1 flower; bracts eglandular. Staminate
120. Discoclaoxylon flowers pedicellate; calyx closed in bud, splitting
into 2–4 valvate segments; petals 0; disk of inter-
staminal segments; stamens (10–)20–30(200),
117. Mareya Baillon filaments distinct; anthers extrorse, muticous,
Mareya Baillon, Adansonia 1: 73 (1860); Adam, Mém. connective not enlarged, anther sacs distinct
Mus. Nat. Hist. Nat. Bot. 20: 495, fig. 184 (1971); Radcl.- from connective and erect; pollen grains spheroi-
Sm., Fl. E. Trop. Afr. Euphorb.:1: 216 (1987); J. Léonard, dal, 3–5-colporate, colpi granulose, sexine finely
Fl. Afr. Centr. Euphorb. 3: 18, t. 3 (1996).
tectate-perforate, microverrucate; pistillode 0.
Monoecious trees or shrubs; indumentum sim- Pistillate flowers pedicellate; sepals (2)3(4), dis-
ple. Leaves abaxially glandular-puncticulate, tinct, imbricate, entire, persistent in fruit; petals
entire to denticulate, adaxially with 2 or more 0; disk cupular or deeply lobed; ovary 2–3(4)-
embedded glands near base; petiole sometimes locular, glabrous or pubescent; ovules anatro-
adaxially stipellate at junction with blade, the pous, inner and outer integuments thin [4–6 cell
stipels deciduous; stipules deciduous. Inflores- layers]; stylodia nearly distinct, unlobed, papil-
cences axillary and sometimes terminal as well, lose. Fruits capsular or indehiscent, 3-lobed,
spiciform, slender; bisexual; with proximal stami- cocci reticulate; columella sometimes persistent.
nate and distal bisexual glomerules; bracts Seeds globose, exotesta fleshy, endotesta hard
eglandular, persistent. Staminate flowers subses- and rugose or foveolate. n ¼ 18, 31, 54.
Euphorbiaceae 121
About 75 spp., all Asiatic and Australasian 2-lipped, segments petaloid; ovary 2-locular,
except for 10 species in Madagascar. Claoxylopsis pubescent; stylodia abbreviated, dilated, lacerate.
was accepted as distinct by Webster (1994) and Fruits capsular, deeply 2-lobed, loculicidal; colu-
Radcliffe-Smith (1988, 2001), but it does not mella not persistent. Seeds globose, exotesta
appear to have any convincing differences. It fleshy, endotesta smooth or reticulate.
might be included in Claoxylon sect. Parviflora. Four spp., W African, three of them endemic
to the São Tomé Islands.
119. Lobanilia Radcl.-Sm.
121. Micrococca Benth.
Lobanilia Radcl.-Sm., Kew Bull. 44: 334, t. 1 (1989);
Schatz, Generic Tree Fl. Madagascar: 157 (2001). Micrococca Benth., Niger Fl.: 503 (1849), Gen. Pl.: 309
Claoxylon sect. Luteobrunnea Pax & K. Hoffm. (1914). (1880); Radcl.-Sm., Fl. E. Trop. Afr. Euphorb. 1: 260, fig.
52 (1987), Fl. Zambesiaca 9(4): 176, t. 37 (1996); Sagun &
Dioecious trees; indumentum stellate, sometimes van Welzen, Blumea 47: 149, fig. 1 (2002).
simple as well. Leaves glandular-serrate to entire, Claoxylon sect. Micrococca (Benth.) M€ ull. Arg. (1865).
densely stellate-pubescent abaxially; stipules Monoecious or dioecious shrubs or herbs; buds
deciduous. Inflorescences axillary, racemiform, not perulate; indumentum simple. Leaves
the staminate elongate, pistillate contracted. Sta- minutely puncticulate adaxially, entire or
minate flowers pedicellate; calyx closed in bud, dentate; stipules persistent. Inflorescences axil-
splitting into 3 valvate segments; petals 0; disk lary, sometimes fasciculate, unisexual or some-
segments interstaminal; stamens 15–30, filaments times bisexual, spiciform; bracts eglandular,
distinct; anthers extrorse, basifixed, 2-celled, the- glumaceous. Staminate flowers pedicellate;
cae erect; pollen grains subspheroidal, 3-colpo- calyx closed in bud, splitting into 3 valvate seg-
rate, colpi inoperculate, sexine finely tectate- ments; petals 0; disk of interstaminal segments, or
perforate and microverrucate; pistillode 0. Pistil- 0; stamens 5–50, filaments distinct; anthers
late flowers pedicellate; sepals 3, open in bud, extrorse, basifixed, muticous, pollen grains sphe-
reflexed; petals 0; disk annular or of 3 segments; roidal, 3–4-colporate, colpi inoperculate, sexine
ovary 3–4-locular; stylodia nearly distinct, finely tectate-reticulate, microverrucate; pistil-
unlobed, reflexed, papillate-plumose. Fruit cap- lode 0. Pistillate flowers pedicellate; sepals 3 or
sular, loculicidally dehiscent. Seeds subglobose, 4, imbricate, persistent in fruit, petals 0; disk
extotesta fleshy. segments 3, linear; ovary 3-locular, pubescent;
Eight spp., endemic to Madagascar. ovules anatropous, inner integument thin, outer
integument thinner; stylodia distinct, unlobed,
120. Discoclaoxylon (M€
ull. Arg.) Pax & K. Hoffm. plumose-laciniate. Fruits capsular, 3-lobed, scab-
Discoclaoxylon (M€ ull. Arg.) Pax & K. Hoffm., Wiss. ridulous, sometimes purplish; columella persis-
Ergebn. Deutsche Zentral-Afr. Exped. 2: 452 (1912), Pflan- tent. Seeds subglobose, exotesta thin and fleshy,
zenr. 147, VII: 137, fig. 19 (1914); Radcl.-Sm., Fl. E. Trop. endotesta rugulose.
Afr. Euphorb. 1: 279, fig. 54 (1987). Twelve spp., distributed from South Africa
Claoxylon sect. Discoclaoxylon M€ ull. Arg. (1864).
and tropical Africa to Madagascar, Arabia, south-
Dioecious trees and shrubs; indumentum simple. ern Asia, and Malesia.
Leaves abaxially minutely puncticulate, margins
glandular-dentate, without laminar glands; sti- 122. Erythrococca Benth.
pules deciduous. Inflorescences axillary, staminate Erythrococca Benth., Niger Fl.: 506 (1849); Gilbert, Kew
racemose, flowers several per glomerule, pistillate Bull. 42: 363, fig. 5 (1987); Radcl.-Sm., Fl. E. Trop. Afr.
spiciform, flowers 1 per node; bracts small, persis- Euphorb. 1: 265, fig. 53 (1987), Fl. Zambesiaca 9(4): 165,
figs. 35, 36 (1996).
tent, eglandular or obscurely glandular. Staminate
flowers pedicellate; calyx closed in bud, splitting Dioecious shrubs; buds perulate; indumentum
into 3 or 4 valvate segments; petals 0; disk urceo- simple. Leaves short-petiolate, without laminar
late or dissected; stamens 6–12, filaments distinct, glands, margins glandular-dentate to subtentire;
shorter than anthers; anthers muticous; pollen stipules entire, persistent, sometimes spinose.
grains subprolate, 3-colporate, exine tectate-perfo- Inflorescences axillary, glomerular or racemoid,
rate, microverrucate; pistillode 0. Pistillate flowers glomerules sessile or capitellate-pedunculate;
subsessile; sepals 4, entire, persistent in fruit; disk bracts minute. Staminate flowers pedicellate,
122 G.L. Webster
pedicels basally or mid-articulate; calyx closed in Pojarkova, Fl. URSS 14: 295, t. 17.5 (1949); Vindt, Trav.
bud, splitting into 3–5 valvate segments; petals 0; Inst. Sci. Chérifien 6:13, fig. 7 (1953); Webster, J. Arnold
Arb. 48: 366 (1967); Tutin, Fl. Europaea 2: 212 (1968);
disk segmented, sometimes segments interstam- Radcl.-Sm., Fl. Iraq 4(1): 222, fig. 60 (1980); Correll, Fl.
inal; stamens (2)10–60, filaments distinct; anthers Bahama Arch.: 831, fig. 345 (1982); G€ uemes, Fl. Iberica 8:
extrorse, 2-celled, locules discrete, erect; pollen 201, t. 49, 50 (1997).
grains spheroidal, 3–5-colporate, narrowly oper-
Dioecious (monoecious) perennial or annual
culate, sexine tectate-reticulate, obscurely verru-
herbs; rootstocks with purplish pigment; indu-
cate; pistillode 0. Pistillate flowers articulate-
mentum simple. Leaves opposite, dentate, with
pedicellate; sepals 2(4), valvate, entire, persis-
minute paired basal glands at junction with petiole;
tent in fruit; disk segments 2 or 3, rarely entire;
stipules scarious, sometimes glandular, persistent.
ovary 2(3)-locular, glabrous or pubescent; ovules
Inflorescences axillary, unisexual, the staminate
anatropous; stylodia distinct or basally connate,
spiciform with flowers in glomerules, the pistillate
unlobed, smooth to laciniate. Fruits capsular,
glomerular or on brachyblasts; bracts scarious,
deeply lobed, without a persistent columella.
persistent. Staminate flowers sessile or subsessile;
Seeds subglobose, exotesta fleshy, endotesta
petals and disk 0; calyx closed in bud, splitting into
foveolate to nearly smooth.
3 valvate segments; petals and disk 0; stamens 8–15
Forty spp., from southern Africa to Ethiopia,
(20), filaments distinct; anthers extrorse, locules
1 sp. extending into Arabia.
distinct, diverging from connective; pollen grains
subprolate, 3-colporate; pistillode 0. Pistillate flow-
7c. SUBTRIBE MERCURIALINAE PAX (1890).
ers pedicellate; sepals 3, imbricate, scarious, entire,
Monoecious or dioecious herbs; indumentum sim- persistent, sexine finely tectate-perforate and
ple; leaves alternate or opposite, without embed- densely microverrucate; petals 0; disk segments
ded laminar glands; inflorescences axillary, mostly [staminodia] 2, elongated; ovary 2-locular, gla-
unisexual, spiciform, glomerular, or of solitary brous or pubescent, sometimes appendiculate;
flowers; stamens (2–)4–20, filaments distinct; ovules anatropous, inner integument thin, outer
anthers extrorse, muticous; pollen grains 3-colpor integument thick; stylodia distinct, unlobed, papil-
(oid)ate, sexine tectate-perforate and finely reticu- lose. Fruits capsular; columella membranous-
late; ovary 2-locular, smooth or muricate; stylodia winged, persistent. Seeds ovoid or globose; testa
unlobed; fruit capsular; seeds ecarunculate. smooth or papillate. x ¼ 8.
A problematic alliance of three Old World Eight spp. of temperate Eurasia (Macaronesia
genera: Mercurialis shows a number of characters to eastern Asia). The genus has been intensely
in common with taxa in the Claoxylinae and is studied in Europe since the 17th century, and
resolved in the same molecular clade (A2), but biosystematics studies of the European taxa
Seidelia and Leidesia appear in a different lineage have been made by Durand (1963) and
(A5). The pollen grains of the three genera of Kr€ahenb€ uhl et al. (2002). Mercurialis resembles
Mercurialinae are very similar among themselves genera of Claoxylinae in its leaves and flowers
and to those of Dysopsis (Takahashi et al. 2000). with bluish pigment, but its pollen grains are
different, and more suggestive of the Ricininae.
KEY TO THE GENERA OF SUBTRIBE MERCURIALINAE
1. D ioecious; stamens 8–20; pistillate disk segments 124. Seidelia Baill.
(staminodia) 2; seeds carunculate 123. Mercurialis Seidelia Baill., Étude Gén. Euphorb.: 465 (1858); Prain,
– Monoecious; stamens 2–7; pistillate disk 0 or rudimen- Ann. Bot. 27: 398 (1913), Fl. Capensis 5(2): 464 (1920);
tary; seeds ecarunculate 2 Dyer, Gen. S. Afr. Fl. Pl., ed. 3: 316 (1975); Radcl.-Sm.,
2. Capsule smooth; pistillate sepals 3; leaf blades entire Gen. Euphorb.: 206 (2001).
or denticulate 124. Seidelia
– Capsule setose; pistillate sepals obsolete; leaf blades Monoecious (dioecious) annual herbs; indumen-
crenate-dentate 125. Leidesia tum simple. Leaves alternate or opposite below,
subentire or denticulate, purplish, colliculose,
123. Mercurialis (Tournefort) L. eglandular; stipules minute, deciduous. Inflores-
Mercurialis (Tournefort) L., Sp. Pl. 2: 1035 (1753), cences axillary, glomerular, unisexual or bisexual,
Gen. Pl. ed. 5: 437 (1754); Zimmermann et al. in Hegi, with the pistillate at proximal axils, the staminate
Ill. Fl. Mitteleur. 5(1): 126, figs. 1746–1750 (1925); distal. Staminate flowers pedicellate, pedicels
Euphorbiaceae 123
mid-articulate; calyx splitting into 3 valvate seg- 3–6; pollen grains globose, colpate, finely reticu-
ments; petals and disk 0; stamens (1) 2–5, fila- late; ovary 3-locular, stylodia unlobed, lacerate;
ments nearly distinct; anthers extrorse, muticous, fruit capsular; seeds with obsolete caruncle.
2-locular, with locules distinct, opening 4-valved; Monogeneric, the single genus Dysopsis
pollen grains prolate spheroidal, 3-lobed, sexine restricted to Central and South America. It
finely tectate-perforate, microverrucate, muri shows clear resemblances with the Mercurialinae
crested; pistillode 0. Pistillate flowers pedicellate, in both habit and flowers, but these appear to be
articulate above the middle; sepals 3 (4), imbri- superficial, because in the molecular analysis it is
cate, entire, persistent; petals and disk 0; ovary 2- resolved as sister to Caperonia.
locular, glabrous or pubescent, smooth; ovules
anatropous, inner and outer integuments thin; 126. Dysopsis Baill.
stylodia distinct, unlobed. Fruits capsular, 2- Dysopsis Baill., Étude Gén. Euphorb.: 435 (1858); Pax & K.
lobed; columella slender, persistent. Seeds Hoffm., Pflanzenr. 147, VII (Heft 63): 286, fig. 45 (1914);
ovoid; testa smooth or reticulate. Burger & Huft, Fieldiana Bot. n.s. 36: 113 (1995); Radcl.-
Two spp., South Africa. Sm., Gen. Euphorb.: 207, fig. 26 (2001).
Molina Gay (1851; nom. illeg.).
125. Leidesia M€
ull. Arg. Monoecious herbs; indumentum simple.
Leidesia M€ ull. Arg., DC. in Prodr. 15(2): 792 (1866); Leaves palmately veined, eglandular; stipules
Benth., Hook. Ic. Pl. 13: 66, t. 1284 (1879); Dyer, Gen. S. deciduous. Flowers axillary, mostly solitary;
Afr. Fl. Pl., ed. 3: 316 (1975); Radcl.-Sm., Fl. Zambesiaca bracts minute. Staminate flowers pedicellate;
9(4): 159, t. 33 (1996). sepals 3 (4), connate; petals and disk 0; stamens
Monoecious annual herbs; indumentum simple, 3, or 6 in 2 whorls, the inner filaments connate;
very sparse. Leaves alternate or subopposite, pin- anthers introrse, thecae adnate to slender connec-
nately veined or triplinerved, crenate-dentate, tive; pollen grains subprolate, 3-colpate, colpi
minutely puncticulate, eglandular; stipules min- narrow, sexine tectate-reticulate; pistillode 0. Pis-
ute, subulate. Inflorescences terminal, bisexual, tillate flowers pedicellate; sepals 3, subvalvate,
racemoid-spiciform, pistillate flower solitary and entire, persistent; petals and disk 0; ovary 3-locu-
basal, staminate glomerules distal; occasional lar, pubescent; ovules anatropous, inner and outer
axillary pistillate flowers produced as well; bracts integuments thin; stylodia unlobed, laciniate.
entire, gland-tipped. Staminate flowers subsessile Fruits capsular. Seeds subglobose, minutely car-
or pedicellate; calyx closed in bud, splitting into 3 unculate, testa smooth; embryo linear.
valvate segments; petals and disk 0; stamens 4–7, A single polymorphic sp., D. glechomoides
filaments distinct; anthers introrse, muticous, (A. Rich.) Muell. Arg., Costa Rica to Chile and
subglobose, discrete and pendulous; pollen Juan Fernandez.
grains oblate spheroidal, 3-colporoidate, sexine
tectate-perforate; pistillode 0. Pistillate flower 7e. SUBTRIBE AVELLANITINAE G.L. Webster, subtr.
subsessile; calyx obsolete; disk 0; ovary 2-locular, nov.1
hispid-muricate; ovules anatropous, inner and A monotypic subtribe endemic to Chile.
outer integuments thin; stylodia nearly distinct,
unlobed. Fruits capsular, setose; columella mem-
branous-winged, subpersistent. Seeds subglo- 127. Avellanita Phil.
bose; testa smooth. Avellanita Phil., Linnaea 33: 237 (1864); Benth., Gen. Pl. 3:
289 (1880); Radcl.-Sm., Gen. Euphorb.: 229, fig. 29 (2001);
A single sp., L. procumbens (L.) Prain, in Barrera et al., Bol. Mus. Nac. Hist. Nat. Chile 30: 7 (2001).
southern Africa, in habit somewhat resembling
Dysopsis.
7d. SUBTRIBE DYSOPSIDINAE Hurus. (1954).

1
Subtribe Avellanitinae G.L. Webster, subtr. nov., arbuscu-
Monoecious herbs; indumentum simple; leaves lae monoicae; indumentum simplex; folia alterna, integra,
alternate, crenate; flowers axillary, mostly soli- purpurea; dichasia terminales; flores ♂ subsessiles, stamina
tary; staminate calyx gamophyllous; stamens > 50; flores ♀ sepalis 6, valvatis, stylodiis integris; fructus
capsularis; semina earunculata. Typus: Avellanita Philippi.
124 G.L. Webster
Monoecious shrubs; indumentum simple. Leaves Monoecious (dioecious) trees, shrubs, or herbs;
alternate or distally pseudo-verticillate, entire, indumentum simple or glandular (stellate).
eglandular, purplish-tinged; stipules persistent. Leaves alternate, pinnately or palmately veined,
Inflorescences terminal, pedunculate, dichasial, entire or dentate, sometimes gland-dotted but not
with 1 central pistillate flower and 2 or 3 lateral with embedded laminar glands, rarely stipellate at
staminate flowers. Staminate flowers subsessile; base; stipules usually persistent. Inflorescences
sepals 5, valvate; petals and disk 0; stamens > 50, terminal or axillary, unisexual or bisexual, the
filaments distinct; anthers small, subglobose, staminate spiciform with flowers in glomerules,
dehiscing horizontally, together forming a glo- the pistillate spiciform, sometimes racemose or
bose mass; pollen grains subspheroidal, 3-colpo- paniculate, the bisexual with proximal pistillate
rate, sexine tectate-rugulose, microverrucate; and distal staminate glomerules; staminate bracts
pistillode 0. Pistillate flowers subsessile or pedi- minute, subtending several flowers, pistillate
cellate; sepals 6, valvate, entire, persistent in fruit; bracts foliose, entire or dentate, subtending
ovary 3-locular, hirtellous; stylodia distinct, erect, 1–3 flowers, usually accrescent in fruit. Staminate
elongated, unlobed, papillose. Fruits capsular; flowers subsessile; calyx closed in bud, splitting
columella persistent. Seeds spheroidal, ecaruncu- into 4 valvate segments; petals and disk 0;
late; testa smooth, dry. stamens 4–8(16), filaments distinct or basally
A single sp., A. bustillosii Philippi, endemic to connate; anthers 2-celled, theca discrete,
central Chile. It seems to be correctly placed in elongated, twisted, vermiform; pollen grains
tribe Acalypheae (Radcliffe-Smith 2001), but its about 10–14 x 12–15 mm, oblate-spheroidal,
exact position within this tribe remains to be 3–5-porate, sexine rugulose-tectate, verrucate;
determined. It shares characters such as the pur- pistillode 0. Pistillate flowers sessile or subsessile
plish foliar pigment with subtribe Mercurialinae, (pedicellate); sepals 3–4(5), basally connate,
but differs from that group in its high stamen imbricate, entire, persistent in fruit; petals and
number, 3-locular ovary, and ecarunculate disk 0; ovary (1)2–3-locular, smooth or muricate,
seeds. It also shares purplish foliar pigments often pubescent or papillose; ovules anatropous,
with the Claoxylinae, but differs in stamen mor- inner and outer integuments thin; stylodia nearly
phology and seeds with a non-fleshy testa. Bar- distinct, laciniate (bifid or entire). Fruits capsu-
rera et al. (2001), on the basis of epidermal lar; columella persistent. Seeds ovoid or ellipsoid,
characters, suggest an affinity with Chiropetalum carunculate or not, testa smooth or foveolate.
(Chrozophoreae), but this is contradicted by the n ¼ 7, 10, 14, 20.
palynological evidence. Over 450 spp., pantropical, with a few extra-
tropical spp. in the Americas and E Asia, rela-
7f. SUBTRIBE ACALYPHINAE Griseb. (1859). tively stereotyped morphologically but one
section, Linostachys, differing in its pedicellate
A monogeneric subtribe, distinctive for its pistillate flowers.
unusual anthers and accrescent fruiting calyx. It
appears in some respects most similar to the
Claoxylinae, but differs in its very distinctive pol- 7g. SUBTRIBE BLUMEODENDRINAE G.L. Webster
len grains (Nowicke and Takahashi 2002), inflo- (1975).
rescence structure, remarkably modified Dioecious trees; indumentum simple or minutely
stamens, and flowers lacking a disk. stellate; leaves alternate, opposite, or verticillate,
long-petiolate; stipules minute or 0; inflores-
128. Acalypha L. cences axillary, spiciform to racemose or panicu-
Acalypha L., Sp. Pl.: 1003 (1753); Pax & K. Hoffm., late; staminate flowers pedicellate; calyx splitting
Pflanzenr. 147, XVI: 12, figs. 1–3 (1924); Wilson, Hook. into 3–5 valvate segments; disk of interstaminal
Ic. Pl. 36: t. 3588 (1962); P.I. Forster, Austrobaileya segments; stamens 15–70, anther connectives
4:209–226 (1994); Cardiel, Fl. Colombia, Mon. 15: 23,
figs. 1–25 (1995); Qiu Huaxing & Gilbert, Fl. China 11: enlarged; pollen grains 3-colporate, colpi short,
251–255 (2008); Sagun et al., Blumea 55: 21–60 (2010), rev. sexine tectate-perforate or coarsely reticulate,
Malesian spp. tectum thick, with crotonoid sculpture; pistillode
Euphorbiaceae 125
0; pistillate flowers sessile or pedicellate; sepals interstaminal; stamens 15–50, filaments distinct,
3–5, distinct or basally connate; disk annular or flexed in bud; anthers introrse or latrorse, locules
pulviniform; ovary 2- or 3-locular; ovules with adnate to the enlarged connective, pollen grains
thick integuments, the outer vascularized; stylo- oblate spheroidal, 3-colporate, colpi short and
dia unlobed; fruits large, thick-walled, indehis- narrow, costate, inoperculate; sexine thick, tec-
cent or tardily dehiscent; seeds ecarunculate, tate-perforate, microverrucate; pistillode 0. Pistil-
with fleshy testa. late flowers pedicellate, articulate; sepals 3–5,
This group of four Asiatic genera was earlier imbricate, entire, deciduous in fruit; disk annu-
(Webster 1994) included as a subtribe of the lar; ovary 2- or 3-locular, smooth, glabrous; inner
Pycnocomeae. Radcliffe-Smith (2001) suggested integuments very thick, outer integuments thin-
that they might represent a separate tribe, and ner, vascularized; stylodia unlobed, papillate.
this is supported by the pollen evidence of Now- Fruits capsular, dehiscing tardily; columella not
icke et al. (1999). Here they are shifted from the persistent. Seeds large, compressed, testa fleshy.
Pycnocomeae, from which they differ in the long Five spp., ranging from Burma and the Anda-
petiolate leaves, paniculate inflorescences and mans through Indonesia to the Bismarck Archi-
large indehiscent fruits, to the Acalypheae, pelago, several spp. in Borneo.
which is also supported by the thick vascularized
outer integuments shared by Blumeodendron, 130. Ptychopyxis Miq.
Mallotus and Macaranga and by the molecular
Ptychopyxis Miq., Fl. Ned. Ind., Suppl.: 402 (1861); Hook.
data (Wurdack et al. 2005). f., Hook. Ic. Pl. 18: t. 1703 (1887); Croizat, J. Arnold Arb.
23: 47 (1942); Airy Shaw, Kew Bull. 14: 363 (1960), Kew
KEY TO THE GENERA OF SUBTRIBE BLUMEODEN- Bull. Add. Ser. 4: 188 (1975), Kew Bull. 36: 340 (1981).
DRINAE
Dioecious trees; indumentum simple or 0. Leaves
1. Anther connective moderately enlarged, not umbraculi- sometimes clustered at branch tips, entire;
form; pistillate disk tenuous or 0; stylodia elongated 2
petioles pulvinulate at both ends; stipules minute
– Anther connective greatly enlarged, umbraculiform;
pistillate disk massive; stylodia stigmatiform
or 0. Inflorescences axillary, racemoid or panicu-
132. Botryophora late; bracts persistent, eglandular. Staminate
2. Anther locules adnate to connective; fruits not flowers pedicellate, pedicel articulate above the
glandular 3 base; calyx closed in bud, splitting into 3–5 val-
– Anther locules pendent; fruits glandular 131. Podadenia vate segments; petals 0; disk of many small inter-
3. Anther locules 2; fruits smooth, sometimes carinate, staminal segments; stamens 35–65, filaments
not beaked 129. Blumeodendron
distinct; anthers introrse, connective apiculate;
– Anther locules 4; fruits ribbed or spinose, beaked
130. Ptychopyxis pollen grains oblate spheroidal, 3-colporate,
colpi short, narrow, inoperculate; sexine tectate-
129. Blumeodendron (M€
ull. Arg.) Kurz reticulate; pistillode 0. Pistillate flowers subses-
sile; sepals 4 or 5, basally connate, entire,
Blumeodendron (M€ ull. Arg.) Kurz, J. Asiatic Soc. Bengal,
Nat. Hist. 42: 245 (1873); J.J. Smith, Meded. Dept. Land-
persistent in fruit; disk annular, glabrous or
bouw 10: 458 (1910); Whitmore, Tree Fl. Malaya 2: 68, pubescent; ovary 2–3-locular, pubescent; ovules
figs. 2, 3 (1973); Airy Shaw, Kew Bull. Add. Ser. 4: 57 anatropous, inner integuments very thick, outer
(1975), Kew Bull. 36: 267, fig. 3A (1981). integuments thinner and vascularized; stylodia
Mallotus sect. Blumeodendron M€ ull. Arg. (1866). connate, unlobed. Fruits capsular but tardily
Dioecious; indumentum scanty, simple and stel- dehiscent; beaked, smooth to distinctly ribbed.
late on new growth. Leaves alternate or more Seeds oblong, exotesta fleshy, endotesta bony.
often opposite or verticillate, long-petiolate, pin- Thirteen spp., ranging from Thailand
nately veined or triplinerved, entire, eglandular through Malesia to New Guinea.
(obscurely punctate); stipules minute or 0. Inflor-
escences axillary, the staminate glomerular or 131. Podadenia Thwaites
racemoid, pistillate spiciform; staminate flowers Podadenia Thwaites, Enum. Pl. Zeyl. 4: 273 (1861);
pedicellate, calyx closed in bud, splitting into Trimen, Handb. Fl. Ceylon 4: 62 (1898); Philcox, Fl.
3 or 4 valvate segments, microlepidote abaxially, Ceylon 11: 144 (1997); Radcl.-Sm., Gen. Euphorb.: 170,
petals 0; disk receptacular, convex, segments fig. 19 (2001).
126 G.L. Webster
Dioecious trees; indumentum simple, glandular tardily dehiscent. Seeds somewhat angular,
on inflorescences. Stipules 0. Inflorescences axil- plano-convex, testa smooth and dry.
lary, paniculate (compound thyrsoid), glandular- A single sp., B. kingii Hook. f., ranging from
pubescent; bracts persistent. Staminate flower Burma to Sumatra, Java and Borneo. Airy Shaw
pedicellate; calyx closed in bud, splitting into 3 (1960) provided a taxonomic revision of this
or 4 glandular-pubescent reflexed segments; genus, and was the first investigator to clearly
petals 0; disk of numerous interstaminal seg- relate it to Blumeodendron.
ments; stamens 20–30, filaments distinct; anthers
latrorse, with enlarged apiculate connective; 7h. SUBTRIBE ROTTLERINAE Meisner (1841).
locules pendent; pollen grains 3-colporate, colpi
short and narrow; sexine tectate-punctate, micro- Dioecious (monoecious) trees or shrubs; indumen-
verrucate; pistillode 0. Pistillate flowers pedicel- tum stellate (simple); leaves alternate or opposite;
late; sepals 4–7, imbricate, entire, glandular- blade unlobed or lobed, pinnately or palmately
pubescent, persistent or deciduous in fruit; disk veined; sometimes with embedded laminar glands;
0; ovary 3-locular, glandular-pubescent; ovules stipules deciduous or obsolete; inflorescences ter-
anatropous, inner integuments moderately minal or axillary, racemose or paniculate; stami-
thick, outer integuments thinner, vascularized; nate calyx splitting into valvate segments; petals 0;
stylodia nearly distinct, unlobed, papillate. Fruit disk of interstaminal segments or 0; stamens
indehiscent, beaked, covered with gland-tipped 15–300, filaments distinct; anthers muticous,
setae. Seeds 1 or 2 per fruit, compressed; exo- anther sacs not pendulous; pollen grains 3(4)-col-
testa fleshy, endotesta bony. porate, finely to coarsely tectate-perforate and
A single sp., P. thwaitesii (Baill.) M€
ull. Arg., microverrucate; pistillate sepals 3–6(10), distinct
from Ceylon. Although Philcox (1997) treated it or connate; disk 0; ovary 2–4(8)-locular, some-
as a species of Ptychopyxis, the Ceylonese species times echinate; stylodia unlobed, plumose or
appears sufficiently different from the other lacerate; fruit capsular, baccate, or drupaceous;
known taxa of Ptychopyxis that its placement in seeds ecarunculate (carunculate), testa often fleshy.
a separate genus is justifiable. After the drastic reductions of the satellite
genera around Mallotus, which were found to be
132. Botryophora Hook. f. embedded in this genus, subtribe Rottlerinae
comprises only three genera, although two very
Botryophora Hook. f., Fl. Brit. Ind. 5: 476 (1888; nom. large ones.
cons.); Airy Shaw, Kew Bull. 14: 374 (1960), Hook. Ic. Pl.
36: t. 3576 (1962), Kew Bull. 36: 269, fig. 4 (1981).
KEY TO THE GENERA OF SUBTRIBE ROTTLERINAE
Dioecious trees; indumentum simple. Leaves
1. Anthers 3- or 4-celled; indumentum simple; leaves
sometimes clustered, entire, eglandular; petioles alternate 134. Macaranga
long, pulvinate at both ends; stipules obsolete. – Anthers 2-celled; indumentum simple or stellate; leaves
Inflorescences axillary, the staminate paniculate alternate or opposite with one member of the pair
(compound spiciform or contracted), the pistil- smaller 2
late spiciform; bracts minute. Staminate flowers 2. Indumentum of simple and stellate hairs; leaf blades
subsessile; calyx closed in bud, glabrous, splitting usually granulose-glandular beneath; pollen grains
with tectae-microperforate exine; stylodia not particu-
into 2 spathaceous segments; petals 0; disk glands larly elongate 133. Mallotus
receptacular, numerous, several surrounding – Indumentum not stellate but of simple hairs or gland-
each stamen; stamens c. 30–60, filaments distinct; tipped hairs or sessile peltate-stellate hairs with central
anther connective pileiform-peltate, anthers cells; leaf blades eglandular; pollen grains with areolate
4-locellate; pollen grains spheroidal, 3-colporate, exine sculpture; stylodia often very long [> 20 mm]
135. Hancea
inoperculate and emarginate, sexine tectate-
punctae, microverrucate; pistillode 0. Pistillate
133. Mallotus Lour. Fig. 28
flowers sessile or subsessile; sepals 3 or 4,
minute, hidden under the massive pulviniform Mallotus Lour., Fl. Cochinch.: 635 (1790), nom. cons.; Pax &
segmented disk; petals 0; ovary 3- or 4-locular, K. Hoffm., Pflanzenr. 147, VII: 145, figs. 23–29 (1914); Airy
smooth and glabrous; stylodia unlobed, thick, Shaw, Kew Bull. 21: 379 (1968), Hook. Ic. Pl 38: t. 3715
shorter than ovary, papillate. Fruit capsular but (1974), Kew Bull. Add. Ser. 4: 160 (1975), 8: 162 (1980);
Euphorbiaceae 127
Dioecious (monoecious) trees, shrubs or clim-

bers; indumentum simple and stellate or tufted
and glandular. Leaves alternate or opposite,
unlobed or palmately lobed, entire to dentate,
sometimes peltate, usually with extrafloral nec-
taries adaxially; stipules persistent, deciduous, or
obsolete. Inflorescences terminal or axillary, spi-
ciform to racemoid or paniculate, glomerulate or
umbel-like, usually unisexual; bracts eglandular,
persistent or deciduous; staminate flowers 1–15
per bract; pistillate flowers 1 per bract, several
when umbel-like. Staminate flowers pedicellate;
calyx closed in bud, sepals (2)3–5, valvate; petals
0; interstaminal disk glands sometimes +;
stamens 15–130, filaments distinct or connate;
anthers basifixed, 2-locellate; connective some-
times broadened or apiculate; pollen grains sphe-
roidal, 3(4)-colporate, colpi vestigially operculate,
sexine tectate/microperforate; pistillode 0 (+).
Pistillate flowers pedicellate; sepals (2)3–6, val-
vate, connate at least at base, sometimes calyx
cupular or spathaceous; petals and disk 0; ovary
(1)2–3(9)-locular; ovules anatropous, inner
integuments thick, outer integuments thinner,
sometimes vascularized; style + or 0; stylodia dis-
tinct or basally connate into a common style,
stylodia or their distal tips unlobed, papillose to
plumose. Fruits capsular (drupaceous), 3(2–5)-
locular, very rarely 1-locular [Neotrewia] or 7–9-
locular [Octospermum], smooth or softly spiny;
columella persistent. Seeds globose to ovoid, ecar-
unculate, exotesta sometimes fleshy, endotesta
Fig. 28. Euphorbiaceae-Acalyphoideae. Mallotus panicu-
latus. A Flowering branch. B Base of upper leaf surface hard, smooth to rugose. n ¼ 11, 12, 18.
with extrafloral nectaries. C Staminate flowers and buds. According to Sierra et al. (2010), Mallotus
D Pistillate flower. E Fruit. F Dehisced fruit with seeds and includes about 110 spp. distributed mainly in
apex of column. (Sierra & van Welzen 2005) (sub)tropical Asia, Malesia, Australia, and the
southwestern Pacific islands, and only two spp.
McPherson & Tirel, Fl. Nouv.-Caléd. 14(1): 104, t. 35, 6–9
(1987); Radcl.-Sm., Fl. E. Trop. Afr. Euph. 1: 235 (1987); in Africa/Madagascar, growing in various habi-
McPherson, Adansonia III, 17: 169 (1995); Bollendorf et al., tats ranging from the understorey to swamp for-
Blumea 45: 319, figs. 1–10 (2000); Slik & van Welzen, Blu- est and montane forest, and often in disturbed
mea 46: 3, figs. 1–22 (2001); Sierra & van Welzen, Blumea and secondary vegetation.
50: 249–274 (2005), tax. sect. Mallotus; Kulju et al., Amer. J.
Bot. 94: 1726–1743 (2007), mol. syst.; Kulju et al., Blumea 52:
115–136 (2007), Neotrewia, Octospermum and Trevia 134. Macaranga Thouars
reduced; Qiu Huaxing & Gilbert, Fl. China 11: 225–237 Macaranga Thouars, Gen. Nov. Madag.: 26 (1806);
(2008); Sierra, Kulju, Fiser, Aparicio & van Welzen, Taxon Gagnep., Fl. Indochine 5: 434, figs. 52–54 (1926); Perrry,
59: 101–116 (2010), mol. syst.; van Welzen et al., Blumea 55: J. Arnold Arb. 34: 191 (1953); Whitmore, Tree Fl. Malaya
285–290 (2011), key to Males. spp. 2: 105, figs. 8, 9 (1973); A.C. Smith, Fl. Vitiensis Nova 2:
Trevia L. (1753) (¼ Trewia). 500, figs.131–135 (1981); Coode, Taxon 25: 184 (1976),
Echinus Lour. (1790). Fl. Mascar. 160: 53, t. 10 (1982); McPherson & Tirel,
Rottlera Roxb. (1802). Fl. Nouv.-Caléd. 14: 172–185, t. 37 (1987); Radcl.-Sm.,
Fl. E. Trop. Afr. Euphorb. 1: 239, fig. 49 (1987), Fl.
Coelodiscus Baill. (1858).
Zambesiaca 9(4): 161, t. 34 (1996); McPherson, Adansonia
Neotrewia Pax & K. Hoffm. (1914). III, 18: 275 (1996); Florence, Fl. Polynésie Française 1: 104,
Octospermum Airy Shaw (1965). figs. 17–19 (1997); Slik et al., Gard. Bull. Singapore 52: 12,
128 G.L. Webster
figs. 1–24 (2000); Qiu Huaxing & Gilbert, Fl. China 11:
237–240 (2008).
Mappa Juss. (1824).
Pachystemon Blume (1826).
Dioecious trees and shrubs; indumentum of sim-
ple (fasciculate) and conspicuous, usually colour-
ful glandular hairs; twigs sometimes hollow, often
with reddish exudate. Leaves long-petiolate,
unlobed to palmately lobed, pinnately to pal-
mately veined, often peltate, abaxially granulose-
glandular; stipules minute to large, persistent or
deciduous. Inflorescences axillary, the staminate
spiciform or capitulate, the pistillate racemoid,
sometimes compound; bracts often conspicuous
and glandular. Staminate flowers subsessile or
pedicellate; calyx closed in bud, splitting into
2–4 valvate segments; petals and disk 0; stamens
(1–)3–20(30), filaments distinct or basally con-
nate; anthers muticous, 3- or 4-locellate; pollen
grains spheroidal, 3-colporate, colpi granulate,
sexine tectate-microperforate; pistillode 0. Pistil-
late flower pedicellate; calyx cupular, subentire to
4–6-lobed or -partite, persistent in fruit; petals
and disk 0; ovary 1–6-locular, glandular-granu-
lose and sometimes echinate; ovules anatropous,
inner integument thick, outer integument thicker,
vascularized; stylodia distinct (shortly connate),
unlobed. Fruits capsular, (1)2–3(6)-locular;
smooth or spiny, loculicidal; columella not persis-
tent. Seeds globose, ecarunculate; exotesta fleshy,
endotesta indurate, often rugose. n ¼ 11.
About 260 spp., 26 in Africa, 10 in Madagas-
car, the rest distributed from India and Ceylon
east to Malesia and New Caledonia and the Pacific Fig. 29. Euphorbiaceae-Acalyphoideae. Hancea integrifo-
islands as far east as Polynesia but not Hawaii, lia . A Stipulate leaf. B Part of inflorescence with pistillate
usually as forest plants and often as members of flowers. (Sierra et al. 2006; drawn by A. Walsmit Sachs)
secondary vegetation.
Monoecious or dioecious shrubs or trees; indu-
135. Hancea Seem. Fig. 29 mentum of simple hairs and gland-tipped hairs or
Hancea Seem., Bot. Voy. Herald : 409 (1857); Slik & van sessile peltate-stellate hairs with central cells;
Welzen, Blumea 46: 3–66 (2001), as Mallotus sect. Hancea; resin sometimes +. Leaves opposite [alternate
Sierra et al., Blumea 51: 524–537 (2006), sub Cordemoya; in H. subpeltata], entire, pinnately or palmately
Sierra et al., Blumea 52: 361–366 (2007), new comb. under 3-veined, with one of each leaf of a pair either
Hancea.
Mallotus Lour. sect. Hancea (Seem.) Pax & K. Hoffm.
slightly smaller than the other [subg. Cordemoya]
(1914). or strongly reduced to appear like a stipule
Boutonia Bojer ex Bouton (1846). [sect. Diplochlamys)]; stipules axillary or intrape-
Cordemoya Baill. (1861), Radcl.-Sm., Gen. Euph.: 228 tiolar, persistent. Inflorescences axillary, termi-
(2001). nal, or ramiflorous, uni- or bisexual, racemes or
Mallotus sect. Cordemoya M€ ull. Arg. (1865).
Deuteromallotus Pax & K. Hoffm. (1914).
less often panicles; staminate flowers 1–3 per
Euphorbiaceae 129
bract, pistillate flowers 1 per bract. Staminate KEY TO THE GENERA OF SUBTRIBE LASIOCCINAE
flowers pedicellate; sepals 2–4, valvate, distinct 1. Monoecious; pistillate sepals persistent; indumentum
to basally connate; petals and disk 0; stamens simple; ovary muricate 136. Lasiococca
40–250; filaments distinct; anthers extrorse; pol- – Dioecious; pistillate sepals deciduous; indumentum
len grains oblate spheroidal, 3-colporate, sexine simple or lepidote 2
areolate, scabrate; pistillode 0. Pistillate flowers 2. Lepidote scales 0; ovary tuberculate; pollen sexine not
pedicellate; sepals (3)4–6(7), valvate or imbricate, striate 137. Spathiostemon
distinct; petals and disk 0; ovary 2–3-locular; – Lepidote scales present; ovary smooth; pollen sexine
striate 138. Homonoia
ovules anatropous, inner integuments very
thick, outer integuments thinner, vascularized;
stylodia 2–3, unbranched, erect, 5–25 mm long,
only basally shortly connate. Fruits capsular, 136. Lasiococca Hook. f.
spiny, spines 4–6 or up to 170, sometimes Lasiococca Hook. f., Hook. Ic. Pl. 16: t. 1587 (1887), Fl.
gland-tipped. Seeds subglobose, ecarunculate, Brit. Ind. 5: 456 (1887); Haines, Kew Bull. Misc. Inf. 1920:
testa not fleshy. 70 (1920); Airy Shaw, Kew Bull. 16: 358 (1963), 21: 406
(1968); Whitmore, Tree Fl. Malaya 2: 104 (1973); Thin, J.,
A genus of 17 spp., as enumerated by Sierra Biol. Sinh Hoc 8(3): 36 (1986); van Welzen, Thin & Hoai
et al. (2007), four of them in Madagascar and the Duc, Blumea 43: 141–144, fig. 3 (1998).
Mascarenes (subg. Cordemoya), the rest (subg.
Hancea, with 2 sections) distributed from Hong Monoecious trees or shrubs; indumentum simple
Kong throughout SE Asia to New Guinea. The or glandular. Leaves alternate or subopposite,
whole floral structure and particularly the numer- sometimes pseudo-verticillate, entire, glandular
ous stamens and extremely elongate stylodia are on margins; stipules deciduous. Inflorescences
strongly indicative of wind pollination. axillary, unisexual; staminate flowers in racemes,
bracts uniflorous; pistillate flowers solitary, axil-
7i. SUBTRIBE LASIOCOCCINAE G.L. Webster (1975). lary, bracts deciduous. Staminate flowers pedicel-
late; calyx closed in bud, splitting into 3 valvate
Monoecious or dioecious trees or shrubs; indu- segments; petals and disk 0; stamens > 100, fila-
mentum simple or lepidote; leaves alternate, stip- ments connate and paniculately branching;
ulate; inflorescences axillary, unisexual, racemose anthers introrse, muticous, thecae globose; pollen
or solitary, bracts eglandular; staminate sepals grains spheroidal, 3-colporate, sexine tectate-per-
usually 3; disk 0; stamens many, filaments con- forate, slightly rugulose, microverrucate; pistil-
nate and ramified; pollen grains 3-colporate, not lode 0. Pistillate flowers pedicellate; sepals 5
operculate, sexine tectate-microperforate or stri- (7), foliose, imbricate, entire, persistent in
ate-verruculose; pistillode 0; pistillate sepals 5–8, fruit; petals and disk 0; ovary 3-locular, tubercu-
imbricate, persistent or deciduous; disk 0; ovary late; stylodia basally connate, unlobed. Fruits
3-locular; stylodia unlobed, smooth to papillose capsular, echinate; columella persitent. Seeds
or plumose; fruit capsular; seeds ecarunculate, ovoid, ecarunculate, testa smooth.
testa usually fleshy. Three spp., distributed from India to SE Asia
Three Asiatic genera, originally placed in the and Malesia.
subtribe Ricininae by Pax and Hoffmann (1919)
because of the ramified filaments, but the totality
of characters suggests that the androecial resem- 137. Spathiostemon Blume
blance to Ricinus may be due to convergence. Spathiostemon Blume, Bijdr.: 621 (1816); Airy Shaw, Kew
Airy Shaw (1974) and van Welzen et al. (1998) Bull. 16: 357 (1963), Hook. Ic. Pl. 38: t. 3720 (1974), Kew
thought that the Lasiococcinae may be related to Bull. 36: 345, fig. 9 (1981); van Welzen et al., Blumea 43:
the Malloteae, whereas the molecular signals 145–150, fig. 4 (1998).
Polydragma Hook. f. (1887).
(Wurdack et al. 2005) place Homonoia and Clonostylis S. Moore (1925); still incompletely known;
Spathiostemon with low support close to Acaly- van Welzen (Blumea 43: 150. 1988) argues for its indepen-
pheae and Pycnocomeae. dent generic status.
130 G.L. Webster
Monoecious trees or shrubs; indumentum sim-

ple. Leaves glandular at base; petioles pulvinate;
stipules deciduous. Inflorescences axillary
(pseudo-terminal), racemose, not fasciculate,
unisexual; bracts entire, eglandular, persistent.
Staminate flowers sessile to shortly pedicellate,
articulate above the middle; calyx closed in bud,
splitting into 3 valvate segments; petals and disk
0; stamens > 100, filaments connate into 4–7
ramified phalanges; anthers introrse or latrorse,
muticous, 2-celled; pollen grains spheroidal,
3-colporate, sexine coarsely rugulose-tectate and
verrucate; pistillode 0. Pistillate flowers pedicel-
late; sepals 5 or 6, entire, imbricate, persistent in
fruit; petals and disk 0; ovary 3-locular, smooth
or papillate; ovules anatropous, inner integument
thick, outer integument thin; stylodia distinct,
unlobed, acuminate, adaxially papillate. Fruits
capsular, smooth or echinate, septicidal and loc-
ulicidal; columella sometimes persistent, apically
dilated. Seeds obovoid, ecarunculate, hilum trian-
gular, testa smooth.
Two spp., distributed from peninsular Thai-
land and Indonesia to New Guinea. Pax and Hoff-
mann (1919) combined Spathiostemon with
Homonoia, and both genera are strongly sup-
ported as sisters (rbcL only) in the analysis of
Wurdack et al. (2005), but their pollen exine
ornamentation is very different (Nowicke and
Takahashi 2002).
Fig. 30. Euphorbiaceae-Acalyphoideae. Homonoia riparia.

138. Homonoia Lour. Fig. 30 A Flowering branch. B Lower leaf surface showing marginal
Homonoia Lour., Fl. Cochinch.: 636 (1790); Airy Shaw, gland and scale hairs. C Staminate flower. D Pistillate flower.
Kew Bull. 36: 310 (1981); Philcox, Fl. Ceylon 11: 173 E Fruit. F Fruit column after dehiscence. (van Welzen et al.
(1997); van Welzen, Blumea 43: 136–141, fig. 2 (1998). 1998; drawn by J. van Os)
Dioecious (monoecious) arborescent shrubs; persistent, 3-angled. Seeds ovoid, carinate, ecarun-
indumentum simple and lepidote. Leaves egland- culate, exotesta fleshy, endotesta smooth.
ular; stipules deciduous. Inflorescences axillary, Two spp., distributed from India to China and
solitary, spiciform, usually unisexual; glomerules throughout Malesia to New Guinea, growing as rheo-
uniflorous. Staminate flowers sessile; petals and phytes mainly in and along rivers at low altitudes.
disk absent; calyx splitting into 3 valvate segments; The absence of petals and a disk, the numerous
stamens > 100, filaments connate into a paniculate stamens united into a "staminate tree", and the plu-
androecium; anthers introrse, muticous; pollen mose, stigmatic stylodia suggest wind pollination.
grains spheroidal, 3-colporate, colpi narrow and
operculate, sexine tectate, striate-beaded; pistil- 8. TRIBE PYCNOCOMEAE Hutch. (1969).
lode 0. Pistillate flowers sessile; sepals 5, basally
connate, imbricate, entire, persistent in fruit; Monoecious or dioecious trees or shrubs; indu-
petals and disk 0; ovary 3-locular, pubescent; mentum simple or 0; leaves alternate, sometimes
ovules anatropous, inner and outer integuments stipellate or with laminar glands; stipules decidu-
thin; stylodia unlobed, linear, plumose-laciniate. ous or rudimentary; inflorescences axillary, race-
Fruits capsular, loculicidal; columella sometimes moid or spiciform (compounded); staminate
Euphorbiaceae 131
calyx splitting into 3–5 valvate segments; petals 0; 139. Necepsia Prain
disk of intrastaminal segments or 0; stamens Necepsia Prain, Kew Bull. 1910: 343 (1910); Léonard, Fl.
8–120, filaments distinct; anthers mostly introrse, Afr. Centr. Euphorb. 3: 32 (1996); Radcl.-Sm., Fl. Zambe-
sometimes with enlarged connective; pollen siaca 9(4): 23, t. 27 (1996).
grains subspheroidal, 3-colporate, colpi inoper- Palissya Baill. (1858; nom. illeg.).
culate and lacking costae; sexine tectate-perfo- Neopalissya Pax (1914).
rate, or with granules arranged in areoles; Monoecious (dioecious) trees or shrubs; indu-
pistillode 0; pistillate sepals 3–7, imbricate, per- mentum simple. Leaves denticulate, abaxially
sistent in fruit; disk annular, glabrous or 0; ovary with embedded scattered glands; stipules persis-
3-locular; ovules usually with inner integuments tent or deciduous. Inflorescences axillary, unisex-
thicker than the outer; stylodia bifid to unlobed, ual or bisexual, spiciform, the staminate
sometimes laciniate; fruits dehiscent or indehis- elongated, the pistillate abbreviated; bracts scari-
cent; columella usually persistent; seeds round- ous, persistent. Staminate flowers pedicellate,
ish, ecarunculate, testa smooth and dry. pedicel articulate at base; calyx closed in bud,
The tribe Pycnocomeae is here redefined splitting into (3)4(5) valvate segments; petals 0;
from the treatments of Webster (1994) and Rad- receptacle convex; disk of numerous pubescent
cliffe-Smith (2001) by the expulsion of the Blu- interstaminal segments; stamens numerous,
meodendrinae and the addition of the mostly > 50, filaments distinct; anthers with
Necepsinae, but remains heterogeneous: both enlarged glandular apiculate connective,
Paranecepsia and Necepsia are resolved as sister locules pendent; pollen grains subprolate, 3-col-
taxa to Pseudagrostis (Wurdack et al. 2005; Kulju porate, scarcely marginate, angulaperturate, col-
et al. 2008), whereas Amyrea is resolved in a clade par membrane granulose; sexine tectate-
with Cyttaranthus and Discoglypremna. perforate; pistillode 0. Pistillate flowers subses-
sile; sepals (4)5(6), imbricate, persistent in fruit;
KEY TO THE SUBTRIBES OF PYCNOCOMEAE petals 0; disk annular; ovary 3-locular, verrucu-
1. Stylodia bifid; inflorescences mostly unisexual; leaf lose and pubescent; ovules anatropous, inner
blades with or without discoid glands 8a. Necepsinae integument thick, outer integument thin; stylodia
– Stylodia unlobed; inflorescences usually bisexual; leaf bifid to twice bifid. Fruits capsular, 3-lobed, ver-
blades eglandular or minutely punctuate ruculose; columella winged, persistent. Seeds
8b. Pycnocominae subglobose, ecarunculate, testa smooth, hilum
lunate, conspicuous.
8a. SUBTRIBE NECEPSINAE G.L. Webster (2004). Three spp., tropical Africa. In Webster (1994)
Monoecious or dioecious; leaves alternate, vena- and Radcliffe-Smith (2001), Necepsia was
tion mostly eucamptodromous, with scattered included in Bernardieae but here has been trans-
laminar glands [except in Paranecepsia]; bisexual ferred to Pycnocomeae because of the palynolog-
inflorescences lacking terminal pistillate flower; ical similarities with that tribe. See also the
stamens 10–120; connective sometimes enlarged comments under the tribe.
or apiculate; pistillate disk annular or lobed; sty-
lodia bifid, sometimes lacerate. 140. Paranecepsia Radcl.-Sm.
Five genera, all African, one extending to Paranecepsia Radcl.-Sm., Kew Bull. 30: 684 (1976), Fl. E.
Madagascar. Trop. Afr. Euphorb. 1: 220, fig. 43 (1987); Fl. Zambesiaca 9
(4): 145, t. 28 (1996).
KEY TO THE GENERA OF NECEPSINAE Dioecious trees; indumentum simple. Leaves
1. Pistillate disk 8–10-lobed; anthers muticous; pollen clustered at branch tips, serrate, eglandular, sti-
sexine tectate-striate 141. Amyrea pellate at base; stipules entire, pubescent. Inflores-
– Pistillate disk annular to crenulate; anthers apiculate; cences axillary, solitary, lax, racemoid, often
pollen sexine tectate-perforate 2
2. Ovary verruculose; leaf lamina not stipellate at base
aggregated paniculately; staminate glomerules
139. Necepsia 1–5-flowered, pistillate flowers solitary at each
– Ovary smooth; leaf lamina stipellate at base bract. Staminate flowers pedicellate, pedicels artic-
140. Paranecepsia ulate; calyx closed in bud, splitting into 3–5 valvate
132 G.L. Webster
segments; petals 0; disk segments numerous, intras- rarely so in Argomuellera, the staminate glomer-
taminal; stamens 25–40, filaments distinct; anthers ules proximal; stamens 15–80, anther connective
dorsifixed, introrse, connective apiculate but not not enlarged; pollen sexine tectate-perforate or
enlarged, thecae pendulous; pollen grains spheroi- gemmate; pistillate disk + or 0; stylodia distinct
dal, 3-colporate, scarcely marginate, sexine tectate- or connate, unlobed, apically dilated.
perforate; pistillode 0. Pistillate flowers pedicellate; Three genera of Africa and Madagascar.
sepals 5–7, distinct, unequal, imbricate, accrescent
and persistent in fruit; petals 0; disk annular; ovary KEY TO THE GENERA OF PYCNOCOMINAE
3-locular, pubescent; stylodia basally connate, bifid, 1. Pistillate disk +; pistillate flowers usually not terminal
papillose. Fruits 3-lobed, septicidally and loculicid- in inflorescence 142. Argomuellera
ally dehiscent; columella persistent. Seeds globose, – Pistillate disk 0; pistillate flowers terminating inflores-
ecarunculate, testa smooth. cences 2
A single sp., P. alcheornifolia Radcl.-Sm., East 2. Terminal bud not perulate; pistillate flower solitary and
Africa: Tanzania and Mozambique. The stipellate terminal at apex of inflorescence; filaments arcuate in
bud; ovary 6-horned, pubescent 143. Pycnocoma
leaf blades suggest a relationship with the Alchor-
– Terminal bud perulate; pistillate flowers not solitary
neae, but the inoperculate pollen grains rule out and terminal at apex of inflorescence; filaments straight
assignment to that tribe. For its possible relation- in bud; ovary unappendaged, glabrous
ship with Necepsia, see above under the tribal 144. Droceloncia
description.
142. Argomuellera Pax
141. Amyrea Leandri Argomuellera Pax, Bot. Jahrb. 19: 90 (1894); Prain, Fl.
Amyrea Leandri, Notul. Syst. 9: 168, t. 1, figs. 16–19 Trop. Afr. 6(1): 925 (1912); Léonard, Bull. Soc. Roy.
(1941); Radcl.-Sm., Kew Bull. 53: 438, t. 1–6 (1998), Gen. Belg. 91: 274 (1959), Fl. Afr. Centr. Euphorb. 3: 56, t. 10,
Euphorb.: 161 (2001); Schatz, Generic Tree Flora Madag.: 11 (1996); Radcl.-Sm., Fl. Zambes. 9(4): 147, t. 29 (1996).
143, fig. 131 (2001). Pycnocoma sect. Wetriaria M€ ull. Arg. (1866).
Neopycnocoma Pax (1909).
Dioecious shrubs or trees; indumentum simple or
Monoecious trees or shrubs; indumentum sim-
0. Leaves entire or dentate, eglandular; stipules
ple. Leaves sometimes apically clustered, simple,
deciduous. Inflorescences axillary, subterminal,
subsessile or short-petiolate, entire or denticu-
or terminal, spiciform or racemoid, bracts con-
late, sometimes with abaxial glands; stipules
vex, glumiform, subtending solitary flowers. Sta-
deciduous. Inflorescences axillary, spiciform
minate flowers subsessile to pedicellate; calyx
(compound); cymules usually bisexual; bracts
closed in bud, splitting into 3–5 valvate segments;
persistent. Staminate flowers pedicellate, pedicel
petals 0; disk segments interstaminal, pubescent;
articulated near base; calyx closed in bud,
stamens 20–30, filaments distinct; anthers muti-
splitting into 2–5 valvate segments; petals 0; disk
cous, latrorse, subpendulous; pollen grains 3-
segments interstaminal, pilose; stamens (15–)
angled, 3-colporate, colpi narrow and not bor-
25–120, filaments distinct, erect in bud; anthers
dered; sexine tectate-striate; pistillode 0. Pistillate
with enlarged connective, muticous, locules pen-
flowers pedicellate; sepals 5, imbricate, entire,
dent; pollen grains spheroidal, angulaperturate,
usually persistent; disk fleshy, 8–10-lobed; ovary
3-colporate, emarginate, colpar membrane gran-
3-locular, smooth and glabrous; stylodia bifid,
ulose, sexine tectate-reticulate; pistillode usually
papillose. Fruit capsular; columella persistent.
0. Pistillate flowers pedicellate; sepals usually (3–)
Seeds ovoid, ecarunculate; testa smooth, blackish.
5–6(9), imbricate, entire, persistent in fruit;
Eleven spp. endemic to Madagascar. In
disk annular; ovary 3(4)-locular, pubescent;
describing the genus, Leandri noted resemblance
ovules pachychazal, inner integument thick,
to Neopalissya (Necepsia) and to Mareya, which
outer thinner; stylodia basally connate, unlobed,
furnished the basis of the anagrammatic generic
tips recurved. Fruits capsular, 3-lobed; columella
name. See also above under the tribe.
persistent, apically dilated. Seeds globose, ecar-
8b. SUBTRIBE PYCNOCOMINAE G.L. Webster (1994). unculate, smooth.
Eleven spp., Africa and Madagascar. The
Monoecious; leaves alternate, lacking dispersed group was first treated by M€ uller (1866) as a
laminar glands, venation brochidodromous; pistil- section of Pycnocoma. Pax originally regarded
late flowers solitary, terminating inflorescences, Argomuellera as related to Mallotus, but noted a
Euphorbiaceae 133
resemblance in habit to Pycnocoma, and Rad- segments; petals 0; disk receptacular, convex, gla-
cliffe-Smith (2001) has included Argomuellera in brous; stamens c. 40, filaments distinct and
the subtribe Pycnocominae. straight in bud; anther connective not enlarged;
pollen grains subprolate, 3-colporate, inopercu-
143. Pycnocoma Benth. late and emarginate, sexine tectate-perforate; pis-
Pycnocoma Benth., Niger Fl.: 508 (1849); Léonard, Bull.
tillode 0. Pistillate flowers pedicellate; sepals 5 or
Jard. Bot. Nat. Belg. 65: 38 (1996), Fl. Afr. Centr. Euphorb. 6 (?), imbricate; petals and disk 0; ovary 3-locular,
3: 36, fig. 2, t. 6–9 (1996); Radcl.-Sm., Fl. E. Trop. Afr. smooth and glabrous; stylodia distinct, unlobed,
Euphorb. 1: 228; fig. 46 (1987). apically dilated. Fruit capsular.
Wetriaria (M€ ull. Arg.) Kuntze (1903). A single sp., D. rigidifolia (Baill.) Léonard,
Monoecious trees or shrubs; indumentum sim- Madagascar and Comoro I. Although originally
ple. Leaves sometimes pseudo-verticillate, sub- included in Pycnocoma, its pollen is very different
sessile, entire or dentate, rarely lobed, from that genus (Nowicke et al. 1999).
eglandular but minutely punctate abaxially; sti-
pules deciduous or 0. Inflorescences axillary, 9. TRIBE AMPEREEAE M€
ull. Arg. (1864).
bisexual, racemiform or spiciform, staminate
Monoecious (dioecious) herbs or subshrubs;
flowers in glomerules at proximal nodes, pistillate
indumentum simple; leaves alternate, narrow, eri-
solitary and terminal; staminate bracts concave.
coid, stipulate. Inflorescences glomerular, axillary
Staminate flowers pedicellate, pedicel articulate at
or terminal; staminate flowers pedicellate; sepals
base; calyx closed in bud, splitting into 2–5 val-
3–5, basally connate, valvate; petals + or 0; disk
vate segments; petals 0; disk receptacular, usually
dissected or obsolete; stamens 3–10, filaments
glabrous; stamens > 50, filaments distinct, flexu-
distinct; anthers biseriate, locules pendulous; pol-
ous in bud, long-exserted; anthers introrse, muti-
len grains subprolate to suboblate, 3-colporate,
cous, connective not enlarged; pollen grains 3-
colpi inoperculate; exine reticulate, tectum psi-
angled, 3-colporate, emarginate, inoperculate
late; pistillode present or 0; pistillate flowers sub-
but colpar membrane granulose; sexine minutely
sessile on a 1(3)-flowered peduncle; sepals (4) 5,
tectate-punctate, slightly rugulose, with coarse
imbricate, entire to fimbriate; disk cupular, cren-
reticulum of granular projections; pistillode 0.
ulate; ovary 3-locular, integuments thin; stylodia
Pistillate flowers pedicellate; sepals (4)5–8, imbri-
distinct or nearly so, spreading, bifid; fruits cap-
cate, entire, persistent in fruit; petals and disk 0;
sular, septicidal; seeds carunculate.
ovary 3-locular, 6-horned or -winged, pubescent;
The two Australian genera of Ampereeae
ovules anatropous, inner and outer integuments
have been associated in the taxonomic literature
moderately thick; stylodia connate into a column,
since the treatment of M€ uller (1866), who placed
unlobed, apically dilated. Fruits capsular, 3-
them in separate subtribes of the tribe Amper-
lobed; columella persistent. Seeds globose, ecar-
eeae. Henderson (1992), on the basis of suppo-
unculate, testa minutely puberulent.
sedly intermediate characters in Amperea spicata
Eighteen spp., tropical Africa. The pollen of
Airy Shaw, has suggested that Monotaxis should
Pycnocoma is unique in its areolate tectum, which
probably be combined with Amperea. However,
somewhat simulates the pattern in Croton.
although Tokuoka and Tobe (2003) did not find
any significant differences in ovule and seed anat-
144. Droceloncia Léonard
omy, Nowicke et al. (1998), on the basis of pollen
Droceloncia Léonard, Bull. Soc. Roy. Bot. Belge 91: 279 characters, have indicated that there are clear
(1959); Webster, Ann. Missouri Bot. Gard. 81: 76 (1994); palynological distinctions between the genera.
Radcl.-Sm., Gen. Euphorb.: 167, fig. 18 (2001).
Furthermore, since there are distinct differences
Monoecious shrubs; indumentum simple; termi- in stipules, inflorescences, anthers, and stylodia,
nal buds perulate. Leaves subentire or dentate, amalgamation of the two genera seems prema-
eglandular; stipules deciduous. Inflorescences ture. The finding of Wurdack et al. (2005) that
axillary, spiciform, bisexual, staminate flowers Adriana appears in a clade with the two Amper-
in glomerules, pistillate solitary and apical; bracts eeae genera needs attention; also Nowicke et al.
large, persistent. Staminate flowers pedicellate; (1998) had observed a palynological resemblance
calyx closed in bud, splitting into several valvate between Amperea and Adriana.
134 G.L. Webster
KEY TO THE GENERA OF AMPEREEAE

1. Staminate petals +; sepals imbricate; anther connective
not glandular, locules disjunct; pollen grains 3-angled,
sexine reticulate; stylodia fimbriate; cymes terminal
145. Monotaxis
– Staminate petals 0; sepals valvate; anther locules
pendulous from a glandular connective; pollen grains
3-lobed, sexine perforate-foveolate; stylodia usually
entire; flowers axillary 146. Amperea
145. Monotaxis Brongn. Fig. 31

Monotaxis Brongn., Voy. Coq. Bot.: 223 (1829); Airy
Shaw, Muelleria 4: 239 (1980); Halford & Henderson,
Austrobaileya 6: 273–292 (2002), rev.
Monoecious (dioecious) perennial or annual herbs,
sometimes suffruticose; indumentum 0. Leaves
alternate (subopposite or verticillate), entire (den-
tate), one-veined, lateral veins obscure; stipules Fig. 31. Euphorbiaceae-Acalyphoideae. Monotaxis linifo-
entire or lobed, persistent. Inflorescences glomeru- lia. A Section of twig with stipulate leaf. B Inflorescence.
lar, terminal, each usually with 1–3 pistillate and C Staminate flower. D Auriculate petal. E Antepetalous
several staminate flowers subtended by small scar- stamen, abaxial view. F Fruit. (Halford & Henderson 2002;
drawn by W. Smith)
ious bracts. Staminate flowers pedicellate; sepals 4
or 5, often petaloid; petals 4 or 5, clawed, basally
auriculate; disk segments 4–5; stamens 8–10(11), Inflorescences [except in A. spicata] axillary,
filaments distinct; anthers 2-celled; anther cells dis- flowers in dense unisexual or bisexual glomer-
tinct, divergent to pendent on transverse connec- ules, subtended by dissected persistent bracts.
tive; pollen grains subprolate, angulaperturate, 3- Staminate flowers pedicellate; sepals 3–5(6),
colporate, colpi marginate, exine very thick, sexine basally connate, entire, petaloid; petals 0; disk
reticulate, transitional to tectate-perforate; pistil- rudimentary or 0; stamens 3–10, filaments dis-
lode trifid to subulate or 0. Pistillate flowers pedi- tinct, exserted; anthers with enlarged minutely
cellate; sepals 4 or 5, imbricate, persistent in fruit; glandular connective, thecae pendulous; pollen
petals + or 0; disk segments 3–10, truncate or bifid; grains oblate spheroidal, 3-colporate and 3-
ovary 3-locular; ovules anatropous, inner and outer lobed in polar view, colpi inoperculate, with
integuments thin; stylodia shortly connate at base, broad margo, sexine tectate-perforate and
distally distinct, deeply bifid, branches fimbriate. verrucate; pistillode 0. Pistillate flowers sub-
Fruit capsular; columella slender, persistent. Seeds sessile; sepals 4 or 5, nearly distinct, imbricate,
oblong, smooth, carunculate; endosperm copious; entire, persistent in fruit; petals 0; disk annular,
embryo cylindrical, straight or curved; cotyledons tenuous; ovary 3-locular; ovules anatropous,
much longer than radicle. inner and outer integuments thin; stylodia basally
Eleven spp., endemic to tropical and temper- connate, bifid. Fruits capsular, valves sometimes
ate Australia. with distal appendages; columella persistent.
Seeds ellipsoidal, somewhat compressed, carun-
culate; testa smooth (minutely foveolate-striate);
146. Amperea A. Juss. embryo cylindrical, cotyledons narrow and lon-
Amperea A. Juss., Euphorb. Tent.: 35 (1824); Henderson, ger than radicle.
Austral. Syst. Bot. 5: 1, figs. 1–4 (1992); Radcl.-Sm., Gen. Eight spp., all Australian, 6 of these confined
Euphorb.: 130 (2001). to Western Australia. The 3-lobed pollen with
Monoecious or dioecious perennial herbs; indu- broad margo and verrucate exine of Amperea
mentum scanty or 0. Leaves sessile or petiolate, resembles that of Bernardia. However, that
one-veined, laterals obscure, entire or dentate, genus differs in its larger number of pistillate
often revolute; sometimes scale-like; stipules sepals and ecarunculate seeds; the pollen similar-
entire to fimbriate, persistent to deciduous. ity may be homoplasious.
Euphorbiaceae 135
10. TRIBE EPIPRINEAE (M€

ull. Arg.) Hurus. (1954). 4. Pistillate flower sessile, calyx involucrate, accrescent in
fruit; stylodia connate into a distinct column; stamens
Monoecious; indumentum stellate; leaves alter- mostly 8–15; leaf blades with paired basal glands at
nate (opposite), petiole sometimes with apical junction with petiole 147. Epiprinus
paired glands; stipules sometimes glandular; – Pistillate flowers pedicellate, calyx not involucrate; sty-
lodia scarcely connate; stamens (3) 4 or 5; leaf blades
inflorescences terminal or axillary, paniculate, eglandular [except Cleidiocarpon] 5
racemoid, or the staminate capitulate; staminate 5. Filaments straight in bud; staminate sepals connate;
flowers sessile or subsessile; calyx splitting into stipules persistent 150. Koilodepas
2–6 valvate segments; petals and disk 0; stamens – Filaments inflexed in bud; staminate calyx of distinct
4–15(50); pollen grains 3-colporate, colpi inop- segments; stipules deciduous or obsolete 6
erculate and scarcely marginate, endoaperture 6. Monoecious; fruits capsular; leaves without laminar or
petiolar glands 148. Symphyllia
large, sexine tectate-perforate or coarsely reticu-
– Dioecious; fruits indehiscent; petiole with paired distal
late; pistillode +; pistillate flowers sessile or pedi- glands 149. Cleidiocarpon
cellate; sepals (4)5–8, imbricate, entire to lacerate, 7. Inflorescences axillary; stamens (2–)4(5); pistillate
usually persistent in fruit; petals and disk 0; ovary flowers subsessile, sepals unlobed, accrescent; leaf
3(4)-locular, stellate-tomentose; stylodia distinct blades white-tomentose beneath 151. Cladogynos
or connate, bifid to multifid (unlobed); fruits – Inflorescences terminal; stamens 6–10; pistillate flow-
ers pedicellate, sepals pinnately or bipinnately lobed
capsular (indehiscent); columella winged, usually
(entire); leaf blades not white-tomentose beneath
persistent; seeds subglobose, ecarunculate; testa 152. Cephalocroton
smooth (rugose).
Seven genera, all Asiatic except Cephalocro- 10a. SUBTRIBE EPIPRININAE M€
ull. Arg. (1865).
ton. Except for the enigmatic Rockinghamia,
which is added only tentatively, the Epiprineae Leaves pinnately or palmately veined; staminate
appear to be a monophyletic group, also in sepals distinct; stamens mostly 6–8 or more, fila-
the light of the molecular analysis (Wurdack ments distinct; pollen sexine not spinulose; pis-
et al. 2005), although the sampling (4 genera) is tillate sepals mostly persistent; capsules smooth.
limited. Rockinghamia has a habit suggestive Six paleotropical genera.
of Mallotus, but the studies of Nowicke et al.
(1999) show that its pollen differs from that of 147. Epiprinus Griffith
all Acalypheae in the lack of microverrucae, Epiprinus Griffith, Notul. Pl. Asiat. 4: 487 (1854); Croizat,
whereas it shares with Epiprineae the deeply J. Arnold Arb. 23: 52 (1942); Thin, Tâp Chi Sinh Vât Hoc
puntate-microreticulate exine sculpture. In the 10(2): 30 (1988); Radcl.-Sm., Gen. Euphorb.: 176 (2001).
rbcL tree of Wurdack et al. (2005), it is resolved
(without support) as basal to the genera of the Monoecious shrubs or trees; indumentum stellate.
Epiprineae. Leaves alternate or subopposite, clustered at ends
of branches, subsessile, subentire, with sparse
glands; petiole biglandular at apex; stipules entire,
KEY TO THE SUBTRIBES AND GENERA OF EPIPRI-
glandular at base, deciduous. Inflorescences ter-
NEAE
minal, spiciform, bisexual, with 1 or 2 proximal
1. Anthers 4-celled; leaves opposite; staminate disk as pistillate flowers and distal glomerules or spike-
interstaminate pubescent disk glands; pistillate disk lets of staminate flowers. Staminate flowers ses-
annular. Subtribe 10c. Rockinghamiinae
154. Rockinghamia
sile; calyx splitting into mostly 3 or 4 valvate
– Anthers 2-celled; leaves alternate; staminate and pistil- segments; petals and disk 0; stamens (5–)8–15,
late disk 0 2 filaments distinct, inflexed in bud; anthers
2. Pistillate sepals persistent in fruit; staminate calyx of introse, minutely apiculate; pollen grains oblate
distinct segments; pollen sexine not spinulose. Subtribe spheroidal, 3-colporate, colpi narrow, inopercu-
10a. Epiprininae 3 late and emarginate; sexine tectate-perforate tran-
– Pistillate sepals deciduous; staminate sepals connate; sitional to reticulate, with microverrucae at angles
pollen sexine spinulose. Subtribe
10b. Cephalomappinae 153. Cephalomappa of lumina; pistillode clavate, pubescent. Pistillate
3. Staminate flowers in racemoid or spiciform inflores- flowers pedicellate; calyx involucrate, bracts glan-
cences 4 dular at base, deciduous; sepals 5 or 6, imbricate,
– Staminate flowers in pedunculate capitula 7 entire, accrescent and persistent in fruit; ovary
136 G.L. Webster
3-locular, stellate-tomentose; ovules with inner introrse, 4-locellate; pollen grains prolate spheroi-
integument thin, outer thick; stylodia bifid or dal, 3-colporate, colpi inoperculate and emargin-
unlobed, papillose-lacerate. Fruits capsular, ate, sexine tectate-punctate; pistillode +. Pistillate
valves thick and woody; columella persistent. flowers subsessile; sepals 4–8, valvate, entire, per-
Seeds roundish, ecarunculate, testa smooth. sistent in fruit; petals and disk 0; ovary 2-locular,
Four spp., southeast Asia and Indonesia. stellate-tomentose; stylodia basally connate, dis-
tally distinct, multifid. Fruits indehiscent, drupa-
148. Symphyllia Baill. ceous, rugose, beaked. Seed solitary, rugose.
Two spp. from northern Burma, China, west-
Symphyllia Baill., Étude Gén. Euphorb.: 473 (1858);
Gagnep., Fl. Indochine 5: 477, fig. 60, 10–12 (1926); Web-
ern Thailand and Vietnam. Thin (1988) has pro-
ster, Ann. Missouri Bot. Gard. 81: 78 (1994); Radcl.-Sm., posed a new subtribe Cleidiocarpinae for it, and
Gen. Euphorb.: 177, fig. 21 (2001). the genus is indeed unusual in the drupaceous
fruit.
Monoecious trees or shrubs; indumentum stel-
late. Leaves eglandular; stipules glandular, decid- 150. Koilodepas Hassk.
uous. Inflorescences terminal or axillary,
spiciform or paniculate, usually bisexual; bracts Koilodepas Hassk., Versl. Med. Afd. Natuurk. Kon. Akad.
Wetensch. 4: 139 (1856), Flora 40: 531 (1857, Coelodepas);
inconspicuous. Staminate flowers subsessile; Croizat, J. Arnold Arb. 23: 50 (1942); Airy Shaw, Kew Bull.
calyx closed in bud, splitting into 3–6 valvate 14: 382 (1960), Kew Bull. 36: 310 (1981); Radcl.-Sm., Gen.
segments; petals and disk 0; stamens 3–6, mostly Euphorb.: 180, fig. 22 (2001).
4 or 5, filaments distinct, inflexed in bud; anthers
Monoecious trees; indumentum stellate. Leaves
muticous, 2-locellate; pollen grains prolate sphe-
entire or crenate-dentate, abaxially biglandular
roidal, 3-colporaate, colpi inoperculate and emar-
or with scattered glands; stipules dentate, per-
ginate, sexine tectate-peroratae, heterobrachate;
sistent. Inflorescences axillary, spiciform, usually
pistillode +. Pistillate flowers subsessile, not
bisexual, with 1 or few proximal pistillate flowers
involucrate; sepals 5–7, entire, valvate, persistent
and distal staminate or bisexual glomerules;
in fruit; petals and disk 0; ovary 3-locular, stel-
bracts ovate, dentate apically, eglandular. Sta-
late-tomentose; stylodia connate into a column
minate flowers sessile; calyx closed in bud,
longer than the ovary, bifid or twice bifid. Fruits
splitting into 3 or 4 valvate segments; petals and
capsular. Seeds smooth.
disk 0; stamens 4–5(8), filaments basally con-
Three spp., eastern Asia: India to Hainan and
nate, dilated above the middle, anthers introrse,
Malaya. Croizat (1942) combined Symphyllia
connective not enlarged, 2-locellate, locules
with Epiprinus, which was followed by Govaerts
divergent; pollen grains subspheroidal, 3-colpo-
et al. (2000); however, it differs from that genus in
rate, colpi narrow, inoperculate and emarginate;
its eglandular leaves and lack of a pistillate invo-
sexine tectate-punctate, psilate; pistillode +. Pis-
lucre. Symphyllia is therefore provisionally
tillate flowers sessile; sepals 4–10, basally con-
retained as a distinct genus.
nate, entire; persistent in fruit; petals and disk 0;
ovary (2)3-locular, stellate-tomentose; ovules
149. Cleidiocarpon Airy Shaw with thick integuments; stylodia bifid to multifid,
Cleidiocarpon Airy Shaw, Kew Bull. 19: 313 (1965), 32: 410 laciniate. Fruits capsular; columella persistent,
(1978); Thin, Tâp Chi Sinh Vât Hoc 10(2): 32 (1988); broadly winged. Seeds subglobose, testa smooth.
Webster, Ann. Missouri Bot. Gard. 81: 79 (1994); Rad- Eleven spp., distributed from India to Borneo
cliffe-Smith, Gen. Euphorb.: 179 (2001); Qiu Huaxing &
Gilbert, Fl. China 11: 250 (2008).
and New Guinea. Airy Shaw (1960) proposed 2 sec-
Sinopimelodendron Tsiang (1973). tions, one (sect. Hyalodepas Airy Shaw) including
2 spp. with accrescent fruiting calyx suggestive of
Monoecious trees; indumentum stellate. Leaves Epiprinus, and referring the other 9 spp. to sect.
apically clustered on branches, eglandular; petiole Koilodepas, with non-accrescent calyx.
stipellate at apex; stipules deciduous. Inflores-
cences terminal, spiciform-paniculate, bisexual or 151. Cladogynos Zipp. ex Span.
sometimes unisexual; bracts and prophylls con-
Cladogynos Zipp. ex Span., Linnaea 15: 349 (1841); Pax &
spicuous. Staminate flowers subsessile; sepals 4 K. Hoffm., Pflanzenr. 147, VII: 264, fig. 41 (1914); Gagnep.,
or 5, valvate; petals and disk 0; stamens 4 or 5, Fl. Indochine 5: 478, fig. 61 (1926); Airy Shaw, Kew Bull. 26:
filaments distinct, inflexed in bud; anthers 232 (1972); Radcl.-Sm., Gen. Euphorb.: 182 (2001).
Euphorbiaceae 137
Monoecious shrubs; indumentum stellate. Leaves sular, 3-lobed, glandular-verrucate; columella

pinnately veined or triplinerved, double-dentate, persistent, broadly winged. Seeds roundish,
peltate or subpeltate at base, eglandular; stipules smooth or sparsely pubescent.
minute, deciduous. Inflorescences axillary, bisex- About seven spp. in tropical Africa, Mada-
ual, of 1 proximal pedicellate pistillate flower and gascar, Socotra and Ceylon.
a distal capitulum of staminate flowers. Staminate In Webster (1994), Adenochlaena and Cepha-
flowers sessile; petals and disk 0; stamens (3)4(5), locrotonopsis were accepted as distinct genera.
filaments distinct, inflexed in bud; anthers Radcliffe-Smith (2001) also accepts both genera,
introrse, muticous, locules; pollen grains 3-colpo- even though he earlier sank Cephalocrotonopsis
rate, colpi narrow, sexine tectate-perforate; pistil- into Cephalocroton. However, the pollen grains of
lode +. Pistillate flowers pedicellate; sepals 5–7, these taxa are identical (Takahashi et al. 2000),
subfoliaceous, basally contracted, subentire, per- and it now seems preferable to treat both Adeno-
sistent in fruit; petals and disk 0; ovary 3(4)- chlaena and Cephalocrotonopsis as sections of an
locular, pubescent; ovules with inner integument enlarged genus Cephalocroton, distinguished by
thick, outer thin; stylodia basally connate, twice its unique inflorescence type.
bifid or more, papillose. Fruits capsular, colu-
mella persistent, apically dilated. Seeds spheroi- 10b. SUBTRIBE CEPHALOMAPPINAE G.L. Webster
dal, testa smooth. (1975).
A single sp., C. orientalis Zipp. ex Span.,
widespread in southeast Asia and Malesia. A monotypic subtribe.
152. Cephalocroton Hochst. 153. Cephalomappa Baill.

Cephalocroton Hochst., Flora 24: 370 (1841); M€ ull. Arg. in Cephalomappa Baill., Adansonia 11: 130 (1874); Pax,
DC., Prodr. 15(2): 760 (1866); Pax, Pflanzenr. 147, II: 7, Pflanzenr. 147, II: 16. 1910; Airy Shaw, Kew Bull. 14: 378
figs. 3, 4 (1910); Prain, Fl. Trop. Afr. 6(1): 843 (1912); (1960); Kosterm., Reinwardtia 5: 413 (1961); Backer &
Radcl.-Sm., Kew Bull. 28: 123 (1973); Fl. E. Trop. Afr. Backh., Fl. Java 1: 486 (1963); Whitmore, Tree Fl. Malaya
Euphorb. 1: 282, fig. 55 (1987); Gilbert, Kew Bull. 42: 2: 75, fig. 4 (1973); Airy Shaw, Kew Bull. 36: 274 (1981);
365, fig. 6 (1987); Webster, Ann. Missouri Bot. Gard. 81: Widuri & van Welzen, Reinwardtia 11: 153, figs. 1–16
79 (1994); Radcl.-Sm., Gen. Euphorb.: 183 (2001). (1998); Radcl.-Sm., Gen. Euphorb.: 186, fig. 23 (2001).
Adenochlaena Baill. (1858). Monoecious trees or shrubs; indumentum sim-
Cephalocrotonopsis Pax (1910).
ple, stellate, and lepidote. Leaves entire to den-
Monoecious shrubs; indumentum stellate and tate, eglandular or sometimes with abaxial
glandular-setose. Leaves palmately veined or tri- glands; petioles pulvinate; stipules deciduous.
plinerved, usually dentate, eglandular; stipules Inflorescences terminal or axillary, racemiform,
persistent, sometimes minute or obsolete. Inflor- sometimes compounded and corymbose; stami-
escences terminal (axillary), the staminate flow- nate flowers in pedunculate capitula, pistillate
ers in distal capitula, the pistillate flowers flowers 1–3 at proximal axils; bracts deciduous.
proximal, separated by a long internode. Stami- Staminate flowers sessile; calyx turbinate, trun-
nate flowers pedicellate (sessile), articulate at cate, 2–5-lobed, lobes valvate; petals and disk 0;
base of calyx; sepals closed in bud, splitting into stamens (2)3–4, filaments connate into a column,
3–6 valvate lobes; petals and disk 0; stamens (4–) tips inflexed in bud; anthers latrorse, connective
6–8(10), filaments distinct, inflexed in bud; glandular, muticous, 2-celled; pollen grains
anthers dorsifixed, introrse, muticous, 2-locellate; oblate spheroidal, 3-brevicolporate, sexine
pollen grains prolate spheroidal, 3-colporate, coarsely reticulate and muri spinulose; pistillode
colpi narrow, inoperculate and emarginate, sex- clavate, sometimes apically lobed. Pistillate flow-
ine tectate-perforate, muri microverrucate; pistil- ers pedicellate; sepals 4–6(8), imbricate, con-
lode 2- or 3-lobed. Pistillate flowers pedicellate, nate at base, deciduous; petals and disk 0; ovary
the pedicel usually greatly elongating in fruit; 3-locular, stellate-tomentose or lepidote; ovules
sepals 4–6, pinnately or bipinnately lobed anatropous, inner integuments very thick, outer
(entire), persistent and accrescent in fruit; petals integuments thick; stylodia connate, tips bifid
and disk 0; ovary 3-locular, stellate-tomentose; or palmately incised. Fruits capsular, verrucose,
ovules with inner integuments thicker than 3-lobed, lepidote. Seeds spheroidal, hilum ovate,
outer integuments; stylodia multifid. Fruits cap- testa smooth.
138 G.L. Webster
Five spp., southern China, Malay Peninsula, escences axillary or terminal, mostly racemoid
Sumatra and Borneo. The pollen of Cephalo- or spicate; staminate flowers subsessile or pedi-
mappa is so distinctive that Takahashi et al. cellate; calyx splitting into 3–5 valvate sepals;
(2000) suggest the genus should be placed in a petals (0)4–5(10); disk dissected or 0; stamens
separate tribe. However, in other features it 5–many, filaments distinct or connate; connec-
appears similar to genera of Epriprineae, and in tives generally not enlarged and thecae not pen-
the molecular analysis of Wurdack et al. (2005) it dulous; pollen grains mostly 3–6-colporate, colpi
is resolved in a clade with Koilodepas and Cepha- inoperculate, sexine reticulate and often dis-
locroton. tinctly heterobrochate; pistillode + or 0; pistil-
late flowers usually pedicellate; sepals 5 or 6,
10c. SUBTRIBE ROCKINGHAMIINAE G.L. Webster imbricate or valvate, usually entire, persistent
(2004). in fruit; petals 5 (reduced or 0); disk annular
or dissected (0); ovary (2)3-locular; stylodia
A monogeneric Australasian subtribe. unlobed to bifid or twice bifid, sometimes lacer-
ate; fruit capsular; columella usually persistent;
154. Rockinghamia Airy Shaw seeds ecarunculate.
Rockinghamia Airy Shaw, Kew Bull. 20: 29 (1966), 35: 667, With 11 genera and 125 species or more, tribe
fig. 6 (1980); Hyland & Whiffin, Austral. Trop. Rain For. Chrozophoreae is widespread and diverse in
Trees: 150 (1993); Radcl.-Sm., Gen. Euphorb.: 232 (2001). warm temperate to tropical areas. As the tribal
Monoecious trees; indumentum simple, sparse. description indicates, the morphological diversity
Leaves mostly opposite, often pseudo-verticillate is extreme, and the molecular data of Wurdack
at end of branches, entire or crenate, eglandular; et al. (2005) (still with incomplete sampling)
petioles pulvinate; stipules deciduous. Inflores- point to placements in different clades of the
cences terminal (axillary), racemoid or panicu- acalyphoids: Ditaxinae as sister to Adelieae, and
late, bisexual; bracts glandular, entire, persistent. Speranskiinae and Chrozophorinae into a clade
Staminate flowers pedicellate; calyx closed in together with Epiprineae. Available data permit,
bud, splitting into 4 valvate segments; petals 0; however, the transfer of Caperonia from Sperans-
disk segments numerous, interstaminal, pubes- kiinae to Ditaxinae, and to merge Doryxylinae
cent; stamens 25–50, filaments distinct; anthers with monotypic Chrozophorinae.
4-celled, muticous; pollen grains prolate spheroi-
dal, 3-colpor(oid)ate, inoperculate and emargin- KEY TO THE SUBTRIBES OF CHROZOPHOREAE
ate, endoapertures indistinct; sexine perforate-
tectate to reticulate with narrow muri and deep 1. Indumentum stellate; leaves with basal laminar glands
11c. Chrozophorinae
lumina, sometimes heterobrochate; pistillode + – Indumentum simple or malpighiaceous, or if stellate
or 0. Pistillate flowers pedicellate; sepals 5, imbri- then leaves without basal laminar glands 2
cate, subentire, persistent in fruit; petals 0; disk 2. Inflorescence terminal; indumentum simple; stems not
annular; ovary 3-locular, pubescent; ovules anat- spinose; ovary muricate 11a. Speranskiinae
ropous, inner integument thick, outer thin; stylo- – Inflorescence axillary; indumentum malpighiaceous or
dia distinct or basally connate, bifid or unlobed. stellate (entirely simple and stems spinose in Philyra)
11b. Ditaxinae
Fruits capsular, 3-lobed, muricate; columella per-
sistent, winged-dilated. Seeds roundish or
oblong, hilum deltoid, testa smooth. 11a. SUBTRIBE SPERANSKIINAE G.L. Webster (1975).
Two spp., tropical Australia (Queensland).
Monoecious herbs; indumentum simple; leaves
11. TRIBE CHROZOPHOREAE (M€
ull. Arg.) Pax & glandular-dentate; inflorescences terminal, spici-
K. Hoffm. (1919). form; sepals and petals usually 5 in both sexes;
stamens 10–15, filaments distinct; pollen grains
Monoecious or dioecious trees, shrubs, or herbs; 3-colporate; pistillate flowers pedicellate or sub-
indumentum stellate, lepidote, malpighiaceous, sessile; disk cupular or 0; ovary usually muricate;
or simple; leaves alternate, unlobed or lobed, inner and outer ovular integuments thin; stylodia
entire or dentate, sometimes with laminar distinct, branches lacerate; seed coat dry.
glands; stipules persistent or deciduous; inflor- A monogeneric Asian subtribe.
Euphorbiaceae 139
155. Speranskia Baill. – Leaves not as above; disk +, at least in staminate flow-
ers; pollen 3–4-colporate; pistillode 0 or scarcely devel-
Speranskia Baill., Étude Gén. Euphorb.: 388 (1858); Pax & oped; stylodia bifid to dissected 2
K. Hoffmann, Pflanzenr. 147,VI: 14, fig. 3 (1912); Hwang 2. Dioecious; tepals scarious; shrubs or trees, branches
Shu-mei, Bull. Bot. Res. Harbin 9 (4): 37–40 (1989); often spiny 156. Philyra
Radcl.-Sm., Gen. Euphorb.: 138 (2001); Qiu Huaxing &
– Monoecious; tepals not scarious; undershrubs or herbs,
Gilbert, Fl. China 11: 223 (2008).
not spiny 3
Monoecious perennial herbs; indumentum sim- 3. Petals 3–7-lobed or -partite; stylodia once bifid; stellate
hairs sometimes present; pollen grains 3-colporate
ple. Leaves petiolate or sessile, glandular-dentate, 159. Chiropetalum
without laminar glands, stipellate at base; stipules – Petals entire; stylodia twice bifid to multifid; stellate
deciduous or persistent. Inflorescences terminal, hairs 0 4
spiciform; proximal cymules bisexual or pistil- 4. Stamens 8–10, anthers in 2 whorls; pollen grains mainly
late, distal staminate; bracts eglandular, persis- 3-colporate, bilaterally symmetrical 158. Ditaxis
tent. Staminate flowers pedicellate; calyx – Stamens 4 or 5, anthers in 1 whorl; pollen grains ste-
splitting into 5 valvate sepals; petals 5; disk seg- phanocolporate, not bilaterally symmetrical
157. Argythamnia
ments 5; stamens 10–15, filaments distinct;
anthers in 2 (3) whorls, dorsifixed, extrorse; pollen
156. Philyra Klotzsch
grains subprolate, 3-colporate, colpi operculate,
endoaperture lalongate, exine tectate-punctate to Philyra Klotzsch, Arch. Naturgesch. 7(1): 199 (1841);
tectate-reticulate; pistillode 0. Pistillate flowers Pax & K. Hoffm., Pflanzenr. 147,VI: 49, fig. 10 (1912);
O’Donell & Lourteig, Lilloa 8: 60 (1942); Radcl.-Sm.,
pedicellate; sepals 5, distinct, lanceolate, entire, Gen. Euphorb.: 141 (2001).
deciduous or persistent; petals 5 or 0; disk cupular; Argythamnia sect. Philyra (Klotzsch) M€
ull. Arg. (1865).
ovary 3-locular, verruculose or tuberculate; inner
and outer integuments thin; stylodia distinct, bifid, Dioecious trees or shrubs; indumentum 0 except
branches lacerate. Fruits capsular, 3-lobed; colu- for simple hairs on inflorescences. Leaves alternate;
mella deciduous or persistent. Seeds globose, testa stipules persistent, indurate, often associated with
smooth or foveolate to rugulose. long infrastipular spines. Inflorescences axillary,
Two spp., endemic to China. brachyblastic, or the pistillate flowers on expanded
tenuous racemes; bracts indurate, persistent. Sta-
11b. SUBTRIBE DITAXINAE Griseb. (1859). minate flowers pedicellate; sepals 3–5, fused in bud,
splitting into valvate segments; petals 5, distinct;
Monoecious (dioecious) trees, shrubs, or herbs; disk segments 5, adnate to staminal column;
indumentum simple, glandular, stellate or mal- stamens (8) 10, filaments connate below into a
pighiaceous; leaves entire or serrate, stipulate; column; anthers in 2 superposed whorls, extrorse;
inflorescences axillary, racemoid or glomerulate; pollen grains prolate spheroidal to subprolate, 3-
staminate sepals mostly 4 or 5; petals 5, distinct colporate, colpi exoperculate and emarginate,
or adnate to staminal column; disk dissected, endoaperture circular; sexine tectate-punctate; pis-
reduced, or 0; stamens 4–15, filaments connate tillode trifid, at apex of staminal column. Pistillate
(almost distinct); anthers introrse, pollen grains flowers long-pedicellate; sepals 5, linear-lanceolate,
3–4-colporate; pistillode + or 0; pistillate flowers entire, persistent in fruit; petals 5, longer than
sessile or pedicellate; sepals 5 (6), distinct, imbri- sepals; disk thickened as gynophore; ovary 3-locu-
cate; petals 5 or 0; disk dissected or 0; ovary lar; ovules with inner integument moderately thick,
3-locular, smooth or muricate; stylodia bifid to outer thin; stylodia bifid, branches multifid or
laciniate; seeds with dry testa, smooth or orna- lobed. Fruits capsular; columella persistent, trifid
mented. at apex. Seeds globose; testa smooth.
As here delimited, subtribe Ditaxinae includes A single sp., P. brasiliensis Klotzsch, from
5 mostly neotropical genera with 110–115 species. Argentina and Paraguay to northeastern Brazil
(Bahia), in deciduous or semi-deciduous tropical
KEY TO THE GENERA OF DITAXINAE woodlands. Although appearing in a clade with
three adelioid genera (Wurdack et al. 2005), this
1. Leaves finely serrate with straight parallel lateral
nerves; disk 0; pollen 3–6-colporate; pistillode +; stylo- petaloid genus differs significantly from them,
dia dissected 160. Caperonia and the pollen is very distinct.
140 G.L. Webster
157. Argythamnia P. Browne filaments connate into a column; anthers biseriate,

Argythamnia P. Browne, Civ. Nat. Hist. Jamaica: 338
introrse, dehiscing vertically; pollen grains
(1756); Ingram, Gentes Herb. 10(1): 1, figs. 3–19 (1967); oblate, asymmetrically 3-colporate; colpi inop-
Correll, Fl. Bahama Arch. 777, fig. 325 (1982); Howard, Fl. erculate; sexine tectate-punctate (superreticulate);
Lesser Ant. 5: 14, fig. 5 (1989); Webster, Ann. Missouri pistillode at apex of staminal column, divided (0).
Bot. Gard. 81: 72 (1994); L.J. Gillespie, Fl. St. John 203 Pistillate flowers pedicellate; sepals 5, distinct,
(1996); Radcl.-Sm., Gen. Euphorb.: 143 (2001).
imbricate, entire or denticulate, persistent in fruit;
Monoecious (dioecious) shrubs or subshrubs; petals 5, imbricate, entire, often persistent in fruit;
indumentum malpighiaceous. Leaves entire or disk segments 5, often confluent; ovary 3-locular;
denticulate, pinnately veined (triplinerved), with glabrous or pubescent; ovules with inner integu-
steeply ascending laterals; stipules entire, ment moderately thick, outer thin; stylodia bifid,
persistent. Inflorescences axillary, glomerular branches sometimes emarginate or shortly bifid.
or racemiform, pistillate flowers proximal and Fruits capsular; columella persistent, 3-pronged.
staminate distal in bisexual inflorescences; bracts Seeds smooth to rugose or foveolate.
1-flowered, persistent. Staminate flowers subses- About 40–50 spp. ranging from the USA
sile; sepals connate in bud, dehiscing into 4(5) south to Argentina, primarily in subarid regions.
valvate segments; petals 4(5), shorter than sepals, The distinct pollen grains furnish the best diag-
imbricate, entire; disk segments 4(5), adnate to nostic character (Takahashi et al. 1995; Nowicke
base of staminal column; stamens 4–6, filaments et al. 1999).
connate at the base, sometimes forming a short
column; anthers introrse; pollen grains 4-colpo- 159. Chiropetalum A. Juss.
rate, colpi inoperculate, sexine tectate-perforate; Chiropetalum A. Juss., Ann. Sci. Nat. (Paris) 25: 21 (1832);
pistillode rudimentary. Pistillate flowers subses- O’Donell & Lourteig, Lilloa 8: 38, figs. 1–3, t. II (1942);
sile; sepals 5(6), distinct, valvate, entire, persis- Ingram, Gentes Herb. 11(7): 437, figs. 4–9 (1980); L.B.
tent in fruit; petals 5, distinct, entire, often Smith et al., Fl. Ilustr. Catar. EUFO: 154 (1988); Radcl.-
reduced or rudimentary; disk divided into 5 seg- Sm., Gen. Euphorb.: 144 (2001).
Argythamnia subg. Chiropetalum (A. Juss.) Ingram (1980).
ments; ovary 3-locular; ovules with inner and
outer integuments thin; stylodia bifid or twice Monoecious shrubs or subshrubs; indumentum
bifid. Fruits capsular; columella persistent. Seeds malpighiaceous, sometimes also simple or stel-
round, testa dry, roughened (alveolate). late. Leaves alternate, serrate (entire), 3–5-veined
About 18 spp., most of which occur in the from base; stipules entire, persistent. Inflores-
West Indies, and 8 from S Mexico to Honduras. cences axillary, racemiform, pistillate flowers
proximal, staminate flowers distal; bracts 1-flow-
158. Ditaxis Vahl ex A. Juss. ered, persistent. Staminate flower pedicellate;
sepals connate in bud, dehiscing into 5 valvate
Ditaxis Vahl ex A. Juss., Euphorb. Tent.: 27 (1824);
O’Donell & Lourteig, Lilloa 8: 62, figs. 5–10, t. I, III–VII
segments; petals 5, distinct, imbricate, unguicu-
(1942); Wiggins, Fl. Sonoran Desert: 784 (1964); Webster, late, 3–7-lobed or -partite; disk segments 5;
Jepson Man. Pl. Calif.: 572 (1993); Diggs et al., Fl. North stamens 5 (6), filaments connate into a column;
Central Texas: 600 (1999); Radcl.-Sm., Gen. Euphorb.: 142 anthers stipitate atop column; pollen grains sphe-
(2001). roidal to prolate spheroidal, 3-colpate, colpi oper-
Argythamnia sect. Ditaxis (P. Browne) M€ ull. Arg. (1865).
culate, emarginate; exine finely tectate; pistillode
Monoecious (dioecious) shrubs, perennial herbs, rudimentary. Pistillate flower pedicellate; sepals
and annual herbs; indumentum malpighiaceous. 5, valvate, entire, persistent in fruit; petals 5,
Leaves entire or dentate, pinnately veined (tripli- shorter than sepals, entire (lobed), sometimes
nerved), often with steeply ascending laterals; sti- obsolete; disk segmented into 5 lobes; ovary 3-
pules entire, persistent. Inflorescences axillary, locular; ovules with inner and outer integuments
glomerular or racemiform, typically bisexual. Sta- thin; stylodia distinct or basally connate, bifid.
minate flowers pedicellate; sepals connate, separat- Fruits capsular; columella persistent. Seeds sphe-
ing into usually 5 segments at anthesis; petals 5, roidal; testa rugose.
imbricate, distinct but each petal unguiculate and About 22 spp., 2 in Mexico and 20 in South
adnate to base of staminal column; stamens (7–)10, America: Peru and Chile to S Brazil. Nowicke
Euphorbiaceae 141
et al. (1999) point out that the 3-colpate opercu- ual; staminate sepals 3–5; petals 5–10 or 0; disk
late pollen grains of Chiropetalum are very differ- reduced or 0; stamens (4–)30–250, distinct or
ent from the 4-colporate inoperculate grains of partially connate; pollen grains 3(6–9)-colporate,
Argythamnia, contradicting the merging of the angulaperturate, sexine reticulate; pistillate sepals
two genera by Ingram (1967). 5–6, imbricate or valvate, petals rudimentary or 0;
disk annular or obsolete; ovary 2–3-locular; sty-
160. Caperonia St. Hil. lodia bifid or unlobed; seeds with fleshy exotesta,
Caperonia St. Hil., Pl. Remarq. Brésil: 244 (1826); Fawc. & foveolate endotesta.
Rendle, Fl. Jam. 4: 288, fig. 93 (1920); O’Donell & Lourteig, An Old World, mainly Asiatic, subtribe of
Lilloa 8: 54, fig. 4 (1942); Léonard, Fl. Congo 8(1): 166, 5 genera.
t. 11 (1962); Webster, J. Arnold Arb. 48: 363 (1967),
Ann. Missouri Bot. Gard. 54: 265, fig. 10 (1968); Radcl.-
Sm., Fl. E. Trop. Afr. Euphorb. 1: 163, fig. 29 (1987); KEY TO THE GENERA OF CHROZOPHORINAE
Webster, Fl. Venez. Guayana 5: 102, fig. 100 (1999), Fl. 1. Petals 0; ovary usually 2-locular; stamens 200 or more
Nicaragua: 856 (2001); Radcl.-Sm., Gen. Euphorb.: 140 164. Melanolepis
(2001).
– Petals +; ovary 3-locular; stamens 4–100 2
Monoecious annual or perennial herbs, some- 2. Stamens connate; pollen grains stephanocolporate;
times rhizomatous or fruticose, stems often hol- herbs or subshrubs 165. Chrozophora
– Stamens distinct [except in Thyrsanthera]; pollen
low; indumentum simple, sometimes prickly or grains 3-colporate; shrubs or trees 3
glandular. Leaves alternate, serrate, pinnately and 3. Inflorescences axillary; stylodia unlobed, basally con-
strikingly parallel-veined; stipules scarious, per- nate; stamens distinct 161. Doryxylon
sistent. Inflorescences axillary, spiciform, usually – Inflorescences terminal; stylodia bifid, distinct 4
bisexual, with 1–5 basal pistillate flowers; bracts 4. Stamens distinct; staminate disk 0; pistillate sepals
entire, persistent, subtending solitary flowers. imbricate 162. Sumbaviopsis
Staminate flowers with articulate pedicels; calyx – Stamens connate; staminate disk +; pistillate sepals
valvate 163. Thyrsanthera
of 5 valvate lobes fused in bud; petals 5, often
unequal, basally adnate to the staminal column;
161. Doryxylon Zoll.
disk 0; stamens 10, filaments partly connate into a
column, anthers in 2 superposed whorls; pollen Doryxylon Zoll., Tijdschr. Ned.-Indië 14: 172 (1857);
grains subspheroidal, 3- or 6-colporate, colpi N.P. Balakr., Bull. Bot. Surv. India 9: 56, figs. 1–7 (1967);
Webster, Ann. Missouri Bot. Gard. 81: 72 (1994);
operculate, emarginate, endoapertures obscure, van Welzen, Blumea 44: 422, fig. 3 (1999).
germ pores large, sexine tectate-punctate to -
reticulate; pistillode at apex of staminal column. Monoecious trees or shrubs; twigs often with
Pistillate flowers subsessile; calyx deeply 5-lobed, axillary spines; indumentum simple and stellate.
with 0–5 smaller supernumerary lobes, persistent Leaves palmately veined, dentate, whitish-tomen-
in fruit; petals 5, often unequal or reduced; disk 0; tose abaxially; stipules minute, deciduous. Inflor-
ovary 3-locular, usually muricate; ovules anatro- escences terminal, racemoid, often unisexual;
pous, inner and outer integuments thin; stylodia bracts uniflorous. Staminate flowers pedicellate
distinct, laciniate. Fruit capsular, valves thin, usu- or subsessile; sepals 5, valvate; petals 5, shorter
ally verrucose; columella persistent, apically than sepals; disk 0; stamens 30–130, filaments
dilated. Seeds globose; testa minutely foveolate. distinct, inserted on hairy receptacle; anthers
n ¼ 11. dorsifixed, latrorse; pollen grains oblate spheroi-
About 35 spp., mostly American, but a few dal, angulaperturate, 3-colporate, colpi inopercu-
spp. in Africa and Madagascar, widely spread in late and emarginate, sexine reticulate,
marshy habitats. heterobrochate; pistillode 0. Pistillate flowers
pedicellate; sepals 5 or 6, imbricate, entire; petals
11c. SUBTRIBE CHROZOPHORINAE M€
ull. Arg. (1865). 0; disk annular, glabrous; ovary 3(4)-locular,
tomentose; stylodia unlobed, connate below.
Doryxylinae G.L. Webster (1975).
Fruits capsular; columella persistent. Seeds with
Monoecious trees, shrubs or herbs; indumentum fleshy exotesta, foveolate endotesta.
stellate or lepidote; leaves with laminar glands; A single Malesian sp., D. spinescens Zoll.,
inflorescences terminal or axillary, usually bisex- from the Philippines and Lesser Sunda Islands.
142 G.L. Webster
162. Sumbaviopsis J.J. Sm. 3-locular, tomentose; stylodia distinct, unlobed,

Sumbaviopsis J.J. Sm., Meded. Dept. Landb. Ned.-Indië
papillose. Fruits capsular, thin-walled; columella
10: 356 (1910); Airy Shaw, Kew Bull. Add. Ser. 4: 197 slender, persistent, apically 3-angled. Seeds
(1975); van Welzen, Blumea 44: 426, fig. 4 (1999); obovoid, angular, with thin fleshy exotesta, meso-
Radcl.-Sm., Gen. Euphorb.: 148, fig. 13 (2001). testa vascularized.
Monoecious trees or shrubs; indumentum mainly A single sp., Th. suborbicularis Pierre ex
stellate. Leaves peltate, palmately veined or tripli- Gagnep., recorded only from Thailand and
nerved, subentire or dentate, densely stellate- Cambodia.
tomentose abaxially, with small embedded
glands; stipules deciduous. Inflorescences termi- 164. Melanolepis Rchb. f. & Zoll.
nal (pseudo-axillary), unisexual, staminate thyr- Melanolepis Rchb. f. & Zoll., Verh. Natuurk. Ver. Ned.
soid, pistillate racemoid; bracts subtending 1–3 Ind. 1: 22 (1856); Gagnep., Fl. Indochine 5: 347 (1925);
staminate flowers or solitary pistillate flowers. Airy Shaw, Kew Bull. Add. Ser. 8: 174, t. 5 fig. 3 (1980); van
Welzen, Blumea 44: 438, fig. 1 (1999).
Staminate flowers pedicellate; calyx closed in
bud, splitting into 3–5 valvate segments; petals 4 Dioecious trees or shrubs; indumentum mainly
or 5, shorter than sepals; disk represented by a stellate or dendritic. Leaves long-petiolate, del-
convex pubescent torus; stamens 35–45(75?), toid, palmately veined, coarsely dentate, often 3-
inserted on torus, filaments distinct; anthers basi- lobed, with raised glandular area at junction with
fixed, introrse; pollen grains suboblate, 3-colpo- petiole; stipules inconspicuous, deciduous.
rate, colpi inoperculate, emarginate; sexine Inflorescences terminal, paniculate with 2 or 3
coarsely reticulate; pistillode 0. Pistillate flowers elongated racemoid or spiciform axes, usually
pedicellate; sepals 5(6), imbricate, entire, persis- unisexual; bracts entire, subtending several sta-
tent in fruit; petals 0; ovary 2- or 3-locular, minate flowers or 1(2) pistillate flower. Staminate
tomentose; ovules anatropous; stylodia distinct flowers pedicellate; sepals (4)5, valvate; petals 0;
or nearly so, bifid. Fruits capsular, valves laterally disk 0; stamens 200–250, filaments distinct,
corrugated; columella persistent, apically dilated. inserted on convex receptacle; anthers introrse,
Seeds spheroidal, exotesta fleshy, endotesta smooth. with glandular connective; pollen grains subo-
A single sp., S. albicans (Blume) J.J. Sm., blate, angulaperturate, 3-colporate, colpi inoper-
distributed from Thailand and Laos to Java, culate but with granular membranes, emarginate,
Borneo, and Palawan. endoaperture ovate to lalongate, exine tectate-
punctate to reticulate; pistillode 0. Pistillate flow-
163. Thyrsanthera Pierre ex Gagnep. ers pedicellate; sepals 5, basally connate, valvate,
Thyrsanthera Pierre ex Gagnep., Bull. Soc. Bot. France 71:
entire, persistent in fruit; petals 0; disk annular;
878 (1924); Gagnep., Fl. Indochine 5: 299, figs. 32, 33 ovary 2–3-locular, tomentose; ovules anatropous;
(1925); Backer & Backh., Fl. Java 1: 477 (1963); van Wel- stylodia unlobed or emarginate. Fruits capsular;
zen, Blumea 44: 431, fig. 5 (1999); Radcl.-Sm., Gen. leathery, valves and columella persistent. Seeds
Euphorb.: 148, fig. 14 (2001). roundish, with thin sarcotesta, foveolate endo-
Monoecious shrubs; indumentum stellate. Leaves testa.
palmately veined, entire, at margin with reddish Two Asiatic spp., one endemic to Cambodia,
indumentum, stipellar abaxially at junction with the other distributed from Thailand to Malesia
petiole; stipules deciduous. Inflorescences termi- and the Pacific islands (Ryukyus, Micronesia).
nal, racemoid, bisexual, pistillate flowers proxi-
mal; bracts pubescent, 1-flowered (?). Staminate 165. Chrozophora Necker ex A. Juss. Fig. 32
flowers pedicellate; sepals 5, valvate, tomentose Chrozophora Necker ex A. Juss., Euphorb. Tent.: 27 (1824,
abaxially; petals 5, imbricate; disk 0; stamens nom. cons.); Prain, Kew Bull. Misc. Inf. 1918: 49 (1918);
40–60, filaments connate into a column; anthers Pojark., Fl. SSSR 14: 288, t. 17, 1–4 (1949); Vindt, Monogr.
stipitate, basifixed, latrorse; pollen grains subo- Euphorb. Maroc. 10, t. 6 (1953); Rech. f. & Schiman-
Czeika, Fl. Iranica 6: 5 (1964); Zohary, Fl. Palaestina 2:
blate, 3-colporate, exine reticulate; pistillode 0. 266, t. 384–387 (1972); Radcl.-Sm., Fl. Iraq 4(1): 318, t. 59
Pistillate flowers sessile or subsessile; sepals 5, (1980), Fl. Zambesiaca 9(4): 141, t. 26 (1996); van Welzen,
narrow, entire, valvate; disk annular; ovary Blumea 44: 418, fig. 2 (1999).
Euphorbiaceae 143
fruit; disk annular; ovary 3-locular, smooth or

muricate, with stellate or lepidote hairs; ovules
anatropous; stylodia distinct except at base, bifid.
Fruits capsular, valves thin; columella persistent.
Seeds ovoid to subglobose, ecarunculate, with
thin fleshy sarcotesta, mesotesta vascularized.
n ¼ 11, 22.
About 10 spp., central and southern Asia,
southern Europe to East Africa. The genus
appears very different in habit from the other
genera of the subtribe, but resembles them by
characters of indumentum and seeds, and in the
molecular analysis of Wurdack et al. (2005) is
resolved in the same clade. The ovules of Chrozo-
phora have thin inner and outer integuments, as
opposed to the much thicker inner and outer
integuments of the remaining genera of the sub-
tribe (Tokuoka and Tobe 2003).
12. TRIBE ADELIEAE G.L. Webster (1975).

Dioecious (monoecious) trees or shrubs; indu-
mentum simple or stellate; leaves alternate, sim-
ple; lamina pinnately or palmately veined, entire
or dentate, eglandular; stipules small; flowers in
axillary glomerules or racemes; staminate flowers
pedicellate; calyx closed in bud, splitting into 4 or
5 valvate segments; petals 0; disk annular or obso-
lete; stamens 8–18(30), filaments distinct or
connate at base; anthers introrse, muticous, ver-
Fig. 32. Euphorbiaceae-Acalyphoideae. Chrozophora rot- satile; pollen grains 3(4)-colporate, spheroidal,
tleri. A Flowering/fruiting branchlet. B Abaxial side of colpi emarginate, operculate, opercula stratified;
leaf base with extrafloral nectaries. C, D Staminate flower. sexine tectate-perforate or reticulate, microverru-
E Staminate petal. F Pistillate flower. G Fruit. H Seed. cate, the sculpture crotonoid; pistillode usually +;
I Fruit column after dehiscence. (van Welzen 1999;
drawn by J. van Os) pistillate flowers pedicellate; sepals 5 or 6, valvate
or open at anthesis, persistent in fruit; petals 0;
disk annular; ovary 3-locular, pubescent; ovules
Monoecious herbs or subshrubs; indumentum with inner integuments thick, outer thin to thick;
stellate or lepidote. Leaves subentire or crenate, stylodia bifid to multifid; fruits capsular; colu-
palmately veined, with paired sessile patelli- mella persistent, apically 3-pronged; seeds round-
form glands near junction with petiole; stipules ish, ecarunculate, exotegmic, testa smooth.
deciduous or persistent. Inflorescences terminal This American tribe is here expanded to
or pseudo-lateral, bisexual, racemiform; pistillate include Garciadelia. Crotonogynopsis and Enri-
flowers proximal, staminate flowers distal; bracts quebeltrania, which were added by Webster
subtending solitary flowers. Staminate flowers (1994) and Radcliffe-Smith (2001), are excluded
subsessile; calyx dehiscing into 5 valvate lobes; and treated as unplaced core acalyphoid genera.
petals 5; disk segments 5; stamens 4–16, filaments
connate into a column; anthers in 1–3 superposed
series, extrorse; pollen grains oblate, 6–9-colpo- KEY TO THE GENERA OF ADELIEAE
rate, colpi short, inoperculate and emarginate, 1. Indumentum simple; stipules deciduous; stylodia lac-
sexine coarsely reticulate, sometimes heterobro- erate 166. Adelia
chate; pistillode 0. Pistillate flowers pedicellate; – Indumentum stellate or stellate-lepidote; stipules 0;
sepals and petals 5, narrow, entire, persistent in stylodia bifid to lacerate 2
144 G.L. Webster
2. Pistillate disk 0; stylodia aplanate, undivided, apically ments distinct, purplish, pubescent; anthers
incised 169. Garciadelia introrse, versatile, muticous; pollen grains oblate
– Pistillate disk conspicuous; stylodia usually not apla- spheroidal, 3-colpate, colpi operculate; sexine
nate, bifid or lacerate 3
3. Pistillate disk entire; stylodia bifid or subentire; fila-
tectate-perforate, microverrucate; pistillode min-
ments distinct; pollen 3-colporate; leaves triplinerved, ute or 0. Pistillate flowers pedicellate, mid-articu-
indumentum stellate 167. Lasiocroton late; sepals 5, aestivation open, persistent in fruit;
– Pistillate disk lobed; stylodia lacerate; filaments disk annular-pulvinate, tomentose; ovary 3(4)-
connate; pollen grains 4-colporate; leaves pinnately locular, pubescent; stylodia bifid or unlobed,
veined, indumentum stellate-lepidote tips laciniate. Fruits capsular; columella persis-
168. Leucocroton
tent, slender, apically trifid. Seeds roundish, testa
166. Adelia L. smooth.
Six spp., Greater Antilles and Bahamas.
Adelia L., Syst. Nat. ed. 10, 2: 1298 (1759, nom. cons.);
Lourteig & O’Donell, Lilloa 8: 285, fig. 3 (1942); Webster,
Ann. Missouri Bot. Gard. 54: 272, fig. 11 (1968); De-Nova, 168. Leucocroton Griseb.
Sosa & Steinmann, Syst. Bot. 32: 583–595 (2007), rev.
Leucocroton Griseb., Mem. Amer. Acad. Sci. 8: 160 (1860),
Dioecious trees or shrubs; indumentum simple; ottingen 9: 20 (1861); Borhidi, Acta Bot.
Abh. Ges. Wiss. G€
twigs often spinescent. Leaves often clustered on Acad. Hungar. 36: 13, figs. 1–16 (1990); Jestrow et al.,
short shoots, entire or sometimes crenate, beneath Taxon 59: 1801–1814 (2010).
domatiiferous; stipules minute, deciduous. Inflor- Dioecious (monoecious) shrubs; indumentum
escences axillary, glomerulate; bracts persistent. stellate or stellate-lepidote. Leaves pinnately
Staminate flowers pedicellate; sepals 4 or 5, val- veined or triplinerved, eglandular; stipules 0.
vate; petals 0; disk annular or 5-lobed, fleshy; Inflorescences axillary, pedunculate, staminate
stamens 6–30, filaments basally connate; anthers flowers in glomerules on spiciform thyrses, pistil-
versatile, muticous; pollen grains spheroidal, late flowers solitary or paired on shorter bracteate
angulaperturate, 3–4-colp(oroid)ate, colpi opercu- axes; bracts prominent. Staminate flowers subses-
late, operculum fusiform, sexine tectate-perforate, sile or pedicellate; calyx closed in bud, splitting
microverrucate; pistillode minute, trifid. Pistillate into 3–5 valvate segments; petals 0; disk annular
flowers long-pedicellate; sepals 5–7, narrow, per- or lobed; stamens (5–)7–15(28), filaments dis-
sistent and reflexed in fruit; petals 0; disk annu- tinct or basally connate; anthers introrse, muti-
lar, pubescent; ovary (2)3(4)-locular, pubescent; cous; pollen grains prolate-spheroidal, 3-colpate,
stylodia 3, multifid-lacerate. Fruits capsular; colu- colpi operculate, sexine tectate-perforate, micro-
mella persistent, apically 3-pronged. Seeds sphe- verrucate; pistillode pubescent. Pistillate flowers
roidal to oblate, testa smooth. pedicellate; sepals 5 or 6, persistent in fruit; disk
Nine spp., ranging from North America (N massive, annular or lobed, pubescent; ovary 2–3-
Texas) through Mexico and the Greater and Lesser locular, stellate-pubescent; ovules with inner inte-
Antilles to South America (Bolivia/Argentina). guments thick, outer thin; stylodia plicate, multi-
fid. Fruits capsular, columella slender, 3-pronged,
167. Lasiocroton Griseb. persistent. Seeds roundish, testa smooth.
Lasiocroton Griseb., Fl. Brit. W. Ind.: 46 (1859), Abh. About twenty-six spp., all endemic to Cuba
K€onigl. Ges. Wiss. G€ottingen 9: 20 (1861); Fawc. & where it grows exclusively in serpentine-rich
Rend., Fl. Jamaica 4: 293, fig. 96 (1920); Alain, Fl. Cuba areas. The genus is unique insofar that all
3: 87 (1953); Correll & Correll, Fl. Bah. Arch.: 825, fig. 342
spp. have the ability for hyperaccumulation of
nickel, having radiated along the fragmented
Dioecious shrubs; indumentum simple and stel- patches of serpentine soils across the island
late. Leaves triplinerved or palmately veined, (Jestrow et al. 2010).
entire; stipules 0. Inflorescences axillary, race-
mose; staminate flowers in glomerules, pistillate
flowers 1 per bract. Staminate flowers pedicellate; 169. Garciadelia Jestrow & Jiménez Rodr.
calyx closed in bud, splitting into 5 valvate seg- Garciadelia Jestrow & Jiménez Rodr., Taxon 59: 1809
ments; petals 0; disk annular; stamens 7–18, fila- (2010).
Euphorbiaceae 145
Dioecious treelets or shrubs; indumentum stel- 170. Bernardia Houston ex Mill.

late. Leaves cordate, eglandular; stipules 0. Inflor- Bernardia Houston ex Mill., Gard. Dict. abr. edn (1754);
escences axillary, the staminate subpaniculate, Fawc. & Rend., Fl. Jam. 4: 290, fig. 94 (1920); Lourteig &
the pistillate racemes. Staminate flowers 5-mer- O’Donell, Lilloa 8: 275, figs. 1, 2 (1942); Allem, Rev. Brasil.
ous, sepals triangular, valvate; petals 0; stamens Biol. 39: 529, figs. 1–9 (1979); Correll, Fl. Bahama Arch.
twice the number of sepals; anthers dorsifixed; 781, fig. 327 (1982); Liogier, Fl. Española 4: 85, t. 116–7
(1986); Webster, Fl. Venez. Guayana 5: 101, fig. 99 (1999).
disk weakly lobed; pistillate flowers with perianth Traganthus Klotzsch (1841).
as in males; disk 0; ovary 3-locular; stylodia 3,
aplanate, undivided but apically incised. Fruit a Monoecious or dioecious shrubs or subshrubs
3-lobed schizocarp, covered with stellate tri- (herbs); indumentum of simple or fasciculate
chomes. Seeds orbicular. hairs. Leaves often with basal embedded glands,
Four spp., endemic to Hispaniola. usually dentate; stipules small, persistent. Inflor-
escences axillary, [pistillate ones sometimes
13. TRIBE BERNARDIEAE G.L. Webster (1975). pseudo-terminal], spiciform or racemose; bracts
eglandular. Staminate flowers sessile or pedicel-
Monoecious or dioecious shrubs or herbs; indu- late; calyx splitting into 3 or 4 valvate lobes; petals
mentum simple or stellate; leaves alternate, entire, 0; disk of many interstaminal pubescent seg-
often with laminar glands; stipules deciduous; ments; stamens (2–)5–20(45), filaments dis-
inflorescences terminal to axillary, racemiform or tinct; anthers 4-locellate, introrse to latrorse,
spiciform; flowers pedicellate, the staminate sev- connective emarginate to enlarged; pollen grains
eral per bract, the pistillate solitary; staminate 3-lobed, 3-colporate, colpi marginate and inoper-
calyx closed in bud, splitting into 3–4 valvate seg- culate; endoaperture lalongate, sexine tectate-
ments; petals 0; disk intrastaminal, pulviniform, perforate; pistillode rudimentary or 0. Pistillate
dissected or obsolete; stamens 3–60, filaments dis- flowers sessile or subsessile; sepals 4–6, imbri-
tinct; anthers sometimes appearing 4-locellate; cate, entire, persistent in fruit; petals 0; disk
connective sometimes apiculate; pollen grains subannular or dissected; ovary 3-locular, pubescent;
spheroidal, 3-lobed, 3-colporate, colpi inopercu- ovules pachychalazal, inner integument thick,
late, sexine perforate-tectate; pistillode outer thin; stylodia bifid to flabellate. Fruits cap-
rudimentary or 0; pistillate sepals 4–6, imbricate; sular (indehiscent), 3-angled or -lobed; collu-
disk annular or dissected; ovary 3-locular, ovules mella slender, persistent. Seeds roundish,
with thick inner and thin outer integuments; sty- distinctly carinate, ecarunculate, testa smooth or
lodia bifid, the branches sometimes lacerate; fruit rugulose; endosperm copious; cotyledons longer
capsular; seeds roundish, carunculate or ecarun- and broader than receptacle. n ¼ 26.
culate, exotegmic, testa smooth. A genus of over 50 spp., mostly neotropical,
The circumscription of this tribe is still prob- one sp. in southwestern USA. Govaerts et al.
lematic: in the molecular analysis (Wurdack et al. (2000) list 68 species, some of which may be
2005) Bernardia and Adenophaedra are resolved synonyms. The genus is variable in both vegeta-
with Caryodendron in a sister position to Pluke- tive and floral characteristics; Pax & Hoffmann
netieae, whereas Discocleidion, differing, i.a., in a (1914) recognized 7 sections. Section Traganthus
vascularized outer integument, appears (without is distinctive in its herbaceous habit, but has the
support) in a clade with Ricinus and Speranskia. pollen and seed morphology of other sections; its
recognition as a genus would clearly make Ber-
KEY TO THE GENERA OF BERNARDIEAE nardia excessively paraphyletic.
1. Leaf blades stipellate, palmately veined; seeds caruncu-
late, testa rugose 172. Discocleidion 171. Adenophaedra (M€
ull. Arg.) M€
ull. Arg.
– Leaf blades not stipellate, pinnately veined; seeds ecar- Adenophaedra (M€ ull. Arg.) M€
ull. Arg., Fl. Brasil. 11(2):
unculate, testa smooth 2 385, t. 101 (1874); Pax & K. Hoffm., Pflanzenr. 147, VII:
2. Stylodia bifid or trifid; staminate disk of interstaminal 261 (1914); Webster & Huft, Ann. Missouri Bot. Gard. 75:
segments; seeds carinate; stamens (3)4–25; leaf blades 1099 (1988).
usually with abaxial embedded glands 170. Bernardia Bernardia sect. Adenophaedra M€ ull. Arg. (1865).
– Stylodia unlobed and dilated; staminate disk 0; seeds
not carinate; stamens 2(3); leaf blades lacking abaxial Dioecious trees or shrubs; indumentum simple.
glands 171. Adenophaedra Leaves dentate, abaxial laminar glands + or 0;
146 G.L. Webster
stipules deciduous. Inflorescences axillary or bifid, densely papillose. Fruit capsular; columella
the pistillate sometimes appearing terminal, slender, persistent. Seeds ovoid, carunculate, car-
spiciform, the staminate sometimes fasci- uncle deciduous; exotesta rugulose, purplish-
culate or paniculate-clustered; bracts eglandular. tinged; mesotesta vascularized.
Staminate flowers pedicellate; calyx splitting into Two spp, China, Ryukyu islands.
3 valvate lobes; petals and disk 0; stamens 2 (3);
anthers longer than filaments, introrse, connec- 14. TRIBE PLUKENETIEAE (Benth.) Hutch. (1969).
tive enlarged; pollen grains prolate spheroidal, 3-
lobed, 3-colporate, colpi marginate and inoper- Monoecious (dioecious), woody or herbaceous;
culate, endoaperture lalongate; sexine tectate- leaves alternate, unlobed to palmately lobed or
perforate; pistillode 0. Pistillate flowers pedicel- parted, pinnately to palmately veined, sometimes
late; sepals 6, biseriate, imbricate, persistent in glandular or stipellate at base; stipules usually
fruit; petals 0; disk 3-lobed; ovary 3-locular; persistent; inflorescences terminal or axillary,
ovules anatropous, inner integument very thick, usually bisexual, mostly racemoid or spiciform;
outer integument thin; stylodia contracted into bracts eglandular; staminate calyx closed in bud,
dilated sessile stigmas. Fruits capsular, 3-lobed; splitting into 3–5 valvate segments; petals 0; disk
columella apically dilated, 3-winged, persistent. + or 0; stamens (2)3–100, filaments usually dis-
Seeds roundish, ecarunculate, testa smooth. tinct; pollen grains mostly 3-colpate with irregu-
Three neotropical spp., occurring from Costa lar colpar margins, sometimes 3-porate or
Rica to Brazil. Although Adenophaedra has inaperturate but 3-colporate in Dalechampia,
highly reduced flowers and differs from Bernar- sexine intectate to tectate, microverrucate; pistil-
dia in a number of other characters, the similarity lode 0; pistillate sepals 3–6, imbricate; petals and
in pollen noted by Punt (1962) and Nowicke et al. disk 0; ovary 3- or 4-locular; stylodia unlobed,
(1999) supports its inclusion in the Bernardieae. usually connate into an elongated column, tips
smooth to lacerate; fruit capsular; columella per-
172. Discocleidion (M€
ull. Arg.) Pax & K. Hoffm. sistent; seeds ecarunculate, testa usually smooth
and dry.
Discocleidion Pax & K. Hoffm., Pflanzenr. 147, VII: 45, fig.
6 (1914); Webster, Ann. Missouri Bot. Gard. 81: 75 (1994);
Sixteen genera in three subtribes, both in the
Qiu Huaxing & Gilbert, Fl. China 11: 241 (2008). New World and the Old, but the majority of taxa
neotropical. Although the tribe appears to be
Dioecious trees or shrubs; indumentum simple. monophyletic (nine genera included in the
Leaves glandular-serrulate, 2–4-glandular abaxi- molecular analysis of Wurdack et al. 2005), paly-
ally above the stipellate base; stipules persistent. nologically it is the most diverse tribe in subfam-
Inflorescences mostly terminal, the staminate ily Acalyphoideae (Gillespie 1994). For the
paniculate, pistillate spiciform or racemoid position of Dalechampia within the tribe, see
(branched); staminate bracts lobed, persistent, under Subtribe Dalechampiinae.
eglandular. Staminate flowers pedicellate; calyx
splitting into 3–5 valvate segments; petals 0; disk KEY TO THE SUBTRIBES OF PLUKENETIEAE
segments interstaminal, on the convex receptacle;
stamens 35–60, filaments distinct, longer than 1. Inflorescence pseudanthial, bibracteate; pollen grains
prolate, coarsely reticulate, with pronounced costae
anthers; anthers apicifixed, supertrorse to latrorse, equatoriales
muticous, thecae discrete and pendulous; pollen 14c. Dalechampiinae
grains spheroidal, 3-colporate, colpi with broad – Inflorescence racemoid or spiciform; pollen grains sub-
margins, inoperculate, sexine tectate-perforate, globose, 3-colpate, not coarsely reticulate, lacking cos-
sometimes with prominent excrescences; pistil- tae equatoriales 2
lode 0. Pistillate flowers pedicellate; sepals 5, 2. Stinging hairs 0; fruit winged, crested, carinate, or
indehiscent; pollen grains 3-colporate
entire, imbricate, persistent in fruit; petals 0; 14a. Plukenetiinae
disk annular, crenulatae, pubescent; ovary 3(4)- – Stinging hair +; fruit usually appendaged; pollen grains
locular, sericeous; ovules with inner integument 3-colporate or -porate (inaperturate), sexine tectate-
thick, outer thinner and vascularized; stylodia perforate to rugulose 14b. Tragiinae
Euphorbiaceae 147
14a. SUBTRIBE PLUKENETIINAE Benth. (1880). very irregular margins; sexine tectate-rugulose,
microverrucate; pistillode 0. Pistillate flowers
Trees, shrubs, lianas, or herbaceous vines; indu- pedicellate; sepals usually 4, imbricate, narrowly
mentum simple, sometimes glandular, never urti- lanceolate, entire, persistent in fruit; petals and
cant. Leaves alternate, often with embedded disk 0; ovary 3-locular, glabrous, 3-horned; sty-
laminar glands, sometimes stipellate; inflores- lodia connate into a clavate hollow column with
cences axillary or terminal; stamens 4–50, fila- lobulate rim. Fruits capsular, cocci horned; colu-
ments distinct; pollen grains tectate-perforate to mella massive, 3-angled, persistent. Seeds turbi-
semitectate-reticulate; ovary 3–4-locular, often nate-truncate, testa smooth.
appendaged; stylodia partly to entirely connate A single sp., A. cornutus Benth., Amazonian
into a common style; fruit capsular (indehiscent). Brazil and Venezuela.
Five genera, all neotropical except for a few
species of Plukenetia in Africa.
174. Haematostemon (M€
ull. Arg.) Pax &
KEY TO THE GENERA OF PLUKENETIINAE K. Hoffm. Fig. 33
1. Ovary 3-locular; pistillate sepals 5 or 6; trees, shrubs, or Haematostemon (M€ ull. Arg.) Pax & K. Hoffm., Pflanzenr.
vines 2 147, IX: 31, fig. 11 C, D (1919); Sandwith, Kew Bull. 1950:
– Ovary 4-locular; pistillate sepals 4; twining vines or 133 (1951); Jablonski, Mem. N. Y. Bot. Gard. 17: 143
lianas 177. Plukenetia (1967); L.J. Gillespie & Armbruster, Smiths. Contr. Bot.
2. Stamens 4; staminate disk segments 4 or filament bases 86: 28, fig. 9 (1997); Radcl.-Sm., Gen. Euphorb.: 241, fig. 31
forming a pseudo-disk; leaves pinnately veined; trees (2001).
or shrubs 3 Astrococcus sect. Haematostemon M€ ull. Arg. (1865).
– Stamens 10 or more; staminate disk 0 or of numerous Monoecious trees or shrubs; indumentum sim-
minute segments 4 ple. Leaves crenulate or serrulate, eglandular or
3. Disk exstrastaminal; ovary horned 173. Astrococcus
– Disk 0, the filament bases forming a pseudo-disk; ovary
obscurely glandular adaxially; stipules minute.
verrucate 174. Haematostemon Inflorescences axillary, bisexual, spiciform;
4. Trees or shrubs; flowers in axillary glomerules; leaves lower nodes with solitary pistillate flowers or
pinnately veined; stamens c. 20; style urceolate bisexual cymules, distal with glomerules of stami-
175. Angostyles nate flowers, terminal flower pistillate; bracts
– Twining vines; flowers in axillary racemes; leaves eglandular. Staminate flowers pedicellate; calyx
palmately veined; stamens 10; style apically 3-lobed
176. Romanoa
splitting into 4 valvate segments; petals 0;
stamens 4, filaments basally dilated and connate
173. Astrococcus Benth. into a pseudo-disk; anthers basifixed, introrse,
muticous; pollen grains suboblate, 3-colpate,
Astrococcus Benth., Hook. J. Bot. Kew Gard. Misc. 6: 327 colpi sometimes occluded by sexine, tectum
(1854); M€ull. Arg. in DC., Prodr. 15(2): 766 (1866), Fl. foveolate-rugulose, microverrucate; pistillode
Brasil. 11(2): 330, t. 49 (1874); Benth., Gen. Pl. 3: 326
(1880); L.J. Gillespie, Fl. Venez. Guayana 5: 100, fig. 98 0 or incorporated into pseudo-disk. Pistillate
(1999). flowers pedicellate; sepals 4–6, imbricate, entire;
Monoecious trees or shrubs, branches rib-angled disk 0; ovary 3-locular, tuberculate; stylodia con-
distally; indumentum simple. Leaves subentire nate into a cup-shaped hollow column with mul-
(minutely glandular-serrulate), eglandular; sti- tidentate rim. Fruits capsular.
pules minute, deciduous. Inflorescences axillary, Two spp. in northern South America: Vene-
spiciform, bisexual; pistillate flowers solitary zuela (Guayana Highland) and Guyana.
at the 4 or 5 proximal nodes, staminate in 1- or
2-flowered glomerules at distal nodes, terminal 175. Angostylis Benth.
flower pistillate; bracts entire, eglandular, Angostylis Benth., Hook. J. Bot. Kew Gard. Misc. 6: 328
recurved. Staminate flowers short-pedicellate; (1854, as Angostyles); M€ ull. Arg. in DC., Prodr. 15(2): 767
calyx closed in bud, splitting into 4 valvate seg- (1866), Fl. Brasil. 11(2): 331, t. 50 (1874); Radcl.-Sm., Gen.
Euphorb.: 244 (2001).
ments; petals 0; disk extrastaminal, 4-lobed, lobes
enclosing anthers; stamens 4, filaments basally Monoecious trees or shrubs; indumentum sim-
confluent; anthers basifixed, introrse, muticous; ple, non-urticant. Leaves subsessile, clustered at
pollen grains oblate spheroidal, 3-colpate with branch-tips, denticulate, pluriglandular; stipules
148 G.L. Webster
176. Romanoa Trevis.

Romanoa Trevis., Saggio Algh. Coccot.: 99 (1848, nom.
cons.); Pax & K. Hoffm., Pflanzenr. 147, IX: 28, fig. 9
(1919, under Anabaenella); Radcl.-Sm., Kew Bull. 34:
589 (1980); Radcl.-Sm., Gen. Euphorb.: 244 (2001).
Anabaena A. Juss. (1824, nom. illeg.).
Anabaenella Pax & K. Hoffm. (1919).
Monoecious vines; indumentum simple. Leaves
palmately veined, biglandular on margins at
base; stipules deciduous. Inflorescences axillary,
usually bisexual with 1 or 2 basal pistillate flowers
and several glomerules of staminate flowers;
bracts entire, eglandular. Staminate flowers sus-
bsessile or short-pedicellate, pedicel medially
articulate; calyx closed in bud, splitting into 5
segments; petals 0; receptacle glandular;
stamens 10, filaments distinct; anthers extrorse
or introrse, 2-celled but opening into 4 valves;
pollen grains oblate spheroidal, angulaperturate,
3-colpate, colpi broad with uneven margins, sex-
ine tectate-perforate, microverrucate; pistillode
subulate. Pistillate flowers pedicellate; sepals 5
or 6, narrow, entire, persistent in fruit but not
accrescent; petals and disk 0; ovary 3-locular,
glabrous; stylodia connate into a clavate, apically
3-lobed column. Fruits capsular, valves not veiny;
columella persistent, slender, 3-pronged. Seeds
carinate-angled, testa smooth.
A single sp., R. tamnoides (A. Juss.) Radcl.-
Sm., eastern and southern Brazil, Paraguay.
Although Romanoa appears similar to Plukenetia
Fig. 33. Euphorbiaceae-Acalyphoideae. Haematostemon
in many respects, the pollen is distinctive (Gille-
guianensis. A Flowering branch. B Base of leaf blade, spie 1994).
adaxial surface. C Pistillate flower, with verrucate ovary
lobes and massive truncate style. D Staminate flower.
(Gillespie & Armbruster 1997; drawn by Cathy Pasquale) 177. Plukenetia L.
Plukenetia L., Sp. Pl.: 1192 (1753), Gen. Pl. ed. 5: 438
subulate. Inflorescences axillary, glomerular, the (1754); M€ ull. Arg. in DC., Prodr. 15(2): 768 (1866), Fl.
staminate 2 or 3 per cymule, the pistillate solitary. Brasil. 11(2): 232 (1874); Pax & K. Hoffm., Pflanzenr.
Staminate flowers pedicellate; calyx closed in 147. IX: 12 (1919); Webster, Ann. Missouri Bot. Gard.
bud, splitting into 3 (4) valvate segments; petals 54: 293, fig. 15 (1968), ibid. 81: 93 (1994); Huft, Ann.
0; stamens c. 20, filaments distinct; anthers dor- Missouri Bot. Gard. 75: 1105 (1989); Gillespie, Syst. Bot.
18: 575 (1993), Fl. Venez. Guayana 5: 207, figs. 196, 197
sifixed, extrorse, muticous; pollen grains subo- (1999); L.J. Gillespie & Armbruster, Smiths. Contr. Bot.
blate, angulaperturate, 3-brevicolpate, colpar 86: 30, figs. 10–12 (1997).
margins irregular, tectum foveolate-rugulose, Vigia Vell. (1832).
microverrucate; pistillode 0. Pistillate flowers Apodandra Pax & K. Hoffm. (1919).
pedicellate; sepals 5, imbricate, entire, adaxially Eleutherostigma Pax & K. Hoffm. (1919).
Elaeophora Ducke (1925).
glandular; petals and disk 0; ovary 3-locular,
muricate; stylodia connate into a hollow infundi- Monoecious (dioecious) lianas or twining vines;
biliform column with 6-dentate rim. Fruits cap- indumentum simple. Leaves pinnately or palmately
sular, muricate. Seeds subglobose, testa smooth. veined, subentire to serrulate, adaxially with
A single sp., A. longifolia Benth., Brazilian embedded laminar basal glands and often stipellate
Amazon. at base; stipules deciduous. Inflorescences axillary
Euphorbiaceae 149
or terminal on short shoots, bisexual, racemose; Ten genera with c. 150 species. Tragia is
pistillate flowers at proximal nodes, staminate glo- pantropical, Acidoton and Platygyna are neotro-
merules distally; bracts minute, eglandular. Stami- pical, and the other genera paleotropical.
nate flowers pedicellate; calyx splitting into 4 or 5
valvate segments; petals 0; disk interstaminal or KEY TO THE GENERA OF TRAGIINAE
extrastaminal, sometimes 0; stamens 15–40, fila-
ments distinct, sometimes shorter than anthers, 1. Stylodia usually slender; sepals not adaxially inflexed;
anthers extrorse or introrse; stamens 2–many; pollen
inserted on convex or conical receptacle; anthers grains tectate-perforate 2
extrorse or introrse, muticous, thecae divergent; – Stylodia massive; staminate sepals uually inflexed to
pollen grains suboblate, angulaperturate, 3-colpate, form a pseudo-disk; stamens 3 or 4, anthers introrse 7
colpi broad with uneven margins, sexine tectate- 2. Anthers 2, subsessile; staminate calyx flat with
perforate to semitectate-reticulate; pistillode 0. Pis- lobes reflexed; stylodia distally recurved and proxi-
mally connate into a broad conic-cylindrical base
tillate flowers pedicellate; sepals 4 (5), imbricate, 183. Pachystylidium
entire, persistent in fruit but not accrescent; – Anthers 3 or more, rarely 2, but then not subsessile;
petals and disk 0; ovary 4-locular, angled or lobed, staminate calyx concave; stylodia distinct to partly
glabrous; ovules anatropous, inner integument connate into a columnar style 3
moderately thick, outer thin; stylodia connate into 3. Anther connective with tuft of stinging hairs; stamens
a column, the stigmatic tips entire to bilobed. Fruits 25–60; dioecious shrubs 182. Acidoton
capsular or baccate; columella persistent, apically – Anther connective lacking tuft of stinging hairs;
stamens (2)3–50; monoecious subshrubs, herbs, or
dilated. Seeds lenticular to subglobose, testa vines 4
smooth. 4. Stylodia thickened, apically bifid; inflorescences uni-
In the concept of Gillespie (1993), Plukenetia sexual, contracted [< 1 cm]; pollen grains inaperturate
includes 13 spp. scattered from tropical America 181. Platygyna
to Africa, Madagascar, and southeast Asia. The – Stylodia slender, entire; inflorescences bisexual, elon-
genus is very heterogeneous in vegetative, floral, gated [> 1 cm]; pollen grains 3-colporate 5
5. Stamens 8–40; inflorescences bifurcate, staminate and
and palynological characters, and it is not pistillate flowers on separate axes; pollen sexine
surprising that Pax and Hoffmann (1919) recog- coarsely tectate-perforate 178. Bia
nized a number of segregate genera. Gillespie has – Stamens 2–4 (or else pistillate sepals pinnatifid); inflor-
in effect included all Plukenetiinae with 4-locular escences not bifurcate, usually bisexual; pollen sexine
ovaries within Plukenetia; although Eleutheros- finely tectate-perforate-rugulose 6
6. Stamens 2–4, anthers not linear, often shorter than
tigma was maintained as distinct by Webster filaments; pistillate calyx entire or pinnatifid
(1994) and Radcliffe-Smith (2001), it was merged 179. Tragia
with Plukenetia by Gillespie (1993, 1994). The – Stamens 30–50, anthers linear and much longer than
reduction of Eleutherostigma and Vigia, accepted filaments; pistillate calyx pinnatifid 180. Ctenomeria
as generically distinct by Webster (1994), is fol- 7. Pistillate sepals entire or dentate; pistillode 0 8
lowed here, but it still appears possible that – Pistillate sepals pinnatifid; pistillode present in stami-
further analysis will result in redefinition of nate flower; stylodia connate into a conical or infun-
dibular column 187. Tragiella
the generic boundaries of Plukenetia and other 8. Pistillate sepals entire; anther connective enlarged,
Plukenetiinae. sometimes caudate 9
– Pistillate sepals toothed; inflorescences terminal and
14b. SUBTRIBE TRAGIINAE G.L. Webster (1975). leaf-opposed; leaves subglabrous 185. Sphaerostylis
9. Stylodia distinct or nearly so, adaxially papillose;
Monoecious shrubs or herbs, often scandent; inflorescences terminal and leaf-opposed; leaves with
indumentum of simple and urticant hairs; leaves stinging hairs 184. Cnesmone
simple (lobed), without embedded laminar glands, – Stylodia connate into a clavate or globose column, not
not stipellate; stipules persistent or deciduous; adaxially papillose; inflorescences axillary; leaves sub-
glabrous, not urticant 186. Megistostigma
inflorescences axillary or terminal and opposite
leaves; stamens (2) 3–50, filaments distinct or con-
nate; pollen grains 3-colpate, 3-porate, or inaper- 178. Bia Klotzsch
turate, sexine mostly tectate-perforate and Bia Klotzsch, Arch. Naturgesch. 7(1): 189 (1841).
microverrucate; ovary 3-locular; stylodia nearly Zuckertia Baillon (1858).
distinct to completely connate; fruit capsular. Tragia sect. Bia (Klotzsch) M€
ull. Arg. (1865).
150 G.L. Webster
Monoecious vines; indumentum simple (unicellu- only in its coarsely tectate-perforate exine, but
lar) and urticant. Leaves palmately veined and also in other traits such as the larger stamen
usually cordate at base, dentate, eglandular; sti- number (see key and description above). With
pules persistent. Inflorescences axillary, bisexual, the recognition of Bia and Ctenomeria as distinct
eglandular, bifurcate, pistillate branch spiciform, genera, Tragia becomes less heterogeneous and
with 5–20 subsessile flowers, staminate branch easier to define, although much more study needs
racemoid, with c. 20–30 glomerules of 1–3 pedicel- to be given to identifying monophyletic groups.
late flowers; bracts entire, eglandular. Staminate
flowers pedicellate; calyx splitting into 3–6 valvate 179. Tragia (Plumier) L. Fig. 34
segments; petals 0; disk of interstaminal segments Tragia (Plumier) L., Sp. Pl.: 980 (1753), Gen. Pl. ed. 5: 421
or 0; stamens (5–)8–40, filaments distinct; anthers (1754); M€ ull. Arg. in DC., Prodr. 15(2): 927 (1866); Pax &
introrse, muticous, sagittate; pollen grains subo- Hoffm., Pflanzenr. 147, IX: 32, figs. 12–22 (1919); Fawc. &
blate or spheroidal, 3-colpate or inaperturate, tec- Rendle, Fl. Jam. 4: 304, fig. 100 (1920); Lourteig & O’Do-
tate-perforate to semitectate-reticulate; pistillode nell, Lilloa 6: 347 (1941); Miller & Webster, Rhodora 69:
241, figs. 17–29 (1967); Webster, J. Arnold Arb. 48: 376
0. Pistillate flowers subsessile; sepals 6, imbricate, (1967), Ann. Missouri Bot. Gard. 54: 291, fig. 14 (1968);
entire, persistent in fruit; disk 0; ovary 3-locular, Leandri, Adansonia II, 11: 437 (1971); Radcl.-Sm., Fl. E.
pubescent with simple and urticant hairs; stylodia Trop. Afr. Euphorb. 1: 291, figs. 57–59 (1987); Radcl.-Sm.,
unlobed, adaxially papillose with complex papillae, Fl. Zambesiaca 9(4): 216, figs. 46–48 (1996), Gen.
basally connate. Fruits capsular; columella persis- Euphorb.: 253 (2001).
tent. Seeds spheroidal, testa smooth, without dis- Monoecious (dioecious) shrubs or herbs, often
tinct microsculpturing. twining; indumentum simple and urticant, some-
Five neotropical spp. The genus Bia is here times glandular. Leaves usually petiolate, mostly
reinstated from the synonymy of Tragia and con- palmately veined or triplinerved, entire to dentate
strued to include two sections1. The work of Now- (lobed or divided), often cordate; stipules usually
icke and Takahashi shows great palynological persistent. Inflorescences usually terminal or
heterogeneity in Tragia, and Bia appears mor- opposite leaves, bisexual, spiciform or racemoid,
phologically distinct from Tragia (s. str.) not with 1–3 basal pistillate flowers and distal stami-
nate flowers; bracts subfoliose, eglandular, mostly
uniflorous. Staminate flowers pedicellate; pedicel
1
A. Bia sect. Bia. Type: Bia sellowiana Klotzsch ex Baillon articulated just above base to mid-length; calyx
(¼ B. alienata Didrichsen).
Staminate flowers with dissected disk; stamens introrse, closed in bud, splitting into 3–6 valvate segments;
8–20, anthers muticous; style column slender. Four petals and disk 0; stamens (2)3–4(40), filaments
spp.: distinct or basally connate; anthers extrorse
1. B. alienata Didrichsen (1857) (¼ B. sellowiana or introrse, muticous; pollen grains suboblate
Klotzsch ex Baill. 1858). to spheroidal, 3-colpate or inaperturate, colpi
2. Bia fallax (M€ ull. Arg.) G.L. Webster, comb. nov. narrow to broad, sometimes circular; sexine
Basionym: Tragia fallax M€ ull. Arg., Linnaea 34: 179 reticulate to tectate-rugulose, sometimes micro-
(1865). Type: Peru, Pavon (holotype G-Less., photo- verrucate; pistillode laminar to globose. Pistillate
graph F7155). – Tragia japurensis M€ ull. Arg.
3. Bia fendleri (M€ ull. Arg.) G.L. Webster, comb. nov. flowers pedicellate; sepals mostly 6–8, imbricate,
Basionym: Tragia fendleri M€ ull. Arg., Linnaea 34: 179 entire to pinnatifid, persistent and sometimes
(1865). Type: Venezuela, Biscaina, 3,000 feet, Fendler accrescent in fruit; petals and disk 0; ovary
1208 (holotype G; photograph F7156). 3-locular, glabrous or urticant-pubescent; ovules
4. Bia lessertiana Baillon (1858) [lesseriana]. – Tragia pachychalazal, inner integument thick, outer
lessertiana (Baill.) M€
ull. Arg.
B. Bia sect. Zuckertia (Baill.) M€ull. Arg., Linnaea 34: 178 thin; stylodia unlobed, smooth or papillose,
(1865) (as Tragia sect. Zuckertia). basally connate. Fruits capsular; columella persis-
Zuckertia Baill., Étude Gén. Euphorb.: 495 (1858). tent, apically 3-fid. Seeds globose, testa smooth.
Type: Zuckertia cordata Baill. About 170 spp., most in America and Africa
1. Bia cordata (Baill.) G.L. Webster, comb. nov. Basio- but extending to Asia and Australia (Govaerts
nym: Zuckertia cordata Baill, Étude Gén. Euphorb.: et al. 2000). The genus is quite variable in all
496, t. 10, figs. 10–13 (1858). Type: Mexico, Tabasco:
Teapa, Linden (holotype P).
Euphorbiaceae 151
above the base; calyx closed in bud, splitting

into 5 valvate segments; petals and disk 0; recep-
tacle convex; stamens 30–60, filaments distinct;
anthers longer than filaments, apiculate; pollen
grains oblate spheroidal, 3-aperturate, apertures
elliptic, occluded by thin exine layer, sexine
tectate-perforate, microverrucate; pistillode 0.
Pistillate flowers pedicellate; sepals 6, pinnatifid,
imbricate, persistent and accrescent in fruit;
ovary 3-locular, pubescent; stylodia unlobed,
densely papillate-laciniate. Fruits capsular. Seeds
globose, testa smooth.
Two spp., southern Africa. Although treated
as a synonym of Tragia by most recent authors,
including Webster (1994) and Radcliffe-Smith
(2001), the palynological studies of Gillespie
(1994) support the concepts of Prain and Dyer
in recognizing Ctenomeria as generically distinct.
181. Platygyna P. Mercier

Fig. 34. Euphorbiaceae-Acalyphoideae. Tragia lesserti- Platygyna P. Mercier, Bull. Bot. 1: 168 (1830); M€ull. Arg.
ana. A Part of stem with leaves and staminate flowers. in DC., Prodr. 15(2): 913 (1866, as Platygyne); Pax & K.
B Part of staminate inflorescence with detail of bract on Hoffm., Pflanzenr. 147, IX: 26 (1919); Alain, Fl. Cuba 3:
left. C Staminate flower. D Part of pistillate inflorescence. 100, fig. 28 (1953), Mem. N. Y. Bot. Gard. 21: 132 (1971);
E Lateral view and medial section of pistillate flower. Borhidi, Ann. Hist.-Nat. Mus. Nat. Hung. 64: 89 (1972);
F Part of infructescence. G Capsule. (Reproduced with Radcl.-Sm., Gen. Euphorb.: 255, fig. 33 (2001).
kind permission of the artist Bobbi Angell) Acanthocaulon Klotzsch (1850).
Monoecious woody vines; indumentum simple
characters and has presented a continuing chal- and urticant. Leaves dentate, eglandular; stipules
lenge to systematists. Pax and Hoffmann (1919) persistent. Inflorescences terminal, axillary or
recognized 9 sections, and 9 subsections in the leaf-opposed, glomerular or contracted-spiciform
African sect. Tagira; Gillespie (1994) recognized (< 1 cm long). Staminate flowers pedicellate;
5 sections for the Americas. Pollen evidence calyx closed in bud, splitting into 4 or 5 valvate
presented by Gillespie (1994) indicates great segments; petals 0; disk pulviniform, strigose-his-
heterogeneity within the genus, and suggests pid; stamens 5 (4–14), filaments distinct; anthers
that Tragia may be an unnatural taxon. extrorse, muticous; pollen grains spheroidal, ina-
perturate, sexine tectate-perforate, sometimes
180. Ctenomeria Harv. vermiculate, not verrucate; pistillode 0. Pistillate
Ctenomeria Harv., Hook. London J. Bot. 1: 29 (1842); flowers subsessile or pedicellate; sepals 6 (5–9),
Baill., Étude Gén. Euphorb.: 494 (1858); Prain, Fl. Capen- imbricate, entire, persistent in fruit; petals and
sis 5(2): 500 (1920); Dyer, Gen. S. Afr. Fl. Pl. 1: 318 (1975). disk 0; ovary 3-locular, tomentose; ovules pachy-
Tragia sect. Ctenomeria (Harv.) Benth. (1880). caulous, inner integument thick, outer integu-
Monoecious twining herbs; indumentum simple ment thin; stylodia 3 (4), bifid or emarginate,
and scantily urticant. Leaves palmately veined, contiguous in a thick, infundibuliform stylar col-
sometimes palmately lobed or subpeltate, den- umn. Fruits capsular; columella 3-pronged, per-
tate, eglandular; stipules reflexed, persistent. sistent. Seeds globose, testa smooth.
Inflorescences terminal or leaf-opposed, mostly According to the revision of Borhidi (1972)
bisexual, with 1 or 2 proximal pistillate flowers seven spp., endemic to Cuba. Alain (1971)
and distal glomerules of staminate flowers; bracts reduced Platygyna to a synonym of Tragia, but
1-flowered, eglandular. Staminate flowers pedi- the pollen evidence of Gillespie (1994) does not
cellate, pedicel articulate and bracteate at or support this.
152 G.L. Webster
182. Acidoton Sw. 183. Pachystylidium Pax & K. Hoffm.

Acidoton Sw., Prodr. 6: 83 (1788; nom. cons.), Fl. Ind.. Pachystylidium Pax & K. Hoffm., Pflanzenr. 147, IX: 108
Occid. 2: 952, t. 18 (1800); M€ull. Arg. in DC., Prodr. 15(2): (1919); Backer & Bakh., Fl. Java 1: 491 (1963); Airy Shaw,
914 (1866); Urb., Symb. Ant. 7: 513 (1913); Pax & Kew Bull. 23: 115 (1969), 26: 310 (1971); Webster, Ann.
K. Hoffm., Pflanzenr. 147, IX: 24, fig. 8 (1919); Fawc. & Missouri Bot. Gard. 81: 95 (1994); Radcl.-Sm., Gen.
Rendle, Fl. Jam. 4: 303, fig. 99 (1920); Webster, Ann. Euphorb.: 260, fig. 36 (2001).
Missouri Bot. Gard. 54: 191 (1967), 289, fig. 13 (1968);
Liogier, Fl. Española 4: 71 (1986); Webster, Ann. Missouri Monoecious vines; indumentum simple and urti-
Bot. Gard. 81: 95 (1994); Burger & Huft, Fieldiana Bot. 36: cant. Leaves cordate, palmately veined, dentate,
56 (1995); Govaerts et al., World Checkl. Bibl. Euporb. 1: eglandular; stipules persistent. Inflorescences ter-
111 (2000); Radcl.-Sm., Gen. Euphorb.: 257, fig. 34 (2001); minal or leaf-opposed, pedunculate, bisexual, with
Webster, Fl. Nicaragua 1: 852 (2001).
Gitara Pax & Hoffm. (1924). proximal pistillate flowers and distal glomerules of
staminate flowers; bracts concave, ovate, eglandu-
Monoecious or dioecious shrubs, sometimes lar. Staminate flowers pedicellate; calyx closed in
arborescent; indumentum simple and urticant. bud, splitting into 4 or 5 valvate segments; petals 0;
Leaves petiolate or subsessile, entire or dentate, disk or pseudo-disk with 2 cavities; stamens 2,
eglandular; stipules aristate, persistent. Inflores- filaments distinct; anthers extrorse, longer than
cences axillary, glomerular or racemoid; unisex- filaments, muticous; pollen grains oblate spheroi-
ual, the pistillate much longer than the staminate; dal, 3-porate, apertures ill-defined with sexinous
bracts eglandular, persistent, subtending solitary islands; sexine tectate-punctate, microverrucate;
flowers. Staminate flowers pedicellate; calyx pistillode 0. Pistillate flowers pedicellate; sepals 6,
closed in bud, splitting into 3–5 valvate segments; narrow, entire, imbricate; ovary 3-locular, pubes-
petals 0; disk 0 or obscure; stamens 20–60, fila- cent; stylodia distally recurved, proximally con-
ments distinct; anthers extorse, much shorter nate into a broad conic-cylindrical base. Fruits
than filaments, connective with apical tuft of urti- capsular; columella persistent, 3-pronged. Seeds
cant hairs; pollen grains spheroidal or subprolate, globose, testa smooth.
3-colpate or inaperturate, colpi with uneven mar- A single sp., P. hirsutum (Blume) Pax & K.
gins and sexinous islands, sexine tectate-perfo- Hoffm., widespread from India east as far as
rate and microverrucate or rugulose and smooth; Sulawesi. Pachystylidium is similar in habit to
pistillode 0. Pistillate flowers pedicellate; sepals 5 Cnesmone, and a close relationship is supported
or 6, imbricate, entire, persistent in fruit; petals by Gillespie’s palynological evidence.
and disk 0; ovary 3-locular, urticant-pubescent;
ovules pachycaulous, inner integument moder- 184. Cnesmone Blume
ately thick, outer integument thin; stylodia dis- Cnesmone Blume, Bijdr.: 630 (1826, as Cnesmosa); Croi-
tally recurved, papillose, and basally connate into zat, J. Arnold Arb. 22: 427 (1941); Airy Shaw, Kew Bull. 26:
acommon style. Fruits capsular, covered with 240 (1972); Balakr. & Nair, Gardener’s Bull. Singapore 31:
urticant hairs; columella persistent, 3-pronged 49 (1978); Radcl.-Sm., Gen. Euphorb.: 250 (2001).
apically. Seeds spheroidal, testa smooth. Cenesmon Gagnep. (1924).
Six spp., 5 in the Greater Antilles, 1 on main- Monoecious shrubs or lianas; indumentum sim-
land (Nicaragua to Venezuela). Radcliffe-Smith ple and urticant. Leaves pinnately or palmately
(2001) has accepted the segregate genus Gitara veined, often cordate, entire or serrate, eglandu-
as distinct, based largely on the palynological lar; stipules entire, reflexed, subpersistent. Inflor-
evidence of Gillespie (1994). It is true that the escences terminal or pseudo-axillary leaf-
coarsely dentate leaves and colpate pollen of the opposed, spiciform, bisexual, proximal nodes
mainland Acidoton (Gitara) nicaraguensis con- with solitary pistillate flowers, distal with glomer-
siderably differ from the entire leaves and inaper- ules of staminate flowers; bracts persistent,
turate pollen of the West Indian species. appearing trifid due to prominent bracteoles,
However, the inflorescences, flowers, and fruits eglandular. Staminate flowers pedicellate, pedicel
are very similar overall. It seems preferable, articulate at base; calyx closed in bud, splitting
therefore, in accordance with the opinion of into 3 valvate segments; petals and disk 0;
Govaerts et al. (2000), to maintain Acidoton in stamens 3, filaments basally connate; anthers
the broader sense, but retaining Gitara at the basifixed, introrse, connective enlarged and
sectional level. apiculate, thecae discrete; pollen grains
Euphorbiaceae 153
spheroidal, weakly 3-colpate, colpi broad and Monoecious shrubs or lianas; indumentum sim-
covered with sexinous granules; sexine tectate- ple and urticant. Leaves palmately veined, often
perforate and microverrucate; pistillode 0. Pistil- peltate or cordate; stipules persistent or decidu-
late flowers pedicellate; sepals 3–6, imbricate, ous. Inflorescences axillary, racemoid, unisexual
entire, foliose, persistent in fruit; petals and or bisexual, bracts eglandular, persistent, with
disk 0; ovary 3-locular, not angled, pubescent prominent bracteoles. Staminate flowers subses-
with simple and urticant hairs; ovules with very sile or pedicellate; calyx closed in bud, splitting
thick integuments; stylodia distally spreading, distally into 3 valvate lobes, proximally connate
unlobed, papillate, basally connate. Fruits capsu- into a cup; petals 0; receptacular disk 0, pseudo-
lar, endocarp thick and woody; columella persis- disk at rim of calycine cup; stamens 3, filaments
tent. Seeds globose, testa smooth. distinct; anthers muticous, introrse, ovate,
Eleven spp., Assam and Andaman Is. to China obtuse, connective enlarged, apiculate; pollen
and Malesia. grains spheroidal, irregularly aperturate or ina-
perturate; sexine tectate-perforate, fissured,
185. Sphaerostylis Baill. microverrucate; pistillode 0. Pistillate flowers
Sphaerostylis Baill., Étude Gén. Euphorb.: 466 (1858), in pedicellate; sepals 3–5, imbricate, lanceolate,
Grandidier, Hist. Phys. Nat. Madag. 4 (29): t. 4 (1891); entire, persistent in fruit; petals and disk 0;
Leandri, Bull. Soc. Bot. France 85: 527 (1939); Croizat, ovary 3-locular, pubescent; stylar column globose
J. Arnold Arb. 22: 430 (1941); Radcl.-Sm., Gen. Euphorb.: or clavate, broadly 3-lobed distally. Fruits capsu-
252 (2001); Schatz, Gen. Tree Fl. of Madagascar: 165 (2001). lar; columella persistent, 3-pronged. Seeds glo-
Monoecious shrubs to vines; indumentum simple bose, exotesta fleshy, endotesta hard, smooth.
and urticant, scanty. Leaves 3-palmatinerved at Five spp., distributed from Burma to Malesia.
the base and penninerved above, cordate or pel-
tate at base, entire or dentate; stipules entire, 187. Tragiella Pax & K. Hoffm.
auriculiform, deciduous. Inflorescences terminal
or leaf-opposed, unisexual or bisexual, racemoid; Tragiella Pax & K. Hoffm., Pflanzenr. 147, IX: 104, fig. 24
(1919); Radcl.-Sm., Kew Bull. 35: 777 (1981), Fl. E. Trop.
bracts uniflorous, eglandular. Staminate flowers Afr. Euphorb. 1: 318, fig. 60 (1987), Fl. Zambesiaca 9(4):
pedicellate, pedicel articulate at base; calyx closed 212, t. 45 (1996), Gen. Euphorb.: 252 (2001); Webster,
in bud, splitting into 3 valvate segments; calyx Ann. Missouri Bot. Gard. 81: 94 (1994).
segments inflexed into a pseudo-disk; petals and
Monoecious herbs to twining vines; indumentum
disk 0; stamens 2 or 3, filaments distinct, much
of simple and urticant hairs. Leaves palmately
shorter than anthers, connate; anthers introrse,
veined, denticulate, eglandular; stipules persis-
connective enlarged and apiculate, 2-celled,
tent. Inflorescences terminal or leaf-opposed,
locules divergent, dehiscing longitudinally/hori-
racemose, bisexual, with 1 or 2 basal pistillate
zontally; pollen grains oblate spheroidal, 3-col-
flowers, distal nodes with glomerules of staminate
pate, exine tectate-perforate, microverrucate;
flowers; bracts 1-flowered. Staminate flowers ped-
pistillode 0. Pistillate flowers subsessile; sepals 5
icellate; calyx closed in bud, distally splitting into
or 6, valvate or slightly imbricate, dentate; petals
3 valvate lobes introrsely projecting at top of
and disk 0; ovary 3-locular, pubescent; stylodia
calyx tube; petals and disk 0; stamens 3 (4), fila-
connate into a globose column, tips emarginate.
ments distinct or basally connate; anthers dorsi-
Fruit capsular. Seeds globose, testa smooth.
fixed, introrse, muticous, connective enlarged,
Two spp., Madagascar, in humid forest.
thecae discrete; pollen grains oblate spheroidal,
3-colpate, colpi narrow with uneven margins;
186. Megistostigma Hook. f.
sexine semi-tectate-reticulate, microverrucate;
Megistostigma Hook. f., Hook. Ic. Pl. 16: t. 1592 (1887), Fl. pistillode +. Pistillate flowers pedicellate; sepals
Brit. Ind. 5: 466 (1888); Merr., Phil. J. Sci. 16C: 563 (1920): 6, imbricate, pinnatifid, persistent in fruit; petals
Croizat, J. Arnold Arb. 22: 425 (1941); Backer & Bakh. f.,
Fl. Java 1: 491 (1963); Airy Shaw, Kew Bull. 23: 119 (1969),
and disk 0; ovary 3-locular, pubescent and some-
36: 330 (1981); Radcl.-Sm., Gen. Euphorb.: 251 (2001). times urticant-setose; ovules pachycaulous, inner
Clavistylus J.J. Sm. (1910). integument thick, outer integument thin; stylodia
united into a conical or infundibuliform column
154 G.L. Webster
or spherical mass. Fruits capsular, endocarp

woody; columella 3-pronged, not persistent.
Seeds globose; testa smooth.
Five spp. from east Africa. The genus appears
to be linked to Tragia by its urticant hairs and
pollen morphology, but resembles Cnesmone and
Sphaerostylis in its distinctive staminate flowers
and enlarged stylodia. There is also a resemblance
to Ctenomeria in the pinnatifid pistillate sepals,
but the pollen of the two genera is very different.
14c. SUBTRIBE DALECHAMPIINAE (M€

ull. Arg.)
G.L. Webster (1994).
Includes only the genus Dalechampia, whose
relationships to the Plukenetiinae and Tragiinae
remain unclear. The inflorescence of Dalecham-
pia has no counterpart in other Plukenetieae, and
Gillespie (1994) and Nowicke and Takahasi
(2002) have emphasized that its distinctive pollen
is strikingly different from that of either Plukene-
tiinae or Tragiinae. The primitive Dalechampia
species in sect. Rhopalostylis (Armbruster 1996)
have the stylar apparatus of Plukenetiinae (espe-
cially Romanoa) but the persistent stipules
and bipartite inflorescence of Tragiinae (Bia).
Urticant trichomes, which are present in the
Tragiinae, from Dalechampia are recorded in
the more derived sect. Dioscoreifoliae (on both
vegetative and reproductive parts) but not in the
basal sect. Rhopalostylis. Obviously, the subtribal Fig. 35. Euphorbiaceae-Acalyphoideae. Dalechampia
classification of the Plukenetieae needs a pro- dioscoreifolia. A Part of stem with leaves and inflores-
found revision. The molecular data available cence; note detail of adaxial leaf blade base with stipels.
(Wurdack et al. 2005) indicate that Dalechampia B Inflorescence subtended by bracts. C Staminate flower;
three sepals removed. D Pistillate flower. E Same, medial
might be embedded in the Plukenetieae (sister to section. F Fruit. G Sepals and column after fruit dehis-
Astrococcus). cence. H Fruit segment with seed and isolated seed.
(Reproduced with kind permission of the artist Bobbi
188. Dalechampia [Plumier] L. Fig. 35 Angell)
Dalechampia [Plumier] L., Sp. Pl.: 1054 (1753), Gen. Pl.
ed. 5: 473 (1754); M€ ull. Arg. in DC., Prodr. 15(2): 1232 Monoecious undershrubs or vines; indumentum
(1866), Fl. Brasil. 11(2): 633, t. 88–91 (1874); Pax & K. simple and urticant, sometimes scanty. Leaves
Hoffm., Pflanzenr. 147, XII: 3, figs. 1–9 (1919); Gagnep., entire or dentate, unlobed and pinnately veined
Fl. Indochine 5: 342, t. 39, 40 (1926); Léonard, Fl. Congo 8 to lobed and palmately veined, sometimes palma-
(1): 194 (1962); Webster & Armbruster, Brittonia 31: 352
(1979); Armbruster, Syst. Bot. 9: 272 (1984), 13: 303
tisect, often stipellate at base; stipules persistent.
(1988); Radcl.-Sm., Fl. E. Trop. Afr. Euphorb. 1: 285, fig. Inflorescences pseudanthial, pedunculate, axil-
56 (1987); Webster, Brittonia 41: 1, figs. 1–3 (1989); Web- lary or terminating short shoots, mostly bilater-
ster & Armbruster, Bot. J. Linn. Soc. 105: 137, figs. 1–5 ally symmetrical with a pair of subopposite
(1991); Armbruster, Syst. Bot. 21: 209, figs. 5–14 (1996), involucral bracts, bisexual; pistillate cymule
Smiths. Contr. Bot. 86: 14, figs. 4–8 (1997); Radcl.-Sm.,
Gen. Euphorb.: 262 (2001).
proximal, 3-flowered, bracteolate; staminate cym-
Cremophyllum Scheidw. (1842). ule terminal, involucellate, mostly of 8–12 flow-
Rhopalostylis Klotzsch ex Baill. (1858). ers, bractlets subtending staminate flowers
Megalostylis S. Moore (1916). resiniferous or odoriferous. Staminate flowers
Euphorbiaceae 155
pedicellate, pedicels articulate; calyx closed in pollen grains spheroidal, tricolporate-lalongate,

bud, splitting into mostly 3–6 valvate segments; tectate-microreticulate; pistillode 0. Pistillate
petals and disk 0; stamens 5–90; filaments con- flowers: sepals 5–6, imbricate?; petals 0; disk
nate into a thick column; anthers latrorse, 2- annular, thick; ovary 3-locular; stylodia 2. Fruits
celled, muticous; pollen grains prolate, 3-colpo- lobed capsules, dehiscing septicidally, septifrag-
rate, with costae equatoriales of two thickened ally and loculicidally; column 0?. Seeds subglo-
bands, sexine coarsely reticulate with very large bose; hilum triangular to heart-shaped; testa
lumina and high muri; pistillode 0. Pistillate flow- not fleshy; cotyledons broad, flat.
ers subsessile or pedicellate; sepals 5–12, imbri- One sp., A. kamerunica Pax & K. Hoffm.,
cate, basally connate, entire or pinnatifid; disk 0; Cameroon and Gabon.
ovary 3-locular, glabrous or pubescent; ovules In the molecular analysis of Kulju et al. (2008),
anatropous, inner integument thick, outer integ- Afrotrewia falls into the core acalyphoid clade
ument thinner, vascularized; stylodia completely basal to clades A2 and A3 of Wurdack et al. (2005).
connate into an elongate, sometimes clavate or
peltate column, stigmatic surface extending down
190. Crotonogynopsis Pax
much of the stylar surface. Fruits capsular, usu-
ally enclosed by the accrescent urticant calyx, Crotonogynopsis Pax, Bot. Jahrb. 26: 328 (1899); Prain, Fl.
dehiscing explosively by elastic twisting of the Trop. Afr. 6(1): 924 (1912); Pax & K. Hoffm., Planzenr.
147, VII: 14 (1914); Radcl.-Sm., Fl. Trop. E. Afr. Euphorb.
dry cocci; columella persistent. Seeds globose, 1: 213, fig. 40 (1987); Webster, Ann. Missouri Bot. Gard.
smooth or rugose. 2n ¼ 44, 46, 72, 138. 81: 80 (1994); Léonard, Fl. Afr. Centr. Euph. 3: 28, t. 4
About 120 spp., mainly in tropical regions of (1996); Radcl.-Sm., Gen. Euphorb.: 188 (2001).
the Americas, also in tropical Africa/Madagascar
Monoecious or dioecious trees or shrubs; indu-
and S and SE Asia. The genus was divided into 6
mentum simple. Leaves sessile or subsessile, den-
sections by Webster & Armbruster (1991), and
ticulate, with scattered glands abaxially; stipules
Armbruster (1996) has added an additional sec-
deciduous. Inflorescences axillary (ramiflorous),
tion. Dalechampia had been placed in a separate
unisexual, racemoid, the pistillate shorter and
tribe since M€uller (1866) until treated by Webster
few-flowered; bracts persistent. Staminate flowers
(1994), who demoted the Dalechampieae to a
pedicellate; calyx closed in bud, splitting into 2–5
subtribe of the Plukenetieae.
valvate segments; petals 0; disk of many inter-
Three unplaced acalyphoid genera: staminal glabrous segments; stamens 10–15, fila-
ments distinct; anthers dorsifixed, introrse or
extrorse, connective apiculate, not enlarged; pol-
189. Afrotrewia Pax & K. Hoffm. len grains 3-angled, 3-colporate, colpi granulate,
Afrotrewia Pax & K. Hoffm. in Engl., Pflanzenreich 147. sexine tectate-perforate; pistillode 0. Pistillate
VII: 14 (1914); Radcliffe-Smith, Gen. Euphorb.: 420 (2001); flowers pedicellate; sepals 4 or 5, imbricate,
Kulju, van der Ham & Breteler, Taxon 57: 137–143 (2008). persistent in fruit; petals 0; disk annular-lobed;
Shrubs to small trees, presumably dioecious; ovary 3(4)-locular, smooth and glabrous; ovules
indumentum minute, stellate to simple. Leaves anatropous, inner integument thick, outer integ-
subentire to shallowly serrate, with extrafloral ument thin; stylodia nearly distinct, bifid,
nectaries on both sides, petiole apically pulvinate; branches laciniate. Fruit capsular; columella per-
stipules deciduous to somewhat persistent. sistent. Seeds subglobose, ecarunculate, testa
Inflorescences unisexual, with prophylls at the smooth.
bases of the inflorescence branches and pedicels; Two spp., tropical Africa. Radcliffe-Smith
staminate inflorescences axillary to terminal (2001) referred the genus to the Adelieae, from
panicles, pistillate inflorescences 2–3-flowered which it differs in its 3-colporate pollen grains
racemes. Staminate flowers: sepals 2 or 3, valvate; with non-operculate colpi and a complex infra-
petals 0; receptacle conical; disk interstaminal, of tectum with double-layered columellae, otherwise
hairy glands; stamens 30–40, distinct; anthers known only from Alchorneinae (Alchornea,
basifixed, thecae 2, pendulous, unequally bilobed, Orfila) (Takahashi et al. 2000). The molecular
latrorse or introrse; connective horizontally data (Wurdack et al. 2005) resolve it in the Aca-
enlarged, partly covering the thecae, papillose; lypheae, close to Mareya and Acalypha.
156 G.L. Webster
191. Enriquebeltrania Rzedowski + or 0. Pistillate flowers with (2) 3–6 imbricate or

Enriquebeltrania Rzedowski, Bol. Soc. Bot. México 38: 75
valvate sepals; petals and disk + or 0; ovary
(1979); Webster, Ann. Missouri Bot. Gard. 81: 80 (1994); mostly 3-locular; outer integuments of ovules
Radcl.-Sm., Gen. Euphorb.: 189 (2001); De-Nova et al., sometimes vascularized; stylodia bifid to multifid,
Syst. Bot. 31: 533–546 (2006). less often unlobed; ovules solitary in each locule,
Beltrania Miranda (1957; nom. illeg.). anatropous. Fruit usually capsular or rarely
Dioecious shrubs; branches often spinescent; indehiscent. Seeds carunculate or ecarunculate;
indumentum simple. Leaves fasciculate, distally exotegmen palisadal, testa sometimes fleshy;
crenate to dentate, pellucid-punctate; stipules endosperm usually copious, often oily.
persistent. Flowers in axillary glomerules. Stami- This subfamily, subfam. Crotonoideae s.l.,
nate flowers pedicellate; sepals 3 or 4, valvate; which comprises 68 genera in twelve tribes,
petals and disk 0; stamens 20–30, filaments dis- contains lactiferous taxa with crotonoid pollen
tinct; anthers extrorse, apiculate; pollen grains sculpture and has been resolved to consist of
oblate spheroidal, 3-colporoidate, colpi inopercu- four clades (definition and topology of clades
late, with obscure margins, sexine tectate-pun- after Wurdack et al. 2005): (I) the Adenoclineae
cate, microverrucate; pistillode 0. Pistillate s.l. (tribe 1), (II) the Gelonieae (tribe 2), (III)
flowers pedicellate; sepals 4 or 5, entire, persistent the articulated crotonoids (tribes 3–6), and (IV)
in fruit; disk 0; ovary 2-locular; inner integu- the inaperturate crotonoids (¼ Crotonoideae s.
ments very thick, outer integuments thin; stylodia str.), comprising the remaining tribes 7–12.
distinct, unbranched. Fruits capsular, glabrous. Members of clades I–III share apetalous flowers,
Seeds subglobose, carunculate, testa minutely mostly thin integuments (thick in I), and aper-
rugulose. turate pollen, whereas clade IV has apetalous
Two spp., Mexico, Yucatan and Pacific coast. flowers, inaperturate pollen, and thick inner
Formerly this genus was included in Adelieae, but integuments that contain vascular bundles.
Takahashi et al. (2000) found the inoperculate These clades form a grade, in which (I), (II)
pollen grains of Enriquebeltrania very different and (III) subsequently are unsupported sister
from the operculate grains of Adelia. By molecu- clades to (IV), the latter comprising two sub-
lar studies (Wurdack et al. 2005; De-Nova et al. clades, C1, with tribes 7 and 8, and subclade C2,
2006), no precise placement within the New with tribes 9–12. Subclade C1 is from the New
World acalyphoids could be indicated either. World and the Old, whereas C2 is (nearly) con-
fined to the Old World. Molecular support for
subclade C2 is strengthened by a large trnL-F
spacer deletion.
VI. SUBFAM. CROTONOIDEAE Beilschm. (1833).
Monoecious or dioecious trees, shrubs, or herbs; KEY TO THE TRIBES OF SUBFAM. CROTONOIDEAE
laticifers articulated or non-articulated, latex col- 1. Pollen grains aperturate; petals 0; laticifers articulated
ored or 0; indumentum simple, stellate, or lepi- or inarticulated 2
dote; leaves alternate or opposite (whorled), – Pollen grains inaperturate; petals usually present, at
stipulate or estipulate, lamina simple to palmately least in staminate flowers; laticifers inarticulated
lobed or compound, often with basal laminar (articulated) 5
2. Laticifers articulated; staminate calyx gamophyllous;
glands. Inflorescences axillary or terminal, dicha- pollen grains 3-nucleate; plants monoecious or dioe-
sial to thyrsoid or spiciform. Staminate flowers cious; chromosome base number x ¼ 9 3
with imbricate or valvate sepals; petals and disk + – Laticifers non-articulated; staminate calyx mostly not
or 0; stamens (3–)5–many, filaments distinct or gamophyllous; pollen grains 2-nucleate; chromosome
connate; pollen grains binucleate or trinucleate, base number x ¼ 11 4
tricolpate, porate or inaperturate, sexine reticu- 3. Leaves palmately compound; inflorescences cymose-
late to more commonly tectate with triangular or paniculate, pistillate flowers terminal on lateral axes;
calyces gamophyllous; pollen grains colpate; stylodia
rounded-triangular processes [pillars] frequently stigmatiform; seeds ecarunculate, endosperm oily
forming continuous [triangular] arrays; pistillode 6. Heveeae
Euphorbiaceae 157
– Leaves simple to deeply lobed but not compound; 1. TRIBE ADENOCLINEAE (M€
inflorescences without terminal pistillate flowers on (1975).
lateral branches; calyces of distinct sepals; pollen
grains pantoporate; stylodia mostly multifid; stylodia Monoecious or dioecious; laticifers non-articu-
mostly multifid; seeds carunculate; endosperm lated, latex clear, often colored; indumentum
starchy 4. Manihoteae
simple, malpighiaceous, or stellate; leaves alter-
4. Pollen grains pantoporate; leaves pellucid-punctate,
stipules connate; inflorescences of leaf-opposed nate (opposite), simple; inner integument usually
glomerules 2. Gelonieae lacking vascular bundles and < 6 cells thick
– Pollen grains colp(or)ate; leaves not pellucid- [thick and vascularized in Klaineanthus]; stylodia
punctate, stipules distinct; inflorescences terminal bifid or stigmatiform, rarely completely connate
or axillary 6 into a column; fruits capsular or drupaceous;
5. Pollen exine lacking distinct Croton pattern, colpi seeds ecarunculate.
operculate; monoecious (dioecious); pistillate flowers
subsessile 5. Micrandreae
Unlike the Micrandreae and Manihoteae,
– Pollen exine with distinct Croton pattern, colpi inop- tribe Adenoclineae is represented in both the
erculate; mostly dioecious; pistillate flowers subsessile Neotropics and Paleotropics. Subtribe Endosper-
to pedicellate 1. Adenoclineae minae differs markedly in its stellate indumen-
6. Leaves simple, unlobed to palmately lobed; inflores- tum, connate stamens, and multilocular ovary,
cences terminal or axillary; seeds carunculate or but is linked with Omphalea and Suregada by
ecarunculate 7 the possession of alkaloidal glycosidase inhibitors
– Leaves palmatisect [except Givotia]; inflorescences and/or by being host plants for diurnal uraniine
axillary; seeds ecarunculate 11. Ricinodendreae
7. Staminate sepals distinct or connate but not fused and
moths (Kite et al. 1991; Wurdack et al. 2005).
enclosing petals in bud [except in Anomalocalyx];
seeds carunculate or ecarunculate; stipules persistent KEY TO THE SUBTRIBES AND GENERA OF ADENO-
or deciduous 8
CLINEAE
– Staminate sepals joined in bud, splitting into seg-
ments valvately or irregularly; seeds ecarunculate 1. Indumentum stellate; stamens connate; fruits baccate,
12. Aleuritideae 1–7-locular. Subtribe 1b. Endosperminae
8. Indumentum simple or malpighiaceous; pollen grains 197. Endospermum
binucleate; seeds carunculate or ecarunculate 9 – Indumentum simple or malpighiaceous; stamens
– Indumentum stellate or lepidote; pollen grains binu- distinct, rarely connate; fruits dehiscent or indehiscent;
cleate or trinucleate; seeds mostly carunculate [except 2- or 3-locular. Subtribe 1a. Adenoclininae 2
in Paracroton] 11 2. Stylodia completely connate into an obtuse or shortly
9. Seeds nearly exalbuminous; petals 0; stylodia 2–3-lobate stylar column; stamens 2–3, connate into a
unlobed; fruits indehiscent 3. Elateriospermeae short, slender column 195. Omphalea
– Seeds albuminous; petals usually present; stylodia – Stylodia distinct or largely so; stamens > 3, filaments
mostly bifid; fruits usually dehiscent 10 distinct 3
10. Inflorescences mostly dichasial, terminal, at least in 3. Pistillate disk 0; leaves glandular-dentate, without lam-
part; usually monoecious; indumentum often glandu- inar glands 193. Ditta
lar; leaf blades often palmately lobed or parted; – Pistillate disk +, at least as staminodia; leaves some-
stamens 5–12, filaments distinct or connate; seeds times with laminar glands 4
carunculate 7. Jatropheae 4. Herbs; seed coat not fleshy; staminate disk segments
– Inflorescences racemoid or spiciform to paniculate, ter- interstaminal 192. Adenocline
minal or axillary; monoecious or dioecious; indumen- – Trees or shrubs; seed-coat fleshy; staminate disk extra-
tum rarely glandular; leaf blades rarely palmately lobed; staminal or 0 5
stamens 7–35, filaments mostly distinct or nearly so; 5. Stamens 8–10; anthers 2-locular, not peltate;
seeds carunculate or ecarunculate 9. Codiaeae pollen grains tricolporate; staminate disk of 4 or 5 seg-
11. Branches and leaves with resinous glands, or else ments; leaves eglandular, stipules deciduous; indumen-
leaves opposite or verticillate; filaments erect in bud; tum simple; endotesta smooth 196. Klaineanthus
cotyledons as narrow as radicle or much broader – Stamens 3; anthers 4-locular, peltate; pollen grains tri-
10. Ricinocarpeae colpate; staminate disk 0; leaves usually with laminar
– Branches and leaves lacking resinous glands; leaves glands, stipules persistent; indumentum [at least in part]
alternate; filaments erect or inflexed in bud; cotyledons malpighiaceous; endotesta foveolate or echinulate
much broader than radicle 8. Crotoneae 194. Tetrorchidium
158 G.L. Webster
1a. SUBTRIBE ADENOCLININAE Arg. (1865). Dioecious shrubs; stems with resinous exudate;
indumentum simple, scanty. Leaves rigid, suben-
Trees, shrubs, or herbs; indumentum simple or tire or obscurely crenate with embedded marginal
malpighiaceous; stamens 3–30, filaments distinct; glands, revolute; stipules persistent, incrassate,
ovary 2- or 3-locular; stigmas distinct; fruit dark, resiniferous. Flowers axillary, staminate in
capsular or drupaceous. few-flowered glomerules, pistillate solitary. Sta-
This subtribe comprises five genera, which minate flowers sessile; sepals 3, distinct; petals
are found in the Neotropics and in tropical and and disk 0; stamens 3, filaments suppressed;
temperate Africa. anthers sessile, dorsifixed; pollen grains spheroi-
dal, 3-colpate, sexine tectate with apiculate pil-
192. Adenocline Turcz. lars; pistillode 0. Pistillate flowers sessile; sepals
Adenocline Turcz., Bull.Soc. Imp. Nat. Moscou 16: 59 suppressed; petals and disk 0; ovary 2- or 3-locu-
(1843); Prain, Ann. Bot. 27: 404 (1913), Fl. Cap. 5(2): 488 lar; ovules anatropous; stylodia distinct, unlobed,
(1920); Dyer, Gen. S. Afr. Fl. Pl., ed. 3, 1: 315 (1975); incrassate, stigmatiform. Fruits capsular, reddish.
Radcl.-Sm., Gen. Euphorb. 279 (2001).
Seeds ecarunculate. n ¼ 11.
Dioecious (monoecious) annual or perennial Two spp. described from the Greater Antilles
herbs, sometimes scrambling; laticifers non-artic- (Cuba, Hispaniola, and Puerto Rico), but Liogier
ulated, latex not apparent; indumentum 0. Leaves (1986) doubts that D. maestrensis Borhidi is dis-
alternate or opposite, petiolate or sessile, entire or tinct from D. myricoides Griseb. Ditta is morpho-
dentate, triplinerved to 1-veined, eglandular or logically distant from the other genera of
stipellate at apex of petiole; stipules persistent, Adenoclineae in its resinous axes and highly
lanceolate or subulate, dentate or lacerate, reduced flowers. However, the pollen is consis-
sometimes foliaceous. Flowers axillary, the stami- tent with the Adenoclineae (Nowicke 1994), and
nate in umbellate cymes or glomerules at upper in the molecular analysis Ditta is sister to
axils, sometimes aggregated into terminal pani- Tetrorchidium.
cles; pistillate flowers solitary, leaf-opposed; bracts
dissected, persistent, eglandular. Staminate flow- 194. Tetrorchidium Poepp.
ers pedicellate; sepals 5, imbricate; petals 0; disk Tetrorchidium Poepp., in Poepp. & Endl., Nov. Gen. Sp. 3:
segments interstaminal, sometimes stipitate; 23, t. 227 (1842); M€ ull. Arg. in DC., Prodr. 15(2): 1132
stamens mostly (6–)10(12), biseriate; filaments (1866); Pax & K. Hoffm., Pflanzenr. 147, IV: 29, figs. 8, 9
distinct; anthers basifixed, thecae discrete, glo- (1912); Cuatrecasas, Brittonia 9: 76, figs. 1–4 (1957); Léo-
bose, dehiscing vertically; pollen grains oblate nard, Fl. Congo 8(1): 133, t. 9 (1962); Radcl.-Sm., Fl. E.
Trop. Afr., Euphorb. 1: 373, fig. 70 (1987); Breteler, Adan-
spheroidal, 3-colpate, colpus margins irregular; sonia III, 21: 97 (1999).
sexine tectate, finely clavate; pistillode 0. Pistillate Hasskarlia Baill. (1860).
flowers pedicellate; sepals 5, entire, persistent in Tetrorchiopsis Rauschert (1982).
fruit; petals 0; disk of 3 petaloid segments; ovary
Dioecious trees or shrubs; latex whitish, often
3-locular; ovules anatropous; stylodia nearly dis-
scanty or 0; indumentum simple or malpighiac-
tinct, bipartite. Fruits capsular. Seeds ecaruncu-
eous. Leaves entire to dentate, eglandular [Afri-
late, testa smooth, rugulose, or foveolate.
can spp.] or with raised or stipitate glands at base
Eight spp. in temperate and subtropical
or laterally on petiole; stipules glandular, per-
South Africa (Dyer 1975; Radcliffe-Smith 2001),
sistent. Inflorescences axillary or leaf-opposed;
but only three in the enumeration of Govaerts
staminate spiciform, pistillate racemoid or panic-
et al. (2000).
ulate; bracts persistent, entire, sometimes glandu-
lar. Staminate flowers subsessile, several per
193. Ditta Griseb.
bract; sepals 3, distinct, imbricate, adaxially
Ditta Griseb., Mem. Amer. Acad. Arts Sci. II. 8: 160 ribbed; petals and disk 0; stamens 3, distinct,
(1861); Urban, Symb. Ant. 7: 261 (1912); Alain, Fl. Cuba filaments shorter than anthers; anthers extrorse,
3: 112, fig. 36 (1953); Little et al., Trees Puerto Rico &
Virgin I. 2: 400, t. 436 (1974); Liogier, Fl. Española 4: 135, peltate, 4-locular; pollen grains spheroidal,
348, fig. 116–13 (1986), Descr. Fl. Puerto Rico 2: 383, fig. 3-colpate, sexine tectate with rounded or angular
59–13 (1988); Radcl.-Sm., Gen. Euphorb.: 280 (2001). pillars; pistillode small or obsolete. Pistillate
Euphorbiaceae 159
flowers subsessile or pedicellate; sepals 3, dis-

tinct, imbricate, entire, persistent in fruit; petals
0; disk cupular or 3-lobed; ovary 2- or 3-locular;
ovules anatropous; stylodia distinct, bifid, style
branches sometimes dilated. Fruits capsular,
thin-walled. Seeds rounded, ecarunculate, exo-
testa fleshy, endotesta hard, foveolate.
Nineteen spp. in the Neotropics, and four in
Africa (Govaerts et al. 2000). In agreement with
Radcliffe-Smith (2001), the African Hasskarlia is
treated as a section of Tetrorchidium.
195. Omphalea L. Fig. 36

Omphalea L., Syst. Nat. ed. 10: 1264 (1759; nom. cons.);
M€ull. Arg. in DC., Prodr. 15(2): 1134 (1866), Fl. Brasil. 11
(2): 513, t. 72 (1874); Hemsl., Hook. Ic. Pl. 26: t. 2537
(1897); Croizat, Bull. Jard. Bot. Buit. III, 17: 204 (1941);
Alain, Fl. Cuba 3: 109 (1953); P.I. Forst., Austrobaileya 4:
381, t. 1 (1995); Gillespie, Novon 7: 127, t. 1 (1997),
Smiths. Contr. Bot. 86: 6, t. 1 (1997).
Monoecious trees, shrubs, or lianas; sap red or
pinkish; indumentum simple; foliage evergreen
or deciduous. Leaves pinnately or palmately
veined or lobed, with paired glands at base or
apex of petiole; stipules mostly entire,
deciduous. Inflorescences terminal, racemoid
or paniculate, bisexual or distal nodes staminate;
bracts foliose, generally glandular, stipulate. Sta-
minate flowers pedicellate; sepals (3)4–5, dis- Fig. 36. Euphorbiaceae-Crotonoideae. Omphalea dia-
crete, imbricate; petals 0; disk extrastaminal, ndra. A Flowering branch. B Partial inflorescence with
annular or segmented; stamens 2 or 3, filaments central pistillate flower and outer staminate flower buds.
C Androecium above annular nectary. D Staminate flower
connate into a apically dilated column; anthers at anthesis. E Fruit. (Gillespie 1997)
latrorse; pollen grains oblate spheroidal to oblate,
3-colpate, sexine tectate-foveolate, microverru-
cate; pistillode 0. Pistillate flowers subsessile or iospermum, and other genera now assigned to the
pedicellate; sepals 4 or 5, discrete, imbricate, Crotonoideae. Pax and Hoffmann first (1919)
deciduous; disk 0; ovary 3(4)-locular, glabrous; placed Omphalea into the Hippomaneae, and
ovules pachycaulous, inner and outer integu- later in a subtribe of the Gelonieae. Croizat
ments moderately thick; stylodia completely con- (1941) suggested an affinity with the Plukene-
nate into a stylar column, distally obtuse or very tieae. The molecular data (Wurdack et al. 2005)
shortly 2–3-lobate. Fruits capsular or baccate; place Omphalea in the Adenoclineae. Interest-
columella not persistent. Seeds subglobose, ingly, Omphalea, Endospermum, Tetrochidium
sometimes compressed or angular, ecarunculate; and Suregada (Gelonieae) accumulate alkaloidal
exotesta fleshy to papery, endotesta hard, smooth. protease inhibitors, and are host plants for spe-
Seventeen spp. (Gillespie 1997; 22 according cialist diurnal uraniine moths (Kite et al. 1991;
to Govaerts et al. 2000), scattered through neo- Wurdack et al. 2005).
tropical and paleotropical regions, except the
196. Klaineanthus Pierre ex Prain
Pacific islands. In terms of systematics, Omphalea
is one of the most problematic genera of the Klaineanthus Pierre ex Prain, Kew Bull. Misc. Inf. 1912:
family. M€ uller (1866) assigned it to his subtribe 105 (1912), Fl. Trop. Afr. 6(1): 963 (1913), Hook. Ic. Pl. 30:
t. 2985 (1913); Keay, Fl. W. Trop. Afr., ed. 2, 1: 413 (1958);
Gelonieae, associated with Endospermum, Elater- Léonard, Fl. Congo 8(1): 130 (1962).
160 G.L. Webster
Dioecious trees; latex not recorded; indumentum open in bud; petals 0; disk extrastaminal, 4–5-
simple. Leaves long-petiolate, eglandular; stipules angled; stamens (3–)6–10, filaments connate,
minute, deciduous. Inflorescences terminal and anthers nearly sessile on staminal column, did-
axillary, the staminate paniculate, the pistillate ymous and 4-locellate; pollen grains suboblate, 3-
racemose; bracts deciduous. Staminate flowers colpate, sexine tectate with clavate pillars; pistil-
pedicellate; sepals (3) 4 or 5, basally connate, lode minute or 0. Pistillate flowers subsessile or
imbricate; petals 0; disk segments 4 or 5; stamens pedicellate; calyx 5-angled or -lobed; persistent
8–10, biseriate, filaments distinct; anthers basi- in fruit; petals 0; disk patelliform or cupular;
fixed, introrse; pollen grains suboblate, 3-colpo- ovary 2–6-locular, pubescent; ovules anatropous;
rate, sexine tectate, with clavate pillars; pistillode stylodia stigmatiform, coalescing into a disk.
2–3-lobed. Pistillate flowers pedicellate; sepals 5 Fruits separting into indehiscent, sometimes
(4), distinct, imbricate, entire, deciduous in fruit; fleshy, cocci; columella not persistent. Seeds
petals 0; disk annular, with 4 or 5 staminodes; spheroidal, ecarunculate, testa rugose or verru-
ovary 3-locular, glabrous; ovules anatropous, cose. n ¼ 24.
inner integuments with 9–11 cell layers, vascular- About 10 spp., distributed from China to
ized; stylodia almost distinct, bipartite, the arms northern Australia, Melanesia and Fiji. All Male-
2-lobulate at their apices. Fruits capsular, with sian Endospermum have extrafloral nectaries on
thin papery walls; columella persistent, slender; the lower leaf surface, and two New Guinean
seeds with fleshy exotesta, endotesta smooth. species are myrmecophilous, of which one,
A single sp., K. gaboniae Pierre ex Prain from E. moluccanum (Teijsm. & Binn.) Kurz, produces
West Africa (Nigeria to Gabon). It is anomalous food bodies on which the ants feed (Guerrero and
in the Adenoclineae in having a thick, vascular- van Welzen 2011).
ized integument.
2. TRIBE GELONIEAE (M€
1b. SUBTRIBE ENDOSPERMINAE Pax & K. Hoffm. Dioecious trees or shrubs, without evident latex;
(1931). indumentum simple, usually scanty or 0; leaves
alternate, simple; inflorescences leaf-opposed;
Dioecious trees or shrubs, indumentum stellate; flowers in glomerules; disk receptacular or extra-
staminate calyx gamophyllous; stamens 6–10, staminal; stamens 6–60; pollen grains spheroidal,
filaments connate; ovary 2–6-locular; inner inte- pantoporate, sexine with Croton pattern; ovules
guments thin, not vascularized; stylodia stigmati- with inner integuments thin, not vascularized.
form, confluent; fruit capsular. Two genera, Africa, Madagascar, and tropical
Monotypic, including only the paleotropical Asia.
genus Endospermum.
KEY TO THE GENERA OF GELONIEAE
197. Endospermum Benth. 1. Inflorescences leaf-opposed; pistillode 0; stylodia bifid
or multifid, branches terete, not dilated; leaves petio-
Endospermum Benth., Fl. Hongkong.: 304 (1861; nom.
late, lamina pellucid-punctate, stipules deciduous
cons.); Beccari, Malesia 2: 45, t. 2 (1884); Pax & K.
198. Suregada
Hoffm., Pflanzenr.147, IV: 33 (1912); Radcl.-Sm., Gen.
Euphorb.: 281 (2001); Guerrero & van Welzen, Edinb. J. – Inflorescences terminal; pistillode present; stylodia
Bot. 68:443–482 (2011), rev. dilated, stigmatiform; branches flattened, dilated;
lamina not pellucid-punctate; stipules foliaceous,
Dioecious trees; latex white, often not evident; persistent 199. Cladogelonium
indumentum stellate-fasciculate, sometimes
scanty. Leaves long-petiolate, sometimes peltate, 198. Suregada Roxb. ex Rottl.
palmately veined or triplinerved, with abaxial Suregada Roxb. ex Rottler, Ges. Naturf. Freunde Berlin,
paired sessile patelliform or conical glands at Neue Schriften 4: 206 (1803); Baillon, Étude Gén. Euphorb.:
junction with petiole; stipules entire, deciduous. 395 (1858); Croizat, Bull. Bot. Gard. Buitenzorg III, 17: 212
Inflorescences axillary, staminate paniculate, pis- (1942); Léonard, Bull. Jard. Bot. Brux. 28: 79 (1958), Fl.
Congo 8(1): 124 (1962); Radcl.-Sm., Fl. E. Trop. Afr.,
tillate spicate or racemose, sometimes narrowly Euphorb. 1: 376, fig. 71 (1987), Fl. Zambes. 9(4): 249, t. 53
paniculate; bracts entire or unidentate, persistent (1996); Radcl.-Sm. et al., Kew Bull. 58: 965–970 (2003), key
or deciduous, eglandular. Staminate flowers sub- Malagasy spp.; Li-Bingtao & Esser, Fl. China 11: 276 (2008).
sessile, basally articulate; calyx 2–4-lobed, often Gelonium Roxb. ex Willd. (1806; nom. illeg.).
Euphorbiaceae 161
Dioecious trees or shrubs, without evident latex; Tetrorchidium; they suggest the transfer of the
indumentum simple, usually scanty or 0. Leaves genus to Adenoclineae.
usually pellucid-punctate, without excavated
glands; stipules deciduous. Inflorescences leaf- 3. TRIBE ELATERIOSPERMEAE G.L. Webster (1975).
opposed, glomerular, sessile or subsessile. Stami-
nate flowers subsessile or pedicellate; sepals 5(6), Monoecious laticiferous trees; leaves biglandular
distinct, broadly imbricate; petals 0; disk recepta- at base; pollen inaperturate; fruits capsular; seeds
cular or extrastaminal, annular or dissected; ecarunculate.
stamens (6)10–30(60), filaments distinct; A monotypic paleotropical tribe. Although
anthers dorsifixed, introrse, muticous; pollen articulated laticifers are not yet documented for
grains spheroidal, 2-nucleate, 3–6-porate; sexine this genus (and not for Glycydendron either), the
tectate with clavate pillars in Croton pattern; pis- lack of petals, the presence of a vascularized
tillode 0. Pistillate flowers subsessile or pedicel- inner integument, and the molecular data safely
late; sepals (4)5(8), distinct, imbricate, entire, place them into the articulated crotonoids. For
persistent in fruit; petals 0; disk annular, tenuous, the problematic "inaperturate" pollen grains, see
sometimes with staminodes; ovary (2)3(4)-locu- Nowicke (1994).
lar; ovules pachychalazal, inner integuments thin;
stylodia bifid or multifid, branches slender. Fruits 200. Elateriospermum Blume Fig. 37
capsular or sometimes indehiscent; columella
Elateriospermum Blume, Bijdr.: 620 (1826); Whitmore,
persistent. Seeds ellipsoid, ecarunculate, testa Tree Fl. Malaya 2: 91, fig. 6 (1973); Hoang Van Sam &
smooth. 2n ¼ 22. van Welzen, Blumea 49: 427–436 (2004).
About 30 spp., Africa/Madagascar and tropi-
cal Asia. Monoecious trees; stems with white latex; indu-
mentum simple or 0. Leaves long-petiolate,
199. Cladogelonium Leandri biglandular at junction with petiole; stipules
deciduous. Inflorescences axillary, dichasial,
Cladogelonium Leandri, Bull. Soc. Bot. France 85: 530, bisexual, pistillate flower central; bracts minute.
t. 1.15–19bis (1938); Webster, Ann. Missouri Bot. Gard. Staminate flowers pedicellate; sepals 4–5, dis-
81: 131 (1994); Radcl.-Sm., Gen. Euphorb.: 284, fig. 38
(2001). tinct, imbricate; petals 0; disk massive, lobed,
pubescent; stamens 10–20 or more, filaments
Monoecious shrubs; branches sympodial, flat- distinct; anthers introrse, connective apiculate;
tened into platyclades similar in texture to the pollen grains spheroidal, exine with Croton pat-
leaves; latex not recorded; indumentum 0. Leaves tern, muri broad, pillars angular; pistillode min-
not pellucid-punctate, obscurely dentate and with ute or 0. Pistillate flowers pedicellate; sepals (4)5
abaxial excavated marginal glands; stipules folia- (7), distinct, entire, deciduous (?); disk annu-
ceous, persistent. Inflorescences terminating lar, with subulate staminodia; ovary 2–4-locular,
sympodial axes, glomerulate, unisexual, sessile. villose; ovules with inner integuments thin, vas-
Staminate flowers pedicellate; sepals 5, imbricate, cularized; stylodia 2–4, massive, stigmas thick,
distinct, each with an abaxial gland; petals 0; dilated. Fruits capsular, endocarp thick and
stamens 10, filaments distinct; anthers dorsifixed, woody; columella not persistent. Seeds over 3
extrorse; pistillode 3-fid. Pistillate flowers soli- cm long, testa smooth; endosperm scanty, coty-
tary, pedicellate; sepals 5, distinct, imbricate, ledons massive.
each with an abaxial gland; petals 0; disk tripartite A single sp., E. tapos Blume, distributed in
[staminodial?]; ovary 3-locular; stylodia stigmati- lowland rainforests from peninsular Thailand
form. Fruit capsular. Seeds ellipsoid, with minute through Malaya to Sumatra, Java and Borneo.
caruncle, testa foveolate.
A single very rare sp., C. madagacariense 4. TRIBE MANIHOTEAE (M€
Leandri, in dry deciduous forest of Madagascar.
It appears to be a satellite genus of Suregada, Monoecious (dioecious); laticifers articulated,
from which it differs in details of inflorescence latex white; indumentum simple (urticant); leaves
and floral structure. Lobreau-Callen and Suarez mostly palmately lobed or dissected, sometimes
Cervera (1997) report 3- or 6-colpate pollen from unlobed; staminate calyx synsepalous; disk
Cladogelonium and a nexine structure similar to intrastaminal or extrastaminal; stamens 8–10
162 G.L. Webster
– Stinging hairs +, very rarely 0; stamens connate, very

rarely distinct; staminate disk extrastaminal, perianth
whitish; leaf blades glandular at base; inflorescence
dichasial-paniculate 202. Cnidoscolus
201. Manihot Miller

Manihot Miller, Gard. Dict. ed. 4 (1754); Pohl, Pl. Bras. Ic.
1: 17 (1827); Croizat, J. Arnold Arb. 23: 216 (1942); Web-
ster, J. Arnold Arb. 48: 345 (1967); Rogers & Appan, Fl.
Neotrop. 13: 1, figs. 9–124 (1973); Allem, Rev. Brasil. Biol.
49: 1 (1989), Genet. Res. Crop Evol. 41: 133 (1994).
Manihotoides C.J. Rogers & Appan (1973).
Monoecious trees or shrubs, sometimes scandent
or nearly herbaceous, often with tuberous roots;
latex white; indumentum simple or 0. Leaves
mostly long-petiolate, simple, mostly palmately
lobed, entire (serrulate), stipellate at junction
with petiole; stipules mostly deciduous. Inflores-
cences terminal or pseudo-axillary, racemoid
or paniculate, pistillate flowers generally basal,
staminate in distal glomerules; bracts entire to
laciniatae, mostly deciduous. Staminate flowers
subsessile to pedicellate; sepals 5, imbricate,
connate basally; petals 0; disk central, intras-
taminal, entire or 5-lobed, lobes bifid; stamens
10, biseriate, filaments distinct; anthers introrse;
pollen spheroidal, pantoporatae; pistillode rudi-
mentary or 0. Pistillate flowers distinctly pedicel-
late; sepals 5, imbricate, deciduous in fruit; disk
annular, fleshy, scarcely lobed; ovary 3-locular,
glabrous; ovules anatropous, thick, vascularized;
stylodia basally connate, branches dilated to lac-
Fig. 37. Euphorbiaceae-Crotonoideae. Elateriospermum erate. Fruits capsular, sometimes ribbed or
tapos. A Flowering branch. B Basal leaf glands. C Part of winged; columella often persistent. Seeds carun-
inflorescence with one pistillate and six staminate flower
buds. D Staminate flower with sepals removed. E Pistillate culate, testa smooth. 2n ¼ 36.
flower with sepals removed, showing pistil and disk lobes. About 100 spp. in the Neotropics, divided by
F Fruit. G Seed. H Fruit valves with complete septicidal Rogers and Appan (1973) into 19 sections, plus
dehiscence and partly loculicidal dehiscence. (Hoang Van
Sam & van Welzen 2004; drawn by J. van Os)
the proposed segregate genus Manihotoides,
which could easily be accommodated in Manihot
adjacent to sect. Parvibracteatae.
(25); pollen grains 3-nucleate, pantoporate;
inner integuments thick, vascularized.
A small neotropical tribe of only two genera
202. Cnidoscolus Pohl
but with over 100 spp., mostly found in deciduous
or thorn forests in arid regions of both North and Cnidoscolus Pohl, Pl. Bras. Icon. Descr. 1: 56 (1827); León,
South America. Mem. Soc. Cubana Hist. Nat. 15: 235, t. 23, 24 (1941);
Lourteig & O’Donell, Lilloa 9: 105, figs. 5–9 (1943);
McVaugh, Bull. Torrey Bot. Club 71: 457 (1944); Breckon,
KEY TO THE GENERA OF MANIHOTEAE Brittonia 31: 125, figs. 3–6 (1979); Fernández Casas, Font-
1. Stinging hairs 0; stamens distinct, staminate disk queria 55: 69 (2002).
intrastaminal, perianth usually yellowish or greenish; Victorinia León (1941).
leaf blades stipellate at base; inflorescence racemoid or Monoecious trees, shrubs, or perennial herbs;
racemose-paniculate 201. Manihot
stems with septate pith; latex white; indumentum
Euphorbiaceae 163
simple and urticant. Leaves palmately (pinnately) KEY TO THE GENERA OF MICRANDREAE
veined or lobed or sometimes dissected, glandu- 1. Sepals connate; staminate disk 0; stamens 8–10; floral
lar at junction with petiole; stipules entire to bracts large 204. Cunuria
laciniate, glandular, mostly persistent. Inflor- – Sepals distinct; staminate disk +; stamens 5–8; floral
escences terminal or sometimes pseudo-axillary, bracts small 2
pedunculate, dichasial, pistillate flowers at proxi- 2. Indumentum simple; anthers elliptical; pollen muri
mal nodes, staminate in clusters at distal nodes; smooth 203. Micrandra
bracts entire to laciniate, sometimes foliaceous, – Indumentum stellate; anthers linear; pollen muri irre-
gularly appendaged 205. Micrandropsis
eglandular. Staminate flowers subsessile or short-
pedicellate; sepals imbricate, connate, the calyx 203. Micrandra Benth.
salverform; disk annular, extrastaminal;
stamens 8–10(25), filaments connate into a col- Micrandra Benth., Hook. Kew J. Bot. 6: 371 (1854; nom.
umn or those in the outer whorl distinct [all cons.); Schultes, Bot. Mus. Leafl. 15: 201, t. 66–73 (1952);
Webster, Ann.. Missouri Bot. Gard. 81: 98 (1994); Radcl.-
filaments distinct in C. urnigerus]; anthers in Smith, Gen. Euphorb.: 268 (2001); Berry & Wiedenhoeft,
2–3 (4–5) whorls, basifixed to dorsifixed, introrse; Syst. Bot. 29: 125–133 (2004).
pollen grains spheroidal, 3-nucleate, pantoporate,
sexine with angular pillars; pistillode of 3 filiform Monoecious trees, often buttressed or with stilt
processes atop the column. Pistillate flowers sub- roots at base; laticifers non-articulated; latex
sessile or pedicellate; sepals 5, distinct or connate, copious, white; indumentum simple. Leaves
entire, deciduous in fruit; petals 0; disk annular, eglandular at base [except M. elata]; stipules
sometimes with staminodia; ovary 3-locular, mostly deciduous. Inflorescences terminal on
often with urticant hairs; ovules anatropous, principal and lateral shoots, pedunculate; stami-
inner integuments thick, vascularized; stylodia nate flowers in glomerules, pistillate solitary at
distinct or nearly so, multifid (bifid). Fruits cap- tips of lateral axes; bracts entire, eglandular,
sular or less often fleshy and tardily dehiscent; deciduous. Staminate flowers subsessile or pedi-
columella slender, persistent. Seeds carunculate, cellate; sepals 5, valvate or slightly imbricate;
testa smooth. 2n ¼ 36. petals 0; disk-segments 5; stamens usually 4 or
About 70 spp. in the Neotropics. McVaugh 5, filaments distinct, apically inflexed in bud;
(1944) recognized 5 sections that have not yet anthers subglobose, dehiscing laterally; pollen
been evaluated by a phylogenetic analysis. Speci- grains spheroidal, 3-colpate, colpi operculate,
ation has been most pronounced in Mexico; sexine [except in M. elata] tectate-reticulate,
Breckon (1975) revised sect. Calyptosolen and with smooth muri; pistillode much shorter than
recognized 20 species in Mexico and Central filaments or obsolete. Pistillate flowers subsessile;
America. sepals 5, entire, distinct, deciduous; petals 0; disk
annular; ovary 3-locular, sericeous, beaked;
5. TRIBE MICRANDREAE (M€
ull. Arg.) G.L. Webster ovules anatropous, inner integument thick, vas-
(1975). cularized; stylodia distinct, bifid. Fruits capsular;
columella slender, subpersistent. Seeds large, > 1
Monoecious or dioecious; stems with inarticu- cm long, ecarunculate or with rudimentary car-
lated laticifers and usually whitish latex; indu- uncle, testa smooth.
mentum simple or stellate; leaves unlobed, Five or six neotropical spp. from lowland
entire; disk dissected, lobed, or 0; pollen grains Amazonian forests; following Webster (1994)
3-nucleate, 3-colpate, colpi operculate; inner and in contrast to Govaerts et al. (2000) and
integument thick, vascularized. Berry and Wiedenhoeft (2004), Micrandra is
This tribe resembles various taxa of subfam. here delimited to exclude Cunuria. Even in this
Acalyphoideae; Micrandra in particular shows limited sense, it is heterogeneous: M. siphonioides
many acalyphoid characters, resembling Cheilo- Benth. and M. minor Benth. have 3-colpate
seae in its androecium and Alchorneae in its operculate pollen grains with tectate-reticulate
operculate pollen grains. exine very suggestive of pollen grains in the
164 G.L. Webster
Acalyphoideae, but M. elata has pollen grains with imbricate; disk 5-lobed; stamens 5(7), filaments
Crotonoid ornamentation distinct from all other distinct; anthers linear; pollen grains spheroidal,
Micrandrinae. M. inundata P.E. Berry & A.C. Wie- 3-colpate, tectate-reticulate with irregularly spi-
denhoeft is known from the banks of seasonally nose muri; pistillode 0. Pistillate flowers subses-
flooded blackwater rivers in southwestern Vene- sile; sepals 5, distinct, entire, deciduous in fruit;
zuela; its trunks consist of very lightweight wood. disk annular, tenuous; ovary 3-locular, sericeous.
Fruits capsular, valves rugose; seeds carunculate.
204. Cunuria Baill. A single sp., M. scleroxylon (W.A. Rodrigues)
Cunuria Baill., Adansonia I, 4: 287 (1864); Baldwin &
W.A. Rodrigues from the lowland rainforest in
Schultes, Bot. Mus. Harvard Univ. 12: 325, t. 42–46 the vicinity of Manaus, Amazonas, Brazil, but
(1947); Webster, Ann. Missouri Bot. Gard. 81: 98 (1994); later reported from Amazonian Colombia by
Radcl.-Sm., Gen. Euphorb.: 270 (2001). Murillo & Franco (1995). The pollen grains of
Monoecious or dioecious trees; laticifers non-artic- Micrandropsis are quite distinct from both
ulated, latex white or yellow; indumentum simple. Micrandra and Cunuria in their unique irregu-
Leaves biglandular at basal juncture with petiole; larly ornamented muri.
stipules deciduous. Inflorescences terminal or axil-
lary, pedunculate, dichasially paniculate; staminate 6. TRIBE HEVEEAE (M€
ull. Arg.) G.L. Webster,
flowers in glomerules, pistillate solitary, terminal stat. nov.1
on axes; bracts entire, eglandular, deciduous. Sta-
Monoecious; leaves palmately compound; inflo-
minate flowers sessile or subsessile; sepals 5, dis-
rescence with terminal pistillate flower; staminate
tinct, imbricate; petals and disk 0; stamens 7–10,
sepals connate, valvate; stamen filaments connate
distinct; anthers dorsifixed, muticous; pollen
into a column; pollen grains 3-colpate, colpi
grains spheroidal, 3-colpate, colpi operculate, sex-
operculate, sexine tectate-baculate.
ine with raised muri and usually regular or irregu-
In Webster (1994), Hevea was assigned to
lar processes; pistillode trifid. Pistillate flowers
subtribe Heveinae in the Micrandreae. However,
subsessile; sepals 5, imbricate, entire, deciduous;
Hevea differs from the Micrandreae in a number
disk cupular or annular, sometimes lobed; ovary 3-
of characters, such as its articulated laticifers,
locular, glabrous or pubescent; stylodia bifid,
gamophyllous calyx, connate stamens, very fine
branches thickened. Fruits capsular. Seeds ecarun-
pollen exine ornamentation, and palmately com-
culate; testa smooth and shiny.
pound leaves. It appears that Hevea has more in
Six neotropical spp. This genus was generally
common with the Manihoteae, despite differences
accepted by botanists until Schultes (1952) decided
in pollen apertures.
to combine it with Micrandra, in which he is
followed by Berry and Wiedenhoeft (2004). How-
206. Hevea Aubl. Fig. 38
ever, if Micrandra elata is transferred to Cunuria,
the remainder of the Micrandra species differ from Hevea Aubl., Hist. Pl. Guiane Fr. 2: 871, t. 335 (1775);
Cunuria in having leaf blades without basal glands Ducke, Arch. Inst. Biol. Veg. Rio Janeiro 2: 217 (1935);
Schultes, Bot. Mus. Leafl. Harvard Univ. 25: 243, t. 51, 52
and very different pollen ornamentation. (1977); Bot. Review 36: 197 (1970); Malaysian Rubber Res.
Dev. Board Mon. 14: 5 (1990); Murillo & Franco, Euforb.
205. Micrandropsis W.A. Rodrigues Reg. Araracuara 78, figs. 17, 18 (1995); Hoang Van Sam &
van Welzen, Blumea 49: 427–435 (2004).
Micrandropsis W.A. Rodrigues, Acta Amazonica 3(2): 5
(1973); Webster, Ann. Missouri Bot. Gard. 81: 98 (1994); Monoecious trees; laticifers articulated, latex
Murillo & Franco, Euphorb. Reg. Araracuara 117, fig. 119 whitish; indumentum simple. Leaves palmately
(1995); Radcl.-Sm., Gen. Euphorb.: 270, fig. 37 (2001). compound, with 1–3 raised glands at apex of
Monoecious trees; latex scanty; indumentum petiole, leaflets entire; stipules entire, deciduous.
minute, stellate. Leaves with 1 or 2 sessile glands
at juncture with petiole. Inflorescences terminal
on principal and lateral axes, pedunculate, stami-
nate flowers in few-flowered dichasia, pistillate 1
Tribus Heveeae (M€ ull. Arg.) G.L. Webster, stat. nov. Basio-
flowers solitary; bracts glandular or eglandular. nym: Subtribus Heveinae M€ uller Argoviensis, Linnaea 34:
Staminate flowers subsessile; sepals 5, distinct, 202 (1865). Type: Hevea Aubl.
Euphorbiaceae 165
basally connate, stigmatiform. Fruits capsular; col-

umella fragile, semipersistent. Seeds ellipsoid,
smooth, ecarunculate; endosperm oily. n ¼ 18.
According to Schultes (1970), Hevea includes
nine spp. with a total of five varieties that are
essentially confined to the Amazon basin, although
Hevea brasiliensis (A. Juss.) M€ull. Arg. is widely
cultivated circumtropically. In its compound
leaves and connate sepals and stamens, Hevea is
distinct from the taxa of tribe Micrandreae; how-
ever, the pollen grains show some similarities to
species such as Micrandra lopezii.
Unplaced genus with a strong affinity to the

articulated crotonoids:
207. Glycydendron Ducke
Glycydendron Ducke, Arq. Jard. Bot. Rio Janeiro 3: 199
(1922), 4: 107, t. 10 figs. a–i (1925); Pax & K. Hoffm., Nat.
Pflanzenfam. ed. 2, 19c: 181 (1931, as Glycynodendron);
Webster, Ann. Missouri Bot. Gard. 81: 100 (1994); Vás-
quez Martı́nez, Fl. Res. Biol. Iquitos, Peru: 287, t. 35A
Dioecious trees; latex whitish, translucent; indu-
mentum simple. Leaves long-petiolate, deciduous
before flowering, simple, entire, triplinerved, with
paired adaxial excavated basal glands; stipules
minute, deciduous. Inflorescences axillary, stami-
nate also subterminal, sometimes fasciculate, the
staminate cymose-paniculate, pistillate racemose;
bracts minute, eglandular. Staminate flowers ped-
Fig. 38. Euphorbiaceae-Crotonoideae. Hevea brasiliensis. icellate; sepals 4, distinct, imbricate; petals 0; disk
A Flowering branchlet. B Part of inflorescence with a large glands intrastaminal on the pilose receptacle;
terminal pistillate flower and smaller staminate flowers. C stamens 25–30, filaments distinct; anthers
Staminate flower, part of calyx removed. D Pistillate
flower, part of calyx removed. E Fruit. F Seed, ventral introrse, basifixed, muticous; pollen grains
and dorsal view. (Hoang Van Sam and van Welzen 2004; oblate, 3-colpate, sexine tectate with Croton pat-
drawn by J. van Os) tern; pistillode 0. Pistillate flowers pedicellate;
sepals 4, imbricate, distinct, entire, deciduous;
Inflorescences axillary, paniculate with dichasial petals 0; disk annular with staminodia; ovary 2-
subunits, the pistillate flowers solitary and termi- locular, sericeous; ovules anatropous, inner inte-
nal on lateral axes; bracts small, entire, deciduous. guments 8–10 cells thick, vascularized; stylodia
Staminate flowers pedicellate; sepals connate with distinct, bipartite. Fruits drupaceous, 1-seeded,
valvate lobed; disk segments distinct or connate; endocarp woody. Seeds ecarunculate.
stamens 6–9 (10) in 2 irregular whorls; anthers Probably monotypic, G. amazonicum Ducke
sessile; pollen grains 3-colpate, colpi inoperculate, appears to be common and widespread from
sexine tectate-reticulate, muri hexagonal or pen- Pará in eastern Brazil west to Ecuador, Peru, and
tagonal, with rounded or angular projections; pis- Bolivia, with an outlying population in the mata
tillode represented by terminal appendage of atlántica of eastern Brazil (Espı́rito Santo). In the
staminal column. Pistillate flowers pedicellate, molecular analyses (Wurdack et al. 2005; Tokuoka
sepals 5, connate, deciduous in fruit; disk rudi- 2007), it is resolved in the articulated crotonoids;
mentary or 0; ovary 3-locular; ovules anatropous, in its larger stamen number and drupaceous fruit,
inner integuments thick, vascularized; stylodia it comes close to paleotropical Elateriospermum.
166 G.L. Webster
7. TRIBE JATROPHEAE (Meisn.) Pax (1890). pedicellate; sepals 5, imbricate, distinct to basally
connate, entire to dentate, sometimes foliaceous;
Monoecious (dioecious); stems with articulated petals 5, distinct to coherent or connate; disk
and/or non-articulated laticifers, the latex white entire to lobed or dissected; stamens (6–)8–10
or reddish; indumentum simple, sometimes glan- (12), filaments distinct or connate; anthers
dular; leaves simple to deeply palmately lobed or mostly biseriate, dorsifixed; pollen grains sphe-
compound; pollen grains spheroidal, binucleate, roidal, binucleate, inaperturate, exine with Cro-
inaperturate, sexine with Croton pattern; inner ton pattern, pillars rounded or angular, smooth
integuments thick, vascularized. or sulcate; pistillode 0. Pistillate flowers pedicel-
Three genera, two of which are neotropical late; sepals 5, nearly distinct, imbricate, entire or
whereas one, Jatropha, is well developed in Amer- dentate, persistent in fruit; petals 5, distinct to
ica and Africa and extends to Madagascar and coherent or connate, imbricate; disk annular or
Asia. Among the genera previously placed in this deeply lobed; ovary (1–)3-locular, glabrous or
tribe, Leeuwenbergia safely belongs to the inaper- pubescent; ovules anatropous, inner integuments
turate crotonoid clade C2 with the large deletion in thick, vascularized; stylodia bifid, sometimes
the trnL-F spacer (Wurdack et al. 2005), whereas dilated (multifid). Fruits capsular, sometimes
molecular data for the Old World Deutzianthus with fleshy exocarp and tardily dehiscent, (1–)
and Oligoceras are lacking (provisionally placed 3-seeded; columella generally not persistent.
in Aleuritideae-Grosserinae). Seeds ellipsoidal to spherical, carunculate; testa
thin and smooth; endosperm oily; cotyledons
KEY TO THE GENERA OF JATROPHEAE much longer and broader than radicle. 2n ¼ 22.
1. Leaves simple, entire, elobate; stipules infra-axillary, Over 180 spp., widely distributed in the tro-
deciduous; seeds ecarunculate 210. Vaupesia pics and subtropics of America and Africa, rare in
– Leaves palmately lobed or palmately compound; sti- Madagascar and Asia. The circumscription of
pules not infra-axillary or 0 2 Jatropha in the 19th and early 20th century
2. Leaves 3- to more palmatilobed or -partite; sepals dis-
tinct, covering the petals in bud; fruit capsular; seeds was confused due to the mistaken inclusion of
carunculate 208. Jatropha Cnidoscolus by M€ uller (1866) and Pax (1910),
– Leaves compound; staminate calyx open in bud, but McVaugh (1944) demonstrated that the two
not covering the petals; fruit drupaceous; seeds genera are not closely related.
ecarunculate 209. Joannesia
209. Joannesia Vell.
208. Jatropha L.
Joannesia Vell., Alogr. Alkalis: 199 (1798); Ducke, Arch.
Jatropha L., Sp. Pl.: 1006 (1753); M€
ull. Arg. in DC., Prodr. Jard. Bot. Rio de Janeiro 3: 198, t. 21 (1922); Schultes, Bot.
15(2): 1076 (1866); Pax, Pflanzenr. 147, I: 21 (1910); Mus. Leafl. Harvard Univ. 17: 25 (1955); Radcl.-Sm., Gen.
McVaugh, Bull. Torrey Bot. Club 72: 271, figs. 1–24 Euphorb.: 292 (2001).
(1944); Webster, J. Arnold Arb. 48: 340 (1967); Dehgan
& Webster, Univ. California Publ. Bot. 74: 35, t. 1–33 Monoecious trees; laticifers non-articulate,
(1979); Radcliffe-Smith, Fl. Trop. E. Afr., Euphorb. 1: branches sometimes with viscid reddish latex;
343, figs. 65–67 (1987).
indumentum simple. Leaves palmately com-
Monoecious (dioecious or gynodioecious) trees, pound; leaflets entire; petioles with 2 apical
shrubs, or herbs often with thickened caudices or glands; stipules gland-tipped or obsolete. Inflor-
rhizomes; stems with articulate, inarticulate, or escences terminal or subterminal, bisexual,
idioblastic laticifers producing yellowish to red cymose-paniculate; bracts entire, deciduous. Sta-
latex; indumentum simple, sometimes glandular. minate flowers pedicellate; calyx cupular-trun-
Leaves petiolate to subsessile, simple to palmately cate, the 5 sepals represented by minute teeth;
3–7-lobed or divided, entire to serrate, lacking petals 5, distinct, imbricate, much longer than
paired basal glands; stipules entire to dissected, calyx, pubescent on both faces; disk segments 5;
sometimes glandular or spinose (obsolete). stamens 7–10, filaments biseriate, the inner
Inflorescences axillary to terminal, often long- longer; anthers basifixed-cordate, introrse or
pedunculate, of simple to paniculate cymes, latrorse; pollen grains spheroidal, inaperturate,
sometimes solitary and axillary; pistillate flowers exine with Croton pattern; pistillode usually 0.
central in bisexual cymules. Staminate flowers Pistillate flowers subsessile, calyx and petals as
Euphorbiaceae 167
in the staminate; disk dissected into 5 segments; is here construed to comprise several genera that
ovary 2-locular, pubescent; ovules anatropous, formerly were included in tribe Codiaeae (Sago-
inner integuments thick, vascularized; stylodia tia, Acidocroton with Ophellanthe) and tribe
short, stigmas lobate. Fruits drupaceous or sub- Aleuritideae (Sandwithia). The close relationship
dehiscent, verrucose, exocarp 1 cm thick. Seeds between Sagotia and Sandwithia had first been
ecarunculate, endosperm oleaginous. recognized by Secco (1988).
Three neotropical spp., from Venezuela and
Amazonian and coastal Brazil. KEY TO THE GENERA OF CROTONEAE
1. Indumentum simple 2
210. Vaupesia R.E. Schultes – Indumentum stellate and/or lepidote 4
Vaupesia R.E. Schultes, Bot. Mus. Leafl. Harvard Univ. 17: 2. Inflorescences axillary; stipules transformed into
27, t. 12 (1955); Murillo & Franco, Euforb. Reg. Arara- spines 213. Acidocroton
cuara 158, fig. 48 (1995); Radcliffe-Smith, Gen. Euphorb.: – Inflorescences mostly terminal; stipules not trans-
290 (2001). formed into spines 3
3. Staminate sepals 2–3, completely connate in bud; disk
Monoecious trees; latex of trunk whitish, scanty, +; stylodia connate into a common style with distal
of branches reddish. Leaves with conspicuous bifid or entire style branches 211. Sandwithia
basal glands; stipules deciduous. Inflorescences – Staminate sepals 5(6), distinct; disk 0; stylodia distinct
or nearly so, bifid 212. Sagotia
terminal, bisexual, paniculate; bracts entire, per- 4. Stamen filaments not inflexed in bud; stylodia deeply
sistent. Staminate flowers pedicellate; sepals 5, bifid 215. Brasiliocroton
imbricate, entire; petals 5, entire; disk segments – Stamen filaments distinctly inflexed in bud;
5, at base of staminal column; stamens 8, fila- stylodia bifid to multifid 5
ments connate into a column; anthers biseriate; 5. Stamens (3)8–20(400); receptacle usually pilose; seeds
pollen grains spheroidal, inaperturate, exine with terete to compressed; stylodia various 216. Croton
hexagonal pillars; pistillode 0. Pistillate flowers – Stamens 8–15; receptacle usually glabrous; seeds qua-
drangular; stylodia multifid 214. Astraea
pedicellate; sepals 5, distinct, imbricate, margins
fimbriate; petals 5, entire; disk 5-lobed, reddish; 211. Sandwithia Lanj.
ovary 3-locular; stylodia proximally connate,
bifid. Fruits capsular, thick-walled; columella Sandwithia Lanj., Kew Bull. 1932: 184 (1933); Jablonski,
Mem. N. Y. Bot. Gard. 17: 152 (1967); Secco, Bull. Mus.
not persistent. Seeds ecarunculate, subprolate, Par. Emilio Goeldi Bot. 3: 157 (1987), 4: 177 (1988); Fl.
compressed, dorsally carinate, with conspicuous Venez. Guayana 5: 217, fig. 203 (1999).
hilum.
One sp., V. cataractarum R.E. Schultes, from Monoecious or dioecious trees; latex reddish;
the upper Rio Negro region in SE Colombia and indumentum simple. Leaves alternate, petiolate
adjacent Brazil. (slightly pulvinate at apex), entire, eglandular;
stipules deciduous. Inflorescences terminal, flow-
8. TRIBE CROTONEAE Dumort. (1829). ers in cymose racemes or clusters; bracts decidu-
ous. Staminate flowers pedicellate; sepals 2 or 3,
Monoecious (dioecious) trees, shrubs, or herbs; completely connate in bud; petals 3 or 4, distinct,
laticifers non-articulated, latex clear to yellowish imbricate, s.t. reduced; disk segments 2–4, recep-
or reddish, sometimes 0; indumentum simple, tacle pilose; stamens 15–25, filaments distinct;
lepidote, or stellate; leaves simple to palmately pollen grains spheroidal, inaperturate, exine
lobed with or without basal paired glands; granular, with Croton pattern, pillars smooth;
stamens 3–400; anthers extrorse; pollen grains pistillode 0. Pistillate flowers pedicellate; sepals
trinucleate, spheroidal, inaperturate, with Croton 4, connate, tips imbricate in bud, erect; petals
pattern; ovules with thick vascularized inner inte- 4, minute, deciduous; disk annular; ovary 3-locu-
guments; stylodia distinct or basally connate, lar, pilose; stylodia connate into a common style
bifid to multifid; seeds mostly carunculate; coty- with bifid or entire style branches. Fruits capsu-
ledons mostly broader than radicle. lar; columella not persistent? Seeds carunculate,
Following the molecular work of Berry et al. testa smooth.
(2005) and Wurdack et al. (2005), tribe Crotoneae Two spp., Amazonian South America.
168 G.L. Webster
212. Sagotia Baill. Pistillate flowers pedicellate; sepals 5 or 6,

Sagotia Baill., Adansonia I, 1: 53 (1860; nom. cons.);
imbricate, persistent in fruit and sometimes
Jablonski, Mem. N. Y. Bot. Gard. 17: 151 (1967); Secco, accrescent; petals rudimentary; ovary 3-locular,
Acta Amazonica 15 (1–2, suppl.): 81 (1985); Rev. Gen. glabrous or pubescent; ovules anatropous, inner
Anomalocalyx, etc.: 99, figs. 29–34 (1990); Webster, Ann. integuments thick, vascularized; stylodia unlobed
Missouri Bot. Gard. 81: 107 (1994). or bifid, dilated and petaloid. Fruits capsular;
Monoecious trees or shrubs; latex clear, yellow- sepals 5 or 6, distinct, imbricate, entire, persistent
ish, or reddish; indumentum simple. Leaves peti- in fruit; columella persistent in fruit. Seeds trigo-
olate (pulvinate), entire, eglandular; stipules nous, carunculate, testa smooth.
deciduous, leaving annular scars. Inflorescences Thirteen spp., of which ten in sect. Acidocro-
terminal, racemoid or thyrsoid-paniculate, uni- ton are confined to the West Indies, whereas the
sexual or bisexual; pistillate flowers proximal; three of sect. Ophellantha occur from Mexico to
bracts deciduous. Staminate flowers pedicellate; Colombia. Radcliffe-Smith (2001) rejects the
sepals 5 (6), distinct, imbricate; petals 5 (7), dis- combining of Ophellantha with Acidocroton by
tinct, imbricate, longer than the sepals; disk not Webster (1994). However, Radcliffe-Smith’s char-
evident; stamens 20–30, filaments distinct; acters based on higher stamen number and lower
anthers subsessile, basifixed, latrorse, connective carpel number in Ophellantha do not hold, and
broad; pollen grains spheroidal, inaperturate, the only remaining distinction is in the bifid
exine granular, with Croton pattern, pillars stylodia and somewhat more accrescent pistillate
acute or spinose; pistillode 0. Pistillate flowers sepals of Ophellantha. Acidocroton and Ophel-
pedicellate; sepals 5 (6), distinct, recurved, persis- lantha form a monophyletic group (Berry et al.
tent and accrescent in fruit; petals and disk 0; 2005), and there is little to gain in generic subdi-
ovary 3-locular, pilose; ovules pachychalazal, vision.
inner integuments thick, vascularized; stylodia
distinct or early so, bifid. Fruits capsular; colu- 214. Astraea Klotzsch
mella not persistent. Seeds ellipsoidal, caruncu-
Astraea Klotzsch, Arch. Naturgesch. 7: 194 (1841); Baill.,
late, testa smooth. Étude Gén. Euphorb.: 363 (1858, as section); Caruzo &
Two spp., Central America and northern Cordeiro, Hoehnea 34: 572 (2007); Cavalari De-Paula
South America, in lowland rainforest. et al., Pl. Syst. Evol. 292: 1–14 (2011), floral morph.
213. Acidocroton Griseb. Monoecious subshrubs or herbs; indumentum

stellate; laticifers non-articulated, latex scanty
Acidocroton Griseb., Fl. Br. W. Ind.: 42 (1859; nom. cons.); or apparently 0. Leaves deeply palmately lobed
M€ull. Arg. in DC., Prodr. 15(2): 1042 (1866); Pax, Pflan-
zenr. 147, I: 13 (1910); Urban, Symb. Ant. 7: 513 (1913);
(simple); stipules entire or dissected, sometimes
Fawc. & Rend., Fl. Jam. 4: 315, fig. 104 (1920); Alain, Fl. glandular, persistent. Inflorescences terminal,
Cuba 3: 73 (1953), Fl. Española 4: 69 (1986); Webster, bisexual, racemoid; pistillate flowers solitary at
Ann. Missouri Bot. Gard. 81: 107 (1994); Fernández- proximal nodes; bracts entire, persistent, eglandu-
Alonso & Jaramillo-Mejı́a, Caldasia 17: 389 (1995); lar. Staminate flowers pedicellate; sepals 5, imbri-
cate; petals 5, imbricate, densely pilose at the basis;
Ophellantha Standl. (1924).
receptacle glabrous; disk 5-lobate; stamens 8–15,
Monoecious shrubs; latex not recorded; indu- distinct; filaments glabrous, inflexed in bud;
mentum simple. Leaves petiolate to subsessile, anthers basifixed, muticous; pollen grains spheroi-
entire, eglandular; stipules transformed into dal, with Croton pattern; pistillode 0. Pistillate
spines. Inflorescences unisexual, axillary and glo- flowers subsessile, elongating in fruit; sepals 5,
merular or pistillate flowers solitary and subter- valvate, glandular-dentate; disk segments 5;
minal. Staminate flowers pedicellate; sepals 5–6, ovary glabrous or hispidulous with stellate or
imbricate; petals 5–7, distinct, imbricate, gla- simple hairs; stylodia distinct, multifid. Fruits
brous, longer than the sepals; disk annular, capsular; columella persistent. Seeds quad-
pubescent; stamens 20–50(100), filaments dis- rangular, rugose, carunculate, caruncle reniform-
tinct; anthers introrse, with enlarged apiculate peltate, testa rugulose-costate. 2n ¼ 18.
connective; pollen grains spheroidal, inapert- About 10 spp. in the Neotropics. This
urate, with Croton pattern; pistillode obsolete. genus has been treated as a section of Croton by
Euphorbiaceae 169
nearly all 20th century authors. Webster (1967)

remarked that Astraea had the best claim of any
Croton section to generic status, and molecular
studies (Berry et al. 2005) have resoled Astraea
as the sister group of Acidocroton in the grade
leading from Jatropha to Croton s.str.
215. Brasiliocroton P.E. Berry & I. Cordeiro

Brasiliocroton P.E. Berry & I. Cordeiro, Syst. Bot. 30: 357,
fig. 2 (2005).
Monoecious tree; indumentum stellate. Leaves
with two stipitate glands at the junction with the
petiole; stipules filiform, deciduous. Inflores-
cences terminal bisexual panicles, the distal flow-
ers pistillate and opening earliest. Staminate
flowers: sepals 5(6), valvate, slightly connate at
base; petals 5(6); disk of 5 distinct antesepalous
lobes; stamens (20–)25(30); filaments erect in
bud; anthers bilobed, latrorse; pollen inapertu-
rate with Croton pattern, the subunits striate.
Pistillate flowers: sepals 5(6), valvate, basally
connate for 1/2 to 2/3 their length; petals 0; disk
5-lobate; ovary 3(4)-locular; ovule 1 per locule;
stylodia 3, deeply bifid and basally connate into a
short column. Fruits schizocarpic, the exocarp
separating from the woody cocci; columella
8–12 mm long. Seeds ellipsoid, dorsally angled,
with a small caruncle.
A single sp., B. mamoninha P.E. Berrry & Fig. 39. Euphorbiaceae-Crotonoideae. Croton arnhemi-
cus. A Flowering branchlet. B Undersurface of leaf. C
I. Cordeiro, lowland forests in N and E Brazil. Base of leaf lamina showing extrafloral nectaries. D Node
with stipules. E Inflorescence with pistillate flowers in
216. Croton L. Fig. 39 lower half and staminate flowers in upper. F Pistillate
flower. G Staminate flower. H, I Fruits. J Seed. (P.I. Forster
Croton L., Sp. Pl. 2: 1004 (1753); M€
ull. Arg. in DC., Prodr. 2003; drawn by W. Smith)
15(2): 512 (1866); Ferguson, Rep. Missouri Bot. Gard. 12:
33, t. 4–31 (1901); Hutchinson, Fl. Trop. Afr. 6(1): 746
(1912); Gagnepain, Fl. Indochine 5: 256, figs. 28, 29 (1925); Crotonopsis Michx. (1803).
Leandri, Ann. Inst. Bot.-Geol. Colon. Marseille V, 7(1): 1 Julocroton Mart. (1837, nom. cons.).
(1939); Webster, J. Arnold Arb. 48: 358, fig. 2 (1967); Eremocarpus Benth. (1844).
Leandri, Adansonia II, 10: 191 (1970); Liogier, Fl. Espa- Colobocarpos Esser & van Welzen (2001).
ñola 4: 108 (1986); Chakrabarty & Balakrishnan, Bull. Bot.
Survey India 34: 1–88, figs. 1–16 (1992); Webster, Novon Monoecious (dioecious) trees, shrubs, or herbs;
2: 270 (1992), Taxon 42: 793 (1993); Ann. Missouri Bot. laticifers non-articulated (0), latex clear to red or
Gard. 81: 111 (1994), Fl. Nicaragua 1: 864–875 (2001), yellow, sometimes resinous; indumentum stel-
Contr. Univ. Michigan Herb. 23: 353, fig. 1 (2001); Martı́-
late or lepidote; sessile or stipitate glands some-
nez Gordillo, Contr. Herb. UNAM 2: 9 (1995); Berry, Fl.
Venez. Guayana 5: 111, figs. 108–125 (1999); Radcl.-Sm., times present. Leaves alternate (opposite),
Gen. Euphorb.: 319 (2001); Berry et al., Amer. J. Bot. 92: simple or lobate, petiolate (sessile), entire, den-
1520–1534 (2005) (mol. systematics); Forster, van Ee & tate or serrate, usually 2-glandular at the junc-
Berry, Syst. Bot. 35: 151–167 (2010), mol. syst. sect. tion of the petiole with the blade; stipules entire
Heptallon; Riina et al., Syst. Bot. 34: 360–374 (2009),
or dentate to dissected, persistent or deciduous,
mol. syst. sect. Cyclostigma; Riina et al., Taxon 59:
1147–1160 (2010), mol. syst. sect. Luntia; van Ee et al., sometimes 0. Inflorescences terminal (axillary),
Taxon 60: 791–823 (2011), rev. class. of New World taxa. usually bisexual, mostly racemoid with solitary
Tridesmis Lour. (1790). pistillate flowers at proximal nodes and
170 G.L. Webster
glomerules of staminate flowers distally, stami- 216b. Croton subg. Moacroton (Croizat) van Ee &
nate flowers sometimes 1 per bract; bracts entire P.E. Berry, Bot. Rev. 74: 158 (2008).
to dissected, sometimes glandular. Staminate
Moacroton Croizat (1945); Borhidi, Acta Bot. Acad. Sci.
flowers pedicellate; sepals (4) 5 (6), essentially Hung. 36: 7 (1990), rev.
distinct to connate, imbricate to valvate; petals Cubacroton Alain (1960).
usually 5(0), distinct, imbricate, glabrous or Croton sect. Corylocroton G.L. Webster (1993).
pubescent; disk entire or dissected; receptacle
usually pilose (glabrous); stamens (3–)8–20 Indumentum usually lepidote; stylodia bifid or
(400), distinct, filaments usually inflexed in simple; sepals of pistillate flowers usually connate
bud, glabrous or pubescent; anthers extrorse in at the base but not valvate.
bud; pollen grains spheroidal, inaperturate, About 28 or more spp., distributed in
exine with Croton pattern, pillars rounded or North America, Mesoamerica, the Caribbean,
angular, mostly sulcate; pistillode 0. Pistillate and South America, where they generally show
flowers sessile to pedicellate; sepals (4)5–7 a preference for mesic habitats. In addition to
(10), distinct to connate, imbricate, entire to the taxa synonymized above, the group com-
dentate, usually persistent in fruit and often prises a number of South American species of
accrescent; petals 0 (+); disk annular (dissected), Croton hitherto not assigned to a section; they
sometimes with staminodia [reduced petals?]; have been revealed as a basal clade of Croton
ovary 3(1)-locular; ovules anatropous, inner by van Ee et al. (2008). The six spp. formerly
integuments thick, vascularized; stylodia distinct included in Moacroton are endemic to Cuba,
or basally connate, bifid to multifid. Fruits cap- where they grow on serpentine outcrops.
sular (indehiscent); columella persistent, usually
slender. Seeds terete to compressed, carunculate 9. TRIBE CODIAEAE (Pax) Hutch. (1969).
(arillate), testa smooth. 2n ¼ 20, 28, 64.
Monoecious or dioecious trees or shrubs; laticifers
Over 1,200 spp., circumglobal in warm-
non-articulated, latex clear or sometimes reddish;
temperate to tropical regions, 712 spp. recognized
indumentum simple or malpighiaceous; leaves
for the New World. Croton is the second
alternate, pinnately veined or triplinerved, usually
largest genus of Euphorbiaceae and is highly
without laminar glands; stipules deciduous or
diverse morphologically and cytologically. Not
obsolete; inflorescences terminal or axillary, race-
surprisingly, many generic segregates had been
moid-thyrsoid or paniculate; staminate sepals 4–6,
proposed in the past but have been brought
distinct or connate, imbricate or valvate; petals 5,
back to Croton again, and those upheld until
distinct, usually imbricate; disk dissected or lobed;
recently (Crotonopsis, Eremocarpus, Julocroton,
stamens (5–)10–100 or more, distinct or basally
Moacroton) have been reduced on the basis
connate; pollen grains binucleate, spheroidal, ina-
of extensive molecular studies. There is also
perturate, exine with Croton pattern; pistillode 0;
evidence pointing to a New World origin of
pistillate sepals 4 or 5, usually imbricate, some-
Croton, with a subsequent divergence in the Old
times accrescent; disk mostly annular; ovary
World, followed, back in the New World, by the
mostly 3-locular; ovules anatropous, inner integu-
principal morphological diversification of the
ments thick, vascularized; stylodia unlobed to
genus (Berry et al. 2005).
bipartite; fruits capsular; columella usually persis-
tent; seeds carunculate or ecarunculate, testa
Two subgenera:
sometimes fleshy; endosperm present.
This pantropical but predominantly Asiatic
216a. Croton subg. Croton
tribe shows the greatest generic diversity in the
Indumentum mostly stellate; stylodia usually Crotonoideae. Further study since the treatment
multifid; sepals of pistillate flowers usually valvate. of Webster (1994) has led to the recognition
Contains the bulk of the Croton species, of four subtribes. The geographic distribution of
which generally show a preference for xeric the Codiaeae is distinctive in its primarily South
habitats. American/Asian concentration (10 genera), with a
Euphorbiaceae 171
secondary concentration in Australasia (4 genera), glands; stipules entire, persistent, sometimes

and a single genus (Pantadenia) disjunct from minute. Inflorescences terminal or axillary, usu-
southeast Asia to Madagascar; continental Africa ally bisexual, racemoid; bracts entire, eglandular,
is entirely excluded. The treatment of the neotropi- usually persistent. Staminate flowers pedicellate;
cal taxa adopted here is indebted to the careful sepals 5, distinct, imbricate; petals 5, yellow to
work of Secco (1990) on the Amazonian taxa of orange or red, distinct, imbricate, usually longer
Codiaeae. than calyx; disk dissected or urceolate; stamens 3
(5), filaments connate into a column; anthers
KEY TO THE SUBTRIBES OF CODIAEAE extrorse, basifixed, connective enlarged and
1. Petals 0 or rudimentary in pistillate flowers
often apiculate or bifid; pollen grains spheroidal,
9d. Codiaeinae inaperturate, exine with Croton pattern, pillars
– Petals well-developed in pistillate flowers 2 spinulose; pistillode 0. Pistillate flowers pedicel-
2. Stamens 3 (5), filaments all connate into a column late; sepals 5, imbricate, entire, persistent in fruit,
9a. Trigonostemoninae sometimes accrescent; petals 5, yellow to orange
– Stamens 7 or more, filaments not all connate into a or red, distinct, imbricate, mostly glabrous; disk
column 3 annular or 5-lobed; ovary 3-locular, glabrous or
3. Inflorescences mostly terminal; monoecious (except
Hylandia); anthers extrorse; seeds carunculate or fruit pubescent; stylodia distinct or nearly so, unlobed
drupaceous 9b. Baloghiinae to bifid or bipartite. Fruits capsular, cocci thin-
– Inflorescences mostly axillary; anthers mostly introrse; walled; columella not persistent. Seeds ecaruncu-
seeds ecarunculate, fruits capsular 9c. Ostodeinae late, testa smooth.
Nearly 100 spp., distributed from India and
9a. SUBTRIBE TRIGONOSTEMONINAE (G.L. Webster) China south and east to the Philippines, northern
G.L. Webster1 Australia (Queensland), New Guinea, and Fiji.
Monoecious; latex reddish; indumentum simple;
anthers with enlarged connective; stylodia bifid 9b. SUBTRIBE BALOGHIINAE G.L. Webster1
or twice bifid; fruits capsular. Seeds ecarunculate. Dioecious (monoecious); basal laminar glands
A monogeneric paleotropical subtribe with present or 0; inflorescences terminal or subtermi-
nearly 100 species. In the molecular analysis nal; pistillate flowers petaliferous; petals longer
(Wurdack et al. 2005), Trigonostemon is resolved than sepals; ovary often pubescent; columella
as the sister group to the rest of the genera form- usually persistent.
ing clade C2. Four Australasian genera. The subtribe is
possibly related to subtribe Cocconerioninae in
217. Trigonostemon Blume the Ricinocarpeae.
Trigonostemon Blume, Bijdr. Fl. Ned. Ind. 600 (1825; nom.
cons.); M€
ull. Arg. in DC. Prodr. 15(2): 1105 (1866); Pax & KEY TO THE GENERA OF BALOGHIINAE
K. Hoffm., Pflanzenr. 147, III: 85 (1911); Gagnepain, Fl.
1. Indumentum stellate; inflorescences covered with resin
Indochine 5: 309, fig. 35, 3–10 (1925); Airy Shaw, Kew
221. Alphandia
Bull. Add. Ser. 4: 201 (1975), Kew Bull. 35: 352 (1981);
Webster, Ann. Missouri Bot. Gard. 81: 108 (1994); Milne, – Indumentum mainly simple; inflorescences not
Kew Bull. 50: 23, figs. 1–3 (1995); Radcl.-Sm., Gen. resinous 2
Euphorb.: 307 (2001). 2. Staminate calyx truncate; fruits drupaceous
Kurziodendron Balakr. (1966). 220. Fontainea
– Staminate calyx distinctly lobed; fruits capsular 3
Monoecious shrubs or trees; reddish latex some- 3. Ovary 3-locular; petals glabrous adaxially
times present; indumentum simple. Leaves alter- 218. Baloghia
nate or opposite, entire, without paired basal
1
Subtr. Baloghiinae G.L. Webster, subtr. nov.: plantae dioi-
1
Subtr. Trigonostemoninae (G.L. Webster) G.L. Webster, cae inflorescentiis terminalibus, staminibus 10–100, liberis,
stat. nov., based on Tribe Trigonostemoneae G.L. Webster, antheris extrorsis, seminibus carunculatis vel ecarunculatis.
Taxon 24: 599 (1975). Type: Trigonostemon Blume. Type: Baloghia Endlicher.
172 G.L. Webster
– Ovary 2-locular; petals sericeous abaxially and abaxially; disk of 5 lobes or segments; ovary
219. Hylandia 2-locular, densely hispid-sericeous; stylodia bifid,
branches somewhat dilated. Fruits indehiscent,
218. Baloghia Endl. ribbed, exocarp somewhat fleshy. Seeds spheroi-
Baloghia Endl., Prodr. Fl. Norf.: 84 (1833); Airy Shaw, dal, apparently ecarunculate, testa hard, smooth.
Kew Bull. 35: 598 (1980); McPherson & Tirel, Fl. Nouv.- A single sp., H. dockrillii Airy Shaw, tropical
Caléd. 14(1): 43–72, t. 8–13 (1987); Hyland & Whiffin, rainforests of Queensland. Hylandia appears to
Austral. Trop. Rain For. Trees 2: 119 (1993).
be closely related to the more widely distributed
Monoecious or dioecious trees or shrubs; laticifers genus Baloghia.
non-articulated, latex yellowish to reddish; indu-
mentum simple or 0. Leaves alternate (opposite), 220. Fontainea Heckel
pinnately veined, brochidodromous, basal laminar Fontainea Heckel, Études sur Fontainea pancheri (1870);
glands mostly 0 [+ on margins near base]; stipules Baillon, Adansonia I, 11: 80 (1873); Airy Shaw, Kew Bull.
0. Inflorescences terminal, thyrsoid, unisexual; 35: 632 (1980); Jessup & Guymer, Austrobaileya 2:
bracts entire, persistent, uniflorous. Staminate 112–125, figs. 1–6 (1985); McPherson & Tirel, Fl. Nouv.-
Caléd. 14(1): 74–78 (1987); Forster, Austrobaileya 5:
flowers pedicellate; sepals (4)5(6), slightly connate 29–37, figs. 1–3 (1997); Forster & van Welzen, Blumea
at base, imbricate; petals (4)5(6), white, distinct, 44: 101, fig. 2 (1999).
usually longer than sepals; disk annular or dis-
sected (0); stamens (10–)40–50(100), distinct or Dioecious (monoecious) trees or shrubs; latex
connate on a convex receptacle; anthers dorsi- reddish to brown; indumentum simple, scanty.
fixed, extrorse; pollen grains spheroidal, inapertu- Leaves alternate (opposite), usually with basal or
rate, sexine with Croton pattern, pillars obtuse and subbasal marginal glands; stipules 0. Inflores-
slightly sulcate; pistillode 0. Pistillate flowers pedi- cences terminal, sometimes axillary, compound
cellate; sepals (4)5(6), imbricate, entire, persistent cymose; bracts entire, inconspicuous. Staminate
in fruit; petals (4)5(6), pubescent adaxially; disk flowers pedicellate; sepals pubescent, connate
annular, sometimes lobed; ovary 3(4)-locular, into a shallowly 3–6-toothed or angled cup; petals
sericeous; ovules anatropous, inner integuments 5, white, distinct, densely pubescent, exserted
very thick, vascularized; stylodia bifid to multifid. well beyond the calyx; disk obscure; receptacle
Fruits capsular; columella persistent. Seeds sphe- densely woolly; stamens 18–32 (40), filaments
roidal or ellipsoidal, mostly carunculate, testa distinct or basally connate; anthers dorsifixed,
smooth, exotesta parenchymatous. extrorse, connective sometimes apiculate; pollen
Fifteen spp., Australasia, 12 of them endemic grains spheroidal, inaperturate, exine with Cro-
to New Caledonia. ton pattern, pillars acute, sulcate; pistillode 0.
Pistillate flowers pedicellate; sepals connate,
219. Hylandia Airy Shaw dehiscing irregularly, deciduous in fruit; petals
5, white, distinct, exserted from calyx; disk annu-
Hylandia Airy Shaw, Kew Bull. 29: 329 (1974); ibid. 35:
lar, glabrous; ovary (2)3(6)-locular, pubescent;
643, fig. 4 (1980); Hyland & Whiffin, Austral. Rain For.
Trees 2: 140 (1993). ovules anatropous, inner integuments thick, vas-
cularized; stylodia bifid. Fruits drupaceous; endo-
Dioecious trees; latex viscid, pith reddish; indu- carp bony, angled, sometimes beaked, smooth or
mentum simple. Leaves with or without raised rugose. Seeds ecarunculate.
glands at junction with petiole; stipules 0. Inflor- Nine or more spp., distributed from Australia
escences terminal, thyrsoid-paniculate; bracts not to New Guinea, New Caledonia, and Vanuatu.
evident. Staminate flowers pedicellate; sepals 5,
distinct, imbricate, unequal; petals 5, white, dis- 221. Alphandia Baill.
tinct, imbricate, obovate to spathulate, pubescent Alphandia Baill., Adansonia I, 11: 86 (1873); Pax, Pflan-
adaxially, appressed-sericeous abaxially; disk zenr. 147, III: 22 (1911); McPherson & Tirel, Fl. Nouv.-
segments 5, massive, glabrous; stamens 10–16, Caléd. 14(1): 86–90, fig. 17 (1987); Airy Shaw, Kew Bull.
filaments basally connate; anthers extrorse; pol- Add. Ser. 8: 27 (1980).
len grains inaperturate, with Croton pattern; pis- Monoecious trees or shrubs; stems with yellowish
tillode 0. Pistillate flowers with pedicel articulate or reddish resinous latex, branches and leaves
near base; sepals 5, distinct, imbricate, entire; with resinous exudate; indumentum stellate.
petals 5, distinct, whitish, pubescent adaxially Leaves with paired glands at base; stipules 0.
Euphorbiaceae 173
Inflorescences terminal or subterminal, unisexual – Staminate calyx distinctly lobed; fruiting calyx not
or bisexual, thyrsoid-paniculate; bracts entire, accrescent 2
eglandular. Staminate flowers pedicellate; sepals 2. Indumentum partly malpighiaceous; flowers subsessile,
monoecious 227. Pausandra
5, connate, not closed in bud; petals 5, yellowish,
– Indumentum simple; pistillate flowers distinctly pedi-
distinct, imbricate, pubescent abaxially; disk cellate; seeds ecarunculate 3
segments 5, distinct or confluent; stamens 20–35, 3. Leaf blades dentate, with basal paired glands; anthers
inner filaments connate at base, outer distinct, not glandular 222. Ostodes
apically deflexed; anthers extrorse; pollen grains – Leaf blades mostly entire, lacking basal paired glands 4
spheroidal, inaperturate, with Croton pattern; pis- 4. Anthers glandular; leaf blades with dispersed laminar
glands, inflorescences leaf-opposed 226. Pantadenia
tillode 0. Pistillate flowers pedicellate; sepals 5,
– Anthers not glandular; leaf blades with or without dis-
connate at base, not imbricate; petals 5, distinct, persed laminar glands; inflorescences terminal or axil-
imbricate, deciduous; disk annular; ovary 3-locu- lary 5
lar, pubescent and resinous; ovules anatropous, 5. Staminate and pistillate flowers externally glabrous;
inner integuments thick; stylodia bifid. Fruits cap- stamens 20+, inserted on convex receptacle; staminate
sular, thin-walled; columella persistent. Seeds disk and ovary glabrous; stylodia 6-branched
224. Anomalocalyx
oblong, apiculate, carunculate or ecarunculate;
– Staminate and pistillate flowers externally pilose;
cotyledons much broader than radicle. stamens 7–16, inserted on an almost plane receptacle;
Three spp., Melanesia, 1 in New Guinea and staminate disk pilose, ovary densely pilose; stylodia
2 in New Caledonia/Vanuatu. Pax (1911) referred 10–12-branched 225. Dodecastigma
Alphandia to the Codiaeinae, where it could be
placed except for the stellate indumentum.
Although the stellate indumentum of Alphandia 222. Ostodes Blume
suggests a relationship with Ricinocarpeae sub-
tribe Cocconeriinae, the pollen exine (Lobreau- Ostodes Blume, Bijdr.: 619 (1825); M€
ull. Arg. in DC. Prodr.
15(2): 1114 (1866); Gagnepain, Fl. Indochine 5: 322, fig.
Callen in McPherson and Tirel 1987) shows the 33, 7–9 (1925); Airy Shaw, Kew Bull. 20: 409 (1967), 35:
typical Crotonoid sculpturing of the Baloghiinae 334 (1981); Grierson & Long, Fl. Bhutan 1(3): 795, fig. 49
and other Codieae, which is very different from p–r (1987); Chakrabarty & Balakr., Bull. Bot. Surv. India
the reduced sculpturing of the Cocconeriinae. It 27: 259 (1987).
seems possible that Alphandia may provide a Dioecious trees; latex not recorded; indumentum
connecting link between the Baloghiinae and simple. Leaves crenate-serrate, with 2 or more
Cocconeriinae, and that these Australasian taxa adaxial basal glands at junction with petiole; sti-
may belong to a single clade. pules gland-tipped, entire, deciduous. Inflores-
cences axillary, unisexual, the staminate thyrsoid,
9c. SUBTRIBE OSTODEINAE G.L. Webster1 pistillate paniculate; bracts entire, eglandular, per-
Dioecious; leaves with or without basal paired sistent. Staminate flowers pedicellate; sepals 5,
glands or dispersed laminar glands; stamens connate, imbricate, glabrous; petals 5 or 6, white
5–35, filaments distinct; anthers mostly introrse; or pinkish, imbricate, concave, glabrous, longer
stylodia bifid or multifid; seeds ecarunculate than sepals; disk segments 8–12, massive, puberu-
(except in Pausandra). lent; stamens 30–35, filaments distinct, pilose;
A heterogeneous subtribe of six genera, three anthers dorsifixed, introrse; pollen grains spheroi-
paleotropical and three neotropical. Pausandra in dal, inaperturate, with Croton pattern, pillars
particular appears aberrant and may prove not to rounded; pistillode 0. Pistillate flowers pedicellate,
belong to this subtribe. articulate near base; sepals 5, connate, imbricate,
entire, deciduous in fruit; petals 5, distinct, imbri-
KEY TO THE GENERA OF OSTODEINAE cate, longer than sepals; disk annular-cupulate,
1. Staminate calyx cupular, barely lobed; fruiting calyx puberulent; ovary 3-locular, sericeous; ovules
accrescent 223. Dimorphocalyx anatropous, inner integuments thick, vascular-
ized; stylodia bifid. Fruits capsular, exocarp fleshy,
verrucose, endocarp bony; columella persistent.
1
Subtr. Ostodeinae G.L. Webster, subtr. nov.: plantae dioicae Seeds angular, ecarunculate, exotesta fleshy, tenu-
inflorescentiis terminalibus, staminibus 5–35, liberis, ous, endotesta bony; cotyledons much longer and
antheris plerumque introrsis, seminibus plerumque ecar-
unculatis. Typus: Ostodes Blume.
broader than radicle. 2n ¼ 20.
174 G.L. Webster
A single polytypic sp., O. paniculata Blume, abaxially with embedded glands; stipules entire,
extending from Assam through southeast Asia to deciduous. Inflorescences terminal; staminate
Malaya and Java. Partly because of the jumbled flowers thyrsoid-paniculate, pistillate flowers soli-
treatment of Pax and Hoffmann (1911), the genus tary; bracts entire, eglandular, deciduous. Stami-
has been confused with Paracroton, which now is nate flowers pedicellate; sepals 3 or 4, glabrous,
referred to the Aleuritideae. connate in the bud, opening in 2 lobes at anthesis;
petals 5, distinct, imbricate, adaxially pubescent at
base; disk annular, glabrous; stamens 23–30, fila-
223. Dimorphocalyx Thwaites
ments distinct from the convex receptacle; anthers
Dimorphocalyx Thwaites, Enum. Pl. Zeyl.: 278 (1861); Pax introrse; pollen spheroidal, inaperturate, sexine
& K. Hoffm., Pflanzenr. 147, III: 31, fig. 8 (1911); Airy with Croton pattern, pillars rounded, sulcate; pis-
Shaw, Kew Bull. 23: 123 (1969); Chakrabarty & Balakrish-
nan, Proc. Indian Acad. Sci. 100: 286, figs. 1–4 (1990);
tillode 0. Pistillate flowers pedicellate; sepals 3–5,
Philcox, Fl. Ceylon 11: 107 (1997); Naithani et al., Forest glabrous, connate in bud, splitting into (2) 3 lobes
Fl. Goa 552, t. 118 (1997). at anthesis, persistent in fruit; petals 5, scantily
pilose adaxially at base; disk annular, glabrous;
Dioecious trees or shrubs; latex not evident;
ovary 3-locular, glabrous; stylodia bifid. Fruit cap-
indumentum simple, scanty or 0. Leaves entire
sular, endocarp woody; columella stout, persistent.
or denticulate, without basal glands; stipules
Seeds rounded, carinate, ecarunculate, testa
entire, persistent or deciduous. Inflorescences
slightly rugose.
terminal or subterminal, sometimes axillary,
A single sp., A. uleanus (Pax) Ducke, Amazo-
pedunculate, cymose or racemoid, pistillate some-
nian Brazil, from near Manaus to Amapá.
times reduced to solitary flowers; bracts entire,
Although it closely resembles Dodecastigma in
persistent, eglandular. Staminate flowers pedicel-
habit, Secco notes that it is immediately distin-
late; sepals 5, connate into a dentate or lobed cup;
guishable by its glabrous inflorescences.
petals 5, white, distinct, glabrous, equaling or
longer than the calyx; disk segments 5, glabrous; 225. Dodecastigma Ducke
stamens (5)8–20(100), the inner 3–10 filaments
Dodecastigma Ducke, Notizbl. Bot. Gart. Berlin 11: 343
connate; anthers basifixed or dorsifixed, introrse; (1932); Arq. Jard. Bot. Rio Jan. 6: 58, t. 5 (1933); Sandw.,
pollen spheroidal, inaperturate, exine with Kew Bull. 1950: 134 (1951); Jablonski, Mem. N. Y. Bot.
Croton pattern; pistillode 0. Pistillate flowers ped- Gard. 17: 154 (1967); Secco, Rev. Gen. Anomalocalyx
icellate; sepals 5, distinct, imbricate, entire, et al.: 42, figs. 10–12 (1990); Webster, Ann. Missouri Bot.
persistent and accrescent in fruit; petals 5, Gard. 81: 106 (1994).
distinct, imbricate, shorter than sepals; disk Dioecious trees or shrubs; latex turning reddish;
annular or cupular; ovary pubescent, sometimes indumentum simple and malpighiaceous, flowers
rugose; ovules anatropous, inner integuments pilose. Leaves petiolate (pulvinate), cuspidate,
thick, vascularized; stylodia connate at base (dis- entire with cartilaginous border, abaxially with
tinct), bifid, branches slender. Fruits capsular; small dispersed embedded discoid glands but
columella persistent. Seeds ellipsoidal, ecaruncu- lacking paired basal glands, stipules minute,
late, testa smooth. deciduous. Staminate inflorescences axillary,
Seventeen spp., distributed from India and thyrsoid-paniculate; pistillate inflorescences
Ceylon to the Philippines, New Guinea, and racemoid, terminal and axillary; bracts entire,
northern Australia. inconspicuous, persistent, subtending glomerules
of staminate flowers or solitary pistillate flowers.
224. Anomalocalyx Ducke Staminate flowers pedicellate; calyx 3- or 4-lobed,
Anomalocalyx Ducke, Notizbl. Bot. Gart. Berlin 11: 344 lobes imbricate; petals 3 or 4, green or yellow-
(1932), Arq. Jard. Bot. Rio Jan. 6: 60, figs. 7–9 (1933); green, distinct, imbricate, pilose abaxially; disk
Secco, Rev. Gen. Anomalocalyx et al.: 39, figs. 7, annular-crenulate, pilose; stamens 7–16, fila-
8 (1990); Webster, Ann. Missouri Bot. Gard. 81: 115 ments distinct; anthers introrse; pollen grains
(1994); Radcl.-Sm., Gen. Euphorb.: 338, fig. 43 (2001).
spheroidal, inaperturate, exine with Croton pat-
Dioecious trees; latex white; indumentum mal- tern; pistillode 0. Pistillate flowers long-pedicel-
pighiaceous but flowers glabrous. Leaves petiolate late; sepals (2) 3–4, imbricate, entire, abaxially
(pulvinate), adaxially with paired basal glands, pubescent, deciduous in fruit; petals mostly 3 or
Euphorbiaceae 175
4, green or yellow-green, imbricate, pilose abaxi- etc.: 58, figs. 14–25 (1990); Webster, Ann. Missouri Bot.
ally, subpersistent in fruit; disk annular, pilose; Gard. 81: 105 (1994); Murillo & Franco, Euforb. Reg.
Araracuara: 124, fig. 34 (1995); Radcl.-Sm., Gen.
ovary 3-locular; densely sericeous; ovules anatro- Euphorb.: 296 (2001).
pous, inner integuments thick, vascularized; sty-
lodia multifid. Fruits capsular, endocarp wood; Dioecious trees or shrubs; laticifers non-articu-
columella massive, persistent. Seeds elliptic, ecar- lated, latex reddish; indumentum malpighia-
unculate, testa smooth and mottled. ceous. Leaves with paired basal glands at
Three spp. in Amazonian Brazil and the Gui- junction with petiole; stipules entire, eglandular,
anas. The genus appears to be closely related to subpersistent or deciduous. Inflorescences axil-
Anomalocalyx. lary, spiciform-thyrsoid, staminate flowers in glo-
merules, pistillate solitary at nodes; bracts
226. Pantadenia Gagnep. inconspicuous, eglandular. Staminate flowers
Pantadenia Gagnep., Bull. Soc. Bot. France 71: 873 (1925);
subsessile; sepals 5, distinct, imbricate; petals 5
Airy Shaw, Kew Bull. 23: 122 (1969), 26: 312 (1972); (6), imbricate, basally connate, adaxially villous;
Webster, Ann. Missouri Bot. Gard. 81: 106 (1994); disk extrastaminal, urceolate-lobate, glabrous;
Radcl.-Sm., Gen. Euphorb.: 298 (2001). stamens (3–)5–7, filaments distinct; anthers
Parapantadenia Capuron (1972). introrse; pollen grains spheroidal, inaperturate,
Dioecious trees; latex not recorded; indumentum exine with Croton pattern, pillars distinctly sul-
simple. Leaves petiolate or subsessile, abaxially cate; pistillode 0. Pistillate flowers subsessile;
with dispersed scutelliform glands; stipules subu- sepals 5, imbricate, entire, persistent in fruit;
late, deciduous. Inflorescences leaf-opposed, petals 5, distinct, imbricate, adaxially villous;
the staminate thyrsoid-racemose, pistillate flow- disk urceolate, sometimes lobate, glabrous;
ers mostly solitary; bracts entire, eglandular. Sta- ovary 3-locular, pubescent; ovules anatropous,
minate flowers pedicellate; sepals 5, distinct, inner integuments thick, vascularized; stylodia
imbricate; petals 5, distinct, imbricate, with 3 distinct, bifid. Fruits capsular; columella persis-
apical marginal glands; disk cupular, undulate; tent. Seeds carunculate, testa smooth.
stamens 13–15, filaments distinct; anthers Six neotropical spp., extending from Nicara-
introrse, dorsifixed, connective glandular at gua to Bolivia and southern Brazil (Secco 1990).
apex; pollen grains spheroidal, inaperturate, The status of some species awaits verification.
exine with Croton pattern, lumina large, pillars Pausandra is isolated within neotropical Codiaeae
rounded; pistillode rudimentary. Pistillate flow- by a number of features such as the massive
ers pedicellate; sepals 5 or 6, distinct, imbricate, petiolar glands or stipels, the subsessile flowers,
entire, persistent in fruit; petals 2 or 3, apically and the malpighiaceous indumentum; its place-
glandular; disk annular; ovary 2- or 3-locular, ment needs reevaluation.
hispidulous; stylodia bifid. Fruits capsular or
indehiscent and 1-seeded; columella 3-pronged, 9d. SUBTRIBE CODIAEINAE Pax (1911).
persistent. Seeds spheroidal, ecaruncular, testa Monoecious (dioecious); leaf blades without
smooth. basal laminar glands; inflorescences terminal
Two spp., widely disjunct: Madagascar and or axillary; bracts eglandular; pistillate flowers
Vietnam. Govaerts et al. (2000) and Radcliffe- apetalous or petals rudimentary; seeds mostly
Smith (2001) uphold Parapantadenia as a distinct carunculate; columella persistent.
genus, based on its 2-locular ovary and indehis- Four genera, all paleotropical.
cent 1-seeded fruit. The question of generic status
is problematic, but there is no doubt of a close KEY TO THE GENERA OF CODIAEINAE
relationship between the two species despite the
considerable geographical disjunction. 1. Inflorescences axillary; ovary glabrous 2
– Inflorescences terminal; ovary pubescent 3
2. Staminate flowers apetalous; leaf blades with basal
227. Pausandra Radlk. laminar glands; inflorescences bisexual, long thyrsopa-
Pausandra Radlk., Flora 53: 92, t. 2 (1870); Jablonski, niculate 231. Baliospermum
Mem. N. Y. Bot. Gard. 17: 153 (1967); Secco, Bol. Mus. – Staminate flowers petaliferous; leaf blades without
Par. Emilio Goeldi, Bot. 3: 59 (1987); Huft, Ann. Missouri basal laminar glands; inflorescences unisexual, race-
Bot. Gard. 75: 1115 (1989); Secco, Rev. Anomalocalyx, moid 228. Codiaeum
176 G.L. Webster
3. Pistillate sepals glandular-fimbriate; stipules alternate (subopposite), eglandular; stipules decid-

persistent; seeds carunculate 230. Strophioblachia uous or persistent. Inflorescences terminal [or pis-
– Pistillate sepals not glandular-fimbriate; stipules decid- tillate terminal and axillary], racemoid, sometimes
uous; seeds ecarunculate 229. Blachia
umbellate; bracts inconspicuous or 0 above
basal nodes. Staminate flowers pedicellate; sepals
228. Codiaeum Rumph. ex A. Juss. 4 or 5, distinct, concave, membranous; petals 4
Codiaeum Rumph. ex A. Juss., Euphorb. Tent.: 33 (1824; or 5, distinct, imbricate, shorter than sepals; disk
nom. cons.); Airy Shaw, Kew Bull. Add. Ser. 4: 88 (1975); a convex receptacle; disk segments 5; stamens
8: 62 (1980); A.C. Smith, Fl. Vitiensis Nova 2: 549 (1981); 10–20(40), filaments distinct; anthers basifixed,
McPherson & Tirel, Fl. Nouv.-Caléd. 14(1): 95–101, t. 19
(1987); Howard, Fl. Lesser Antilles 5: 32, fig. 13 (1989);
extrorse, connective broad, muticous; pollen
Radcl.-Sm., Gen. Euphorb.: 301 (2001). grains spheroidal, inaperturate, exine with Croton
pattern; pistillode 0. Pistillate flowers pedicellate;
Monoecious trees or shrubs; laticifers non-artic- sepals 4 or 5, imbricate, entire, usually accrescent
ulated, latex clear; indumentum simple. Leaves and persistent in fruit; petals rudimentary or 0;
alternate (opposite); stipules 0. Inflorescences disk annular; ovary 3–5-locular, glabrous or
axillary, unisexual, racemoid; bracts entire, per- pubescent; ovules anatropous, inner integuments
sistent, eglandular. Staminate flowers pedicellate; thick (18–20 cell layers), vascularized; stylodia
sepals mostly (3–)5(6), distinct, imbricate; petals bifid, branches slender. Fruits capsular; columella
4–6, minute, distinct, imbricate; disk segments persistent or not. Seeds subspheroidal, ecaruncu-
4–6; stamens 10–35, filaments distinct; anthers late; testa smooth. 2n ¼ 36.
basifixed, latrorse, muticous; pollen grains Eleven spp., from India to China and Malesia,
spheroidal, inaperturate, exine with Croton pat- south to the Andaman Islands. Balakrishnan and
tern, pillars smooth and rounded; pistillode 0. Chakrabarty (1989) regard Pantadenia as the
Pistillate flowers pedicellate; sepals mostly 5, dis- closest relative of Blachia.
tinct, imbricate, entire, usually deciduous in fruit;
petals 0; disk cupular; ovary 3-locular, glabrous; 230. Strophioblachia Boerl.
ovules anatropous, inner integuments thick, vas-
Strophioblachia Boerl., Handl. Fl. Ned. Ind. 3(1): 235
cularized; stylodia slender, unlobed. Fruits cap- (1900); Pax, Pflanzenr. 147, III: 35, fig. 10 (1911); Gagne-
sular; columella mostly persistent. Seeds pain, Fl. Indochine 5: 408, figs. 47, 8–14, 48, 1–4 (1926);
carunculate, testa smooth. 2n ¼ 48, 72, 96, 120. Airy Shaw, Kew Bull. 25: 544 (1971); Thin et al., Blumea
Seventeen spp., distributed from Java and 43: 484, t. 1–4 (1998).
Borneo to the Philippines, New Guinea, tropical Monoecious shrubs; latex not recorded; indu-
Australia and New Caledonia. M€ uller (1866) con- mentum simple. Leaves eglandular; stipules
strued Codiaeum in a very broad sense, to include entire, pilose, deciduous or persistent. Inflores-
species of Austrobuxus, Baloghia, Blachia, Fon- cences terminal, unisexual, racemoid, few-flow-
tainea, and Trigonostemon, but Bentham (1880) ered. Staminate flowers pedicellate; sepals 4 or 5,
reconstituted it in the circumscription that is distinct, imbricate; petals 5, white, dentate; disk
accepted at present. segments 5; stamens 15–30, filaments distinct;
anthers extrorse, rounded, muticous; pollen
229. Blachia Baill. grains spheroidal, inaperturate, exine with Cro-
Blachia Baill., Étude Gén. Euphorb.: 385 (1858); Pax, ton pattern; pistillode 0. Pistillate flowers pedicel-
Pflanzenr. 147, III: 36 (1911); Gagnep., Fl. Indochine 5: late; sepals 5, distinct, imbricate, persistent and
410, fig. 48, 5–13 (1926); Airy Shaw, Kew Bull. 23: 121 with glandular-fimbriate margins in fruit; petals
(1969), 26: 223 (1972); Balakrishnan & Chakrabarty, Proc.
Indian Acad. Sci. 99 568, figs. 1–4 (1989); Thin, Tap Chi 0; disk annular; ovary 3-locular, glabrous; ovules
Sinh Hoc 11(3): 16 (1989); Webster, Ann. Missouri Bot. anatropous, inner integuments thick, vascular-
Gard. 81: 107 (1994); Philcox, Fl. Ceylon 11: 105 (1997). ized; stylodia connate at base, bifid. Fruits capsu-
Bruxanellia Dennst. ex Kostel. (1830; nom. rej.). lar, thin-walled; columella persistent. Seeds
subspheroidal, carunculate, testa smooth.
Monoecious shrubs or trees; latex not recorded;
Two spp., southern China to Vietnam, the
indumentum simple, often 0 or nearly so. Leaves
Philippines and Sulawesi.
Euphorbiaceae 177
231. Baliospermum Blume KEY TO THE SUBTRIBES AND GENERA OF RICINO-

CARPEAE
Baliospermum Blume, Bijdr.: 603 (1826); Decaisne in Jac-
quemont, Voy. Inde Atlas 2: 155 (1844); M€ ull. Arg. in DC. 1. Embryo with cotyledons much broader than the radi-
Prodr. 15(2): 1125 (1866); J.J. Sm., Meded. Dept. Landb. 10: cle; pollen exine with reduced pillars; leaves alternate,
599 (1910); Pax & K. Hoffm., Pflanzenr. 147, IV: 24, figs. 6, 7 not ericoid. 10a. Cocconeriinae 2
(1912); Gagnepain, Fl. Indochine 5: 429, figs. 51, 6–19, 52, – Embryo with narrow cotyledons no broader
1–6 (1926); Airy Shaw, Kew Bull. 36: 267 (1981); Long, Fl. than radicle; pollen exine with massive pillars (except
Bhutan 1(3): 809, fig. 50 m–o (1987); Chakrabarty & Balak- Bertya); leaves opposite or verticillate; shrubs or trees,
rishnan, Bull. Bot. Survey India 32: 3, figs. 1–8 (1992); often with reddish latex. 10b. Bertyinae 4
Webster, Ann. Missouri Bot. Gard. 81:108 (1994). 2. Leaves opposite; inflorescences terminal, racemoid;
Monoecious or dioecious shrubs, sometimes stylodia multifid 232. Myricanthe
scandent; latex not recorded; indumentum sim- – Leaves verticillate; staminate inflorescences subcapitate
or flowers axillary; stylodia bifid or twice bifid 3
ple. Leaves pellucid-punctate, glandular-dentate 3. Sepals 4–7; ovary (2)3-locular; stylodia bifid or twice
or serrate, biglandular at base or subbasal on bifid; flowers axillary, solitary 233. Cocconerion
margins; stipules minute, glandular, persistent. – Staminate sepals 0, pistillate sepals 3; ovary
Inflorescences mostly axillary, long-pedunculate, 2-locular; stylodia bifid; staminate inflorescences ter-
thyrsoid-paniculate; bracts entire, eglandular. minal, subcapitate 234. Borneodendron
4. Stylodia dilated, connivent into a pseudo-stigma over
Staminate flowers pedicellate; sepals 4–5(6), dis- the top of the ovary, discoid or rarely with 2 appressed
tinct, imbricate, orbicular; petals 0; disk annular, entire limbs or shallowly 3-lobulate 236. Beyeria
lobed, or segmented; stamens 10–25, filaments – Stylodia 3 spreading and divergent limbs, entire or
distinct; anthers extrorse, muticous, connective 2–5-lobed 5
narow; pollen grains spheroidal, inaperturate, 5. Stylodia entire; flowers in racemes 237. Shonia
exine with Croton pattern; pistillode 0. Pistillate – Stylodia deeply 2(5)-lobed; flowers usually not in
flowers pedicellate; sepals 5 or 6, distinct, imbri- racemes 6
6. Petals 0 or rudimentary; calyx strongly recurved at
cate, entire or denticulate, persistent in fruit and anthesis; disk 0 238. Bertya
often accrescent; petals 0; disk annular; ovary 3 – Petals conspicuous, or if 0 then calyx spreading
(4)-locular, glabrous or pubescent, ovules pachy- at anthesis; disk + 235. Ricinocarpos
chalazal, inner integuments thick, vascularized;
stylodia bifid. Fruits capsular; columella persis- 10a. SUBTRIBE COCCONERIINAE G.L. Webster, subtrib.
tent, distally alate. Seeds carunculate, caruncle nov.1
apical, discoidal; testa smooth. 2n ¼ 44.
Five spp., distributed from the Himalayas and Subtribe Bertyinae sensu Webster (1994), exclud-
Yunnan south to Sumatra and Java. Pax (1912) ing the genus Bertya.
followed M€ uller (1866) in placing Baliospermum Monoecious trees or shrubs; latex reddish or 0;
next to Suregada; however, the pollen study of leaves opposite or verticillate; stipules deciduous
Punt (1962) demonstrated that Baliospermum has or 0; inflorescences terminal or axillary, racemoid
pollen much more similar to genera of Codiaeae or flowers solitary; petals and disk 0; stamens
than to Suregada. 30–100, filaments connate into a column, anthers
extrorse, pubescent; pollen grains with minute cla-
10. TRIBE RICINOCARPEAE M€
ull. Arg. (1864).
vae instead of pillars; ovary 2- or 3-locular; fruits
Monoecious (dioecious); latex reddish, often capsular, columella persistent; seeds carunculate,
scanty or 0; indumentum stellate, sometimes cotyledons broader than radicle.
glandular; sepals 4–6, distinct, imbricate, often Three Melanesian genera; these appear to be
petaloid; petals 5 (0); disk dissected or 0; stamens related to Alphandia, and also to the Bertyinae,
15–100, filaments mostly connate; pollen grains
inaperturate, exine with Croton pattern; stylodia
unlobed to multifid; seeds carunculatae.
This strictly Australasian tribe includes seven 1
Subtrib. Cocconeriinae G.L. Webster, subtrib. nov., plantae
genera. It has certain characters in common with monoicae foliis oppositis vel verticillatis, antheris pilosis,
Alphandia, a possible sister group, which is here filamentis connatis, pollinis grana exinio microclavato,
referred to the Codiaeae. cotyledonibus quam radicula latioribus. Typus: Cocconer-
ion Baillon
178 G.L. Webster
although the phylogenetic connections are not yet 234. Borneodendron Airy Shaw
very clear. In Webster (1994), these genera were Borneodendron Airy Shaw, Kew Bull. 16: 359 (1963),
included in subtribe Bertyinae. Hook. Icon. Pl. 7(2): t. 3633 (1967), Kew Bull. Add. Ser.
4: 60 (1975); Webster, Ann. Missouri Bot. Gard. 81: 110
232. Myricanthe Airy Shaw (1994); Radcl.-Sm., Gen. Euphorb.: 315 (2001).
Myricanthe Airy Shaw, Kew Bull. 35: 390 (1980); McPher- Monoecious tree; latex reddish; indumentum
son & Tirel, Fl. Nouv.-Caléd. 14(1): 72–73, t. 14, figs. 1–5 stellate. Leaves verticillate (ternate), petiolate,
(1987). without basal paired glands; stipules deciduous.
Monoecious shrubs; latex not recorded; indu- Staminate inflorescences terminal, subcapitate,
mentum stellate. Leaves opposite; stipules 0. pedunculate, bracts verticillate; pistillate flowers
Inflorescences terminal, unisexual or bisexual, solitary, axillary. Staminate flowers without peri-
racemoid; bracts deciduous. Staminate flowers anth or disk; stamens 25–30, filaments connate
pedicellate; sepals 3, distinct, imbricate; petals into a column, anthers basifixed, anther sacs
and disk 0; stamens 60–80, filaments connate pubescent; pistillode 0. Pistillate calyx 4–5-
into an elongated column; anthers subsessile on lobed; petals and disk 0; ovary 2-locular; stylodia
column, minutely pilose; pollen grains spheroi- bifid. Fruits capsular, endocarp woody; columella
dal, inaperturate, with Croton pattern, clavae persistent. Seeds ellipsoid, carunculate, hilum
minute and spinulose; pistillode 0. Pistillate flow- smooth; cotyledons much broader than radicle.
ers pedicellate; sepals 6, distinct, entire, subper- A single sp., B. aenigmaticum Airy Shaw,
sistent in fruit; petals and disk 0; ovary 3-locular, endemic to northeastern Borneo, in hill forest or
stellate-tomentellous; ovules anatropous, inner Casuarina forest on ultrabasic substrate. Rad-
integuments very thick, vascularized; stylodia cliffe-Smith (2001) refers to several traits in
multifid (palmatifid). Fruits capsular; columella which it resembles Oldfieldioid genera such as
persistent. Seeds cylindrical, carunculate. Aristogeitonia and Mischodon; hence, knowledge
A single sp., M. discolor Airy Shaw, confined to of its pollen structure would be highly desirable.
ultrabasic substrates in northern New Caledonia. The strongly reduced flowers suggest a shift to
wind pollination. Unpublished sequence data on
233. Cocconerion Baill. Borneodendron mentioned by Hoffmann and
Wurdack (2007) have not been accessible to us.
Cocconerion Baill., Adansonia I, 11: 87 (1873); Airy Shaw,
Kew Bull. 25: 503 (1971); McPherson & Tirel, Fl. Nouv.-
Caléd. 14(1): 38–43, t. 7 (1987); Webster, Ann. Missouri 10b. SUBTRIBE BERTYINAE M€
ull. Arg. (1865).
Bot. Gard. 81: 110 (1994).
Subtribe Ricinocarpinae G.L. Webster (1975).
Monoecious trees or shrubs; latex translucent, yel-
lowish to reddish; indumentum stellate-lepidote. Monoecious or dioecious; stems and foliage often
Leaves verticillate, 6–10 per node, tapering gradu- resinous or with reddish latex; flowers
ally to an ill-defined petiole, abaxially densely petaliferous; pollen grains with large pillars
brownish appressed-stellate; stipules 0. Flowers [except in Bertya]; embryo with cotyledons as
solitary, axillary. Staminate flowers pedicellate; narrow as radicle.
sepals 4–7, imbricate, pubescent on both faces; An entirely Australian subtribe of four genera.
petals and disk 0; stamens 30–100, filaments con-
nate into a column; anthers pubescent; pollen 235. Ricinocarpos Desf.
grains spheroidal, inaperturate, reticulate with Ricinocarpos Desf., Mém. Mus. Hist. Nat. Paris 3: 459, t. 22
micro-clavae; pistillode 0. Pistillate flowers pedi- (1817); M€
ull. Arg. in DC., Prodr. 15(2): 203 (1866); Hal-
cellate; sepals 5–7, imbricate, entire, pubescent on ford & Henderson, Austrobaileya 7: 387–449 (2007), rev.
both faces, persistent in fruit; petals and disk 0, Monoecious (dioecious) shrubs or small trees;
staminodes sometimes present; ovary (2)3-locular; laticifers non-articulated; latex 0 but stems and
ovule anatropous, inner integuments thick, vascu- foliage sometimes resinous; indumentum stellate
larized; stylodia bifid or twice bifid, branches slen- and simple. Leaves linear, 1-veined, entire,
der. Fruits capsular; columella persistent. Seeds often revolute; stipules 0. Inflorescences terminal
ellipsoidal, carunculate, testa smooth. or pseudo-axillary, glomerulate or of solitary
Two spp. endemic to New Caledonia. flowers in upper axils; bracts persistent.
Euphorbiaceae 179
Staminate flowers pedicellate; sepals (4)5(6),

nearly distinct; petals (4)5(7) or 0; disk of dis-
tinct alternipetalous segments or forming a con-
tinuous ring; stamens numerous, > 15, filaments
connate into a column; anthers dorsifixed,
extrorse; pollen grains spheroidal, inaperturate,
with Croton pattern, pillars massive and spinulose;
pistillode 0. Pistillate flowers pedicellate; sepals
persistent or deciduous in fruit; petals often mar-
cescent, deciduous or sometimes 0; disk annular
or urceolate; ovary 3-locular; ovule 1/locule, inner
integument very thick, vascularized; stylodia
basally very shortly connate, the distinct branches
spreading, deeply 2(3–5)-fid. Fruits capsular,
smooth or echinate; columella persistent. Seeds
ellipsoidal, carunculate, testa smooth; endosperm
copious; cotyledons longer than radicle.
According to the revision of Halford and
Henderson, 28 spp., all endemic to Australia.
236. Beyeria Miq. Fig. 40. Euphorbiaceae-Crotonoideae. Shonia tristigma

subsp. borealis. A Flowering branchlet. B Transverse sec-
Beyeria Miq., Ann. Sci. Nat. III, 1: 350, t. 15 (1844); M€
ull. tion of leaf. C Side view of staminate flower. D Stamen. E
Arg. in DC., Prodr. 15(2): 201 (1866); Halford & Hender- Side view of pistillate flower. F Face view of fruit. G
son, Austrobaileya 7: 577–635 (2008), rev. Ventral view of seed. (Henderson & Halford 2005; drawn
by W. Smith)
Monoecious (dioecious) shrubs or small trees;
stems and foliage resinous; laticifers non-articu-
lated, latex not recorded; indumentum stellate. Monoecious (dioecious) shrubs or small trees;
Leaves petiolate or subsessile, 1-veined, often rev- latex and resin not recorded; indumentum stel-
olute; stipules 0. Inflorescences axillary, glomerulate. Leaves shortly petiolate, abaxially pubescent;
lar (racemoid) or the pistillate flowers solitary. stipules 0. Inflorescences racemose (paniculate),
Staminate flowers pedicellate; sepals (4) 5 (6), terminal or apparently axillary, proximally with
distinct, imbricate, often petaloid; petals (4) 5, 1–2 pistillate flowers and few to several staminate
sometimes or 0; disk segments mostly 5 or disk flowers distal to them. Staminate flowers with
0; stamens 15–40, filaments distinct, shorter than calyx 5(4)-lobed; petals +; disk of distinct alter-
anthers; anthers glabrous, extrorse; pollen grains nipetalous glands or a continuous ring; stamens
spheroidal, inaperturate, exine with Croton pat- 10–30, erect; filaments distinct; anthers extrorse;
tern, pillars massive; pistillode 0. Pistillate flowers pistillode 0. Pistillate flowers with calyx lobes
pedicellate; sepals 4 or 5, persistent and some- persistent and appressed to ovary; petals +; disk
times accrescent in fruit; petals 4 or 5; disk circular; ovary 3-locular; stylodia distinct, entire.
obsolete; ovary 2–3-locular; ovules anatropous, Fruit trilobate, capsular, separating into three
inner integuments very thick; stylodia connivent 2-valved cocci; columellla persistent. Seeds car-
into a conical pseudo-stigma. Fruits capsular; unculate; endosperm copious; cotyledons nar-
columella persistent. Seeds oblong, carunculate, rower than radicle.
testa smooth; embryo cylindrical, cotyledons as Four spp., endemic to Australia (Northern
narrow as radicle. Territory, Queensland).
Twenty-four spp. endemic to Australia
recorded by Halford and Henderson (2008).
237. Shonia R. Henderson & Halford Fig. 40 238. Bertya Planch.

Shonia R. Henderson & Halford, Austrobaileya 7: 218 Bertya Planch., Hook. London J. Bot. 4: 472 (1845); M€
ull.
(2005). Arg. in DC., Prodr. 15(2): 208 (1866); Halford & Hender-
son, Austrobaileya 6: 187–245 (2002), rev.
180 G.L. Webster
Monoecious or dioecious shrubs (trees); laticifers KEY TO THE GENERA OF RICINODENDREAE

non-articulated, latex not recorded; stems often 1. Leaves simple, unlobed to 5-lobed; stipules minute or 0
resinous; indumentum stellate. Leaves alternate 239. Givotia
(opposite), abaxially densely tomentose, margins – Leaves palmatisect 2
revolute; stipules 0. Inflorescences axillary, flow- 2. Stipules flabelliform; indumentum stellate
ers solitary or in glomerules; bracts forming an 240. Ricinodendron
involucre at base of calyx. Staminate flowers ped- – Stipules not flabelliform; indumentum malpighiaceous
icellate; sepals 5 (4), basally connate, imbricate, or 0 3
3. Stylodia slender, bifid; ovary 3-locular; stamens 15–25,
sometimes petaloid; petals and disk 0; stamens anthers muticous; staminate disk dissected; fruit capsu-
15–70; filaments connate into a column, shorter lar; foliage glabrous 242. Annesijoa
than anthers; anthers dorsifixed, extrorse; pollen – Stylodia dilated, subentire to lacerate; ovary 2-locular;
grains spheroidal, inaperturate, exine with Cro- stamens 18–33; anthers apiculate; staminate disk annu-
ton pattern, clavae reduced; pistillode 0. Pistillate lar and intrastaminal; fruit drupaceous; indumentum
flowers pedicellate or subsessile; sepals 5, dis- malpighiaceous 241. Leeuwenbergia
tinct, imbricate, entire, persistent and sometimes
239. Givotia Griff.
accrescent in fruit; petals 0 or rudimentary; disk
0; ovary (2)3(5)-locular, glabrous or stellate- Givotia Griff., Calcutta J. Nat. Hist. 4: 88 (1843); M€ull. Arg.
pubescent; ovules anatropous, inner integument in DC., Prodr. 15(2): 1112 (1866); Pax & K. Hoffm., Pflan-
zenr. 147, III: 44, fig. 15 (1911); Radcl.-Sm., Kew Bull. 22:
very thick, vascularized; stylodia 3(4), proximally 493, fig. 2, t. 5 (1968), Fl. Trop. E. Afr. Euphorb. 1: 329, fig.
shortly connate, distally 2- to several-lobed. 62 (1987).
Fruits capsular, usually 1-seeded; columella not
persistent. Seeds oblong or rounded, carunculate, Dioecious trees or shrubs; latex clear; indumen-
testa smooth; embryo cylindrical, cotyledons as tum stellate. Leaves simple to 3–5-lobed, entire to
broad as radicle. coarsely dentate with sessile discoid glands, base
Twenty-eight spp. distinguished by Halford & of lamina with or without paired glands; stipules
Henderson (2002), widely distributed in Australia coarsely glandular-toothed or 0. Staminate inflor-
except for the Northern Territory. escences axillary, racemoid, with scattered glo-
merules; pistillate inflorescences terminal, of
11. TRIBE RICINODENDREAE (Pax) Hutch. (1969). solitary or clustered flowers; bracts subulate,
entire, deciduous. Staminate flowers pedicellate;
Dioecious or monoecious trees or shrubs; latici- sepals 5, distinct, imbricate; petals 5, greenish-
fers non-articulated; reddish latex + or not; indu- yellow, distinct at first, later becoming partially
mentum stellate, malpighiaceous, or 0; leaves adnate; disk segments 5, sometimes confluent;
alternate or opposite, simple to lobed or palma- stamens (3–)8–20, filaments connate at base
tisect, petiole usually glandular at apex; stipules into a column shorter than the filaments; anthers
entire, lobed, or 0; inflorescences axillary or the dorsifixed, extrorse, muticous; pollen grains
pistillate terminal, cymose-paniculate, glomeru- spheroidal, inaperturate, with Croton pattern;
lar, or reduced to solitary flowers; sepals 4 or 5, pistillode 0. Pistillate flowers pedicellate; sepals
distinct or connate; petals 5; disk dissected or (4) 5, distinct, imbricate, entire, deciduous in
lobed; stamens (3–)5–35; pollen inaperturate, fruit; petals (4) 5, greenish-yellow, distinct,
with Croton pattern; pistillode 0; ovary 1–3-locu- imbricate; disk annular; ovary 1–3-locular,
lar; stylodia bifid; fruits drupaceous or capsular; pubescent; ovules anatropous, inner integuments
seeds ecarunculate. thick, vascularized; stylodia bifid, compressed.
Four paleotropical genera, entirely African/ Fruits drupaceous, 1-seeded; endocarp thin.
Madagascan except for the New Guinean Annesi- Seeds ecarunculate, smooth.
joa and one Indian species of Givotia. In Webster Four spp., one in Africa, two in Madagascar,
(1994) and Radcliffe-Smith (2001), Leeuwenber- and one in India and Ceylon.
gia and Annesijoa were part of the broadly cir-
cumscribed Jatropheae, but in the molecular 240. Ricinodendron M€
ull. Arg.
analysis of Wurdack et al. (2005), Leeuwenbergia Ricinodendron M€ ull. Arg., Flora 47: 533 (1864), in DC.,
is resolved (rbcL only) in clade C2 together with Prodr. 15(2): 1111 (1866); Benth., Hook. Icon. Pl. 13: t.
Ricinodendron, Schinziophyton and Givotia. 1300 (1879); Léonard, Fl. Congo Belge 8(1): 116 (1962);
Euphorbiaceae 181
(1987), Fl. Zambesiaca 9(4): 294, t. 59, 60 (1996); Webster,

Ann. Missouri Bot. Gard. 81: 113 (1994); Radcl.-Sm., Gen.
Euphorb.: 328 (2001).
Schinziophyton Hutch. ex Radcl.-Sm. (1990).
Dioecious trees; laticifers non-articulated, latex

clear; indumentum stellate. Leaves palmatisect,
3–7-foliolate, pellucid-punctate, denticulate; peti-
ole glandular at apex; stipules flabellately dentate,
persistent or deciduous. Inflorescences axillary or
subterminal, the staminate cymose-paniculate,
the pistillate contracted; bracts entire, deciduous,
the pistillate foliaceous. Staminate flowers pedi-
cellate; sepals 4 or 5, distinct, imbricate; petals 5,
greenish or whitish, imbricate, coherent to form a
tube; disk segments 4–6, glabrous; stamens (7–)
10–20, filaments connate basally into a short col-
umn, exserted beyond calyx; anthers dorsifixed,
versatile, introrse; pollen grains spheroidal, ina-
perturate, exine with Croton pattern; pistillode 0.
Pistillate flowers pedicellate; sepals (4) 5, distinct,
imbricate, deciduous in fruit; petals 5, greenish or
whitish, imbricate, coherent; disk annular, crenu-
late, glabrous; ovary 1–3-locular, pubescent,
ovules anatropous, inner integuments very
thick, vascularized; stylodia bifid. Fruits drupa-
ceous, exocarp fleshy, endocarp woody. Seeds
ecarunculate, subspheroidal, testa irregularly
ridged. 2n ¼ 22.
Two spp. widely distributed in tropical
Africa. Originally, six African spp. had been
Fig. 41. Euphorbiaceae-Crotonoideae. Leeuwenbergia
described in Ricinodendron, of which only a sin- africana. A Leaf, seen from below. B Glands on petiole
gle polymorphic species, R. heudelotii, with 3 apex. C Flowering branch. D Pistillate flower. E Corolla
subspecies was recognized by Govaerts et al. aestivation. F Ovary, side view. G Same, transversal section.
(2000); an additional species was transferred to H Same, longitudinal section. I Ovule with and without
obturator. (Letouzey & N. Hallé 1974; drawn by N. Hallé)
the segregate genus Schinziophyton. However, the
key to the Congo spp. of Ricinodendron by Léo-
nard contrasts two species that are very distinct, bracts deciduous. Staminate flowers pedicellate;
but which in my opinion can easily be accommo- sepals connate, irregularly dehiscent into 2 or 3
dated in a single genus; Schinziophyton is therefore lobes, closed in bud; petals 5, white, orbiculate,
here relegated to synonymy, and Ricinodendron is distinct; disk interstaminal, receptacular; stamens
considered to include at least 2 species. 18–33, distinct, inserted in cavities of the disk;
filaments distinct; anthers introrse, apiculate;
241. Leeuwenbergia Letouzey & Hallé Fig. 41 pollen grains inaperturate, with Croton pattern;
Leeuwenbergia Letouzey & Hallé, Adansonia II, 14: 379, pistillode 0. Pistillate flowers pedicellate; calyx
figs. 2, 3 (1974); Webster, Ann. Missouri Bot. Gard. 81: 104 and petals as in staminate; disk annular-cupular,
(1994). hirsute; ovary 2–3-locular, hirsute; ovules anatro-
Dioecious trees; latex reddish; indumentum mal- pous, inner integuments thick, vascularized; sty-
pighiaceous. Leaves palmatisect, petiolate with lodia foliose-stigmatiform, crenulate. Fruits and
1 or 2 large apical glands; leaflets entire; stipules seeds unknown.
deciduous. Inflorescences axillary or subtermi- Two spp. of west African rainforests: Gabon
nal, unisexual, racemoid-thyrsoid or paniculate; to Cameroon and Zaire.
182 G.L. Webster
242. Annesijoa Pax & K. Hoffm. The five subtribes contain a total of 13 genera
Annesijoa Pax & K. Hoffm., Pflanzenr. 147, XIV: 9 (1919);
and 45 species. van Welzen and Stuppy (1999)
Airy Shaw, Kew Bull. 16: 345 (1963), Hook. Icon. Pl. have shown in a cladistic analysis of the
38: t. 3713 (1974), Kew Bull. Add. Ser. 8: 27, t. 7 (1980); tribe that subtribe Grosserinae as delimited by
Hoang Van Nam & van Welzen, Blumea 49: 427–437 Webster (1994) is unnatural. Paracroton, for-
(2004). merly doubtfully included in the Codiaeae, in
Monoecious trees; latex not recorded; indumen- the combined and the partial molecular analyses
tum 0. Leaves palmatisect, petiolate with apical (Wurdack et al. 2005) always goes together with
paired stipelliform deciduous glands; leaflets genera of the Aleuritideae and here is shifted to
entire; stipules rudimentary. Staminate inflores- this tribe.
cences axillary, dichasial-paniculate, pistillate
flowers few and subterminal; bracts inconspicu- KEY TO THE SUBTRIBES OF ALEURITIDEAE
ous. Staminate flower pedicellate; calyx cupular, 1. Petals 0; leaves gland-dotted 12e. Neoboutoninae
shallowly lobed, of 5 connate sepals, open in bud; – Petals +, at least in staminate flowers; leaves not gland-
petals 5, white, distinct; disk segments 5; stamens dotted 2
15–25, filaments distinct or coherent in part; 2. Petals 6–13; stamens 30–100, filaments distinct; inflor-
anthers ellipsoidal; pollen grains spheroidal, ina- escences terminal, glomerular; leaf blades pinnately
veined 12b. Garciinae
perturate, with Croton pattern; pistillode 0. Pis-
– Petals 4 or 5; stamens 6–40, filaments distinct or
tillate flowers pedicellate; sepals 5, distinct or connate 3
nearly so, entire; petals 5, distinct, imbricate; 3. Stamens 7–20, filaments connate; monoecious; inflores-
disk segments 5; ovary 2-locular, glabrous; ovules cences terminal; leaf blades palmately lobed or veined;
anatropous, inner integuments moderately thick, monoecious; stems with latex 12a. Aleuritinae
vascularized; stylodia bipartite. Fruits capsular; – Stamens 6–40, filaments distinct or connate; dioecious
(monoecious); inflorescences terminal or axillary; leaf
endocarp woody; columella not persistent. Seeds blades pinnately or palmately veined; stems mostly
spheroidal or angled, ecarunculate (?), testa without latex 4
smooth. 4. Indumentum simple or 0; inflorescences mostly termi-
One sp., A. novoguineensis Pax & K. Hoffm., nal [axillary in Tapoides] 12c. Grosserinae
of rainforests in New Guinea. – Indumentum stellate or lepidote; inflorescences termi-
nal or axillary 5
5. Staminate petals usually coherent or connate; leaves
12. TRIBE ALEURITIDEAE Hurus. (1954). entire; caruncle 0; dioecious 12d. Crotonogyninae
Monoecious trees or shrubs; laticifers non-artic- – Staminate petals distinct; leaves serr(ul)ate; caruncle +
or 0; dioecious or monoecious 12f. Paracrotoninae
ulated, latex white or reddish; indumentum sim-
ple or stellate; leaves alternate, simple, entire 12a. SUBTRIBE ALEURITINAE (Hurus.) G.L. Webster
(dentate), pinnately to palmately veined or pal- (1975).
mately lobed, eglandular or with basal laminar
glands; stipules present or 0; inflorescences ter- Monoecious trees or shrubs; indumentum simple
minal or axillary, cymose-paniculate or reduced or stellate; leaves palmately veined or lobed,
to glomerules; staminate calyx closed in bud, with glands at apex of petiole; inflorescences
splitting into valvate segments; petals 5(6–13), paniculate; bracts eglandular, deciduous; sta-
distinct, imbricate; disk dissected or interstam- minate calyx segments 2 or 3; petals 5, distinct,
inal; stamens 7–20(100); pollen grains spheroi- glabrous; disk-segments 5; stamens 7–20, inner
dal, inaperturate, with Croton pattern; sepals and filaments connate; ovary 2–3-locular; fruit drupa-
petals as in staminate; disk lobed, dissected, or ceous or capsular.
obsolete; ovary 2–5-locular; ovules anatropous, This small paleotropical subtribe includes all
inner integuments thick, vascularized; stylodia of the species traditionally referred to Aleurites.
bifid; fruit drupaceous or capsular; seeds ecarun- The fractionation into 3 genera proposed by Airy
culate (carunculate). Shaw has been widely accepted, but despite the
As here circumscribed, the Aleuritideae are critical study of the Aleuritinae by Stuppy et al.
an exclusively paleotropical group, except for (1999), it seems preferable to combine Reutealis
Garcia, the affinities of which are dubious. with Vernicia.
Euphorbiaceae 183
KEY TO THE GENERA OF ALEURITINAE tum simple, bifid, or stellate. Leaves unlobed to 3
1. Fruits drupaceous; indumentum stellate; stamens (5)-lobed, palmately veined, entire, basally with
17–32, anthers in 2 or 4 whorls; leaf blades triplinerved, paired glands at junction with petiole; stipules
brochidodromous 243. Aleurites deciduous. Inflorescences terminal, usually bisex-
– Fruits capsular; indumentum simple or stellate; ual, thyrsoid-paniculate, bracts persistent or
stamens 8–12(14), mostly in 2 whorls; leaf blades deciduous. Staminate flowers pedicellate; calyx
palmately veined, eucamptodromous 244. Vernicia splitting into 2 or 3 valvate lobes; petals 5, dis-
tinct, contorted-imbricate, adaxially pubescent;
243. Aleurites J.R. & G. Forst. disk segments 5, glabrous; stamens 7–12(15),
Aleurites J.R. & G. Forst., Charact. Gen. Pl.: 111, t. 56 biseriate, those of the outer stamens distinct to
(1776); M€ ull. Arg. in DC., Prodr. 15(2): 722 (1866); Airy basally connate, of the inner connate halfway or
Shaw, Kew Bull. 20: 393 (1967); Webster, J. Arnold Arb. more; anthers basifixed, extrorse or introrse; pol-
48: 342 (1967); Walker, Fl. Okinawa: 644, figs. 96–98 len grains spheroidal, inaperturate, exine with
(1976); A.C. Sm., Fl. Vitiensis Nova 2: 547 (1981); Croton pattern; pistillode 0. Pistillate flowers ped-
(1987); P.I. Forster, Muelleria 9: 6, figs. 1, 2 (1996); Stuppy icellate; perianth and disk as in staminate flowers;
et al., Blumea 44: 79, fig. 1 (1999). ovary 3(5)-locular, pubescent; ovules anatro-
pous, inner integuments moderately thick; stylo-
Monoecious trees; latex not obvious; indumen- dia bifid. Fruits capsular. Seeds trigonous,
tum simple and stellate. Leaves shallowly 3–5- ecarunculate, testa thick and woody.
palmately lobed or unlobed, triplinerved, entire, Four spp., SE Asia, Malesia, and extending to
with paired basal glands at junction with petiole; Japan.
stipules deciduous. Inflorescences terminal,
cymose-paniculate, uni- or bisexual; bracts
deciduous. Staminate flowers pedicellate; calyx 12b. SUBTRIBE GARCIINAE M€
ull. Arg. (1865).
splitting valvately or irregularly into 2 or 3 Monoecious; indumentum simple; leaves entire,
lobes; petals 5(6), distinct, imbricate, exserted without glands at apex of petiole, estipulate;
from calyx; disk segments 5; stamens 17–32, 4- inflorescences terminal, glomerular; petals 6–13;
seriate, the filaments of the outer ones distinct, of staminate disk intrastaminal; stamens 30–100,
the inner connate into a column; anthers dorsi- filaments distinct; ovary 3-locular, pubescent;
fixed, introrse, muticous, with dilated connective; ovules anatropous, inner integuments thick; sty-
pollen grains spheroidal, inaperturate, exine with lodia bifid; fruits capsular; seeds ecarunculate.
Croton pattern; pistillode 0. Pistillate flowers ped- In accord with Webster (1975, 1994), subtribe
icellate; perianth as in staminate flowers; disk Garciinae is restricted to Garcia, which is very
annular, 5-lobed; ovary 2–3(4)-locular, tomen- isolated within the Aleuritideae. There are
tose; stylodia 2 or 3, connate at the base, bilobed. striking resemblances between Garcia and Sago-
Fruits drupaceous; exocarp fleshy, endocarp tia and Sandwithia, so that Garcia might belong
thin-walled; seeds massive, subspheroidal, ecar- to the Crotoneae.
unculate; endosperm copious. 2n ¼ 22.
Three spp., distributed from Ceylon to SE
Asia, Australia, Melanesia and Polynesia, where 245. Garcia Vahl
it is commonly introduced. Garcia Vahl, Skriv. Naturh. Selsk. Kj€ obenh. 2: 217, t. 9
(1792); M€ull. Arg. in DC. Prodr. 15 (2): 721 (1866); Pax,
244. Vernicia Lour. Pflanzenr. 147, I: 14 (1910); Lundell, Wrightia 1: 1 (1945);
Webster, Ann. Missouri Bot. Gard. 54: 238, fig. 6 (1968);
Vernicia Lour., Fl. Cochinch.: 586 (1790); Hemsley, Hook. Burger & Huft, Fieldiana Bot. n. s. 36: 122, fig. 21 (1995).
Icon. Pl. 29: t. 2801, 2802 (1906, under Aleurites); Airy
Shaw, Kew Bull. 20: 394 (1967); Radcl.-Sm., Fl. E. Trop. Monoecious shrubs or trees; laticifers non-
Afr. Euphorb. 1: 178, fig. 35 (1987); Webster, Ann. Mis- articulated; latex not recorded; indumentum
souri Bot. Gard. 81: 114 (1994); Stuppy et al., Blumea 44: simple. Leaves petiolate (pulvinate), cartilagi-
88, fig. 3 (1999); Radcl.-Sm., Gen. Euphorb.: 331 (2001). nous, without paired basal glands; stipules 0.
Reutealis Airy Shaw (1967).
Inflorescences terminal, glomerular, bisexual,
Monoecious trees; laticifers non-articulated, latex each with 1 or 2 pistillate and several staminate
whitish or reddish, often not apparent; indumen- flowers; bracts entire, inconspicuous. Staminate
184 G.L. Webster
flowers pedicellate; calyx splitting into 2 or 3 5. Bracts small, persistent; inflorescence paniculate; leaf
valvate segments; petals 6–13, reddish, distinct, blades denticulate; stipular scars minute; pistillate
sepals medianly thickened 247. Grossera
narrow, sericeous on both faces, exserted from
– Bracts large, imbricate, deciduous; inflorescence cone-
calyx; disk ill-defined, mainly represented on the like; leaf blades entire; stipular scars subannular, con-
convex pilose and glandular receptacle; stamens spicuous; pistillate sepals not thickened 246. Cavacoa
30–100, filaments distinct; anthers basifixed,
minutely apiculate; pollen grains spheroidal, ina- 246. Cavacoa Léonard
perturate, exine with Croton pattern; pistillode 0.
Cavacoa Léonard, Bull. Jard. Bot. Brux. 25: 320, fig. 54
Pistillate flowers pedicellate; calyx splitting into (1955), Fl. Congo Belge 8(1): 191, fig. 16 (1962); Elffers &
2 or 3 valvate segments, deciduous; petals as in Taylor, Hook. Icon. Pl. 36: t. 3561 (1956); Radcl.-Sm., Fl.
the staminate flowers; disk deeply lobed; ovary 3- E. Trop. Afr. Euphorb. 1: 174, fig. 33 (1987); Radcl.-Sm.,
locular, sericeous; ovules anatropous, inner inte- Fl. Zambesiaca 9(4): 304, t. 63 (1996).
guments thick, vascularized; stylodia thick, Dioecious trees or shrubs; branches terminating in
reflexed, bifid. Fruits capsular; columella persis- perulate buds; latex not recorded; indumentum
tent. Seeds subspheroidal, ecarunculate; testa simple. Leaves pellucid-punctate, entire, usually
smooth. 2n ¼ 66. with basal paired glands and scattered embedded
Two spp., Mexico south to Colombia. glands; stipules deciduous. Inflorescences termi-
nal, racemoid; bracts large, covering flowers in
12c. SUBTRIBE GROSSERINAE G.L. Webster (1975). bud, deciduous. Staminate flowers pedicellate;
Dioecious (monoecious) trees or shrubs; latex usu- calyx splitting into 2 segments; petals 4 or 5, dis-
ally not recorded; indumentum simple or 0; leaves tinct, imbricate, glabrous; disk segments 4 or 5,
pinnately veined or triplinerved, with or without glabrous; stamens 15–30, filaments connate into a
basal laminar glands; stipules deciduous or 0; column [outer filaments distinct]; anthers with
inflorescences terminal, racemoid, or paniculate; enlarged connective; pollen grains spheroidal,
staminate petals 4 or 5, distinct or basally connate; inaperturate, exine with Croton pattern; pistillode
staminate disk dissected; stamens 6–40, filaments 0. Pistillate flowers pedicellate; sepals 4 or 5, dis-
distinct or connate; pistillate petals 4 or 5, distinct; tinct, imbricate, entire, deciduous in fruit; petals
ovary 3–5-locular; stylodia (twice) bifid; fruit cap- 4 or 5, distinct, imbricate, glabrous; disk annular,
sular; seeds ecarunculate (carunculate). glabrous; ovary 3-locular, glabrous; ovules anat-
Six paleotropical genera, three African/Mada- ropous, inner integuments thick, vascularized;
gascan and three SE Asian/Malesian. Deutzianthus stylodia bifid. Fruits capsular; columella persis-
and Oligoceras, included by Webster (1994) and tent. Seeds ecarunculate, testa smooth.
Radcliffe-Smith (2001) in the Jatropheae, are here Three spp. of tropical Africa south to Natal.
brought to tribe Aleuritidae, in consonance with
Thin (1995), and provisionally included in sub- 247. Grossera Pax
tribe Grosserinae. Molecular data for this place- Grossera Pax, Bot. Jahrb. 33: 281 (1903); Pax & K. Hoffm.,
ment are still lacking. Pflanzenr. 147, VI: 105, fig. 21 (1912); Leandri, Bull. Soc.
Bot. France 85: 524 (1939); Cavaco, Bull. Mus. Hist. Nat.
KEY TO THE GENERA OF GROSSERINAE Paris 21: 272 (1949); Léonard, Bull. Jard. Bot. Brux. 25: 316
(1955), 28: 118 (1958), Fl. Congo Belge 8(1): 188 (1962);
1. Pollen grains with echinate exinous pillars; leaf blades Keay, Fl. W. Trop. Afr. ed. 2, 1: 398 (1958); Radcl.-Sm.,
triplinerved; bracts glandular 249. Tannodia Gen. Euphorb.: 337, fig. 42 (2001).
– Pollen grains with rounded exinous pillars; leaves pin-
nately veined; bracts eglandular 2 Dioecious shrubs; latex not recorded; indumen-
2. Leaf blades not pellucid-punctate 3 tum simple. Leaves entire or denticulate, pellu-
– Leaf blades pellucid-punctate 5 cid-punctate, usually with paired basal glands at
3. Staminate petals adaxially pubescent 248. Tapoides junction with petiole; stipules deciduous. Inflor-
– Staminate petals glabrous 4 escences terminal, cymose-paniculate; bracts
4. Monoecious; glabrous; stylodia 3, connate at the base, entire, eglandular, the pistillate deciduous. Stami-
bifid 250. Oligoceras
nate flowers pedicellate; calyx closed in bud,
– Dioecious; indumentum simple, largely confined to
inflorescences and fruits; stylodia 3, distinct, twice splitting into 2 or 3 valvate segments; petals 5,
bifid, or deeply branched and distally bifid, hence white, distinct, imbricate, glabrous; disk seg-
appearing as 6 stylodia 251. Deutzianthus ments 5, glabrous; stamens 13–40, filaments
Euphorbiaceae 185
irregularly connate at the base; anthers with Holstia Pax (1909; nom. illeg.).
enlarged connective; pollen spheroidal, grains Domohinea Leandri (1941).
Neoholstia Rauschert (1982).
inaperturate, exine with Croton pattern, pillars
acute, smooth; pistillode 0. Pistillate flowers ped- Monoecious or dioecious trees; latex not
icellate; sepals 4 or 5, distinct, imbricate, entire, recorded; indumentum simple, scanty [inflores-
persistent in fruit; petals 4 or 5, distinct, imbri- cences only]. Leaves eglandular; stipules persis-
cate; disk cupular, lobed, glabrous; ovary 3-locu- tent or deciduous. Inflorescences terminal,
lar, glabrous; ovules anatropous, inner racemoid or spiciform, or the staminate in con-
integuments thick, vascularized; stylodia bifid. tracted glomerules, unisexual or bisexual; bracts
Fruits capsular; columella triquetrous, persistent. entire to glandular-lacerate. Staminate flowers
Seeds ecarunculate, testa smooth. pedicellate; calyx closed in bud, splitting into
Eight spp., C and W Africa and (1) Madagascar. 2–5 valvate segments; petals 4 or 5, white, dis-
tinct, imbricate, longer than calyx; disk segments
248. Tapoides Airy Shaw 4 or 5, glabrous; stamens 7–12, filaments connate
Tapoides Airy Shaw, Kew Bull. 14: 473 (1960), 20: 412 below, biseriate; anthers dorsifixed, the outer
(1966), Hook. Icon. Pl. 37: t. 3632 (1967), Kew Bull. Add. extrorse, inner introrse, connective broad; pol-
Ser. 4: 200 (1975); Radcl.-Sm., Gen. Euphorb.: 337 (2001). len grains spheroidal, inaperturate, exine with
Croton pattern; pistillode 0. Pistillate flowers
Dioecious trees; latex reddish; indumentum sim-
pedicellate; sepals 4 or 5, basally connate, imbri-
ple, scanty. Leaves petiolate (pulvinate), crowded
cate, entire, persistent in fruit; petals 4 or 5,
at ends of branches, entire, lacking basal glands;
distinct, imbricate, exserted beyond sepals; disk
stipules subulate, minute. Inflorescences axillary
annular; ovary 3-locular, pubescent; ovules anat-
or subterminal, the staminate thyrsoid-panicu-
ropous, inner integuments thick, vascularized;
late, the pistillate glomerular; bracts entire. Sta-
stylodia basally connate, erect, bifid. Fruits
minate flowers pedicellate; calyx closed in bud,
capsular. Seeds ellipsoidal, ecarunculate, testa
splitting into 3 valvate segments; petals 5, dis-
smooth.
tinct, imbricate, adaxially pubescent; disk seg-
Nine spp., three in Africa and six in Mada-
ments 5; stamens 6–8, filaments distinct,
gascar (including the Comoros).
pubescent; pistillode 0. Pistillate flowers pedicel-
late; calyx splitting into 3 valvate segments, per-
250. Oligoceras Gagnep.
sistent in fruit; petals and disk unknown; ovary 3-
locular, pubescent. Fruits capsular. Mature seeds Oligoceras Gagnep., Bull. Soc. Bot. France 71: 872 (1925);
not recorded. Fl. Indochine 5: 467, fig. 58, 3–9 (1926); Airy Shaw, Kew
Bull. 14: 392 (1960).
A single sp., T. vilamilii (Merr.) Airy Shaw,
endemic to Borneo (Sabah). Airy Shaw (1967) Monoecious trees; latex not recorded; foliage
suggested that it is most closely related to Aleur- glabrous. Leaves long-petiolate, lamina
ites and Elateriospermum but also pointed out deltoid, with 2 discoid glands at apex of peti-
possible affinities with Omphalea. However, the ole; stipules 0. Inflorescences terminal, panicu-
reddish latex, mainly axillary inflorescences, late, bisexual; bracts scale-like, apically
and distinct stamens of Tapoides set it apart fimbriate. Staminate flowers pedicellate; calyx
from the other Grosserinae; this suggests that it campanulate, sepals 5, connate, each with a
may be misplaced. cylindric-cornute truncate appendage; petals 5,
distinct, imbricate, unguiculate; disk segments 5,
249. Tannodia Baill. connate around the base of the staminal column;
Tannodia Baill., Adansonia I, 1: 251 (1861); M€ ull. Arg. in stamens and staminodes connate in 2 whorls
DC., Prodr. 15(2): 728 (1866); Pax & K. Hoffm., Pflanzenr. into a cylindrical column, outer whorl of 5
147, VI: 110 (1912); Léonard, Bull. Jard. Bot. Brux. 25: 300 stamens, inner of 3 staminodes; anthers introrse;
(1955), Fl. Congo Belge 8(1): 186 (1962); Radcl.-Sm., Fl. E. pollen grains spheroidal, inaperturate, exine
Trop. Afr. Euphorb. 1: 172, fig. 32 (1987); Webster, Ann. with Croton pattern; pistillode 0. Pistillate flow-
Missouri Bot. Gard. 81: 115 (1994); Radcl.-Sm., Fl. Zam-
besiaca 9(4): 306, t. 64 (1996), Kew Bull. 53: 173 (1998), ers pedicellate; calycine appendages obtuse, oth-
Gen. Euphorb.: 338 (2001). erwise perianth as in staminate flowers; disk
Tandonia Baillon (1861). annular; ovary 3-locular, stylodia nearly distinct,
186 G.L. Webster
bifid, stigmatic portion coiled. Fruits drupa- connate; anthers often apiculate; pistillate sepals
ceous. Seeds unknown. 4 or 5, imbricate or valvate; ovary 3-locular;
A single sp., O. eberhardtii Gagnep., known stylodia bifid or multifid; fruits capsular; seeds
only from Annam, Vietnam. ecarunculate.
A subtribe of three African genera.
251. Deutzianthus Gagnep.
KEY TO THE GENERA OF CROTONOGYNINAE
Deutzianthus Gagnep., Bull. Soc. Bot. France 71: 139
(1924), Fl. Indochine 5: 296, figs. 31, 3–9, 32, 1 (1925); 1. Leaf blades palmately veined; petioles with inflated
Airy Shaw, Kew Bull. 14: 362 (1960), 16: 346 (1963); hairs; stylodia bifid; lianas 254. Manniophyton
Webster, Ann. Missouri Bot. Gard. 81: 104 (1994); – Leaf blades pinnately veined; petioles lacking inflated
Radcl.-Sm., Gen. Euphorb.: 291 (2001). hairs; stylodia bifid or multifid; trees or shrubs 2
Loerzingia Airy Shaw (1963). 2. Stylodia bifid; inflorescences terminal, paniculate; sta-
minate petals distinct; staminate disk receptacular and
Dioecious trees; latex yellowish-orange; indumen- extrastaminal, of > 10 segments 252. Cyrtogonone
tum simple. Leaves long-petiolate, with 2 adaxial – Stylodia multifid; inflorescences axillary, racemoid
disciform glands at junction with petiole; stipules or spiciform; staminate petals usually coherent
deciduous. Inflorescences long-pedunculate, ter- or connate; staminal disk extrastraminal, of 5–8
segments 253. Crotonogyne
minal or subterminal, of compound unisexual
dichasia, bracts linear, eglandular, persistent. Sta- 252. Cyrtogonone Prain
minate flowers pedicellate; sepals 5, connate,
the calyx 5-dentate, valvate or subimbricate; petals Cyrtogonone Prain, Kew Bull. 1911: 231 (1911),
5, distinct, entire, adaxially pilose; disk glands 5; Hook. Icon. Pl. 31: t. 3009 (1915); Pax & K. Hoffm., Pflan-
zenr. 147, VI: 111, fig. 23 (1912); Keay, Fl. W. Trop. Afr.,
stamens 7 or 8, biseriate, 5 outer distinct, 2 or 3 ed. 2, 1: 399 (1958); Webster, Ann. Missouri Bot. Gard. 81:
inner connate to halfway; anthers cordulate or 116 (1994); Radcl.-Sm., Gen. Euphorb.: 342 (2001).
sagittate at the base; pollen grains spheroidal, ina-
perturate, exine with Croton pattern; pistillode Dioecious trees; latex not recorded; indumentum
0 or rudimentary. Pistillate flowers pedicellate; lepidote. Leaves entire or dentate, silvery-lepidote
perianth as in staminate; disk annular or 5-lobed; abaxially, with paired basal glands at junction
ovary 3-locular, sericeous; stylodia 3, distally twice with petiole; stipules minute. Inflorescences ter-
bifid, or deeply branched and apically bifid, hence minal, cymose-paniculate; bracts inconspicuous.
appearing as 6 stylodia. Fruits indehiscent or Staminate flowers pedicellate; calyx closed in
tardily dehiscent. Seeds unknown. bud, splitting into 2–4 valvate segments; petals 5
Two species, disjunct in Vietnam and Suma- (6), distinct, contorted, glabrous; disk segments c.
tra. Radcliffe-Smith (2001) maintains Loerzingia 10, receptacle glandular; stamens 12–30, fila-
as a distinct genus on the basis of its deciduous ments distinct, glabrous; anthers dorsifixed,
foliar glands, imbricate sepals, and presence of introrse; pollen grains spheroidal, inaperturate,
pistillode in the staminate flower. The type spe- exine with Croton pattern; pistillode 0. Pistillate
cies, D. tonkinensis Gagnep., does indeed differ in flowers pedicellate; sepals 4 or 5, imbricate,
these characters, but the similarities seem more entire; petals 5, distinct, imbricate; disk annular;
important than the differences. ovary 3-locular, pubescent; stylodia bifid. Fruits
capsular, cocci dorsally verrucate. Seeds sub-
spheroidal, ecarunculate.
12d. SUBTRIBE CROTONOGYNINAE G.L. Webster A single sp., C. argentea (Pax) Prain, W Africa.
(1975).
Trees or shrubs, sometimes scandent; latex 253. Crotonogyne M€
ull. Arg.
apparently not produced; indumentum stellate Crotonogyne M€ ull. Arg., Flora 47: 535 (1864), in DC.,
or lepidote; leaves pinnately or palmately Prodr. 15(2): 720 (1866); Pax & K. Hoffm., Pflanzenr.
veined, biglandular at junction with petiole, stip- 147, VI: 111, fig. 24 (1912); Prain, Hook. Icon. Pl. 31: t.
3019 (1915); Keay, Fl. W. Trop. Afr., ed. 2, 1: 399 (1958);
ulate; inflorescences axillary, spiciform or race- Léonard, Fl. Congo Belge 8(1): 174, fig. 14 (1962); Web-
moid to paniculate; staminate sepals connate in ster, Ann. Missouri Bot. Gard. 81: 116 (1994); Radcl.-Sm.,
bud, valvately dehiscent; petals coherent or con- Gen. Euphorb.: 342 (2001).
nate; disk dissected; stamens 7–40, distinct or Neomanniophyton Pax (1912).
Euphorbiaceae 187
Dioecious (monoecious); latex not recorded; indu- enlarged; pollen grains spheroidal, inaperturate,
mentum simple and stellate or lepidote. Leaves exine with Croton pattern; pistillode 0. Pistillate
with basal laminar glands at junction with petiole; flowers pedicellate; sepals 5, slightly connate
stipules entire, persistent. Inflorescences axil- at base, imbricate, entire, persistent and some-
lary, racemoid or spiciform, sometimes branched; what accrescent in fruit, petals 5, yellowish-green,
bracts biglandular at base. Staminate flowers pedi- distinct, imbricate, adaxially pubescent; disk
cellate; calyx closed in bud, splitting into 2–4 val- annular, pubescent; ovary 3-locular, sericeous-
vate segments; petals 5, mostly connate; disk hispid; ovules anatropous, inner integuments
segments 5–8, glabrous; receptacle glabrous; moderately thick, vascularized; stylodia bipartite.
stamens (7–)10–28, filaments distinct; anthers Fruits capsular, cocci thick and woody; columella
apiculate; pollen grains spheroidal, inaperturate, persistent. Seeds compressed, ecarunculate, thin
exine with Croton pattern; pistillode 0. Pistillate and shiny.
flowers pedicellate; sepals (4)5, slightly connate A single sp., M. africanum M€ ull. Arg., West
basally, imbricate, abaxially lepidote, persistent in Africa, Liberia to Angola.
fruit; petals (4)5(6), distinct, imbricate, glabrous;
disk annular-lobed, glabrous; ovary 3-locular, stel- 12e. SUBTRIBE NEOBOUTONINAE (Hutch.)
late-pubescent or lepidote; ovules anatropous, G.L. Webster (1975).
inner integuments thick, vascularized; stylodia
multifid. Fruits capsular; columella persistent, Dioecious trees or shrubs; indumentum stellate
slender, apically dilated. Seeds somewhat com- or lepidote; leaves unlobed, pinnately or pal-
pressed, ecarunculate, testa smooth. mately veined, glandular-dotted, stipulate; inflor-
Sixteen spp., W Africa, from Sierra Leone to escences terminal or axillary, racemoid to
Angola. Pax (1912) separated Neomanniophyton paniculate; calyx segments 2 or 3; disk dissected
from Manniophyton on the basis of leaf venation or 0; stamens 15–40, filaments distinct, shorter
and division of the stylodia. However, further study than anthers; anthers with glandular connective;
has shown that Neomanniophyton is closer to Cro- ovary 3-locular; stylodia bifid; fruit capsular;
tonogyne in its leaf venation and stylar branching. seeds carunculatae or ecarunculate.
Although generic boundaries in the Crotogyninae Two genera, African and Malagasy.
need further study, it appears best to follow Webster
(1994) and Radcliffe-Smith (2001) in combining KEY TO THE GENERA OF NEOBOUTONINAE
Crotonogyne and Neomanniophyton. 1. Leaf blades palmately veined; stipules persistent; seeds
carunculate; indumentum stellate 255. Neoboutonia
254. Manniophyton M€
ull. Arg. – Leaf blades pinnately veined; stipules deciduous; seeds
ecarunculate; indumentum lepidote 256. Benoistia
Manniophyton M€ ull. Arg., Flora 47: 530 (1864), in DC.,
Prodr. 15(2): 719 (1866); Benth., Hook. Icon. Pl. 13: t. 1267 255. Neoboutonia M€
ull. Arg.
(1878); Pax & Hoffm., Pflanzenr. 147, VI: 120, fig. 25
(1912); Keay, Fl. W. Trop. Afr. et. 2, 1: 400 (1958); Léo- Neoboutonia M€ ull. Arg., J. Bot. 2: 336 (1864), in DC.,
nard, Fl. Congo Belge 8(1): 171, fig.13 (1962); Radcl.-Sm., Prodr. 15(2): 892 (1866); Benth., Hook. Icon. Pl. 13: t.
Gen. Euphorb.: 343 (2001). 1298, 1299 (1879); Pax & K. Hoffm., Pflanzenr. 147, VII:
71 (1914); Radcl.-Sm., Fl. Zambesiaca 9(4): 310, t. 66
Dioecious lianas; latex reddish; indumentum (1996).
simple to stellate. Leaves simple, sometimes
3–5-lobed, palmately veined, with paired stipels Dioecious trees and shrubs; latex not recorded;
and bottle-shaped glands abaxially near junction indumentum tufted-stellate and simple. Leaves
with petiole; stipules deciduous. Inflorescences palmately veined, cordate, with basal laminar sti-
terminal and axillary, often in pairs, thyrsoid- pellate glands and minute dispersed discoid
paniculate; bracts biglandular at base. Staminate glands; stipules foliaceous, entire, persistent.
flowers pedicellate; calyx closed in bud, splitting Inflorescences terminal and axillary, cymose-
into 2 or 3 valvate segments; petals 5, glabrous, paniculate; staminate bracts entire, eglandular,
connate into an urceolate corolla [lobes much persistent. Staminate flowers pedicellate; calyx
shorter than tube]; disk segments 5 or 6, pubes- closed in bud, splitting into 2 or 3 segments;
cent; receptacle pilose; stamens 10–20, filaments petals 0; disk segments 8–10, minute; stamens
distinct; anthers basifixed, apiculate, connective 15–40, filaments distinct; anthers basifixed,
188 G.L. Webster
introrse, glandular-apiculate; pollen grains sphe- 10–32, outer filaments distinct, the inner connate
roidal, inaperturate, exine with Croton pattern; into a column; inflorescences terminal or axillary;
pistillode 0. Pistillate flowers pedicellate; sepals 5, seeds carunculate or ecarunculate.
distinct, imbricate, entire, persistent, sometimes Mildbraedia and Paracroton appear to repre-
accrescent; petals 0; disk annular; ovary 3-locular, sent sister genera united by a distinctive synapo-
stellate-pubescent or lepidote; ovules anatropous, morphy of the androecium with the outer
inner integuments very thick, vascularized; stylo- filaments distinct and the inner connate.
dia bipartite, branches slender. Fruits capsular;
columella persistent. Seeds subspheroidal, carun-
cle small, adpressed; testa smooth. KEY TO THE GENERA OF PARACROTONINAE
Three spp., widespread in tropical Africa. Pax 1. Dioecious; inflorescences axillary; leaf blades without
and Hoffmann (1914) spectacularly misplaced paired basal glands; seed coat dry 257. Mildbraedia
Neoboutonia into the Acalypheae-Mercurialinae, – Monoecious; inflorescences terminal; leaf blades with
assigning it to series Neoboutoniiformes. This paired basal glands; seed coat fleshy 258. Paracroton
was negated by Punt (1962), who showed that
the pollen of Neoboutonia has the typical Croto- 257. Mildbraedia Pax Fig. 42
noid pattern of exine ornamentation.
Mildbraedia Pax, Bot. Jahrb. 43: 319 (1909); Pflanzenr.
256. Benoistia H. Perrier & Leandri 147, III: 11 (1911), 147, VII: 403 (1914); Léonard, Fl.
Congo Belge 8(1): 85 (1962); Radcliffe-Sm., Fl. E. Trop.
Benoistia H. Perrier & Leandri, Bull. Soc. Bot. France 85: Afr., Euphorb. 1: 340, fig. 64 (1987), Fl. Zambesiaca 9(4):
528 (1938); Radcl.-Sm., Kew Bull. 43: 632 (1988). 273, t. 56 (1996).
Neojatropha Pax (1910).
Dioecious trees; latex not recorded; indumentum
simple and glandular-lepidote. Leaves pellucid- Dioecious trees or shrubs; latex not recorded; indu-
punctate, lacking basal glands at junction with mentum stellate. Leaves simple, sometimes lobed,
petiole; stipules minute, deciduous. Inflorescences triplinerved or pinnately veined, dentate, eglandu-
axillary, racemoid or paniculate; pistillate bracts lar; stipules subulate, deciduous. Inflorescences
foliose. Staminate flowers pedicellate; calyx closed axillary or slightly supraaxillary, pedunculate,
in bud, splitting into 2 or 3 valvate lobes; petals 0; dichasial, the pistillate flowers central; bracts
disk segments interstaminal or 0; stamens 28–30, minute, deciduous. Staminate flowers pedicellate;
filaments distinct, shorter than anthers; anthers sepals 5 (6), distinct, imbricate; petals 5 (6),
basifixed, introrse, connective apically glandular; distinct, imbricate, pubescent abaxially; disk seg-
pollen grains spheroidal, inaperturate, with Croton ments 5; stamens 10–25, outer filaments distinct
pattern; pistillode 0. Pistillate flowers pedicellate; but inner connate into a column, not inflexed
sepals 5–7, imbricate, persistent and accrescent in bud; anthers dorsifixed, introrse, muticous;
in fruit; petals 0; disk annular, pubescent; ovary pollen grains spheroidal, inaperturate, eine
mainly (2)3(4)-locular, stellate-tomentose; ovules with Croton pattern; pistillode 0. Pistillate flowers
anatropous, inner integuments very thick, vascu- pedicellate; sepals 5 (6), distinct, imbricate,
larized; stylodia bifid. Fruits smooth to tubercu- persistent in fruit; petals 5 (6), distinct, imbricate;
late, dehiscing septicidally; columella trigonous, disk annular, slightly lobed, glabrous; ovary
persistent. Seeds ellipsoidal, ecarunculate, testa 3-locular, pubescent; ovules pachychalazal, inner
smooth; endosperm oily. integuments thick, vascularized; stylodia nearly
Three spp., endemic to Madagascar. distinct, deeply bifid, branches slender. Fruits
capsular, cocci thin-walled; columella persistent.
12f. SUBTRIBE PARACROTONINAE G.L. Webster1 Seeds subspheroidal, caruncle subspherical,
Monoecious trees or shrubs; leaves dentate; appressed; testa crustaceous, smooth.
sepals and petals 5(6), distinct; staminate disk of Three or four spp., tropical Africa, from
distinct glands, pistillate disk annular; stamens Liberia and Gabon to Kenya and Mozambique.
258. Paracroton Miq.

1
Subtr. Paracrotoninae G.L. Webster, subtrib. nov., monoici;
inflorescentiis axillaribus; fructibus capsularis; sepalis foe-
Paracroton Miq., Fl. Ind. Batav. 1(2): 382 (1859); M€ ull.
mineis persistentibus; petalis connatis vel liberis. Typus:
Arg. in DC., Prodr. 15(2): 1112 (1866); J.J. Sm., Meded.
Paracroton Miquel.
Dept. Landb. 12: 585 (1910); Pax, Pflanzenr. 147, III: 12
Euphorbiaceae 189
narrowly thyrsoid; pistillate inflorescences termi-

nal, thyrsoid, sparsely branched, flowers 1–3 per
node; bracts deciduous. Staminate flowers subses-
sile; sepals 5, slightly connate, imbricate, dorsally
often with knob- or horn-like appendage; petals 5,
distinct; disk segments 5, 7 or 10; stamens 12–32,
outer filaments distinct, inner ones connate into a
column; anthers dorsifixed, extrorse, muticous;
pollen grains spheroidal, inaperturate, with Cro-
ton pattern. Pistillate flowers pedicellate; calyx as
in staminate, persistent in fruit, petals 5, coherent,
glabrous, deciduous; disk annular, lobed, pilose;
ovary 3-locular, pubescent; stylodia bifid. Fruits
capsular. Seeds spheroidal, ecarunculate, testa
smooth or striate, slightly fleshy.
Four spp., tropical Asia from Sri Lanka to
W Malesia, the Philippines, and New Guinea.
Balakrishnan and Chakrabarty (1993) believe
that the closest allies of Paracroton are Ostodes
and Dimorphocalyx.
Unplaced genus of inaperturate crotonoids:
259. Radcliffea P. Hoffm. & K. Wurdack

Radcliffea P. Hoffm. & K. Wurdack, Kew Bull. 61: 194
(2006).
Dioecious shrub or tree, probably latescent; indu-
mentum stellate. Leaves long-petiolate, domatii-
ferous, at base palminerved, at junction with
petiole provided on both sides with minute
glands; stipules apparently sometimes replaced
by minute glands. Inflorescences terminal, panic-
ulate, with 3–5 orders of branching. Staminate
Fig. 42. Euphorbiaceae-Crotonoideae. Mildbraedia carpi- flowers pedicellate, sepals 5, distinct, imbricate,
nifolia var. strigosa. A Distal portion of fruiting branch.
B Distal portion of branch with staminate flowers.
outermost bearing a small abaxial gland; petals 5,
C Staminate flower. D Petal. E Pistillate flower. F Pistil. distinct, imbricate; disk glands 5, distinct;
G Fruit. (Radcliffe-Smith 1996; drawn by J.M. Fothergill) stamens 5, with the filaments connate for more
than half their length into a column; pollen grains
spheroidal, inaperturate, with Croton pattern;
(1911); N.P. Balakr. & Chakrab., Kew Bull. 48: 716, figs. pistillode +. Pistillate flowers pedicellate, sepals
1–3 (1993); Radcl.-Sm., Gen. Euphorb.; 318 (2001). 5, distinct, imbricate, all with abaxial gland; petals
Fahrenheitia Rchb. f. & Zoll. (1857). 4–7, distinct, imbricate; disk-glands 5, distinct or
Desmostemon Thwaites (1861). irregularly connate; ovary 1-locular, asymmetri-
cal; ovule 1, anatropous, inserted subapically;
Monoecious trees; latex not recorded; indumen- stylodia 3, inserted excentrically, simple, unequal.
tum stellate or lepidote. Leaves petiolate (pulvi- Fruits and seeds unknown.
nate), glandular-dentate or serrate, biglandular at A single sp., R. smithii P. Hoffm. &
junction with petiole; stipules represented by ses- K. Wurdack, from deciduous forests on limestone
sile glands. Inflorescences terminal, unisexual; in W Madagascar.
staminate inflorescences terminal and axillary,
190 G.L. Webster
VII. SUBFAM. EUPHORBIOIDEAE – Inflorescences pseudanthial, i.e., cyathial, usually with a

single terminal pistillate flower, and 4 or 5 lateral sta-
Monoecious (dioecious) trees, shrubs, or herbs minate monochasia or dichasia; perianth reduced or 0;
stylodia mostly bifid; colpi bordered by two conspicu-
(scandent); laticifers non-articulate, latex mostly ous intine thickenings 3. Euphorbieae
whitish; indumentum simple or 0, dendritic in
Mabea and Senenefelderopsis. Leaves alternate 1. TRIBE STOMATOCALYCEAE (M€
ull. Arg.)
or opposite, simple, unlobed, entire or dentate, G.L. Webster (1975).
usually pinnately veined, often with basal glands
at junction with petiole; stipules often reduced or Dioecious trees, shrubs, or lianas; latex yellowish,
0. Inflorescences terminal or axillary, spiciform to often scanty; indumentum simple or 0; leaves
racemoid or paniculate or condensed into the alternate, unlobed, pinnately veined, without
pseudanthial cyathium; bracts often biglandular glands at base of blade; stipules small and cadu-
at base. Flowers apetalous, erect or inclinate in cous or 0; inflorescences axillary, racemoid, bracts
bud; staminate sepals (1)3–6, imbricate to val- eglandular; staminate sepals 4–8, distinct and
vate, mostly open in bud, commonly minute or imbricate or connate; stamens 10–30, distinct, fila-
0; disk 0; stamens 1–20(80), filaments distinct ments shorter than anthers; pollen grains coarsely
or connate; pollen grains binucleate or trinucle- reticulate or reticulate-perforate; pistillate sepals
ate, 3-colporate, colpi usually marginate, exine 5 or 6, distinct or connate; ovary 2–10-locular;
usually tectate-perforate; pistillode 0; pistillate stylodia unlobed, sometimes dilated or stigmati-
sepals 3–6, distinct or connate, imbricate or form; fruit capsular or indehiscent, 1–3-seeded;
open in bud, sometimes reduced to obsolete; seeds ecarunculate; endosperm copious, oily.
disk 0; ovary 2–3(20)-locular; stylodia distinct Four genera and about 12 spp., pantropical.
or connate, nearly always unlobed. Fruits capsu-
lar (drupaceous). Seeds carunculate or ecaruncu- KEY TO THE GENERA OF STOMATOCALYCEAE
late; testa dry or fleshy, exotegmen palisadal; 1. Fruits indehiscent; stylodia abbreviated; ovary 1–10-
endosperm copious. locular; pollen grains finely reticulate-perforate 2
Molecular analyses (Wurdack et al. 2005; – Fruits capsular; stylodia elongated; ovary 2–3-locular;
Tokuoka 2007) confirm the broad lines of the pollen grains coarsely reticulate 3
classification of this subfamily and the inclusion 2. Sepals 6–8, distinct; ovary 1-locular 262. Plagiostyles
of the Stomatocalyceae, which had been sug- – Sepals connate, calyx 2-lipped; ovary 2–10-locular
263. Pimelodendron
gested by Webster (1975) and accepted by Rad- 3. Ovary 3-locular; stylodia erect; stamens 18–20; pistil-
cliffe-Smith (2001); the Stomatocalyceae are late sepals not glandular; stems scandent
resolved as sister to the rest of the subfamily. 261. Hamilcoa
This is divisible into two subclades, which con- – Ovary 2-locular; stylodia spreading; stamens 10; pistil-
tain the Hippomaneae and the Euphorbieae, the late sepals with large basal glands; stems not scandent
latter broadened by inclusion of the Hureae and 260. Nealchornea
Pachystromateae.
260. Nealchornea Huber
Nealchornea Huber, Bol. Mus. Goeldi 7: 297 (1913);
KEY TO THE TRIBES OF SUBFAM. EUPHORBIOI- Ducke, Arch. Jard. Bot. Rio de Janeiro 3: 201, t. 10
DEAE ofer, Linzer Biol. Beitr. 23(2): 777 (1991);
(1922); Walln€
1. Pollen exine tectate-reticulate; bracts eglandular, dis-
tinct from rachis; stylodia unlobed; seeds ecarunculate;
dioecious trees or lianas 1. Stomatocalyceae Dioecious trees; latex whitish or yellowish; indu-
– Pollen exine tectate-perforate; bracts glandular, often mentum simple, scanty. Leaves remotely glandu-
adnate to rachis; stylodia bifid or unlobed; seeds carun- lar-crenate, with small abaxial basal or
culate or ecarunculate; monoecious (dioecious) trees, submarginal glands; stipules obsolete. Inflores-
shrubs, or herbs, rarely scandent 2 cences axillary and subterminal, the staminate
2. Inflorescences mostly racemoid or spiciform, some- paniculate, the pistillate racemose; bracts egland-
times capitate but never cyathial; flower buds usually
inclinate; staminate calyx usually well developed; ular. Staminate flowers pedicellate; sepals 4, biseri-
stylodia mostly unlobed; intine thickenings along the ate, slightly imbricate, open in bud; stamens 8–15
colpi 0 2. Hippomaneae s. l. (18), distinct; anthers sessile, erect, dehiscing
Euphorbiaceae 191
longitudinally; connective enlarged, conspicuously Pistillate flowers pedicellate; sepals 5, unequal,

appendiculate terminally; pollen grains oblate imbricate, entire, persistent in fruit; ovary
spheroidal, 3-colporate, sexine coarsely reticulate. 1(2)-locular; ovules anatropous, inner integu-
Pistillate flowers pedicellate; sepals 4, basally con- ments moderately thick; stylodia lateral with
nate, slightly imbricate, glandular, persistent in discoid stigma. Fruit drupaceous; endocarp
fruit; ovary 2-locular; ovules anatropous, inner membranaceous, adherent to seed. Seeds ecarun-
integuments moderately thick, outer integuments culate, transversally veined; endosperm yellow-
vascularized; stylodia unlobed, thickened, slightly ish, oily; cotyledons reniform.
connate at base. Fruits baccate, not regularly Only one sp., P. klaineana Pierre, from the
dehiscent, thin-walled. Seeds ecarunculate. rainforests in Nigeria, Gabon, and Congo.
Two spp., lowland Amazonian forests of
Brazil and adjacent Peru and Colombia. 263. Pimelodendron Hassk.
Pimelodendron Hassk., Versl. Med. Afd. Natuurk. Kon.
261. Hamilcoa Prain Akad. Wetensch. 4: 140 (1856); J.J. Sm., Bull. Jard. Bot.
Hamilcoa Prain, Kew Bull. 1912: 107 (1912), Fl. Trop. Afr. Buit. III, 6: 100 (1924); Airy Shaw, Kew Bull. 36: 339
6(1): 1000 (1913); Stapf, Hook. Icon. Pl. 31: t. 3990 (1915); (1981).
Keay, Fl. W. Trop. Afr., ed. 2, 1: 413 (1958).
Dioecious trees; latex whitish or yellowish; indu-
Dioecious lianas; latex white; indumentum 0. mentum 0 on twigs. Leaves subentire or crenate,
Leaves simple or occasionally lobed, entire or eglandular; stipules minute, deciduous. Inflores-
subentire, eglandular; stipules minute, caducous. cences axillary, sometimes cauliflorous, race-
Inflorescences racemoid, staminate axillary, pistil- moid; bracts eglandular. Staminate flowers
late terminal; bracts eglandular, 1-flowered. pedicellate; sepals connate into a 2-lipped calyx;
Staminate flowers pedicellate; sepals 5, distinct, stamens 10–16, distinct or coherent, shorter than
imbricate; stamens 18–30, distinct, filaments anthers; anthers basifixed, extrorse, dehiscing
much shorter than anthers; anthers basifixed, longitudinally; pistillode 0. Pistillate flowers
the exterior extrorse, dehiscing longitudinally; pedicellate; calyx cupular, 2–3-lobed, persistent
pollen grains coarsely reticulate. Pistillate flowers in fruit; ovary 2–10-locular; ovules anatropous,
with thick reflexed pedicels; sepals 6, biseriate, inner integuments thick; outer integuments vas-
imbricate, subentire; ovary 3-locular; stylodia cularized; stylodia coalescent into a stigmatoid
connate at base, erect, dilated, unlobed. Fruits apex. Fruit indehiscent, fleshy. Seed solitary, car-
capsular. Seeds with spongy testa. unculate, subspheroidal, testa striate-reticulate.
Only one sp., H. zenkeri Prain, from lowland Six to eight spp. from tropical Asia and
forests in Nigeria and Cameroon. Australia.
262. Plagiostyles Pierre 2. TRIBE HIPPOMANEAE A. Juss. ex Spach (1834).

Plagiostyles Pierre, Bull. Mens. Soc. Linn. Paris 2: 1326 Hureae Dumort. (1829).
(1897); Prain, Fl. Trop. Afr. 6(1): 1001 (1913); Stapf, Pachystromateae (Pax & Hoffm.) Pax (1924).
Hook. Icon. Pl. 31: t. 3010 (1915); Léonard, Fl. Congo 8
Monoecious, less commonly dioecious trees,
(1): 131 (1962); Radcl.-Sm., Gen. Euphorb.: 348, fig. 44
(2001). shrubs, or herbs; latex usually milky, sometimes
toxic; indumentum simple or 0, dendritic in
Dioecious trees or shrubs; latex white, scanty; Mabea; leaves alternate (opposite), simple and
indumentum simple. Leaves eglandular or with unlobed; lamina pinnately (palmately) veined,
small paired adaxial glands; stipules entire, decid- commonly with laminar or petiolar glands;
uous. Inflorescences axillary, paniculate or race- stipules sometimes reduced or 0; inflorescences
moid; bracts eglandular, 1-flowered. Staminate terminal or axillary, racemoid or spiciform
flowers pedicellate; sepals 5–8, unequal, imbricate (paniculate), usually bisexual with 1 or 2 pistillate
but not covering the stamens in bud; stamens flowers at proximal nodes; bracts mostly glandu-
15–32, distinct, aggregated-capitulate, filaments lar; flowers usually inclinate in bud; staminate
much shorter than anthers; anthers extrorse, calyx usually open in bud, sepals 0–3, distinct or
reddish, dehiscing longitudinally; pistillode 0. connate, imbricate to valvate; stamens 1–70,
192 G.L. Webster
filaments distinct or connate proximally; anthers 3. Ovary 5–20-locular; stamens 10–80, connate; tips of
extrorse; pollen grains 3-colporate, colpi usually stylodia radiating from apical disk of column; seeds
marginate, exine tectate-perforate: pistillate strongly compressed 279. Hura
sepals 3–6, imbricate, sometimes reduced or 0; – Ovary 3-locular; stamens 1–3, distinct or connate; sty-
lodia connate but tips not radiating from a disk 4
ovary 2–3(10)-locular; stylodia distinct or con- 4. Stylodia connate 1/3 to 2/3 their length, not distally
nate, unlobed, in one genus the stigmas connate clavate; tips of staminal sepals distinct, imbricate;
into an umbrella-shaped disk; fruits capsular or staminate spikes mostly terminal
drupaceous; seeds carunculate or ecarunculate, 280. Algernonia
testa dry or fleshy; endosperm copious, not oily. – Stylodia connate their entire length, distally clavate with
A mainly neotropical tribe of about 30 genera, deltoid apices; staminal sepals completely connate into
a cup; staminate spikes axillary
broadened here to include tribes Pachystroma- 281. Ophthalmoblapton
teae and Hureae. In the analysis of Wurdack 5. Flowers erect in bud; staminate calyx closed in bud,
et al. (2005), the monotypic Pachystromateae splitting valvately into 2 segments; filaments and
are embedded in the Hureae, and the Hureae anthers connate; seeds ecarunculate; leaves spinose-
form part of one of the two strongly suppor- dentate 282. Pachystroma
ted subclades representing the Hippomaneae. – Flowers inclinate in bud; staminate calyx open in bud;
stamens distinct or filaments connate; seeds caruncu-
This confirms earlier expectations by Webster
late or ecarunculate; leaves mostly not spinose-den-
(1994b), who doubted the justification of tribal tate [Hippomaneae s. str.] 6
distinctness of Pachystroma and pointed to the 6. Staminate calyx laterally compressed or zygomorphic
closeness of the Hureae to the Hippomaneae inclinate, with 1 or 2 distinct lobes; stylodia apically
(which had been included in that tribe as subtribe glandular and often bifid 271. Homalanthus
Hurinae by previous authors). The subtribal divi- – Staminate calyx radially symmetric or rarely zygo-
sion of the Hippomaneae by Webster (1994) and morphic-inclinate, with 2 or more lobes or completely
connate; stylodia apically undivided and eglandular
Esser (2001) is not confirmed by the molecular 7
data and their subtribes are not maintained 7. Staminate flowers with 5(6) sepals; stylodia usually
here, but in the molecular analysis two subclades connate into a long common style with slender
(H1 and H2) are recognized that cannot be char- branches; leaves and stems with dendritic hairs
acterized morphologically. The classification of 275. Mabea
the Hippomaneae presents some of the most – Staminate flowers with 0–3(4) sepals, calyx with more
or less connate lobes, or without distinct lobes, or
intractable problems in the family, and both
calyx lacking at all 8
the delimitation of some larger genera such as 8. Ovary 6–10-locular; bracts glandular
Excoecaria, Gymnanthes, Sapium, and Sebastiana 283. Hippomane
and the justification of several recently described – Ovary 2- or 3-locular; bracts glandular or eglandular
small genera remain controversial. Overall, the 9
preponderance of small genera (five monotypic, 9. Seed testa dry [at most with thin fleshy layer] 10
five others with only two or three species) is – Seed testa fleshy [arillate] 28
striking and appears as the work of excessive 10. Columella 3-horned at base 11
splitting. – Columella not 3-horned at base 12
11. Staminate calyx 2-lobed; petiole glandular near apex
289. Stillingia
– Staminate calyx 0; petiole eglandular
KEY TO THE GENERA OF TRIBE HIPPOMANEAE 287. Adenopeltis
1. Leaves present only on young branchlets, plants later 12. Leaf blades palmately 3–11-lobed; staminate flower
leafless, succulent, thorny shrubs, flowering when with 1 stamen 270. Dalembertia
leafless 288. Spegazziniophytum – Leaf blades simple; staminate flower at least with
– Leaves present on older branches; flowering with 2 stamens 13
leaves or, if flowering when leafless, then not a succu- 13. Staminate flowers in a glomerulate head; upper part of
lent thorny shrub 2 seed covered by an enlarged caruncle; leaves long-
2. Floral bracts eglandular, peltate or adnate to the pedicellate 264. Maprounea
rachis and covering the flowers; seeds ecarunculate – Staminate flowers in an elongated inflorescence; car-
3 uncle 0 or small on top of seeds 14
– Floral bracts 2-glandular at the base or eglandular, not 14. Pistillate flowers sessile or subsessile, pedicel usually
peltate nor adnate to the rachis 5 < 5 mm long in fruit; inflorescences mostly terminal,
sometimes axillary as well 15
Euphorbiaceae 193
– Pistillate flowers pedicellate, pedicel usually > 5 mm – Inflorescence buds sheathed by the stiff, scaly floral
long in fruit, or else ovary appendiculate; inflores- bracts, not by sterile scales; stamens 2–6
cences mostly axillary 27 269. Gymnanthes
15. Inflorescences axillary; leaves alternate or opposite; 28. Inflorescences unisexual; fruits irregularly dehiscent;
calyx lobes of staminate flowers distinct or nearly so; seeds whitish arillate 290. Falconeria
mostly dioecious 293. Excoecaria – Inflorescences bisexual 29
– Inflorescences terminal or axillary; leaves mostly alter- 29. Seed aril reddish; flowers sessile or subsessile; leaf
nate, opposite in Microstachys and Colliguaja, calyx blades pinnately veined 291. Sapium
lobes connate; mostly monoecious 16 – Seed aril whitish; flowers pedicellate; leaf blades tri-
16. Inflorescences mostly opposite leaves and with stami- plinerved 268. Triadica
nate flowers distichous; ovary with 2 vertical rows of
appendages on each locule; seeds cylindric, truncate, 264. Maprounea Aubl.
with a stipitate caruncle 272. Microstachys
– Inflorescences terminal; staminate flowers spiral on Maprounea Aubl., Hist. Pl. Guiane: 895, t. 342 (1775); Pax &
inflorescence axis; ovary not with 2 rows of K. Hoffm., Pflanzenr. 147, V: 175, fig. 32, 33 (1912);
appendages on each locule; seeds neither cylindric Léonard, Fl. Congo 8(1): 142, fig. 9 (1962); Radcl.-Sm., Fl.
nor truncate, mostly ecarunculate 17 Trop. E. Africa, Euphorb. 1: 395, fig. 75 (1987); Webster &
17. Floral bracts with basal glands, or else stipules cadu- Huft, Ann. Missouri Bot. Gard. 75: 1131 (1988); Esser,
cous; fruits capsular or indehiscent 18 Novon 9: 32, fig. 1 (1999).
– Floral bracts without basal glands; stipules persistent, Monoecious (dioecious) trees or shrubs; latex
ovate, ciliate; fruits capsular 23
18. Staminate calyx completely connate without distinct whitish; indumentum 0. Leaves usually with
lobes; stamens 4–17 267. Senefeldera prominent subbasal laminar glands abaxially;
– Staminate calyx of 3 basally connate calyx lobes; petiole eglandular; stipules entire, eglandular,
stamens 2–3 19 persistent. Inflorescences terminal, usually bisex-
19. Staminate flowers 5–9 per bract 20 ual, with 1–5 proximal pistillate flowers separated
– Staminate flowers 1–3 per bract 22 from an ament-like staminate subflorescence;
20. Sepals and stamens 2 per flower; ovary 2-locular, not
appendaged; fruit drupaceous 278. Balakata
bracts biglandular at base, the staminate subtend-
– Sepals 3, stamens 2–3 per flower; fruit capsular 21 ing 1–3(5) flowers. Staminate flowers sessile to
21. Leaves eglandular above; ovary and fruit with 3 pairs subsessile; sepals 2–3, connate into a cupular
of appendages 274. Sclerocroton calyx; stamens (1)2–3, filaments connate into
– Leaves above often with a pair of petiolar glands at a column exserted beyond the calyx; anthers
the junction with the blade; ovary and fruit not extrorse, dehiscing longitudinally. Pistillate flow-
appendaged 284. Pleradenophora ers pedicellate, spreading or recurving; sepals 3,
22. Stylodia distinct or nearly so; staminate flowers sessile
to subsessile; leaves serrate 276. Sebastiana discrete, persistent in fruit; ovary 3-locular,
– Stylodia proximally connate into a style, style smooth; ovules anatropous, inner and outer inte-
branches undivided; staminate flowers shortly but guments thin (5 or 6 cell layers); stylodia connate
distinctly pedicellate; leaves entire 273. Ditrysina into a common style [except in M. amazonica],
23. Lobes of staminate calyx 2–3, evident though basally style branches unlobed. Fruits capsular; colu-
connate 24 mella slender, not persistent. Seeds ellipsoid,
– Lobes of staminate calyx 1 or 2 and minute or with a cap-like caruncle covering the upper half,
completely reduced 26
24. Inflorescence axillary; ovary 2–3-locular and with 4 or testa smooth or foveolate.
6 appendages 277. Anomostachys Five spp., three in South America and two in
– Inflorescence terminal; ovary 3-locular, unappen- Africa.
daged 25
25. Staminate flowers 6–20 per bract; stamens 2–5, fila- 265. Actinostemon Mart. ex Klotzsch
ments distinct; indumentum simple, often colored
266. Senefelderopsis Actinostemon Mart. ex Klotzsch, Arch. Naturg. 7: 184
– Staminate flowers 2–4 per bract; stamens 3, filaments (1841); Pax & K. Hoffm., Pflanzenr. 147, V: 57, figs.
connate into a column; plants glabrous 10–14 (1912); Jablonski, Phytologia 18: 213–240 (1969);
286. Grimmeodendron Burger & Huft, Fieldiana Bot. II, 36: 57 (1995); Berry &
26. Inflorescences axillary; leaves (sub)opposite Esser, Fl. Venez. Guayana 5: 87, fig. 84 (1999).
292. Colliguaja
Monoecious trees or shrubs; latex scanty; indu-
– Inflorescences on lateral short shoots; leaves alternate
285. Bonania mentum simple, often scanty. Leaves alternate or
27. Inflorescence buds covered by caducous bud-scales; pseudo-verticillate, with dispersed glands abaxi-
floral bracts irregular and weak, often nearly 0; ally; stipules deciduous. Inflorescences terminal
stamens 4–16 265. Actinostemon and sometimes axillary, racemoid, and in bud
194 G.L. Webster
ament-like, enclosed by imbricate bud scales, axis thrix does not appear to be strongly different
glabrous or hirtellous; pistillate flowers proximal, from Senefelderopsis s. str., and there seems to
1–3 per bract; staminate flowers distal, 2 or 3 per be no reason why it cannot be accommodated
bract; bracts biglandular (eglandular) at base, as a section of that genus.
lamina minute or obsolete. Staminate flowers
pedicellate; calyx rudimentary or 0; stamens 267. Senefeldera Mart.
4–15 or more, filaments distinct; anthers basi- Senefeldera Mart., Flora 24 (Beibl. 2): 29 (1841); Jablonski,
fixed, extrorse, dehiscing longitudinally; Mem. N. Y. Bot. Gard. 12: 171 (1965); Webster, Ann.
pollen grains 3-lobed, 3-colpate, exine tectate- Missouri Bot. Gard. 75: 1127 (1989); Murillo & Franco,
perforate; pistillode 0. Pistillate flowers pedicel- Euforb. Reg. Araracuara 149, fig. 45 (1995); Esser in
late, pedicel elongating in fruit; sepals minute Radcl.-Sm., Gen. Euphorb.: 389 (2001).
Rhodothyrsus Esser (1999).
or 0; ovary 3-locular, sometimes appendaged; Pseudosenefeldera Esser (2001).
stylodia unlobed, connate basally. Fruits capsu-
lar; columella persistent. Seeds spheroidal, sub- Monoecious trees; latex white; indumentum sim-
apically carunculate, testa smooth. ple, dibrachiate or 0. Leaves sometimes pseudo-
About 15 spp. ranging from the Caribbean verticillate, abaxially with scattered (sometimes
to South American. The genus was combined marginal) glands; stipules caducous. Inflorescences
by Webster (1994) with the superficially similar axillary, pedunculate, usually bisexual, compound
Gymnanthes but this is not supported by the thyrsoid, floral bracts with a pair of elliptic glands;
molecular evidence. pistillate flowers solitary at 1–4 proximal nodes,
glomerules of 1–3 staminate flowers at distal
266. Senefelderopsis Steyerm. nodes. Staminate flowers pedicellate to sessile or
subsessile; calyx completely connate without dis-
Senefelderopsis Steyerm., Bot. Mus. Leafl. Harvard Univ.
15: 45, t. 16 (1951); Jablonski, Mem. N. Y. Bot. Gard. 12: tinct lobes, sometimes zygomorphic; stamens (2)
174 (1965); Gillespie, Brittonia 45: 92 (1993); Murillo & 4–17, filaments distinct; anthers dehiscing longitu-
Franco, Euforb. Reg. Araracuara 153, fig. 46 (1995); Esser dinally. Pistillate flowers pedicellate to subsessile;
in Radcl.-Sm., Gen. Euphorb.: 376 (2001). sepals 3, distinct or basally connate; ovary 3-locu-
Dendrothrix Esser (1993). lar, smooth, glabrous; ovules anatropous, inner
Monoecious trees or shrubs; latex white; indu- integuments moderately thick (8–10 cell layers),
mentum simple, on inflorescence axes sometimes outer integuments thin (3–5 cell layers), non-vas-
dendritic. Leaves with paired adaxial or abaxial cularized; stylodia proximally usually connate into
glands; stipules caducous. Inflorescences bisex- a short style with 3 distal stigmatic branches. Fruits
ual, terminal, compound, axes thyrsoid; floral spheroidal, capsular; columella triquetrous, dis-
bracts eglandular, subtending proximal solitary tinctly alate, persistent. Seeds subspheroidal, ecar-
pistillate flowers and distal cymules of 6–20 sta- unculate, testa smooth or sculptured.
minate flowers on each axis. Staminate flowers Six spp., tropical South America from Panama
subsessile to pedicellate; sepals 2 or 3, basally south to Peru. The two segregate genera proposed
connate; stamens 2–5, filaments distinct or by Esser have some distinctive characters, but
connate, as long as anthers. Pistillate flowers they share a very similar Gestalt and it does not
short-pedicellate; sepals 3(6), basally connate; seem necessary to dismember a genus as small
ovary 3-locular, pubescent; ovules anatropous, as Senefeldera. With some modification, the seg-
inner integuments moderately thick, outer inte- regate taxa could be fitted into the sectional
guments thick, non-vascularized; stylodia undi- arrangement of Pax and Hoffmann (1912).
vided, basally connate into a short common style.
Fruits capsular, sometimes partly fleshy; colu- 268. Triadica Lour.
mella persistent, distally alate; seeds carunculate Triadica Lour., Fl. Cochin. 2: 598, 610 (1790); Small, Man.
or ecarunculate. Southeastern Flora 789 (1933); Hurus., J. Fac. Sci. Univ.
Five spp., centered on the Guayana highlands Tokyo, III. 6: 315 (1954); Kruijt, Bibl. Bot. 146: 7 (1996);
Esser, Harvard Papers Bot. 7: 17–21 (2002).
and extending to its southern and eastern
foreland. Except for the number of staminate Monoecious trees; latex white; indumentum 0.
sepals and the dendritic indumentum, Dendro- Leaves entire with submarginal glands and adaxial
Euphorbiaceae 195
paired glands at junction with petiole; stipules or connate; ovary 3-locular, smooth or appendi-
entire, persistent. Inflorescences terminal, bisex- culate; ovules anatropous, inner integuments
ual, spiciform; bracts biglandular at base, subtend- moderately thick (8–10 cell layers), outer integu-
ing solitary pistillate flowers and distal cymules of ments thin (4–6 cell layers); stylodia distinct or
3–8 staminate flowers. Staminate flowers pedicel- basally connate, simple. Fruits capsular (indehis-
late; sepals 3, basally connate; stamens 2 or 3, cent); columella triquetrous, alate, persistent.
filaments distinct; anthers basifixed, extrorse, Seeds elliptic, smooth, caruncle small or 0.
dehiscing longitudinally; pollen grains 3-nucleate. About 45 spp., most in the New World from
Pistillate flowers pedicellate; sepals 3, basally con- the southern USA through the Antilles to South
nate, deciduous in fruit; ovary 3-locular; ovules America, few spp. in Africa, S and SE Asia, Malesia,
anatropous, inner integuments moderately thick, and extending to Micronesia, Melanesia and Japan.
outer integuments thick; stylodia connate at base, Apart from the exclusion of Actinostemon,
unlobed. Fruits capsular, dehiscing into 6 valves; Esser (2001) has modified the circumscription of
columella persistent, alate. Seeds ecarunculate, Gymnanthes by including Sarothrostachys,
adherent to columella, sarcotesta whitish. 2n ¼ 44. Duvigneaudia, and Adenogyne. In the treatment
Three spp., India, China, and Malesia. The presented here, Gymnanthes is only enlarged by
genus has usually been included in Sapium, but the inclusion of his segregate genera Neoshirakia
differs in its pedicellate flowers and the whitish and Shirakiopsis (see Esser, Blumea 44: 165–172.
(not reddish) sarcotesta. 1999). The boundaries of Gymnanthes have not
yet been established satisfactorily, and it remains
269. Gymnanthes Sw. to be seen whether they will be decreased to
Gymnanthes Sw., Prodr. Veg. Ind. Occ.: 95 (1788); Grise-
exclude Neoshirakia.
bach, Fl. Brit. W. Ind.: 50 (1859); Bentham, Gen. Pl. 3: 337
(1880); Sargent, Silva N. Amer. 7: t. 309 (1995); Pax & 270. Dalembertia Baill.
K. Hoffm., Pflanzenr. 147, V: 81 (1912); Fawc. & Rend., Fl.
Jam. 4: 329, fig. 111 (1920); Webster, J. Arnold Arb. 48: 387 Dalembertia Baill., Étude Gén. Euphorb.: 545 (1858);
(1967), Taxon 32: 304 (1983), Ann. Missouri Bot. Gard. 75: M€ull. Arg. in DC., Prodr. 15(2): 1225 (1866); Standley,
1129 (1989), 81: 122 (1994); Howard, Fl. Lesser Ant. 5: 52, Contr. U. S. Nat. Herb. 23: 646 (1923); Standley &
figs. 16, 18 (1989); Esser in Radcl.-Sm., Gen. Euphorb.: 382 Steyerm., Fieldiana Bot. 24 (6): 86 (1949); Webster, Ann.
(2001); Burger & Huft, Fieldiana n.s. 36: 123 (1995); Berry Missouri Bot. Gard. 81: 122 (1994); Esser in Radcl.-Sm.,
& Esser, Fl. Ven. Guayana 5: 149 (1999); Webster, Fl. Gen. Euphorb.: 391 (2001).
Nicaragua 1: 884 (2001). Monoecious herbs or subshrubs, often with
Shirakia Hurus. (1954), nom. illeg.
Neoshirakia Esser (1998). tuberous roots; latex white; indumentum simple,
Shirakiopsis Esser (1999). multiseriate. Leaves usually palmately 3–11-
lobed, entire or distantly serrate; petioles egland-
Monoecious or dioecious trees or shrubs, ular; stipules persistent. Inflorescences terminal,
branches sometimes spinose; latex whitish, some- bisexual, with pedicellate proximal pistillate flow-
times not evident; indumentum simple or 0. ers and a spiciform ament of staminate flowers;
Leaves entire or serrate, egandular adaxially, bracts biglandular, subtending solitary pistillate
abaxially with marginal (scattered) glands; sti- flowers and cymules of 1–3 staminate flowers.
pules entire, sometimes glandular. Inflorescences Staminate flowers pedicellate; calyx completely
usually bisexual, racemoid, terminal and axillary, connate without distinct lobes, zygomorphic; sta-
solitary, sometimes with minor proximal men solitary, filament long-exserted; anthers lon-
branches; bracts sometime pedunculate, usually gitudinally dehiscent; pistillode 0. Pistillate
biglandular at base; pistillate flowers solitary at flowers with elongated recurved pedicels;
proximal 1–3 nodes, staminate flowers in (1)3–5- sepals 3, distinct, entire, deciduous in fruit;
flowered cymules. Staminate flowers pedicellate; ovary 3-locular, smooth; ovules anatropous,
sepals 1–2, often reduced or obsolete; stamens (2) inner integuments moderately thick, outer inte-
3–6, filaments often longer than anthers; anthers guments thin; stylodia connate into a common
basifixed, extrorse, longitudinally dehiscent; pol- style with 3 apical unlobed, recurved tips. Fruits
len grains 3-nucleate. Pistillate flowers pedicel- capsular; columella 3-angled, persistent. Seeds
late, often expanding in fruit; sepals 3, distinct smooth, ecarunculate.
196 G.L. Webster
Four or five spp. native to Mexico and Gua-

temala, where they grow in deciduous woodland.
Their inflorescences, with basal recurved long-
pedicellate pistillate flowers subtending an
ament of staminate flowers, are strikingly similar
to those of Maprounea. However, no relationship
between the two genera is discernible in the
molecular analysis of Wurdack et al. (2005).
271. Homalanthus A. Juss.

Homalanthus A. Juss., Tent. Euphorb.: 50 (1824) (Oma-
lanthus, orth. rej.); Airy Shaw, Kew Bull. 21: 409 (1968),
Kew Bull. Add. Ser. 8: 115 (1980); McPherson & Tirel, Fl.
Nouv.-Caléd. 14(1): 25, t. 4 (1987); Forster, Telopea 6: 169
(1994); Florence, Fl. Polynes. Franc. 1: 115, fig. 20 (1997).
Monoecious or dioecious trees or shrubs; exudate
watery, innocuous; indumentum simple or 0.
Leaves often peltate, without laminar glands;
petioles with paired glands at junction with lam-
ina; stipules caducous. Inflorescences terminal
and sometimes axillary as well, bisexual or uni-
sexual, spiciform with pistillate flowers at base or
at separate axils; bracts uniflorous, biglandular at
base. Staminate flowers pedicellate; calyx laterally
compressed, sepals 2, cordate, rarely 1, subimbri-
cate; stamens (5–)10–50, filaments distinct;
anthers extrorse, dehiscing longitudinally; pollen
grains 2-nucleate; pistillode 0. Pistillate flowers
long-pedicellate; sepals 2 or 3, deciduous in
fruit; ovary 2(3)-locular; ovules anatropous, the
inner integuments thin, the outer thick and non-
vascularized; stylodia distinct or basally connate, Fig. 43. Euphorbiaceae-Euphorbioideae. Microstachys
unlobed, often glandular at apex. Fruits capsular chamaelea. A Flowering branchlet. B Detail of leaf margin
or indehiscent; columella persistent. Seeds with with densely packed glandular teeth. C Stipulate node with
hypertrophied arilloid caruncle. 2n ¼ 36, 44. young fruit and inflorescence. D Subsessile staminate
flower with large bract glands and distinct sepals.
About 23 spp., distributed from SE Asia to E Infructescence. F Fruit with excrescences. G Partly opened
Australia, New Caledonia, New Zealand, and fruit with alate columella. H Mericarp with the large septa.
Polynesia. The resemblance of this genus with I Carunculate seed. (Esser 1999; drawn by J. van Os)
the Stomatocalyceae has been noted since the
time of M€ uller (1866), who included Homa- Monoecious herbs or subshrubs; latex not evident;
lanthus in Pimelodendron (as Carumbium sect. indumentum simple or dendritic. Leaves entire or
Pimelodendron), but in the light of the molecular serrulate; stipules scarious, usually obscure. Inflor-
findings (Wurdack et al. 2005), this similarity escences spiciform, mostly opposite the leaves,
appears homoplasious. sometimes terminal or axillary; pistillate flowers
proximal, solitary at 1 (2) basal nodes; staminate
272. Microstachys A. Juss. Fig. 43 flowers distichous (spiral), 1–3 per node; stami-
nate bracts biglandular (eglandular) at base. Sta-
Microstachys A. Juss., Euphorb. Tent.: 48 (1824); Griseb., minate flowers sessile or subsessile; sepals 3,
Fl. Brit. W. Ind. 1: 49 (1859); Esser, Blumea 44: 173–179
(1999), Gen. Euphorb.: 365 (2001). distinct or basally connate; stamens 3, filaments
Cnemidostachys Mart. & Zucc. (1824). distinct; anthers basifixed, extrorse, dehiscing lon-
Sebastiania sect. Microstachys (A. Juss.) M€
ull. Arg. (1866). gitudinally; pistillode 0. Pistillate flowers sessile or
Euphorbiaceae 197
subsessile; sepals 3, distinct, eglandular or basally submarginal glands; stipules deciduous. Inflores-
minutely glandular; ovary 3-locular, usually with 6 cences mostly terminal, unbranched, bisexual, 1
vertical rows of appendages; ovules anatropous, or 2 proximal nodes with pistillate flowers, dis-
inner and outer integuments thin; stylodia distally with cymules of 5–9 staminate flowers;
tinct, unlobed. Fruits capsular; columella persis- bracts rounded to acuminate, biglandular at
tent, alate. Seeds elliptic to cylindrical, often base. Staminate flowers pedicellate; sepals 3,
truncate, caruncle discoid, testa smooth. basally connate; stamens 2–3, filaments distinct;
Approximately 15 spp., mainly neotropical, anthers dehiscing longitudinally; pollen grains 3-
3–4 disjunct in Africa and 1 in southwest Asia colporate, exine tectate-reticulate; pistillode 0.
and Australia. Microstachys has previously been Pistillate flowers pedicellate; sepals 3, distinct,
classified as part of Sebastiania (Webster 1994), often with basal glands; ovary 3(4)-locular, each
but Esser (1994) insisted on generic status for locule with 2 appendages; stylodia basally con-
Microstachys. In the molecular analysis it is nate into a short common style with 3 short
resolved as sister to Ditrysinia. undivided style branches. Fruits capsular, some-
times with fleshy exocarp, endocarp woody; colu-
273. Ditrysinia Raf. mella persistent, alate. Seeds ellipsoid to
spheroidal, mostly ecarunculate or with small
Ditrysinia Raf., Neogenyton: 2 (1825); Esser in Radcl.-
Sm., Gen. Euph.: 385 (2001).
apical caruncle.
Gymnanthes sect. Ditrysina M€ ull. Arg. (1865). Six spp., Africa and Madagascar.
Sebastiana sect. Stillingiopsis (M€ ull. Arg.) G.L. Webster
(1967). 275. Mabea Aubl. Fig. 44
Monoecious shrubs; indumentum simple or 0. Mabea Aubl., Hist. Pl. Guiane 2: 867, t. 334 (1775); M€ ull.
Arg. in DC., Prodr. 15(2): 1148 (1866), Fl. Bras. 11(2): 515,
Leaves entire; stipules undivided. Inflorescences t. 73, 74 (1874); Pax & K. Hoffm., Pflanzenr. 147, V: 26,
terminal, solitary; bracts with paired elongate kid- figs. 4–6 (1912); Jablonski, Mem. N. Y. Bot. Gard. 17: 164
ney-shaped glands, 2–6 proximal ones with 1 (1967); Huft, Phytologia 62: 339 (1987), Ann. Missouri
female flower, numerous distal ones with 1 male Bot. Gard. 75: 1125 (1989); Esser, Syst. Hippom.:
flower; prophylls 0. Staminate flowers shortly pedi- 118–221, figs. 7–10 (1994).
cellate; calyx 3-lobed, lobes partly connate; stamens Monoecious trees or shrubs; latex whitish; indu-
3, filaments short, distinct, anthers basifixed, mentum mostly dendritic. Leaves eglandular
extrorse, longitudinally dehiscent; pistillode 0. Pis- [except marginally], entire or denticulate; petiole
tillate flowers shortly to moderately pedicellate; eglandular; stipules subpersistent or deciduous.
calyx 3-lobed, lobes nearly distinct; ovary glabrous, Inflorescences terminal or axillary, mostly bisex-
3-locular; stylodia connate into a short common ual racemoid thyrses, sometimes distinctly
style with 3 undivided branches. Fruits spheroidal, branched; bracts mostly biglandular, subtending
capsular, columella triquetrous, alate, persistent. pedunculate glomerules of mostly 3 staminate
Seeds elliptic, smooth, with small caruncle. flowers or solitary pistillate flowers. Staminate
A single sp., D. fruticosa (Bartram) Govaerts flowers long-pedicellate; sepals 5(6), partly con-
& Frodin (¼ D. ligustrina (Michx.) Raf.), eastern nate; stamens (3–)10–80, inserted on the
USA. Formerly subsumed under Gymnanthes and convex receptacle; filaments 0 to as long as the
Sebastiana, this genus is resolved as sister to extrorsely dehiscent anthers; pollen grains 2-
Microstachys (Wurdack et al. 2005). nucleate; pistillode 0. Pistillate flowers distinctly
pedicellate; sepals 5(6); ovary 3-locular, finely
274. Sclerocroton Hochst. and densely tomentose; inner integuments
Sclerocroton Hochst. in C. Krauss, Flora 28: 85 (1845); moderately thick, outer integuments thick, non-
Kruijt & Roebers, Bibl. Bot. 146: 16–27 (1996); Esser in vascularized; stylodia connate into a short to long
Radcl.-Sm., Gen. Euphorb.: 378 (2001). common style, the distal tips simple, recurved.
Excoecaria sect. Sclerocroton M€
ull. Arg. (1866). Fruits capsular; columella alate, persistent. Seeds
Sapium subg. Sclerocroton Pax in Pflanzenr. IV, 147, V: carunculate.
213–249 (1912).
About 40 spp., restricted to the (sub)humid
Monoecious shrubs or trees; latex not evident; neotropical lowlands from Mexico to São Paulo,
indumentum 0. Leaves abaxially with basal or Brazil. Esser (1994) recognized 2 sections: sect.
198 G.L. Webster
? Adenogyne Klotzsch (1841).

Dendrocousinsia Millsp. (1913).
Monoecious (dioecious) trees, shrubs, or herbs;

latex whitish, often not evident; indumentum
simple or 0. Leaves eglandular except sometimes
for marginal glands; margins entire to serrulate
or dentate; stipules minute and often dissected.
Inflorescences terminal or leaf-opposed, solitary,
bisexual; bracts biglandular at base, glands usu-
ally flat and elongated or sometimes divided or
rugulose; pistillate flowers solitary at proximal
nodes, often separated from the distal staminal
cymules of mostly 1–3 flowers. Staminate flowers
sessile or subsessile; sepals 3, slightly connate at
base, sometimes unequal; stamens 3, distinct,
filaments longer than anthers; anthers basifixed,
extrorse, dehiscing longitudinally; pollen grains
3-nucleate. Pistillate flowers sessile to short-
pedicellate; sepals 3, discrete, denticulate,
persistent in fruit; ovary 3-locular, sometimes
appendiculate; ovules anatropous, inner and
outer integuments thin; stylodia distinct or nearly
so, undivided. Fruits capsular; columella persis-
tent, alate, often with adherent caruncles. Seed
smooth, carunculate, caruncles stipitate and
deciduous.
About 25 spp., distributed from Mexico south
into tropical South America. The circumscription
of this genus has been one of the most conten-
Fig. 44. Euphorbiaceae-Euphorbioideae. Mabea speciosa. tious issues in the taxonomic history of
A Inflorescence and part of stem with leaves (right); tribe Hippomaneae. Pax and Hoffmann (1912)
note that inflorescence is pendent on plant. B Proximal followed M€ uller (1866) in adopting a very
portion of inflorescence with pistillate flowers at base
and staminate ones above; note glands on bracts sub- broad and patently unnatural circumscription of
tending staminate flowers. C Cluster of staminate flowers Sebastiania. Esser (1994) has provided a radical
subtended by biglandular bract. D Medial section of solution to the problems of circumscribing Sebas-
staminate flowers (right) and two views of anthers (left).
E Pistillate flower. F Immature capsules. (Reproduced
tiania by transferring sections Sarothrostachys
with kind permission of the artist Bobbi Angell) and Adenogyne from Sebastiania to Gymnanthes.
However, the species of Adenogyne assigned to
Spiculigerae Pax & K. Hoffm., with 2 spp. (one of Sebastiania by M€ uller (1866) have terminal
them the widespread Brazilian M. fistulifera inflorescences and pedicellate pistillate flowers,
Mart.); and sect. Mabea, which includes the and the species of Sarothrostachys have axillary
other sections of Pax and Hoffmann (1912). inflorescences and sessile or subsessile pistillate
flowers. Both species groups have character com-
276. Sebastiania Sprengel binations that conflict with the characters of
Gymnanthes and Sebastiania, and it is obvious
Sebastiania Spreng., Neue Entd. Pflanzenk. 2: 118, t. 3
(1820); M€ ull. Arg. in DC., Prodr. 15(2): 1164 (1866), Fl.
that the current generic boundaries are unsatis-
Bras. 11(2): 544 (1866); Pax & K. Hoffm., Pflanzenr. 147, factory. Sarothrostachys has axillary branched
V: 88 (1914); figs. 19–22, 26–28 (1912), Nat€url. Pflanzen- inflorescences, and its transfer to Anomostachys
fam. ed. 2, 19c: 192, fig. 103 (1931); L.B. Sm. et al., Fl. should be considered. Dendrocousinsia is treated
Illustr. Catar. EUFO 295, fig. 40 (1988); Webster, Ann. by Esser as a highly distinct genus, but in fact the
Missosuri Bot. Gard. 81: 121 (1994); Burger & Huft, Fieldi-
ana n.s. 36: 155 (1995); Esser, Syst. Hippom.: 55 (1994), in
diagnostic character—adaxial ciliae or glands in
Radcl.-Sm., Gen. Euphorb.: 363 (2001). the staminate and pistillate calyx—was described
Euphorbiaceae 199
by M€ uller for a considerable number of species of ers pedicellate; sepals 2, connate; ovary 2-locular;
Sebastiania, including the type (S. brasiliensis stylodia 2, basally connate, unlobed. Fruits dru-
Spreng.). The molecular data of Wurdack et al. paceous. Seeds oblate-spheroidal, ecarunculate,
(2005) fail to indicate the position of this genus with thin sarcotesta. 2n ¼ 44.
within the Hippomaneae, because the signals of Two spp., India to China and New Guinea.
the two species included in the analysis repre- Balakata appears to be closely related to Anom-
senting the core of Sebastiana (S. klotzschiana ostachys, and the two genera should perhaps be
M€ull. Arg. and S. pavoniana M€ ull. Arg.) point to united.
different subclades of the Hippomaneae.
279. Hura L. Fig. 45
277. Anomostachys (Baill) Hurus. Hura L., Sp. Pl.: 1008 (1753); M€ ull. Arg. in DC., Prodr. 15
(2): 1228 (1866), Fl. Brasil. 11(2): 632, t. 86 (1874); Pax &
Anomostachys (Baill.) Hurus., J. Fac. Sci. Univ. Tokyo, III. K. Hoffm., Pflanzenr. 147, V: 271 (1912); Standl. & Steyer-
Bot. 6: 311 (1954); Kruijt, Biblioth. Bot. 146: 8–12, fig. mark, Fieldiana Bot. 24(6): 124 (1949); Burch, Ann. Mis-
2 (1996); Esser in Radcl.-Sm., Gen. Euphorb.: 385 (2001). souri Bot. Gard. 54: 330, fig. 24 (1968); Burger & Huft,
? Sarothrostachys Klotzsch (1841). Fieldiana n.s. 36: 126, fig. 32 (1995); Esser in Radcl.-Sm.,
Duvigneauia Léonard (1959); Kruijt & Roebers, Biblioth. Gen. Euphorb.: 397 (2001).
Bot. 146: 12–15 (1996), rev.
Monoecious trees, trunk spiny; latex white; indu-
Monoecious trees and shrubs; latex whitish; indu-
mentum simple. Leaves deciduous, dentate, with
mentum 0. Leaves below with marginal glands,
paired sessile glands at the junction with petiole;
otherwise eglandular; stipules caducous. Inflores-
stipules caducous. Inflorescences unisexual, the
cences axillary, solitary, ramified, branches race-
staminate terminal, pedunculate, compound-spi-
moid; bracts eglandular, the proximal 2–5
cate, bracts numerous, united into a membranac-
subtending single pistillate flowers and the distal
eous, tunica-like sheath; pistillate flowers solitary,
ones subtending cymules of 2–8 staminate flow-
at base of staminate spike or axillary. Staminate
ers. Staminate flowers sessile; sepals 3, basally
flowers pedicellate; sepals connate into a crenulate
connate; stamens 3; anthers dehiscing longitudi-
cup; stamens 10–80, in 2 or more whorls of 12–15
nally; pistillode 0. Pistillate flowers pedicellate;
sessile anthers on a stout column formed by the
sepals 3, distinct, often bifid; ovary 2- or 3-locu-
stamen filaments; pistillode 0. Pistillate flowers
lar, appendiculate when young but smooth at
pedicellate; sepals 5, completely connate into a
maturation; ovules anatropous, inner integu-
truncate calyx; ovary 5–20-locular; stylodia connate
ments thick [10–20 cell layers], outer integu-
into a long columnar style with a distal 5–20-lobed
ments thin [3 or 4 cell layers], non-vascularized;
discoid stigma. Fruits capsular, dehiscing explo-
a common style short but evident, with 3 undi-
sively; columella not persistent. Seeds laterally
vided distal branches. Fruits indehiscent. Seeds
compressed, suborbicular, ecarunculate. 2n ¼ 44.
carunculate or not.
Two neotropical spp., common in lowland
Three (or more?) spp., tropical Africa, Mada-
deciduous forests. Hura polyandra Baill. of Mex-
gascar.
ico and Central America is replaced in Nicaragua
by H. crepitans L., which is widely distributed
278. Balakata Esser
from Nicaragua to the West Indies, Peru, and
Balakata Esser, Blumea 44: 154 (1999), in Radcl.-Sm., Brazil. Hura crepitans is cultivated throughout
Gen. Euphorb.: 378 (2001). most of the Neotropics, and has been introduced
Sapium sect. Pleurostachya Pax & K. Hoffm. (1912).
into paleotropical regions with monsoonal cli-
Monoecious trees or shrubs; latex white; indu- mates. The unusual floral structure, at first glance
mentum 0. Leaves abaxially with marginal or very different from that in Algernonia and
submarginal glands; stipules entire, eglandular. Ophthalmoblapton, evidently reflects adaptation
Inflorescences terminal and axillary, compound, for bat pollination (Steiner 1982).
bisexual; bracts biglandular at base, at proximal
nodes subtending solitary pistillate flowers, at 280. Algernonia Baill.
distal nodes cymules with 5–9 staminate flowers. Algernonia Baill., Ann. Sci. Nat. IV, 9: 198 (1858), Étude
Staminate flowers pedicellate; sepals 2, connate; Gén. Euphorb.: 546 (1858); M€ ull. Arg. in DC., Prodr. 15
stamens 2, filaments distinct; anthers basifixed, (2): 1230 (1866); Fl. Bras. 11(2): 533, t. 87 (1874); Benth.,
extrorse, dehiscing longitudinally. Pistillate flow- Gen. Pl. 3: 339 (1880); Pax & Hoffm., Pflanzenr. 147, V:
200 G.L. Webster
About 10 spp., ranging from E Brazil to Peru.

Baillon (1858) described both Algernonia and
Tetraplandra simultaneously, and Bentham
(1880) selected the former on joining the two
genera, in which he is followed here. Pax and
Hoffmann (1912) distinguished the genera by
the more gamophyllous staminate calyx in Alger-
nonia, but in both floral and vegetative structures
the two taxa appear similar overall. Emmerich
(1981) provided detailed descriptions and excel-
lent illustrations of both genera, and the entire
reliance on vegetative characters in her generic
key is revealing.
281. Ophthalmoblapton Allem.

Ophthalmoblapton Allem., Pl. Novas Brasil: 4 (1849), Ann.
Sci. Nat. Bot. III, 13: 119 (1849); M€ull. Arg. in DC., Prodr.
15(2): 1155 (1866), Fl. Bras. 11(2): 531 (1874); Benth., Gen.
Pl. 3: 333 (1880); Pax & K. Hoffm., Pflanzenr. 147, V: 278,
fig. 58 (1912); Emmerich, Bol. Mus. Nac. Rio de Janeiro,
Bot. 62: 1, t. 1 (1981); L.B. Sm. et al., Fl. Illustr. Catar.
EUFO 325, t. 41 figs. h–m (1988); Howard, Fl. Lesser Ant.
5: 57 (1989).
Monoecious trees; latex white, highly toxic; indu-
Fig. 45. Euphorbiaceae-Euphorbioideae. Hura crepitans.
A Stem with leaves and staminate inflorescences (left) and mentum 0. Leaves entire or dentate, adaxially with
pistillate flower (right). B Pistillate flower. C Androecium basal glands at junction with petiole; stipules cadu-
with staminal column bearing two rows of anthers. D cous. Inflorescences axillary, bisexual or stami-
Ovary in transverse section (left) and medial section of nate, spiciform; pistillate flowers basal and
pistillate flower (right). E Stem with capsule. F Segment of
capsule and seed. G Prickles on trunk. (Reproduced with solitary, separated from crowded staminate
kind permission of the artist Bobbi Angell) flowers; bracts eglandular, staminate bracts sub-
tending 3–12 flowers. Staminate flowers sessile,
276, fig. 56 (1912); Emmerich, Arq. Mus. Nac. Rio Janeiro
sometimes laterally confluent; calyx closed in
56: 91–110, t. 1–11 (1981). bud, splitting into 2 or 3 lobes or segments; stamen
Tetraplandra Baill. (1858); Emmerich, l.c. solitary, exserted from calyx; anther dehiscing lon-
gitudinally. Pistillate flowers sessile or subsessile,
Monoecious trees or shrubs; latex white; indumen- becoming somewhat pedunculate in fruit; sepals 5
tum 0. Leaves entire or obscurely crenulate and or 6, discrete, imbricate, persistent in fruit; ovary
occasionally with small abaxial glands, biglandular 3-locular; stylodia basally connate into a columnar
at base; stipules caducous. Inflorescences terminal, style with distal stigmatic style branches. Fruits
the staminate ones occasionally axillary, bisexual, capsular; endocarp woody; columella deciduous.
spiciform; bracts adnate to rachis, eglandular; pis- Seeds ovoid, ecarunculate, testa smooth.
tillate flowers solitary at distal nodes, staminate Four spp. of the mata atlântica region in east-
cymules with mostly 3 flowers. Staminate flowers ern Brazil (Bahia to Santa Catarina).
subsessile; sepals 3–5, distinct or connate; stamens
1–3, filaments mostly connate; anthers basifixed, 282. Pachystroma M€
ull. Arg. Fig. 46
extrorse, dehiscing longitudinally. Pistillate flowers
sessile or subsessile; sepals distinct or connate, Pachystroma M€ ull. Arg., Linnaea 34: 177 (1865), in DC.,
Prodr. 15(2): 893 (1866), Fl. Bras. 11(2): 387, t. 54 (1874);
persistent in fruit; ovary 3-locular, exappendicu- Pax, Pflanzenr. 147, II: 99 (1910); Senna, Bradea 3 (48):
late; stylodia connate c. 1/3 to 2/3 of length into a 421 (1983).
columnar common style, tips terete, papillose.
Fruits capsular; trigonous base of columella persis- Monoecious trees or shrubs; latex white; indu-
tent. Seeds subspheroidal, smooth, ecarunculate. mentum 0. Leaves eglandular, usually spinose-
Euphorbiaceae 201
although noting its resemblance in habit to Hip-

pomaneae. Pax and Hoffmann (1931) created a
tribe for it, but still placed it between the Acaly-
pheae and Dalechampieae. In the molecular anal-
ysis of Wurdack et al. (2005), Pachystroma is
embedded within the Hureae, which are part of
one of the hippomanoid subclades, as foreseen by
Webster (1994a).
283. Hippomane L.
Hippomane L., Sp. Pl.: 1191 (1753); M€ ull. Arg. in DC.,
Prodr. 15(2): 1199 (1866); Pax & K. Hoffm., Pflanzenr.
147, V: 261, fig. 51 (1912); Fawc. & Rend., Fl. Jam. 4: 327,
fig. 110 (1920); Webster, J. Arnold Arbor. 48: 393 (1967);
Webster, Ann. Missouri Bot. Gard. 81: 124 (1994).
Mancanilla Miller (1754).
Monoecious trees or shrubs; latex white, toxic;
indumentum 0. Leaves cordate at base, entire
or obscurely crenulate to spinose-dentate, with a
single adaxial gland at junction with petiole;
stipules lanceolate, caducous. Inflorescences ter-
minal, bisexual, spiciform, bisexual; bracts
biglandular at base, subtending 1 or 2 proximal
solitary pistillate flowers and distal cymules of
8–15 staminate flowers. Staminate flowers subses-
sile; sepals 2 or 3, connate; stamens 2, filaments
Fig. 46. Euphorbiaceae-Euphorbioideae. Pachystroma connate into a column; anthers basifixed, extrorse,
longifolium. A Flowering branchlet. B Staminate flower.
C Same, longitudinally sectioned. D Transversal section of dehiscing longitudinally. Pistillate flowers sessile
androecium. E Pistillate flower. F Ovary medially sec- or subsessile; sepals 3, distinct, persistent in fruit;
tioned. G Fruit locule. H Columella after seed dehiscence. ovary 5–10-locular; stylodia basally connate into a
(Pax 1910)
short style, branches unlobed, densely papillate
adaxially. Fruits drupaceous with bony endocarp.
dentate; petioles eglandular; stipules caducous. Seeds smooth, ecarunculate. 2n ¼ 22.
Inflorescences terminal, spiciform, bisexual; Three Caribbean spp., two of which are
bracts biglandular, basally up to 2 with 1 pistillate endemic to Hispaniola, while H. mancinella L.,
flower, apically numerous ones with cymules of the noted highly toxic "manchineel", is widely
1–3 staminate flowers. Staminate flowers sessile; distributed along littoral zones from Florida to
calyx cupular, 2-lobed; stamens 3, filaments con- northern South America and the Galapagos. In
nate into a column; anthers basifixed, extrorse, the molecular analysis of Wurdack et al. (2005),
dehiscing vertically, exserted from calyx. Pistil- Hippomane is resolved in a clade with Plerade-
late flowers subsessile; sepals 3, distinct, imbri- nophora, Bonania, and Grimmeodendron.
cate, persistent in fruit; ovary 3-locular; stylodia
connate into a columnar style, style branchlets 284. Pleradenophora Esser
unlobed, compressed. Fruits capsular, valves
Pleradenophora Esser in Radcl.-Sm., Gen. Euph.: 377
woody and thick-walled; trigonous base of colu- (2001).
mella persistent. Seeds ovoid, somewhat com-
pressed, cordate at base, ecarunculate. Monoecious shrubs or trees; indumentum 0; latex
A single sp., P. longifolium (Nees) I.M. John- white. Leaves serrate, below sometimes with mar-
ston of SE Brazil, Bolivia and Peru. M€ uller (1866) ginal glands, often with a pair of disk-shaped
misplaced the genus among the Acalyphoideae, petiolar glands on the junction of the petiole
and Pax (1910) placed it in the Adrianeae, with the blade; stipules small. Inflorescences
202 G.L. Webster
terminal or laterally displaced, solitary; floral Monoecious trees or shrubs; latex white; indu-
bracts with several pairs of disk- or cup-shaped mentum 0. Leaves serrulate, adaxially biglandular
glands, 1–3 basal ones with 1 pistillate flower, at base at junction with petiole; stipules deltoid or
numerous apical ones with 5–10 staminate flow- lunate, the fimbriate scarious tip deciduous, the
ers. Staminate flowers shortly pedicellate; calyx 3- lunate base persistent. Inflorescences terminal,
lobed, partly connate; stamens 2–3(5), filaments bisexual; bracts eglandular, subtending 1 or 2 soli-
distinct, anthers basifixed, extrorse, longitudi- tary proximal pistillate flowers and several distal
nally dehiscent; pistillode 0. Pistillate flowers staminate cymules with 2 or 3 flowers. Staminate
(sub)sessile; calyx 3-lobed, lobes distinct; ovary flowers subsessile; sepals 3, valvate, distinct or
smooth, 3-locular; stylodia undivided, basally basally connate, entire; stamens 3, filaments con-
connate or not into a short common style. Fruit nate into a column exserted from the calyx;
subglobose, smooth, dehiscing septicidally into 3 anthers extrorse, dehiscing longitudinally. Pistil-
mericarps, columella 3-quetrous, alate, persis- late flowers sessile; sepals 3, deltoid, entire, persis-
tent. Seeds ovoid-globose, smooth, often macu- tent in fruit; ovary 3-locular; stylodia united at
late, ecarunculate. base into a short style, tips unlobed, recurving.
Three spp., Mexico, Guatemala, Belize; for- Fruits capsular; columella alate distally, persistent.
merly treated as part of Sebastiana (S. longicuspis Seeds ovoid-subspheroidal, smooth, ecarunculate.
Standl.). Two spp., Greater Antilles except Puerto Rico
and Bahamas.
285. Bonania A. Rich.
Bonania A. Rich., Hist. Fis. Cuba 2, 11: 201, t. 68 (1853);
287. Adenopeltis Bert. ex A. Juss.
Benth., Gen. Pl. 3: 335 (1880); Pax & K. Hoffm., Pflanzenr. Adenopeltis Bert. ex A. Juss., Ann. Sci. Nat. I, 25: 24
147, V: 259 (1912); Borhidi, Acta Bot. Acad. Sci. Hungar. (1832); M€
ull. Arg. in DC., Prodr. 15(2): 1164 (1866); Pax
22: 305 (1976); Webster, Ann. Missouri Bot. Gard. 81: 123 & K. Hoffm., Pflanzenr. 147, V: 264, fig. 52 (1912); Esser in
(1994); Esser in Radcl.-Sm., Gen. Euphorb.: 369 (2001). Radcl.-Sm., Gen. Euphorb.: 369 (2001).
Monoecious shrubs; latex not recorded; indu- Monoecious shrubs; latex not recorded; scabrid
mentum 0. Leaves distichous, subsessile, egland- emergences on stems and inflorescence axes.
ular, margins glandular-crenate; stipules Leaves short-petiolate or subsessile, eglandular,
persistent. Inflorescences terminal on axillary glabrous, margins prominently glandular-denti-
branches, unbranched, bisexual, pistillate flower culate; stipules small, divided into several ciliae,
solitary at base, staminate flowers in numerous eglandular. Inflorescences terminal and axillary,
distal 1–3-flowered cymules; bracts eglandular. solitary, bisexual, spiciform; bracts ovate, entire,
Staminate flowers subsessile; calyx nearly with a pair of stipitate, disk-shaped glands, the
completely connate, without distinct lobes; 0–2 basal ones subtending solitary pistillate flow-
stamens 2 or 3, filaments about as long as anthers, ers, and numerous apical ones with 1–3 staminate
distinct or basally connate; anthers basifixed, flowers. Staminate flowers subsessile; calyx 0;
extrorse, dehiscing longitudinally. Pistillate flow- stamens 2 or 3, connate at base; anthers basifixed,
ers sessile, calyx urceolate, 2–3-lobed or toothed; dehiscing longitudinally; pistillode 0. Pistillate
ovary 3-locular, smooth; ovules anatropous, flowers pedicellate; sepals 0; ovary 3-locular; sty-
inner integuments moderately thick [8–10 cell lodia distinct or basally connate, slender. Fruits
layers], outer integuments thin [3–5 cell layers]; capsular, 3-lobed; columella not persistent, base
stylodia incrassate, recurved, papillate adaxially. of cocci persistent as three lobes. Seeds subsphe-
Fruits capsular; columella persistent, alate. Seeds roidal, ecarunculate, testa smooth.
spheroidal, ecarunculate, testa smooth. A single sp., A. serrata (Ait.) G.L. Webster,
Seven spp., endemic to the Greater Antilles Chile and Peru.
and Bahamas but not in Jamaica.
288. Spegazziniophytum Esser
286. Grimmeodendron Urb.
Spegazziniophytum Esser in Radcl.-Sm., Gen. Euporb.:
Grimmeodendron Urb., Symb. Ant. 5: 397 (1908); Pax & K. 371 (2001).
Hoffm., Pflanzenr. 147, V: 258, fig. 50 (1912); Borhidi,
Acta Bot. Acad. Sci. Hungar. 22: 305 (1976); Esser in Monoecious, succulent shrubs, branches trans-
Radcl.-Sm., Gen. Euphorb.: 368 (2001). formed into thorns. Leaves present only on
Euphorbiaceae 203
young branches, subsessile. Inflorescences axil- World species ascribed to Stillingia would merit
lary, solitary, catkin-like; bracts with a pair of critical revision. Also the inclusion of Adenopeltis
cup- to disk-shaped glands; pistillate flowers 1 and Spegazziniophytum in Stillingia should seri-
per bract, staminate flowers 2 or 3 per bract. ously be considered; it is not formally proposed
Staminate flowers sessile; calyx 2-lobed, connate; here only because not all new combinations are
stamens 2, filaments distinct; pistillode 0. Pistil- available.
late flowers subsessile; calyx 3-lobed, lobes nearly
distinct; ovary smooth; stylodia entire. Fruit 2- 290. Falconeria Royle
locular, dehiscent; columella alate but lacking a Falconeria Royle, Ill. Bot. Himal.: 354 (1839); Wight, Ic.
carpidiophore. Seeds 2 per fruit, subglobose, with Pl. Ind. Or. 5(2): 20, t. 1866 (1853); Esser, Blumea 44:
minute apical caruncle. 160–165 (1999), rev.
A single sp., S. patagonicum (Speg.) Esser, Sapium sect. Falconeria (Royle) Hook. f. (1888); Pax & K.
Argentinian Patagonia. Formerly included in Col- Hoffm., Pflanzenr. IV: 147, V: 241, fig. 45 (1912).
liguaja, but in the molecular analysis (Wurdack Monoecious trees; latex white; indumentum 0.
et al. 2005) resolved in a clade with Stillingia and Leaves deciduous, sometimes apically clustered,
Adenopeltis, from which it differs by the lack of a serrate, abaxially with paired basal glands at junc-
carpidiophore. tion with petiole; stipules dissected. Inflorescences
terminal, unisexual, spiciform; bracts biglandular
289. Stillingia Garden ex L. at base with discoid glands, subtending solitary
Stillingia Garden ex L., Syst. Nat. ed. 12, 2: 637 (1767); pistillate flowers or cymules of 10–15 staminate
M€ull. Arg. in DC., Prodr. 15(2): 1155 (1866); Fl. Bras. 11 flowers. Staminate flowers subsessile; sepals 2,
(2): 537 (1874); Pax & K. Hoffm., Pflanzenr. 147, V: 180 connate; stamens 2, filaments distinct; anthers
(1912); Rogers, Ann. Missouri Bot. Gard. 38: 207, figs. 1–3, basifixed, extrorse, dehiscing longitudinally. Pistil-
6–16 (1951); van Steenis, Blumea Suppl. 5: 302 (1966);
Webster, J. Arnold Arb. 48: 388, fig. 5 (1967); A.C. Sm., late flowers subsessile [short-pedicellate in fruit];
Fl. Vit. Nov. 2: 565, fig. 161 (1981); Esser in Radcl.-Sm., sepals 2 or 3, connate below, eglandular; ovary 2-
Gen. Euphorb.: 370 (2001). or 3-locular, smooth; stylodia distinct or basally
Gymnostillingia M€ ull. Arg. (1863). connate, unlobed. Fruits spheroidal, dehiscing
Monoecious arborescent shrubs, subshrubs, or irregularly, pericarp fleshy becoming papery; col-
herbs, sometimes with succulent stems; latex umella 2- or 3-angled, alate, caducous. Seeds sphe-
white; indumentum 0. Leaves alternate or oppo- roidal, ecarunculate, testa pale, arillate. 2n ¼ 32.
site, entire to glandular-serrulate, with paired A single sp., F. insignis Royle, E Asia from
basal glands at junction with petiole; stipules India to SW China and peninsular Malaysia. Fal-
laciniate-dissected. Inflorescences terminal, coneria is distinctive due to its unisexual spikes
unbranched, spiciform, bisexual; bracts bigland- and irregularly dehiscing fruits enclosing seeds
ular at base; with distal cymules of 1–3 staminate with whitish aril. It seems to be closer morpho-
flowers. Staminate flowers subsessile; sepals 2, logically to Sapium (s. str.) than to other Asiatic
connate; stamens 2 (3), filaments distinct, Hippomaneae.
exserted from calyx; anthers basifixed, extrorse,
dehiscing longitudinally; pollen grains 2-nucle- 291. Sapium Jacq.
ate. Pistillate flowers subsessile; sepals 3, some- Sapium Jacq., Enum. Syst.: 9 (1760), nom. cons.; Pax & K.
times reduced or 0; ovary 2- or 3-locular; Hoffm., Pflanzenr. 147, V: 199 (1912); Webster, J. Arnold
ovules anatropous, inner integuments moderately Arb. 48: 391 (1967), pro parte; Kruijt & Zijlstra, Taxon 38:
320 (1989); Kruijt, Bibl. Bot. 146: 27–91, figs. 10–31, 33–35
thick, outer integuments thin, vascularized; sty- (1996), rev.; Esser in Radcl.-Sm., Gen. Euphorb.: 373 (2001).
lodia connate basally, unlobed. Fruits capsular;
columella alate, usually persistent, with 3 horn- Monoecious or dioecious trees or shrubs; latex
like projections from base; seeds subspheroidal, white, often viscous; indumentum 0. Leaves
carunculate (caruncle reduced). 2n ¼ 22, 30, 36. entire to serrulate; petioles usually biglandular;
About 30 spp., most of them in the New stipules ovate, persistent. Inflorescences terminal
World (North, Central and South America), a and axillary, solitary or clustered, spiciform;
few spp. with scattered distribution in the Old bracts biglandular, discoid to oblong, subtending
World (2 in Madagascar, 1 in Malesia, and 1 or solitary proximal pistillate flowers and distal
2 on Pacific islands extending to Fiji). The Old cymules of (2–)5–10(15) staminate flowers.
204 G.L. Webster
Staminate flowers mostly subsessile; sepals 2, 268 (1971), Add. Ser. 4: 112 (1975), 8: 88 (1980); Radcl.-
connate; stamens 2, distinct or basally connate; Sm., Fl. E. Trop. Afr. Euphorb. 1: 382, fig. 72 (1987);
Chakrab. & Gangop., J. Econ. Tax. Bot. 18: 193 (1994);
anthers basifixed, extrorse, dehiscing longitudi- Radcl.-Sm., Fl. Zamb. 9(4): 315, t. 68 (1996); Esser in
nally. Pistillate flowers sessile or subsessile; sepals Radcl.-Sm., Gen. Euphorb.: 360 (2001).
2 or 3, basally connate; ovary (1)2–3-locular, Commia Loureiro (1790).
unappendaged; ovules anatropous, inner and Spirostachys Sond. (1850).
outer integuments moderately thick; stylodia dis- Conosapium M€ ull. Arg. (1863).
Taeniosapium M€ ull. Arg. (1866).
tinct to basally connate into a common style, Glyphostylus Gagnep. (1925).
distally unlobed. Fruits capsular; columella per-
sistent. Seeds spheroidal, ecarunculate, testa cov- Dioecious or monoecious trees or shrubs; latex
ered with red aril. 2n ¼ 22, 44. milky, toxic; indumentum 0. Leaves alternate or
About 25 spp. from the Neotropics. The clas- opposite, entire or crenulate, eglandular, rarely
sical circumscription of the genus of Pax and with basal glands; stipules deciduous or
Hoffmann (1912), which has been followed in persistent. Inflorescences axillary or terminal on
most floristic works until recently, was much brachyblasts, solitary, spiciform; bracts usually
more inclusive, but Kruijt (1996) has excluded biglandular at base, subtending 1–3 staminate
all taxa lacking an aril, so that Sapium appears flowers or single pistillate flowers. Staminate flow-
restricted to the New World. ers sessile or subsessile; sepals 3(2), distinct;
stamens 3(2), filaments distinct; anthers extrorse,
292. Colliguaja Molina dehiscing longitudinally; pistillode 0. Pistillate
flowers subsessile or short-pedicellate; sepals 3,
Colliguaja Molina, Saggio Chile: 158 (1781); M€ ull. Arg. in
DC., Prodr.15(2): 1226 (1866), Fl. Bras. 11(2): 630, t. 85 distinct or basally connate; ovary 3-locular; ovules
(1874); Pax & K. Hoffm., Pflanzenr. 147, V: 265, fig. 53 anatropous, inner integuments moderately thick,
(1912); L.B. Smith et al., Fl. Illustr. Catar. EUFO: 326 (1988). outer integuments thin; stylodia distinct or slightly
connate at base. Fruits capsular; columella persis-
Monoecious shrubs, sometimes arborescent; latex
tent, alate. Seeds subglobose, ecarunculate or the
not recorded; indumentum 0. Leaves mostly oppo-
caruncule persistent on top of the columella.
site or subopposite, short-petiolate, glandular-
About 40 spp. distributed from Africa to
denticulate or entire; stipules entire, minute,
Australia and Melanesia. Esser (2001) has re-
deciduous. Inflorescences terminal, bisexual, spi-
defined the genus by excluding some elements
ciform, bracts ovate, eglandular, subtending 1 or
retained by Pax and Hoffmann (1912) and syno-
2 proximal solitary pistillate flowers and several
nymizing Glyphostylus. There is still considerable
cymules of 3 staminate flowers. Staminate flowers
uncertainty about the monophyly of Excoecaria;
subsessile; sepals 1 or 2 and minute or 0; stamens
for example, the study of ovular structure by
2–4 (12–18 in C. brasiliensis), filaments distinct or
Tokuoka and Tobe (2002) indicates that E. bussei
connate; anthers basifixed, extrorse, longitudinally
differs greatly from other species and may
dehiscent. Pistillate flowers subsessile; sepals 3,
not be congeneric. Although Esser maintains Spir-
minute; ovary 2–3(4)-locular; ovules anatropous,
ostachys as a distinct genus, it is included within
inner integuments moderately thick, outer integu-
Excoecaria here (and was resolved as the strongly
ments thin; stylodia basally connate into a short
supported sister to E. agallocha; Wurdack et al.
style, the distinct branches recurved, adaxially
2005), where it will probably survive as a well-
papillate. Fruits capsular, trigonous; columella tri-
marked subgenus or section. Conosapium from
quetrous, alate apically, persistent. Seeds sub-
Madagascar, although generically accepted by
spheroidal, ecarunculate, testa smooth. 2n ¼ 36.
Esser (2001) and Schatz (2001), can probably be
Four spp., three in C Chile and one in S Brazil
accommodated as a distinctive section.
and adjacent Paraguay and Uruguay; C. brasilien-
sis is rheophytic.
3. TRIBE EUPHORBIEAE
293. Excoecaria L. Monoecious (dioecious) trees, shrubs, or herbs,
Excoecaria L., Syst. Nat. ed. 10: 1288 (1759); F.M. Bailey, sometimes succulent; latex milky, sometimes
Queensl. Fl. 5: 1456 (1902); Pax & Hoffm., Pflanzenr. IV, toxic; indumentum simple or 0; leaves alternate,
147, V: 157, figs. 29–31 (1912); Airy Shaw, Kew Bull. 16: opposite, or whorled, mostly unlobed and
Euphorbiaceae 205
pinnately veined, sometimes reduced, without glands commissural; bracts of staminate dichasia
laminar or petiolar glands; stipules + or 0; inflor- large, imbricate, enclosing staminate flowers;
pistillate sepals 5 or 6, large, imbricate. 3b. Neoguil-
escences pseudanthial with connate bracts lauminiinae 4
forming a cyathium that encloses a single terminal – Petaloid appendages interbracteal [opposite cyathial
pistillate flower and 4 or 5 lateral staminate dicha- glands], alternate with staminate monochasia;
sia or monochasia; cyathia often aggregated in cyathial glands on rim of cyathium; bracts of stami-
compound inflorescences; staminate flowers with nate monochasia small or obsolete; pistillate sepals +
calyx reduced or 0, the solitary stamen articulating or 0. 3c. Euphorbiinae (298. Euphorbia s. l.) 5
with the pedicel; pollen grains 3-colporate, colpi 4. Cyathial glands 8 or 10, massive; petaloid involucral
bracts c. 1 cm long; leaves spiral
bordered by conspicuous intine bands, exine per- 296. Neoguillauminia
forate-tectate; pistillate flowers pedicellate, – Cyathial glands 4, small; petaloid bracts < 1 cm long;
calyx 3–6-lobed or reduced or 0; ovary (2)3(4)- leaves opposite 297. Calycopeplus
locular; stylodia distinct or basally connate, bifid, 5. Cyathia bilaterally symmetrical; glands enclosed
rarely entire; fruit capsular, rarely drupaceous; within a nectar spur; stylodia connate into a long
seeds carunculate or ecarunculate; testa dry. column. 17 spp., most in Mexico
Euphorbia ’Pedilanthus’
This tribe includes 3 subtribes grouped into 5
– Cyathia radially symmetrical, glands not enclosed
genera and > 2,000 spp., the vast majority in the within a nectar spur; stylodia distinct or nearly so 6
genus Euphorbia s.l. In many molecular analyses 6. Involucral glands distinct, (1–)4–5(8), alternating
(e.g., Steinmann and Porter 2002; Wurdack et al. with involucral lobes on rim of cyathium; bracts sub-
2005; Horn et al. 2012), Euphorbieae are resolved tending cyathium distinct or connate 7
as monophyletic with the Anthosteminae as the – Involucral glands not 4 or 5 alternating with lobes;
earliest diverging group, followed by the Neoguil- bracts subtending cyathium often connate into a cup
lauminiinae and Euphorbiinae in a sister position. 8
7. Leaves alternate, opposite, or whorled, but if opposite
Genera until recently kept separate from Euphor- then not inaequilateral at base; veins of leaves not
bia s. str. but now found to be deeply embedded chlorenchyma-sheathed; stipules + or 0; main
and sunk into it (see below under Euphorbia s. l.) [embryonic] axis not abortive Euphorbia s.str.
are not formally accepted here but are keyed out – Leaves opposite, stipulate, inaequilateral at base; veins
in order to demonstrate on which characters they chlorenchyma-sheathed; main axis abortive above
had been based; this is also because they have cotyledons. About 250 spp., mostly in trop. America
and Africa Euphorbia ’Chamaesyce’
been in wide use in general and phytogeographic
8. Pistillate flower calyculate; involucral lobes
writings until recently, and may still be so until a connate; glands 2, connate into a shield; seeds ecar-
full infrageneric classification of Euphorbia s. l. unculate; stems not succulent. 3 spp., Cuba, Hispa-
becomes available and accepted. niola Euphorbia ’Cubanthus’
– Pistillate flower naked; involucral lobes and glands
connate into a ring; seeds mostly carunculate; stems
KEY TO THE SUBTRIBES AND GENERA OF EUPHOR- succulent 9
BIEAE 9. Cyathium somewhat bilaterally symmetrical; gland-
ring with a gap on one side. 70 spp., trop. Africa
1. Staminate calyx +; involucre of 4 partially or fully Euphorbia ’Monadenium’
connate bracts; cyathial glands commissural, not on
– Cyathium radially symmetrical, without a gap on one
rim of cyathium. 3a. Anthosteminae 2
side 10
– Staminate calyx 0; involucral bracts usually 5, connate 10. Involucral glands connate, conspicuous; ovary with-
into a cup or tube; cyathial bracts different 3 out angular crests. 19 spp., E and S trop. Africa
2. Inflorescences axillary; cyathia bisexual; involucral Euphorbia ’Synadenium’
bracts partially connate into an open 4-lobed involu- – Involucral glands distinct, inconspicuous; ovary with
cre; cyathial glands at margins of involucral bracts; prominent double angular crests. 1 sp., Angola
ovary 3-locular 294. Anthostema Euphorbia ’Endadenium’
– Inflorescences terminal; cyathia mostly unisexual;
involucral bracts connate into a closed cup; cyathial
glands connate by pairs into 4 lobes alternating with
3a. SUBTRIBE ANTHOSTEMINAE (Klotzsch & Garcke)
the involucral bracts; ovary 4-locular G.L. Webster (1975).
295. Dichostemma
3. Petaloid appendages bracteal [dilated involucral
Monoecious or dioecious; leaves with dispersed
lobes], opposite the staminate dichasia; cyathial and submarginal glands; stipules caducous;
206 G.L. Webster
cyathia in axillary inflorescences; cyathial involucellate; calyx cupular or obscurely 4-lobed; ovary
cre of 4 bracts subtending or enclosing 4 stami- 4-locular; stylodia distinct, entire or obscurely
nate partial inflorescences; staminate flowers in bifid. Fruits capsular; endocarp woody but thin;
dichasia enclosed by prophylls; pistillate flower columella 4-angular, persistent. Seeds trigonous-
solitary, central and terminal or apparently lat- rounded, ecarunculate, testa smooth.
eral; cyathial glands projecting inward from Two spp. from W Africa.
infolded margins of involucral bracts; staminate
and pistillate flowers with gamophyllous calyx; 3b. SUBTRIBE NEOGUILLAUMINIINAE Croizat (1938).
ovary 3–4-locular; stylodia emarginate to bifid;
fruit capsular; seeds carunculate or ecarunculate. Monoecious; indumentum 0; leaves alternate or
Two genera of Africa and Madagascar. opposite, entire; stipules 0; cyathia in axillary
or subterminate pedunculate cymes; involucral
bracts 4(6), basally connate, distally dilated or
294. Anthostema A. Juss.
petaloid; staminate dichasia opposite the involu-
Anthostema A. Juss., Euphorb. Tent.: 56 (1824); Pax & K. cral bracts; nectary glands in 4 or 5 pairs, some-
Hoffm., Nat€ url. Pflanzenfam. ed. 2, 19c: 207, fig. 11 a, b times small, or 0; pistillate flower central.
(1931); Keay, Fl. W. Trop. Afr. ed. 2, 1: 416 (1958); Ber-
haut, Fl. Ill. Sénégal 3: 379 (1975); Schatz, Gen. Tree Fl.
Staminate flowers in bracteate dichasia; stami-
Madag.: 145, fig. 134 (2001). nate calyx 0; pistillate sepals 5 or 6, imbricate;
ovary 3-locular; stylodia unlobed, dilated; seeds
Monoecious trees; latex white; indumentum 0. smooth, carunculate.
Leaves distichous, with minute punctiform This Australasian subtribe of 2 genera (1 in
glands and larger abaxial submarginal glands. Australia and the other in New Caledonia) appears
Inflorescences axillary, of contracted cymes of to bridge the gap between the Anthosteminae and
cyathia; cyathia with involucre of 4 bracts con- Euphorbiinae.
nate laterally; each bract with 2 marginal glands.
Staminate flowers enclosed by 4 prophylls; calyx
296. Neoguillauminia Croizat Fig. 47
3- or 4- dentate; pollen grains oblate-spheroidal,
3-colporate, colpi marginate, exine reticulate; pis- Neoguillauminia Croizat, Phil. J. Sci. 64: 398 (1938), Bull.
tillate flowers pedicellate; calyx 3- or 4-dentate; Jard. Bot. Buit. III, 17: 206 (1941); Guillaumin, Fl. Anal.
Syn. Nouv-Caléd.: 182 (1948); McPherson & Tirel, Fl.
ovary 3-locular, sessile; stylodia for lower half Nouv.-Caléd. 14(1): 22–25, t. 3 (1987).
connate into a column, distally clavate or bifid.
Fruits capsular; endocarp thick and woody; Monoecious trees or shrubs; latex white; indu-
columella trigonous-alate, clavate, persistent. mentum 0, young stems pruinose. Leaves alter-
Seeds laterally compressed, carunculate. 2n ¼ 22. nate or opposite, eglandular. Inflorescences
Three spp., 2 in west Africa and 1 in Mada- axillary and subterminal, long-pedunculate;
gascar. cyathia mostly 3 in a terminal dichasium; involu-
cral bracts 4–6, basally connate into a cup and
distally with 4–6 petaloid appendages reddish
295. Dichostemma Pierre
without and white within; staminate dichasia
Dichostemma Pierre, Bull. Mens. Soc. Linn. Paris 1 (159): opposite the involucral bracts, bracteate, each
1259 (1896); Pax & K. Hoffm., Nat€ url. Pflanzenfam. ed. 2, surrounded by broad, erose, rectangular bracts,
19c: 207 (1931); Keay, Fl. W. Trop. Afr. ed. 2, 1: 416 (1958),
Trees Nigeria 151 (1989); Radcl.-Sm., Gen. Euphorb.: 400, 10–28-flowered; nectary glands 8–12, stipitate,
fig. 46 (2001). fleshy, pair-wise partitioning the staminate par-
tial inflorescences; pistillate flower central. Stami-
Monoecious or dioecious trees, sometimes climb- nate flowers pedicellate; perianth 0; pollen grains
ing; latex white; indumentum pruinose on inflo- 2-nucleate; pistillate flower subsessile; calyx
rescence axes. Leaves eglandular. Inflorescences annular, persistent in fruit; anther introrse with
terminal with the cyathia in a panicle; cyathia respect to the cyathium; ovary 3-locular; stylodia
mostly unisexual, bilateral, with involucre of recurved, basally connate, distally bifid. Fruits
paired connate bracts; glands connate in pairs capsular, endocarp woody; columella apically
alternating with involucral bracts. Staminate dilated, deciduous except for conical base. Seeds
flowers 5–12; calyx 4-lobed; pistillate flower pedi- ellipsoid, carunculate, testa smooth.
Euphorbiaceae 207
paired, subsessile; involucral bracts 4, connate,

their distinct lobes small, petaloid, white or green-
ish; nectary glands 4 between the lobes, sometimes
reduced; staminate partial inflorescences opposite
the lobes in 3–7-flowered bracteate dichasia; pis-
tillate flower solitary, central. Staminate flowers
pedicellate; perianth 0; anther introrse in relation
to pistillate flower. Pistillate flower pedicellate;
calyx cupular, 4–6-lobed; ovary 3-locular, sessile
or subsessile; stylodia distinct or basally connate.
Fruit capsular, 3-lobed, endocarp thin and crusta-
ceous; columella persistent, clavate. Seeds sub-
spheroidal or oblong, carunculate, testa smooth.
Five spp., all endemic to Australia, which
differ by a xeromorphic habit from Neoguillau-
minia but share with it the cyathial structure.
3c. SUBTRIBE EUPHORBIIINAE

Monoecious (dioecious) trees, shrubs, or herbs;
latex milky; indumentum simple or 0; leaves
alternate, opposite, or whorled, usually unlobed
and pinnately veined, stipulate or exstipulate;
inflorescences terminal or axillary, cyathia often
subtended by paired bracts; cyathium distinctly
cupulate with (1–2)4–5 interbracteal glands on
the rim; glands with or without petaloid appen-
dages; pistillate flower central, staminate flowers
in 4 or 5 monochasia [usually cincinni] (rarely
Fig. 47. Euphorbiaceae-Euphorbioideae. Neoguillaumi-
nia cleopatra. A Flowering branchlet. B Young cyathium. 1-florous) with reduced prophylls; staminate
C Cyathium at anthesis. D Nectar gland seen from the side calyx 0; stamen 1; pollen grains 2- or 3-nucleate;
and from below. E Involucral bract. F Bracteate staminate pistillate calyx 3–6-lobed, often vestigial or 0;
flowers. G Fruit. H Carunculate seed. (McPherson & Tirel ovary 3-locular; stylodia distinct or connate,
1987; drawn by D. Storez)
bifid (entire); fruit capsular (drupaceous); seeds
carunculate or ecarunculate.
Only one sp., N. cleopatra (Baill.) Croizat, Only one genus, Euphorbia (s. l.).
endemic to New Caledonia. The showy and fra-
grant cyathia resemble those of species of 298. Euphorbia L.
Euphorbia, but the petaloid appendages may not
Euphorbia L., Sp. Pl.: 450 (1753); Boissier in DC., Prodr.
be homologous. 15(2): 7 (1862), Icon. Euphorb.: t. 27–119 (1866); Carter,
Fl. E. Trop. Afr., Euphorb. 2: 409 (1988); Carter & Leach,
297. Calycopeplus Planch. Fl. Zambesiaca 9(5): 339, t. 71–84 (2001); Radcl.-Sm., Gen.
Euphorb.: 405, figs. 47–50 (2001).
Calycopeplus Planch., Bull. Soc. Bot. France 8: 30 (1861); Tithymalus Gaertn., Fruct. 2: 115 (1790), nom. cons.
Boissier, Ic. Euphorb.: t. 120 (1866); Airy Shaw, Kew Pedilanthus Necker ex Poit., Ann. Mus. Nat. Hist. 19: 388
Bull. 35: 603 (1980); P.I. Forster, Austrobaileya 4: 418, (1812), nom. cons.; Dressler, Contr. Gray Herb. 182: 1–188
t. 1, 2 (1995). (1957).
Euphorbia sect. Calycopeplus (Planch.) Boiss. (1862). Chamaesyce S. F. Gray, Nat. Arrang. Brit. Pl. 2: 260 (1821);
Monoecious shrubs, sometimes arborescent, or Koutnik, S. Afr. J. Bot. 3: 262 (1984); Ya Yang & Berry,
Amer. J. Bot. 98: 1486–1503 (2011).
subshrubs; stems and branches virgate; latex Poinsettia Graham, Edinburgh New Phil. J. 20: 412 (1836).
white. Leaves opposite or verticillate, petiolate to Synadenium Boiss. in DC., Prodr. 15(2): 187 (1862); S.
sessile, eglandular, soon deciduous. Inflorescences Carter, Fl. E. Trop. Afr., Euphorb. 2: 534, fig. 101 (1988),
axillary, rarely terminal, the cyathia solitary or Fl. Zambes. 9(5): 434, t. 85 (2001).
208 G.L. Webster
Monadenium Pax, Bot. Jahrb. 19: 126, fig. (1895); P.R.O. angiosperm genera. It is subcosmopolitan in dis-
Bally, Genus Monadenium: 14, figs. 1–34, t. I–XXXII (1961). tribution and is extremely variable in habit; Hallé
Elaeophorbia Stapf, Hooker’s Icon. Pl. 29: t. 2823 (1906).
Cubanthus (Boiss.) Millsp., Field Mus. Nat. Hist. Bot. 2:
et al. (1978), in their architectural analysis,
371 (1913); Steinmann et al., Ann. Jard. Bot. Madrid 64: reported from it 12 of the 23 recognized struc-
123–133 (2007). tural types ("models"). Horn et al. (2012) ana-
Endadenium Leach, Garcia de Orta 1: 31 (1973). lyzed the major structural traits in a
phylogenetic context, which revealed an extreme
Monoecious (dioecious) trees, shrubs, or herbs, parallelism in the evolution of the xeromorphic
often succulent and/or spiny; stems erect to pro- growth forms within the genus.
cumbent, rarely plants acaulescent; latex whitish; Delimitation of the genus Euphorbia always
indumentum simple or 0. Leaves alternate, oppo- has been strongly contentious, and in the 260
site, or whorled, persistent or deciduous, simple years that elapsed since Linnaeus treated the
(deeply lobed), sessile to petiolate, entire or vari- then-known 56 spp. of Euphorbia, more than 75
ously toothed, mostly pinnately veined, eglandu- satellite genera have been proposed. Most of
lar; stipules small and inconspicuous or 0, often them, however, have fallen into synonymy, and
glandular, sometimes modified into spines. Inflor- until very recently, only six or seven generally
escences bisexual (unisexual), contracted into have been maintained, as by Webster (1994) and
pseudanthial cyathia; cyathia in terminal or axil- Radcliffe-Smith (2001). Molecular phylogenetic
lary dichasia or pleiochasia, sometimes solitary, research by Steinmann and Porter (2002) demon-
each subtended by 2 variously colored distinct or strated that Chamaesyce, Pedilanthus, Monade-
connate modified leaves, composed of a single nium, Synadenium, and Endadenium are all
central terminal pistillate flower and 5 or 4 stami- deeply nested within paraphyletic Euphorbia,
nate cincinnate monochasia with their associated and that most of the historically recognized sub-
prophylls, the monochasia alternating with 5 or 4 genera and sections of Euphorbia s.str. are either
involucral bracts connate into a radially or isobi- paraphyletic or polyphyletic. Further work
laterally symmetrical campanulate to hemispheri- (Steinmann 2003; Bruyns et al. 2006; Steinmann
cal involucre; the tips of the involucral bracts et al. 2007; Zimmermann et al. 2010; Horn et al.
alternating with (0)1–5(20) glands situated on the 2012) confirmed and extended these findings, and
rim of the opening (enclosed in a zygomorphic- led to the broadly construed generic concept of
spurred involucre); glands sometimes forming a Euphorbia that is gaining increasing acceptance.
continuous rim on the outer wall of the involu- Steinmann and Porter (2002) also recovered four
cre but usually distinct and entire, bicornute, cup- major lineages within Euphorbia, designated as
ular, bilabiate, or sometimes highly divided, clades A–D; with one exception, these do not
usually with dorsal appendages, these often flat correspond to any previously known taxon but
and petaloid, sometimes glandular or branched. are composed of various sections or subgenera of
Staminate flowers monandrous; perianth usually Euphorbia; only a single clade, by and large, is
0 or very rarely developed; anthers dehiscing lon- congruent with subg. Esula. Bruyns et al. (2006),
gitudinally and introrsely in relation to the vertical in their study of southern African Euphorbia,
axis of the cyathium; pollen grains 2- or 3-nucle- confirmed the existence of these clades; they
ate, subspheroidal, tricolporate, reticulate to finely used the names of some of the existing subgenera
tectate-perforate; pistillate flower pedicellate, ped- with which the clades showed partial overlap for
icel often becoming reflexed; sepals (0)3–6, designating and formally treating them as the
connate basally; ovary (2)3(4)-locular, glabrous four subgenera of Euphorbia: A ¼ subg. Rhi-
or pubescent, often angled or carinate; stylodia zanthium, B ¼ subg. Esula, C ¼ subg. Euphorbia,
distinct or connate at base, 2-fid (unlobed). Fruits and D ¼ subg. Chamaesyce. This proposal has
capsular (drupaceous); columella persistent. also been taken up by, i.a., Horn et al. (2012),
Seeds carunculate or ecarunculate, testa smooth, who have provided the hitherto most comprehen-
ridged, pitted, or tuberculate. x ¼ 6–10; many spp. sive overall phylogenetic analysis of the genus
based on x ¼ 7 or x ¼ 10. and its structural traits. Although these clades
With > 2,000 spp., this is the largest genus in are strongly supported and, as such, valuable for
the Euphorbiaceae and one of the five largest further phylogenetic understanding of
Euphorbiaceae 209
Euphorbia, they seem hardly acceptable to us as of stipular spines accompanied by supernumerary

taxa because they are very heterogeneous, are not spines associated with the leaf bases (Uhlarz
distinguishable on the basis of morphology, and 1974). Inflorescences terminal or more often lat-
rather are the mere product of the algorithm. A eral with concomitant transition from sympodial
taxonomic subdivision of Euphorbia, which is a to monopodial growth. Cyathial glands 5(4); sti-
great desideratum, favorably would have to be pules + or 0; caruncles often +. The extreme
built upward from clearly circumscribed entities xerophytic subg. Euphorbia is nested within this
of lower rank such as sections. This has been large clade. About 700 spp., pantropical, but with
accomplished for the Chamaesyce Clade by greatest diversity in Africa/Madagascar.
Yang et al. (2012)1. D. Chamaesyce Clade. Herbs, subshrubs, or shrubs,
A brief characterization of the four clades sometimes succulent; leaves alternate or oppo-
reads as follows: site; stipules + or 0; inflorescences terminal;
cyathial glands 5, 4, or 2–1, glands sometimes
A. Esula Clade. Annual and perennial herbs, shrubs,
with petaloid appendages; caruncles rarely +.
and trees, some are stem succulents; inflorescence
About 600 spp. with mostly C3 or CAM photo-
terminal, pseudo-umbellate; branching sympo-
synthesis in the Old World including Africa/
dial; cyathial glands 4; stipules 0; caruncles +.
Madagascar and Australia, and 350 New World
About 480 spp., mainly in temperate Northern
spp. of mostly C4 photosynthetic herbs.
Hemisphere, outside this area woody as trees or
stem succulents in the Canary Isl., Africa, Mada-
gascar, and Arabia; very few in tropical America.
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Euphroniaceae Marcano-Berti in Pittiera 18: 16 (1989).
K. K U B I T Z K I
Medium-sized savannah trees or shrubs with con- pitting; wood parenchyma paratracheal-aliform;
spicuous arachnoid tomentum of unicellular rays uniseriate and biseriate, heterocellular Type II
hairs; branches terete, lanuginose, glabrescent. and III, containing gum; fibre tracheids thick-
Leaves alternate, simple, entire or revolute, pin- walled, non-septate (Espinoza de Pernia 1989).
nately veined; stipules minute, caducous. Inflores- The indumentum consists of unicellular lignified
cences terminal and subterminal racemes or and unlignified hairs, the latter dense and woolly,
thyrses; bracts minute; prophylls 0. Flowers bisex- and oxalate druses and tanniniferous tissue is found
ual, perigynous, obliquely monosymmetric; calyx in all floral parts (Matthews and Endress 2008).
with 5 subequal, imbricate sepals connate at the
base and inserted on a campanulate or campanu- F L O W E R S T R U C T U R E . The turbinate-campanulate
late-turbinate cup, the two outer sepals shorter floral tube seems to be glandular inside and may
and narrower than the inner ones, the inner sur- secrete nectar (Marcano-Berti 1989). The fertile
face of the floral cup lined by a smooth unlobed stamens appear to belong to two different cycles.
nectary; petals 3, distinct, all of same size, con- The ovules are bitegmic, incompletely tenuinucellar,
torted in bud, unguiculate; androecium with a anatropous and apotropous. The obliquely mono-
large anterior pointed staminode with sterile symmetric floral plan fits well with that of related
anther flanked by two stamen pairs and two pos- families (Chrysobalanaceae, Dichapetalaceae, Trigo-
terior groups of 2–3 tooth-like staminodes; each niaceae; see Matthews and Endress 2008, the INTRO-
stamen pair with a longer outer antesepalous sta- DUCTION TO MALPIGHIALES, and Fig. 48).
men and a shorter inner antepetalous stamen,
these fused with the dilated bases of their fila-
ments; anthers dorsifixed, dithecal, slightly apicu-
late, introrse, dehiscing with longitudinal slits;
filaments of fertile stamens apically glabrous, of
the large staminode retrorsely pilose throughout;
gynoecium posteriorly positioned, syncarpous;
ovary partly inferior, 3-locular; ovules apotropous,
bitegmic, incompletely tenuinucellar, 2 per locule,
axile, nearly collateral but one pendant, the other
erect and shorter; style antrorsely pilose; stigma
capitate. Fruit a 3-valved capsule with fleshy exo-
carp and persisting calyx and androecium, dehisc-
ing septicidally from apex towards base. Seeds 1
per locule, with thin endosperm, slightly winged.
A single genus comprising 3 spp., distributed
on white sand savannahs and heath forests in
northern South America (Brazil, Venezuela).
A N A T O M Y . Year rings obvious; vessels solitary, Fig. 48. Euphroniaceae. A Euphronia hirtelloides. B
with simple perforations and alternate intervascular E. acuminatissima. (Steyermark & Marcano Berti 1999;
drawn by Bruno Manara)
218 K. Kubitzki
P O L L E N M O R P H O L O G Y . The pollen of Euphronia

Selected Bibliography
is tricolporate-oblate with a triangular amb and a
rugulate exine (Lleras 1976).
Espinosa de Pernia, N. 1989. Estudio xilologico del genero
Euphronia. Pittieria 18: 57–61.
A F F I N I T I E S . Originally placed in Rosaceae or Litt, A., Chase, M.W. 1999. The systematic position of
Vochysiaceae, more recently the genus has been Euphronia, with comments on the position of Bala-
considered a member of Trigoniaceae. Lleras nops: an analysis based on rbcL sequence data. Syst.
Bot. 23: 401–409.
(1976, 1978) recognised some differences between Lleras, E. 1976. Revision and taxonomic position of the
Euphronia and Trigoniaceae and, in spite of the genus Euphronia Martius ex Martius & Zuccarini
lack of a well-developed spur in Euphronia, opted (Vochysiaceae). Acta Amazonica 6: 43–47.
for its vochysiaceous affinity. Although both Tri- Lleras, E. 1978. Trigoniaceae. Flora Neotropica Mono-
graph 19: 1–73.
goniaceae and Vochysiaceae traditionally were Marcano-Berti, L. 1989. Euphroniaceae: una nueva
included in the order Polygalales, molecular stud- familia. Pittieria 18: 15–19.
ies did not confirm a link between them. Litt and Matthews, M.L., Endress, P.K. 2008. Comparative floral
Chase (1999) provided strong support for a clade structure and systematics in Chrysobalanaceae s. l.
(Chrysobalanaceae, Dichapetalaceae, Euphronia-
of Euphronia + Chrysobalanaceae sister to Dicha- ceae, Trigonaceae; Malpighiales). Bot. J. Linn. Soc.
petalaceae + Trigoniaceae, a relationship that 157: 249–309.
since has been confirmed by several multigene Steyermark, J.A., Marcano-Berti, A. 1999. Euphroniaceae.
In: Berry, P. et al. (eds.) Flora of the Venezuelan
analyses and the comparative floral morphologi- Guayana, vol. 5. St. Louis: Missouri Bot. Gard. Press.
cal study by Matthews and Endress (2008).
Thus, the notion of Marcano-Berti (1989),
who had disagreed with the view of Lleras, is
fully justified.
A single genus:
Euphronia Mart. Fig. 48

Euphronia Mart. ex Mart. & Zucc., Flora 8: 32 (1825);
Steyermark, Ann. Missouri Bot. Gard. 74: 89–94 (1987);
Steyermark & Marcano-Berti, Fl. Venezuel. Guayana 5:
228–230 (1999). Lightia Schomb., Linnaea 20: 753 (1847).
Characters as for family.
Goupiaceae
Goupiaceae Miers, Ann. Mag. Nat. Hist. III, 9: 292 (1862).
K. K U B I T Z K I
Evergreen aluminium-accumulating trees. Leaves The nodes are unilacunar. The wood contains
alternate, distichous, petiolate, subtriplinerved vasicentric tracheids; vessel perforation is always
and transversely venose, sometimes domatiifer- scalariform with 3–10 thick bars, and the fibres
ous; stipules narrow, rather long, caducous. Inflo- have distinctly bordered pits. The rays are het-
rescence an axillary, umbel-like, pedunculate erogeneous and have elongate ends. The axial
raceme. Flowers bisexual, regular, (4)5-merous; parenchyma is apotracheal and paratracheal.
pedicels slender; bracts short and triangular,
hairy; sepals connate, lobes imbricate; petals very F L O W E R S T R U C T U R E . The following account of
long and subulate, concave, induplicate-valvate, the flower structure of Goupia reviews the excel-
the upper third sharply inflexed in bud and some- lent, well-illustrated description given by Miers
times geniculate or sigmoid at anthesis; nectary (1862) and his interpretation. The cup-shaped
disk large, cupular, thin-walled, sinuate on the calyx is deeply divided into five acute lobes that
margin; stamens 5, alternating with petals, have an imbricate aestivation. The petals are linear-
inserted on the inner edge of the nectary disk, oblong and fleshy and more than six times the
filaments very short, anthers small, ovoid, length- length of the calyx. Their straight margins are
wise dehiscing, with a thickened and apically introflexed and valvate in bud; their appendiciform
setose-pilose connective and ellipsoidal, separate, apices, measuring half their length, are inflected
introrse locules; pollen grains small, 3-colporate; and united together by their valvate margins. At
gynoecium syncarpous, partly enclosed by disk anthesis, the petals become horizontally expanded,
but free from it, ovary depressed-globose, 5-locu- with the inflected apices standing erect at right
lar; stylodia 5, subulate, stellately divergent; stig- angles with them. The disk forms a notable struc-
mas diminutive; ovules several in each locule, ture, being nearly the size of the calyx, and has five
basal, axile, ascending. Fruit a small, globose very short teeth that alternate with the stamens.
drupe, 1–3-locular, hard. Seeds 1–4 per fruit, The five stamens are erect and stand within the
erect, obovoid, with axile straight embryo in fleshy disk, free from it as well as from the ovary; their
endosperm; testa reticulate, pitted inside. n ¼ ? linear connectives extend beyond the anthers and
A monogeneric family with four or perhaps are truncated at their summit, where they are furn-
five spp. from tropical America. ished in front with a horizontal tuft of long hairs,
whereas their margins, behind the anthers, are
M O R P H O L O G Y A N D A N A T O M Y . Goupia glabra, ciliated with similar hairs. The anthers are bilobed
the most frequent species, is a vast, buttressed and introrse, each theca dehiscing by a longitudinal
tree corresponding with Roux’s architectural and somewhat oblique fissure. One half of each
model. The upper leaf epidermis is underlain by anther rises above the margin of the disk, and
a hypodermis. Stomata are laterocytic or rarely their long apical horizontal tufts of hairs meet in
anisocytic. The palisade tissue is two-layered. The the middle of the ovary, passing between the stylo-
spongy tissue is crossed by branched scelerench- dia and serving as collectors to convey their pollen
ymatous elements and veinlets, the ends of which to them. An illustration of Baillon (1877, p. 10)
are in close contact with vesiculose sclereids that shows a Goupia flower at the beginning of the
do not contain any visible contents (Rao and anthesis, in which the petals not yet spread and
Bhattacharya 1975). Cork originates superficially. the setose hairs at the anther tips are directed
220 K. Kubitzki
testa is about six cells thick and the subepidermal

layer is sclerified. The outer epidermis of the
tegmen is moderately thickened, but otherwise it
is crushed (Melikian and Savinov 2000).
A F F I N I T I E S . Traditionally incorporated in
Celastraceae, Miers (1862) raised this aberrant
taxon to family rank, whereas Loesener (1892)
retained it in Celastraceae as a distinct subfamily.
Molecular work (e.g. Simmons et al. 2001) con-
firmed the exclusion of Goupia from Celastraceae,
and subsequent work resolved it as member of the
broadly circumscribed order Malpighiales, as sis-
ter to Achariaceae (Davis et al. 2005), in an unsup-
ported position in the ’parietal clade’ (Violaceae/
Passifloraceae) (Wurdack and Davis 2009; Soltis
et al. 2011), and finally, with moderate support, in
a sister position with Violaceae (Xi et al. 2012),
with which it shares appendages on the stamen
connectives and haplostemony, among other
traits, but the differences are significant.
D I S T R I B U T I O N A N D H A B I T A T S . Goupia is a
genus of rain forest trees from northern South
America and Central America (Guatemala).
U S E S . Goupia glabra is a frequent and impor-

tant timber tree in the Guyanas and Pará, Brazil.
Fig. 49. Goupiaceae. Goupia glabra. A Flowering branch. Its light red-brown wood has a disagreeable smell
B Leaf of young plant and stipule. C Umbellate inflores- and has been used in carpentry and for railway
cence. D Floral bud. E Apical (left) and lateral (right, with sleepers.
two petals removed) views of flowers. F Petals. G Flower,
petals removed, nectary disk surrounding the stamens.
H Adaxial and lateral views of stamens. I Nectary disk A single genus:
surrounding pistil with distinct stylodia, petals removed.
J Same, medial section. K Leaf with infructescence. L Fruit Goupia Aubl. Fig. 49
and seed. (Reproduced with the kind permission of the
artist Bobbi Angell) Goupia Aubl., Hist. Pl. Gui. 1: 295, tab. 116 (1775); Miers,
Contr. Bot. 2: 131–137, t. 74 (1862); Oliver, Ic. Pl. III, 6,
upwards. This strange floral structure certainly t. 1590 (1897).
calls for live observations of the floral biology. Description as for family.
Four or perhaps five spp. from northern
P O L L E N M O R P H O L O G Y . The pollen is suboblate, South and Central America.
very small (11 x 12–13 mm), and tricolporate, the
amb is circular to convex triangular, and both the
ectoapertures and the transversal endoapertures Selected Bibliography
are very straight; the ectexine is microreticulate
or finely perforate to foveolate (Lobreau-Callen Araujo, P.A.M., Filho, A.M. 1973. Estrutura da madeira de
1977; Furness 2011). Goupia glabra Aubl. (Goupiaceae). Arq. Jard. Bot.
Rio de Janeiro 19: 149–153.
Baillon, H. 1877. Histoire des plantes, vol. 6. Paris:
F R U I T A N D S E E D . The fruits are probably bird- Hachette.
dispersed. The endocarp is heavily sclerified. The Davis, C.C. et al. 2005. See General References.
Goupiaceae 221
Furness, C.A. 2011. Comparative structure and develop- Metcalfe, C.R., Chalk, L. 1950. See General References.
ment of pollen and tapetum in Malpighiales, with a Miers, J. 1862. Contributions to Botany 2: 131–137. Lon-
focus on the parietal clade. Int. J. Pl. Sci. 172: 980–1011. don, Edinburgh: Williams and Norgate (repr. in Ann.
Lobreau-Callen, D. 1977. Les pollen des Celastrales: illus- Nat. Mag. Hist. Soc. III, 9: 289).
trations, commentaires. École Pratique des Hautes Rao, T.A., Bhattacharya, J. 1975. On foliar terminal vesi-
Études, Institut de Montpellier. culose sclereids in Goupia glabra Aubl. Curr. Sci. 44:
Loesener, T. 1892. Celastraceae. In: Engler & Prantl, Nat. 132–134.
Pflanzenfam, III, 5. Leipzig: W. Engelmann, Simmons, M.P., Clevinger, C.C., Savolainen, V., Archer,
pp. 189–222. R.H., Mathews, S., Doyle, J.J. 2001. Phylogeny of
Lundell, C.L. 1986. Goupia guatemalensis (Celastraceae), the Celastraceae inferred from phytochrome B
a genus and species new to Mesoamerica. Phytologia gene sequence and morphology. Am. J. Bot. 88:
57: 238–239. 313–325.
Melikian, A.P., Savinov, I.A. 2000. Goupiaceae. In: Takhta- Soltis, D.E. et al. 2011. See General References.
jan, A. (ed.) Anatomia seminum comparativa, vol. 6. Wurdack, K.J., Davis, C.C. 2009. See General References.
St. Petersburg: Nauka, pp. 135–136. Xi, Z. et al. 2012. See General References.
Humiriaceae
Humiriaceae Juss. in St.-Hil., Fl. Bras. mérid. 2: 87 (1827), nom. cons.
K. K U B I T Z K I
Evergreen trees or shrubs; wood often with bal- from pulpy to fibrous; endocarp woody, usually
samic juice; cork subepidermal; indumentum of very hard, compact or containing many round,
simple hairs, usually sparse or 0. Leaves alternate, resin-filled cavities, rarely spongious-woody, (4)5
simple, involute, often distichous, penninerved, (7)-septate, with 1–2(5) seeds developed;
entire, crenulate or slightly serrate, petiolate or dehiscence germinal with longitudinal valves at
rarely sessile, sometimes decurrent along the back of each carpel; subapical foramina alter-
branches; stipules very small, lateral, caducous nating with valves often present. Seeds oblong,
or 0. Inflorescences axillary or terminal, panicu- germinating enclosed in the fruit, retaining some
late, often corymbiform, dichotomous or trichot- nucellar remnants; embryo straight or slightly
omously branched, branchlets often articulate; curved; endosperm fleshy and oily. x ¼ 12.
bracts and prophylls small, amplectant, persistent A family of 8 genera and about 50 species,
or caducous. Flowers hermaphroditic, actino- from Costa Rica through tropical America to
morphic, pedicellate; sepals 5, connate, quin- southern Brazil, one sp. in coastal West Africa.
cuncially arranged or imbricate, all of same size
or 2 outer ones smaller; petals 5, caducous or VEGETATIVE MORPHOLOGY AND ANATOMY. Branching
sometimes persistent, distinct, white (red), con- is from the previous flush. The leaves are often
torted, cochlear or quincuncial; stamens numer- distichously arranged; the buds on growing
ous or in definite number of 10–30 and 1–2- points are long-pointed and elongate while still
seriate, alternating in length, sometimes the 5 rolled up and often have longitudinal lines down
antepetalous ones trifurcate at apex; filaments at the blade (Stevens 2001). The leaf blade is usually
base connate in a long tube; anthers dorsifixed coriaceous, and the margin varies from indis-
or subbasifixed; thecae 2, bilocular and each cell tinctly entire to crenulate and dentate.
dehiscing by longitudinal slit, or else 4 or 2, Stomata are anomocytic in Endopleura,
unilocular, dehiscing by detachment or some- Humiria, Humiriastrum and Sacoglottis, and
times by a slit; connective thick, fleshy, most paracytic in Vantanea. Sclerenchymatous idio-
commonly produced in an apiculum or lingui- blasts extending from the upper to the lower
form appendix, or obtuse; some stamens occa- epidermis are found in Sacoglottis. Clustered
sionally staminodial; disk intrastaminal, girding and solitary crystals are found in the mesophyll.
the ovary, membranaceous or subcoriaceous, Vessels are solitary and their elements have sim-
tubular or cupular, dentate, lobate, laciniate or ple or scalariform perforations and their mem-
composed of 10–20 distinct scales; gynoecium bers are extremely long. Axial parenchyma is
syncarpous, (4)5(7)-carpellate, carpels opposite predominantly diffuse and somewhat vasicentric
the sepals or rarely alternating with them; ovary or ranging to confluent. Rays are uniseriate or 2
ovoid or ellipsoid, (4)5(8)-locular; placentation (4) cells wide and have conspicuous marginal
axile; ovules 1 or 2 per cell, pendulous, anatro- rows of upright or square cells which are filled
pous, epitropous, with the micropyle facing with a gum-like substance. Fibres are very long
upwards and the raphe ventral, when 2 then and have bordered pits (Metcalfe and Chalk 1950;
superposed and the lower hanging from the lon- Queiroz de Vilhena 1978).
ger funicle; style entire; stigma capitate, 5-lobate The sieve element plastids contain protein
or 5-radiate. Fruit drupaceous; exocarp varying crystalloids and starch (P5cs type, Behnke 1991).
224 K. Kubitzki
F L O W E R S T R U C T U R E . Floral structure including

anatomy has been studied by Narayana and Rao
(1969–1977). Sepals are 3-traced; aestivation is
quincuncial throughout, and their degree of
fusion varies; in one Vantanea (V. parviflora),
the limbs of the sepals are completely reduced.
Aestivation of the corolla varies from quincuncial
to contorted; petals are 3-traced in Vantanea but
1-traced in probably all other genera. The stamen
filaments are basally more or less connate into a
shorter or longer tube, which in Humiria extends
up to one third or a half of their entire length.
From its peripheral part the staminal traces sepa-
rate, while the inner part of the tube forms the
disk. In the fully differentiated flowers, however,
vascular bundles have only been observed in the
disks of Vantanea, Humiria and Schistostemon.
The multistaminate androecium of Vantanea
has been interpreted as 3-whorled by Cuatrecasas
(1961) but the floral anatomical studies of Nar-
ayana and Rao (1969) show that the densely
packed stamens do not form distinct whorls. In
genera such as Humiria and Humiriastrum, the
biseriate androecium consists of 20 stamens with
an outer antesepalous whorl of 15 stamens
arranged in five groups each of three and an
Fig. 50 Humiriaceae. Vantanea guianensis. A Flowering
inner whorl of five antepetalous stamens. The branchlet. B Leafy apex of flowering stem. C Flower bud.
bundle supply of the triplets indicates that they D Flower. E Vertical section of base of flower, petals and
are stamen fascicles. In Sacoglottis there are ten stamens removed. F Longitudinal section of flower with
stamens in two whorls, and in Duckesia among the petals removed; note connate stamen filaments and cup-
ular disk. G Adaxial and lateral view of stamens showing
numerous stamens the five antepetalous ones are expanded connective. H Entire fruit and endocarp.
usually fertile and in Schistostemon the antesepa- (Reproduced with the kind permission of the artist
lous stamens are trifurcate. Thus, to my mind, the Bobbi Angell)
androecium of the family appears to be built upon
a basically (ob)diplostemonous plan extended by usually antesepalous; only in Humiria are they
lateral dédoublement and further reduction, constantly antepetalous, as first observed by
rather than being basically multistaminate. Urban (1877); there is no explanation for this
The complete 4-sporangiate anther is only anomaly. There are two superposed ovules per
found in Vantanea. In the two superposed pairs carpel in Vantanea and Humiria, each of the mar-
of thecae of Endopleura, the outer pair of spor- ginal placentae bearing one ovule; locules of the
angia is sterile in the upper thecae and the inner remaining genera are uniovulate. The common
part in the lower (Rao and Narayana 1965). Fur- style has a stylar canal lined by transmitting tissue
ther reduction leading to the complete loss of and is traversed by the dorsal carellary bundles; in
sporangia is met with in the rest. The outer pair Endopleura, the common median lateral bundles
of sporangia is sterile in Humiria and Schistoste- extend into the style (Rao and Narayana 1965).
mon, whereas in Humiriastrum and Sacoglottis it
is the inner ones (Narayana and Rao 1969–1977). E M B R Y O L O G Y . The ovule is anatropous and
Whereas the androecium shows a great varia- epitropous with the micropyle facing upwards,
bility, the ovary is nearly always 5-carpellate and bitegmic and crassinucellate (Netolitzky 1926).
always syncarpous. Orientation of the carpels is In Humiria balsamifera, which has been studied
Humiriaceae 225
by Boesewinkel (1985), the nucellus is large and outer epidermis develops into a fibrous exoteg-
persistent and an endothelium is lacking; the men with heavily thickened, lignified cells (Boe-
outer integument is mainly 2-layered and sewinkel 2000). The development of the seed coat
contains tannins. The inner layer of the 3-layered of Humiria shows much resemblance to that of
inner integument contains also tannins and is not Linaceae and Erythroxylaceae, which differ, how-
endothelial. The micropyle is formed by both ever, in having less strongly fortified seed coats,
integuments. The embryo sac is 8-nucleate and lacking nucellar remnants in the mature seed and
the endosperm in Vantanea is probably nuclear having an endothelium.
(Mauritzon 1934).
P H Y T O C H E M I S T R Y . The composition of the bal-
P O L L E N M O R P H O L O G Y . Pollen is suboblate to samic juice is unknown. Bergenin and gallate are
subprolate, 26–58 x 25–47 mm and 3–4-colporate; present in the bark of Sacoglottis. The seeds of
in LM, the tectum appears punctitegillate or more Sacoglottis gabonensis contain considerable
rarely faintly reticulate, the colpi are provided amounts of palmitinic, oleic and linolic acids
with sexinous thickenings, and the ora are lalon- (Hegnauer 1966, 1990).
gate to lolongate. Pollen of the Vantanea type
(all genera except Humiria) has long ectocolpi AFFINITIES AND SUBDIVISION OF THE FAM-
and a thick columellar infratectum, whereas the ILY. Humiriaceae were considered as part of
Humiria type (Humiria only) has short ectocolpi Engler’s Geraniales and Hutchinson’s Malpigh-
and a thin columellar infratectum (Erdtman 1952; iales, in both groupings together with Linaceae
Suryakanta 1974; Bove and Melhem 2000). (including Ixonanthes and Ctenolophon) and
Erythroxylaceae. Of the two large-scale molecu-
K A R Y O L O G Y . The chromosome number of lar analyses of Wurdack and Davis (2009) and
Sacoglottis gabonensis was determined as 2n ¼ 12. Soltis et al. (2011), the former placed the family
in an unresolved position in Malpighiales,
F R U I T A N D S E E D . The drupaceous fruit varies in whereas the latter found strong support for its
size from the size of a pea to that of a hen’s egg. sister relationship with the rest of the Malpigh-
The coriaceous, juicy or fibrous mesocarp encloses iales clade. In the analysis of Xi et al. (2012),
the woody fruit stone (pyrene), which in the five Humiriaceae are resolved in a sister position to
locules contains usually only 2–5 seeds. Humiria the families of the pariatal clade.
has two fertile seeds per locule, whereas all other A cladistic analysis of 40 morphological char-
genera have single-seeded locules. The endocarp is acters (Herrera et al. 2010) confirmed the obvious
usually very hard and in some genera contains placement of Vantanea as sister to the rest of the
many round, resin-filled cavities which may con- family, followed by Humiria and the other
tribute to the buoyancy of the fruits. At germina- genera, and with Sacoglottis and Schistostemon
tion the seedlings seem to push away the valves in the latest diverging subclade, as also antici-
which cover every locule or alternatively may seek pated by Cuatrecasas (1961).
exit through the apical foramina of the pyrene,
which are present in the endocarp of some genera. DISTRIBUTION, HABITATS AND DISPERSAL. Humiria-
After fertilisation, a nuclear endosperm ceae are members of the neotropical rain forests
develops and continues to resorb the nucellus and sometimes of woodland and savannahs up to
and eventually becomes cellular, fleshy and oily. an elevation of 1,400 m. They grow mostly in
In the mature seed there remains always a thin upland (non-flooded) forest, some species also
layer of nucellar tissue (perisperm) bordering the along creeks. In the Guyana Highland, Humiria
seed coat. The outer integument remains 2- or balsamifera is rich in local varieties and is an
3-layered but its outer epidermis (exotesta) is important constituent of scrub, savannah and
fortified by thick-walled cells with lamellate and woodland communities on rocks and yellow
lignified, pitted cell walls. The inner integument and white sand and grows up to an altitude of
divides further and becomes 4–6-layered, and its 2,300 m.
226 K. Kubitzki
Only one species, Sacoglottis gabonensis – Most stamens fertile; endocarp not resinous-lacunose,
(Baill.) Urb., is found outside tropical America, at the base with 5 elevated prominent ribs each divided
on the West African coast, where it is a tall tree into 2 higher up 3. Endopleura
5. Stamens 20–30, only 5–15 fertile; endocarp promi-
growing in inundated forest. nently 5-costate, not resinous 5. Hylocarpa
Humiriaceous fruits or endocarps are often – Stamens 10–20 6
found in drifts and drift deposits of the Amazon 6. Stamens 10; endocarp resinous, often bullate, shallowly
and Magdalena rivers. Particularly those of Saco- or inconspicuously furrowed; valves broad, alternating
glottis amazonica have been carried by currents ribs thin, inconspicuous 8. Sacoglottis
from the Orinoco and Amazon deltas to the – Stamens 20 7
shores of the West Indies and Central America 7. Stamens all with a single anther, 10 longer ones alter-
nating with 10 shorter ones; thecae basally attached;
without having been established there, and even endocarp with 5 elliptic, short valves in the upper half
across the Atlantic to the British Islands (Cuatre- of fruit alternating with small apical foramina
casas 1961). The age and means of the establish- 6. Humiriastrum
ment of Sacoglottis gabonensis in West Africa are – Five antesepalous stamens distally trifurcate and tri-
unknown. antheriferous, 5 antepetalous stamens shorter, with a
single anther, and the remaining 10 shortest, between
sepals and petals; endocarp resinous-lacunose, valves
F O S S I L R E C O R D . The fossil record of the family broad, apical foramina usually lacking
was evaluated by Cuatrecasas (1961), who 7. Schistostemon
excluded from it all finds from outside of South
America. More recently, Herrera et al. (2010)
reinvestigated and listed all fossil endocarps
which, with the exception of one from Bahia GENERA OF HUMIRIACEAE
(Selling 1945), are from the Andean countries
between Costa Rica and Bolivia. The majority 1. T R I B E V A N T A N E E A E Cutrec. (1961).
are of Neogene age (Miocene/Pliocene); Vanta-
nea dates back to the Eocene. The fossil pollen of Stamens numerous; anthers with 2 basal-lateral
Psilabrevitricolporites devriesii (Lorente) Silva- bilocular thecae; anther locules longitudinally
Caminha et al., which is similar to extant dehiscing; carpels opposite sepals, 2-ovulate.
Humiria, has been observed in beds dating back
to the Early Miocene along the northern Andes
and from western Amazonia (Herrera et al. 2010). 1. Vantanea Aubl. Fig. 50
Vantanea Aubl., Pl. Guiane 1: 572, t. 229 (1775).
U S E S . Local use is made of the oil content of the
seeds. Schultes (1979) gives details of interesting Trees. Bracts caducous; stamens 50–120(180),
uses of the family by indigenous people in the in 3 indistinct series, anthers 2-thecate, bilocu-
north-western Amazon. lar. Drupe medium-sized to large, ovoid or
ellipsoid, the exocarp carnose; endocarp
woody, not resinous, dehiscent at seed germina-
KEY TO THE GENERA tion by oblong valves. Seeds usually only 1 per
1. Stamens numerous (30–180) and pluriseriate; anthers fruit.
with 2 bilocular thecae dehiscing longitudinally About 15 spp., from Costa Rica and through-
1. Vantanea out northern South America southwards to Boli-
– Stamens 10–30; anthers with unilocular, distinct thecae via and Brazil (Santa Catarina).
dehiscing by detachment 2
2. Carpels 2-ovulate, ovary locules opposite petals; thecae
of anthers hirsute 2. Humiria
– Carpels 1-ovulate, ovary locules opposite sepals; thecae 2. T R I B E H U M I R I E A E Cuatrec. (1961).
of anthers glabrous 3
3. Anthers 4-thecate; stamens 20–30 4
– Anthers 2-thecate; stamens 10–30 5 Stamens 10–30, in 1–2 series; anthers with 2–4
4. Usually only 5 stamens fertile; endocarp spongy-lacu- unilocular thecae; carpels opposite sepals or
nose, evenly costate with long valves 4. Duckesia alternating with them, 2- or 1-ovulate.
Humiriaceae 227
2. Humiria St.-Hil. Trees. Bracts persistent; stamens biseriate, 20–25,

Humiria St.-Hil., Exp. Fam. 2: 374 (1805); Cuatrecasas, plus occasionally some shorter staminodes;
Contr. US Nat. Herb. 35: 67–122 (1961), rev. anthers glabrous, usually only 5 fertile; connec-
tive lanceolate, carnose; thecae 4, 2 attached at
Trees or shrubs. Leaves often decurrent on base of connective, 2 laterally in the middle; disk
branchlets. Bracts persistent; stamens 20, uniseri- of 10 subulate distinct scales surrounding the
ate, united into a tube for about halfway up from ovary. Drupe large, ovoid; exocarp thick, coria-
base, longer and shorter alternating; filaments ceous; endocarp verrucose, resinous-lacunose;
glandular; thecae inserted at base, hirsute; disk germinal valves 5, alternating with 5 apical
annular, of 20 linear, thick, united scales; carpels foramina. Seeds few, oblong.
opposite petals; ovary cells with 2 superposed One sp., D. verrucosa (Ducke) Cuatrec., Bra-
pendulous ovules. Drupe small (<16 mm), zilian Amazonia.
ovoid, ellipsoid or oblong; exocarp thin, meso-
carp fleshy, palatable, endocarp woody, with 10
equidistant striae marking 5 narrow germinal 5. Hylocarpa Cuatrec.
valves alternating with 5 small apical foramina. Hylocarpa Cuatrec., Contr. US Nat. Herb. 35: 84 (1961).
Seeds commonly 4–1.
Four spp. in rain forests, woodland and Trees. Bracts caducous; stamens 30, biseriate,
savannah vegetation of tropical South America, only 5–15 with fertile anthers; connectives thick,
H. balsamifera (Aubl.) St.-Hil. widespread with club or hammer shaped; thecae 2, basal; disk of 10
numerous varieties and forms, an important distinct, thick scales. Drupe large, fusiform, exo-
constituent of scrub or thicket communities carp thick; endocarp woody, not resinous, prom-
in savannahs and the Guayana sandstone massif. inently 5-costate, with germinal valves between
the furrows. Seeds oblong, 1–3(5) per fruit.
Only one sp., H. heterocarpa (Ducke) Cua-
All other Humiriaceae: trec., in the upper Rio Negro region of Brazil.
Anther sacs glabrous; carpels opposite sepals;
ovary cells 1-ovulate.
6. Humiriastrum (Urb.) Cuatrec.
Humiriastrum (Urb.) Cuatrec., Contr. US Nat. Herb. 35:
3. Endopleura Cuatrec. 122–146 (1961).
Sacoglottis subg. Humiriastrum Urb.
Endopleura Cuatrec., Contr. US Nat. Herb. 35: 80 (1961).
Trees. Bracts persistent; stamens biseriate, 20–30, Trees. Stamens 20, of alternating length, in 1 series;
united in lower half or third, the largest opposite thecae 2, basal; disk a dentate ring or of distinct
the petals; anthers dorsifixed; thecae 4, 2 basal scales. Drupe medium sized to small, ellipsoid to
and 2 apical or lateral, sometimes 2 thecae or subglobose, exocarp carnose; endocarp woody,
rarely all 4 sterile; disk of 10 triangular-ovate usually without resinous cavities; germinal valves
scales, united at base. Drupe large, ellipsoid, exo- in upper half of fruit, oblong, alternating with 5
carp coriaceous; mesocarp fleshy; endocarp apical foramina. Seeds 1–2(5) per fruit.
woody, not resinous, with 5 strongly elevated About 16 spp. spread from Costa Rica through-
ribs at the base each divided into two higher up; out tropical South America from the eastern to the
germinal valves inconspicuous between the western coast southwards to Rio de Janeiro.
unbranched ribs; apical foveola 5. Seeds usually
2–3 per fruit. 7. Schistostemon (Urb.) Cuatrec.
One sp., E. uchi (Huber) Cuatrec., Brazilian
Schistostemon (Urb.) Cuatrec., Contr. US Nat. Herb. 35:
and Venezuelan Amazonia. 146–161 (1961).
Sacoglottis subg. Schistostemon Urb.
4. Duckesia Cuatrec. Trees. Bracts persistent; stamens 20, in one series,
Duckesia Cuatrec., Contr. US Nat. Herb. 35: 76 (1961). the longest 5 antesepalous, trifurcate at apex and
228 K. Kubitzki
triantheriferous, 5 a little shorter, antepetalous, Hallier, H. 1923. Beitr€age zur Kenntnis der Linaceae. 9.
entire and with single anthers, 10 short, between Die Humiriaceen. Beih. Bot. Centralbl. 39, Abt. 2:
56–62, 174.
sepals and petals, also with single anthers; disk Hegnauer, R. 1966, 1990. See General References.
cupular, dentate, or rarely of 10 distinct scales. Herrera, F., Manchester, S.R., Jaramillo, C., MacFadden,
Drupe rather large, exocarp carnose, endocarp B., Silva-Caminha, S.A. da 2010. Phytogeographic
woody, resinous-lacunose, the surface rugose or history and phylogeny of the Humiriaceae. Int.
J. Pl. Sci. 171: 392–408.
bullate, valves broad, separating ribs thin, apical Mauritzon, J. 1934. Etwas u €ber die Embryologie der Zygo-
foveola usually 0. phyllaceae sowie einige Fragmente u €ber die Humir-
About nine spp. in northern South America iaceen. Bot. Notiser 1934: 409–422.
southwards to Peruvian and Brazilian Amazonia. Metcalfe, C.R., Chalk, L. 1950. See General References.
Narayana, L.L., Rao, D. 1969. Contributions to the floral
anatomy of Humiriaceae 1. J. Jap. Bot. 44: 328–335.
Narayana, L.L., Rao, D. 1973. Contributions to the floral
8. Sacoglottis Mart. anatomy of Humiriaceae 2. J. Jap. Bot. 48: 143–146.
Sacoglottis Mart., Nov. Gen. Spec. Pl. 2: 146 (1827). Narayana, L.L., Rao, D. 1973. Contributions to the floral
anatomy of Humiriaceae 3. J. Jap. Bot. 48: 242–246.
Trees. Bracts persistent or deciduous; stamens 10, Narayana, L.L., Rao, D. 1976a. Contributions to the floral
5 antesepalous and longer than the 5 antepetalous anatomy of Humiriaceae 4. J. Jap. Bot. 51: 12–15.
Narayana, L.L., Rao, D. 1976b. Contributions to the floral
ones; thecae 2, attached dorsally near base; disk anatomy of Humiriaceae 5. J. Jap. Bot. 51: 42–44.
cupular, dentate; drupe of medium to large size, Narayana, L.L., Rao, D. 1977. Contributions to the floral
exocarp carnose, endocarp woody, globose, bul- anatomy of Humiriaceae 6. J. Jap. Bot. 52: 145–153.
late, resinous-lacunose, valves broad, apical Netolitzky, F. 1926. See General References.
Queiroz de Vilhena, R.C. 1978. Anatomia foliar de très
foramina rare. espécies da famı́lia Humiriaceae. Acta Amazonica 8:
About eight species, seven in northern South 25–43.
America and one, S. gabonensis (Baill.) Urb., in Rao, D., Narayana, L.L. 1965. Vascular anatomy of
coastal West Africa. Humiriaceae. Curr. Sci. 34: 383–384.
Record, S. 1944. Random observations on tropical Amer-
ican timbers; Humiriaceae in Central America. Trop.
Woods 77: 8–9.
Selected Bibliography Reiche, K. 1890. Humiriaceae. In: Engler & Prantl,
Die nat€ url. Pflanzenfam. III, 4: 35–37. Leipzig:
W. Engelmann.
Behnke, H.-D. 1991. See General References. Schultes, R.E. 1979. Interesting uses of the Humiriaceae in
Boesewinkel, H.-D. 1985. The ovule and seed of Humiria the northwestern Amazon. J. Ethnopharmacol. 1:
balasamifera (Aubl.) St. Hil. Acta Bot. Neerl. 34: 89–94.
183–191. Selling, O.E. 1945. Fossil remains of the genus Humiria.
Boesewinkel, F.D. 2000. Semeistwo Humiriaceae, pp. Sv. Bot. Tidskr. 39: 258–269, figs. 1–24.
16–18. In: Anatomia Seminum Comparativa, vol. 6. Soltis, D.E. et al. 2011. See General References.
St. Petersburg: Nauka. Stevens, P.F. 2001 onwards. See General References.
Bove, C.P. 1997. Phylogenetic analysis of Humiriaceae Suryakanta 1974. Pollen morphological studies in the
with notes on the monophyly of Ixonanthaceae. Humiriaceae. J. Jap. Bot. 49: 112–122.
J. Comp. Biol. 2: 19–24. Urban, I. 1877. Humiriaceae. In: Martius, Flora Brasilien-
Bove, C.P., Melhem, T.S. 2000. World Pollen and Spore sis XII, 2: 425–454, pl. 92–96.
Flora 22: 1–35. Humiriaceae Juss. Urban, I. 1878. Die Begrenzung der Gattungen in der
Cuatrecasas, J. 1961. A taxonomic revision of Humiria- Familie der Humiriaceae. Sitz.-Ber. Ges. Naturf.
ceae. Contr. US Nat. Herb. 35, 2: 24–214. Freunde 1878: 2–5. Berlin.
Cuatrecasas, J., Huber, O. 1999. Humiriaceae, pp. Winkler, H. 1931. Unterfam. IV, Humirioideae. In: Engler
623–641. In: Berry, P.E., Yatskievych, K., Holst, B.E. & Prantl, Die nat€ url. Pflanzenfam. ed.2, 19a, pp.
(eds.) Flora of the Venezuelan Guayana, vol. 5. St. 126–129, figs. 58, 59. Leipzig: W. Engelmann.
Louis: Missouri Bot. Gard. Press. Wurdack, K.J., Davis, C.C. 2009. See General References.
Erdtman, G. 1952. See General References. Xi, Z. et al. 2012. See General References.
Irvingiaceae
Irvingiaceae Exell & Mendonça (1951), nom. cons.
K. K U B I T Z K I
Glabrous trees; wood extremely hard; leaves some- intrapetiolar and folded to form a cone around
times papillate underneath; mucilage cells in leaf the terminal bud; they are early caducous and
and stem epidermis; secretory canals containing leave circular scars on the twig. In Klainedoxa
mucilage in leaves and stems. Leaves alternate, they are of enormous size, fall almost at once
simple, entire, petiolate, pinnately veined, coria- and then are found in large numbers beneath
ceous; stipules very large, unequal, intrapetiolar, the tree; in Desbordesia, they are usually less
encircling the terminal bud, early caducous and than 1 cm long but in Irvingia grandifolia occa-
leaving a very distinct scar. Inflorescences panicu- sionally up to 8 cm long.
late, axillary or terminal. Flowers small, hermaph- The leaves are completely glabrous but some-
roditic, regular, pedicels articulated; sepals 5, times papillate on the undersurface, and mucilage
small, imbricate; petals 5, free, imbricate, exceed- cells are present in the leaf and stem epidermis.
ing the sepals; stamens 10 (9), distinct, exceeding Stomata are paracytic. Mucilage cells, mucilage
the petals, inserted below the large intrastaminal spaces and cristarque cells are found in the leaf
nectary disk, filaments plicately folded in mesophyll and the stem. Vessel elements have
bud; anthers subbasifixed, 2-locular, globular, simple perforations; lateral pitting is alternate;
dehiscing by slits; gynoecium syncarpous, 5–4(2)- axial parenchyma is in continuous bands; rays
carpellate; ovary superior, 5–4(2)-locular; ovule 1 are homogeneous.
per locule, epitropous, anatropous, bitegmic and
crassinucellar, pendulous from the top of the cen- F L O R A L S T R U C T U R E A N D B I O L O G Y . The flowers
tral axis; obturator present; style terminal, short; of Irvingiaceae have a prominent, fleshy nectary
stigma punctiform. Fruit a drupe with 1 or (5)4 disk, which surrounds the base of the ovary and
pyrenes or a broadly winged samara. Seeds with pre-soaks among the bases of the filaments.
large embryo; cotyledons flattened, cordate; endo- Along a circular line on the crest of the disk of
sperm fatty, almost 0 to copious. Klainedoxa, Link (1992) observed 10–15 stomata
Three genera and about 11 spp. mainly from deeply sunken each at the bottom of a trumpet-
W and C Africa, one of them from SE Asia and W shaped duct and underlain by a voluminous cav-
Malesia. ity. The disk in an Irvingia was similar but the
stomata were fewer in number and less concealed.
VEGETATIVE MORPHOLOGY AND ANATOMY. Most spe- In all three genera the disks attract attention
cies are vast, buttressed forest trees often with through their size relative to the dimensions of
fluted boles; Desbordesia glaucescens is reported the flowers, and their bright yellow colour and
to grow up to 55 m tall and to develop buttresses strong nectar production (Harris 1996); however,
up to 8 m high. The young stem of Klainedoxa is details about pollinators and the diurnal rhythm
often heavily armed with blunt spines up to 12 cm of nectarial secretion remain unknown.
long. In Klainedoxa the bark is dark brown, and Matthews and Endress (2011) gave a detailed
there are no fibres at the centre of the slash. In description of the floral morphology of one spe-
Desbordia and Irvingia the bark is white to pale cies of Irvingia, and compared it with that of
brown, and in the slash of the latter genus promi- putatively related families, but could not find
nent fibres are present towards the centre of good support from floral morphology for the
the slash. Leaves are involute. Stipules are clade Linaceae, Irvingiaceae and Caryocaraceae.
230 K. Kubitzki
P O L L E N M O R P H O L O G Y . Pollen is quite uniform, prises up to 20 layers of thickened, but unlignified,

spheroidal, 17–23 x 18–24 mm, tricolporate, mostly tangentially elongated cells; also the epider-
prominently triangulaperturate with plane or mis is unlignified. The mesocarp is very broad and
concave intercolpia and emarginate apertural mostly composed of thin-walled parenchyma with
angles, ectoaperture 6 (7.5) mm long, endoaper- numerous large secretory canals, isolated fibres,
ture nearly rectangular; tectum continuous, fibre bundles and vascular bundles ensheathed by
appearing granular in LM but either verrucose fibres. The endocarp consists of a complex pattern
or striate-rugulate in SEM (Sabatier 1974). of elongated sclereids in tangential or longitudinal
orientation and an inner epidermis that is strongly
K A R Y O L O G Y . For Irvingia gabonensis 2n ¼ 28, lignified. The pericarp structure of Allantosper-
and for Klainedoxa 2n ¼ 26 have been reported. mum (see Ixonanthaceae, this volume) was found
to be very different.
F R U I T A N D S E E D . The pericarp anatomy of The seeds (Desbordesia and Klainedoxa stud-
Irvingia and Klainedoxa has been studied by ied by Netolitzky 1926, Irvingia by Tobe and
Fernando and Quinn (1992). The exocarp com- Raven 2011) have a fibrous exotegmen, whilst
the mature testa is sclerotic throughout and tra-
versed by post-chalazal branches of the raphe
bundle. Tobe and Raven (2011) found no similar-
ity with the seed structure of other Malpighiales
families but rather with Huaceae and Connara-
ceae (Oxalidales).
Germination is epigeal and phanerocotylar
throughout.
D I S P E R S A L . The following interesting details are

taken from Harris (1996). The fruits of Klainedoxa
gabonensis and various Irvingia are swallowed
whole by elephants. The pyrenes separate in their
digestive tract and are passed out in the dung,
from where they have been observed germinating.
Lowland gorillas usually eat the mesocarp but do
not swallow the pyrenes. Yellow backed duikers
(Cephalophus sylvicultor) also swallow these fruits
and regurgitate the pyrenes after digesting the
mesocarp. Squirrels and other rodents occasion-
ally open pyrenes and eat the seeds. The fresh
fruits of Irvingia smithii, which grows in season-
ally flooded and riparian forest, develop slimy air
bubbles in the meocarp and are then carried by
the current; when the mesocarp starts to rot, the
fruits lose their buoyancy and sink. The fruits are
also eaten by a wide variety of fish. The samaras of
Desbordesia are blown by the wind.
P H Y T O C H E M I S T R Y . Irvingia contains gallic acid

and ellagic acid (Nooteboom 1967).
Fig. 51. Irvingiaceae. Klainedoxa gabonensis. A Flower-
ing branch, the shoot tips with stipules clasping the ter-
A F F I N I T I E S . Mainly on account of the presence
minal bud. B Flower. C Disk and ovary. D Fruit. E Same, of the mucilage spaces and mucilage cells mis-
transverse section. F Pyrene after remainder of fruit has taken as secretory ducts, Irvingiaceae have been
rotted. (Harris 1996; drawn by Rosemary Wise) considered close to, or included in,
Irvingiaceae 231
Simaroubaceae, which differ profoundly in having – Ovary 2-celled; fruit a drupe with a single pyrene
compound, estipulate leaves, scales on the fila- or fruit a samara; stipules, covering terminal buds,
ments, distinct stylodia and more or less distinct to 4 (very rarely 8) cm long 2
2. Fruit a 1-seeded drupe 2. Irvingia
carpels, and in containing the rutalean nortriter-
– Fruit a 2-seeded samara, broadly winged all around
penoid bitter principles. Hallier (1923) included 3. Desbordesia
Irvingia in his broadly construed Linaceae, which
comprised, i.a., Erythroxylaceae, Ixonanthaceae
and Humiriaceae. Hutchinson (1973) placed the 1. Klainedoxa Pierre ex Engler Fig. 51
latter three families with Irvingiaceae in his Mal- Klainedoxa Pierre ex Engler in Engler & Prantl, Nat.
pighiales. Forman (1965) supported the inclusion Pflanzenfam. III, 4: 227 (1896); Harris, Bull. Jard. Bot.
of the Ixonanthoideae into the Irvingiaceae. In Nat. Belg. 65: 152–166 (1996), rev.
DNA sequence analyses, Irvingia was resolved as Tall trees; stipules massive, ribbed, very long.
sister to the Linaceae, although with low support Leaves not papillose on undersurface. Inflores-
(Tokuoka and Tobe 2006; Korotkova et al. 2009), cences terminal. Flowers congested on lateral
and without support as sister to Ixonanthaceae in branches of the panicles; ovary 5-locular; ovule 1
the clusoid clade (Soltis et al. 2011). In a 4-gene per locule. Fruit a hard drupe; exocarp thin, fleshy;
analysis, Ruhfel et al. (2011) found Irvingia highly mesocarp firm but mucilaginous, fibrous, pyr-
supported as sister to Bruguiera (Rhizophoraceae) enes 5, oblong with very hard endocarp, fibrous
and Cyrillopsis (Ixonanthaceae). In the phyloge- outside. Seeds oblong, with copious endosperm.
nomic analysis of Xi et al. (2012), the Irvingiaceae Two spp. are recognised by Harris, widely
are placed, although with moderate support, in a distributed in the forested regions of W and C
clade together with the Pandaceae; see there (this Africa.
volume) for possible synapomorphies.
D I S T R I B U T I O N A N D H A B I T A T S . The family is 2. Irvingia Hook. f.

restricted to the humid zone of West and Central Irvingia Hook. f. in Trans. Linn. Soc., Bot. 23: 167 (1860);
Africa with the exception of one species (Irvingia Harris, Bull. Jard. Bot. Nat. Belg. 65: 166–196 (1996), Afric.
malayana) in SE Asia/W Malesia. Most of the spp., rev.
African species and Irvingia malayana grow in Trees, often vast. Leaves sometimes papillose on
non-inundated lowland forest; some enter into under surface; stipules large. Inflorescences axil-
dry or flooded forest, savannah or secondary forest. lary and terminal panicles or compound racemes.
Flowers inserted at regular intervals or in fasci-
P A L A E O B O T A N Y . A fossil wood, Irvingiaceoxylon cles on main branches of inflorescence; ovary 2-
dechampsii, is reported from the upper Pliocene/ locular; ovules 1 per locule. Drupes large, edible,
lower Pleistocene of Ethiopia (Gros 1983), testify- thick and fibrous, rounded-oblong in outline,
ing to the extension of humid forests to this region compressed, 1(2)-locular and 1(2)-seeded; pyr-
by that time. Irvingioxylon taibaense, described by ene(s) with thick wall covered with fibres. Seeds
Koeniguer (1970) from the Eocene of Senegal, has oblong, embryo fatty, straight with flat cotyle-
been excluded from Irvingiaceae. dons; endosperm copious to 0.
Seven spp., six in the humid zone of W and C
U S E S . The seeds of species of all three genera Africa, and one in SE Asia/Malesia.
are eaten by humans, either raw or cooked. The
juicy mesocarp of Irvingia gabonensis is eaten as
a dessert fruit; the tree is often planted (Harris 3. Desbordesia Pierre ex van Tiegh.
1996). Desbordesia Pierre ex van Tiegh., Ann. Sci. Nat. IX, 1: 289
(1905); Harris, Bull. Jard. Bot. Nat. Belg. 65: 147–151
(1996), rev.
KEY TO THE GENERA
1. Ovary 5-celled; fruit a drupe with 5 pyrenes; Tall trees. Leaves matt on undersurface. Inflores-
stipules, covering terminal buds, 3.5–12(25) cm long cences usually much branched. Flowers inserted
1. Klainedoxa on short lateral branches of the panicle
232 K. Kubitzki
(occasionally lateral branches fasciculate); ovary Korotkova, N. et al. 2009. See General References.
flattened, 2-locular; ovules 1 per locule. Fruit a Link, D. 1992. The floral nectaries in the Irvingiaceae. Pl.
Syst. Evol. 180: 235–242.
samara with (1)2 fatty seeds; endosperm scanty. Matthews, M.L., Endress, P.K. 2011. See General References.
A single sp., D. glaucescens (Engler) van Netolitzky, F. 1926. See General References.
Tiegh., confined to the lower Guinean block of Nooteboom, H.P. 1967. The taxonomic position of Irvin-
the Central African forest from Nigeria to Zaire. gioideae, Allantospermum Forman, and Cyrillopsis
Kuhlm. Adansonia II, 7: 161–168.
Rojo, J.P. 1968. The wood anatomy of Allantospermum
borneense Forman and Allantospermum multicaule
Selected Bibliography (Capuron) Nooteboom. Adansonia II, 8: 73–83.
Ruhfel, B.R. et al. 2011. See General References.
Sabatier, B. 1974. Contribution de la palynologie à l’étude
Fernando, E.S., Quinn, C.J. 1992. Pericarp anatomy and des Irvingiacées d’Afrique tropicale. Adansonia II,
systematics of the Simaroubaceae sensu lato. Austral. 14: 277–289.
J. Bot. 40: 263–289. Soltis, D.E. et al. 2011. See General References.
Forman, L.L. 1965. A new genus of Ixonanthaceae with Tobe, H., Raven, P.H. 2011. Embryology of the Irvingia-
notes on the family. Kew Bull. 19: 517–526. ceae, a family with uncertain relationships among
Gros, J.-P. 1983. Irvingiaceoxylon dechampsii n.g. et n.sp. the Malpighiales. J. Plant Res. 124: 577–591.
du Cénozoique d’Éthiopie, et Simaroubaceoxylon Tokuoka, T., Tobe, H. 2006. See General References.
(Irvingioxylon) taibaense n.g. et n.comb. de l’Ypré- Verdcourt, B. 1984. Ixonanthaceae. In: Polhill, R.M. (ed.)
sien du Sénégal. Rev. gén. Bot. 90: 153–171. Flora of Tropical East Africa. 9 pp. Rotterdam: Balkema.
Hallier, H. 1921. Beit€age zur Kenntnis der Linaceae van Welzen, P.C., Baas, P. 1984. A leaf anatomical contri-
(DC. 1819) Dumort. Beih. Bot. Centralbl. 39, II: bution to the classification of the Linaceae complex.
1–178. Blumea 29: 453–479.
Hallier, H. 1923. See General References. Winkler, H. 1931. Linaceae. In: Engler & Prantl, Nat.
Harris, D.J. 1996. A revision of the Irvingiaceae in Africa. Pflanzenfam., ed. 2, 19a. Leipzig: W. Engelmann.
Bull. Jard. Bot. Nat. Belg. 65: 143–196. Wurdack, K.J., Davis, C.C. 2009. See General References.
Hutchinson, J. 1973. The families of flowering plants. 3rd Xi, Z. et al. 2012. See General References.
edn. Oxford: Clarendon Press.
Koeniguer, J.-C. 1970. Sur deux bois fossiles de l’Éocène
du Sénégal. 95ème Congr. Soc. Sav., Reims 1970, Sci.,
3: 39–77.
Ixonanthaceae
Ixonanthaceae Planch. ex Miquel (1858), nom. cons.
Ixonantheae Benth. (1862).
K. K U B I T Z K I
Trees or shrubs. Leaves spiral, usually involute, arranged platelets. Sieve-element plastids are S-
entire or glandular-serrate; stipules small, lateral type.
or rarely intrapetiolar. Inflorescences lateral, cor- Vessels have simple perforations. Allantos-
ymbose. Flowers small, mostly hermaphroditic, permum differs from both Ixonanthaceae and
regular or nearly so, commonly 5-merous; sepals Irvingiaceae in its heterogeneous rays and min-
connate at the base or free, imbricate; petals free, ute, half-bordered vessel-ray pits (Rojo 1968).
imbricate or contorted; stamens 5, 10 or 20; fila- Extrafloral epithelial nectaries were described
ments folded in bud, widened at the base, free or from the ventral side of the sepals of Ixonanthes
basally adnate to the conspicuous annular or icosandra (Narayana and Rao 1966; Link 1992)
cupular nectary disk (lacking in neotropical and from the leaf margins of the same species
Ochthocosmus); gynoecium 5(2)-carpellate; style (Belin-Depoux 1978), where they are functional
slender; stigma capitate or discoid; ovary some- in juvenile stages of leaf development.
times apically unilocular and sometimes the
locules divided into locelli by incomplete second- F L O W E R S T R U C T U R E . Articulate pedicels with
ary septa; style and filaments folded in bud; prophylls are reported from Cyrillopsis. Narayana
ovules 2 or rarely 1 per locule, apical. Fruits and Rao (1966) and Link (1992) studied the
septicidal capsules, sometimes also loculicidal prominent intrastaminal disk of Ochthocosmus
by secondary septa; columella persistent or not; africanus and Ixonanthes; the margin of the latter
seeds with obvious basal wing or suprahilar aril- was found to bear 20–25 nectarial stomata over
lode; embryo straight with large cotyledons; stomatal cavities. In Ochthocosmus barrae, a disk
endosperm scanty or 0. is lacking but the filament bases are connected by
A family comprising four genera including a glandular staminal tube secreting nectar
the controversial Allantospermum and about 23 through the numerous open nectarial stomata.
spp. from tropical America, tropical Africa, the
Himalayas and NE India, SE Asia and Malesia to P O L L E N M O R P H O L O G Y . Pollen is (spheroidal-)
New Guinea. subprolate-prolate, tricolporate and 27–52 mm
long. The tectum and nexine are thin, and the
VEGETATIVE MORPHOLOGY AND ANATOMY. Ixo- infratectal layer is columellate. The sculpture is
nanthaceae differ from Irvingiaceae in the usually scabrate, in Cyrillopsis rugulate or striate.
absence of mucilage cells and mucilage cavities The pollen of Ixonanthes is spheroidal, large (up
and the occasional presence of mesophyll scler- to 45 mm diameter) and has distinct supratectal
eids (Ochthocosmus) and tracheoidal idioblasts; spines (Metcalfe et al. 1968).
cristarque cells, which are present in both the
ground tissue and the bundle sheath, are found F R U I T A N D S E E D . The seeds of Ixonanthes are
in Ochthocosmus and, more prominently, in exotegmic (Corner 1976; Boesewinkel and Bouman
Irvingiaceae and Linaceae-Hugonioideae. Sto- 2000). The American and African species of
mata are paracytic, as usual in the Linaceae com- Ochthocosmus (O. s.str versus ‘Phyllocosmus’) dif-
plex; petiole bundles are arcuate (van Welzen and fer in seed structure: the former have seeds with an
Baas 1984). Cuticle waxes are present as variously apical wing pointing upwards in fruit and the latter
234 K. Kubitzki
(funicular?) aril attached to the seed between the

micropyle and the hilum, whereas the two other
species of the genus (sectionally different) have
seed wings that are said to be inserted at the
chalazal end and vascularized, and having a texture
like the rest of the testa (Corner 1976; Kool 1988).
P H Y T O C H E M I S T R Y . Seed oils contain predomi-

nantly lauric and myristic acids, in which they
differ from those in Simaroubaceae. In hydroly-
sates of Allantospermum, Nooteboom (1967)
found ellagic and gallic acid, which are lacking
from the other three genera.
C L A S S I F I C A T I O N . Ochthocosmus and Phyllocos-

mus have sometimes been kept separate (see
Forman 1965) but, as Hallier (1923) and Kool
(1988) have explained, if this course is to be
followed, then a third genus must be distinguished.
Ochthocosmus, Ixonanthes and Cyrillopsis are
clearly related but the position of Allantospermum
is problematic because the intrapetiolar stipules, the
caducous petals and the solitary ovules seem to fit
better with Irvingiaceae than with Ixonanthaceae,
where Cleistanthopsis (a synonym of Allantosper-
mum) originally had been placed. Forman (1965)
considered the central columella of Allantosper-
mum as equivalent to five splinters into which the
central tissue of the fruit in Ixonanthaceae separates
and which remain attached to the edges of the
valves. After considering all characters, however,
Forman was inclined to place the genus into Ixo-
nanthaceae. A different interpretation of the fruit
Fig. 52. Ixonanthaceae. Ochthocosmus longipedicellatus. structure of Allantospermum led Nooteboom (1967)
A Flowering and fruiting branchlet. B Mature capsule. C, to keep Allantospermum in Irvingiaceae. Wood
D Flowers during progressive stages of anthesis. O. ror- anatomically, Allantospermum differs from both
aimae var. parvifolius. E Winged seed. (Reproduced with
kind permission of the artist Bobbi Angell) Irvingiaceae and Ixonanthaceae (Rojo 1968).
A F F I N I T I E S . Forman (1965), Corner (1976) and

an apically attached aril(lode) that is turned down others combined Irvingiaceae and Ixonanthaceae
and clasps the upper part of the seed. Both the in spite of their morphological differences, and
wings and arillodes are anatomically very similar; Takhtajan (2009) placed Ixonanthaceae in the
they develop from the micropylar region and Linales, together with, i.a., Erythroxylaceae/Rhi-
appear homologous (Kool 1988), just following zophoraceae and Ctenolophonaceae. Molecular
the reasoning of Corner in his durian theory datasets have led to a confusing picture. Initially,
(1954) "that all winged seeds are prima facie indi- Ochthocosmus and Ixonanthes remained
cations of arillate ancestry". (See also the com- unplaced or in vacillating positions within the
ments on winged vs. arillate seeds in Malpighiales. Wurdack and Davis (2009)
Rhizophoraceae under that family, this vol.) In provided support for a clade of (Ixonanthes
Ixonanthes one species, I. icosandra Jack, has a (Ochthocosmus + Cyrillopsis)) but left it unplaced
Ixonanthaceae 235
D I S T R I B U T I O N A N D H A B I T A T S . The family is
bound to the tropical lowland forests including
heath forests and swamp forests of Africa and
Asia/Malesia, and in South America mainly
occurs in upland white sand savannahs of the
Roraima formation in southern Venezuela.
Ixonanthes, today considered a typical Malesian

element, appeared in the fossil pollen record of
India in the Palaeocene, along with other origi-
nally African elements such as Durio and Gonys-
tylus (Kar 1985). These are thought as having
rafted from Africa on the Indian Plate and having
dispersed eastwards with subsequent range
reduction in Africa and India.
KEY TO THE GENERA

1. Capsule valves splitting away from a central column;
ovules 1 per cell; stamen filaments distinct from disk;
stipules intrapetiolar; stamens 10 4. Allantospermum
– Capsule without a central column; ovules 2 per cell;
stamen filaments attached at the base to the disk; sti-
pules lateral; stamens 5–20 2
2. Ovary 2-celled; capsule 2-valved; pedicel articulated
and provided with prophylls; stamens 5
3. Cyrillopsis
– Ovary 5(or spuriously 10)-celled; capsule 5-valved;
pedicel not articulated; prophylls 0; stamens 5–20 3
3. Trees of SE Asia and Malesia; stamens 10, 15 or 20;
(sepals with extrafloral nectaries) 1. Ixonanthes
– Trees and shrubs of N South America and tropical
Africa; stamens 5 or 10 2. Ochthocosmus
Fig. 53. Ixonanthaceae. Allantospermum borneense. A

Leaf. B Leaf-bud with stipules (the lower scars are those 1. Ixonanthes Jack
of bud-scales). C Flowering branch. D Flower. E Disk with Ixonanthes Jack, Mal. Misc. 2, 7: 51 (1822); Kool, Blumea
base of filaments (central one removed). F Pistil. G Verti- 26: 191–204 (1988).
cal section of ovary. H Capsule. I Valve of capsule. J
Columella of capsule bearing arilloid processes. K Seed. Evergreen, buttressed trees or treelets. Inflores-
L Embryo. (Forman 1965) cences pedunculate, pleiochasial-corymbose.
Flowers pedicellate; sepals and petals quincun-
within Malpighiales. Ruhfel et al. (2011) reported cial, the sepals with extrafloral nectaries on the
a relationship between Cyrillopsis and Bruguiera ventral side; stamens 10 or (15–)20, apparently in
(Rhizophoraceae), which in turn are sister to one whorl, filaments inserted outside and against
Irvingia (Irvingiaceae). The 7-gene analysis of the disk; anthers (basi-)dorso-versatile; disk
Soltis et al. (2011) resolved Ochthocosmus in an annular or cup-shaped; ovary 5-locular; ovules
unsupported clade with Irvingiaceae as part of 2 per locule, collateral, pendulous; style simple;
the clusioid clade. Strong support for the family stigma discoid. Capsule septicidal and septifragal,
pair Linaceae and Ixonanthaceae was provided by 5-valvate. Seeds 1 or 2 per locule, either with a
Xi et al. (2012), which makes sense in view of their basal wing or a supra-hilar arillode; endosperm
common possession of mostly diplostemonous oily; embryo straight.
flowers, a contorted corolla, carpels with false Three spp., SE Asia and Malesia but lacking
septa, a placental obturator, etc. from Java, the lesser Sunda Islands and the
236 K. Kubitzki
Moluccas. The arillode of I. icosandra Jack (sect. late; sepals and petals 5, imbricate, caducous;
Brewstera) is attached between the hilum and the stamens 10; disk intrastaminal; ovary 5-locular,
micropyle, whereas the remaining two species with 1 pendulous ovule per locule; style simple;
have chalazal wings. stigma capitellate. Capsule 5-valvate, septicidal,
with persistent columella bearing near apex 5
arilloid processes from which the seeds break
2. Ochthocosmus Benth. Fig. 52
away. Seeds cylindric-ellipsoid, up to 2.5 cm
Ochthocosmus Benth. in Hook., London J. Bot. 2: 266 long, shining and waxy, distally with a arillode
(1843); Robson, Fl. Zambes. 2, 1: 100–102; Badré in Fl.
Cameroun 14: 57–63 (1972) and Fl. Gabon 21: 55–62
sometimes adherent to the columella.
(1963); Steyermark & Luteyn, Brittonia 32: 128–143 Two spp., one in Sarawak, the other in Mada-
(1980), rev. neotr. spp. gascar.
Phyllocosmus Klotzsch (1857).
Trees. Inflorescences in racemes. Flowers pedicel-
late or sessile; sepals lacking extrafloral
nectary; petals imbricate or contorted; stamens Belin-Depoux, M. 1978. Contribution à l’étude des glan-
des foliaires d’Ixonanthes isosandra Jack (Ixo-
5 or 10; anthers dorsifixed; ovary 5-locular; nanthaceae). Rev. gén. Bot. 85: 371–382.
locules with 2 ovules, sometimes with incomplete Boesewinkel, F.D., Bouman, F. 2000. Ixonanthaceae. In:
false septum; style simple, stigmas 5, free or Takhtajan, A. (ed.) Anatomia seminum comparativa,
vol. 6. Nauka: St. Petersburg. (In Russian).
fused. Capsule septicidal, 1–3(–5)-seeded. Seeds Corner, E.J.H. 1954. The durian theory extended. 2. Phy-
obliquely oblong, either with a conspicuous thin tomorphology 4: 152–165.
wing or an arillode attached to the distal end. Corner, E.J.H. 1976. See General References.
About 16 spp., 7 of them in northern South Forman, L.L. 1965. A new genus of Ixonanthaceae with
notes on the family. Kew Bull. 19: 517–526.
America and 9 in tropical Africa. Three sections Hallier, H. 1923. See General References.
were distinguished by Hallier (1921); see also Kar, R.K. 1985. The fossil floras of Kachchh - IV. Tertiary
Kool (1988). palynostratigraphy. Palaeobotanist 34: 1–279.
Kool, R. 1988. A taxonomic revision of the genus Ixo-
nanthes (Linaceae). Blumea 26: 191–204.
3. Cyrillopsis Kuhlm. Link, D.A. 1992. The floral nectaries of the Geraniales and
their systematic implications: VI. Ixonanthaceae
Cyrillopsis Kuhlm., Arch. Jard. Bot. Rio de Janeiro 4: 356, Exell and Mendonça. Bot. Jahrb. Syst. 114: 81–90.
t. 29 (1925); Ramı́rez & Berry, Fl. Venez. Guayana 5: Metcalfe, C.R., Lescot, M., Lobreau, D. 1968. A propos de
665–666 (1999). quelques caractères anatomiques et palynologiques
comparés d’Allantospermum borneensse Forman et
Small trees. Stipules very small, fugacious. Inflo- d’Allantospermum multicaule (Capuron) Noote-
rescence a fascicle of racemes. Pedicels articulate, boom. Adansonia II, 8: 337–351.
Narayana, L.L., Rao, D. 1966. Floral morphology of Lina-
with prophylls beneath articulation. Flowers pen- ceae. J. Jap. Bot. 41: 1–10.
tamerous; sepals and petals imbricate, persistent; Nooteboom, H.P. 1967. The taxonomic position of Irvin-
stamens 5, alternipetalous; anthers basifixed, gioideae, Allantospermum Forman, and Cyrillopsis
introrse; ovary 2-locular; ovules 2 per locule, Kuhlm. Adansonia II, 7: 161–168.
Ramı́rez, N., Berry, P.E. 1999. Ixonanthaceae, pp.
pendent from apex of axile placenta; style fili- 665–666. In: Flora of the Venezuelan Guayana, vol.
form. Capsule 2-valved. Seeds with a distal flat 5. St. Louis: Missouri Bot. Gard. Press.
triangular arillode. Robson, N.K.B., Airy Shaw, H.K. 1962. A note on the
Two spp., northern South America, in wood- taxonomic position of the genus Cyrillopsis Kuhl-
mann. Kew Bull. 15: 387–388.
land on white sand and in altitudinal savannahs. Rojo, J.P. 1968. The wood anatomy of Allantospermum
borneense Forman and Allantospermum multicaule
(Capuron) Nooteboom. Adansonia II, 8: 73–83.
4. Allantospermum Forman Fig. 53 Ruhfel, B.R. et al. 2011. See General References.
Allantospermum Forman, Kew Bull. 19: 517 (1965); Noo- Soltis, D.E. et al. 2011. See General References.
Takhtajan, A. 2009. See General References.
teboom, Adansonia II, 7: 161–168 (1967).
van Welzen, P.C., Baas, P. 1984. A leaf anatomical contri-
Cleistanthopsis R. Capuron (1965). bution to the classification of the Linaceae complex.
Blumea 29: 453–479.
Buttressed or polycormic trees. Leaves entire; sti- Wurdack, K.J., Davis, C.C. 2009. See General References.
pules intrapetiolar. Inflorescences thyrso-panicu- Xi, Z. et al. 2012. See General References.
Linaceae
Linaceae DC. ex Perleb, Vers. Arzneikr. Pfl.: 107 (1818), nom. cons.
Hugoniaceae Arn. (1834).
S. D R E S S L E R , M. R E P P L I N G E R , AND C. B A Y E R
Herbs, shrubs, trees, or lianas, sometimes with in Linoideae absent or sometimes reduced to
climbing hooks. Leaves alternate (more rarely glands. In contrast to Ctenolophonaceae, tufted
opposite or whorled), sometimes distichous, sim- hairs are absent.
ple, sessile or petiolate, the lamina usually invo-
lute; stipules present or reduced, sometimes
dentate or incised. Inflorescences terminal or
axillary thyrsoids or botryoids, exceptionally
flowers solitary. Flowers perfect, actinomorphic,
5(4)-merous; sepals quincuncially imbricate,
distinct or connate at the very base, sometimes
persistent; petals usually contorted, often clawed,
distinct or almost so, usually caducous; stamens
usually twice the number of petals or antesepa-
lous stamens and staminodia [these sometimes
reduced] as many as petals, connate at base, usu-
ally with glands outside the tube; ovary superior,
(2)3–5(–8)-carpellate; ovules 2 per carpel, some-
times more, anatropous, epitropous; locules
sometimes divided into 2 one-seeded portions
by complete or incomplete false septae; stylodia
as many as carpels, usually distinct; stigma capi-
tate to filiform. Fruit a septicidal capsule, some-
times schizocarpic or a drupe. Seeds sometimes
arillate; seed coat often mucilaginous; endosperm
usually scanty, sometimes copious; embryo
straight or slightly curved.
A family of 13 genera with ca. 255 species,
mostly from northern temperate to tropical
regions.
V E G E T A T I V E M O R P H O L O G Y . The Old World

Hugonioideae are woody climbers with hooks
that correspond to modified shoots. Such hooks
are often found as lowermost branches of inflor-
Fig. 54. Linaceae. Hugonia villosa. A Flowering branch-
escences. Rhizomes are unknown in Linaceae; let. B Detail of lower leaf surface. C Stipule, adaxial side.
perennial herbs form persistent primary roots D Flower bud. E Petal. F Androecium and gynoecium.
(Troll and Weberling 1989). Cliococca has an G Gynoecium. H Vertical section of ovary. I Part of
extensively branched underground rootstock. fruiting branch. J Fruit. K Pyrene. L Same, transverse
section. M Seed. (Badré 1973, drawn by Hélène Lamour-
Stipules are sometimes pectinate (Hugonioideae), dedieu)
238 S. Dressler et al.
V E G E T A T I V E A N A T O M Y . Winkler (1931) and and Nooteboom 1984). Sieve element plastids are
Metcalfe and Chalk (1950) provide anatomical of the Ss-type.
overviews of the family. Leaf anatomy was stud-
ied in detail by van Welzen and Baas (1984). R E P R O D U C T I V E S T R U C T U R E S . Although little is
The indumentum usually consists of narrow known on the woody Hugonioideae, it is likely
and unicellular trichomes, in Hesperolinon com- that the family has basically monotelic synflores-
bined with glandular hairs with multiseriate stalk cences (see Troll and Weberling 1989). Terminal
(van Welzen and Baas 1984). The leaves are flowers are easy to detect in herbaceous Linaceae,
generally dorsiventral and hypostomatic. Espe- and the basic structure appears to be a thyrsoid.
cially herbaceous members (e.g. Linum, Radiola, Further branching of the inflorescence is restricted
Hesperolinon) are sometimes amphi- or even to the distal portion (e.g. Linum spp., but see also
epistomatic. Stomata are of the paracytic type. Anisadenia; Troll and Weberling 1989). The para-
(Sub)epidermal cells are often mucilaginous (all cladia can be restricted to a single flower with both
Linoideae and most Hugonioideae); their absence prophyllar nodes, which may lead to cymose
is reported for Hebepetalum neblinae (Jardim and branching (e.g. dichasia in Radiola; Troll 1964),
Berry 1999) and three Roucheria spp. studied by or produce more nodes, leading to thyrso-panicu-
van Welzen and Baas (1984), whereas Metcalfe late systems. Reduced ramification leads to deter-
and Chalk (1950) mentioned them for the latter minate raceme-like inflorescences (botryoids, e.g.
genus. Hugonioideae are well defined by regular Anisadenia), cincinnate sympodia (e.g. Linum
sinuously lobed subsidiary cells and lignified austriacum) or single-flowered inflorescences
guard cells (missing in Indorouchera). Mesophyll (some Linum spp.; Troll and Weberling 1989).
contains idioblasts in Hugonia (Metcalfe and The bases of stamens and, where present,
Chalk 1950, not confirmed by van Welzen and staminodia are connate to form a short tube.
Baas 1984) and Roucheria (van Welzen and Baas Nonetheless, the androecium is originally diplos-
1984). Crystals are present as druses or solitary temonous or obdiplostemonous (Str€obl 1925;
ones (rarely clustered) in woody members of the Narayana and Rao 1978). The exterior side of the
family only. Hugonioideae are well defined by the tube often bears distinct glands, which in some
occurrence of cristarque cells in the ground tissue Hugonioideae may form a continuous disk (Nar-
of petiole and midrib (van Welzen and Baas 1984). ayana and Rao 1973). In some Linoideae, small
Stems of Linum have large, well-defined bast nectary glands external to the stamens or at the
fibres in the pericycle or phloem (see Economic inner base of the petals are reported to exist.
Importance), where they sometimes form a con- Floral anatomy of several Linaceae has been
tinuous ring, but these are absent from Reinward- studied by Narayana and Rao (1978, summarizing
tia trigyna (Metcalfe and Chalk 1950). The pith and discussing numerous previous works). Mat-
disintegrates in Linum. thews and Endress (2011) described floral mor-
The wood is characterised by mostly solitary phology and anatomy for several Linaceae, and
vessels (but a few radial multiples of 2–3 occur in reported gynophores in Linum and Reinwardtia.
Hugonia or Linum), scalariform to simple or only Some species of Hugonia, Linum, Tirpitzia
simple perforations, predominantly apotracheal and Reinwardtia are heterostylous; dimorphic
parenchyma, multiseriate rays, and fibres with pollen and stigmas are reported for some species
distinctly bordered pits (whole family). Crystals (Heitz et al. 1971; Dulberger 1973, 1974; Bahadur
are reported in Hebepetalum. Linoideae are inter- et al. 1984, 1996; Lloyd et al. 1990; Sugawara et al.
preted as being more advanced than Hugonioi- 2002; see also McDill et al. 2009). Hugonia serrata
deae in having reduced axial parenchyma, is reported to be tristylous (Thompson et al. 1996).
homogeneous rays, short vessel elements and a The stigma is wet or dry with unicellular papillae
tendency towards vessel aggregation but they (Sutter and Endress 1995). In Linum pratense, the
show vestiges of primitive characters such as tra- persistent sepals move inwards after the fall of the
cheidal fibrous elements and vestiges of scalari- petals, leading to self-pollination (Uno 1984).
form vessel perforations in Linum spp. (Heimsch Some Linum have been reported to be polli-
and Tschabold 1972). Indorouchera has the most nated by bees or flies; bee pollination is also
primitive wood with exclusively scalariform known for Hugonia mystax (Heitz 1980; Kearns
perforations and spiral thickenings (van Hooren and Inouye 1994; Aluri et al. 1997).
Linaceae 239
structure of the embryo sac of Linum, see Vazart

and Vazart (1966), Vazart (1969) and D’Alascio
Deschamps (1973, 1981).
P O L L E N M O R P H O L O G Y . Erdtman (1952) consid-

ered Linaceae to be eurypalynous, which was
based on a broad circumscription of the family,
including genera now placed in Humiriaceae,
Erythroxylaceae, Rhizophoraceae, Ixonanthaceae
and Ctenolophonaceae. Palynological characters
agree with the narrow circumscription of Lina-
ceae followed here. Pollen grains are suboblate
to prolate and 3-colpate or -colpor(oid)ate, occa-
sionally with more apertures (Erdtman 1952;
Rogers and Xavier 1971). Anisadenia, Tirpitzia
and Reinwardtia, earlier reported to have inaper-
turate pollen grains (Erdtman 1952), proved to
have pantoporate ones with similar bacula cover-
ing pores and exine alike; Indorouchera, however,
is reported to have inaperturate pollen ("tritenu-
ate"; Saad 1962). Unlike representatives of
the other families mentioned above, Linaceae-
Linoideae have more or less verrucose or spinose
pollen, sometimes with verrucae of different sizes
(Erdtman 1952; Saad 1961; Xavier et al. 1980;
Fig. 55. Linaceae. Hebepetalum humiriifolium. A Branch
with flower buds. B Lower leaf surface showing punctua- Rogers 1985); in Hugonioideae, the exine is bacu-
tions. C Opening flower bud. D Flower. E Petals. F Flower late or tectate(-reticulate) (Saad 1962).
with perianth removed. G Stamens with basal nectaries. Dimorphic pollen is known from heterosty-
H Pistil, with detail of stigmas and cross section of ovary.
I Part of infructescence. J Fruit. (Reproduced with the
lous species of Linum (Rogers 1980; Dulberger
kind permission of the artist Bobbi Angell) 1981; Grigorieva 1989) and Reinwardtia (Suga-
wara et al. 2002).
E M B R Y O L O G Y . The anther wall is 5-layered with K A R Y O L O G Y . Diploid chromosome numbers

an innermost layer of secretory tapetum of have been reported for Hugonia as 2n ¼ 12, 24,
binucleate cells. Pollen is 3-cellular when shed 26, for Radiola linoides 18, for Reinwardtia trigyna
(Narayana 1964). Most embryological informa- 20, 22, for Cliococca selaginoides 36 (Bolkhovskikh
tion refers to Linum usitatissimum and few et al. 1969; Robertson 1971), for Hesperolinon 34,
other species (for Linum, Hugonia and Reinward- 36 (Rogers et al. 1972) and for Sclerolinon as 12
tia, see especially Narayana 1964). Ovules are (Rogers et al. 1972) or 16 (Raven 1959).
bitegmic, crassinucellar to tenuinucellar, and epi- Within the diverse genus Linum, diploid
tropous. The endothelium is one- or two-layered chromosome numbers range mostly between 16
(Sutter and Endress 1995); its ultrastructure was and 60 (Rogers et al. 1972; Diederichsen and
described by Vazart and Vazart (1965). The Richards 2003; Muravenko et al. 2010). There
micropyle is formed by the inner integument; an are, however, also reports of counts of 2n ¼ 12
obturator is often present (Narayana 1964; Mat- (L. hypericifolium), 68 (L. bahamense), 72
thews and Endress 2011; absent in Radiola: Crété (L. suffruticosum) and 84, 86? (L. monogynum;
1937). Embyro sac development conforms to the all from Bolkhovskikh et al. 1969). Basic numbers
Polygonum type. One of the synergids degener- of n ¼ 9 and 15 predominate (Chennaveeraiah
ates (Vazart 1971). The endosperm develops and Joshi 1983).
according to the Nuclear type; embryogeny
follows the Solanad type (Souèges 1937; Narayana F R U I T A N D S E E D . Drupaceous fruits are found
1964; Erdelská 1967; Johri et al. 1992). For ultra- in Hugonioideae; in some species, the fruits
split into indehiscent mericarps. Most Linoideae

have capsular fruits; each carpel is usually
divided into two one-seeded portions. Capsules
in Linum usitatissimum are described in detail by
Roth (1977).
The seed coat is formed by both integuments
with thick-walled cells in the outer epidermis of the
inner integument (Narayana 1964). Structure and
development of seeds have been studied in detail
in Linum (Boesewinkel 1980). The mature seed
coat consists of five layers: an innermost, pigmen-
ted layer derived from the inner cells of the inner
integument; a layer of compressed cells; a scler-
enchymatic layer derived from the outer layer of
the inner integument; the cells of the inner layer of
the outer integument; and the outermost layer of
mucilage cells. Upon soaking, the mucilage swells,
ruptures the cell walls of the epidermis and pene-
trates through lesions in the cuticle. For produc-
tion and composition of linseed mucilage, see
Heinze and Amelunxen (1984). In contrast to
Linum and Reinwardtia, the seeds of Radiola
lack mucilage in the outer epidermis (Crété 1937).
Roucheria and Hugonia with hard endocarp
in their drupaceous fruits seem to deviate from
the exotegmic state of the family by modification
of this layer from slightly lignified to mostly
unlignified (Corner 1976). Instead, there are
abundant sclerotic cells. Furthermore, no muci-
lage is formed.
The straight to slightly curved embryo with
flat cotyledons is embedded in a nuclear, oily
endosperm.
P H Y T O C H E M I S T R Y . Knowledge on the phyto-

chemistry of the family is limited. Known pheno-
lic compounds include C-glycosyl flavones (only
in Linum usitatissimum), lignans, and cinnamic
acid derivatives. Roucheria griffithiana contains a
saponin; pyrrolizidin alkaloids are known from
Hugonia; Linum spp. and Reinwardtia are cyano-
genic, containing linamarin and lotaustralin
monoglucosides. Mucilage of Linum usitatissi-
mum is restricted to the seed coat epidermis of
mature seeds; it consists of a mixture of neutral Fig. 56. Linaceae. A–C Linum grandiflorum. A Plant with
and acid heteropolysaccharides mixed with pep- flowers and young fruit. B Stamens and pistil. C Pistil. D–G
tides (Hegnauer 1966, 1989). Linum usitatissimum. D Plant with flowers and a capsule.
E Dehiscing capsule. F, G Different growth forms of
cultivated flax. H Tirpitzia sinensis, flower. I Anisadenia
S U B D I V I S I O N A N D A F F I N I T I E S . Linaceae can saxatilis, flowering branch with rhizome. (Takhtajan 1980)
readily be subdivided into subfamilies Hugonioi-
deae (trees, climbers or shrubs, stamens twice the
Linaceae 241
number of petals, staminodia 0, fruit drupeacous) E C O N O M I C I M P O R T A N C E . Linum usitatissimum

and Linoideae (herbaceous to shrubby, stamens and has been under cultivation for more than 7,000
staminodia as many as petals, fruit capsular or years in Europe, the Near East and western Asia
schizocarpic). Linoideae are poorly resolved so far. (Pengilly 2003). Today, special varieties are
Recent molecular studies conducted by McDill and grown for their oil seeds or their excellent fibre
colleagues (McDill et al. 2009; McDill and Simpson (Marchenkov et al. 2003). Linseed (in American
2011) indicate that Linum is paraphyletic in relation English flaxseed) oil is used for paints, varnishes
to Radiola, Hesperolinon, Cliococca and Sclerolinon. and coatings, as well as an edible condiment. The
At present five sections are recognized within seed itself provides a dietary complementary
Linum: Linum, Dasylinum, Syllinum, Linopsis and food, its mucilage facilitating digestion and
Cathartolinum, with Linum and Dasylinum being excretion. Several linseed compounds (e.g.
sister to the remaining sections/genera. Hesperoli- a-linolenic acid, lignans) have proven to have
non, Cliococca and Sclerolinon are nested within a positive pharmacological effects on cardiovascu-
paraphyletic sect. Linopsis. This clade is sister to the lar diseases and cancer (Cunnane 2003; Prasad
monotypic sect. Cathartolinum, being again sister 2003; Rickard-Bon and Thompson 2003). Flax
to the monophyletic sect. Syllinum. Radiola is sister fibre is flexible, strong, durable and absorbs
to all this. Taxonomic consequences, however, have water, making it very useful for towels and
not yet been addressed. table cloths. Usually the fibres are 2–4 cm long,
Some genera formerly thought to be related to in rare instances they may reach up to 12 cm.
Linaceae had to be excluded and are now referred Some other Linum species are cultivated as orna-
to Erythroxylaceae and Ixonanthaceae (van Hoo- mentals.
ren and Nooteboom 1984; APG II 2003; Davis and
Chase 2004). Linaceae belong to Malpighiales KEY TO THE GENERA
(eurosid I clade sensu APG II 2003). Within Mal-
1. Trees or lianas with hooks, rarely shrubs, all ligneous;
pighiales, Linaceae are resolved in an unsupported petals not or hardly clawed; stamens twice as many as
sister position with the chrysobalanoid clade petals; ovary (1–)3–5-celled; fruit a drupe, rarely
(Davis and Chase 2004; Soltis et al. 2011), whereas splitting finally into indehiscent mericarps (subfam.
the analysis of Xi et al. (2012) places Linaceae in a Hugonioideae) 2
well-supported clade with Ixonanthaceae, this lat- – Erect herbs or small shrubs; petals usually long-
ter clade being sister to the phyllanthoids. clawed; stamens as many as petals, alternating with
the same number of staminodes; ovary 6–10-celled (4
in Sclerolinon); fruit usually a capsule (subfam.
D I S T R I B U T I O N A N D H A B I T A T S . Hugonioideae Linoideae) 6
occur in rain forests or periodically inundated 2. Trees or shrubs of the Neotropics 3
savannahs in the tropical belt of northern South – Lianas with hooks, rarely shrubs (New Caledonia) of
America, Africa, southern India and Papuasia. The the Old World tropics 4
neotropical representatives are trees or shrubs, 3. Secondary veins arching near the margin (brochido-
whereas lianas or more rarely shrubs predominate dromous) and reticulately veined in between; petals
in the palaeotropics. The range of the mostly herba- villous on inner surface and basally clawed; stylodia
usually 5 5. Hebepetalum
ceous Linoideae extends into the temperate zones
– Secondary veins fine and closely parallel, running into
of both hemispheres, except for the large deserts. a marginal vein, no reticulate veins in between; petals
Many species are typical of steppe vegetation. glabrous on inner surface; stylodia 3
4. Roucheria
P A L A E O B O T A N Y . Megafossils formerly assigned 4. Indumentum present, at least on the calyx; stylodia 5(6);
to this family (Wetherellia, Decaplatyspermum) ovary 5(6)-celled, locules all distinct in the fruit;
proved to belong elsewhere (Mazer and Tiffney drupe with 2–4(5) developed seeds 1. Hugonia
1982; Collinson et al. 1993). Pollen of the Linum – All parts glabrous; stylodia 3–4(5); ovary 3(4)-celled,
only one, rarely two locules distinct in fruit; drupe
type was found from the upper Miocene of Spain usually one-seeded 5
and the Pliocene of Germany (Muller 1981). 5. Buds and stipules often covered with resin; leaves
Boesewinkel (1984) compared recent with fossil distichous; flowers in axillary fascicles, subtended by
linseed samples, which are frequently found in often densely packed, imbricate bracts
archaeological remains. 3. Indorouchera
– Resin absent; leaves spiral; flowers in rather lax somewhat unequal, distinct; disk + or 0; ovary
racemes or panicles 2. Philbornea 5(6)-locular, stylodia 5(6); drupe (pseudo-)
6. Leaves petiolate 7 indehiscent, stone 5(6)-locular with as many
– Leaves sessile 9 interlocular sterile cavities (sect. Hugonia) or
7. Low shrubs or subshrubs; sepals without stalked splitting into 5 mericarps (sect. Durandea). Seeds
glands; fruit a 6–10-seeded capsule 8 1 per locule, exarillate, but with woody chalaza;
– Perennial herbs; sepals with conspicuous long-stalked embryo straight; endosperm fleshy. 2n ¼ 12,
glands; fruit indehiscent, 1-seeded nut 24, 26.
6. Anisadenia
About 40 spp. from Africa, Madagascar, tro-
8. Flowers white or pink; capsule splitting into 4–5 two-
seeded valves 7. Tirpitzia
pical Asia eastwards to Solomon Isl., Queensland,
– Flowers yellow; capsule splitting into 6–8 one-seeded New Caledonia and Fiji. Hugonia including
valves 8. Reinwardtia Durandea is paraphyletic with respect to Indo-
9. Locules, stylodia and stigmas 2 or 3 10 rouchera and Philbornea (McDill and Simpson
– Locules, stylodia and stigmas 4 or 5 11 2011), but taxonomic consequences remain to
10. Leaves alternate; stipular glands present (sometimes be addressed. Resurrection of the generic status
minute); petal base with appendages; stigmas as wide of Durandea might be the appropriate action.
as stylodia; stylodia 2 or 3 12. Hesperolinon
– Leaves opposite; stipules 0; petals without well-
defined appendages; stylodia wider than stigma; sty- 2. Philbornea Hallier f.
lodia and stigmas 2 11. Sclerolinon Philbornea Hallier f., Arch. Néerl. Sci. Exact. Nat. III, 1:
11. Petals and stylodia 4 12 110 (1912); van Hooren & Nooteboom, Fl. Males. 10(3):
– Petals and stylodia 5 13 614–615 (1988).
12. Flowers small, white 10. Radiola
– Flowers yellow (Linum keniense) 9. Linum Liana with hooks, glabrous. Leaves spiral; stipules
13. Flowers few, terminal on densely leaved branches; entire. Inflorescences axillary or terminal,
petals imbricate, shorter than sepals, whitish or pale raceme-like [probably botryoids], bracts and
pink 13. Cliococca prophylls present; sepals somewhat unequal, con-
– Inflorescense cymose; petals contorted, usually larger nate at the very base; exterior side of connate
than sepals 9. Linum
filament bases glandular; ovary 3-locular, stylodia
3; drupe little fleshy, 1-seeded. Seeds arillate;
embryo straight; endosperm fleshy, scarce.
GENERA OF LINACEAE One sp., P. magnifolia (Stapf) Hallier f.,
Sumatra, Borneo, Palawan.
I. SUBFAM. HUGONIOIDEAE Planch. ex Hooren &
Noot. (1984). 3. Indorouchera Hallier f.
Indorouchera Hallier f., Beih. Bot. Centralbl. 39(2): 50
Trees or lianas, rarely shrubs; leaves alternate; (1921); van Hooren & Nooteboom, Fl. Males. 10(3):
petals without claw; stamens twice the number 615–619 (1988), rev.
of sepals; fruit drupaceous, rarely splitting into
indehiscent mericarps; pantropical. Lianas with hooks, glabrous, young leaves and
buds often covered with resin. Leaves distichous;
stipules entire or toothed. Flowers in dense axil-
1. Hugonia L. Fig. 54 lary clusters or solitary, pedicel with numerous
Hugonia L., Sp. Pl.: 675 (1753); Badré, Adansonia II, 11: bracts; nectaries 0; ovary 3(4)-locular; stylodia
95–106 (1971), reg. rev.; van Hooren & Nooteboom, Fl. 3–5, connate at base or distinct; stigmas discoid.
Males. 10(3): 609–613 (1988), reg. rev. Drupe 1(2)-seeded. Seeds arillate; endosperm
Durandea Planch. (1847). oily, copious; embryo straight to curved.
Two spp., SE Asia, mostly in rain forests.
Lianas, rarely shrubs, often hooked, indumentum
present. Leaves spiral; stipules entire to palmatifid.
Inflorescences axillary or terminal, racemes or 4. Roucheria Planch.
panicles, or axillary cymes, rarely solitary flowers; Roucheria Planch., London J. Bot. 6: 141 (1847); Jardim,
bracts and prophylls present or missing; sepals M.Sc. Thesis Univ. of Missouri, St. Louis (1999), rev.;
Linaceae 243
Ramı́rez et al., Fl. Venez. Guayana 5: 620–623 (1999), reg. 5, with 1 or 2 rows of glandular hairs near margin
rev.; Jardim & Berry, Novon 9: 520–523 (1999), key. and numerous parallel nerves; petals 5, contort,
Shrubs or trees. Leaves alternate, secondary veins purple or white; stamens 5; nectaries 3, at base of
parallel; petiole sometimes indistinct; stipules staminal tube; staminodia 5, filiform; ovary
small. Inflorescences axillary or terminal, panicu- 3-locular without false septae; stylodia 3; stigmas
late or fasciculate; sepals connate at base; petals discoid. Fruit a 1-seeded nut. Seeds oblong.
glabrous or pubescent on margins; stamens Two spp., C China, Himalaya, Thailand.
10(15); nectaries 10; ovary 1–5-locular, ovule
1–2 per locule, subapical; stylodia (2)3(–5); stig- 7. Tirpitzia Hallier f. Fig. 56H
mas discoid. Fruits drupaceous, subglobose, 1–5-
Tirpitzia Hallier f., Beih. Bot. Centralbl. 39(2): 5 (1921).
locular, little fleshy, endocarp 3–5-angled. Seeds 1
or 2 per locule, embryo somewhat curved. Densely branched shrubs. Leaves alternate, petio-
Seven spp., tropical South America. late, obovate-spathulate, nerves pinnate; stipules
minute, caducous. Inflorescences cymose; sepals
5. Hebepetalum Benth. Fig. 55 5; petals 5, white, claw about three times as long
as limb; stamens 5; 5 nectaries at base of staminal
Hebepetalum Benth. in Bentham & Hooker, Gen. Pl. 1: 244 tube present; staminodia 5; ovary 4–5-locular;
(1862); Jardim, M.Sc. Thesis Univ. of Missouri, St. Louis
(1999), rev.; Ramı́rez et al., Fl. Venez. Guayana 5: 618–620 false septae incomplete; stylodia 4–5, connate at
(1999), reg. rev.; Jardim & Berry, Novon 9: 520–523 base; stigmas pear-shaped. Capsule splitting into
(1999), key. 4–5 two-seeded valves.
Three spp., S China, Thailand, Vietnam.
Small to tall trees. Leaves alternate, pedicel dis-
tinct; stipules small. Inflorescences terminal,
paniculate; sepals somewhat connate at base; 8. Reinwardtia Dumort.
petals with short claw, adaxially pubescent; Reinwardtia Dumort., Comment. Bot.: 19 (1822).
androecium with 5 nectaries; ovary 4–5-locular, Erect or prostrate, glabrous shrubs or subshrubs;
stylodia 5, connate at base, with thickened branches erect and prostrate, the latter with
stigmas; locules 1–2-ovulate. Drupe ovoid, adventitious roots. Leaves alternate, petiolate,
1–5-locular, mesocarp black. Seeds 1(2) per base attenuate, elliptic-oblong to lanceolate-
locule; embryo somewhat curved. obovate, nerves pinnate; stipules minute,
Three spp. from tropical South America, in caducous. Inflorescences thyrso-paniculate,
lowland evergreen forests, often on white sand. umbelliform, rarely flowers solitary; flowers he-
terostylous; sepals 5; petals 5, yellow; stamens 5;
II. S U B F A M . L I N O I D E A E Arn. (1832). nectaries at base of staminal tube 2 or 3; stami-
nodia 5; ovary 3–4-locular; false septae present;
Herbs, subshrubs or shrubs; leaves alternate or stylodia 3–4, connate at base; stigmas capitate.
opposite; stipules sometimes reduced to glands Capsule splitting into 6–8 one-seeded valves.
or 0; petals clawed; stamens as many as sepals, Seeds compressed, reniform. 2n ¼ 20, 22.
connate at base; staminodia usually present, ante- One sp., R. indica Dumort. [syn. R. trigyna
sepalous; ovules 2 per locule, locules usually (Roxb.) Planch.], N India, China, SE Asia.
divided into 1-seeded portions; fruit usually a
capsule; mostly northern hemisphere. 9. Linum L. Fig. 56A–G
Linum L., Sp. Pl.: 277 (1753); Rogers, Brittonia 15: 97–122
6. Anisadenia Wall. ex Meisn. Fig. 56I (1963), rev. E North Am.; Rogers, Brittonia 20: 107–135
(1968), rev. C & W North Am.; Mildner & Rogers, Phyto-
Anisadenia Wall. ex Meisn., Pl. Vasc. Gen. 2: 96 (1838). logia 39: 343–390 (1978), rev. South Am.; Rogers, Nord. J.
Bot. 1: 711–722 (1981), rev. South Afr.
Perennial herbs; stems simple or little branched.
Leaves alternate, petiolate, lanceolate, entire or Perennial or annual herbs or rarely shrubs, gla-
slightly denticulate; stipules present, ovate-lance- brous or hairy. Leaves alternate, rarely opposite,
olate. Inflorescences thyrsoid or botryoid; sepals sessile, often lanceolate or sometimes spathulate at
base, entire, nerves parallel; stipular glands present Seeds narrowly ovate, wedge-shaped in cross-
or absent. Inflorescences cymose. Flowers homo- section. 2n ¼ 12, 16.
stylous or heterostylous; sepals 5, sometimes with One sp., S. digynum (A. Gray) C.M. Rogers,
glandular hairs; petals 5, contort, usually yellow, W North America.
blue, white or pinkish, distinct or connate at base;
stamens 5; nectaries at base of staminal tube 0–5;
12. Hesperolinon (A. Gray) Small
staminodia 5 if present; ovary 5-locular; false
septae incomplete or complete; stylodia 5, distinct Hesperolinon (A. Gray) Small in N. Amer. Fl. 25: 84 (1907);
Sharsmith, Univ. Calif. Publ. Bot. 32: 235–314 (1961), rev.
or connate; stigmas linear or capitate. Capsule
splitting into 10 one-seeded valves, 5 two-seeded Ephemeral erect annuals. Leaves alternate,
valves or indehiscent with 10 one-seeded segments. opposite or whorled, sessile, filiform to linear,
Seeds compressed. 2n ¼ 12, 16, 18, 20, 24, 26, 28, caducous; stipular glands present or absent.
30, 32, 34, 36, 40, 42, 52, 54, 60, 62, 68, 72, 84, 86(?). Inflorescences cymose. Flowers homostylous;
About 180 spp., widely distributed in temper- sepals 5, glabrous or hairy; petals 5, yellow to
ate and subtropical regions, esp. Mediterranean orange or white to pink/lavender, base with adaxial
region; five sects. Cathartolinum, Dasylinum, (glandular?) appendages; stamens 5; staminodia
Linopsis, Linum, Syllinum. Linum is paraphyletic absent; nectaries present (?); ovary 2–3-locular;
with respect to Radiola, Hesperolinon, Cliococca false septae incomplete; stylodia 2-3, distinct;
and Sclerolinon (McDill et al. 2009; McDill and stigmas minute. Capsule splitting into 4–6 one-
Simpson 2011), but taxonomic consequences seeded valves. Seeds oblong to clavate, triangular
remain to be addressed. or wedge-shaped in cross-section. 2n ¼ 34, 36.
Thirteen spp., California and Oregon.
10. Radiola Hill
Radiola Hill, Brit. Herb.: 227 (1756). 13. Cliococca Bab.
Small annual, glabrous herbs; leaves opposite, ses- Cliococca Bab., Proc. Linn. Soc. Lond. 1: 90 (1841), Trans.
sile, ovate to lanceolate, 1-nerved; stipules 0. Linn. Soc. 19: 33 (1842); Rogers & Mildner, Rhodora 73:
Inflorescences dichasial. Flowers homostylous; 560–565 (1971), rev.
sepals 4, apically (2)3(4)-toothed; petals 4, Perennial, glabrous herbs with extensively
contort, white, small; stamens 4; nectaries indis- branched underground rootstock. Leaves alter-
tinct; staminodia minute or absent; ovary nate, linear, 1-nerved; stipular glands absent. So-
4-locular; false septae incomplete; stylodia 4, dis- litary flowers terminal on branches, homostylous;
tinct; stigmas capitate. Capsule splitting into sepals 5, margins entire; petals 5, imbricate, whi-
8 1-seeded valves. Seeds irregularly ovate. 2n ¼ 18. tish, shorter than sepals; stamens 5; staminodia
One sp., R. linoides Roth, temperate Asia, sometimes present; nectaries present (?); ovary 5-
Europe, the Mediterranean region and montane locular; false septae complete; stylodia 5, distinct,
tropical Africa. very short; stigmas capitate. Capsule indehiscent,
10 one-seeded segments. Seeds elliptic. 2n ¼ 36.
11. Sclerolinon C.M. Rogers One sp., C. selaginoides (Lam.) Rogers &
Sclerolinon C.M. Rogers, Madroño 18: 182 (1966). Mildner, temperate South America.
Annual, glabrous herbs, simple or occasionally

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Grigorieva, V.V. 1989. O dimorfizme pyl’tsy nekotorykh of flaxseed. In: Muir, A.D., Westcott, N.D. (eds.) Flax
distil’nykh vidov roda Linum (Linaceae) [On pollen – The genus Linum. London, New York: Taylor &
dimorphism in some distylous species of the genus Francis, pp. 252–267.
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Hegnauer, R. 1966. See General References. cular diseases. In: Muir, A.D., Westcott, N.D. (eds.)
Hegnauer, R. 1989. See General References. Flax – The genus Linum. London, New York: Taylor
Heimsch, C., Tschabold, E.E. 1972. Xylem studies in the & Francis, pp. 204–213.
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Heinze, U., Amelunxen, F. 1984. Zur Schleimbildung in plants. Madroño 15: 49–52.
Linum-Samen. Elektronenmikroskopische und che- Rickard-Bon, S.E., Thompson, L.U. 2003. The role of
mische Analysen. Ber. Deutsch. Bot. Ges. 97: flaxseed lignans in hormone-dependent and inde-
451–464. pendent cancer. In: Muir, A.D., Westcott, N.D.
(eds.) Flax – The genus Linum. London, New York: Troll, W. 1964. Die Infloreszenzen. Typologie und Stel-
Taylor & Francis, pp. 181–203. lung im Aufbau des Vegetationsk€ orpers, I. Jena:
Robertson, K.R. 1971. The Linaceae in the southeastern Fischer.
United States. J. Arnold Arbor. 52: 649–665. Troll, W., Weberling, F. 1989. Infloreszenzuntersuchungen
Rogers, C.M. 1980. Pollen dimorphism in distylous spe- an monotelen Familien. Stuttgart, New York: Fischer.
cies of Linum sect. Linastrum (Linaceae). Grana 19: Uno, G.E. 1984. The role of persistent sepals in the repro-
19–20. ductive biology of Linum pratense (Linaceae). South-
Rogers, C.M. 1985. Pollen morphology of the monotypic west. Nat. 29: 429–433.
genus Cliococca (Linaceae). Grana 24: 121–123. Vazart, J. 1969. Organisation et ultrastructure du sac
Rogers, C.M., Xavier, K.S. 1971. Pollen morphology as an embryonnaire du Lin (Linum usitatissimum L.).
aid in determining relationships among some widely Rev. Cytol. Biol. Veg. 32: 227–240.
separated Old World species of Linum. Grana 11: Vazart, J. 1971. Dégénérescence d’une synergide et pénétra-
55–57. tion du tube pollinique dans le sac embryonnaire de
Rogers, C.M., Mildner, R., Harris, B.D. 1972. Some addi- Linum usitatissimum L. Ann. Univ. A.R.E.R.S. 9: 89–97.
tional chromosome numbers in the Linaceae. Britto- Vazart, B., Vazart, J. 1965. Infrastructure de l’ovule de lin,
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Roth, I. 1977. Fruits of angiosperms. Handbuch der Pflan- thélium. Compt. Rend. Hébd. Séances Acad. Sci.
zenanatomie, ed. 2, vol. 10(1). Berlin, Stuttgart: Paris 261: 2927–2930.
Borntr€ager, pp. 233–235. Vazart, B., Vazart, J. 1966. Infrastructure du sac embry-
Saad, S.I. 1961. Pollen morphology and sporoderm strati- onnaire du lin (Linum usitatissimum L.). Rev. Cytol.
fication in Linum. Grana 3: 109–129. Biol. Vég. 29: 251–266.
Saad, S.I. 1962. Palynological studies in the Linaceae. van Hooren, A.M.N., Nooteboom, H.P. 1984. Linaceae and
Pollen Spores 4: 65–82. Ctenolophonaceae especially of Malesia, with notes
Soltis, D.E. et al. 2011. See General References. on their demarcation and the relationships with Ixo-
Souèges, R. 1937. Développement de l’embryon chez le nanthaceae. Blumea 29: 547–563.
Radiola linoides Roth. Bull. Soc. Bot. France 84: van Welzen, P.C., Baas, P. 1984. A leaf anatomical contri-
297–306. bution to the classification of the Linaceae complex.
Str€obl, F. 1925. Die Obdiplostemonie in den Bl€ uten. Bot. Blumea 29: 453–479.
Archiv 9: 210–224. Winkler, H. 1931. Linaceae. In: Engler, A., Harms, H.
Sugawara, T., Tanaka, N., Murata, J., Zaw, K.M. 2002. (eds.) Die nat€ urlichen Pflanzenfamilien, ed. 2, 19a.
Dimorphism of pollen grains and stigmas in the het- Leipzig: Engelmann, pp. 82–130.
erostylous subshrub, Reinwardtia indica (Linaceae) in Xavier, K.S., Mildner, R.A., Rogers, C.M. 1980. Pollen
Myanmar. Acta Phytotax. Geobot. 53: 173–180. morphology of Linum, sect. Linastrum (Linaceae).
Sutter, D., Endress, P.K. 1995. Aspects of gynoecium Grana 19: 183–188.
structure and macrosystematics in Euphorbiaceae. Xi, Z., Ruhfel, B.R., Schaefer, H., Amorim, A.M., Sugu-
Bot. Jahrb. Syst. 116: 517–536. maran, M., Wurdack, K.J., Endress, P.K., Matthews,
Takhtajan, A.L. (ed.) 1980. See General References. M.L., Stevens, P.F., Mathews, S., Davis, C.C. 2012.
Thompson, J.D., Pailler, T., Strasberg, D., Manicacci, D. Phylogenomics and a posteriori data partitioning
1996. Tristyly in the endangered Mascarene Island resolve the Cretaceous angiosperm radiation Mal-
endemic Hugonia serrata (Linaceae). Amer. J. Bot. pighiales. Proc. Natl. Acad. Sc. (PNAS) 109(43):
83: 1160–1167. 17519–17524.
Lophopyxidaceae
Lophopyxidaceae (Engl.) H.H. Pfeiffer in Rev. Sudamer. Bot. 10: 4 (1951).
Lophopyxidoideae Engler in Engler & Prantl, Nat. Pflanzenfam. III, 5: 257 (1893) (in Icacinaceae).
K. K U B I T Z K I
Monoecious climbing shrubs or small trees with bundles; the vascular system of the midrib forms
watch-spring tendrils; branches with lateral bud at a closed flattened cylinder. Stomata are paracytic.
the base; cork superficial. Leaves spiral, simple, Crystals are present as solitary rhomboids and
serrulate to crenulate, domatiiferous; stipules clusters. Hairs are unicellular. The young stem
small, knob-like. Inflorescence a loose axillary pan- has five ribs with a continuous xylem cylinder,
icle with flowers in distant glomerules, the basal which encloses a pentagonal pith with a central
bracts often transformed into completely coiled portion of thick-walled parenchyma cells and a
tendrils. Flowers regular, sessile, small, 5-merous; marginal area of thin-walled cells; Engler (1893)
sepals basally connate, valvate, persistent; petals erroneously interpreted it as intraxylary phloem.
distinct, much smaller than sepals; disk yellowish; Between the ribs the secondary xylem has numer-
staminate flowers: stamens 5(6), antesepalous, with ous vessels; in the ribs the vessels are narrower
filiform filaments and subglobose, introrse, almost and scarcer. Vessel element perforation is simple.
basifixed anthers, alternating with 5(6) antepetalous Through anomalous activity of the cambium, five
cordate glands adnate to the petals; pollen grains phloem strands become enclosed within the
subprolate, 3-colporate, exine reticulate; pistillode xylem in the young shoot. Later intraxylary
subglobose, hairy; pistillate flowers: glands concres- phloem is formed as continuous bands alternat-
cent into a 5(6)-lobed disk with the glands or lobes ing with secondary xylem. The phloem is strati-
opposite the ovary cells; gynoecium 5(4)-carpellate, fied into soft and fibrous portions. Axial
carpels antepetalous; ovary superior, 5(4)-locular, parenchyma is scarce and paratracheal. The
ovoid-oblong, shallowly ribbed; ovules 2 per locule, ground tissue of the xylem consists of fibres
pendulous, apical-axile, anatropous, epitropous, with numerous minutely bordered pits. Rays
each surmounted at the micropyle by a small vary from 1–6-seriate in the young stem. The
obturator-like appendage originating from the funi- outer phloem is surrounded by a cylinder of
cle; stylodia 5(4), minute, subulate, stigmatic. Fruit fibres and stone cells. Cork arises in the subepi-
indehiscent, fusiform, 1-locular and 1-seeded, with dermal layer.
5 broad stramineous wings. Seed oblong, albumin-
ous; embryo erect, with oblong cotyledons and a E M B R Y O L O G Y . The ovule is bitegmic and weakly
short erect radicle. crassinucellate (Mauritzon in Sleumer 1942).
A single genus and sp., Lophopyxis maingayi
Hook. f., in forests from the Malay Peninsula P O L L E N M O R P H O L O G Y . Pollen is 3–4-colporate,
through Malesia to the Solomon and Caroline subprolate, 26 mm long (Erdtman 1952) or foveate,
islands. 39 30 mm (Dahl 1955).
VEGETATIVE MORPHOLOGY AND ANATOMY E C O L O G Y . Sleumer (1971) reports Lophopyxis

(Information from Handa 1940 and Baas as a straggling climber at the edge or in the
in Sleumer 1971). Axillary branchlets are meta- canopy of primary lowland forests, both in well-
morphosed into strong woody tendrils, which drained and swamp forest, in littoral forest and
distally are coiled and often bear a bud. The even sometimes in mangroves, from sea level up
petiole shows a strongly incurved arc of separate to ca. 300 m.
248 K. Kubitzki
with which they share paracytic stomata, uni-

sexual 5-merous flowers with a disk or disk
segments, 2 apical epitropous ovules per carpel,
and 1-seeded fruits (Stevens 2001 onwards).
The clade of Lophopyxidaceae and Putranjiva-
ceae, the “putranjivoids”, has now been
resolved as part of a well-supported tetratomy
that otherwise includes the chrysobalanoids,
the malpighioids and the Caryocaraceae (Xi
et al. 2012).
A single genus:
Lophopyxis Hook. f. Fig. 57

Lophopyxis Hook. f. in Hook., Icon. Pl. 18: t. 1714 (1887).
Description as for family.
Dahl, A.O. 1955. The pollen morphology of several genera
excluded from the family Icacinaceae. J. Arnold
Arbor. 36: 159–163, 1 pl.
Engler, A. 1893. S.-B.Preuss. Akad. Wiss. Berlin 18:
265–266 (a note on Lophopyxis in an article about
anatomical characters of Icacinaceae).
Erdtman, G. 1952. See General References.
Handa, T. 1940. Anomalous secondary growth in the axis
of Lophopyxis pentaptera (K. Schum.) Engler. Bot.
Fig. 57. Lophopyxidaceae. Lophopyxis maingayi. A Mag. Tokyo 54: 41–47, figs. 1–6.
Habit. B Staminate flower buds. C Staminate flower. D Pfeiffer, H. 1926. Das abnorme Dickenwachstum. In: Lins-
Stamens, front and back view. E Pistillate flower. F Pistil bauer, H., Handbuch der Pflanzenanatomie, vol. 9.
vertically sectioned. G Ovary, cross section. H Infructes- Berlin: Borntraeger.
cence. I Fruit. (Sleumer 1971) Pfeiffer, H.H. 1951. Lophopyxis als Typus einer eigenen
Familie. Rev. Sudamer. Bot. 10: 3–6.
A F F I N I T I E S . Formerly (and sometimes still Sleumer, H. 1942. Lophopyxis. In: Engler & Prantl, Nat.
recently) Lophopyxis was placed in or close to Pflanzenfam. ed. 2, 20b, pp. 392–396. Leipzig: W.
Euphorbiaceae, Icacinaceae or Celastraceae, Engelmann.
Sleumer, H. 1968. The genus Lophopyxis Hook.f. (Lopho-
plus four further families (see Sleumer 1968 pyxidaceae). Blumea 16: 321–323.
for details), but Pfeiffer (1951) found neither Sleumer, H. 1971. Lophopyxidaceae. In: Flora Males. I,
position defendable and suggested family rank 7: 89–92. Leyden: Nordhoff.
Soltis, D.E. et al. 2011. See General References.
for it. Lophopyxidaceae are resolved as the Stevens, P.F. 2001 onwards. See General References.
strongly supported sister to Putranjivaceae Wurdack, K.J., Davis, C.C. 2009. See General References.
(Wurdack and Davis 2009; Soltis et al. 2011) Xi, Z. et al. 2012. See General References.
Medusagynaceae
Medusagynaceae Engler & Gilg (1924), nom. cons.
W.C. D I C K I S O N { AND K. K U B I T Z K I 1
Small tree with a rounded crown. Leaves oppo- altitudes in pockets of soil between granite
site, simple, elliptic or elliptic-oblong, glabrous, masses (Hemsley 1905).
with a retuse apex and apical sinus, an attenuate
base, and fine, widely spaced marginal glandular VEGETATIVE MORPHOLOGY AND ANATOMY. Medusagyne
teeth, apical sinus contains a colleter on either grows to about 10 m in height; its trunk is up to
side of the depression and a single one at the 20 cm in diameter with dark, fibrous, striated
base of the cavity; stipules 0. Inflorescences bark (Robertson et al. 1989). Vegetative anatomy
lax, multiflowered, paniculate. Flowers regular, was described by Beauvisage (1920) and Dickison
hypogynous, bisexual and male, the plants andro- (1990a). Leaves are subcoriaceous with pinnate
monoecious; staminate flowers without evidence and brochidodromous venation. Veins are sur-
of an abortive gynoecium; sepals 5, connate at rounded by a double sheath, a dense inner fibrous
the base, quincuncial in their distinct apices; zone and an outer, parenchymatous layer. The
petals 5, contort, at first spreading, later reflexed; petiole contains numerous small collateral bun-
stamens numerous, filaments distinct, slender, dles having various orientations and arranged in
anthers basifixed, bithecate and tetrasporangiate, an arc. The mesophyll contains mucilaginous and
latrorse, dehiscing by slits; connective shortly crystalliferous cells. Cristarque cells have not
protruding as apical extension; carpels numer- been found. Nodal anatomy is multilacunar, mul-
ous, 16–25, attached to central axis, each on its titrace, an infrequent pattern in dicotyledons with
outer shoulder with a short stylodium and capi- opposite leaves (Dickison 1990a). As described
tate stigma; ovules anatropous, epitropous, biteg- by Dickison (1990a), the wood contains solitary
mic, 2 per locule, on separate axile placentas, one vessel elements with simple perforations. Inter-
ascending, the other descending; endosperm vascular pitting is opposite to predominantly
cellular. Fruit a septicidal capsule, each carpel alternate. Tracheids are present. Rays are hetero-
separating acropetally from the central column geneous with both uniseriates and multiseriates;
along its entire margin, the dehisced carpels axial parenchyma is diffuse and diffuse-in-
only maintaining a distal connection to the per- aggregates, also paratracheal scanty. Tannini-
sistent columella. Seeds winged, with a thin layer ferous tissue is present throughout the plant body.
of endosperm and a straight embryo. Sieve-tube plastids are of the S-type (H.-D. Behnke,
A single genus and sp., Medusagyne oppositi- pers. comm.).
folia J.G. Baker, endemic to Mahé of the
Seychelles Islands, where it occurs at middle
F L O R A L M O R P H O L O G Y A N D A N A T O M Y . The
actinomorphic flowers are either bisexual or
staminate (Robertson et al. 1989; Dickison
1990a). Nothing is known of the pollination
1
In his paper on Medusagyne (Dickison 1990a) and in biology or breeding system. Bisexual flowers
the manuscript to this contribution, the late Dr. Dickison possess numerous, distinct stamens that
had ascribed spiral phyllotaxis to the androecium of surround the gynoecium in layer-like groups
Medusagyne, which definitely does not exist there; I
have deleted this from the manuscript, and also added that differ in length and are vascularized by
some recent information on the family. the breakup of individual stamen fascicle traces
K. Kubitzki (stamen trunk bundles). The biovulate carpels
250 W.C. Dickison and K. Kubitzki
Fig. 58. Medusagynaceae. Medusagyne oppositifolia. vertical section showing biovulate condition and ovulary
A Fruiting branching. B Flower bud. C Bisexual flower, vascular supply (OVB). G Stamen. H Fruit. I Open fruit.
vertical section. D Staminate flower, vertical section. J Seed. (A original; B–D Robertson, Wise & White 1989;
E Bisexual flower, transverse section. F Carpel locule, E, F Dickison 1990a)
occur in a single whorl and are borne on the and curved. The upper ovule of the pair is
sides of a broad, elongated torus that represents ascending or erect, micropyle below, raphe
the upward extension of the receptacle. In a dorsal. The micropyle is straight. The inner
manner somewhat comparable to Dillenia (Dil- integument is 3–4 cells thick, and the outer
leniaceae), carpels show pronounced ontoge- epidermal cells of the inner integument differ-
netic adaxial deformation, during which the entiate into a fibrous layer in the seed. Ovules
upper ventral surfaces become expanded, are weakly crassinucellate and vascularized by a
greatly extending the ventral margins and form- single bundle that extends unbranched from the
ing a domelike cushion of carpel-derived tissue funiculus to the chalaza. Embryo sac and
over the summit of the torus (Dickison 1990b). embryo development are unknown.
Details of floral vascularization were presented
by Dickison (1990a, b). P O L L E N . Pollen is subisopolar, i.e. the distal and
proximal faces are unequal, one being very con-
E M B R Y O L O G Y . The anther wall contains a sub- vex, the other less convex, triporate, semiangular,
dermal endothecial layer with cells having well- small (13–15 x 14–17 mm). Apertures protrude.
developed bands of secondary thickenings and Sculpture striate, intertwined at different levels.
an inner tapetal layer. Each carpel locule con- At the TEM level, pollen is tectate-columellate
tains a pair of superimposed ovules attached with a massively developed foot layer (Dickison
separately on axile placentae. Funiculi are long 1990a).
Medusagynaceae 251
F R U I T A N D S E E D . As the fruit matures, carpels dam construction. Attempts to cultivate Medusa-

unfold on a radial frame from the base upwards gyne have met with somewhat limited success.
in a manner resembling the ribs of an umbrella.
In mature pericarps an inner region of thick- One monotypic genus:
walled, lignified cells lines the locules. Seeds are
small, c. 3.0 mm in length, compressed, winged Medusagyne J.G. Baker Fig. 58
and exarillate with reduced endosperm. The
Medusagyne J.G. Baker, Fl. Mauritius Seych.: 17 (1877).
mature seed coat is exotegmic-exotestal with 4
or 5 layers of cells with thickened and suberized Characters as for family.
walls and a tegmic layer of thick-walled, lignified
fibres. The endosperm is represented by a nar-
row, cellular zone without starch. The embryo is
straight. See the detailed study of the fruit and Selected Bibliography
seed anatomy by Doweld (1998).
Baker, J.B. 1877. Flora of Mauritius and the Seychelles: a
A F F I N I T I E S . The totality of the structural evi- description of the flowering plants and ferns of those
islands. London: L. Reeve & Co.
dence confirms the view that Medusagyne is a Beauvisage, L. 1920. Contribution a l’étude anatomique de
very distinct genus. Cladistic analyses based on la famille des Ternstroemiacées. Doctoral diss., Univ.
the plastid gene rbcL by Fay et al. (1997) place de Poitiers. Tours: E. Arrault et Cie.
Medusagyne in a clade with Quiinaceae and Och- Dickison, W.C. 1990a. The morphology and relationships
of Medusagyne (Medusagynaceae). Pl. Syst. Evol.
naceae, and Xi et al. (2012) resolve it with moder- 171: 27–55.
ate support in a sister position with Quiinaceae. Dickison, W.C. 1990b. An additional note on the floral
With these families, Medusagyne shares various morphology and affinities of Medusagyne oppositifo-
characters including multilacunar nodes, muci- lia (Medusagynaceae). Brittonia 42: 191–196.
Doweld, B. 1998. On the phylogenetic relationships of
lage cells and dentate leaves, but differs in, i.a., Medusagyne (Medusagynaceae) as evidenced by the
the organisation of the petiolar vascular supply. structure of its fruits and seeds. Bot. Zhurn. 83:
54–68.
D I S T R I B U T I O N A N D H A B I T A T S . Medusagyne Fay, M.F., Swensen, S.M., Chase, M.W. 1997. Taxonomic
affinities of Medusagyne oppositifolia. (Medusagyna-
oppositifolia is endemic to Mahé, the largest ceae). Kew Bull. 52: 111–120.
island in the Seychelles group. It occurs as a Hemsley, W.B. 1905. Medusagyne oppositifolia J. G.
very rare component of the intermediate forest Baker. Hook. Icon. Plant. IV, 8: 1–3, pl. 2790.
Hickey, L.J., Wolfe, J.A. 1975. The bases of angiosperm
zone. phylogeny: vegetative morphology. Ann. Missouri
Bot. Gard. 62: 538–589.
C O N S E R V A T I O N . The genus was once feared Matthews, M.L. et al. 2012. See General References.
extinct, and even today only very few individual Robertson, A., Wise, R., White, F. 1989. Medusagyne
oppositifolia. Kew Mag. 6: 166–171.
plants remain alive in scattered populations. One Soltis, D.E. et al. 2011. See General References.
locality is now a national park nature reserve, Wurdack, K.J., Davis, C.C. 2009. See General References.
although the area is threatened by a proposed Xi, Z. et al. 2012. See General References.
Ochnaceae
Ochnaceae DC., Nouv. Bull. Sci. Soc. Philom. Paris 2: 209 (1811), nom. cons.
M.C.E. A M A R A L AND V. B I T T R I C H
Trees, shrubs or shrublets, generally evergreen, V E G E T A T I V E M O R P H O L O G Y . The great bulk of

rarely herbs, mostly glabrous. Leaves alternate, gen- the Ochnaceae are evergreen shrubs or small trees.
erally simple, often coriaceous, entire, serrate, ciliate Annual or perennial herbs are restricted to a few
or with persistent or caducous teeth, secondary Sauvagesia spp., larger trees occur in Lophira and
veins often numerous and densely parallel; stipules some species of Brackenridgea. The leaves are
distinct or more rarely basally intrapetiolarly con- often coriaceous and glossy, petiolate, alternate
nate, persistent or caducous, entire or laciniate. and stipulate. Phyllotaxy is generally distichous on
Inflorescences terminal or axillary, cymose or race- plagiotropic branches and spiral on orthotropic
branches. The young shoots of Perissocarpa are
mose, more rarely flowers solitary in leaf axils.
densely covered with cataphylls (Walln€ofer 1998).
Flowers on articulated pedicels, generally bisexual,
The stipules are distinct or more rarely at base
actinomorphic or zygomorphic; sepals generally 5,
intrapetiolarly connate (Campylospermum, Rhabdo-
mostly distinct, imbricate or nearly valvate, cadu- phyllum, Idertia, Testulea), entire, toothed or
cous or persistent, sometimes accrescent, envelop- laciniate, persistent or caducous, leaving small or
ing the mature bud or not; petals generally 5, mostly conspicuous scars. Stipules in Ouratea sometimes
distinct, imbricate, contort (cochlear or quincun- form extrafloral nectaries (Oliveira and Leitão Filho
cial), caducous; stamens 5–10–1, rarely 1, with 1987).
sometimes persistent filaments, generally distinct, The leaves are simple, with the exception of
anthers basifixed, bithecate, tetrasporangiate, Rhytidanthera. The margins are straight, crenate,
dehiscing latrorsely with longitudinal slits or with serrate or serrulate, and mostly denticulate, ciliate
one or two apical or subapical pores; staminodes in some Luxemburgia spp. Many genera of all sub-
present or absent, distinct or connate, petaloid or families show leaves with rather closely spaced,
not, sometimes completely enveloping stamens and rigidly parallel secondary veins, with the tertiary
gynoecium; nectary absent; ovary superior, (2)3(4) veins perpendicular or more rarely nearly parallel
or 5(–15)-carpellate, syncarpous at least at the basal to these. Secondary veins curving strongly near
part of the ovary, and either entire with one apical the margin and continuing more or less parallel
style or deeply and apparently completely divided to it are typical for Ouratea and related genera
into five to several uniovulate lobes with one gyno- (Fig. 59A).
basic style; style distinct from or continuous with
V E G E T A T I V E A N A T O M Y . Most information of
the ovary apex; androgynophore and gynophore
the literature is summarized in Solereder (1899
usually present, often inconspicuous; placentas (2)
and 1908), Metcalfe and Chalk (1950), Carlquist
3(4) or 5(–15) basal, axile or parietal, stigmas dis-
(1988) and Amaral (1991). Cristarque cells
tinct or connate. Fruit generally a septicidal capsule, with a solitary sphaerocrystal are a typical feature
rarely a nut or drupe, or separating into 2 to several of the family (Solereder 1899), and not only for
blackish drupelets on an accrescent reddish recep- the Ochnoideae as assumed by Van Tieghem
tacle. Seeds exarillate, albuminous or exalbuminous, (1902a) and Metcalfe and Chalk (1950). Details
winged or not; embryo straight or curved. about petiole vascularization were reported (partly
A pantropical family of 27 genera with about contradictory) by Metcalfe and Chalk (1950),
500 spp., divided into three subfamilies. Decker (1967) and Schofield (1968). A simple arc
254 M.C.E. Amaral and V. Bittrich
with two lateral traces occurs in Sauvagesia p.p. sometimes even starting in the epidermis
and Tyleria. The bundles may form a solid sipho- (e.g. Ochna, Godoya). Euthemis, Sauvagesia and
nostele in the upper part of the petiole or in the Schuurmansia are characterized by large mucilage
blade (e.g. Elvasia, Lophira, Luxemburgia, Ochna, cells in the cortex and medulla (data from Decker
Wallacea), sometimes with medullary bundles 1966; Schofield 1968).
(Schofield 1968), or they remain separate through- Wood anatomy in Sauvagesioideae is rather
out the leaf partly forming a dissected siphonostele homogeneous, with the exception of Tyleria and
with some medullary bundles (e.g. Ochna, Oura- Testulea, while the Ochnoideae are more hetero-
tea, Sauvagesia p.p.). According to Decker geneous. The pericycle is formed by small isolated
(1967), the latter condition is also characteristic fibre bundles, rarely the cells between the fibre bun-
for Euthemis and a group (probably a clade) dles become sclerified (Ouratea). Vasicentric trac-
formed by Cespedesia, Godoya, Rhytidanthera heids are rare (Lophira), septate fibre tracheids are
and Krukoviella. According to Metcalfe and common, and libriform fibres were found in a few
Chalk (1950), however, the petiole of Lophira genera (Tyleria, Wallacea). Vessels elements occur
shows isolated bundles. Hairs are very rare solitary or rarely in multiples, their perforation in
(Ouratea spp., Elvasia sp.), they are unicellular most species is simple or more rarely both simple
or uniseriate multicellular. The epidermis has and scalariform. Diagonal aggregation of vessel ele-
commonly some mucilage cells (e.g. Elvasia, ments was observed in Lophira. As regards vestured
Ochna, Ouratea, Luxemburgia), not rarely the cell pits, first reports indicated that they are characteris-
walls become lignified. Stomata are paracytic, tic only for the Ochnoideae and absent in the Sau-
sometimes with three subsidiary cells (Sauvagesia, vagesioideae (Bailey 1933). Jansen et al. (2001),
Lophira). They are mostly restricted to the abaxial however, found vestured pits also in various genera
side, more rarely they occur on both sides of Sauvagesioideae s.str. (e.g. Testulea, Wallacea)
(Ouratea spp., Lophira lanceolata), in Cespedesia and in Luxemburgia (Luxemburgioideae). They
they are restricted adaxially to the midrib (Sastre were absent only in Sauvagesia (one sp. investi-
1975a). Hydathodes were reported by Sastre (1975b) gated). Helical thickenings in vessels are sometimes
from Sauvagesia erecta and Cespedesia. Thick glan- present. According to Decker (1966), unilateral
dular hairs, probably colleters, occur on the inner compound pitting between parenchyma and vessel
stipule base in Cespedesia. The sepals of Godoya elements (which show the large pits) is typical for
and Rhytidanthera have very similar glandular the Sauvagesioideae (except Testulea) and absent
hairs at their inner base, while marginal glands are in the Ochnoideae. It is unclear, however, in how
found in several genera of Sauvagesioideae s.str. many genera this was actually observed. Rays are
The mesophyll is generally dorsiventral. Cris- mostly of the heterogeneous type (mainly Kribs type
tarque cells with druses are common in the region IIA), but homogeneous in Lophira and Testulea;
of the midrib and the lateral veins. A hypodermis they are (1)2–5(–8)-seriate. Axial parenchyma is
was observed in Lophira and Elvasia. Spicular cells scanty, it is paratracheal to vasicentric (Blaste-
occur as a layer under the upper epidermis in a few manthus, Cespedesia) or diffuse apotracheal (e.g.
genera (e.g. Blastemanthus, Cespedesia). Distinct most Ochnoideae), while in Lophira it forms
fibres in the mesophyll occur in Cespedesia and broad distinctive metatracheal bands (Decker
Rhytidanthera, and mucilage-filled cavities in the 1966). A chambered crystalliferous axial paren-
mesophyll of Euthemis (Schofield 1968). chyma is present in Elvasia, Lophira and Cespedesia.
Cortical vascular bundles are distributed The tissue of the pith often becomes early lignified.
throughout the family (lacking in, e.g. Krukoviella That of Lophira contains chambered crystalliferous
and Poecilandra), while medullar bundles are rare fibres.
(e.g. Godoya, Rhytidanthera and Cespedesia).
Nodes are multilacunar with many leaf traces (e.g. INFLORESCENCE AND FLOWER STRUCTURE. Most species
Cespedesia, Rhytidanthera), or trilacunar with gen- have cymose or racemose terminal, determinate
erally three (e.g. Schuurmansiella, Sauvagesia p.p.) or indeterminate inflorescences, only a few have
or, more rarely, many leaf traces (Blastemanthus, single axillary flowers. Inflorescence types com-
Poecilandra, Wallacea). Growth rings are some- prise racemes (e.g. Luxemburgia), panicles (e.g.
times evident. Stone cells are common in the cortex. Schuurmansia) and thyrses (e.g. Cespedesia) with
The initiation of cork development is superficial, dichasial or monochasial branching. The pedicels
Ochnaceae 255
A detailed comparative floral morphological

and anatomical study of the Malpighiales family
clade Ochnaceae/Quiinaceae/Medusagynaceae was
undertaken by Matthews et al. (2012). Flowers
are actinomorphic or zygomorphic. The Ochnoi-
deae are characterized by actinomorphic flowers
while, in Luxemburgioideae and Sauvagesioideae
s.str., 7 of the 14 genera have zygomorphic flowers.
Calyx and corolla, however, are always basically
actinomorphic. In several genera (e.g. Cespedesia,
Blastemanthus) zygomorphy develops only during
anthesis, with stamens shifting to the adaxial side
of the flowers and the ovary to the abaxial side
(Fig. 60G). Only in three genera does zygomorphy
exist already before anthesis: In Luxemburgioideae
the androecium surrounds the ovary only on the
adaxial side of the flowers (Fig. 60H), while Testu-
lea (Sauvagesioideae) has only one adaxial stamen.
Sepals are generally distinct, rarely they are con-
genitally united at the base and in Perissocarpa
connate for most of their length. In Ouratea and
segregated palaeotropical genera and in Bracken-
ridgea nitida, the margins of the inner sepals are
cleaved and clasp the margins of the outer ones
("quinconciale engrainée", Van Tieghem 1902b).
Most genera of the Ochnaceae have five sepals;
only Testulea with four, Elvasia with 2–6 and Blas-
temanthus with 10–15 sepals show deviating num-
bers. The outer sepals of some genera (e.g.
Poecilandra, Godoya) are much smaller than the
inner sepals. In Blastemanthus, the outer sepals
are small and distichous (possibly bracts rather
than sepals) and the inner ones are larger and
spirally arranged. Aestivation of the sepals is
mostly imbricate (quincuncial or more rarely
cochlear) or, more rarely, nearly valvate. Sepals
in most genera envelop the mature bud
completely, in a few genera only its basal part
Fig. 59. Ochnaceae. A–H Subfam. Ochnoideae. A–D (e.g. Euthemis, Schuurmansia, Cespedesia). Sepals
Ouratea spectabilis. A Flowering branch. B Flower.
C Longitudinal section of flower; note gynobasic style. are caducous or persistent. In most Ochnoideae
D Pistil, gynophore and very short filaments. E Ochna with gynobasic styles, the sepals are accrescent and
serrulata. Fruit with drupaceous mericarps on enlarged become red during fruit development like the
receptacle. F Lophira alata. Fruit with two enlarged receptacle. In Lophira the two outer sepals are
sepals. G, H Elvasia calophyllea. G Flower. H Fruit. (A–D
redrawn from Engler 1876, all others orig.) prolonged and form the wings of the samaroid
fruit (Fig. 59F). Marginal glands (Adenarake) and
basally adaxial (Godoya, Rhytidanthera) colleters-
are always articulated, generally near the base, like glands on the sepals are of unknown function.
more rarely in the middle or near the apex of the Petals are usually distinct, but very short sym-
pedicel. The receptacle of the flowers of all genera petaly occurs in Cespedesia and Euthemis. Post-
of Ochnoideae with gynobasic styles enlarges dur- genital fusion of the upper parts of the early
ing fruit development and becomes red-coloured. caducous petals occurs in Perissocarpa (Walln€ofer
1998). The vast majority of genera are character-

ized by five petals. In Elvasia the petal number
varies within a species or even within an inflore-
scence. Aestivation is either imbricate quincuncial
(Blastemanthus), cochlear (Luxemburgia spp.) or
usually contort. For the contort condition there
exists in some genera a variation called "préflorai-
son cloisonnée" (Van Tieghem 1902b), whereby the
margins of the petals enter between the stamens
and spirally enclose the style (Ouratea and segre-
gated genera, Perissocarpa (Ochnoideae) and
Euthemis (Sauvagesioideae)). Petals have one (e.g.
Luxemburgia, Philacra (Luxemburgioideae), Och-
noideae, Sauvagesia) or three (part of Sauvagesioi-
deae s.str., e.g. Blastemanthus, Cespedesia) vascular
traces.
The number of stamens varies between one
(Testulea) and more than 50 (Lophira), common
are five or ten stamens. Filaments are distinct
except in Philacra and Luxemburgia, were they are
congenitally connate and persistent. Persistent fila-
ments are also characteristic for the genera of Och-
noideae with gynobasic styles, in some other genera
the persistent stamens are enclosed by the persistent
sepals (e.g. Sauvagesia). Anthers are basifixed,
bithecate and tetrasporangiate; at the transition
from filament to anther, a complete or incomplete
pit may be formed (Matthews et al. 2012). In Och-
noideae, the dorsal pollen-sacs are sometimes smal-
ler than the ventral ones which, according to Eichler
(1878: 257), led some earlier authors to the errone-
ous conclusion that these anthers are monothecate.
The anthers dehisce by longitudinal slits of variable
length or more commonly by one or two pores. The
latter condition is probably primitive within the
family (Amaral 1991). Generally, in the poricidal
anthers an endothecium is present in the whole
anther wall, not only around the pores. This has
probably facilitated the evolutionary reversal from
poricidal dehiscence to dehiscence by slits. In Sau-
vagesia both dehiscence types occur. In Luxembur-
gia, some Ochnoideae (Elvasia, Campylospermum)
Fig. 60. Ochnaceae. A–G Subfam. Sauvagesioideae. A Sau- and Sauvagesioideae (Blastemanthus, Tyleria), how-
vagesia ericoides, flower. B S. alpestris. Flower with partially
removed petals showing the connate petaloid staminodes. ever, the endothecium is restricted to the level of
C–F S. racemosa. C Habit. D Staminodes. E Capsule the pores. Anthers with one pore are frequently
with persistent sepals, staminodes and stamens. F Separated characterized by uneven growth of their apex.
carpel of a capsule showing the placenta. G Blastemanthus The poricidal anthers of Ouratea spp. show regular
gemmiflorus, flower. H–J Subfam. Luxemburgioideae.
H Luxemburgia ciliosa, flower. I L. octandra, capsule, dehis- circular constrictions or wrinkles along the thecae
cence starts apically. J Philacra auriculata, capsule, dehis- (Fig. 59B, C), a condition typical for various buzz-
cence starts basically. (A–F redrawn from Eichler 1871, G, H pollinated flowers in the angiosperms. In some spe-
redrawn from Engler 1876, all others orig.) cies of Sauvagesia the connective is prolongated at
Ochnaceae 257
the anther apex. The anthers of a few genera are that at least an inconspicuous gynophore is usually
covered with wax crystals (e.g. Wallacea, Poecilan- present (Matthews et al. 2012). The gynophore
dra). In Luxemburgioideae, some Ochnoideae becomes swollen and fleshy in fruit in some Och-
(Elvasia, Ochna) and Sauvagesioideae (e.g. Cespede- noideae (Brackenridgea, Ochna, Ouratea).
sia), the anthers are caducous after anthesis. They In part of the Ochnoideae (Ochna, Campylo-
are generally persistent, however, in flowers with spermum, Brackenrigea and related genera) the
staminodes around the fertile stamens (e.g. Sauva- ovary is deeply lobed and for its major part apo-
gesia). carpous (contra Baum 1951), with each lobe
Staminodes occur in several genera of the containing only one ovule. The degree of apocarpy
Sauvagesioideae s.str., always outside the fertile varies in this subclade of Ochnoideae: the placenta
stamens, both are sometimes united at base (Sauva- may be part of the basal synascidiate region
gesia, Tyleria). Two types, petaloid and filamentous, (Campylospermum), or the synascidiate zone ends
can be distinguished. Both may occur in the same below the placenta and the placenta belongs already
flower, with the smaller filamentous staminodes to the plicate region (Brackenridgea, Ochna). The
surrounding the petaloid ones (Fig. 60D). Petaloid ovary in this group lacks a symplicate region
staminodes are highly variable in size and form. completely. Elvasia (also Ochnoideae), on the
They are distinct or connate at base (Tyleria) or other hand, is synascidiate up to above the placenta,
connate for more or less their whole length (Sauva- then symplicate in the upper part except for the
gesia p.p., Fig. 60B). Often large petaloid staminodes apocarpous stigmas. The gynoecium of the Luxem-
completely envelop the stamens and the gynoecium, burgioideae and Sauvagesioideae s.str. is entirely
leaving only a pore-like opening at the apex syncarpous with a basal synascidiate region that
(Fig. 60B). Filamentous staminodes are either subu- may reach only the mid-region or the basal part of
late (Blastemanthus, Schuurmansiella, Wallacea) or the placenta or end even below it. Most of the ovary
markedly broadened at the apex (Sauvagesia p.p.). is thus symplicate. In the Ochnoideae group with
The latter type may occur combined with petaloid mostly apocarpous gynoecium, the styles are
staminodes, the broadened tips possibly mimicking inserted gynobasically (Fig. 59C) and united
anthers. In Philacra auriculata several small stami- completely postgenitally (e.g. Ouratea, Campylos-
node-like filamentous structures occur on the same permum) or are distinct near the apex (Ochna
side as the fertile stamens, they are slightly broad- spp.). The stigma is punctiform in most genera,
ened apically and possibly glandulous here. either non-papillate (e.g. Elvasia, Luxemburgia, Phi-
The androecial ontogeny of Ochna atropur- lacra, Wallacea) or papillate (Campylospermum,
purea was studied by Pauzé and Sattler (1978). On Blastemanthus, Sauvagesia, Tyleria), but more or
five primary androecial primordia, stamina primor- less suction-cup-shaped in Ochna and Brackenrid-
dia development was observed to follow a centri- gea (and in Medusagynaceae and Quiinaceae; Mat-
petal pattern. Flowers of the Luxemburgioideae and thews et al. 2012). Pollen-tube transmission tissue is
Sauvagesioideae, however, show a typical centri- present in the styles and the ovary. A compitum is
fugal development of the androecium (Amaral and present in the symplicate zone in Luxemburgioideae
Bittrich 1998). An androgynophore occurs in all and Sauvagesioideae, in the uppermost part of the
Ochnaceae (Matthews et al. 2012), it is mostly very unilocular ovary in Elvasia (Ochnoideae), and at a
short, but conspicuous in Indosinia and Adenarake. short one in the transition from the synascidiate to
Detailed studies of the gynoecium morphology the apocarpous zone in the Ochnoideae with a
in the family were provided by Guédès and Sastre mainly apocarpous gynoecium. In Campylosper-
(1981) and Matthews et al. (2012). The ovary is mum additionally a compitum may be present in
superior and syncarpous at least at the base, with the closely associated papillae of the stigmas.
angiospermy type 2, 3 and 4. A short and broad In several genera of Sauvagesioideae s.str. and
gynophore is present in Ouratea (Fig. 59C, D) and in the Ochnoideae with gynobasic styles, the ovary
Elvasia (cf. Sastre 2004); in some other genera (e.g. is plurilocular, the placentation axile. In Elvasia
Blastemanthus and Wallacea) the ovary shows a (Ochnoideae) the ovary is unilocular above the
sterile basal zone in longitudinal section, but with- synascidiate zone (Matthews et al. 2012). In other
out being stipitate. A comparative study of Ochna- genera of Sauvagesioideae s.str. and in Lophira
ceae, Quiinaceae and Medusagynaceae showed (Ochnoideae), the ovary is either more or less
unilocular with parietal (e.g. Sauvagesia p.p., Sauvagesia erecta) as well as self-incompatibility
Schuurmansia) or basal (Sauvagesia p.p., Lophira) (e.g. Ouratea spruceana) was observed. Few species
placentation, or it is bi- to plurilocular at base have unisexual flowers (Sauvagesia serrata, Euthe-
and unilocular or incompletely plurilocular in mis minor, Schuurmansiella angustifolia, Schuur-
the upper part. The latter condition is also typical mansia henningsii).
for the Luxemburgioideae. In these cases the pla-
centation is axile at the base and parietal at the E M B R Y O L O G Y . The tapetum is secretory, division
apex. Sometimes the placenta is inserted on deeply of the pollen mother cells is simultaneous and the
protruding and postgenitally fused septa (e.g. Blas- pollen grains are two-celled when shed (except in
temanthus). In Wallacea the ovary is unilocular Ochna kirkii, Davis 1966; Narayana 1975). Ovules
and the placentation is laminar diffuse. Tanniferous are incompletely tenuinucellate (Luxemburgia,
tissue in floral organs is abundant in all Ochnaceae Philacra, Sauvagesia, Tyleria, Wallacea) or weakly
and in the closely related families Quiinaceae and crassinucellate (Blastemanthus) and anatropous or
Medusagynaceae (Matthews et al. 2012). campylotropous, syntropous (curved into the same
direction as the carpel closure), rarely additionally
F L O W E R B I O L O G Y . The predominance of porici- a few ovules are antitropous. Ovules are bitegmic in
dal anthers or of the "poricidal system", where the Luxemburgioideae and Sauvagesioideae s.str., but
pore is formed by staminodes (Fig. 60B; Kubitzki in Ochnoideae appear as mostly (Campylo-
and Amaral 1991), suggests that pollination by bees spermum, Elvasia, Ochna) or completely (Bracken-
capable of vibrating the anthers or the staminodes ridgea, Lophira) unitegmic due to fusion of the
around the stamens to collect pollen ("buzz-polli- integuments. The micropyle is formed either by
nation") is common in the family. Observations on both integuments and often zigzag (Blastemanthus,
Luxemburgia spp., Ouratea spp., and Poecilandra Luxemburgia, Philacra, Sauvagesia, Tyleria) or
retusa with poricidal anthers and on Sauvagesia more rarely straight (Cespedesia, Wallacea), or
spp. with a poricidal system in Brazil and Venezuela only by the inner (Campylospermum, Ochna, also
confirmed this prediction: Small to large bees of in Wallacea), and by the apparently only integu-
various taxonomic groups were observed to act as ment in Brackenridgea and Lophira (Ochnoideae)
pollinators (pers. obs.; Nadia and Machado 2005). (Narayana 1975; Matthews et al. 2012). The embryo
The apically elongated connective on anthers with sac is of the Polygonum type, endosperm formation
longitudinal dehiscence observed in some Sauva- is nuclear becoming cellular throughout. Apospory
gesia spp. might, similarly as in some Ternstroemia was found in two species of Ochna.
spp., transmit the vibrations from the staminodes
directly to the anthers (cf. Bittrich et al. 1993). In P O L L E N M O R P H O L O G Y . Pollen is tricolporate
Perissocarpa, the inner lobes of the apically bilobed (rarely tetracolporate), the exine is laevigate
petals stick firmly together and the thereby formed (Luxemburgioideae and part of the Sauvage-
cap-like structure is thought to protect the anthers sioideae s.str.) or variously sculpted. A striate-
of the buzz-pollinated flowers from the permanent rugulate exine characterizes the Ochnoideae and
moisture in the natural habitat (Walln€ofer 1998). most Sauvagesioideae s.str., which have petaloid
The wax covering the anthers of some species (Wal- staminodes. The presence of a baculate exine
lacea, Blasthemanthus) seems to have rather a pro- suggests sister-group relationship of the former
tective function. Wax-collecting bees were observed genera Neckia and Indovethia, today both included
much more rarely than pollen-collecting bees and in Sauvagesia (Amaral 1991).
they were less effective as pollinators (Poecilandra
sp., pers. obs.). Nectaries are apparently always K A R Y O L O G Y . Nearly all chromosome counts
absent. The report of a sweet mucilaginous secre- reported are from members of the Ochnoideae:
tion in the flowers (from sepal colleters?) of Rhyti- Idertia: n ¼ 12, Rhabdophyllum: n ¼ 12, Campy-
danthera mellifera attracting bees (Schultes 1949) lospermum: n ¼ 10, 12, 24, Ouratea: n ¼ 13,
begs for further investigations. Reports that the Ochna: n ¼ 12, 14, Lophira: n ¼ 14, 2n ¼ 24.
flowers of Lophira lanceolata are a source of The only count of a member of Sauvagesioideae
honey in Nigeria (Mapongmetsem 2007) also need s.str., Sauvagesia erecta, was reported with
confirmation. Self-compatibility (in the weedy 2n ¼ 38. It seem thus probable that the basic
Ochnaceae 259
chromosome number for the Ochnoideae is vegetation. Anemochory must also be assumed for
n ¼ 12 but, in the absence of more counts from the samaroid fruits of Lophira spp., which form
the Luxemburgioideae and Sauvagesioideae s.str., large trees of rainforests or medium-sized trees in
currently no further conclusions can be drawn. savannas. Species which occur in inundated forests
(igapós or várzeas) show adaptations to hydro-
F R U I T A N D S E E D . Many-seeded capsules are chory: the epidermis of the seeds of Wallacea is
common in the Luxemburgioideae and Sauvage- formed by large, dead, air-filled cells. The indehis-
sioideae s.str. These are always septicidal, but the cent fruits of Elvasia elvasioides have an aeren-
dehiscence may start apically (e.g. Sauvagesia, chyma and are hydrochorous (pers. obs.). The
Luxemburgia, Fig. 60I) or basally (e.g. Cespedesia, white or red drupes of Euthemis contrasting with
Philacra, Fig. 60J). A columella is present or the dark red sepals (Kanis 1971) and the blackish
absent. In Wallacea the two or three valves sepa- drupelets inserted on an accrescent red-coloured
rate completely. Dry indehiscent monospermous receptacle of the group with apparently apocarpous
fruits occur in Elvasia (Fig. 59H), Perissocarpa gynoecia are ornithochorous. Species character-
(nut-like) and Lophira (samaroid, Fig. 59F). Dru- ized by ornithochory occur commonly in more or
paceous fruits with one or two seeds per locule less closed forests, but many species of Ouratea
occur in Euthemis while, in all genera of Ochnoi- occur in savannah-like vegetation.
deae characterized by deeply lobed and for their
major parts apocarpous ovaries (see above), the P H Y T O C H E M I S T R Y . Relevant data are reported in
carpels separate completely during fruit develop- Hegnauer (1969, 1990) and Mbing et al. (2003).
ment ("ecological apocarpy", Baum 1951), form- Condensed tannins are present, ellagi- and gallotan-
ing monospermous drupelets inserted on the nins absent. C-glycoflavones, flavones, biflavones
accrescent fleshy receptacle (Fig. 59E). and other biflavonoids (lophirone and related sub-
The wings of the seeds of many capsular fruits stances in subf. Ochnoideae) were more or less
develop from the outer epidermis of the outer regularly found, triterpenes (Ouratea) and alkaloids
integument. These wings may be restricted to one are rare (Testulea, Lophira). The pericarp of species
(e.g. Luxemburgia) or both ends (e.g. Schuurman- of Ochna and Ouratea and of the seeds of Ochna and
siella) of the seeds or surround the seed completely Lophira are rich in fat oil (see Mapongmetsem 2007
(e.g. Poecilandra). Wings can be much longer than for information about the fatty acid composition of
the seed body (e.g. Cespedesia, Schuurmansia) or the seed oil of L. lanceolata).
much shorter (e.g. Luxemburgia, Schuurman-
siella). Endosperm is absent in the ripe seeds of SUBDIVISION AND RELATIONSHIPS WITHIN THE
Ochnoideae, and present in those of Luxemburgio- F A M I L Y . The subdivision of the Ochnaceae into
ideae and Sauvagesioideae s.str. The cuticle of two monophyletic subfamilies, as already proposed
the testa may be smooth or variously sculpted by Engler (1874), seemed to be well-supported by
(Amaral 1991). The cells of the inner epidermis of several morpho-anatomical characters (Amaral
the outer integument in Luxemburgioideae and 1991). Chloroplast DNA sequence data showed,
almost all Sauvagesioideae contain small crystals. however, that Luxemburgia and Philacra form a
The embryo is straight or curved, isocotylar or basal clade within the family, thus rendering the
rarely heterocotylar. Sauvagesioideae in the traditional circumscription
According to Farron (1985) and Sastre (1975b), paraphyletic (Amaral et al., unpubl. data; Wurdack
germination is epigeal (Sauvagesia, Ouratea, Rhab- and Davis 2009). Consequently, these two genera
dophyllum spp., Campylospermum spp.) or hypo- are transferred to subfamily Luxemburgioideae in
geal (Elvasia, Ochna, Idertia, Campylospermum the present treatment. It is unclear at present which
spp., Rhabdophyllum spp.). morpho-anatomical characters support the clade
of Ochnoideae + Sauvagesioideae s.str. and the
D I S P E R S A L . In several genera with capsular fruits subclade of Sauvagesioideae s.str. Even the detailed
the seeds are provided with wings of various form analysis of flower structure by Matthews et al.
and size. The species with such seeds, which are (2012) did not reveal possible synapomorphies of
probably wind-dispersed, occur generally in open these clades. Lack of knowledge about the
infrafamilial phylogeny currently impedes safe logenetic analyses based on DNA sequence data
conclusions. Thus, possibly the striate-rugulate clearly support the inclusion of the Ochnaceae in
pollen surface is synapomorphic for the former a clade with Medusagynaceae and Quiinaceae, the
clade, but it may also have evolved independently "ochnoids", in which Quiinaceae and Medusagy-
in both subfamilies. The exclusion of Lophira from naceae are sister taxa and Ochnaceae the sister
the family, as suggested by Takhtajan (1997), is group to the clade formed by both (Schneider
supported neither by the cladistic analysis of et al. 2005; Wurdack and Davis 2009; Soltis et al.
Amaral (1991), as the genus shares various derived 2011; Xi et al. 2012). This clade is also morpho-
characters with other members of subfamily Och- logically characterized by stratified phloem,
noideae, nor by molecular data (Wurdack and mucilage cells, contorted petal aestivation, lack
Davis 2009). Within the Ochnoideae the genera of nectaries, and tenuinucellate ovules (Stevens
with a more or less apocarpous gynoecium most 2001 onwards), and several floral characters sup-
probably form a monophyletic subgroup. The port the sister-group relationship of Quiinaceae
relationship of this group to the other genera of and Medusagynaceae (Matthews et al. 2012). APG
this subfamily is still unsettled, however, so that a II (2003) suggested the inclusion of all three
further subdivision of the Ochnoideae is pre- families in an expanded family Ochnaceae as
mature. Also a subdivision of the Sauvagesioideae optional and APG III (2009) prefers this lumping,
s.str. in monophyletic tribes is still not possible, but we see little advantage of it, besides the risk of
although some monophyletic groups within the confounding non-specialists. The phylogeny of Xi
subfamily are well supported by the cladistic anal- et al. (2012) shows the clusioids as the moderately
ysis of Amaral (1991). supported sister of the ochnoids.
A F F I N I T I E S . For some time the Ochnaceae were D I S T R I B U T I O N A N D H A B I T A T S . The Ochnaceae

generally but not unanimously accepted as part of have a pantropical distribution. Fourteen genera
the Theales. Guédès and Sastre (1981) proposed are restricted to the New World. Seven genera
its inclusion together with the major part of the occur in the Indo-Malaysian region and nine
Theales in the Violales. Based on the presence of genera in tropical Africa; three of these, Ochna,
cristarque cells, petiole anatomy and trilacunar Brackenridgea and Sauvagesia, are common to
nodes, Schofield (1968) considered Quiinaceae as both regions. Only one genus, Sauvagesia, occurs
the most closely related family. A few taxono- in all three tropical regions. The weedy Sauvagesia
mists believed that Ochnoideae and Sauvagesioi- erecta occurs in the New World and, probably due
deae are only distantly related and belong to to human transport, in Africa. Central America is
different parts of the dicotyledons. Thus, Corner poor in genera (mainly species of Ouratea and
(1976) in the tradition of Bentham and Hooker Cespedesia spathulata); possibly this region was
(1862) suggested that the Ochnoideae are closely colonized via long-distance dispersal by birds or
related to the Simaroubaceae while the Sauvage- only after the formation of the Isthmus of Panama.
sioideae would be close to the Violaceae. This Vicariant distribution patterns of Ochnaceae in
suggestion was based on a superficial character the Guyanas were studied by Sastre (1992).
analysis (see Eichler 1878; Amaral 1991), how- The majority of the species occurs in open
ever, and not supported by new data. There are vegetation, mainly savannas; a few are pioneers
numerous differences between the subfamilies, (Cespedesia spathulatha, Schuurmansia). Some
including the fact the androecial development in genera with hydrochorous fruits or seeds are
the Ochnoideae and the other two subfamilies known from inundated forests of northern South
seems to be different. The monophyly of the America. Lophira alata is a member of the upper
Ochnaceae, however, is today settled. Possible canopy in lowland rainforests. Species of Ochnoi-
synapomorphies of the family are the poricidal deae with bird-dispersed drupelets occur in the
anthers (reversed in Brackenridgea spp., Ochna understories of forests and in savannas.
spp., Sauvagesia spp., Schuurmansia, Schuur-
mansiella) and the crystal layer in the endotesta E C O N O M I C I M P O R T A N C E . Only Lophira alata
(reversed in the Ochnoideae probably due to the has some economic importance. The hard and
development of indehiscent fruits). Recent phy- durable wood is used for construction purposes.
Ochnaceae 261
The seed oil is used for cooking and soap produc- 10. Petals shortly connate at the base; apex of the
tion, also in L. lanceolata (Mapongmetsem 2007). anthers beaked; locules bi-ovulate; fruits drupa-
The South African Ochna serrulata (Hochst.) ceous, 5-locular 21. Euthemis
Walp. is widely cultivated in gardens in the tropics. – Petals distinct; apex of the anthers not beaked;
locules with 4–1 ovules; fruits 1–5-locular, capsu-
lar or indehiscent 11
KEY TO THE GENERA 11. Outer sepals longer than inner sepals; fruits
1. Styles gynobasic; carpels apparently distinct; indehiscent, winged by the accrescent two
receptacle enlarging during fruit development, outer sepals, 1-seeded 11. Lophira
often becoming red 2 – Outer sepals smaller than inner sepals or of
– Styles apical; carpels completely connate; recepta- equal length; capsules with several to many
cle not enlarging during fruit development 7 seeds 12
2. Anthers usually opening by longitudinal slits; 12. Carpels 5 13
stipules laciniate or striate; drupelets with an – Carpels 2–3 17
internal projection of the endocarp 13. Leaf apex emarginate; secondary veins closely
4. Brackenridgea parallel; marginal teeth absent; stamens 5
– Anthers opening by pores, or if by longitudinal 17. Fleurydora
slits, stipules entire and not striate and drupe- – Leaf apex obtuse; secondary veins separated by
lets without internal projection of the endocarp 7–11 areoles; marginal teeth present; stamens
3 10–1 14
3. Stamens 12–1, filament length at least 1/3 of the 14. Stipules deeply bifid, persistent; sepals basally
length of the anther 3. Ochna connate, much smaller than the flower bud
– Stamens 10, anthers sessile or subsessile 4 15. Cespedesia
4. Sepals persistent; embryo curved 5 – Stipules entire, caducous; sepals distinct, at least
– Sepals caducous or rarely persistent; embryo half as long as the flower bud 15
straight 6 15. Leaves compound; stamens 1; flowers white
5. Leaves with more than 30 secondary veins; 13. Rhytidanthera
petals as long as the sepals 7. Rhabdophyllum – Leaves simple; stamens 10; flowers yellow 16
– Leaves generally with ca. 10–25 secondary veins; 16. Sepals enclosing the mature flower bud, centri-
petals longer than sepals 8. Campylospermum petally becoming larger, adaxial with clavate
6. Leaf margin without bristles; flowers in terminal hairs at the base; anthers biporate 12. Godoya
or rarely axillary generally multi-flowered – Sepals not enclosing the mature flower bud, all of
inflorescences; sepals generally caducous (Neo- the same size; clavate hairs 0; anthers uniporate
tropics) 5. Ouratea 14. Krukoviella
– Leaf margin with persistent bristles; flowers in 17. Stamens 1, rarely 4–10, located adaxially already
1–4-flowered axillary inflorescences; sepals per- in bud 18
sistent (Africa) 6. Idertia – Stamens 5 or 10, regularly distributed around
7. Locules uniovulate; fruits indehiscent, dry, not the ovary or moving only during anthesis to the
winged 8 adaxial side 19
– Locules with 2–1 ovules; fruits capsular, drupa- 18. All secondary veins ending in marginal teeth;
ceous, or samaroid 9 anther apex straight; capsule dehiscence starting
8. Sepals distinct; petals yellow, patent; carpels 2–7 at the apex (Brazil south of the Amazon)
9. Elvasia 1. Luxemburgia
– Sepals distinct or connate for most of their – Only part of the secondary veins ending in mar-
length; petals white or cream-yellow, sticking ginal teeth; anther apex recurved; capsule dehis-
together apically and forming a cap-like struc- cence starting at the base (northern Brazil and
ture; carpels 2(3) 10. Perissocarpa southern Venezuela) 2. Philacra
9. Flowers tetramerous; stamen 1; leaf nervation 19. Sepals 10–15; stamens 10; fruits with 1–2 seed
brochidodromous 16. Testulea per locule 18. Blastemanthus
– Flowers pentamerous; stamens (4–)5–1; leaf – Sepals and stamens 5; locules of fruit many-
nervation craspedodromous 10 seeded 20
20. Flowers becoming zygomorphic during anthesis 1. Luxemburgia A. St.-Hil. Fig. 59H, I
21 Luxemburgia A. St.-Hil., Mém. Mus. Hist. Nat. 9: 352
– Flowers actinomorphic 22 (1822).
21. Petals yellow; placentation parietal; capsule
opening apically by 3 valves; seed winged Shrubs or small trees. Leaves with closely paral-
19. Poecilandra lel secondary veins all ending in marginal,
– Petals white to pink; placentation laminar; cap- sometimes ciliate teeth; stipules deeply
sules splitting completely into 2–3 valves; seed not 3-partite, ciliate. Sepals 5(6); petals 5(6), yellow,
winged 20. Wallacea imbricate; stamens numerous, filaments con-
22. Capsule dehiscence starting at the base; seeds nate at base, persistent, anthers partially con-
with long wings at both ends nate, opening by 2 apical pores; staminodes 0;
22. Schuurmansia ovary 3-carpellate, 1-loculate above, placenta-
– Capsule dehiscence starting at the apex; seeds tion parietal. Capsule with many shortly winged
shortly winged or unwinged 23 seeds.
23. Staminodes filiform; seeds short-winged at both 18 spp., mountainous regions of SE, NE and
ends 23. Schuurmansiella central Brazil.
– Staminodes spathulate, petaloid; seeds on the
whole surface with short irregular wing-like
folds or unwinged 24 2. Philacra Dwyer Fig. 59J
24. Staminodes keeled; seeds winged 24. Tyleria Philacra Dwyer, Brittonia 5: 124 (1944); Sastre, Flora
– Staminodes not keeled; seeds unwinged 25 Venez. Guayana 7: 146–148 (2003).
25. Stipules auriculate with marginal glands; testa
finely tuberculate 25. Adenarake Small trees or shrubs. Leaves with closely parallel
– Stipules triangular, margins ciliate or with long secondary veins with fewer persistent marginal
bristles; testa not tuberculate 26 teeth; stipules persistent. Sepals 5; petals 5, imbri-
26. Stipules shortly ciliate; seeds reniform, longitudi- cate, yellow; stamens numerous, rarely 4–10; fila-
nally canaliculate, testa cells rectangular; carpels ments very short, connate, persistent, anthers
2 26. Indosinia partially connate or distinct, strongly curved at
– Stipules with long bristles; seeds globose or their apex and opening by two pores; staminodes,
ovate, not canaliculate; testa cells hexagonal; if present, very small, broadened at the apex,
carpels (2)3 27. Sauvagesia possibly glandular; ovary 3-carpellate, unilocu-
late above, placentation axile to parietal. Capsule
with many, shortly winged seeds.
GENERA OF OCHNACEAE Four spp. in Venezuela and northern Brazil.
I. S U B F A M . L U X E M B U R G I O I D E A E Planch. ex Endl.
(1850) (’Luxemburgieae’). II. SUBFAM. OCHNOIDEAE Burnett, Outl. Bot.: 886,
1093, 1125 (1835) (‘Ochnidae’).
Pitting between vessel elements and parenchyma Pitting between vessel elements and parenchyma
generally unilaterally compound. Flowers not unilaterally compound. Flowers actinomor-
obliquely zygomorphic already in bud with the phic, filaments distinct, staminodes 0, pollen stri-
stamens surrounding the ovary only adaxially; ate/rugulate. Ripe seeds without endosperm, testa
filaments fused congenitally at least at the base, poorly differentiated, unitegmic.
anthers partially fused postgenitally or rarely dis-
tinct, staminodes 0 or very small (in adaxial posi-
tion outside the fertile stamens); pollen laevigate. 3. Ochna L. Fig. 59E
Seeds albuminous, shortly winged, endotesta with Ochna L., Sp. Pl.: 513 (1753); Kanis, Blumea 16: 22–40
small crystalliferous cells. (1968); Robson, Fl. Zambes. 2(1): 225–251 (1963).
Ochnaceae 263
Trees, shrubs or shrublets. Leaves with serrate, rarely setaceous and persistent. Sepals 2–5, mostly
ciliate or entire margins; stipules entire or not, caducous; petals 5, contort, generally yellow;
caducous. Sepals (4)5, persistent, enlarging and stamens 10, filaments very short, anthers trans-
becoming coloured in fruit; petals 5(–12), conversely wrinkled or rarely smooth, poricidal; ovary
tort, mostly yellow; stamens numerous, distinct, 5–10-carpellate, on a short broad gynophore. Fruit
with persistent filaments; anthers dehiscing by separating into 1–10 black drupelets on a red
longitudinal slits or terminal pores; ovary deeply accrescent receptacle. Embryo straight. n ¼ 13.
divided into (3–)5–10(–15) uniovulate lobes with About 200 spp. in tropical or subtropical
gynobasic style. Fruit separated into one to sev- regions of the New World. The separation of
eral black drupelets on an accrescent reddish Campylospermum, Idertia and Rhabdophyllum
receptacle. Embryo straight or curved. n ¼ 12, 14. from Ouratea is dubious and needs further inves-
About 85 spp. in the tropics and subtropics of tigation.
Africa and Asia. The separation of Brackenridgea is
dubious, and possibly makes Ochna paraphyletic.
6. Idertia Farron
4. Brackenridgea A. Gray Idertia Farron, Ber. Schweiz. Bot. Ges. 73: 212 (1963).
Brackenridgea A. Gray, Proc. Amer. Acad. Arts 3: 51. 1853
(sero) (1853); U.S. Expl. Exped. Bot. Phan. 1: 361, t. 42 Small trees or shrubs. Leaf margin with persistent
(1854); Kanis, Fl. Males. I, 7: 101–104 (1971); Robson, Fl. bristles; stipules membranaceous, intrapetiolarly
Zambes. 2(1): 252–255 (1963). united at base. Inflorescences axillary, short,
1–4-flowered; sepals 5, accrescent and red in fruit;
Trees, shrubs or shrublets often with yellow pig-
petals 5, contort, yellow; stamens 10, anthers
ment under the bark. Leaves with entire or glan-
smooth, subsessile, poricidal; ovary deeply 5–6-
dular serrulate margin; stipules and bracts
lobed with completely connate gynobasic style.
longitudinally striate, laciniate, persistent on the
Fruit separating into 1–6 black drupelets on
first year shoots. Sepals (4)5, white or pink in
an accrescent red receptacle. Embryo straight.
flower, accrescent and red in fruit; petals (4)5,
n ¼ 12.
contort, white to pink, rarely yellow; stamens
Four spp. in West Africa, São Tomé.
usually (8–)13–20(–22), anthers usually dehiscing
by longitudinal slits; ovary deeply divided into (3)
5–10 uniovulate lobes, style gynobasic. Fruit
separated into one to several black drupelets on 7. Rhabdophyllum Tiegh.
an accrescent red receptacle. Seeds with an intru- Rhabdophyllum Tiegh., J. Bot. (Morot) 16 17: 201 (1902);
sion of the endocarp, embryo curved. Sosef, Andansonia III, 30: 119–135 (2008), rev.
About seven spp. in tropical Africa, Madagas-
Small trees or subshrubs. Leaf secondary veins
car, Malaysia, and the Philippines.
numerous, dense, margin sometimes toothed;
stipules intrapetiolarly united at base, caducous
5. Ouratea Aubl. Fig. 59A–D or persistent. Inflorescences axillary, many-
Ouratea Aubl., Hist. Pl. Guiane: 397 (1775), nom. cons.; flowered. Sepals 5, accrescent and reddish in
Sastre, Adansonia III, 1: 47–67 (1988); Maguire & Steyer- fruit; petals 5, yellow, contort, of the same length
mark, Mem. N.Y. Bot. Gard. 51: 56–102 (1989); Sastre, as the sepals; stamens 10, anthers subsessile,
Flora Venez. Guayana 7: 131–143 (2003). transversally wrinkled, poricidal; ovary deeply
Gomphia Schreb., Gen. Pl. ed. 8, 1: 291 (1789); Bittrich & 5-lobed with completely connate gynobasic
Amaral, Taxon 43: 89–93 (1994), nomencl.
style. Fruit separating into 1–5 brown or black
Trees or shrubs, glabrous or rarely with hairs. Leaf drupelets on an accrescent receptacle. Embryo
margin entire, serrate to ciliate, secondary veins curved. n ¼ 12.
generally curving strongly near the margin; sti- Eight spp. in tropical moist forests of West
pules entire, distinct, scale-like and caducous, or Africa.
8. Campylospermum Tiegh. 5(6), persistent, anthers opening by 2 apical

Campylospermum Tiegh., J. Bot. (Morot) 16: 35, 40 (1902). pores; ovary with 2(3) uniovulate locules, placen-
tation axile. Fruit 1-seeded, globular, indehiscent.
Small trees, shrubs or subshrubs. Leaves entire or Three spp. in Venezuela, Peru, and northern
with caducous teeth; stipules intrapetiolarly united Brazil, in mountain forests and savannas.
at base. Inflorescence terminal (axillary). Sepals 5,
accrescent and yellow or reddish in fruit; petals 5,
contort, longer than the sepals; stamens 10, anthers 11. Lophira Banks ex C.F. Gaertn. Fig. 59F
sessile or subsessile, often transversely wrinkled, Lophira Banks ex C.F. Gaertn., Suppl. carp.: 52 (1805).
poricidal; ovary deeply 5-lobed with completely con- Small to large deciduous trees up to 50 m. Leaves
nate gynobasic style. Fruit separating into 1–5 black with closely parallel secondary nerves. Flowers
drupelets on a red accrescent receptacle. Embryo actinomorphic; sepals 5, imbricate, persistent;
curved. n ¼ 10, 12, 24. petals 5, contort, white; stamens numerous,
About 65 spp. in the Old World, the majority anthers opening with 2 subapical pores; ovary
in tropical Africa, Madagascar, Ceylon to SE Asia. 2-carpellate, ovules numerous, placentation basal.
Fruit 1-seeded, winged by the two outer accres-
cent sepals. n ¼ 14, 2n ¼ 24.
9. Elvasia DC. Fig. 59G, H
Two vicariant spp. in lowland rainforests and
Elvasia DC., Ann. Mus. Natl. Hist. Nat. 17: 422, t. 20 savannas of West and Central Africa to Sudan.
(1811); Sastre, Flora Venez. Guayana 7: 129–130 (2003). Lophira alata Banks ex C.F. Gaertn. is the source
Small trees or shrubs, glabrous or with indument on of excellent heavy timber. The seeds of both species
young shoot. Leaves with closely parallel secondary are used for the extraction of an edible oil.
nerves and caducous marginal teeth; stipules del-
toid, persistent. Sepals 2–6, caducous; petals 3–8, III. S U B F A M . S A U V A G E S I O I D E A E Beilschm. (1833)
imbricate, yellow; stamens 5–20, anthers opening (‘Sauvagesieae’).
by two apical pores; ovary on a short broad gyno-
phore, basically unilocular, but strongly 2–5(–7)-
Pitting between vessel elements and parenchyma
lobed with one ovule per carpel; placentation
generally unilaterally compound. Flowers zygo-
median and axile at the ovary base. Fruits indehis-
morphic as regards androecium and ovary (in
cent, coriaceous or woody, globular or star-shaped,
most cases the zygomorphy developing only dur-
1(2)-seeded.
ing anthesis) or actinomorphic; stamen filaments
About 15 spp. in South America, mainly
distinct, staminodes often present, sometimes 2 dif-
Guayana Highland and Amazonia, one sp. in NE
ferent types (filamentose and petaloid) in the same
Brazil, and one in Central America.
flower. Seeds albuminous, often winged, endotesta
generally with small crystalliferous cells.
10. Perissocarpa Steyerm. & Maguire
Perissocarpa, Steyerm. & Maguire, Ann. Missouri Bot.
Gard. 71: 319 (1984); Walln€ofer, Ann. Naturhist. Mus. 12. Godoya Ruiz & Pav.
Wien 100B: 683–707 (1998), rev. Godoya Ruiz & Pav., Fl. Peruv. Prodr. 1: 58 (1794).
Small trees or shrubs. Leaves with closely parallel Trees or shrubs. Leaves with distant secondary
secondary nerves and caducous marginal teeth; nerves, margin strongly crenate and toothed; sti-
stipules deltoid, caducous or persistent. Sepals 5, pules caducous. Flowers becoming zygomorphic
distinct or connate for most of their length, cadu- during anthesis; sepals 5, adaxially with long cla-
cous; petals 5, contort, cream-yellow, white or vate hairs at base; petals 5, imbricate, yellow,
greenish-white, basally distinct, bilobed with the inside basally with long clavate hairs; staminodes
inner lobes inflexed and sticking firmly together, 0, stamens 10, anthers opening by 2 apical pores;
corolla thus forming a cap-like structure; stamens ovary with 5 locules and numerous ovules,
Ochnaceae 265
placentation axile. Capsule with numerous One to six spp. from Nicaragua to Bolivia and
winged seeds. central Brazil.
Two spp., north-western South America.
16. Testulea Pellegr.
13. Rhytidanthera Tiegh.
Testulea Pellegr., Bull. Soc. Bot. France 71: 76 (1924).
Rhytidanthera Tiegh., Ann. Sci. Nat. Bot. VIII, 19: 43
(1904). Trees. Leaves with distant secondary nerves, with-
out marginal teeth; stipules intrapetiolarly con-
Trees or shrubs. Leaves 4–5-pinnate with distant nate, persistent. Flowers zygomorphic; sepals 4,
secondary nerves and marginal teeth; stipules petals 4, imbricate, white to pink; staminodes
caducous. Flowers becoming zygomorphic during filamentose, connate for 2/3 of their length; fertile
anthesis; sepals 5, adaxially with glandular hairs stamen 1, adaxial, anther opening by two apical
(colleters?) at base; petals 5, contort, white; stami- pores; ovary bicarpellate, unilocular in the upper
nodes 0, stamens numerous, anthers opening by part, placentation axile to parietal. Capsule with
2 subapical pores; ovary 5-loculate, pluriovulate, numerous winged seeds.
placentation axile. Capsule with numerous winged One sp., T. gabonensis Pellegr., Gabon. This
seeds. genus shows several aberrant characters and
Five spp. in Colombia and Venezuela. deserves further investigation.
14. Krukoviella A.C. Sm.

17. Fleurydora A. Chev.
Krukoviella A.C. Sm., J. Arnold Arbor. 20: 295 (1939).
Fleurydora A. Chev., Bull. Mus. Hist. Nat. (Paris) II, 5: 158
Scandent shrubs. Leaves with distant secondary (1933).
nerves and marginal teeth; stipules caducous. Flow-
Small trees or shrubs. Leaves with closely parallel
ers becoming zygomorphic during anthesis; sepals
secondary veins, without marginal teeth; stipules
5, caducous; petals 5, contort, yellow; staminodes 0,
caducous, margins glandular. Flowers becoming
stamens 10, anthers opening by 1 apical pore; ovary
zygomorphic during anthesis; sepals 5; petals 5,
5-loculate, pluriovulate, placentation axile. Capsule
imbricate, yellow; staminodes 0, stamens 5,
with numerous winged seeds.
anthers opening by 2 subapical pores; ovary
One sp., K. disticha (Tiegh.) Dwyer, in western
with 5 locules, placentation axile. Capsules with
Amazonia.
numerous winged seeds.
One sp., F. felicis A. Chev., Guinea.
15. Cespedesia Goudot
Cespedesia Goudot, Ann. Sci. Nat. Bot. III, 2: 368 (1844);
Sastre, Cespedesia 4: 191–214 (1975). 18. Blastemanthus Planch. Fig. 60G
Blastemanthus Planch., London J. Bot. 5: 644 (1846).
Trees. Leaves up to 1 m long with distant second-
ary nerves and marginal teeth; stipules persistent, Small trees or shrubs. Leaves with closely parallel
inside with glandular hairs (colleters?) at base. secondary veins and caducous marginal teeth;
Flowers becoming zygomorphic during anthesis; stipules persistent. Flowers becoming zygomor-
sepals 5, small, connate at base; petals 5, contort, phic during anthesis; sepals 10–15, caducous;
yellow, shortly connate at base; staminodes 0, petals 5, imbricate, yellow; staminodes numerous,
stamens numerous, anthers covered with wax linear; stamens 10, anthers covered with wax crys-
crystals, opening by 2 subapical pores; ovary tals, opening with one pore; placentation axile to
5-carpellate, apically incompletely septate, placen- parietal; ovary tricarpellate. Capsule with 1–2
tation axile to parietal. Capsule with numerous unwinged seeds per locule.
winged seeds. About three spp., northern South America.
19. Poecilandra Tul. 22. Schuurmansia Blume

Poecilandra Tul., Ann. Sci. Nat. Bot. III, 8: 342 (1847); Schuurmansia Blume, Mus. Bot. 1: 177 (1850); Kanis,
Sastre, Flora Venez. Guayana 7: 148–150 (2003). Blumea 16: 74–80 (1968), rev.
Shrubs or small trees. Leaves with closely parallel Trees or treelets. Leaves up to 85 cm long, with
secondary veins, caducous marginal teeth, and closely parallel secondary veins, persistent or
caducous stipules. Flowers becoming zygomor- caducous marginal teeth, and caducous stipules.
phic during anthesis; sepals 5, imbricate, cadu- Flowers actinomorphic, bisexual or unisexual;
cous; petals 5, contort, yellow; staminodes sepals 5, imbricate, persistent or caducous; petals
numerous, linear or spathulate; stamens 5, 5, contort, white to pink; staminodes petaloid or
anthers covered with wax crystals, opening by linear, distinct or connate at base, persistent;
one pore; ovary tricarpellate with parietal placen- stamens 5, persistent, anthers opening by slits;
tation. Capsule with numerous winged seeds. ovary 3-carpellate with parietal placentation.
Three spp., northern South America. Capsule opening from the base, seeds numerous
with long wings at both ends.
Three spp., New Guinea to the Philippines.
20. Wallacea Spruce ex Benth. & Hook. f.
Wallacea Spruce ex Benth. & Hook. f., Gen. Pl. 1: 320
(1862); Sastre, Flora Venez. Guayana 7: 160–161 (2003). 23. Schuurmansiella Hallier f.
Small trees. Leaves with closely parallel secondary Schuurmansiella Hallier f., Recueil Trav. Bot. Néerl. 10: 344
(1913); Kanis, Blumea 16: 73–74 (1968), rev.
veins, caducous marginal teeth, and caducous
stipules. Flowers becoming zygomorphic during Small trees or shrubs. Leaves with closely parallel
anthesis; sepals 5, imbricate, caducous; petals 5, secondary veins, persistent marginal teeth and per-
contort, white or pink; staminodes numerous, sistent stipules. Flowers actinomorphic, unisexual;
linear or 0; stamens 5, anthers covered with wax sepals 5, valvate, persistent; petals 5, white to pink,
crystals, opening by longitudinal slits; ovary 2–3- contort; staminodes numerous, linear, connate
carpellate with laminar placentation. Capsule at base, persistent; stamens 5, persistent, anthers
splitting completely into 2–3 valves; seeds numer- opening by apical slits; ovary (2)3-carpellate,
ous, unwinged. placentation parietal. Capsule with numerous
Two spp., northern South America. seeds shortly winged on both ends.
One sp., S. angustifolia (Hook. f.) Hallier f., in
NW Borneo.
21. Euthemis Jack
Euthemis Jack, Malayan Misc. 1(5): 15 (1820); Kanis, Blu-
mea 16: 62–66 (1968), rev. 24. Tyleria Gleason
Shrubs. Leaves with closely parallel secondary Tyleria Gleason, Bull. Torrey Bot. Club. 58: 391 (1931);
Sastre, Flora Venez. Guayana 7: 125, 157–160 (2003).
veins, persistent marginal teeth, and caducous
Adenanthe Maguire, Steyerm. & Wurdack (1961).
stipules. Flowers actinomorphic, bisexual or uni-
sexual; sepals 5, imbricate, persistent, purplish- Trees or shrubs. Leaves with closely parallel
red in fruit; petals 5, connate at base, white or secondary veins, marginal teeth or bristles, and
pink; staminodes 5, linear or absent; stamens 5, persistent or caducous stipules. Flowers actino-
anthers beaked, opening by one pore; ovary morphic; sepals 5, imbricate or valvate, persistent;
5-carpellate, placentation axile with two ovules petals 5, contort, white to pink; staminodes 10,
per locule. Fruit drupaceous, white or red, with dimorphic, those opposite the sepals often lacini-
1(2) seeds per locule. ate, petaloid, basally generally connate, those
Two spp., southeast Asia. opposite the petals keeled adaxially; stamens 5,
Ochnaceae 267
persistent; anthers opening by slits or pores; ovary Vausagesia Baill. (1890).

(2)3(5)-carpellate, with parietal placentation. Cap- Indovethia Boerl. (1894).
sule with numerous short-winged seeds. Sinia Diels (1930).
13 spp., Guayana Highland. Pentaspatella Gleason (1931).
Roraimanthus Gleason (1933).
Treelets, shrubs or herbs, rarely small trees.
25. Adenarake Maguire & Wurdack Leaves with closely parallel secondary veins, per-
Adenarake Maguire & Wurdack, Mem. New York Bot. sistent teeth, and persistent, pectinate stipules.
Gard. 10: 15 (1961). Flowers actinomorphic, bisexual or very rarely
Shrubs. Leaves with closely parallel secondary unisexual; sepals (4)5, imbricate or valvate, some-
veins and persistent marginal teeth; stipules times glandulose-ciliate, persistent; petals (4)5,
auriculate with marginal glands, caducous. Flow- contort, white to pink; 5(10) petaloid staminodes
ers actinomorphic with conspicuous androgyno- generally enclosing the stamens, sometimes con-
phore; sepals 5, valvate, with marginal glands; nate (Lauradia), outside of these 1–2 series of
petals 5, contort, white to pink; staminodes 15, smaller staminodes sometimes present; stamens
the inner 5 larger, enclosing the 5 stamens; 5, persistent; ovary (2)3-carpellate, placentation
anthers opening by apical slits; ovary tricarpel- parietal to basal. Capsule with many unwinged,
late, placentation axile. Capsule with numerous alveolate seeds. 2n ¼ 38.
unwinged, tuberculate seeds. One sp. endemic in SE China, two spp.
Two spp., northern Brazil and southern Vene- endemic in Malaysia, one sp. endemic in Africa,
zuela, at higher elevations. and ca. 35 spp. in the Neotropics; S. erecta L. is a
pantropical weed.
26. Indosinia J.E. Vidal

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Kanis, Blumea 16: 68–69 (1968), rev.
Amaral, M.C.E. 1991. Phylogenetische Systematik der
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de varredura. Rev. brasil. Bot. 21: 269–273.
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Kubitzki, K., Amaral. M.C.E. 1991. Transference of function Soltis, D.E., Soltis, P.S., Chase, M.W., Mort, M.E., Albach,
in the pollination system of the Ochnaceae. Pl. Syst. D.C., Zanis, M. and 10 further authors. 2000. Angio-
Evol. 177: 77–80. sperm phylogeny inferred from 18S rDNA, rbcL, and
Mapongmetsem, P.-M. 2007. Lophira lanceolata Tiegh. atpB sequences. Bot. J. Linnean Soc. 133: 381–461.
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Matthews, M.L. et al. 2012. See General References. Stevens, P.F. 2001 onwards. See General References.
Mbing, J.N., Bassomo, M.Y., Pegnyemb, D.E., Tih, R.G., Takhtajan, A. 1997. See General References.
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Metcalfe, C.R., Chalk, L. 1950. See General References. 266–273.
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gesia L. (Ochnaceae). Rev. Brasil. Bot. 28: 255–265. Walln€ ofer, B. 1998. A revision of Perissocarpa Steyerm. &
Narayana, L.L. 1975. A contribution to the floral anatomy Maguire (Ochnaceae). Ann. Naturhist. Mus. Wien
and embryology of Ochnaceae. J. Jap. Bot. 50: 329–336. 100B: 683–707.
Oliveira, P.S., Leitão Filho, H.F. 1987. Extrafloral nectaries: Wurdack, K.J., Davis, C.C. 2009. See General References.
their taxonomic distribution and abundance in the Xi, Z. et al. 2012. See General References.
Pandaceae
Pandaceae Engl. & Gilg (1913), nom. cons.
K. K U B I T Z K I
Dioecious trees or shrubs; indumentum simple. mainly diffuse or in uniseriate apotracheal

Leaves alternate, simple, entire or dentate, pin- bands (Metcalfe and Parameswaran in Forman
nately veined, eglandular; stipules small, inserted 1966; Hayden and Hayden 2000).
at different levels on the axis, generally persistent.
Inflorescences terminal or cauliflorous and SHOOT MORPHOLOGY INCLUDING INFLORES-
pseudoracemose-thyrsiform, or axillary and CENCES. A very conspicuous feature of the fam-
fasciculate, or flowers solitary; bracts minute. ily are the dorsiventral lateral shoots with a
Flowers unisexual; sepals 5, distinct or connate; distichous leaf arrangement which superficially
petals 5, imbricate or valvate; disk 0; stamens can be mistaken as pinnate leaves. However, the
5–15, filaments distinct; anthers bilocular, usually shoots terminate in a claw-like bud which appears
introrse, dehiscing longitudinally; pollen grains capable of producing a further flush of leaves; they
prolate to oblate spheroidal, 3-colporate, inoper- are borne in the axil of a reduced leaf and bear a
culate (operculate), sexine tectate-punctate or bud in their axil. As a result of the strongly dorsi-
reticulate; pistillode columnar, non-lobate, some- ventral shoot organisation, the petioles are turned
times peltate; gynoecium syncarpous; ovary 2–5- almost 90 and the stipules are inserted at differ-
locular; ovules 1 per locule, pendulous, bitegmic, ent levels. Whereas in Microdesmis the leaves sub-
anatropous and epitropous or less often orthot- tend staminate flower-fascicles or solitary
ropous; obturator 0; stylodia 2–5, short, or 0; pistillate flowers, in Galearia and Panda the leaves
stigmas 2–5, stigmatoid, sometimes branched. have completely lost their axillary buds.
Fruits drupaceous, exocarp fleshy, hard, thick, In contrast to the condensed leaf-axillary
sculpted; endocarp bony, entire to perforate or inflorescences of Microdesmis, in Galearia and
ruminate, containing (2)3(4) one-seeded locules, Panda the inflorescences are thyrsiform, with
dehiscing by valves at germination. Seeds the staminate flowers arranged in cymose clusters
ecarunculate; embryo flattened, truncate apically, along the elongate axis and the pistillate flowers
cordate basally; endosperm copious, oily. 2n ¼ 30 solitary or paired in pseudoracemes. In Galearia
(Microdesmis). subgen. Galearia they are produced at the tip of
Three genera, tropical Africa and SE Asia the leafy shoots, and in G. subgen. Orthopetalum
through Malesia to the Solomon Islands. the inflorescences are borne both on bracteose
dorsiventral shoots and on bosses of old wood
V E G E T A T I V E A N A T O M Y . Stomates are anomo- of branches and trunks. Panda has only cauliflor-
cytic or paracytic, in Panda also encyclocytic. ous inflorescences.
Solitary or clustered crystals of calcium oxalate
are present in parenchymatous tissue of the F L O R A L S T R U C T U R E . The calyx of Microdesmis
shoot; vessel segments have scalariform or both is clearly 5-partite or 5-lobed, whereas in
scalariform and simple perforations; imperforate Galearia and Panda it is increasingly cup-shaped;
trachaeary elements have bordered pits and occa- in Panda, the cup bears only five minute teeth.
sionally some are true tracheids; wood rays are Petal aestivation is imbricate, though weakly so in
heterocellular, 4–6-seriate, some uniseriate, staminate Panda, but usually valvate in Galearia.
mostly of procument cells, the uniseriate of To my knowledge, the structure of the ovule has
upright or square cells; wood parenchyma is directly been observed only in Panda, and
270 K. Kubitzki
Forman (1966) has inferred its anatropy in the 1976; Tokuoka and Tobe 2003). The outer
other genera from the position of the embryo in epidermis is underlain by one or several layers of
the seed. In Microdesmis and most Galearia, the longitudinal, pitted, lignified fibres and an inner
seeds develop with the radicle pointing laterally, epidermis. It is uncertain whether this structure is
i.e. as a result of an anatropous ovule. In one tegmic (as usually interpreted) because there is no
Galearia (G. maingayi) and in Panda the radicle indication of a presumably testal layer.
of the seed points downwards. This suggests that
the ovule is orthotropous. F A M I L Y C I R C U M S C R I P T I O N A N D A F F I N I T I E S . On
the basis of vegetative and reproductive charac-
P A L Y N O L O G Y . The pollen grains have been ters, Forman (1966) proposed the inclusion of the
studied by Nowicke (1984) and Nowicke et al. tribe Galearieae (Microdesmis and Galearia) into
(1998). Those of Galearia and Microdesmis are the then monotypic family Pandaceae. This was
very similar: 3-colporate, prolate spheroidal to accepted by Webster (1987), but later (1994) he
prolate, small, 10–23 mm long, with a punctate- returned to including these three genera as an
perforate or microreticulate tectum; Panda dif- emended tribe Galearieae in subfamily Acalyphoi-
fers in being suboblate, up to 28 mm long, and deae. Since then, however, it has been found that
having a deeply reticulate tectum, with a notable the genera of Galearieae display a number of
range of lumina size and muri height—large and features discordant within the Acalyphoideae
high at the mesocolpia and poles, but smaller and and, indeed, within the Euphorbiaceae as a
shallower at the colpi margins. whole. These include divergent floral and wood
structural features, a seed coat with tracheoidal
S E E D A N D F R U I T . The fruits are drupes of very rather than palisadal mechanical layer, and the
different size and shape, but transverse sections of lack of obturators. Also molecular data (e.g.
them reveal a common pattern (Fig. 61). Those of Wurdack and Davis 2009; Soltis et al. 2011) firmly
Panda are subglobose, provided with a massive support the treatment of the Pandaceae as a sepa-
endocarp, and about 7 cm long, whereas those of rate family. These data are inconclusive, however,
Galearia are bilaterally flattened and transversely in regard to the closest relatives of the family.
elongate or depressed-angular. The drupes of Wurdack and Davis (2009) left Pandaceae unre-
Microdesmis are rounded and rarely more than 5 solved, and in Soltis et al. (2011) they appeared in
mm long, although in M. magellanensis they are an unsupported sister position to Centroplaca-
laterally compressed similar to those of Galearia ceae + Linaceae. Xi et al. (2012) resolve Pandaceae
subgen. Galearia. At germination in all genera, in a clade with Irvingiaceae, though with low sup-
valves break off from the stony endocarp to release port. This would make sense, because Pandaceae
the germinating seed (Hill 1937; Forman 1966). and Irvingiaceae share involute vernation, a single
The seed coat anatomy of the three genera is apical ovule per carpel, axile placentation, inde-
very similar (Vaughan and Rest 1969; Corner hiscent fruits, and palaeotropical distribution
(Stevens 2001 onwards; Xi et al 2012).
KEY TO THE GENERA

1. Inflorescences axillary, flowers solitary or in glomer-
ules; staminate petals imbricate; leaves usually pellu-
cid-punctate 1. Microdesmis
– Inflorescences terminal or cauliflorous, thyrsiform;
staminate petals valvate or imbricate; leaves not
pellucid-punctate 2
2. Endocarp mostly thin-walled; ovules usually anatro-
pous; petals valvate; inflorescences mostly terminal
Fig. 61. Pandaceae. Fruits in transverse section. A Micro- 2. Galearia
desmis magellanensis. B Galearia maingayi. C Panda – Endocarp thick-walled; ovules orthotropous; petals
oleosa. (Forman 1966) imbricate; inflorescences cauliflorous 3. Panda
Pandaceae 271
1. Microdesmis Planch. Fig. 61A flat; disk 0; stamens 10, biseriate, or 15 in 1

Microdesmis Planch., Hook. Ic. Pl. 8: t. 758 (1848); Pax & whorl; filaments basally connate, thickened, the
K. Hoffm., Pflanzenr. 147. III (Heft 47): 105, fig. 34 (1911); exterior inflexed; anthers introrse, muticous;
Léonard, Fl. Congo 8(1): 102, t. 8 (1962); Whitmore, Tree pollen grains prolate spheroidal to prolate,
Fl. Malaya 2: 118 (1973); Radcl.-Sm., Gen. Euphorb.: 126 3-colporate, operculate or inoperculate, sexine
(2001).
tectate-perforate; pistillode apically dilated,
Dioecious trees or shrubs; indumentum simple. pubescent; pistillate flowers pedicellate; sepals 5,
Leaves alternate, distichous, simple, entire or entire, imbricate, persistent; petals 5, imbricate;
dentate, minutely pellucid-punctate; stipules disk 0; ovary 2–5-locular; ovule pendulous, anat-
small, linear, persistent. Inflorescences axillary ropous or orthotropous; stigma sessile, variously
(supraaxillary), glomerular. Staminate flowers lobed or laciniate into several arms. Fruit a drupe,
pedicellate; sepals 5, slightly imbricate; petals 5, small, bilaterally flattened and transversely
contorted or imbricate in bud; disk 0; stamens 5 or elongate, or larger, depressed-(sub)globose, usu-
5 + 5, filaments dilated, broader than anthers, ally 1-seeded; exocarp fleshy or bony, endocarp
distinct from or adherent to lobes of the pistil- crustaceous, thin and entire or thick and much
lode; anthers basifixed, introrse or latrorse, some- perforated and channelled, 1–5-celled. Seeds
times apiculate, dehiscing longitudinally; pollen compressed.
subprolate, angulaperturate, 3-colporate, colpi Six spp., SE Asia, from Myanmar through
operculate or inoperculate, sexine tectate- Malesia to the Solomon Islands. Two subgenera:
perforate; pistillode columnar or pentagonal in subgen. Orthopetalum, petals straight and flat,
the lower half and cylindric at the apex; pistillate fruits 3–4-angled or circular; subgen. Galearia,
flowers pedicellate; sepals and petals 5, imbricate, petals in staminate flowers concave or cucullate,
the sepals persistent in fruit; disk 0; ovary 2–5- fruits laterally compressed.
locular; ovule pendulous, solitary, anatropous;
stylodia 2–5, short, deeply bipartite, papillose, 3. Panda Pierre Figs. 61C, 62
spreading. Fruit a drupe, ovoid-subglobose, not
Panda Pierre, Bull. Mens. Soc. Linn. Paris: 1255 (1896);
or scarcely lobed; mesocarp fleshy, endocarp Engl., Notizbl. Kgl. Bot. Gart. Mus. 5 (49): 274–276, with
woody, tubercled or muricate, (1)2–5-locular. fig. (1912); Aubrév., For. Fl. Côte d’Iv. 1: 300 (1959);
Seeds compressed-ovoid, testa smooth; cotyle- Forman, Kew Bull. 20: 309–321 (1966).
dons broad, flat.
Ten spp., 8 in tropical Africa, and 2 in Asia. Dioecious trees; buds in axils of leafy short-
shoots, not in leaf-axils; indumentum simple.
Leaves alternate, distichous, simple, subentire to
2. Galearia Zoll. & Moritzi Fig. 61B shallowly toothed; stipules minute, deciduous.
Galearia Zoll. & Moritzi, Syst. Verz.: 19 (1846); Pax, Pflan- Inflorescences ramiflorous to cauliflorous, the
zenr. 147. III (Heft 47): 97, figs. 31, 32 (1911); Forman, staminate in up to 25 cm long thyrsiform pseu-
Kew Bull. 26: 153–165, fig. 1, 2 (1971), rev.; Airy Shaw, doracemes fascicled on bosses on the trunk and
Kew Bull. 36: 365 (1981); Radcl.-Sm., Gen. Euphorb.: 127 branches, the pistillate flowers solitary or paired
(2001).
on elongate racemes; bracts minute. Staminate
Dioecious trees or shrubs; buds in shoot-axils, not flowers pedicellate; sepals completely connate
in leaf-axils; indumentum simple. Leaves alter- into a truncate 5-toothed cup open in bud; petals
nate, distichous, simple, entire (crenulate 5, valvate below, weakly imbricate above; disk 0;
towards the apex), eglandular; stipules entire to stamens 10, biseriate; filaments distinct, the outer
pinnatifid, persistent or deciduous. Inflores- much longer than the inner; anthers basifixed,
cences terminal or cauliflorous, pseudorace- introrse, dehiscing longitudinally, connective
mose-thyrsiform (the pistillate ones racemose), somewhat enlarged; pollen grains oblate spheroi-
elongate, often pendulous, sometimes in cymose dal, 3-colporate, sexine coarsely reticulate, dis-
fascicles and then pistillate ones solitary. Stami- tinctly heterobrochate; pistillode columnar;
nate flowers sessile or pedicellate; sepals 5, free or pistillate flowers pedicellate; sepals connate into
connate, entire, imbricate; petals 5, concave or a truncate cup; petals 5, imbricate; disk 0;
272 K. Kubitzki
into (2)3–4 valves, leaving a large capped tri- or

tetrapterygoid columella. Seeds flattened,
obtriangular; cotyledons cordate, truncate.
A single sp., P. oleosa Pierre, restricted to
West African rain forest areas (Côte d’Ivoire to
the Congo). The seed oil is regionally used for
cooking. The fruits are said to be dispersed by
elephants.
Corner, E.J.H. 1976. See General References.
Engler, A. 1912. Panda oleosa, ein Ölsamenbaum Westa-
frikas. Notizbl. K€ onigl. bot. Gart. Mus. Berlin 5:
274–276.
Forman, L.L. 1966. The reinstatement of Galearia Zoll. &
Mor. and Microdesmis Hook.f. in the Pandaceae,
with appendices by C. R. Metcalfe and N. Parames-
waran. Kew Bull. 20: 309–321.
Hayden, W.C., Hayden, S.M. 2000. Wood anatomy of
Acalyphoideae (Euphorbiaceae). IAWA J. 21:
213–235.
Hill, A. W. 1937. The method of germination of seeds
enclosed in a stony endocarp. II. Ann. Bot. II, 1:
239–256, fig. 7.
Nowicke, J.W. 1984. A palynological study of the Panda-
ceae. Pollen et Spores 26: 31–42.
Nowicke, J.W., Takahashi, M., Webster, G.L. 1998. Pollen
morphology, exine structure and systematics of Aca-
lyphoideae (Euphorbiaceae) Part 1. Rev. Palaeobot.
Fig. 62. Pandaceae. Panda oleosa. A Branch with stami- Palynol. 102: 115–152.
nate inflorescences. B Part of staminate inflorescence. Radcliffe-Smith, A. 2001. See General References.
C Flower bud. D Staminate flower. E Outer and inner Soltis, D.E. et al. 2011. See General References.
stamen. F Part of pistillate inflorescence. G Calyx and Stevens, P.F. 2001 onwards. See General References.
pistil of pistillate flower. H Ovary in cross section. I Pistil Tokuoka, T., Tobe, H. 2003. Ovules and seeds in Acaly-
in longitudinal section. J Fruit, part of exocarp removed. phoideae (Euphorbiaceae): structure and systematic
K Endocarp, seen from below. L Cross section of endo- implications. J. Pl. Res. 116: 355–380.
carp and locules. M Seed with cotyledon in longitudinal Vaughan, J.G., Rest, J.A. 1969. Note on the testa structure
section. N Seed in radial length section. (Engler 1912) of Panda Pierre, Galearia Zoll. et Mor. and Micro-
desmis Hook. f. (Pandaceae). Kew Bull. 23: 215–218.
Webster, G.L. 1987. The saga of the spurges: a review of
staminodes 0; ovary (2)3–4-locular; ovule pendu- the classification and relationships of the Euphorbia-
lous, orthotropous; stigmas (2)3–4, entire, erect ceae. Bot. J. Linn. Soc. 94: 3–46.
at first, later reflexed. Fruit a drupe, subglobose, Webster, G.L. 1994. Synopsis of the genera and supra-
generic taxa of the Euphorbiaceae. Ann. Missouri
c. 6.5 cm diam., (2)3–4-seeded; exocarp thick, Bot. Gard. 81: 33–144.
tartareous; endocarp massive, bony, pitted, (2) Wurdack, K.J., Davis, C.C. 2009. See General References.
3–4-sutured, tardily dehiscent along these sutures Xi, Z. et al. 2012. See General References.
Putranjivaceae
Putranjivaceae Endl., Ench. Bot.: 174 (1841).
G. L E V I N
Trees or shrubs, with simple trichomes or some- Wood is ring or diffuse porous with the ves-
times stellate on fruits. Leaves alternate or rarely sels solitary or in radial multiples. Perforation
opposite, spirally arranged but often appearing plates are scalariform, simple, or both; intervas-
distichous, simple, petiolate, pinnately veined, cular and ray/vessel pitting is minute and alter-
base usually oblique, margins entire or dentate, nate. Axial parenchyma is diffuse or apotracheal
the teeth sometimes spinose; stipules deciduous in numerous short, often interrupted bands one
or persistent. Flowers in axillary or cauliflorous cell wide. Rays are uniseriate and multiseriate,
clusters, sometimes solitary, rarely cymes, pedi- heterocellular. Fibers are non-septate and very
cellate, actinomorphic, generally unisexual and thick-walled. Prismatic crystals are found in the
the plants dioecious, but occasionally polygamo- axial and ray parenchyma (Smith and Ayensu
dioecious or monoecious; sepals (3)4–5(6), dis- 1964; Hayden 1980; Mennega 1987).
tinct, imbricate or the pistillate rarely open in
bud; petals 0. Staminate flowers: stamens (2) E M B R Y O L O G Y A N D S E E D A N A T O M Y . Readers
3–20(50), filaments distinct, anthers erect, are referred to the detailed descriptions by
dithecal, introrsely or less frequently latrorsely Stuppy (1996) and Tokuoka and Tobe (1999).
or extrorsely dehiscent by longitudinal slits; disk Systematically significant features include glan-
intrastaminal or absent; pistillode small or dular anther tapetum; two-celled pollen grains
absent. Pistillate flowers: sepals deciduous or per- (when shed); anatropous, bitegmic, crassinucel-
sistent; disk annular or absent; ovary superior, late ovules; a two- or three-celled archesporium; a
syncarpous, 1–3(6)-locular, placentation axile; thin, two cell-layered parietal layer in the nucel-
ovules 2 per locule, anatropus, bitegmic, embed- lus; Polygonum type embryo sac; no nucellar
ded in a massive obturator; stylodia as many as beak or cap; early disintegrating nucellar tissue;
the locules, usually short, stigmas subpeltate, a massive funicular obturator; thick, multiplica-
reniform, discoid, bilobed, or petaloid; fruit dru- tive inner and outer integuments; an endothe-
paceous; seeds 1 per locule or fruit by abortion, lium; Nuclear endosperm formation; abundant
albuminous, embryo large, straight. endosperm in the seeds; a large, straight embryo;
Two genera and c. 225 spp., tropical and and generally a fibrous exotegmen.
subtropical Asia, Africa, America, Australia, and
western Pacific islands. P O L L E N M O R P H O L O G Y . Pollen is spheroidal to
prolate, 3-colporate with transversely elongate
VEGETATIVE ANATOMY. The leaves are dorsiventral. ora and often with elongate colpi, and tectate-
Druses may be found in the mesophyll, and pris- reticulate (Punt 1962; K€ohler 1965). Punt (1962)
matic crystals in the mesophyll and vein bundle assigned the pollen of Drypetes and Putranjiva to
sheathes. Venation is brochidodromous or some- different types based on grain shape and exine
time eucamptodromous; higher order venation thickness, whereas K€ohler (1965), who sampled a
patterns are variable and may be systematically more diverse set of Drypetes species, assigned the
informative. Stomata are restricted to the abaxial two genera to the same type and recognized a
side and are brachyparacytic, with the subsidiary second pollen type in Drypetes based primarily
cells overlying the guard cells (Levin 1986). on aperture length. A broader survey could reveal
274 G. Levin
that variation in shape, apertures, exine thick- Brassicales (Rodman et al. 1998; APG II 2003).
ness, and sculpture is systematically informative. Lingelsheimia has not been included in molecular
studies, but its morphology is quite different
K A R Y O L O G Y . Chromosome numbers have been from Putranjivaceae (Léonard 1962) and consis-
reported for only five Drypetes species, four tent with Phyllanthoideae.
African (Hans 1973) and one southeast Asian Affinities within Malpighiales place Putranji-
(Oginuma et al. 1998), and they are consistently vaceae with strong support as the sister of Lopho-
n ¼ 20. In contrast, counts of n ¼ 7, 19, 20, and 21 pyxis (Wurdack and Davis 2009; Soltis et al.
have been reported from Putranjiva roxburghii 2011), and Xi et al. (2012) have resolved Putran-
(Hans 1973; Sanjappa 1979; Chattopadhyay and jivaceae and Lophopyxidaceae, the "putranji-
Sharma 1988). It is possible that the base number voids", within a larger clade, elsewhere
for the family is x ¼ 7, with polyploidy and comprising Caryocaraceae, "malpighioids"
aneuploidy accounting for the other numbers. It (Malpighiaceae, Elatinaceae, Centroplacaceae)
is also possible that the counts other than n ¼ 20, and the "chrysobalanoids". For morphological
particularly those of n ¼ 7, are in error. Broader traits shared by Putranjivaceae and Lophopyxi-
surveys are indicated. daceae, see under Lophopyxidaceae, this volume.
Various efforts have been made to fragment
F R U I T A N D D I S P E R S A L . Although all fruits of Drypetes into smaller genera. The only commonly
Putranjivaceae are drupaceous, there is consider- accepted segregates are Sibangea and Putranjiva.
able variation in the exocarp. It may be fleshy, The former, which has three African species,
leathery, or somewhat woody, and colors at matu- is distinguished by having the pistillate sepals
rity include red, orange, yellow, brown, and open in bud and persistent in fruit, in contrast
white. Tomlinson (1980) reported that the red to imbricate and deciduous in Drypetes s.s.
fruits of Drypetes lateriflora are removed rapidly, (Radcliffe-Smith 2001). Phylogenetic analysis of
whereas the white fruits of D. alba persist on the DNA sequence data place Sibangea within
trees for long periods. There are almost no Drypetes (Wurdack et al. 2004), a placement that
detailed studies of fruit dispersal in the family, is adopted here. Although Hurusawa (1954)
but various doves and pigeons have been reported reduced Putranjiva to a subgenus of Drypetes, a
to disperse the fruits of D. deplanchei (Floyd 1989; treatment that has been widely followed, molecu-
Forster 1997). lar phylogenetic studies show them to be sister
taxa (Wurdack et al. 2004) and here they are
P H Y T O C H E M I S T R Y . Putranjivaceae is notewor- treated as distinct.
thy for producing glucosinolates (mustard oil
glucosides). Phylogenetic studies demonstrate D I S T R I B U T I O N A N D H A B I T A T S . Drypetes grows
that this evolved independently here and in the primarily in tropical and subtropical lowland for-
Brassicales (Rodman et al. 1998). In addition, ests, woodlands, and savannas. A few species are
sesquiterpene lactones and friedelanes are found in tropical montane forests or warm tem-
known from the family (Wandji et al. 2003). perate areas. Its greatest diversity is in Asia, with
about 120 species. About 75 species grow in
P H Y L O G E N Y . Putranjivaceae traditionally have Africa and Madagascar, with only about 20 spe-
been included in Euphorbiaceae subfamily Phyl- cies, many of them undescribed, found in the
lanthoideae (Phyllanthaceae) as tribe Drypeteae, Americas. Putranjiva is found in forests of tropi-
along with the African genus Lingelsheimia Pax cal Asia from the Indian subcontinent to Indone-
(Webster 1994; Radcliffe-Smith 2001). Meeuse sia, south China, Taiwan, and the Ryukyu Islands.
(1990), focusing on embryology, seed anatomy, Most species are narrowly distributed, but P. rox-
and especially chemistry, removed the tribe from burghii is found in seasonal forests from Pakistan
the Euphorbiaceae and placed it as a family in to Indonesia.
Brassicales. Molecular phylogenies clearly show
that Drypetes and Putranjiva are closely related, E C O N O M I C I M P O R T A N C E . Fruit of some species
belong outside the Euphorbiaceae, and are of Drypetes are eaten locally, and the hard wood is
members of the Malpighiales rather than the locally used in construction. Bark extracts from
Putranjivaceae 275
several Drypetes species and Putranjiva roxbur-

ghii are used medicinally in central Africa and
India; these may have pharmacological value
(Chungag Anye et al. 2003). P. roxburghii is
widely cultivated as an ornamental tree in tropi-
cal to warm temperate regions.
C O N S E R V A T I O N . Although many species of

Putranjivaceae are common, in some cases domi-
nant forest trees, others are of conservation con-
cern. This is particularly true of some island
endemics, notably Drypetes glabra and D. henri-
quesii from São Tomé, D. andamanica from the
Andaman Islands, D. leiocarpa from the Nicobar
Islands, and D. riseleyi from the Seychelles.
KEY TO THE GENERA

1. Disk present; stamens mostly 4 or more; stigmas some-
what swollen and subpeltate, reniform, discoid, or
bilobed, but not petaloid 1. Drypetes
– Disk absent; stamens mostly 2 or 3; stigmas conspicu-
ously dilated and petaloid 2. Putranjiva
1. Drypetes Vahl Fig. 63

Drypetes Vahl, Eclog. Amer. 3: 49 (1810); Pax & K. Hoffm.,
Pflanzenreich 147. XV: 229–279 (1922), rev.; Airy Shaw,
Kew Bull. Addit. Series 4: 97–108 (1975), rev. Borneo spp.;
Airy Shaw, Kew Bull. 36: 286–292 (1981), rev. Sumatra
spp.; Radcl.-Sm., Fl. Trop. E Africa, Euphorbiaceae I:
88–103 (1987), rev. E Afric. spp.; Radcl.-Sm., Fl. Zambes.
9(4): 87–93 (1996), rev. SE Afric. spp.; P.I. Forster, Aus-
trobaileya 4: 477–494 (1997), rev. Australian spp.; T.
Chakrab et al., J. Econ. Taxon. Bot. 21: 251–280 (1997),
rev. S Asian spp.; McPherson, Adansonia III, 22: 205–209 Fig. 63. Putranjivaceae. Drypetes mossambicensis. A Part
(2000), rev. Madag. spp.; L. Phuphathanaphong and of distal branch with immature fruits. B Staminate
K. Chayamarit. Flora of Thailand, 8(1); 231–253 (2005), inflorescence. C Staminate flower. D Disk thereof.
rev. Thai spp. E Pistillate flower. F Fruit. (Radcliffe-Smith 1996; drawn
by J-M. Fothergill)
Sibangea Oliv. (1883).
Trees or shrubs, usually dioecious. Leaves alternate About 220 spp.; subsaharan Africa, southern
or rarely opposite, spiral but sometimes appearing and eastern Asia, Australasia, and tropical
distichous, petiolate, stipulate, pinnately veined, America.
usually oblique at the base, margins entire or
spinose-dentate. Flowers in axillary clusters or
2. Putranjiva Wallich
cauliflorous; sepals 4–5, imbricate or the pistil-
late rarely open in bud; petals none. Staminate Putranjiva Wall., Tent. Fl. Napal.: 61 (1826); Hurus., J.
Fac. Sci. Univ. Tokyo III, Bot. 6: 335–338 (1954), reg. rev.
flowers: stamens 3–20(50); disk intrastaminal;
pistillode small or none. Pistillate flowers: disk Trees, dioecious. Leaves alternate, spiral, petiolate,
annular; ovary 1–3(6)-locular; stigmas subpel- stipulate, pinnately veined, usually oblique at the
tate, reniform, discoid, or shallowly bilobed. Fruit base, margins entire or dentate. Flowers in axillary
drupaceous. Seeds 1 per locule or fruit. n ¼ 20. clusters or the pistillate solitary; sepals 3–6,
276 G. Levin
imbricate; petals none; disk none. Staminate Mennega, A.M.W. 1987. Wood anatomy of the Euphor-
flowers: stamens 2–3(4), anthers extrorsely biaceae, in particular of the subfamily Phyllanthoi-
deae. Bot. J. Linn. Soc. 94: 111–126.
dehiscent; pistillode none. Pistillate flowers: ovary Oginuma, K., Kiaptranis, R., Damas, K., Tobe, H. 1998. A
2–3-locular; stigmas dilated, petaloid. Fruit drupa- cytological study of some plants from Papua New
ceous. Seed 1. n ¼ 20 (and possibly 7, 19, and 21). Guinea. Acta Phytotax. Geobot. 49: 105–114.
Four spp., tropical Asia, from Pakistan and Punt, W. 1962. Pollen morphology of the Euphorbiaceae
with special reference to taxonomy. Wentia 7: 1–116.
Ceylon to Taiwan and Ryukyu Islands. Radcliffe-Smith, A. 1996. Euphorbiaceae. In: Pope, G.V.
(ed.) Flora Zambesiaca 9, Pt. 4. Kew: Royal Botanic
Gardens.
Radcliffe-Smith, A. 2001. Genera Euphorbiacearum. Rich-
Selected Bibliography mond: Royal Botanic Garden, Kew, pp. 48–55.
Rodman, J.E., Soltis, P.S., Soltis, D.E., Sytsma, K.J.,
APG II. The Angiosperm Phylogeny Group. 2003. See Karol, K.G. 1998. Parallel evolution of glucosinolate
General References. biosynthesis inferred from congruent nuclear
Chattopadhyay, D., Sharma, A.K. 1988. Sex difference and and plastid gene phylogenies. Amer. J. Bot. 85:
chromosomes in Putranjiva roxburghii Wall. Curr. 997–1006.
Sci. 57: 1017–1019. Sanjappa, M. 1979. IOPB chromosome number reports
Chungag Anye, N.B., Njamen, D., Wandji, J., Fomum, Z.T., LXIII. Taxon 28: 274–275.
Dongmo, A., Nguelefack, T.B., Wansi, D., Kamanyi, A. Smith, A.C., Ayensu, E.S. 1964. The identity of the genus
2003. Anti-inflammatory and analgesic effects of Calyptosepalum S. Moore. Brittonia 16: 220–227.
Drypemolundein A, a sesquiterpene lactone from Soltis, D.E. et al. 2011. See General References.
Drypetes molunduana. Pharmaceut. Biol. 41: 26–30. Stuppy, W. 1996. Systematische Morphologie und Anato-
Floyd, A.G. 1989. Rainforest Trees of Mainland South- mie der Samen der biovulaten Euphorbiaceen. Ph.D.
eastern Australia. Melbourne & Sydney: Inkata Press. dissertation. Kaiserslautern, Germany: Universit€at
Forster, P.I. 1997. A taxonomic revision of Drypetes Vahl Kaiserslautern.
(Euphorbiaceae) in Australia. Austrobaileya 4: 477–494. Tokuoka, T., Peng, C.-I. 1997. Floral morphology and its
Govaerts, R., Frodin, D.G., Radcliffe-Smith, A. 2000. systematic implications in Drypetes integerrima
World Checklist and Bibliography of Euphorbiaceae (Koidz.) Hosok. (Euphorbiaceae, tribe Drypeteae)
(and Pandaceae). Kew: Royal Botanical Gardens, from Bonin Islands, Japan. Acta Phytotax. Geobot.
pp. 591–614, 1381–1383, 1471–1473. 48: 159–166.
Hans, A.S. 1973. Chromosomal conspectus of the Euphor- Tokuoka, T., Tobe, H. 1999. Embryology of tribe Drype-
biaceae. Taxon 22: 591–636. teae, an enigmatic taxon of Euphorbiaceae. Pl. Syst.
Hayden, W.J. 1980. Systematic anatomy of Oldfieldioideae Evol. 215: 189–208.
(Euphorbiaceae). Ph.D. dissertation. College Park: Tomlinson, P.B. 1980. The Biology of Trees Native to
University of Maryland, pp. 202–219. Tropical Florida. Alston, Massachusetts: Harvard
Hurusawa, I. 1954. Eine nochmalig Durchsicht des University Printing Office.
herk€ommlichen Systems der Euphorbiaceen im weite- Wandji, J., Tillequin, F., Mulholland, D.A., Temgoua, A.D.,
ren Sinne. J. Fac. Sci. Univ. Tokyo III, Bot. 6: 209–342. Wansi, J.D., Seguin, E., Fomum, Z.T. 2003. Phenolic
K€ohler, E. 1965. Die Pollenmorphologie der biovulaten constituents from Drypetes armoracia. Phytochemis-
Euphorbiaceae und ihre Bedeutung f€ ur die Taxono- try 63: 453–456.
mie. Grana Palynol. 6: 26–120. Webster, G.L. 1994. Synopsis of the genera and supra-
Léonard, J. 1962. Notulae systematicae XXXIII. Sur les generic taxa of Euphorbiaceae. Ann. Missouri Bot.
limites entre les genres Drypetes Vahl et Lingelshei- Gard. 81: 33–144.
mia Pax (Euphorbiacées). Bull. Jard. Bot. État 32: Wurdack, K.J., Davis, C.C. 2009. See General References.
513–516. Wurdack, K.J., Hoffmann, P., Samuel, R., de Bruijn, A.,
Levin, G.A. 1986. Systematic foliar morphology of Phyl- van der Bank, M., Chase, M.W. 2004. Molecular phy-
lanthoideae (Euphorbiaceae). I. Conspectus. Ann. logenetic analysis of Phyllanthaceae (Phyllanthoideae
Missouri Bot. Gard. 73: 29–85. pro parte, Euphorbiaceae sensu lato) using plastid
Meeuse, A.D.J. 1990. The Euphorbiaceae auct. plur., an rbcL DNA sequences. Amer. J. Bot. 91: 1882–1900.
unnatural taxon. Delft: Eburon. Xi, Z. et al. 2012. See General References.
Quiinaceae
Quiinaceae Engl. (1888), nom. cons.
K. K U B I T Z K I
Usually (andro)dioecious trees of moderate size unbranched stem bears a rosette of huge leaves
with lysigenous mucilage-containing spaces and and is terminated by the inflorescence; it is
ducts. Leaves opposite or whorled, serrate to unknown whether Froesia is hapaxanthous or,
entire, simple, lobed or pinnately compound by continued growth of the stem apex after fruc-
(often pinnately compound on young plants, tification or by sympodial branching, can flower
simple on adults), usually glabrous, petiolate, repeatedly. Phyllotaxis is generally decussate;
secondary veins pinnate, craspedodromous to whorls with preferably four leaves occur in most
brochidodromous, tertiary parallel or plumose species of Lacunaria and a single Quiina, Q. pter-
reticulate and very closely spaced; stipules inter- idophylla. Leaves on juvenile plants and coppice
petiolar, foliaceous and with midrib and pinnate shoots are often pinnatisect or pinnately lobed
venation or setaceous, distinct or sometimes pair- (Fig. 64); this leaf form persists in adult Froesia
wise connate. Inflorescences axillary or (Froesia) and Touroulia. Some species of Quiina, such as
terminal thyrsoids or botryoids. Flowers bisexual Q. leptoclada, Q. indigofera and Q. obovata, have
or more often unisexual and then dioeciously simple seedling leaves. The leaves have interpe-
distributed (the pistillate with stamens producing tiolar stipules which, by virtue of a midrib and
sterile pollen), hypogynous, regular; sepals 3–5(6), pinnate venation, often appear leaf-like and can
small, unequal, imbricate; petals (3)4–5(–8), con- vary from leafy to setose, and caducous to persis-
torted or imbricate, basally sometimes joining tent; they can be deeply dissected with several
with neighbouring petals and sepals to form a setose lobes (Froesia) or united through pair-
short floral cup; stamens 15–numerous, some- wise fusion as in Touroulia (Foster 1950a, b,
times forming 5 indistinct fascicles; filaments 1951). Leaves can be very large and may reach a
distinct, slender; anthers basifixed to slightly length of 1.5 m, the stipules 10 cm.
dorsifixed, bithecate, dehiscing by slits; gynoe-
cium syncarpous, 2–14-carpellate, longitudinally V E G E T A T I V E A N A T O M Y . Gross venation of
ribbed, with as many locules and stylodia as car- leaves and leaflets is craspedodromous or brochi-
pels, or (Froesia) apocarpous and 3-carpellate; dodromous, but last order veinlets produce a
ovules 2(–4) in each locule, on basal-axile placen- peculiar, closely spaced parallel to plumose-
tas, epitropous, anatropous, ascending. Fruit reticulate pattern characteristic of the family. The
berry-like, fleshy or leathery, but often dehiscent veinlets originating from adjacent lateral veins
at full maturity, longitudinally ribbed. Seeds with anastomose straightway or, as in Touroulia guia-
or without endosperm, villous or (Froesia) gla- nensis, meet to form a "geniculate" arch (Fig. 65;
brous; the embryo straight, with short hypocotyl Foster 1950a, b, 1951). Hallier (1911) believed that
and thick cotyledons, or with thin cotyledons and this regular venation pattern might reflect the
thick endosperm (Lacunaria, Touroulia). product of fusion in an originally bipinnate leaf,
Four genera and about 51 spp., centred in the but there is little evidence in support of this.
humid lowlands of northern South America and Stomata are paracytic. The mesophyll of Froe-
extending to Belize and S Brazil. sia and Touroulia guianensis contains masses of
crystalliferous cells with U-shaped thickening
V E G E T A T I V E M O R P H O L O G Y . Quiinaceae are (cristarque cells). They form more or less contin-
small to midcanopy trees. In Froesia, the uous strands, often one abaxially and another
278 K. Kubitzki
Fig. 65. Quiinaceae. Touroulia guianensis, geniculate

venation pattern of seedling leaflet. (Foster 1950b)
such as diameter and simple perforations of ves-

sels, small vestured intervascular pits, and fibri-
form vessel members. Apart from this general
pattern, some noteworthy special features include
rays containing silica and scalariform vessel
perforations in Lacunaria, and banded paren-
chyma in Froesia. The nodes are trilacunar.
Among the numerous families that have been
compared with Quiinaceae, Ochnaceae and
Medusagynaceae are similar in wood anatomy.
I N F L O R E S C E N C E S T R U C T U R E . The basic type

seems to be a determinate thyrse (i.e. a thyrsoid)
with richly flowered dichasial cymes (Froesia,
Touroulia). Often several basal partial inflores-
cences are present per leaf axil, forming a
pleiothyrsoid. Possible progressions include
Fig. 64. Quiinaceae. Lacunaria crenata. Leaf dimorphism, (a) the transition from decussate to dispersed
a seedling with decussate stipulate pinnatified leaves, and phyllotaxis of bracts subtending partial inflores-
shoot tip of adult tree with simple leaves. (Foster 1951) cences (Quiina, Lacunaria); (b) a reduction in
number of flowers per partial inflorescence down
adaxially, which occur in direct contact with the to 1 (leading to botryoids, as in some Quiina); and
fibres flanking the vascular tissue of the veinlets. (c) the reduction in number of basal special
Quiina negrensis is noteworthy for containing thyrsoids (pleiothyrsoid ! thyrsoid).
abundantly small branched sclereids in the leaf
mesophyll (Pires 1950). FLOWER STRUCTURE AND REPRODUCTIVE SYS-
Lysigenous intercellular spaces and ducts T E M S . Although there is some meristic variation
filled with mucilage are generally found in the in the flowers of Quiinaceae, pentamery and an
medulla and vascular strands of the midrib and imbricate calyx originating in quincuncial
petiole. Their occurrence distinguishes the family sequence and a contorted corolla (as found in
from the Guttiferae in which Quiinaceae formerly Froesia and Touroulia) seem to represent the
were included. An extensive study of the anatomy basic condition. In Touroulia the androecium
of the wood and bark of the family revealed a originates on a ring wall in centrifugal direction
uniform pattern (Gottwald and Parameswaran (C. Bayer, pers. obs.), although fascicled androecia
1967). There is broad agreement in features are known as well (Lacunaria coriacea). The
Quiinaceae 279
origin, and molecular data place Froesia as sister to

the rest of the family (see below).
Froesia is the only genus with bisexual flow-
ers (but see Touroulia amazonica); Lacunaria is
truly dioecious with no vestiges of the opposite
sex in its staminate and pistillate flowers. Tour-
oulia guianensis and all species of Quiina are
also functionally dioecious with probably strictly
staminate or (morphologically) bisexual flowers
on different individuals, with significantly less
stamens in the bisexual than in the staminate
flowers. The bisexual flowers of Quiina are
reported to produce sterile pollen, in contrast
to the staminate ones (Baillon 1877; pers.
comm. of the late J.M. Pires to the author).
This issue needs further investigation because
Schneider (1996) observed in six species pollen
dimorphism with inaperturate grains in the sta-
minate flowers, whereas my own preliminary
tests with carmin acetic acid treatment of two
species of Quiina revealed stainable and, hence,
probably viable pollen in the staminate flowers.
Because nectar does not seem to be produced,
the pollen of these flowers may be a (deceptive?)
attractant; relevant field observations would be
of great interest.
A F F I N I T I E S A N D P H Y L O G E N Y . Traditionally,
Quiinaceae have been considered to form part of
a broadly delineated order Theales (Engler 1925),
where they were supposed to be close to Ochna-
ceae. Both families agree in wood anatomy (Gott-
wald and Parameswaran 1967), the possession of
cristarque cells (Amaral 1991), tenuinucellate
ovules, and the lack of nectaries, but in a revised
ordinal classification (APG 1998) they were
included in Malpighiales. Within this vast alli-
ance, multi-gene sequence analyses (Wurdack
and Chase 2009; Soltis et al. 2011; Xi et al. 2012)
place Quiinaceae sister to Medusagyne and these
Fig. 66. Quiinaceae. Lacunaria jenmani. A Branchlet as sister to Ochnaceae s. str.
with pistillate inflorescences. B Detail of pistillate inflo- The obvious basal position of Froesia within
rescence. C Pistillate flower with one sepal and three
petals removed. D Vertical section of pistil. E Vertical the family is confirmed both by a morphologi-
section of staminate flower. F Stamens. G Fruit. H Same, cally based (Schneider et al. 2002) and a DNA-
transversal section, showing cavities in pericarp. I Verti- based cladistic analysis (Schneider et al. 2006).
cal section of seed. (Reproduced with the kind permission
of the artist Bobbi Angell)
D I S T R I B U T I O N A N D H A B I T A T S . The family is
centred in the lowlands and colline belt of Ama-
apocarpy of Froesia raises the question of whether zonia and Guyana, with some extensions to the
it is a primary or secondary condition, but apply- Atlantic rain forest of E and SE Brazil, the cloud
ing the criteria of Endress (1982) for evolutionarily forests of northern Venezuela, the lowlands of
secondary apocarpy fails to indicate a secondary Central America up to Belize, and Jamaica.
280 K. Kubitzki
KEY TO THE GENERA Seeds 1(2) per cell, compressed, ferrugineous-vil-

lous; endosperm copious.
1. Leaves of adult plants pinnate or pinnatifid 2
Two spp. in the lowlands of northern South
– Leaves of adult plants simple 3
America. T. amazonica Pires differs considerably
2. Stipules distinct, setaceous; gynoecium apocarpous;
seeds glabrous 1. Froesia from T. guianensis Aubl. in larger flowers, dis-
– Stipules connate; gynoecium syncarpous; seeds villous tinct sepals, persistent and not resinous petals,
2. Touroulia non-geniculate tertiary veins, and absence of cris-
3. Leaves mostly verticillate, rarely opposite; pistillate tarque cells in the mesophyll, and time and again,
flowers without staminodes; ovary with 4–14 locules generic status has been suggested for the former.
and stylodia 3. Lacunaria
– Leaves mostly opposite, rarely verticillate; pistillate 3. Lacunaria Ducke Fig. 66
flowers with staminodes; ovary with 1–3(–5) locules
and stylodia 4. Quiina Lacunaria Ducke, Arch. Jardim Bot. Rio de Janeiro 4: 139
(1925), and Arch. Inst. Biol. Veg. 2: 168–171 (1935); Rocha
& Secco, Acta Amazonica 34: 425–433 (2004), rev.
GENERA OF QUIINACEAE Small or medium-sized trees, dioecious. Adult

leaves simple, in whorls often of 4, or rarely
1. Froesia Pires opposite (L. oppositifolia Pires), medium-sized
Froesia Pires, Bol. Técn. Inst. Agron. Norte 15: 22 (1948); to large; stipules linear-subulate or -lanceolate,
Steyermark & Bunting, Brittonia 27: 172–178 (1975), rev.; distinct to connate. Inflorescences axillary thyr-
Schneider & Zizka, Novon 7: 406–412 (1997), key. soids. Flowers unisexual; sepals 4–5, petals 4–8,
imbricate or contort, staminate flowers: stamens
Shrubs or small trees; trunk unbranched, ending in
20–170; pistillate flowers: ovary globose-
a terminal rosette of leaves. Leaves very large (up
pyriform, striate, with 4–14 locules and stylodia,
to 1.5 m long), opposite, imparipinnate, 11–27-
each locule with 2 ovules. Fruit a 5–13-celled,
foliolate, the leaflets opposite to alternate; veins
lignescent berry; pericarp fleshy with well-
prominent, lined with series of cristarque cells;
developed mucilage-containing cavities; septa
stipules distinct, setaceous. Inflorescence terminal,
dissolving. Seeds albuminous, compressed,
thyrsoid. Flowers bisexual; sepals (4)5, petals 5,
embedded in pulp; testa ferrugineous-villous.
yellow to roseate; stamens very numerous; carpels
Eight (–10) spp., humid forests of Central
(2)3, distinct; stylodia filiform. Mericarps 1–3-
(Costa Rica, Panama) and northern South
seeded. Seeds exalbuminous (?), glabrous.
America, one species in C Brazil.
Six spp., lowland of northern South America
and cloud forests of Venezuela.
4. Quiina Aubl.
Quiina Aubl., Hist. Pl. Gui. Suppl.: 19 (1775); Schneider,
2. Touroulia Aubl. Figs. 64, 65 Acta Bot. Venez. 21: 1–74 (1998), Venez. spp.
Touroulia Aubl., Hist. Pl. Gui. 1: 492 (1775); Pires, Trees or shrubs, sometimes climbing, androdioe-
Bol. técn. Inst. Agron. Norte 20: 48–50 (1950); Zizka &
Schneider, Willdenowia 29: 227–234 (1999). cious. Leaves opposite, rarely verticillate, simple,
entire or dentate, stipulate, the stipules leafy or
Small trees, androdioecious (T. amazonica with setose, distinct or connate. Inflorescences axillary
bisexual flowers?). Leaves opposite, imparipin- (pleio)thyrsoids or botryoids. Flowers either
nate, sometimes with alate rachis in distal part, morphologically bisexual or staminate but the
7–17-foliolate, leaflets sessile, serrate-crenate; plants probably largely functionally unisexual;
stipules subulate, caducous or persistent. Inflor- sepals 3–6 and petals 3–5(–7); stamens 14–62
escences terminal, slender thyrsoids. Flowers (staminate flowers) or 12–20 (morphologically
either morphologically bisexual or staminate but bisexual flowers); ovary with 2–3(–5) locules
the plants probably dioecious; sepals 5, distinct and stylodia. Fruit a 1–3-seeded, lignescent
or basally connate; petal (4)5; stamens in stami- berry with fibrous pericarp; cavities in the peri-
nate flowers and staminodes in pistillate flowers carp small to inconspicuous. Seeds ferrugineous-
numerous; ovary with 4–11 locules and stylodia. villous or velutinous; cotyledons thick; endo-
Fruit baccate, 4–8(–11)-celled; stylodia persistent. sperm 0.
Quiinaceae 281
About 34 spp., centred in northern S Amer- Gottwald, H., Parameswaran, N. 1967. Beitr€age zur Ana-
ica, extending with one species to coastal S Brazil tomie und Systematik der Quiinaceae. Bot. Jahrb.
Syst. 87: 361–381.
and with another to Belize. Hallier, H. 1911. L’origine et le système phylétique des
angiospermes exposés à l’aide de leur arbre généalo-
gique. Arch. Néerl. Sci. exact.et nat. Sér. III B,
Selected Bibliography 1: 146–234.
Matthews, M., Amaral, M., Endress, P.K. 2012. See Gen-
eral References.
Amaral, M.C.E. 1991. Phylogenetische Systematik der Pires, J.M. 1948. Quiinaceae. Bol. técn. Inst. Agrón. Norte
Ochnaceae. Bot. Jahrb. Syst. 113: 105–196. 15: 22–31.
APG 1998. The Angiosperm Phylogeny Group. An ordinal Pires, J.M. 1950. Contribuição para a flora Amazónica.
classification for the families of flowering plants. Bol. técn. Inst. Agrón. Norte 20: 41–51, pl. V–XV.
Ann. Missouri Bot. Gard. 85: 531–553. Schneider, J. 1996. Morphologische, anatomische und
Baillon, H.E. 1877. Histoire des plantes, t. 6. Clusiaceées – mikromorphologische Untersuchungen zur Taxono-
Quineae. Paris: Librairie Hachette. mie der Gattung Quiina (Quiinaceae). Diplomarbeit,
Endress, P.K. 1982. Syncarpy and alternative modes of Fachbereich Biologie, Joh. Wolfg. Goethe Uni-
escaping disadvantages of apocarpy in primitive versit€at, Frankfurt/Main.
angiosperms. Taxon 31: 48–52. Schneider, J.V., Swenson, U., Zizka, G. 2002. Phylogenetic
Engler, A. 1925. Quiinaceae. In: Engler & Prantl, Die reconstruction of the neotropical family Quiinaceae
nat€urlichen Pflanzenfamilien, 2nd edn, 21: 106–108. (Malpighiales) based on morphology with remarks
Foster, A.S. 1950a. Morphology and venation of the leaf of on the evolution of an androdioecious sex distribu-
Quiina acutangula Ducke. Amer. J. Bot. 37: 159–171. tion. Ann. Missouri Bot. Gard. 89: 64–76.
[Note that this sp. has been renamed Q. pteridophyl- Schneider, J.V., Swenson, U., Samuel, R., Stuessy, T.,
lea (Radlk.) Pires] Zizka, G. 2006. Phylogenetics of Quiinaceae
Foster, A.S. 1950b. Venation and histology of the leaflets (Malpighiales): evidence from trnL-trnF sequence
of Touroulia guianensis Aubl. and Froesia tricarpa data and morphology. Pl. Syst. Evol. 257: 189–203.
Pires. Amer. J. Bot. 37: 848–862. Soltis, D.E. et al. 2011. See General References.
Foster, A.S. 1951. Heterophylly and foliar venation in Wurdack, K.J., Davis, C.C. 2009. See General References.
Lacunaria. Bull. Torrey Bot. Club 78: 382–400. Xi, Z. et al. 2012. See General References.
Rhizophoraceae
Rhizophoraceae Pers. (1806), nom. cons.
A.E. S C H W A R Z B A C H
Shrubs or trees, sometimes with aerial roots. Fourteen genera and about 145 spp. with a
Leaves opposite or verticillate, decussate or biju- pantropical distribution, dominant in tidal
gate, simple, toothed, crenate, or entire, mar- swamps or growing in non-inundable forests.
ginal teeth of a distinctive Macarisioid type;
stipules interpetiolar, valvate and pubescent or V E G E T A T I V E S T R U C T U R E S . Aerial stilt roots are
imbricate and glabrous, always bearing colleters. well known from the mangrove genera but are
Inflorescences axillary, cymose, dichasial, or fas- found also in inland genera (Gynotroches, Cross-
ciculate. Flowers regular, bisexual or rarely uni- ostylis and Carallia). These roots are initiated
sexual, with articulated pedicels; the sepals from aerial parts of the plants and in Rhizophora
valvate, basally usually congenitally connate develop as a series of sympodial loops, whereas in
and above with (3)4–5(16) lobes postgenitally Ceriopos tagal and species of Bruguiera as well as
connected by interdigitation of their papillose in the inland genera they are formed on the hypo-
margins; petals equalling the number of sepals, cotyl and base of the trunk. In Bruguiera and
contorted or infolded in bud, usually with both a Ceriops there are also knee-like pneumatophores
terminal arista and filiform appendages on the that develop by looping of horizontal roots. In the
two lobes, rarely entire, usually each petal mangrove genera, trunk axes are monopodial but
enwrapping 1–5 stamens; androecium diploste- may branch continuously or diffusely, and the
monous or obdiplostemonous or polyandrous, branches may repeat the structure of the parent
the filaments sometimes connate at base, borne axis (Attim’s model). Little is known in this
around the base of or on an intrastaminal entire respect about the inland genera, but Carallia
or lobed nectary disk; anthers tetrasporangiate and Gynotroches may also correspond with
or (Rhizophora) multisporangiate, dehiscent by Attim’s model.
a longitudinal valve; gynoecium syncarpous, Leaves are opposite and have interpetiolar
ovary superior to inferior, (2)3–5(20)-carpel- stipules, which bear colleters that secrete gummy
late, the locules often incompletely or not at all substances onto the buds. However, strictly
separated by septae at anthesis; ovules 2–6 per decussate phyllotaxis (with right angles between
carpel, epitropous, anatropous or less often successive leaf whorls) is restricted to tribe
hemianatropous or campylotropous, bitegmic Macarisieae and Pellacalyx, whereas in Gynotro-
and crassinucellate or rarely tenuinucellate, cheae (except Pellacalyx) and Rhizophoreae suc-
usually apically inserted; style simple, stigma cessive leaf pairs are offset by angles of 70–80
capitate or with pronounced lobes, generally (bijugate phyllotaxis; Tomlinson and Wheat
papillate. Fruit capsular, baccate, or hard-walled 1979). This arrangement reduces shading and
and indehiscent. Seeds 1–many, nonappendaged, provides more flexibility of orientation in the
arillate, or winged; seed coat exotestal or exotes- branch systems as compared to the decussate
tal-exotegmic or undifferentiated; endosperm arrangement. The stipules of the genera with biju-
well-developed, oily; embryo green, usually gate leaves are imbricate, whereas those of genera
straight, with laminar cotyledons and epigeal with decussate leaves are valvate. Juvenile and
germination, or with thick cotyledons or thick usually also adult leaves are variously toothed or
cotyledonary body and viviparous germination. crenate in most inland Rhizophoraceae, but con-
n ¼ (13), 14, 16, 18, 21. sistently entire in the mangrove genera. The latter
284 A.E. Schwarzbach
have also a succulent hypodermal layer and often Crossostylis and Blepharistemma are reported to
abundant sclereids. Leaf texture varies with habi- be polygamodioecious.
tat, those of tribe Rhizophoreae being leathery Petals are basically bifid, provided with a
to somewhat fleshy, glabrous and with well- prominent terminal arista, and each encloses
developed hypodermis (Keating and Randriana- one or several stamens in a very unusual kind of
solo 1988). Laticiferous cells occur in more or less petal vernation. Many filamentous appendages
continuous subepidermal layers in sepals and appear on the distal lobes of the petals, which
ovaries; in Gynotroches and Pellacalyx, the latici- abort early in development in Crossostylis and
fers appear modified into idioblastic secretory Anopyxis and are never initiated in Rhizophora.
cells. Root hairs are absent from the family, with Petals of Bruguiera and Ceriops are postgenitally
the exception of seedlings of Cassipourea. The connected not only with their neighbours but also
nodes are trilacunar or multilacunar with split- among themselves, and the release of this bond is
lateral traces. instrumental in the explosive pollen dispersal of
Wood anatomically, both the Macarisieae these flowers.
and Rhizophoreae are quite homogeneous, The diplostemonous condition with the initi-
whereas Gynotrocheae appear less uniform ation of antesepalous stamens before the antepe-
(van Vliet 1976). Rhizophoreae stand out in talous ones occurs in genera of each tribe and is
having exclusively scalariform vessel perfora- considered as ancestral (Juncosa and Tomlinson
tions and scalariform inter-vessel pits, while 1988b). Modifications thereof have arisen in all
otherwise vessel perforations are mixed scalari- tribes; for instance, in Carallia and Pellacalyx, the
form/simple, and scalariform inter-vessel pits antepetalous stamens are initiated first. (See the
are nearly always absent; libriform fibres are comments for the occurrence of obdiplostemony
restricted to Rhizophoreae, and fibre tracheids and diplostemony within the same family in
are constantly present in Macarisieae and Gyno- Matthews and Endress 2011: 389.) There are also
trocheae. Rays are heterogeneous with scanty different pathways to polyandry in the family,
uniseriates. with antepetalous position of the supernumerary
The narrow, relatively short vessel elements stamens in Kandelia and Cassipourea, and
and their scalariform perforations of the Rhizo- antesepalous in Crossostylis. The nectarial disk
phoreae are an unexpected trait and may be develops after and independently from the
related to the high negative pressures induced androecium.
by the saline environment of the mangrove The ovaries are typically 3–5-carpellate with
(Tomlinson 1986). 2 ovules per carpel, but both the number of
Sieve element plastids of Rhizophoraceae carpels (Crossostylis, up to 20) or the number of
contain numerous rectangular or irregular pro- ovules per carpel (Gynotroches, up to 8) or both
tein crystals, which represents the P5c-type of the number of carpels and of ovules (Pellacalyx)
Behnke (1988) otherwise known only in is increased. The single style has a stigma that
Erythroxylaceae. sometimes bears rather long stigmatic lobes.
Most Macarisieae have superior ovaries, but two
INFLORESCENCE AND FLORAL STRUCTURE. genera of this tribe and most genera of the other
Inflorescences are basically cymose, in other tribes have half-inferior or inferior ovaries.
words, branching is consistently from prophylls,
and pairs of prophylls also subtend each flower. E M B R Y O L O G Y . Relevant traits can be summar-
Branching is mostly dichasial, usually with ized as follows: Anthers tetrasporangiate, tape-
suppression of the terminal flower ("pseudo- tum glandular, its cells binucleate, cytokinesis
dichotomous" branching), and the cymes are simultaneous or successive (in Gynotroches,
pedunculate or more or less sessile. Open- Pellacalyx and Crossostylis, simultaneous and
branched and condensed fasciculate inflores- successive even in the same anther); pollen
cences occur in all three tribes. tetrads tetrahedral or decussate; pollen binucle-
Flowers are mostly bisexual, but Gynotroches ate; ovules epitropus, bitegmic, crassinucellate
and some Sterigmapetalum are functionally dioe- or (Gynotroches and Pellacalyx) tenuinucellate;
cious (Juncosa and Tomlinson 1988b), and some integuments two cell layers thick at inception or
Rhizophoraceae 285
(Carallia and Rhizophoreae) multiseriate; endo- The genera with a superior ovary (Tribe
thelium absent in Rhizophoreae; ovule arche- Macarisieae and Crossostylis) have arillate or
sporium multicelled in Macarisieae, 1-celled in winged seeds, whereas the baccate Gynotrocheae
Gynotrocheae and Rhizophoreae; embryo genera lack any sort of seed appendage. Both
sac Polygonum type; endosperm initially free- the aril and the wing develop as an outgrowth of
nuclear; mature seeds with abundant the exostome and are regarded as homologous
non-starchy endosperm; embryo chlorophyllous (Tobe and Raven 1987). All inland groups
(Juncosa and Tobe 1988b; Tobe and Raven (Macarisieae and Gynotrocheae) basically have a
1987, 1988). similar mature seed coat structure, including a
These data support the sister position of well-developed exotesta and mostly also a fibrous
Gynotroches and Pellacalyx and add evidence in exotegmen; their seed coat is either exotestal or
favour of the placement of Crossostylis in the exotestal-exotegmic. It is significant that the seed
Gynotrocheae (for more details, see Juncosa and coat of arillate seeds is thicker than that of winged
Tobe 1988). The presence of an endothelium in seeds (c. 0.2 mm vs. less than 0.1 mm thick). The
Macarisieae and Gynotrocheae is unusual for a thinner seed coat of the latter may be adaptive in
moderately crassinucellar and bitegmic lineage facilitating their wind dispersal.
because an endothelium is usually associated The seeds of the Rhizophoreae are included
with unitegmic tenuinucellate ovules. However, in indehiscent, hard-walled fruits and their seed
Matthews and Endress (2011: 401) comment coat comprises only a thick testa that is histologi-
that the nucellus in the Rhizophoraceae is long cally undifferentiated but contains profusely
and slender and almost completely disintegrates branched vascular strands. Although the inner
at the flanks by the expansion of the embryo sac, integument is present at ovular stage, the mature
so that in the absence of an endothelium the seeds completely lack a tegmen.
embryo sac locally would come into direct con-
tact with the inner integument. P O L L I N A T I O N A N D D I S P E R S A L . Little is known
about the floral biology of terrestrial taxa, even
P O L L E N M O R P H O L O G Y (see Vezey et al. 1988). though there is a range from plants with perfect
Pollen in Rhizophoraceae is usually spheroidal, flowers to polygamous or dioecious plants.
(2)3(4)-colporate and has a punctate-psilate or Flowers are generally small and individually
rugulate tectum; in Macarisieae and less so in inconspicuous (but up to 6 cm diameter in Cross-
Gynotrocheae the intercolumellar space contains ostylis grandiflora), although their frequent
granular matrix. There are clearly defined endoa- aggregation and appendaged petals increase
pertures, which particularly in tribe Gynotro- their visibility. They may be associated with gen-
cheae are fused laterally. Palynologically tribe eralist flower visitors (small insects). Members of
Macariseae appears intermediate between Gyno- the mangrove tribe Rhizophoreae, on the other
trocheae and Rhizophoreae. Overall, pollen hand, have been well studied and show a consid-
morphology of the family is so generalized that erable range of floral mechanisms (Tomlinson
it hardly offers clues for assessing relationships et al. 1979). Rhizophora is presumed to be wind-
with other families. pollinated; it has a high pollen-ovule ratio and
pollen is discharged within the bud and presented
F R U I T A N D S E E D . Fruit and seed structure in by expansion of the usually hairy, but ephemeral
the family varies greatly and includes (1) dehis- petals. The style, however, is unelaborated for the
cent capsular fruits with arillate or winged seeds, reception of pollen. In Bruguiera and Ceriops
(2) baccate fruits that contain nonappendaged tagal there is an explosive discharge of pollen,
seeds, and (3) indehiscent hard-walled fruits triggered by the flower visitor of appropriate
that contain viviparous seeds (Tobe and Raven behaviour and size in relation to that of the
1988). The seeds of all Rhizophoreae contain flower. In this process, birds act as visitors of
ample endosperm, which in the mangrove genera the large-flowered Bruguieras, whereas day-flying
(except Bruguiera) is formed so abundantly that insects such as butterflies visit the smaller-
it extrudes from the micropyle and fills up the flowered Bruguieras, and small night-flying
space between the fruit wall and the seed. moths search the nocturnally scented Ceriops
tagal. Kandelia seems generalized in its floral (1814) in his circumscription of the family since
biology (Tomlinson 1986; Kondo et al. 1987). he included both mangal (Rhizophora) and
Direct observations on seed dispersal of the terrestrial taxa (Carallia). Consequently, the
terrestrial taxa are not known to us, but it is monophyly of this group has been accepted inde-
obvious that the arillate genera will be dispersed pendently of the sharp ecological contrasts within
by animals (Tobe and Raven 1988 think of ants it. The place of the family in the natural system
and possibly birds) and the winged ones by wind. has been extensively debated but molecular stud-
The distribution of Crossostylis, which is scat- ies have clarified its relationships (Schwarzbach
tered in Polynesian islands, has very likely been and Ricklefs 2000): Rhizophoraceae are the sister
dispersed by birds from one island to the other. family to the Erythroxylaceae, and both are sister
The small, hard seeds of the capsular Gynotro- to the Ctenolophonaceae, all in the order
cheae (Carallia, Gynotroches and Pellacalyx) Malpighiales (Wurdack and Davis 2009; Soltis
appear endozoochorously dispersed by flying et al. 2011; Xi et al. 2012). Anisophylleaceae that
vertebrates. have been considered close relatives of or even
The most distinctive characteristic of the man- been included in Rhizophoraceae (according to
grove Rhizophoraceae is the viviparous seedling in molecular analyses; Schwarzbach and Ricklefs
which the embryo lacks a dormant phase (Sussex 2000; Zhang et al. 2007) are placed in the Cucur-
1975) and develops on the parent plant, the hypo- bitales.
cotyl protruding from the fruit (Fig. 68H, I). The Traditionally the family has been divided
propagule is thus a seedling, not a seed. In into three tribes that, with slight modifications,
Bruguiera, the seedling disperses initially with the persist to the present day. The molecular ana-
fruit (Tomlinson 1986), whereas in Kandelia, lyses (Setoguchi et al. 1996; Schwarzbach and
Ceriops and Rhizophora only the seedling disperses. Ricklefs 2000) support the monophyly of the
Most of the elongated viviparous seedlings recircumscribed tribes and show Gynotrocheae
of the mangrove Rhizophoraceae are not “self- in a sister position with Rhizophoreae and both
planting” by falling vertically into the soft mud sister to Macarisieae.
below the parent tree but are carried away from it. By and large, the tribes can be characterised
For their survival they need habitats where the by fruit characters (capsular fruits, berries, and
tidal range is small and where they can raise the indehiscent fruits with viviparous seeds, see
plumule above the tidal influence so that gas above under FRUIT AND SEED); their flowers are
exchange becomes possible for the seedling, usually (ob)diplostemonous, and ovaries are
which is hindered by the thick, cutinized and mostly superior in Macarisieae and inferior in
stomata-free epidermis of the hypocotyl. Erection the other two tribes. In view of its arillate seeds,
of the horizontally stranded seedling with the Crossostylis has often been considered as
exposition of the plumule into the air is achieved belonging in Macarisieae but the molecular
by an eccentric secondary growth of the xylem in data resolve it firmly in the Gynotrocheae
the distal pole of the seedling, which produces a clade. Indeed, apart from its arillate seeds (per-
distinct hook at its distal end (Tomlinson and haps a convergent development?), Crossostylis
Cox 2000). shares many characters with Gynotrocheae,
such as the (semi)inferior gynoecium develop-
K A R Y O L O G Y . Chromosomes of Rhizophoraceae ing into a capsule that opens with only short
are very small, usually about 1 mm long. Cross- slits (see Schwarzbach and Ricklefs 2000: 557,
ostylis has n ¼ 14; Anopyxis n ¼ 32; for Cassi- and above under EMBRYOLOGY).
pourea, n ¼ 18 and n ¼ 21 have been reported,
and the Rhizophoreae genera uniformly have n ¼ P H Y T O C H E M I S T R Y A N D U S E S (see Hegnauer
18 (see listing in Juncosa and Tomlinson 1988). 1973). The bark of all mangrove species is rich
in tannin, which can amount up to 45 % of the
A F F I N I T I E S A N D S U B D I V I S I O N . Although some- dry weight. The tannins are mainly of the
times referred to as the "mangrove family", this condensed type, based mainly on procyanidin,
appellation is misleading since the majority of but prodelphinidin is also present; there are
taxa are terrestrial, a point established by Brown also some records of myricetin and ellagic acid
Rhizophoraceae 287
from bark and leaf extracts. The bark of type, but where other genera co-occur there is
Bruguiera, Ceriops and Rhizophora contains often a distinct zonation, with Ceriops the most
also large amounts of mucilage (11–35 %), landward and salt-tolerant, Bruguiera and Kan-
which is based on arabinose, rhamnose and delia less so and Rhizophora the least. Rhizophora
galactose. Alkaloids have been found in some commonly dominates the seaward fringe of the
species from all tribes; they belong to the mangal, sometimes associated with Avicennia
tropin, hygrolin and necin type, and sometimes (Avicenniaceae) and Sonneratia (Sonneratia-
contain sulphur. ceae). In some areas Rhizophora forms extensive
The main use of the family is mangrove tim- forests with trees up to 30 m tall; since there is
ber and bark, the former for fire wood and char- little understorey and the trees are supported by
coal, and the latter for tanning (formerly massive stilt roots, such forests have a distinctive
important) and dyeing (batik), but now seems physiognomy. The ecological status of the Rhizo-
to have been superseded by synthetic products phoreae suggests that they combine features of
or persists rather on a regional/local scale. pioneer and mature phases in vegetation devel-
opment because of the species poverty of the
P A L A E O B O T A N Y . Macrofossils are known only mangal, i.e. they have adapted to establish com-
from the mangrove genera, representing Ceriops munities as pioneers and maintain them as
and a Bruguiera-like form from the early Eocene mature-phase species (Tomlinson 1986). Typi-
in the London Clay (Collinson 1983), followed by cally, such plants are shade intolerant, not estab-
Kandelia from the middle Eocene of Alaska lishing under their own canopy, and Rhizophora
(Wolfe 1972). Pollen ascribed to Rhizophora regenerates most conspicuously in open areas
from strata older than middle Eocene that have like forest gaps and along shifting coastlines.
frequently been reported was rejected by Graham Although it has the reputation of being a land-
(2006) but relatively unambiguous palynological builder, it is probably not efficient in the consoli-
records show the genus widespread in coastal dation of highly mobile substrates shifted by
habitats of the Old and New World tropics from water action and tidal scour.
the late Eocene onwards. For the history of The terrestrial taxa of the family are much
Rhizophora in the New World mangrove, see less well studied ecologically but are typical of
Graham (2006). lowland rain forest. Consequently, the pre-
sumed ecological origins of the Rhizophoreae
D I S T R I B U T I O N A N D H A B I T A T S . The family as a within the family as a whole have been little
whole is virtually limited to the tropics and the discussed.
tribes themselves all have a considerable range,
but each with one relatively large and widely
KEY TO THE GENERA
distributed genus (indicated in parenthesis).
Macarisieae (Cassipourea, 55 spp.) extend from 1. Plants of mangrove forests; seeds germinating in
South America to India, but not to SE Asia; the fruit while this is still attached to the tree;
Gynotrocheae (Carallia, 10 spp.) are Old World hypocotyl protruding from the fruit 2
and range from Madagascar to the South Pacific – Plants of inland habitats; seeds not germinating
(Crossostylis); Rhizophoreae (Rhizophora, 8 spp.) in the fruit when this is still attached to the tree;
are pantropical but concentrated in the Indo- hypocotyl not protruding 5
Pacific region. The most widely distributed 2. Calyx always 4-lobed; petals entire, without
species is probably Bruguiera gymnorrhiza, a appendages; anthers multi-locellate, eventually
common component of the tidal environment dehiscing with a large ventral valve
(mangal) from East Africa to Samoa. 14. Rhizophora
Rhizophoreae, which may collectively be – Calyx 5–16-lobed; petals 2-lobed, multifid or
referred to as "the mangrove Rhizophoraceae", with apical appendages; anthers 4-locular,
are restricted to coastal communities on muddy dehiscing with lengthwise slits 3
substrate away from strong wave action and tidal 3. Flowers polymerous, diplostemonous; calyx
currents. Rhizophora species constitute the typi- 8–16-lobed, lobes subulate-lanceolate, pointed;
cal element of this distinctive tropical vegetation petals bilobed or emarginate 11. Bruguiera
– Flowers 4–5-merous, diplostemonous or poly- – (Polygamo-)dioecious; flowers 4–6-merous,

androus; calyx lobes ovate or linear-oblong, stamens 2 or more times the number of petals
acuminate or obtuse; petals fringed with apical 11
appendages or multifid 4 11. Flowers 4-merous, diplostemonous; ovary
4. Flowers 5-merous, diplostemonous; calyx lobes 3-locular 1. Blepharistemma
ovate, 2.5–3.5 mm long; petals less than 0.5 cm – Flowers 4–6-merous, stamens 2, 3, or more
long; hypocotyl ridged 13. Ceriops times the number of petals; ovary 2–5-locular
– Flowers 4–5-merous, polyandrous; calyx lobes 2. Cassipourea
linear-oblong, about 15 mm long; petals c. 1.5 12. Flowers unisexual, plants dioecious
cm long; hypocotyl smooth 12. Kandelia 6. Sterigmapetalum
5. Fruit baccate; seeds nonappendaged 6 – Flowers perfect, plants hermaphrodite 13
– Fruit capsular; seeds appendaged (arillate or 13. Leaves mostly in whorls of three, stamen
winged) 8 filaments connate into a tube 4. Anopyxis
6. Leaves with indumentum of stellate and simple – Leaves in pairs, stamen filaments distinct
hairs; free part of the calyx forming a distinct 5. Macarisia
tube divided into the calyx lobes only at the
upper part, lower part of tube hairy within; TRIBES AND GENERA OF RHIZOPHORACEAE
disk 0; stamens attached to mouth of the calyx
tube; stipules flat, not imbricate
I. T R I B E M A C A R I S I E A E Baill. (1876).
10. Pellacalyx
– Indumentum not stellate; free part of calyx
Shrubs or trees of moist or dry forest; stilt roots 0 or
divided to the base, inside glabrous in lower
poorly developed. Leaves toothed, verticillate or
part; disk annular or cupular-shaped, sur-
opposite; stipules valvate, pubescent. Inflores-
rounding the base of the style; stamens
cences lax-cymose or fasciculate. Flowers bisexual
attached to disk; stipules imbricate and over-
except in Sterigmapetalum and Blepharistemma
lapping 7
(?), diplostemonous (polyandrous in Cassipourea);
7. Young branches solid; leaves black dotted
ovary usually superior, 2–6-locular. Fruit a capsule,
beneath; stipules accompanied by stalked extra-
sometimes indehiscent. Seeds arillate or winged.
stipular glands; flowers usually in peduncled
A group of six genera, centred in Africa,
(rarely in sessile) cymes, 5–8-merous; petals
extending to Madagascar, Sri Lanka and India
erose or fimbriate; calyx lobes persistent and
and to South and Central America. Detailed infor-
terminating the fruit 7. Carallia
mation lacking for several genera.
– Young branches hollow; leaves not black dotted;
extrastipular glands 0; flowers in fascicles, 4–5-
merous; petals fringed with filamentous appen- 1. Blepharistemma Wall. ex Benth.
dages; calyx lobes persistent at the base of the fruit Blepharistemma Wall. ex Benth., J. Proc. Linn. Soc., Bot. 3:
9. Gynotroches 73, 78 (1858); Gamble, Fl. Madras 1: 460 (1997, re-edn);
8. Seeds arillate, the aril not exceeding the seed Sasidharan, Biodivers. documentation for Kerala, Flower-
ing Plts 6: 169 (2004); Saldanha, Fl. Karnatka 2: 57 (1996).
9
– Seeds winged, the wing usually exceeding the Polygamo-dioecious shrubs or small trees. Leaves
wing 12 opposite, apically crenate; stipules caducous.
9. Ovary inferior or semi-inferior, 3–28-locular; Inflorescences axillary, condensed cymes; pro-
stamens 2, 3 or more times the number of petals phylls 0 (?). Flowers small, white, 4-merous,
8. Crossostylis diplostemonous, somewhat dimorphic; petals
– Ovary superior or semi-inferior, 2–5-locular fringed apically, shorter in pistillate flowers;
10 disk 8-lobed; ovary 3-locular, superior, with
10. Hermaphrodite; flowers 4-merous, diplostemo- 2 ovules in each locule; the stigma somewhat
nous; ovary 2-locular 3. Comiphyton 3-lobed. Fruit subglobose, fleshy, variously
Rhizophoraceae 289
described as indehiscent or not. Seeds with aril Dactylopetalum Benth. (1858).

attached to upper half of seed. Petalodactylis Arènes (1954).
A single sp., B. serratum (Dennst.) Suresh,
Shrubs or trees, dioecious or hermaphrodite.
SW India (Western Ghats, North Malabar and
Leaves decussate or rarely whorled, entire to
South Kanara), in the understorey of wet ever-
serrate in apical half; stipules caducous. Inflor-
green forest up to 700 m.
escences axillary condensed cymes, or flowers
solitary. Flowers subsessile, (4)5(6)-merous;
2. Cassipourea Aubl. Fig. 67 calyx lobes valvate; petals white, inflexed in
Cassipourea Aubl., Pl. Guian.: 528, t. 211 (1775); Alston, bud, linear or spathulate, caducous, with
Kew Bull. 1925: 241–276 (1925), rev.; Floret, Bull. Mus. laciniate-fimbriate margins; stamens 2, 3 or
Natl. Hist. Nat. B, Adansonia 10: 25–45 (1985); Breteler, more times the number of petals, inserted
Edinb. J. Bot. 65: 407–424 (2008), synopsis of and key to outside or on the annular disk; ovary superior
subgenera.
to half-inferior, the base sometimes somewhat
Weihea Spreng. (1825).
fused with the calyx tube, often hairy, 2–5-
celled; ovules 2 per locule, pendulous and api-
cally inserted; style filiform or stout, the stigmas
enlarged and obscurely lobed. Fruit a somewhat
fleshy capsule, with septifragal dehiscence.
Seeds 2–6, compressed or angular, nearly
completely covered by an aril.
About 64 spp., ranging from tropical America
through tropical Africa to Madagascar, Sri Lanka
and S India; most spp. concentrated in West
Africa. Many occur in humid lowland forest, but
some grow in drier habitats and there is also a
considerably altitudinal range (to 2,500 m on Mt.
Ruwenzori).
For the division into seven subgenera, see
Floret (1985) and Breteler (2008). The justifica-
tion of the inclusion of Dactylopetalum in Cassi-
pourea by Alston (1925) has been doubted by
Floret (1976) and the genera are held distinct by
Juncosa and Tomlinson (1988a, b), but the mor-
phological separation is not clear cut and molec-
ular data are contradictory (Setoguchi et al. 1996
vs. Schwarzbach and Ricklefs 2000); see the
discussion in Breteler (2008).
3. Comiphyton Floret
Comiphyton Floret, Adansonia II, 14: 501 (1974), ibid. 16:
39–49 (1976).
Shrubs to tall trees. Leaves opposite, decussate,
Fig. 67. Rhizophoraceae. Cassipourea guianensis. distally denticulate; stipules interpetiolar. Inflor-
A Flowering branchlet. B Flower. C Same, transversal
section with pistil removed. D Petal, adaxial view. E Part escences axillary shortly pedunculate contracted
of androecium and disk in adaxial and abaxial view. cymes. Flowers perfect, small, 4-merous, diplos-
F Abaxial and adaxial views of anthers. G Pistil with temonous; calyx campanulate, with 4 short tri-
vertical section and transverse section of ovary. H Flower
past anthesis, the petals have fallen. I Part of axis with
angular teeth, other floral parts shortly exserted;
immature fruits. J Immature fruit. (Reproduced with kind petals distinct, linear, expanded apically into
permission of the artist Bobbi Angell) 5–6 pointed lobes; stamens inserted on a lobed
disk, inner (antepetalous) stamens longer than lobes persistent; petals slightly longer than the
the outer; anthers bilocular; ovary superior, 2- calyx lobes, caducous, distally with 4–13 spoon-
locular, each locule with 2 pendulous ovules shaped imbricate appendages; disk annular, 10-
inserted near the apex of the septum; style toothed; stamens inserted between the teeth of
long, exserted, with a bilobed capitate stigma. the disk; filaments distinct; ovary superior,
Fruit probably dehiscent, 1(2)-seeded. Seeds shortly stalked, 5-locular but with incomplete
arillate, the aril 2–3-lobed, attached to top of septa; ovules 2 per locule; style filiform; stigma
seed. capitate to obscurely lobed. Fruit a septicidal
A single sp., C. gabonense J.-J. Floret, Equato- capsule, somewhat woody. Seeds usually 10,
rial Africa (Gabon and Zaire), mainly of the forest with a membranous wing on the micropylar top.
understorey. Apparently not sharply differen- Seven species, restricted to Madagascar, in
tiated from Dactylopetalum. humid and semidry vegetation from lowland to
200 m above sea level.
4. Anopyxis (Pierre) Engl.
Anopyxis (Pierre) Engl. in E.-P., Nat. Pflanzenfam.,
6. Sterigmapetalum Kuhlm.
Nachtr. II–IV, 2: 48 (1900). Sterigmapetalum Kuhlm., Arch. Jard. Bot. Rio de Janeiro
Macarisia sect. Anopyxis Pierre (1898). 4: 359 (1925); Steyermark & Liesner, Ann. Missouri Bot.
Gard. 70: 179–193 (1983), rev.
Vast trees to 50 m or more. Leaves ternate or
verticillate, entire, glabrous but young leaves Dioecious trees to 30 m. Leaves decussate or in
sometimes covered with long hairs; stipules whorls of 3–4, entire or serrate; stipules glabrous
small, linear. Inflorescences axillary or terminal or sericeous. Inflorescence pedunculate, con-
contracted cymes on long peduncle. Flowers per- gested cymes with few to many flowers. Flowers
fect, 5-merous, obdiplostemonous; calyx cam- unisexual, 4–6-merous, diplostemonous; calyx
panulate with 5 bluntly pointed lobes; petals campanulate, the lobes valvate; petals linear and
entire or toothed, somewhat longer than the usually with 3 lobes elaborated in various ways;
calyx lobes; the stamen filaments connate over disk 0; stamens in staminate flowers alternately
their entire length into a long, slightly exserted long and short and inserted on a membranous
tube with the 10 small anthers at the top; disk 0; staminal tube within the petals; ovary rudimen-
ovary superior, flask-shaped, 5-locular; each loc- tary; pistillate flowers with reduced staminodial
ule with 2 ovules inserted axially or apically; style tube; ovary superior, 5–6-celled, with 2 ovules per
slender, the stigma slightly enlarged. Fruit a 5- cell; style short, expanded distally into a radially
valved woody hairy capsule with septicidal dehis- symmetrical discoid stigma. Fruit a septicidal
cence. Seeds up to 10, with a membranous wing 5–6-celled capsule. Seeds oblong, with a membra-
on the micropylar top. nous wing on the micropylar top.
A single sp., A. klaineana (Pierre) Engl., West About nine spp., northern South America
Africa from Sierra Leone to Zaire and Angola from E Colombia to Suriname and Brazilian
(Cabinda). Amazonia. Two subgenera, distinguished by the
presence or absence of a resinous exudate on the
buds and flower buds, this feature correlated with
5. Macarisia Thouars whorled versus opposite leaves (Steyermark and
Macarisia Thouars, Hist. Vég. Isles Austr. Afr.: 49, pl. 14 Liesner 1983).
(1806); Arènes, Fl. Madagascar Comores, Rhizophora-
ceae: 21–31 (1954); Schatz, Generic Tree Fl. Madagascar: II. T R I B E G Y N O T R O C H E A E Engl. (1892).
314 (2001).
Trees, with young parts hairy. Leaves entire or Shrubs and trees, some in disturbed vegetation;
glandular dentate; stipules caducous. Inflores- prominent aerial roots in all genera but
cences axillary, few-flowered pedunculate cymes, Pellacalyx. Leaves crenate, bijugate or (Pellaca-
or flowers solitary. Flowers 5(6)-merous, diplos- lyx) decussate. Flowers bisexual except in Gyno-
temonous; calyx at the base short tubular, the troches, diplostemonous; ovary inferior (superior
Rhizophoraceae 291
in Gynotroches), 5- or many-carpellate; ovules cymes with usually few (2–12) flowers. Flowers
2 or many per carpel (locules incompletely perfect or sometimes pistillate, sometimes large
separated). Fruit baccate, 1–many-seeded, (to 6 cm diam.), 4–6-merous, diplostemonous to
unappendaged (arillate in Crossostylis). polyandrous; petals apiculate, entire or slightly
Four genera, centred in Malesia, but extend- lobed or few-toothed; stamens 8–28, apparently
ing to Madagascar, India, northern Australia, arranged in one whorl; disk lobed, the lobes
and Fiji. opposite to or alternate with the stamens
(formerly interpreted as staminodia); ovary half-
inferior or inferior, 3- to 28-locular and incom-
7. Carallia Roxb.
pletely septate; ovules 2 per locule; style exserted,
Carallia Roxb., Pl. Corom. 3: 8, t. 211 (1811), nom cons.; stigma discoid or 4–many-lobed with linear to
Ding Hou, Fl. Males. II, 5: 481–488 (1958).
filiform stigmatose lobes. Fruit a conical capsule,
Shrubs or trees to 50 m, sometimes with small enclosed by the calyx, dehiscence tardy with
buttresses or stilt-roots. Leaves bijugate, black- distal slits or an operculum. Seeds numerous,
dotted beneath, entire to dentate or serrate; sti- arillate, the aril attached to the micropylar top
pules large, lanceolate, caducous, accompanied of the seed.
by large stalked glands. Inflorescences resinous, About 12 spp., six of them in the Solomon
sessile or peduncled, lax-cymose or condensed, Islands, Vanuatu and Fiji, and the others in New
di- or trichotomously branched, or sometimes Caledonia and Polynesia. With emphasis on seed
reduced to 2 or 1 flowers; prophylls distinct and morphology, Tobe and Raven (1988) suggest the
caducous or connate and persistent. Flowers ses- erection of a monotypic tribe to accommodate
sile or pedicelled, small, perfect, 5–8-merous, this genus.
obdiplostemonous; calyx deeply divided into del-
toid lobes; petals caducous or persistent, nar- 9. Gynotroches Blume
rowed below but the margins and apex of the
expanded distal portion lobed or fimbriate; Gynotroches Blume, Bijdr.: 218 (1825); Ding Hou, Fl.
Males. II, 5: 488–489 (1958).
stamens distinct or shortly connate into a tube,
persistent; disk within the stamens, annular, Dioecious shrubs or trees up to 40 m; young
sometimes lobulate, fleshy; ovary inferior, either branches hollow. Leaves bijugate, entire to
5–8-celled and each cell 2-ovulate, or 1-celled denticulate; stipules lanceolate. Inflorescences
with 10–12 ovules; style filiform or slightly coni- fasciculate or flowers solitary. Flowers unisexual
cal, stigma discoid, capitate, or obscurely lobed. by abortion, 4–5-merous, diplostemonous; calyx
Fruit a globose to obovoid berry, 1-celled, 1(5)- deeply lobed; petals unguiculate, with distal fila-
seeded. Seeds ellipsoid or reniform. mentous appendages; stamens distinct, inserted
Eleven spp., Madagascar, India, Nepal, Indo- at the margin of the calyx cup and outside the
china, S China, Malesia, Solomon Islands, and N shallowly lobed disk, sterile in pistillate flowers;
Australia, in (sub)humid forest up into the ovary superior (pistillate flowers), 4–6-celled,
mountains to c. 1,800 m. Carallia brachiata each cell with 3–8 ovules, or half-inferior, with
(Lour.) Merr. has the widest range of the genus sterile ovules (staminate flowers); style simple,
from Madagascar to the Solomons; its timber is very short in staminate flowers, stigma discoid
commercially valuable. or 4–6-lobed; staminate flowers with a reduced
gynoecium. Fruit a globose to oblong berry. Seeds
few to many.
8. Crossostylis J.R. & G. Forst.
One sp., G. axillaris Blume (or perhaps sev-
Crossostylis J.R. & G. Forst., Char. Gen. Pl.: 44 (1775); eral ones: Ding Hou 1958; Juncosa and Tobe
Setoguchi et al., J. Plant Res. 109: 7–19 (1996), floral
morph.
1988), in non-seasonal parts of southeast Asia
from Myanmar through Malesia to northern
Trees up to 20 m or shrubs, sometimes polygamo- Australia, the Solomon Islands and into Micro-
dioecious. Leaves bijugate, glabrous, entire or nesia, in forests and along rivers, occasionally in
serrulate. Inflorescences axillary condensed secondary forests, up to c. 2,000 m.
10. Pellacalyx Korth.

Pellacalyx Korth., Tijd. Nat. Gesch. Phys. 3: 20 (1836);
Ding Hou, Fl. Males. I, 5: 490–493 (1958).
Trees up to 45 m; young branches hollow.
Leaves decussate, entire, serrulate or serrate,
pubescent with stellate or simple hairs, or gla-
brous; stipules flat, valvate. Inflorescences axil-
lary, few-flowered, scarcely pedunculate
glomerules, or flowers occasionally solitary;
pairs of prophylls fused into a toothed cup.
Flowers perfect, 4–5-merous, obdiplostemonous,
small; calyx deeply tubular, hairy inside, apically
with deltoid lobes; petals distinct, inserted on
the margin of the calyx tube, denticulate or
fringed; stamens attached to mouth of calyx
tube with very short filaments in two series or
one; ovary inferior, with 9–10(12) locules, each
locule with 8–25 ovules in a cluster on the axile
placenta; style columnar, not or scarcely Fig. 68. Rhizophoraceae. Bruguiera exaristata. A Flower-
ing branchlet. B–D Petals, lateral, ventral (with 2 stamens),
exserted, with a capitate, slightly lobed stigma. and dorsal. E Stamen. F Flower, vertical section, petals
Fruit a few- to many-seeded berry. Seeds small. and stamens removed. G Ovary, transversal section. H, I
About 7 spp., distributed from Myanmar to S Hypocotyls. (Ding Hou 1958; drawn by Ruth van Crevel)
China and Malesia. The genus seems to fall natu-
rally into one group of species with predomi- Trees with trunks with sturdy basal buttresses and
nantly tetramerous and small flowers (e.g. with knobby pneumatophores produced from hor-
P. lobbii (Hook. f.) Schimp.) and another with izontal roots, and sometimes with aerial roots
predominantly pentamerous and larger flowers when young. Leaves bijugate, entire; stipules
(e.g. P. axillaris Korth.). lanceolate, glabous, to 4 cm long. Inflorescences
cymose with numerous small flowers or reduced to
1–3 large flowers, then often without prophylls.
III. T R I B E R H I Z O P H O R E A E Bartl. (1830). Flowers bisexual, 4–16-merous, diplostemonous;
petals 2-lobed, often with reduced distal appen-
Mangrove shrubs or trees. Aerial stilt roots always dages, each embracing a pair of stamens that are
present but prominent only in Rhizophora. released explosively when stimulated by a probing
Leaves bijugate, entire. Inflorescences cymose. pollinator, the petals being held together by mar-
Flowers 4–16-merous, mostly diplostemonous; ginal interlocking hairs that unzip instantly to
petals usually enclosing one to several stamens; allow pollen discharge; disk cup-shaped, adnate
ovary half to fully inferior, 2–3-carpellate; ovules to calyx tube; ovary deeply inferior, adnate to
2 per carpel. Fruit baccate, fibrous, 1-seeded; lower part of calyx tube, 2–4-celled, each cell with
germination viviparous, the huge seedling axis 2 ovules; style filiform; stigma obscurely 2–4-
(up to 1 m) emerging from both the seed coat lobed. Fruit usually 1-celled. Seeds 1(2); hypocotyl
and the fruit up to 9 months before abscission. terete or obscurely ribbed, perforating the apex of
Four genera, the tribe pantropical. the fruit and falling with it.
Six spp., from south-east Africa through
11. Bruguiera Savigny Fig. 68 Malesia to northern Australia and the Pacific Islands
(Samoa). The genus has been divided into "large-
Bruguiera Savigny, Encycl. (Lamarck) 4: 696 (1798); Ding
Hou, Nova Guinea II, 8: 163–171 (1957), Fl. Males. II, 5: flowered" (e.g. B. gymnorrhiza) and "small-flow-
457–468 (1958); Duke & Ge, Blumea 56: 36–48 (2011), ered" (e.g. B. cylindrical) species, obviously in rela-
hybridization. tion to the size of the pollinators (birds vs. insects).
Rhizophoraceae 293
12. Kandelia (DC.) Wight & Arn. Two spp., both wide-ranging from E Africa to
Kandelia (DC.) Wight & Arn., Prodr. Fl. Ind. Orient.: 310 Queensland, Melanesia and Micronesia north to
(1834); Ding Hou, Fl. Males. II, 5: 472–473 (1958). Hong Kong; they are strongly salt-tolerant and
grow preferentially in the back-mangal, often at
Small trees to 7 m; buttresses and pneumato- the margins of saltpans in the higher tidal
phores 0. Leaves decussate or bijugate, entire; reaches. The species differ in the non-explosive
stipules linear. Inflorescences axillary, long and explosive methods of pollen discharge, the
pedunculate, 4–9-flowered. Flowers perfect, pen- latter presumably evolved independently from
tamerous, polyandrous; calyx deeply lobed, sub- that in Bruguiera.
tended by a cuplike prophyllar involucre, lobes
long and reflexed at anthesis; petals bilobed, with
long seta in the sinus, each lobe multifid; stamens 14. Rhizophora L.
numerous (30–40) and inserted on the rim of the Rhizophora L., Sp. Pl.: 443 (1753), Gen. Pl. ed. 5: 202
calyx cup, the filaments uneven in length and (1754); Salvoza, F.M., Nat. Appl. Sci. Bull. Un. Philipp. 5:
with minute anthers; ovary inferior below a 179–237 (1936); Ding Hou, Fl. Males. II, 5: 448–457 (1958),
well-developed nectar cup with a pronounced Blumea 10: 625–634 (1960).
intrastaminal rim; ovary half-inferior, 1-celled; Trees; stems supported by numerous branched
ovules 6, attached apically to a pronounced stilt roots; most vegetative parts contain tricho-
basal extension of the ovary wall; style filiform, sclereids. Leaves bijugate, with numerous micro-
the stigma minutely 3-lobed. Fruit pedunculate scopic black cork-warts below; stipules
with persistent reflexed sepals. Seed 1, viviparous; lanceolate. Inflorescences peduncled and di- or
hypocotyl up to 40 cm long. trichotomously branched cymes. Flowers 4-
Two spp., one distributed from Bangladesh to merous, stamens 2(3) times the number of petals;
Borneo, the other from China to S Japan calyx deeply lobed, subtended by cup-shaped
(Kyushu), typically growing in back-mangrove prophylls; petals inserted at base of disk, entire,
communities and on banks of tidal rivers. ephemerous, the edges barely enclosing the single
antepetalous stamen; stamens (sub)sessile, multi-
13. Ceriops Arn. locellate, dehiscing introrsely; ovary half-inferior,
2-celled; ovules 2 per cell; style obscure or dis-
Ceriops Arn., Ann. Mag. Nat. Hist. 1: 363 (1838); Ding
Hou, Fl. Males. I, 5: 468–472 (1958).
tinct; stigma 2-lobed. Fruit ovoid. Seeds 1(2, 3),
viviparous; hypocotyl maturing on tree to 15–30
Small to moderate-sized trees with a characteristic (70) cm; cotyledons fused to form an emergent
basal cone of appressed aerial stilt roots and some- collar from which the seedling abscises at matu-
times with knobby pneumatophores produced rity. Wind-pollinated.
from horizontal roots. Leaves bijugate, entire; sti- About 8 spp. of which at least two appear to
pules lanceolate. Inflorescences subsessile to be hybrids. The genus as a whole is pantropical
shortly peduncled trichasial to monochasial but there is an almost complete geographic sepa-
cymes. Flowers 5(6)-merous, diplostemonous, ration between Old World and New World
their prophylls partly connate; calyx deeply species (Tomlinson 1986), with the exception of
lobed; petals keeled, emarginate or truncate, the western Pacific (Fiji and New Caledonia).
fringe-like divided or with 3 clavate appendages,
enwrapping 2 stamens or not; disk shallowly
lobed, lobes antesepalous; stamens inserted in the Doubtfully attached:
sinuations of the disk; ovary half-inferior, 3-celled,
each cell with 2 ovules; style simple; stigma simple 14a. Paradrypetes Kuhlm.
or obscurely 2–3-lobed. Fruits conical, mainly by
Paradrypetes Kuhlm., Arq. Inst. Biol. Veg. Rio Jan. 2: 84
extension of the upper part of ovary. Seed solitary, (1935); Levin, Syst. Bot. 17: 74–83 (1992); Webster, Ann.
viviparous, hypocotyl up to 20 cm long, terete or Missouri Bot. Gard. 81: 56 (1994); Radcl.-Sm., Gen.
ridged, released from the cotyledon at dispersal. Euphorb.: 87 (2001).
Dioecious trees; indumentum 0. Leaves opposite, Bentham, G., Hooker, J.D. 1865. Genera Plantarum, Vol. 1,
simple, spinulose-dentate or subentire; stipules Part 2. London: L. Reeve.
Brown, R. 1814. General remarks, geographical and sys-
sheathing, interpetiolar, with colleters. Inflores- tematical, on the botany of Terra Australis. Appen-
cences cymose, ebracteate, the peduncle in both dix III. Pp. 533–613. In: M.A. Flinders, Voyage to
sexes adnate to subtending petiole or not. Flowers Terra Australis. London: G. & W. Nicol.
apetalous; staminate flowers pedicellate; sepals 3 Collinson, M.E. 1983. Fossil plants of the London Clay. A
field guide to fossils. No. 1. London: The Palaeonto-
or 4, distinct, imbricate; disk 0; stamens 10–12, logical Association.
filaments suppressed; anthers extrorse, dehiscing Ding Hou 1957. A conspectus of the genus Bruguiera
longitudinally; pollen grains oblate, 4-brevicolpo- (Rhizophoraceae). Nova. Guinea, n.s. 8: 163–171.
rate, exine echinate, tectum psilate-wrinkled; Ding Hou 1958. Rhizophoraceae. Flora Malesiana I, 5:
429–493.
pistillode 0; pistillate flowers solitary or in 3-flow- Ding Hou 1960. A review of the genus Rhizophora with
ered dichasia; sepals 4, caducous; disk annular; special reference to the Pacific species. Blumea 10:
ovary 3-locular, ovules anatropous; stigmas 3, 625–634.
Duke, N.C. 2010. Overlap of eastern and western man-
dilated, sessile. Fruits drupaceous with crusta- groves in the south-western Pacific: hybridization of
ceous endocarp, 1-seeded. Seeds ecarunculate; all three Rhizophora (Rhizophoraceae) combinations
testa reticulately veined; endosperm abundant; in New Caledonia. Blumea 55: 171–188.
cotyledons chlorophyllous, plicate, much longer Floret, J.-J. 1974. Comiophyton, genre nouveau gabonais
Rhizophoraceae-Macarisieae. Adansonia II, 14:
and broader than radicle. 499–506.
Two Brazilian spp., one in central Amazonia, Floret, J.-J. 1976. A propos de Comiophyton gabonense
the other in the Atlantic coastal forests. (Rhizophoraceae). Adansonia II, 16: 39–49.
Hutchinson (1969) was the first to refer Graham, A. 2006. Paleobotanical evidence and molecular
data in reconstructing the historical phytogeography
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placement in tribe Podocalcyeae was supported 327–334.
by the palynological study of Levin and Simpson Hegnauer, R. 1973. .See General References.
(1994). Paradrypetes shares with Podocalyx the Hisahi, Y. et al. 1984. Karyomorphological studies in five
species of mangrove genera in the Rhizophoraceae.
unique synapomorphy of the absence of distinct La Kromosomo II, 35/36: 1115–1116.
columellae in the interstitium between endexine Hutchinson, J. 1969. Tribalism in the family Euphorbia-
and ectexine. However, molecular studies by ceae. Amer. J. Bot. 56: 738–758.
Wurdack et al. (2004) and Wurdack and Davis Juncosa, A.M. 1982. Developmental morphology of the
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habit between Paradrypetes and genera such as opment in Cassipourea elliptica (Sw.) Poit. (Rhizo-
phoraceae). Amer. J. Bot. 71: 170–179.
Cassipourea, and the opposite leaves, interpetiolar Juncosa, A.M. 1984b. Embryogenesis and developmental
stipules with colleters and chlorophyllous embryos morphology of the seedling in Bruguiera exaristata
are rhizophoraceous rather than euphorbialean, Ding Hou (Rhizophoraceae). Amer. J. Bot. 71:
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Juncosa, A.M., Tobe, H. 1988. Embryology of the tribe
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and sessile stigmas as Paradrypetes (see Levin Bot. Gard. 75: 1410–1424.
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Trigoniaceae
Trigoniaceae A. Juss. in Orbigny, Dict. univ. hist. nat. 12: 670 (1849), nom. cons.
V. B I T T R I C H
Trees, shrubs, scandent shrubs or lianas, glabrous or hairs on the leaf, stipule, bract and bracteole
or with indumentum of unicellular hairs, some- margins also occur in some Trigonia species
times with glands. Leaves opposite or alternate, (e.g. T. reticulata).
simple, entire, venation pinnate; stipules simple Trichomes are reported mainly as simple,
or bifid, interpetiolar when leaves opposite, often unicellular; sometimes a more or less dense
conspicuous, mostly caducous. Inflorescences arachnoid indumentum of twisted hairs is found,
panicles, racemes, cymes, or thyrses. Flowers her- especially on the lower leaf surface; 2-armed
maphrodite, small, papilionaceous, obliquely unicellular trichomes, similar as in Malpighia-
zygomorphic, 5-merous; sepals unequal, imbri- ceae, were reported by Rao and Sharma (1992).
cate, connate at base or partly distinct; petals dis- Mucilaginous cells are common in the epidermis,
tinct, the posterior forming a saccate or spurred which is uniseriate in Trigonia and two-layered
standard; stamens and staminodes connate basally in Humbertiodendron and Trigoniastrum. The
into a tube that is more strongly developed on the paracytic stomata are restricted to the abaxial
anterior part of the flower by the 4–8 fertile leaf surface. Branched sclereids are distri-
stamens, staminodes 0 or up to 6 in the posterior buted in the mesophyll of Humbertiodendron and
part, anthers opening longitudinally; nectary Trigoniastrum. Rhomboid or irregularly shaped
glands 1–2(–4) opposite to the standard petal or crystals are common.
0; ovary superior or partly inferior, 3(4)-locular, Information on stem anatomy is found in
rarely unilocular, pubescent, each locule with Metcalfe and Chalk (1950), Bridgwater and Baas
1–numerous ovules; style simple. Fruits septicidal (1982), and Carlquist (1988). The cork of the
capsules or indehiscent 3(4)-winged, sometimes stem is derived from a subepidermal phellogen.
easily splitting into 1-winged samaras. Seeds Cortical sclereids are especially common in
hairy or not, embryo straight, endosperm + or 0. Humbertiodendron and Trigoniastrum. Vessels
A pantropical family of five genera and about are solitary or in radial bands, the perforations
30 species, centred in South America, but also in simple or rarely scalariform, the parenchyma
Malesia and Madagascar. generally apotracheal and irregular (Trigonias-
trum) or banded (Trigonia); scanty paratra-
VEGETATIVE MORPHOLOGY AND ANATOMY. The cheal parenchyma was reported from
available information is summarized mainly in Trigoniastrum. Petersen (1896) described
Lleras (1978). The species of Humbertiodendron, medullar vascular bundles with internal phloem
Isidodendron, Trigoniastrum and Trigoniodendron forming an interrupted ring from an unidenti-
form middle-sized or large trees, while treelets, fied Trigonia species. Crystals occur in cham-
shrubs, scandent shrubs or lianas are typical bered axial and ray parenchyma. The vessels
for Trigonia. The branches are generally terete have bordered pits to the ground tissue of the
and lenticellate. Inconspicuous pilose domatia xylem; Carlquist (2007) reported bordered pits
occur on the leaf blades of Isidodendron. Leaf from the axial parenchyma of Trigoniastrum
margin and the abaxial surface of the bracts hypoleucum, evident especially on the cross-
have impressed glands in Trigoniastrum; Stevens walls. True tracheids and fibre tracheids are
(2001) mentions marginal glands towards the leaf present, but libriform fibres and intraxylary
bases for Humbertiodendron; glandular papillae phloem are absent. The rays are 1–5-seriate
298 V. Bittrich
and heterogeneous (Kribs’ type II). Sieve ele- opposite the standard, they are sometimes lobed
ment plastids contain starch grains (type Ss, or laciniate, often puberulous. Such nectary
Behnke 1991). glands are absent in Isidodendron, possibly the
swollen bases of its staminodes secrete nectar.
F L O W E R S T R U C T U R E . The flowers are united in The ovary is superior or partly inferior, hairy,
panicles, racemes, cymes or thyrses, the pedicels 3(4)-locular or 1-locular by reduction of the
are articulate. The buds of the short branches septa, placentation axile or axile-apical or in uni-
bearing the simple racemes of Isidodendron are locular ovaries parietal on more or less deeply
protected by perulae. A detailed analysis of the intruding placentas with a central column absent;
flower structure of Trigonia rugosa and Trigo- the ovules are epitropous or antitropous and
niastrum hypoleucum is provided in Matthews horizontal (Trigonia), pendulous (Trigoniastrum)
and Endress (2008), who also compared the or erect (Humbertiodendron). The style is terminal,
results with their data from closely related simple, sometimes pubescent, the stigma capitate
families. The flowers are obliquely zygomorphic or rarely trilobate or circular, rarely punctate,
with the plane of symmetry through the third its surface papillate. The ovary and lowest part
sepal, hypogynous or slightly perigynous (Eichler of the common style were found to be completely
1878). The sepals are unequal (the inner larger) synascidiate in Trigoniastrum, with the carpel
with quincuncial aestivation, congenitally united tips in the style distinct or postgenitally connate
at base or two sepals free on the posterior side of for most of their length; in Trigonia, however,
the flower, and generally have a strigose indu- only the lower 2/3 of the ovary is synascidiate,
mentum. Their margin is glandular in Trigonio- but symplicate above; in the locules of Trigonia
dendron. The corolla is papilionaceous and an obturator is formed by unicellular papillae
consists of five petals, which are contorted protruding from the funicle, while in Trigonias-
in bud. The posterior (standard) and the wing trum long, unicellular hairs are present on top of
petals are mostly hairy at base, but glabrous the locule (Matthews and Endress 2008).
in Trigoniodendron. The standard petal is sac-
cate or shortly spurred and bulges outwards E M B R Y O L O G Y . Only a few species of Trigonia
between the two free posterior sepals (observed were studied embryologically (Mauritzon 1936;
by Matthews and Endress 2008 in Trigonia and Boesewinkel 1987). The ovule primordium is
Trigoniastrum); the two anterior petals are often trizonate. Endosperm formation of Trigonia is
saccate and form a keel. The petal epidermis is initially nuclear, becoming cellular throughout.
epithelium-like in Trigonia and Trigoniastrum The nucellus becomes resorbed by the ovule
(Matthews and Endress 2008). The filaments of and the mature ovules are bitegmic, anatropous,
the four to eight fertile stamens and the one to six and epitropous or rarely apotropous, and tenui-
staminodes, if present, are fused into an nucellate (the report of crassinucellate ovules
unequally developed tube, much higher so in the by Mauritzon (1936) is dubious and needs confir-
anterior part of the flower, where the fertile mation). The embryo sac is probably of the
stamens are positioned; studying young buds, Polygonum type. The integuments are of dermal
Matthews and Endress (2008) concluded that the origin, the inner is strongly multiplicative.
fertile stamens are basically arranged in two
series; the anthers are tetrasporangiate, nearly P O L L E N M O R P H O L O G Y . Pollen grains are 3–4
basifixed and with a dorsal pit at the insertion of (5)-porate, 25–60 mm long with the pollen size
the filament, they open introrsely by longitudinal being highly variable within the same species;
slits; the endothecium is continuous over the exine is tenui-exinous and psilate (Lleras
the dorsal side of the connective (Matthews 1978).
and Endress 2008). Staminodes are long and
pointed or more rudimental, the latter become K A R Y O L O G Y . Trigonia virens has 2n ¼ ca. 20.
incorporated into the nectary glands; anthers
rudiments are always lacking. The one or two P O L L I N A T I O N . The basically papilionaceous
nectary glands (or up to four according to Eichler flower typical for the family suggests insect-,
1878), possibly of staminodial origin, are inserted probably bee-pollination, but observations are
Trigoniaceae 299
nearly completely lacking. Also the glands in the various anatomical characters in common, form
flowers suggest nectar as reward, but Lleras a monophyletic subgroup.
(1978) was unable to find any traces of it in A couple of families including Polygalaceae,
flowers of Trigonia spp. Chrysobalanaceae, Dichapetalaceae, Sapindaceae
and Vochysiaceae were considered as next related
F R U I T A N D S E E D . Trigonia and Trigonioden- to Trigoniaceae. Trigoniastrum was sometimes
dron have septicidal capsules, which are hairy included in Polygalaceae. Based on ovule and
outside and sometimes inside, the hard endocarp seed structure, Boesewinkel and Venturelli
generally separates from the exocarp at maturity; (1987) suggested a close relationship of Trigonia-
Isidodendron, Humbertiodendron and Trigonias- ceae with the families of Linales (Malpighiales),
trum have 3-winged fruits which easily split into but also Vochysiaceae (Myrtales). Vochysiaceae
1-winged samaras in Humbertiodendron and were most often considered as the closest rela-
Trigoniastrum. The seed coat is mostly covered tives, and various differences in flower morphol-
with long often silky unicellular hairs, but is gla- ogy between the two families were already
brous in Humbertiodendron and probably Isido- discussed by Petersen (1896). Lleras (1978) men-
dendron, the seeds of which are reported to be tioned anatomical evidence, like a simple petiole
alate. Only the seed anatomy of Trigonia and epidermis, general lack of libriform fibres and
Trigoniastrum was studied in detail (Boesewinkel pith without sclereids in Trigoniaceae. The genus
1987; Matthews and Endress 2008). The seeds are Euphronia was formerly often considered a mem-
bitegmic. The outer integument is 2–3 cell layers, ber of Trigoniaceae, but rbcL sequence data
and the inner 3–6 layers thick. The exotegmen is revealed Euphronia as sister to Chrysobalanaceae
fibrous and lignified, the endotegmen tanninifer- (Litt and Chase 1999). The analysis based on com-
ous. The micropyle was described as zigzag for bined rbcL and non-molecular datasets by Nandi
Trigonia cipoensis (Boesewinkel 1987); in Trigo- et al. (1998) depict Chrysobalanaceae and Dicha-
niastrum it is formed by the inner integument petalaceae as the closest relatives of Trigoniaceae
only (Matthews and Endress 2008). The raphal within Malpighiales. The cladogram of Litt and
bundle is amphicribral and complex. An endo- Chase (1999) shows Trigoniaceae as sister group
sperm rich in fat is present in Trigonia but of Dichapetalaceae, both together forming the sis-
absent in Humbertiodendron and Trigoniastrum. ter group of Euphroniaceae + Chrysobalanaceae,
The embryo is straight and positioned longitudi- a topology confirmed by Wurdack and Davis
nally or transversely to the length of the seed, (2009), Soltis et al. (2011) and Xi et al. (2012).
the cotyledons are flat but rather thick in This family group, first proposed by Hallier
Trigoniodendron. (1921), was object of a careful study of floral
structures revealing numerous characters of taxo-
D I S P E R S A L . The species with winged fruits are nomic importance (see Matthews and Endress
probably anemochorous. Nearly all species with 2008). The four families share the occurrence of
capsular fruits have long-haired seeds which may obliquely zygomorphic flowers, sepals congenitally
be adapted to anemochory as well as to hydro- united at base, tenuinucellate ovules, and the
chory. The seeds of two Trigonia species (T. spru- occurrence of a zigzag micropyle (Trigonia). Tri-
ceana and T. hypoleuca) have echinate trichomes goniaceae and Dichapetalaceae also share various
and are possibly hydrochorous. Hydrochory and floral characters, such as special mucilage cells in
anemochory match with the typical habitats of the mesophyll of the sepals, ovary and lower style
Trigonia species. sometimes completely synascidiate (Trigonias-
trum), nectary semi-annular and with distinct
P H Y T O C H E M I S T R Y . The plants are often tanni- scales or lobes.
niferous (stem and seeds); they show no accumu-
lation of aluminium (Hegnauer 1973). D I S T R I B U T I O N A N D H A B I T A T S . The family
shows an interesting amphipacific distribution:
S U B D I V I S I O N A N D A F F I N I T I E S . Most probably Humbertiodendron is restricted to Madagascar,
the palaeotropical genera Humbertiodendron Trigoniastrum to Indonesia and Malaysia, and
and Trigoniastrum, with 3-winged fruits and Isidodendron, Trigonia and Trigoniodendron to
300 V. Bittrich
the Neotropics, with Trigoniodendron being Trees up to 30 m; bark with a yellowish gummy
endemic in coastal Espı́rito Santo (Brazil) and juice. Leaves alternate, acuminate, with margins
Isidodendron in the Rio Magdalena valley in cen- and acumen generally glandular, thinly arachnoid
tral Colombia. Possible scenarios to explain the pubescent below; stipules caducous. Flowers in
distribution pattern of Trigonia species are dis- axillary or terminal panicles, bracts with marginal
cussed by Lleras (1978). With exception of Trigo- glands; standard slightly pilose externally;
nia, the genera occur in moist tropical forests. stamens 6, rudimental staminodes +; glands
Trigonia species are found in periodically flooded 1–2, puberulous; ovary hairy, 3-locular with
riverine forests, gallery forests, edges of wet for- 2 ovules per locule. Fruit 3-winged, splitting into
ests, and disturbed sites. 1-winged samaras. Seeds velutinous.
One sp., T. hypoleucum Miq., from tropical
E C O N O M I C I M P O R T A N C E . The family has little rainforests of Indonesia, Malaysia, Singapore and
economic importance. The hard wood of Trigo- Brunei.
niastrum hypoleucum is used for furniture
making. 3. Trigonia Aubl. Fig. 69
Trigonia Aubl., Hist. Pl. Guiane: 387 (1775); Lleras, Fl.
KEY TO THE GENERA Neotrop. Monogr. 19: 29–62 (1978), rev.
1. Leaves alternate 2 Treelets, shrubs, scandent shrubs, or lianas.
– Leaves opposite 4 Leaves opposite, often with white indument
2. Leaves and bracts with impressed marginal glands; below, stipules interpetiolar, entire or apically
stamens 6. Malesia 2. Trigoniastrum
bifid, caducous or rarely persistent. Flowers in
– Leaves and bracts glandless; stamens 7. South America
thyrses or panicles, bract margins glandular or
3
3. Flowers in panicles; staminodes 0; fruit a capsule eglandular; standard and wing petals mostly
4. Trigoniodendron hairy at base; fertile stamens 4–8, staminodes up
– Flower in racemes; staminodes 4–6; fruit 3-winged, to 6 or 0; glands 2(–4), pilose or glabrous; ovary
indehiscent 5. Isidodendron 3–4-carpellate, sometimes 1-locular due to reduc-
4. Floral gland 1; ovule 1 per locule; fruit 3-winged tion of the septa; ovules few to numerous per
dehiscing into1-winged samaras; seeds glabrous. locule. Fruit a septicidal, often trigonous, outside
Madagascar 1. Humbertiodendron
mostly pilose capsule. Seeds pilose.
– Floral glands 2; ovules more than one per locule,
fruit a 3(4)-valvate capsule; seeds hairy. Neotropics 24 spp., trop. and subtrop. America, tempo-
3. Trigonia rarily inundated forests, gallery forests, open for-
est edges, and disturbed areas.
GENERA OF TRIGONIACEAE
4. Trigoniodendron E.F. Guim. & Miguel
1. Humbertiodendron Leandri Trigoniodendron E.F. Guim. & Miguel, Rev. Brasil. Biol.
47: 559 (1987).
Humbertiodendron Leandri, Compt. Rend. Hebd. Séances
Acad. Sci. 229: 848 (1949). Trees. Leaves alternate, stipules entire, caducous.
Small trees. Leaves opposite, stipules connate. Flowers in terminal panicles; margin of sepals
Flowers in axillary triflorate cymes, bracts glandular, petals glabrous, stamens 7, filaments
eglandular; petals slightly strigose externally; connate in the lower half, staminodes 0, glands 2;
stamens 6; 1 fleshy pilose gland adpressed to the ovary lanuginose. Capsule globose, septicidal, 3-
ovary; ovary pubescent, 3-winged, trilocular, with valvate. Seeds hairy, cotyledons thick.
1 ovule per locule. Fruit 3-winged, dehiscing into One sp., T. spiritusanctense E.F. Guim. &
1-winged samaras. Seeds glabrous. Miguel, known only from the Atlantic rainforest
One sp., H. saboureaui Leandri, from coastal of Espı́rito Santo state, Brazil.
forests of E Madagascar.
5. Isidodendron Fern. Alonso, Pérez-Zab. &
2. Trigoniastrum Miq. Idagarra
Trigoniastrum Miq., Fl. Ned. Ind., Eerste Bijv.: 394 (1861), Isidodendron Fern. Alonso, Pérez-Zab. & Idagarra, Revista
nom. cons. Acad. Colomb. Ci. Exact. 24: 348 (2000).
Trigoniaceae 301
One species, I. tripterocarpum Fern.-Alonso,

Pérez-Zab. & Idagarra, known only from the
moist forests of the Rio Magdalena valley, central
Colombia.
Behnke, H.-D. 1991. See General References.

Boesewinkel, F.D. 1987. Ovules and seeds of Trigoniaceae.
Acta. Bot. Neerl. 36: 81–91.
Boesewinkel, F.D., Venturelli, M. 1987. Ovule and seed
structure in Vochysiaceae. Bot. Jahrb. Syst. 108:
547–566.
Bridgwater, S., Baas, P. 1982. Wood anatomy of Xantho-
phyllum Roxb. IAWA Bull. n.s. 3: 115–125.
Carlquist, S. 1988. Comparative wood anatomy. Berlin,
Heidelberg, New York: Springer.
Carlquist, S. 2007. Bordered pits in ray cells and axial
parenchyma: the histology of conduction, storage,
and strength in living wood cells. Bot. J. Linn. Soc.
153: 157–168.
Chase, M.W. et al. 1993. See General References.
Eichler, A.W. 1878. Bl€ uthendiagramme, 2. Teil. Leipzig:
Wilhelm Engelmann, pp. 343–345.
Hallier, H. 1921. Beitr€age zur Kenntnis der Linaceen
(DC. 1819) Dumort. Beihefte Bot. Centralbl., Abt. 2,
39: 1–178.
Hegnauer, R. 1973. .See General References.
Litt, A.J., Chase, M.W. 1999. The systematic position of
Euphronia, with comments on the position of Bala-
nops: an analysis based on rbcL sequence data. Syst.
Bot. 23: 401–409.
Lleras, E. 1978. Trigoniaceae. Fl. Neotrop. Monogr. 19:
1–73.
Fig. 69. Trigoniaceae. Trigonia villosa. A Flowering Matthews, M.L., Endress, P.K. 2008. See General
branch. B Flower bud. C Flower, oblique-apical view. References.
D Flower after petals have fallen. E Adaxial and lateral Mauritzon, J. 1936. Zur Embryologie und systematischen
views of lateral petal. F Adaxial and lateral views of ante- Abgrenzung der Reihen Terebinthales and Celas-
rior petal. G Three views and medial section of saccate trales. Bot. Notiser (1936): 161–212.
posterior petal. H Abaxial and adaxial views of anthers Metcalfe, C.R., Chalk, L. 1950. See General References.
and entire androecium. I Lateral view of pistil with basal Nandi, O., Chase, M.W., Endress, P.K. 1998. A combined
nectary and medial section thereof. J Fruits. K Seed with cladistic analysis of angiosperms using rbcL and
hairs and with hairs removed. (Reproduced with kind non-molecular data sets. Ann. Missouri Bot. Gard.
permission of the artist Bobbi Angell) 85: 137–212.
Petersen, O.G. 1896. Trigoniaceae. In: Engler & Prantl,
Nat€url. Pflanzenfam. 3(4): 309–311.
Rao, S.R.S., Sharma, V. 1992. Morphology of 2-armed
Trees. Leaves alternate. Flowers in racemes on per- trichomes in relation to taxonomy: Malpighiales.
ulate short branches; petals glabrous, stamens 7, Feddes Repertorium 103: 55–565.
filaments connate in the lower half, staminodes Soltis, D.E. et al. 2011. See General References.
Stevens, P.F. 2001 onwards. See General References.
4–6, carnose at base, glands 0; ovary lanuginose. Wurdack, K.J., Davis, C.C. 2009. See General References.
Fruit 3-winged. Seeds winged. Xi, Z. et al. 2012. See General References.
Violaceae
Violaceae Batsch, Tab. Affin. Regni Veg.: 57 (1802), nom cons.
H.E. B A L L A R D , J. DE P A U L A -S O U Z A , AND G.A. W A H L E R T
Trees, treelets, shrubs or subshrubs, infrequently 1–many, anatropous, bitegmic, crassinucellate;

lianas or herbs, sometimes stoloniferous. Leaves style filiform or sometimes clavate, straight, curved
alternate, sometimes opposite, distichous or or sigmoid; stigma often simple, less often rostrate,
pseudo-whorled, pinnately or rarely palmately orifice porrect or bent downward especially in
veined, entire or rarely lobed or dissected, with zygomorphic flowers. Fruits commonly a capsule
conspicuous persistent or sometimes deciduous dehiscing with (2)3(4–6) coriaceous to woody,
stipules, petiolate or sometimes subsessile, epul- rarely elastic or papery valves, sometimes fleshy,
vinate; plants glabrous or variously pubescent rarely a nut. Seeds in most genera uniform, globose
with unicellular or multicellular hairs. Inflores- to narrowly ellipsoid, compressed to strongly flat-
cences thyrso-paniculate, botryoids (pseudo- tened and often with an intermittent to entire wing
racemes, compound or simple cymes), fascicles in a few genera, rarely of two types (spindle-shaped
or condensed “short shoots”, or flowers solitary; and discoid), in some genera bearing elaiosomes;
inflorescences subtended by a commonly persis- endosperm nuclear, oily; embryo flat, straight.
tent bract; pedicels commonly articulated, almost A predominately tropical family consisting
always bearing a pair of prophylls; flower buds in of up to 1,100 species in 22 currently recognized
Viola and some other strongly zygomorphic genera, plus ten additional generic segregates
genera becoming resupinate at anthesis, with sac- to be extracted eventually from polyphyletic
cate or spurred petal lowermost. Flowers bisexual Hybanthus and Rinorea (Wahlert et al., accepted).
(unisexual), actinomorphic to strongly zygomor- The largest genus, Viola, is cosmopolitan in tem-
phic, hypogynous (slightly perigynous); sepals 5, perate and mountainous regions; the second and
free, typically quincuncially arranged, persistent third largest genera as currently circumscribed,
through fruiting, equal in actinomorphic flowers, Rinorea and Hybanthus, are mainly pantropical.
slightly unequal or rarely strongly unequal in
zygomorphic ones; petals 5, free, rarely persistent CHARACTERS OCCURRING IN RELATIVELY FEW
into fruit, equal in actinomorphic flowers, the G E N E R A A N D S P E C I E S . Roots tuberous in Viola
anterior slightly to much longer and saccate or arborescens, xylopodiferous in a few Hybanthus.
spurred in zygomorphic flowers, aestivation com- Climbing or scrambling habit in Agatea, Anchie-
monly apotact sensu Hekking (1988b), sometimes tea, Calyptrion and Hybanthopsis; entirely herba-
quincuncial or rarely convolute; stamens 5(3), ceous habit in a few Hybanthus and most Viola,
free or filaments weakly to strongly connate into acaulescent habit in some Viola, stolons produced
a tube; anthers dithecal, introrse, rarely extrorse, in some acaulescent Viola; annual duration in a
commonly bearing a dorsal sterile connective few Viola. Thorns in a few Hybanthus and one
appendage, occasionally with two ventral connec- Melicytus. Domatia in some Rinorea. Leaves oppo-
tive appendages; in zygomorphic flowers 2 fila- site in some Rinorea, a few Hybanthus, pseudo-
ments and/or dorsal surfaces of the anthers whorled in Allexis; pellucid-punctate in Leonia.
bearing individual or fused nectariferous glands Leaf blades lobed or dissected in a few Viola;
enclosed by a sac or spur at the base of a differ- stipules lacerate or lobed in some Viola. Flowers
entiated petal; ovary superior, (2)3(4–5)-carpel- cauliflorous in Allexis, one Leonia, rarely in Meli-
late, unilocular with parietal placentas; ovules cytus and Paypayrola, basally cauliflorous in
304 H.E. Ballard et al.
Hekkingia; flowers unisexual in some Anchietea V E G E T A T I V E A N A T O M Y . Extensive detail of

and Melicytus. Inflorescence a spiciform botryoid wood and leaf anatomy are provided by Metcalfe
in Paypayrola, solitary flower in a condensed and Chalk (1972). Leaf laminas are typically dor-
“short shoot” in Isodendrion; flowers consistently siventral (centric in a few Hybanthus) and lack
solitary in some Hybanthus and nearly all Viola. secretory cavities. Most species have sparse to
Some flowers in a spiciform botryoid aborting dense or irregularly distributed vegetative indu-
early and apparently secreting nectar in Paypayr- mentum of unicellular, uniseriate or rarely stel-
ola. Racemoid with large and emarginate bracts late trichomes (these occasionally gland-tipped),
in Hekkingia. Pedicels not articulated in Viola. and also commonly have marginal serrations or
Sepals strongly unequal in size and shape in crenations tipped by a small hard gland. The
Hybanthus heterosepalus, H. lehmannii, and cuticle is usually thin and sometimes striated,
Schweiggeria, auricled at base in Viola. Petal and may lie beneath a thin wax layer. The inner
aestivation convolute in Fusispermum, quincun- walls of the epidermal cells are often mucilagi-
cial in Hekkingia, Leonia, some Gloeospermum, nous, with mucilage cells showing up as transpar-
and rarely in Rinorea. Corolla persistent into ent dots in anatomical cross sections; various
mature fruit in Noisettia. Lateral petals “bearded” species of several genera also produce reddish to
with tufts of hair on inner surfaces near throat in yellow resin cells. The mesophyll varies with 1 to
some Viola, bottom petal pubescent within in a few palisade layers, with layer number being
some Hybanthus and some Viola. Stamens three species-specific and apparently correlated with
in one Leonia species. Filaments fully connate degree of light intensity in Hawaiian Viola species
into a tall tube and anthers slanting or horizontal (C. Havran and H.E. Ballard, unpubl.). Secretory
in Hekkingia, Leonia and Paypayrola. Ring of cells containing a brown unidentified substance
stamens with internal basal finger-like projec- (tannins?) have been noted in the mesophyll but
tions in a few Rinorea. Dorsal connective scales not in the epidermis of some Hawaiian Viola
lacking in Fusispermum and Leonia, minute or species. Vascular bundles in herbaceous species
lacking in Hekkingia, some Isodendrion and Pay- (largely confined to Viola) lack sclerenchyma
payrola. Ventral connective scales large in Allexis while sclerenchyma strands are well developed
and a few Rinorea. Ovary 5-carpellate in some around bundles in woody genera. An adaxial
Leonia, 4-carpellate in some Anchietea, 2-, 4- or hypodermis is rarely present (some Melicytus).
5-carpellate in some Melicytus. Ovary disintegrat- Leaves of all genera contain calcium oxalate crys-
ing early, with seeds maturing naked on placental tals: irregular discoid crystals dispersed through-
traces in Decorsella; fruit a nut in most Leonia and out the cytoplasm (almost entirely restricted to
fleshy in one Leonia and a few Rinorea, all Gloeos- phylogenetically basal genera), uniform prismatic
permum and Melicytus; semidehiscent and crystals restricted to veins (restricted to phylo-
woody in Calyptrion, and dehiscing by six sutures genetically derived genera), and rarely, spinose
in one Agatea species and by one suture in druses (thus far found only in Hybanthus conco-
Hybanthopsis; capsule tuberculate in Amphir- lor, H.E. Ballard, unpubl.). Minor leaf veins lack
rhox, Hekkingia, Orthion, and a few Rinorea; cap- phloem transfer cells, at least in Viola. Stomata
sule thin-walled and elastic in Hybanthopsis and a are primarily or entirely confined to the abaxial
papery “bladder” in Anchietea. Seeds strongly surface, being anisocytic or paracytic.
flattened in Agatea, Anchietea and Calyptrion, Taylor (1972) found considerable variability
winged in Agatea and most Anchietea and with in wood structure across the genera that he inter-
basal expansions in Hybanthopsis; seeds ellipsoi- preted as largely supporting the traditional clas-
dal and with longitudinal ridges and/or foveolae sification, except for endemic island groups. The
in Old World Hybanthus enneaspermus group; nodes of caulescent taxa are trilacunar. Primary
seeds fusiform or discoid in Fusispermum; seeds vascular tissue forms a cylinder, with secondary
pubescent in Hybanthopsis, some Rinorea and a tissue derived from a cylinder of cambium. Libri-
few Viola; seeds bearing elaiosomes in Allexis, form fibers are present or absent associated with
Fusispermum, Hekkingia and Viola, very reduced vessels, depending on the genus. Wood-rays are
in Hybanthus. heterocellular or homocellular, mixed uniseriate
Violaceae 305
and pluriseriate. Crystals are common in ray I N F L O R E S C E N C E S T R U C T U R E . The fundamental

cells. Imperforate tracheary elements are present. inflorescence form, found in the most basal
Wood parenchyma is sparse and paratracheal or genera Fusispermum, Leonia and Rinorea, is the
lacking. Vessel members have angular pores, thin thyrsoid, also loosely referred to as a panicle.
walls and uniform diameter. According to Hekking (1988b), from this type
The more basal genera, specifically Fusisper- depauperation (reduced ramification) leads to
mum in subfamily Fusispermoideae, as well as the thyrsoid, the botryoid and the spiciform
subfamily Leonioideae and subfamily Violoideae botryoid, but also to the compound cyme, found
tribe Rinoreeae, have very elongate vessels with in many genera, and the simple cyme, the latter
tapering end walls and scalariform perforation found principally in some Hybanthus. A fascicle,
plates. The endemic island genera Isodendrion found in several genera, may be derived from a
and Melicytus, traditionally placed in tribe Rinor- compound cyme by condensation of the second-
eeae, and all genera in previously circumscribed ary and primary axes, or from a simple cyme or
tribe Violeae have short vessels with perpendicu- racemoid through drastic shortening of the pri-
lar to oblique end walls and simple perforation mary axis. A short shoot with solitary flowers is
plates. Taylor (1972) argued that the insular found in Isodendrion. Solitary axillary flowers
Rinoreeae experienced accelerated rates of xylem are almost ubiquitous in Viola, frequent in
evolution and were therefore anomalous in their Hybanthus and occasionally occur in otherwise
“non-primitive” vessel morphologies. Recent typically dichasial or fasciculate Gloeospermum.
phylogenetic reexamination of the family using The simple “cymes” in some Hawaiian species of
multiple plastid and nuclear DNA sequences Viola are derived from initially branched stems
(Hodges et al. 1995; Tokuoka 2008; Wahlert through loss of leaves and telescoping of inter-
et al., accepted) has dramatically revised our nodes (Skottsberg 1940).
view of relationships among genera and provided
the foundation for a new intrafamilial classification F L O R A L S T R U C T U R E A N D A N A T O M Y . Pedicels
(see AFFINITIES below). Nevertheless, all more-basal are articulated except in Viola, and always bear
genera in the family within this new phylogenetic a pair of prophylls. Flowers are hypogynous or
framework possess elongate “primitive“ vessels, slightly perigynous and perfect except in some
whereas more derived genera possess shorter Melicytus and rare Rinorea. Petals are spreading
“advanced” vessels as described by Taylor. The or somewhat recurved distally, rarely strongly
anomalous endemic island groups are phylogen- reflexed in Allexis, persistently tightly closed in
etically placed as highly derived among other most Gloeospermum. Flowers have 5 equal or
derived taxa, thus demonstrating the significant slightly unequal sepals, dramatically unequal in
correspondence of vessel element morphology Schweiggeria, with auricles in Viola. Corollas in
with phylogenetic position. approximately half of the genera, namely, in cur-
Taylor (1972) also found similarities between rently recognized tribe Rinoreeae, have been cus-
the three lianescent genera then known, which he tomarily interpreted as actinomorphic, but
interpreted as the product of convergence. He numerous floral dissections across taxa of nearly
believed an affinity between Indo-Pacific Agatea all genera in the family by the authors (unpub-
on the one hand and Latin American Anchietea lished data) have revealed no instances of true
and Calyptrion on the other to be highly unlikely. actinomorphy in which all petals are identical in
Chloroplast DNA data have confidently placed size and shape (Allexis, excluding A. zygomorpha,
these lianescent genera and the more recently approaches this). Conversely, nearly all members
described herbaceous twining genus Hybanthop- of the family express at least weak zygomorphy in
sis into an exceptionally well-supported “lianes- the corolla, or the whole perianth, and often also
cent” clade (Paula-Souza and Souza 2003; in the androecium and gynoecium. While Arnal’s
Wahlert et al., accepted), revealing the unusual (1945) characterization of the “transitional”
scandent to vining habit as a phylogenetically genera Amphirrhox, Decorsella, Paypayrola, and
meaningful trait and supporting the surprising Isodendrion as groups with one petal slightly
southern amphi-Pacific disjunction. to noticeably broadened in the apical portion
relative to the others but essentially the same Current investigations into floral development,
length has remained accurate, about half of the details of sepal and petal aestivation, and androe-
reportedly actinomorphic genera in the family cial morphology (especially staminal gland fea-
nevertheless show noticeably zygomorphic features) are providing many additional overlooked
tures in calyx, corolla, androecium, gynoecium characteristics with which to distinguish segregate
or combinations of these, and the remainder of lineages of polyphyletic Hybanthus and Rinorea as
the family is moderately to strongly zygomorphic separate genera, and to create a more natural clas-
in floral morphology. sification expressing recognizable taxonomic
Zygomorphic flowers within tribe Violeae diversity in the Violaceae (Wahlert et al., accepted).
have the anterior petal moderately to strongly Violets (mainly Viola) provide examples of
differentiated in shape, often into a slender claw the evolutionarily successful mixed chasmoga-
and abruptly widened blade, commonly also lon- mous/cleistogamous breeding system (Ballard
ger than the others, saccate or spurred at the base. et al. 2011). Cleistogamous flowers have been
The sac or spur encloses two free or fused slightly documented in a few Hybanthus and most Viola
or greatly prolonged staminal glands functioning species. The few exceptions in Viola lacking cleis-
as nectaries (Fig. 73G). In Calyptrion, Noisettia, togamous flowers are inhabitants of xeric, desert
Schweiggeria and some Viola the spur is long. In or grassland environments in a number of dispa-
Allexis zygomorpha, some Hybanthus, some Viola rate groups, suggesting multiple independent
and some Rinorea, the bottom petal blade is losses of cleistogamy in these particular cases.
pubescent on the inner surface near the throat, In cleistogamous flowers, stamen and pistil devel-
or over both surfaces. The lateral petals are pubes- opment is accelerated relative to the sepals and
cent on the inner surface near the throat in some petals in the very young bud (Mayers and Lord
Viola. 1983). The reproductive organs remain somewhat
The androecium in perfect flowers is com- rudimentary in form and sometimes fewer
posed of 5 fertile antesepalous stamens (3 in Leo- stamens are formed; comparatively fewer pollen
nia triandra). Anthers and dorsal connective grains are shed than in the typical chasmogamous
appendages are often connivent or tightly coher- flower. The style in some species curves around
ent around the slightly to moderately exserted within the tightly coherent pollen cone, and pol-
style, forming a “pollen cone” especially well len grains, which have germinated within the top
developed in zygomorphic flowers and in some of the anthers, penetrate the anther walls and
species of Melicytus. Filaments are weakly to mod- contact the stigmatic orifice in situ. Thus, cleis-
erately connate in Fusispermum, genera in the togamous capsules are easily distinguishable
traditional tribe Rinoreeae and some Violeae to from chasmogamous ones by the tightly curled,
very strongly so, and form a tall staminal tube in rudimentary style commonly persisting at the tip
Hekkingia, Leonia and Paypayrola. Small glands of one of the valves. In European Viola odorata
are borne on or at the base of the filaments or on the production of chasmogamous vs. cleistoga-
the filament tube in the traditional subfamily Vio- mous flowers is determined by the photoperiod;
loideae, in tribe Violeae two of them enlarged, and plants receiving 11 hours or less of daylight pro-
sometimes fused, to form nectaries. Thecae are duce chasmogamous flowers, whereas those
vertical, slanting in Hekkingia and Paypayrola or receiving 14 hours or more produce cleistoga-
horizontal in Leonia. Stamens terminate in a mous ones (Mayers and Lord 1983). Similar
membranous connective appendage, which is observations hold for New Zealand Viola cunning-
large and conspicuous in most genera but rudi- hamii (Holdsworth 1966). At least some tropical
mentary in some Isodendrion and Rinoreocarpus montane Viola species, however, produce both
and absent or rudimentary in Fusispermum, Hek- types simultaneously.
kingia, Leonia, and Paypayrola. Broadly attached
ventral connective appendages are present in E M B R Y O L O G Y . The vast majority of embryolog-
Fusispermum, and small linear or bicuspidate ven- ical information comes from studies of various
tral processes are found in some Allexis and a few Viola species, in a genus which is anomalous in
Rinorea species. many ways and quite derived in the family (for
Violaceae 307
summary see Johri et al. 1992). The anther is 2000). Similarly, in the Viola nuttallii complex
tetrasporangiate and four- to six-layered; the (Fabijan et al. 1987) and in Viola adunca
endothecium is fibrous, middle layers are ephem- (McPherson and Packer 1974), pollen size has
eral in some Viola species, and Ubisch granules been found to increase with each higher ploidy
stud the inner walls of tapetal cells in certain level.
Hybanthus. Pollen is shed at the two-celled
stage. In Viola riviniana, the generative cell has K A R Y O L O G Y . Most counts in the family are for
been documented dividing either in the pollen Viola (over 1,200), but several other genera have at
grain or in the pollen tube. least one (see Ballard 1996). Relatively few chro-
Ovules are anatropous, bitegmic, and crassi- mosome counts exist for members of tribe Rinor-
nucellate; the inner and outer integuments form- eeae, and none for Fusispermum and Leonia. The
ing a zigzag micropyle in Viola and some lowest regularly confirmed number yet counted is
Hybanthus, whereas only the inner integument n ¼ 5 for Viola mercurii and V. parvula, two small
forms the micropyle in Hybanthus concolor. annual Mediterranean pansies of section Mela-
Embryo sac development follows the Polygonum nium; the largest number is n ¼ ca. 120 for
type. Antipodals are small and ephemeral in V. bubanii, another pansy in Mediterranean Eur-
some, relatively large in other Hybanthus and ope. Counts for some Rinoreeae (e.g., Rinorea) but
Viola. Endosperm development is of the Nuclear also certain Violeae (Viola, African and Australa-
type with centripetal wall formation; the endo- sian Hybanthus, North American Hybanthus con-
sperm is massive and persistent. Embryogeny color) have diploid and polyploid numbers
corresponds to the Asterad type. The mature conforming to a base number of x ¼ 6, whereas
embryo is chlorophyllous, straight and spatulate, Hawaiian Isodendrion, Indo-Pacific Agatea, Aus-
with flat cotyledons embedded in the endosperm. tralian-Oceanic Melicytus and other New World
Hybanthus have diploid and polyploid numbers
P O L L E N M O R P H O L O G Y . As with embryological arising from a base of x ¼ 8 (Beuzenberg 1961;
information, very few other genera besides Viola Bennett 1972; Kellogg and Weitzman 1985). While
have been adequately examined and documented; the affinities suggested by chromosome numbers
information summarized here is derived from do not correspond well with the traditional intra-
Erdtman (1952). In several Viola species as familial classification, they closely match the affi-
well as a few Hybanthus, Leonia, Melicytus and nities of genera portrayed by plastid DNA
Rinorea, pollen grains are 3–4(5)-colporate and sequence data (Wahlert et al., accepted): basal
suboblate to prolate or prolate-spheroidal. The genera possess 2n ¼ 24 or 48 (potentially poly-
exine is tectate, with a psilate tectum. Pollen ploids on x ¼ 6), while more derived genera have
exine micromorphology appears to be approxi- the higher base number (x ¼ 8). However,
mately similar across genera but this demands Marcussen et al. (2012) have argued from Viola
further scrutiny. Preliminary observations of a and other genera with more abundant counts
few other genera indicate greater diversity to be that the basal number in the genus may in fact
found than previously appreciated; e.g., Hekkin- be x ¼ 7. Chromosome numbers support molecu-
gia pollen is cylindrical and monocolpate lar systematic evidence for extreme polyphyly of
(H.E. Ballard, unpubl.). Recent Malpighiales- the genus Hybanthus; unfortunately, little cytolog-
wide studies of pollen morphology and tapetum ical evidence is yet available for the large and
(Furness 2011) revealed potential palynological heterogeneous genus Rinorea. On the contrary,
synapomorphies defining the loosely phenotypi- the genus Viola, largest in the family, is demon-
cally circumscribed order and also suggestive of strably monophyletic and yet shows extensive
shared traits uniting sister families Passifloraceae polyploid speciation (Ballard et al. 1999). Using a
and Violaceae, namely, a thick bilayered or chan- molecular phylogeny of Viola groups worldwide
neled apertural intine, and pollen (in Viola) with based on the nuclear Internal Transcribed Spacer
more than 3 colporate apertures. DNA region, Ballard et al. (1999) proposed that all
Pollen heteromorphism is known from Viola, but one of several aneuploid groups have arisen
where specifically the number of pores generally from euploid progenitors through loss of one or
increases as polyploidy increases (Nadot et al. more chromosomes. Exceptionally, the three
species assigned to the Asian Viola subsect. Diffu- insects instead (Freitas and Sazima 2003; Braun
sae were inferred to have each gained a chromo- et al. 2012).
some relative to their nearest sister group. The generalized pollination process in Viola
Subsequent studies by Marcussen et al. (2012) relies in part on the tightly coherent ring of dorsal
recently applied genetic information from a sin- connective appendages of the stamens surround-
gle-copy nuclear gene, glucose phosphate isomer- ing the style to form a “pollen cone”, and in part
ase (GPI), to demonstrate that all polyploid on the elastic flexibility of the style on the pistil.
groups in North America and Hawaii are allopoly- This is also echoed in some species of Melicytus
ploid, with two to several genomes from different that have tightly coherent stamens and well-
diploid and tetraploid ancestral groups. Results developed anther connectives. The head or pro-
documented a primary allopolyploid 10x level boscis of the insect forcing its way into the flower
common to all five high-polyploid groups, and displaces the style from the center of the pollen
secondary allopolyploidization with additional cone. As the style is displaced, a drop of viscous
tetraploid ancestors in three of the five groups. fluid exudes from the stigmatic orifice, collecting
pollen grains from neighboring conspecific flow-
P O L L I N A T I O N . Most members of the family, ers that are scraped off the insect’s body. Con-
including the most primitive genera, are comitantly, the displacement of the style opens a
undoubtedly entomophilous, displaying chasmo- gap in the pollen cone, permitting pollen of the
gamous floral syndromes appropriate for attract- present flower to fall onto the insect (Beattie
ing butterflies, bumblebees, solitary flees, 1969). When the insect withdraws its head, the
hawkmoths or, in the tiniest flowers of some style snaps back into position, drawing in the
Andean Viola species, perhaps even small flies. viscous drop with collected pollen grains and
A few taxa, including the red-flowered South also closing off the gap in the pollen cone. In
American scandent shrub or liana Viola arguta, some weakly zygomorphic genera (e.g., Amphir-
with flowers bearing an inflated spur, may in fact rhox, Paypayrola), the dorsal connective appen-
be pollinated by birds. A diversity of Viola spe- dages are rudimentary and do not form a “pollen
cies with different corolla color patterns and cone”, and the floral structure (stigmas hidden in
divergent floral morphologies, including mem- the long, narrow corolla tubes and anthers
bers of the same subsection, have been shown to located on short filaments near the base of the
attract largely different classes of insect pollina- corolla) suggests pollination through the mouth-
tors, resulting in a significant degree of pollinator parts of long-tongued insects. Therefore, pollen-
partitioning, not to mention some level of etho- release mechanisms do not occur as observed in
logical or mechanical isolation between closely Viola and possibly most species of the family, in
related species (Beattie 1974). In Viola, a yellow which pollen placement on the insect body is
corolla has been inferred to be a generalist syn- trigged by the manipulation of the style (Braun
drome, whereas a blue or white corolla (depend- et al. 2012). Further study of other zygomorphic
ing also on the petal morphology and flower genera is required, to determine whether the clas-
architecture) solicited a narrower and more spe- sic “Viola” strategy or another pollination mech-
cific suite of pollinators, usually bee and butterfly, anism is employed in other zygomorphic flowers,
or moth, respectively. and what pollination mechanism(s) predominate
The basic flower structure in zygomorphic in genera with actinomorphic flowers.
groups of the Violaceae fits that of “nectar flow- Floral syndromes of certain species have
ers”, the pollination reward for most species become quite specialized for their preferred pol-
being the secretion of nectariferous glands linators. Several studies have been conducted on
located on or at the base of the anther’s filaments, floral variation and reproductive success of the
which is stored inside the anterior petal’s sac or extraordinarily long-spurred Viola cazorlensis,
spur. However, some species produce very scanty narrowly endemic to the limestone hills of the
nectar or none at all (although nectaries are pres- Cazorla region of Spain (see Herrera 1988).
ent, as in some species of Viola sect. Leptidium) Short of exceptional morphological specializa-
and pollination is effected by pollen-collecting tions, closely related species in the same group
Violaceae 309
of Viola have been found to attract modally dif- F R U I T A N D S E E D . Most genera produce a char-
ferent classes of pollinators (Beattie 1970). A sub- acteristic thick-walled, three-valved capsule;
stantial degree of ethological isolation through Hybanthopsis bears a very thin-walled elastic cap-
pollinator partitioning is accomplished among sule. The capsules of most genera have a smooth
simultaneously flowering species by virtue of the to slightly rugulose surface but those of Amphir-
evolution of a few seemingly minor floral differ- rhox, Hekkingia and most Orthion are strongly
ences (Beattie 1974). During nectar feeding in tuberculate. In Rinorea anguifera of tropical Asia,
Viola, flowers with comparatively large bottom as well as a few species of Brazilian Hybanthus,
petals, tufts of hairs (“beards”) on lateral petals the capsule produces a dense indumentum of
and absence of strong corolla color contrast or long shaggy multicellular, multiseriate hairs. Leo-
heavy nectar guides encourage insect approaches nia bears indehiscent nuts in most species and a
in the prone (head up) position for all insect berry-like nut with a thin, edible pericarp in
pollinators, leaving pollen deposited on the top L. glycycarpa. Neotropical Gloeospermum and
of the head and upper thorax. Conversely, flowers Australian-Oceanic Melicytus produce soft fleshy
with small bottom petals, glabrous lateral petals berries, edible in at least some species of the first
and presence of strongly contrasting corolla color genus. The capsule of extra-Amazonian Anchie-
patterns (including heavy nectar guides or eye- tea is paper-thin and bladder-like (Fig. 73). The
spot) foster visitation in the supine (head down) seeds of most genera are narrowly ovoid to
position for some insects, particularly bumble- obovoid or globose; those of the lianescent genera
bees and solitary bees, leaving pollen deposited Agatea, Anchietea and Calyptrion are distinctly to
on the face and lower thorax. strongly flattened and in Anchietea and Agatea
The anthesis in Violaceae is mostly diurnal are typically bordered by an intermittent to con-
(Beattie 1969, 1974; Augspurger 1980; Powlesland tinuous thin wing. The twining herb Hybanthop-
1984; Munzinger and Pauly 2003), although sis, phylogenetically allied with these lianescent
recent studies have reported a nocturnal flower- genera, has a compressed seed with two basal
ing in Anchietea (Hoffmann et al. 2010), Amphir- processes which are presumably homologous to
rhox and Paypayrola (Braun et al. 2012). Two the flattened ridge or marginal wing found in
phases of anthesis have been inferred, at least in related genera. A few genera with ovoid or ellip-
Viola, from changes in petal position and flower soid seeds, namely Allexis, Fusispermum, Hekkin-
orientation (Beattie 1969): shedding of pollen gia, a few Hybanthus and Rinorea species, and all
from anthers into the pollen cone, and deposition Viola species, possess an elaiosome which origi-
of pollen from the cone onto the bottom petal. nates as an outgrowth of the micropyle or the
The first is believed to be effective only for polli- raphe base (in Hybanthus and Viola) or from
nators properly operating this primary pollination the woody annulus surrounding the apex of the
mechanism, whereas the second is a “failsafe” funicle (Rinorea) (Corner 1976). The testa is
secondary pollen presentation mechanism in made of tabular or radially elongate cells with a
which a wider range of insects may accomplish thin to strongly thickened outer wall, sometimes
pollination. with stomata, and scattered sclerotic cells. The
In Hybanthus enneaspermus, flowers not suc- mesophyll is thin- or thick-walled, sometimes
cessfully cross-pollinated roll up their enlarged somewhat lignified and with crystal-cells. The
bottom petal longitudinally and orient the rolled exotegmen consists of longitudinally oriented
petal downward, permitting pollen to fall onto the flat lignified fibers laid down in 1 to 4 layers
style within. Pollen tubes will often germinate, (few and short in Hybanthus and Viola, more
effecting self-fertilization (J. Munzinger, unpubl.). and elongate to sinuous in Rinorea). Seeds of all
Some pansy species of Viola sect. Melanium are genera have well-developed nuclear oily endo-
purportedly primarily or exclusively selfers, per- sperm. The two cotyledons are flat. The embryo
haps aided by the same mechanism. The tightly is chlorophyllous and straight (in several species
closed corollas of most Gloeospermum species of Viola). Seed shape, size and color provide
suggest that selfing might be common in that taxonomically significant traits in Viola sect.
genus as well. Leptidium (Ballard, unpubl.) and seed coat
For cleistogamy, see FLORAL STRUCTURE AND microsculpturing diverges recognizably among
ANATOMY above. species in certain Viola groups, e.g., subsect.
Boreali-Americanae, and provides some taxo- The genera Gloeospermum, Melicytus and the
nomic potential for inferring relationships or species Leonia glycycarpa produce berries and
hybrid origin (Gil-ad 1995, 1998). Sculpturing of are presumably dispersed by birds or other ani-
the seed coat and size, shape and color of the mals. The indehiscent “nuts” of most Leonia spe-
mature seeds have proven to be of great taxonomic cies are likely dispersed by animals as well. The
utility (together with androecial and inflorescence winged seeds of Anchietea and the soft, floating
morphology) in supporting the recognition of sev- seeds of the riverine species of Calyptrion suggest
eral phylogenetically distinct lineages currently that hydrochory and anemochory play a major
lumped under the polyphyletic assemblage role in the distribution patterns within these
Hybanthus. genera (Paula-Souza et al. 2011).
Significant seed predation by infesting micro- The germination ecology of seeds in temper-
lepidoptera and depressed seed set due to ate species of Viola is complex. Many species
reduced pollinator visitation in temporally (e.g., Viola egglestonii) display true dormancy
isolated flowering individuals of Hybanthus pru- immediately following dispersal, and will not ger-
nifolius may have been key factors in selection minate until after a period of cool temperatures
for synchronous flowering time in that species or cold stratification (Baskin and Baskin 1975).
(Augspurger 1982). Contrariwise, the winter annual V. bicolor (as
V. rafinesquii) requires high temperatures such
D I S P E R S A L . Seed dispersal in most capsule- as would be passed during summer dormancy, to
bearing representatives of Viola is of the diplo- foster germination (Baskin and Baskin 1972). The
chorous, or explosive, type (Beattie and Lyons seed ecophysiology of other genera remains
1975), and this is presumably the case with unknown.
other dehiscent capsule genera, too. This is char-
acterized by readily dehiscing capsules, usually R E P R O D U C T I V E B I O L O G Y . All except some
green and commonly on upright, well-elevated dioecious Anchietea and Melicytus species pro-
peduncles. At maturity, the fruit dries and the duce bisexual flowers; most genera are presum-
expanding seeds force open the capsule valves, ably facultatively self-compatible, and perhaps
which spread wide; the valves complete the occasional to frequent selfers through allogamy.
drying process and the side walls squeeze The vast majority of species in Viola and at least
together, forcibly ejecting the seeds up to 2 (or one Hybanthus (H. concolor) produce cleistoga-
more) meters away. Seeds are located by ants and mous flowers in addition to the normal chasmo-
secondarily removed further from the parent gamous ones and are at least intermittently
plant. autogamous. Studies of such species suggest that
Species of some acaulescent groups of Viola cleistogamy serves as an important stopgap mea-
produce small, globose, purple-blotched cleistog- sure to compensate for losses of outcrossed seeds
amous capsules on short, prostrate or burrowing during environmentally stressful periods (Redbo-
peduncles conforming to the myrmecochorous Torstensson and Berg 1995). Contrary to theoret-
mode of seed dispersal. In these species, the cap- ical expectations, fitness in Viola canadensis is
sule dehisces slowly or gradually disintegrates, not significantly reduced in plants derived from
spilling the seeds onto the ground surface or cleistogamous versus chasmogamous seeds, sug-
burying them under the soil. In the former gesting that high levels of selfing are tolerated well
event, ants are attracted to the elaiosome on the by this species (Culley 2000). Slight responses
seeds and may carry them some distance away interpreted as low-level inbreeding depression
from the parent plant. This latter dispersal syn- occur in the offspring of cleistogamous flowers
drome may represent a fairly high degree of of other Viola species but this cost is likely out-
coevolution with particular ant species (Beattie weighed by the lesser effort required to produce
and Lyons 1975). Generally, diplochorous species cleistogamous flowers (Redbo-Torstensson and
occupy open sites with presumably greater Berg 1995).
environmental change, whereas myrmecochor-
ous species are characteristically woodland inha- P H Y T O C H E M I S T R Y . Whereas hydrolysable tan-
bitants. nins are absent from Violaceae, the occurrence
Violaceae 311
of condensed tannins (proanthocyanin and catalyzed intensive scrutiny by phytochemists to

proanthodelphinidin) in the leaves of very many determine their taxonomic distribution across
species of the herbaceous genus Viola is remark- families and lineages of angiosperms as well as
able (Hegnauer 1973). A wide variety of flavones, their overall diversity. These proteins are espe-
flavonols and related flavonoids and their deriva- cially abundant in the Cucurbitaceae, Rubiaceae
tives have been isolated and identified in leaves of and Violaceae (see review by Craik 2010). A pop-
several Viola species particularly of the yellow- ulation- and species-level study of cyclopeptides
flowered subsect. Nuttallianae (Fabijan et al. in Australian Hybanthus demonstrated that the
1987). Various volatile compounds have been distribution of characterized proteins correlated
isolated in leaves of Viola odorata (Cu et al. well with morphological similarity of species and
1992), and violaxanthin esters have been shown characterized 246 new proteins in this class,
to occur in Viola tricolor flowers (Hansmann and bringing the total for the family to more than
Kleinig 1982). Leaves and flowers harbor consid- 9,000 types (Simonsen et al. 2005).
erable quantities of acidic mucilage, which is typ-
ically found in epidermal cells. Many Violaceae S U B D I V I S I O N O F T H E F A M I L Y . The Violaceae
thus far tested (e.g., Hybanthus enneaspermus, are currently subdivided into three subfamilies,
Anago et al. 2011) possess saponins and/or alka- the Fusispermoideae (with Fusispermum), the
loids, flavonoids, and often resinous secretory Leonioideae (with Leonia), and the Violoideae
cells with yellowish to brownish contents. Methyl (with the remaining 20 genera). Subfamily Vio-
salicylate has been identified in rhizomes of the loideae consists of two tribes, with an approxi-
pansies Viola arvensis and V. bicolor (Hayden mately equal number of genera in each. Tribe
and Clough 1990) as well as the branchlets of Rinoreeae bears more or less actinomorphic or
Rinorea, and pentacyclic triterpenes as well as very scarcely zygomorphic flowers, and the stam-
putatively anti-allergic aliphatic hydrocarbons inal glands lack any further elaboration, although
and methyl esters of fatty acids have been they may be fused to each other or to any existing
extracted from Anchietea salutaris (¼ A. pyrifolia) filaments. Tribe Violeae possesses distinctly
(Di Stasi et al. 1999). Unfortunately, the chemistry zygomorphic, saccate or spurred flowers, with
of the Violaceae has never raised much interest; two staminal glands elaborated to form nectaries,
the basalmost woody genera have been neglected which may be free or conjoined. Due primarily to
in this respect, and the nature of the alkaloids of the efforts of Hekking and others with the actino-
the family appears to remain unknown. morphic woody genera, tribe Rinoreeae has been
Aluminum accumulation is known to occur divided further into subtribes, principally on the
in Allexis and Amphirrhox (Chenery 1948), and basis of androecial characters. Relatively little
nickel uptake has been demonstrated in New change has taken place since de Candolle’s time
Caledonian Agatea deplanchei (Jaffré 1980), in (De Candolle 1844), aside from the addition of
Australian and South Pacific Hybanthus species new genera and further grouping of genera into
(Severne and Brooks 1972; Cole 1973; Jaffré and an intrafamilial framework. The only comprehen-
Schmid 1974; Brooks et al. 1974, 1977a; Kelly et al. sive treatment of genera in the family and their
1975; Lee et al. 1977), in Indo-Malayan Rinorea relationships has been that by Melchior (1925a,
bengalensis (Brooks and Wither 1977) and 1925b). The classification presented below fol-
R. javanica (Brooks et al. 1977b), and accumula- lows that of Hekking (1988b), who provided the
tion of zinc and other heavy metals is done by most recent research on many Neotropical woody
Viola calaminaria (Jedrzejczyk et al. 2002) and genera and gave a useful synopsis of the family in
other European pansies, as well as by the New his treatment on Rinorea and Rinoreocarpus.
World V. cuneata (Reeves et al. 1983). Newly described or recognized genera have been
Cyclopeptides were first detected and isolated inserted in their most logical positions based on
as metabolites earlier in the last century and later morphological characters.
ascribed a role in plant defense (Craik 1999). Recently initiated cladistic studies using mor-
Their recently recognized medical promise, with phological and anatomical characters (Ballard
some types showing high levels uterotonic, anti- et al., unpubl.) support basal positions for sub-
HIV, antimicrobial and insecticidal activity, has families Leonioideae and Fusispermoideae, and
place some genera of tribe Rinoreeae basal to and Passifloraceae s.l. The violet family (plus
those of tribe Violeae except for Isodendrion, Goupiaceae) is nevertheless relatively isolated
which is embedded within a clade of Neotropical and divergent from other higher Rosidae from a
Hybanthus, and Melicytus, which is associated morphological and molecular standpoint.
with zygomorphic genera. However, the relation-
ships among genera are evidently more complex D I S T R I B U T I O N A N D H A B I T A T S . The family as a
than the simplistic portrayal implied by the tradi- whole is nearly cosmopolitan, owing primarily to
tional classification. Molecular phylogenies from the widely distributed genus Viola. Generally,
plastid DNA trnL intron, trnL–trnF spacer and phylogenetically basal genera occupy low-
rbcL sequences (Tokuoka 2008; Wahlert et al., elevation wet or dry forests, whereas more derived
accepted) are congruent in revealing several inde- genera range into a greater diversity of habitats,
pendent derivations of zygomorphic-flowered often reaching higher elevations, temperate or
genera from within broader actinomorphic boreal latitudes and more extreme environmental
lineages, dramatic polyphyly in Hybanthus and conditions. Except for a few Hybanthus and Viola,
Rinorea suggesting the need to distinguish an nearly all members of the family are restricted to
additional 10 lineages as recognized genera, and subtropical and tropical regions. The three largest
Gloeospermum and Leonia as sister taxa in a non- genera, Viola, Rinorea and Hybanthus are found
basal clade. Recent comparative anatomical and in both the New and Old Worlds and represent
micromorphological studies of Fusispermum, approximately 93% of the species diversity
Rinorea and other basal groups have supported (ca. 1,020 species) in the family. Whereas species
molecular systematic inferences on relationships of Rinorea worldwide are relatively restricted to
and provided their own additional illumination lower elevation humid or seasonally dry forests,
on the distinctive taxonomic and phylogenetic representatives of Hybanthus and Viola are found
status of Hekking’s “Apiculatae” group as a in a wide range of habitats, with the latter ranging
small transitional lineage between the first-men- from snowmelt lines of permanent glaciers above
tioned genera (Hoyos Gomez 2011). Although 6,000 m down to mid-elevation cloud forests at
macromorphological, micromorphological, ana- 1,200 m or in low-elevation deserts in both the
tomical and palynological research proceeds New and Old World. Centers of species diversity
apace across the family to characterize each dis- lie in the Amazon Basin, Indonesia, western Africa
tinct genus and clade more precisely, it is prema- and Madagascar for Rinorea; in central Mexico,
ture to segregate the several additional genera northern South America and Australia for
deserving of formal recognition. Hybanthus; and in the Andes, central Mexico,
Rocky Mountains and Appalachians of North
A F F I N I T I E S . On morphological and embryolog- America, mountainous Asia and southwestern
ical grounds, among them salicoid teeth, parietal Europe for Viola. Of the remaining genera, five
placentation and a primitive actinomorphic pen- are confined to the Old World: Allexis and Decor-
tamerous floral condition, Violaceae have tradi- sella inhabit seasonally dry forests of western
tionally been allied with a number of other Africa, and Agatea and Melicytus are distributed
“Parietalean” families including Passifloraceae in monsoon forests and coastal or montane
(now including Malesherbiaceae and Turnera- habitats, respectively, in the South Pacific region.
ceae) and Salicaceae (including Flacourtiaceae in The remaining 15 genera are Neotropical, with
part). All recent molecular data (Wurdack and Amphirrhox, Calyptrion, Fusispermum, Gloeos-
Davis 2009; Korotkova et al. 2009; Xi et al. 2012) permum, Leonia, Orthion and Paypayrola found
support this broader assemblage with the addi- in both Mesoamerica and South America (all but
tion of Goupia, now segregated from Celastraceae Orthion best represented in South America);
into its own family (Goupiaceae). Results from Mayanaea restricted to Guatemala in Mesoamer-
numerous plastid sequences by Xi et al. (2012) ica; and Anchietea, Hekkingia, Hybanthopsis,
show an unexpected sister relationship between Noisettia, Rinoreocarpus and Schweiggeria con-
Goupiaceae and Violaceae, with this clade sister fined to South America. Most Neotropical genera
to one containing Salicaceae s.l. (including Scy- inhabit low-elevation rainforests or the low
phostegiaceae, Samydaceae and Lacistemataceae) montane slopes delimiting the Amazonian basin.
Violaceae 313
Exceptionally, Viola in the tropics often reaches anthers sessile on receptacle; connective of stamens
exceedingly high altitudes, and in the Andes of with a rudimentary to well-developed dorsal scale,
this broadly attached to the apex in most genera
South America reaches its greatest species diver- (extending down the sides in most Rinorea) 5
sity and its most divergent morphologies in the 3. Leaves pellucid- or dark-punctate; thecae of anthers
Western Hemisphere. horizontal 2. Leonia
– Leaves lacking punctae. Thecae of anthers slanting
E C O N O M I C I M P O R T A N C E . Certain Viola species 4
are cultivated and bred as ornamentals, but many 4. Inflorescence a basally cauliflorous long-pedicellate
more wild species have been distributed as novel- raceme; bracts subtending pedicels persistent, deeply
ties for specialty gardeners. Flowers of commonly emarginate; capsule tuberculate 11. Hekkingia
cultivated Viola odorata and a few wild species – Inflorescence a ramiflorous (cauliflorous) spicate raceme;
bracts subtending pedicels deciduous, acute; capsule
are used for condiments, jellies and syrups, and smooth 12. Paypayrola
candies (see MacNicol 1967). Young leaves of 5. Corolla actinomorphic or weakly to distinctly zygo-
various Viola species are occasionally eaten as a morphic, all petals similar in size or different but
potherb rich in vitamin C, and syrup from the one not substantially larger and dramatically different
flower has been used medicinally. Fresh leaves of in shape than the others, nor conspicuously saccate
Leonia glycycarpa have been used in bird-lime. or spurred at base; staminal glands never elaborated
into prolonged nectarines 6
C O N S E R V A T I O N . Several polytypic genera (Aga- – Corolla strongly zygomorphic, at least one petal
("bottom" petal) considerably longer or shorter than
tea, Hybanthus, Rinorea and Viola) harbor nar- the others and conspicuously saccate or spurred;
rowly endemic species, some of which are two staminal glands commonly prolonged or differ-
threatened with extinction; in some cases these entiated into nectaries enclosed by sac or spur 13
are protected in nature reserves or on municipal 6. All petals essentially or quite identical 7
or federal lands. Some Rinorea species in Mada- – One petal slightly differently shaped from the others,
gascar are now believed extinct (e.g., R. micro- gradually or abruptly expanded in the apical half,
commonly clawed 12
phylla, R. verticillata). All species in the endemic
7. Fruit a three-valved capsule 8
Hawaiian genus Isodendrion are now imperiled
and are currently listed as "threatened" or – Fruit a berry 11
8. Corolla white to violet or yellow, very rarely pink 9
"endangered" at the state and federal levels. A
few genera are also highly localized in distribu- – Corolla red to red-orange 10
9. Inflorescence various but not a short shoot with a
tion, e.g., Mayanaea has not been collected solitary flower; dorsal connective appendage of stamens
around Lake Izabal in Guatemala for decades, conspicuous, completely covering top, and often sides,
and Hybanthopsis is known from a small area of of stamen 3. Rinorea
Bahia state in easternmost Brazil. – Inflorescence a short shoot with a solitary flower;
dorsal connective appendage of stamens a small
prickle or lacking 9. Isodendrion
KEY TO THE GENERA 10. Petals porrect or weakly spreading; dorsal connective
appendage a tiny linear or deltoid prickle tipping the
1. Flowers and capsules to 2 mm long; connective of stamen; seeds not arillate 4. Rinoreocarpus
stamens lacking a dorsal scale but bearing a broad
ventral scale; seeds of two types, spindle-shaped or
– Petals strongly reflexed; dorsal connective appendage
of stamens large, completely covering top of stamens;
discoid 1. Fusispermum
seeds arillate 6. Allexis (most spp.)
– Flowers or capsules longer than 2 mm; connective of 11. Petals connivent with one petal clasping around
stamens usually bearing a well-developed dorsal scale others, rarely spreading at anthesis; dorsal connective
(rudimentary in Isodendrion, Melicytus and Rinoreo- appendage large and conspicous, commonly covering
carpus, absent in Hekkingia, Leonia and Paypayrola), top of anthers completely; flowers strictly bisexual
lacking a broad ventral scale; (Rinorea sometimes 5. Gloeospermum
with a linear or bicuspidate process); seeds uniformly
globose to obovoid, or strongly flattened 2
– Petals widely spreading to recurved at anthesis; dorsal
connective appendage a tiny slender prickle tipping
2. Filaments fused into a tall tube much longer than the the anthers; flowers bisexual or unisexual
anthers, anthers sessile; connective of stamens lacking 8. Melicytus
a dorsal scale 3 12. Bottom petal emarginate at the apex; dorsal connec-
– Filaments fused into a short continuous or inter- tive appendage a conspicuous linear prickle tipping
mittent tube distinctly shorter than the anthers, the stamens; ovary persistent in maturing fruit,
with anthers borne on a free portion of filament, or enclosing seeds 10. Amphirrhox
– Bottom petal narrowly rounded at apex; dorsal con- – Three sepals much larger and broader than the other
nective appendage ovate, completely covering top of two, ovate-triangular; spur stout, 4–8 mm thick;
anthers; ovary disintegrating early, seeds maturing corolla white, caducous 20. Schweiggeria
naked on spreading placental traces 7. Decorsella
13. Bottom petal saccate or with very short cylindrical
spur, the blade at least one and one-third times longer GENERA OF VIOLACEAE
than lateral and upper petals, commonly much longer,
often distinctly differentiated into a claw and abruptly I. S U B F A M . F U S I S P E R M O I D E A E Hekking (1984).
expanded blade 14
– Bottom petal distinctly spurred or greatly distended
and gibbous at base, the blade slightly longer or Corolla aestivation in bud convolute; stamens 3,
shorter than lateral and upper petals, not strongly filaments weakly connate, dorsal connective
differentiated into a claw and blade 17 appendage lacking, ventral appendage present;
14. Plant a liana or scandent shrub; seeds moderately to fruit a three-valved capsule; seeds discoid or fusi-
strongly flattened, bordered by one or two roughened
ridges or a wing 15
form.
– Plant a tree, treelet, erect shrub or herb; seeds obovoid
to globose, lacking ridges or wings 16 1. Fusispermum Cuatrec.
15. Liana or scandent shrub; capsule thick-walled and Fusispermum Cuatrec., Fieldiana Bot. 27: 94 (1950);
firm, dehiscing by 3 or 6 sutures; seeds strongly Hekking, Proc. Kon. Ned. Akad. Wetensch. C 87:
flattened, encircled by an interrupted or broad mem- 121–130 (1984).
branous wing 16. Agatea
– Twining herb; capsule thin-walled and elastic, dehisc- Trees; leaves oblanceolate. Flowers in axillary
ing by 1 suture; seeds obovoid, with a pair of lateral paniculate cymes or pseudoracemes; corolla
expansions only at base 15. Hybanthopsis white, actinomorphic; stamens 5, filaments
16. Plant a full-sized tree; inflorescence a long-pedunculate
compound cyme; capsule usually tuberculate
weakly connate, dorsal connective appendage
13. Orthion absent; pistil 3(4)-carpellate, style filiform. Fruit
– Plant a herb or shrub (a treelet); inflorescence a a thick-walled 3-valved capsule. Seeds 30–50 per
raceme, simple cyme, fascicle or solitary flower, rarely carpel, fertile seeds fusiform, sterile ones discoid.
a short-pedunculate compound cyme; capsule Three spp., southern Central America, north-
smooth 14. Hybanthus ern South America.
17. Plant a herb, rarely a subshrub or shrub; pedicels
not articulated; sepals auricled at base 22. Viola
– Plant a tree, liana or shrub; pedicels articulated; sepals II. S U B F A M . L E O N I O I D E A E Melch. (1925).
not auricled at base 18
18. Plant a liana or scandent shrub; seeds strongly Corolla aestivation in bud imbricate or quincun-
flattened (with an intermittent or broad, entire or cial per Hekking (1988b); stamens 3 or 5, fused
erose wing in Anchietea) 19
into a tall tube, dorsal connective appendage
– Plant an erect shrub or tree; seeds obovoid to globose,
lacking a wing 20 lacking; fruit a nut or berry; seeds oblong-ovoid.
19. Spur less than one-half as long as petal blades;
capsule very thin-walled, bladder-like; seed margins 2. Leonia Ruiz & Pav. Fig. 70
bearing an intermittent or rudimentary to broad wing
18. Anchietea Leonia Ruiz & Pav., Fl. peruv. 2: 69 (1799).
– Spur at least twice as long as the petal blades; capsule Trees; leaves oblanceolate. Flowers in short-
thick-walled, firm and semi-dehiscent; seed margins
lacking a wing 19. Calyptrion
pedunculate, cauliflorous or ramiflorous com-
20. Plant a tree or treelet; spur under 3 mm long 21 pound cymes or racemoids, actinomorphic;
– Plant a shrub; spur 5–20 mm long 22 corolla yellow to orange; thecae oriented horizon-
21. Flowers ramiflorous, in a long-pedunculate compound tally and dehiscing laterally ("apically"); pistil
cyme, glabrous within; bottom petal with short, cylin- 3–5-carpellate, style filiform. Fruit a nut with
drical spur; capsule unspotted 17. Mayanaea thick woody pericarp, one sp. (L. glycycarpa)
– Flowers cauliflorous, in a fascicle, heavily pubescent with thin juicy pericarp (a "berry"). Seeds numer-
within; bottom petal with hugely distended gibbous
spur; capsule red-spotted 6. Allexis (A. zygomorpha)
ous per carpel, oblong-ovoid. Chromosome num-
22. Sepals uniformly linear-lanceolate; spur slender, less ber unknown.
than 2 mm thick; corolla orange with yellow throat, Five to six spp., southern Central America,
persistent into fruit 21. Noisettia northern South America.
Violaceae 315
Melicytus; seeds narrowly ovoid to globose, or

flattened in a few lianescent genera.
1. Tribe Rinoreeae Reiche & Taub.

Corollas actinomorphic or weakly zygomorphic,
lacking a spur and calcarate stamens; filaments
weakly to strongly connate or stamens fused into
a tube (Hekkingia, Paypayrola), dorsal connec-
tive appendage entire or erose to lacerate, thecae
vertically oriented (diagonally on tube in Hekkin-
gia and Paypayrola), dehiscing introrsely (later-
ally in Hekkingia and Paypayrola).
1a. Rinoreeae subtribe Rinoreinae Melch. (1925).
3. Rinorea Aubl. Fig. 71

Rinorea Aubl., Hist. pl. Guiane 1: 235 (1775), nom. cons.;
Hekking, Fl. Neotrop. 46 (1988b).
Alsodeia Thouars (1806).
Trees, treelets or shrubs; leaves alternate or oppo-
site, linear to orbicular leaves. Flowers in axillary
compound cymes, racemoids, short-pecunculate
compound cymes, fascicles or solitary; corolla
white to yellow, scarcely to notably zygomorphic
with one petal scarcely to somewhat longer,
sometimes this and nearest petals also somewhat
Fig. 70. Violaceae. Leonia glycocarpa var. glycocarpa. A different in shape from the remaining pair; fila-
Branchlet with inflorescence. B Part of inflorescence. C ments weakly to strongly connate, dorsal connec-
Detail thereof. D Flowers, at left two petals removed. tive appendage large, linear-oblong to ovate,
E Lateral, adaxial and abaxial views of stamens. F Pistil
in lateral view and vertically sectioned. G Fruit, entire and entire or erose to lacerate; style filiform to rostel-
sectioned. (Reproduced with kind permission of the artist late. Capsule smooth to tuberculate or scaly
Bobbi Angell) thick-walled. Seeds 1–4 per carpel, globose to
obovoid. 2n ¼ 24, 48.
230–280 spp., pantropical.
III. S U B F A M . V I O L O I D E A E Burnett (1835).
4. Rinoreocarpus Ducke
Leaves alternate or sometimes in pseudo-whorls,
or sometimes opposite in Rinorea; bud Rinoreocarpus Ducke, Arch. Jard. Bot. Rio de Janeiro 4:
144 (1925); Hekking, Fl. Neotrop. (1988b).
aestivation mostly apotact sensu Hekking
(1988b), sometimes imbricate in Gloeospermum; Trees or treelets with oblong-lanceolate leaves.
dorsal connective appendage commonly large, Flowers in axillary, short-pedunculate compound
linear to deltoid-ovate, entire to lacerate, but cymes; corolla caducous, reddish-orange, actino-
obsolete in Paypayrola, inconspicuous or obso- morphic; filaments free, dorsal connective
lete in Isodendrion; fruit a thick-walled 3-valved appendage small, apiculate; style truncate. Fruit
dry capsule in most genera, very rarely a fleshy a thick-walled three-valved capsule. Seeds 2–4 per
3-valved capsule in Rinorea, a paper-thin "blad- carpel, obovoid.
der" in Anchietea, a thin-walled follicle in One sp., R. ulei (Melch.) Ducke, northern
Hybanthopsis, or berry in Gloeospermum and South America.
Fig. 71. Violaceae. Rinorea physiphora. A Flowering F Capsule. G Same, dehisced. Viola cotyledon. H Habit.
branch. B Flower. C Same, vertically sectioned. D Abaxial I Leaf. J Stigma with orifice. (Takhtajan 1981)
view of petal with nectary. E Same adaxially, with stamen.
5. Gloeospermum Triana & Planch. Fig. 72 style filiform. Capsule thick-walled. Seeds 1–2
Gloeospermum Triana & Planch., Ann. Sci. Nat. IV, 17: 128 per carpel, obovoid, arillate.
(1862). Ca. four spp., W Africa.
Trees or treelets; leaves (oblong-)lanceolate. 7. Decorsella A. Chev.
Flowers in axillary short-pedunculate compound
cymes, fascicles or solitary; corolla caducous or Decorsella A. Chev., Bull. Soc. Bot. Fr. 61, Mém. 8: 297
(1917).
briefly persistent, white, essentially actinomor-
Gymnorinorea Keay (1953).
phic; filaments weakly to strongly connate, dorsal
connective appendage large, deltoid to oblong, Treelets or shrubs; leaves oblanceolate. Flowers in
erose to lacerate; style filiform. Fruit a dryish axillary, short-pedunculate compound cymes;
fleshy berry. Seeds 7–22 per carpel, globose. corolla yellow to reddish, slightly zygomorphic,
About 12 spp., Central America, northern one petal slightly longer and distinctly widened
South America. distally; filaments strongly connate, dorsal con-
nective appendage large, ovate, apical and lateral,
6. Allexis Pierre entire; style filiform; ovary wall disintegrating
Allexis Pierre, Bull. Soc. Linn. Paris II, 1: 25 (1898). early, leaving seeds to mature naked on spreading
placental traces. Seeds 1–2 per placenta, drupe-
Treelets or shrubs; leaves pseudo-whorled, oblan- like with thin fleshy testa, globose. 2n ¼ 20.
ceolate. Flowers in compound, cauliflorous One sp., D. paradoxa A. Chev., W Africa.
cymes; floral forms of two dramatically different
types: a) corolla nearly actinomorphic with petals
narrowly lanceolate and reflexed, stamens erect 1b. Rinoreeae subtribe Hymenantherinae Melch.
and exserted, dorsal connective appendages large, (1925).
linear-attenuate, entire; or b) corolla hypocertoid,
stamens short and included, dorsal connective 8. Melicytus J.R. Forst.
appendages short, ovate, erose (A. zygomorpha); Melicytus J.R. Forst., Char. Gen.: 123, t. 62 (1776).
corolla in all cases pink to red; filaments free; Hymenanthera R. Brown (1818).
Violaceae 317
9. Isodendrion A. Gray
Isodendrion A. Gray, Proc. Am. Acad. 2: 324 (1852).
Treelets to shrubs; leaves lanceolate to ovate.
Flowers solitary in axillary short shoots; corolla
caducous, blue, weakly zygomorphic, petals pro-
longed into a "tube" with bottom petal slightly
longer than others and weakly differentiated,
scarcely saccate at base; filaments free, anthers
dehiscing laterally, dorsal connective appendage
a minute prickle or obsolete; style filiform or
rostellate. Capsule thick-walled. Seeds 2(–4) per
carpel, obovoid. 2n ¼ 16.
Four to ten spp., Hawaiian Islands.
1d. Rinoreeae subtribe Paypayrolinae Melch.

(1925).
10. Amphirrhox Spreng.

Amphirrhox Spreng., Syst. veg.: 51 (1827).
Spathularia A. St.-Hil. (1824), non Spathularia Pers. (1797).
Trees, treelets or shrubs; leaves oblanceolate,

Fig. 72. Violaceae. Gloeospermum sphaerocarpum. A alternate at branch tips, opposite below. Flowers
Branchlet with flower buds. B Part of inflorescence with in subterminal or terminal long-pedunculate sim-
flower bud. C Flower, two petals removed. D Adaxial view
of upper part of androecium opened out; note laciniate ple cymes, rarely in compound cymes; corolla
connective extensions on apices of anthers. E Androecium white, actinomorphic, one petal twice as broad
surrounding the gynoecium. F Pistil with medial trans- as others, differentiated into claw and blade and
verse section of the ovary. G Part of branch with dried emarginate; filaments weakly connate, dorsal
fruit. H Fruit in transverse section showing parietal attach-
ment of seeds. (Reproduced with kind permission of the connective appendage large, linear, entire; style
artist Bobbi Angell) filiform. Capsule thick-walled, tuberculate. Seeds
3–6 per carpel, obovoid.
One sp., A. longifolia (A. St.-Hil.) Spreng.,
Treelets or shrubs; leaves alternate or pseudo- Mesoamerica, northern South America.
whorled, linear to ovate; branches occasionally
thorny. Flowers unisexual or bisexual, in rami- 11. Hekkingia H.E. Ballard & Munzinger
florous, rarely caulifolorous racemoids, fascicles Hekkingia H.E. Ballard & Munzinger, Syst. Bot. 28: 345
or solitary; corolla white or yellow to blue, acti- (2003).
nomorphic; filaments strongly connate, dorsal
Trees or shrubs; leaves lanceolate to oblanceolate.
connective appendage large, deltoid to oblong-
Flowers in long-pedicellate racemoids, basally cau-
ovate and lacerate in bisexual flowers (lacking in
liflorous; corolla yellow, actinomorphic, one petal
unisexual ones); style filiform to trilobate. Fruit a
scarcely broader than others, all petals emarginate,
juicy berry. Seeds 2–3 per carpel, obovoid. 2n ¼
spreading to strongly recurving; filaments connate
32, 64, 96.
into a tall tube with anther tissue distinct, dorsal
Ca. 10 spp., Australia, New Zealand, South
connective minute and apiculate, thecae oriented
Pacific islands.
diagonally on tube and dehiscing laterally; style
1c. Rinoreeae subtribe Isodendriinae Melch. filiform. Capsule strongly tuberculate, thick-
(1925). walled. Seeds 5–8 per carpel, obovoid, arillate.
One sp., H. bordenavei H.E. Ballard & Mun- nective appendage large, oblong-ovate, entire;
zinger, northern Brazil, French Guiana. style rostellate. Capsule thick-walled, typically
tuberculate. Seeds 3 per carpel, broadly obovoid.
12. Paypayrola Aubl. Six spp., Mesoamerica and northern South
Paypayrola Aubl., Hist. pl. Guiane 1: 249 (1775). America.
Trees or shrubs; leaves lanceolate to oblanceolate. 14. Hybanthus Jacq.

Flowers in axillary or terminal spiciform race- Hybanthus Jacq., Enum. Pl. Carib.: 2 (1760).
moids; corolla white to yellow, actinomorphic,
one petal twice as broad as others, differentiated Treelets, shrubs or herbs; leaves alternate to oppo-
into claw and blade and emarginate; filaments site, linear to orbicular; branches occasionally
connate into a tall tube continuous with anther thorny or very rarely twining. Flowers in axillary
tissue, dorsal connective minute or obsolete, the- racemoids, simple cymes or solitary, rarely in
cae slanting and dehiscing laterally; style filiform. short-pedunculate compound cymes or fascicles;
Capsule smooth, thick-walled. Seeds 5–8 per car- corolla greenish-white to white, yellow or blue, the
pel, globose. throat commonly yellow to orange within, weakly
Eight spp., southern Central America, north- zygomorphic with bottom petal slightly longer
ern and eastern South America. and scarcely differentiated, or strongly zygomor-
phic with bottom petal much longer and differen-
2. Tribe Violeae Reiche & Taub. tiated into claw and blade, bottom petal saccate at
Corollas weakly to strongly zygomorphic, the base; filaments free or rarely strongly connate,
anterior ("bottom") petal distinctly longer than dorsal connective appendage large, oblong-ovate,
others, commonly shorter in Viola, weakly to entire; style filiform. Capsule thick-walled. Seeds
strongly differentiated from the others, saccate 1–several per carpel, narrowly ovoid to globose,
or inflated into a spur at the base; filaments free rarely with a small (non-functional?) aril. 2n ¼ 8,
or scarcely connate, either all stamens distinct 12, 16, 24, 32, 48.
and with separate glands at the summit of their Ca. 120 spp., most pantropical, desert SW
filaments, or the bottom two stamens only with North America (three spp.), temperate E North
a separate gland each on the filament or the America (one sp.).
bottom of the connective (or both), or the
glands fused to form one ridge or shield-like 15. Hybanthopsis Paula-Souza
structure across the base of the two stamens; Hybanthopsis Paula-Souza, Brittonia 55: 210 (2003).
thecae vertical, dehiscing introrsely; dorsal con-
nective appendage large, entire; seeds narrowly Twining herbs; leaves ovate-lanceolate. Flowers
ovoid to obovoid or globose, strongly flattened solitary; corolla violet, strongly zygomorphic,
in Agatea, Anchietea, Calyptrion and bottom petal very large and differentiated into
Hybanthopsis, then with a membranous wing claw and blade, saccate at base; filaments weakly
or ridge in many species. connate, two lowest anthers weakly calcarate,
dorsal connective appendage large, oblong-
ovate, entire; style filiform-rostellate. Fruit thin-
2a. Violeae subtribe Hybanthinae Melch. (1925).
walled, elastic, "follicle" dehiscing by only one
longitudinal suture. Seeds several per carpel,
13. Orthion Standl. & Steyerm.
obovoid with a pair of basal-lateral expansions.
Orthion Standl. & Steyerm., Publ. Field Mus. Nat. Hist., One sp., H. bahiensis Paula-Souza, eastern
Bot. Ser. 22: 249 (1940)
Brazil.
Treelets to shrubs; leaves lanceolate to oblan-
ceolate. Flowers in terminal, long-pedunculate 16. Agatea A. Gray
compound cymes; corolla white, weakly zygo-
Agatea A. Gray, Proc. Am. Acad. 2: 323 (1852).
morphic, bottom petal slightly longer than others
and somewhat differentiated, bottom petal sac- Lianas or reclining shrubs; leaves lanceolate to
cate at base; filaments weakly to strongly connate ovate. Flowers in terminal pseudo-racemes or
(occasionally entire stamens connate), dorsal con- racemoids; corolla white, strongly zygomorphic,
Violaceae 319
bottom petal very large and differentiated into

claw and blade, saccate at base; filaments weakly
connate, two lowest anthers weakly calcarate,
ovate, entire; style filiform to clavate. Capsule
thin- to thick-walled, 3(6)-valved. Seeds 2–several
per carpel, strongly flattened with broad thin
wing. 2n ¼ 16.
Ca. eight spp., New Guinea, S Pacific islands.
17. Mayanaea Lundell

Mayanaea Lundell, Wrightia 5: 58 (1974).
Trees; leaves lanceolate. Flowers in terminal,
long-pedunculate compound cymes; corolla blue
to white, strongly zygomorphic, bottom petal
much longer than others and differentiated into
a claw and blade, spur well exserted; filaments
weakly connate, two lowest stamens calcarate,
ovate, entire; style filiform. Capsule thick-walled.
Seeds 1–2 per carpel, broadly obovoid.
One sp., M. caudata (Lundell) Lundell, Gua-
temala.
2b. Violeae subtribe Violinae Melch. (1925).
18. Anchietea A. St.-Hil. Fig. 73

Anchietea A. St.-Hil., Ann. Sci. Nat. I, 2: 252 (1824).
Lianas or reclining shrubs; leaves oblong-lanceo-

late to ovate. Flowers unisexual or bisexual, in
axillary racemoids or fascicles; corolla white to
orange, strongly zygomorphic, bottom petal lon-
ger than others and weakly differentiated, spur
well exserted; filaments strongly connate, two
lowest stamens calcarate, dorsal connective
appendage small, ovate, entire; style rostellate.
Capsule very thin-walled, bladder-like. Seeds
many per carpel, orbicular in outline, strongly
flattened and encircled by a low interrupted
ridge or uniformly broad wing.
Five spp., South America.
Fig. 73. Violaceae. Anchietea pyrifolia. A Flowering stamens calcarate. H Adaxial view of stamen. I Capsule,
branch. B Fruiting branchlet with torn capsule valves. with seed maturation continuing when valves are torn.
C Floral bud. D Flower. E The petals: 2 small posterior J Seed. A. pyrifolia var. pyrifolia. K Capsule, the bladder-
ones, 2 clawed lateral ones, and the anterior spurred one. like valves withered but seed maturation continues.
F Gynoecium with calyx. G Androecium, the two anterior (Takhtajan 1981)
19. Calyptrion Ging. to reniform. Flowers axillary and solitary, rarely in

Calyptrion Ging., Mém. Viol.: 5 (1822). cymes; corolla white to yellow, orange or blue or
Corynostylis Mart. & Zucc., Nov. Gen. Sp. Pl. 1: 25–26 multicolored with or without yellow throat, strongly
(1824). zygomorphic, bottom petal slightly to much
shorter than others and weakly differentiated,
Lianas; leaves oblong-lanceolate to ovate. Flowers spur scarcely exserted to very long; filaments free,
in axillary or terminal racemoids, rarely solitary; two lowest stamens calcarate, dorsal connective
corolla white, strongly zygomorphic, bottom appendage large, oblong-ovate, entire; style filiform
petal slightly longer than others and weakly dif- or clavate and globose to rostellate at tip. Capsule
ferentiated, spur very long; filaments free, two thick-walled. Seeds few to many per carpel, obovoid
lowest stamens calcarate, dorsal connective to globose, typically arillate. 2n ¼ 10, 12, 14, 16, 18,
appendage large, oblong-ovate, entire; style fili- 20, 22, 24, 26, 32, 34, 36, 40, 44, 46, 48, 52, 54, 58, 60,
form. Capsule thick-walled, semi-dehiscent. 70, 72, 74, 80, 100, 104, 128.
Seeds numerous per carpel, asymmetrically Ca. 525 spp., temperate regions and montane
orbicular in outline, strongly flattened. areas in the tropics worldwide.
Four spp., Mesoamerica, South America.
20. Schweiggeria Spreng. Selected Bibliography

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Addition to Peridiscaceae
C. B A Y E R AND S. D R E S S L E R
The placement of the problematic genus Medu- Two species, M. richardsiana Brenan and
sandra in Peridiscaceae reported by Wurdack M. mpomiana Letouzey & Satabié, trop. W
and Davis (2009) motivates us to the following Africa.
addition to the treatment of the family in Vol. IX When he described Medusandra, Brenan
of this series, pp. 297–300. (1952) placed it in a unispecific family and
order of its own, which was thought to be
Peridiscaceae related to Olacales or Santalales. In 1954, he
added Soyauxia (formerly in Passifloraceae or
syn. Medusandraceae Brenan, Kew Bull. 1952: 228 (1952), Flacourtiaceae) to that family, a view that was
nom. cons.
followed by many but rejected by Hutchinson
Medusandra Brenan Fig. 74 (1959). Medusandra differs from Soyauxia in
the presence of secretory canals, a more
Medusandra Brenan, Kew Bull. 1952: 228 (1952).
condensed inflorescence, and floral characters
Trees up to 20 m, exuding a yellowish to reddish such as the apert aestivation of the sepals, sta-
sap when cut; young parts pubescent. Leaves spi- men number, the presence of (at anthesis)
ral, simple, elliptic to ovate, pinnatinerved, faintly elongate, finely pubescent staminodia, and the
toothed, minutely pubescent below; petiole long, lack of a disk; the latter traits, together with the
with proximal and distal pulvinus; stipules small, catkin-like inflorescences, call for observations
caducous. Inflorescences axillary, often paired of the mode of pollination. Based on molecular
(rarely 3 per axil), racemose, more or less lax and studies, Soyauxia was placed in Peridiscaceae
pendulous, or condensed, with caducous bracts. (Davis and Chase 2004), and Medusandra
Flowers hermaphroditic, actinomorphic, penta- remained in an unigeneric family related to
merous, small, scented; sepals distinct or almost so, Passifloraceae in the Malpighiales (Soltis et al.
persistent, aestivation apert; petals distinct, 2005). However, Wurdack and Davis (2009)
imbricate, white to greenish; fertile stamens ante- received strong support for its placement in
petalous, distinct but adnate to base of petals, Saxifragales as sister to Peridiscus and
anthers tetrasporangiate, introrse, dehiscing by Soyauxia. This is corroborated by floral fea-
valves; staminodia antesepalous, pubescent, tures (unilocular ovary with central placenta-
folded in bud, at anthesis much longer than tion and pendulous ovules as well as distinct
stamens, with rudimentary anther; ovary supe- stylodia), similar seed coat structure, small
rior, 3–4-carpellate, unilocular, with central col- embryo and an insertion in 18S rDNA consid-
umn and apical placenta; ovules 2 per carpel, ered unique in Saxifragales. Nevertheless, Peri-
pendulous, anatropous; stylodia 3–4, short, dis- discaceae differ from the bulk of Saxifragales
tinct. Fruit a globose, 3–4-valved, one-seeded cap- by syncarpy of three (to five) carpels and
sule. Seed large, albuminous, somewhat ruminate; hypogyny.
embryo small, straight, peripheral.
324 C. Bayer and S. Dressler
Fig. 74. Peridiscaceae. Medusandra richardsiana. A and base of staminode. F Dehisced fertile anther. G Ovary
Flowering twig. B Stipule and bud. C Flower bud with with stylodia. H Mature fruit. I Same, dehisced. J Seed.
bract. D Anthetic flower. E Two fertile dehisced anthers (Brenan 1952)
Metcalfe, C.R. 1952. Medusandra richardsiana Brenan.

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Index
References to accepted names in bold-faced print, to synonyms in upright print, to illustrations in italics.
Aalius, 79 Amyrea, 132 Baccaurea, 86

Acalypha, 124 Anabaena, 148 Bafodeya, 24
Acalypheae, 116 Anabaenella, 148 Balakata, 199
Acalyphinae, 124 Anchietea, 319, 319 Balanopaceae, 6, 9
Acalyphoideae, 106 Andrachne, 68 Balanops, 10, 10
Acanthocaulon, 151 Andrachne sect. Phyllantopsis, 66 Balanopseae, 9
Achariaceae, 6 Andrachne sect. Baliospermum, 177
Acidocroton, 168 Pseudophyllanthus, 67 Baloghia, 172
Acidoton, 152 Androstachydaceae, 92 Baloghiinae, 171
Acioa, 23 Androstachys, 93, 94 Beltrania, 155
Actephila, 66 Aneulophus, 44, 47 Benoistia, 188
Actinostemon, 193 Angostylis, 147 Bergia, 41
Adelia, 144 Anisadenia, 240, 243 Bernardia, 145
Adelieae, 143 Annesijoa, 182 Bernardieae, 145
Adenanthe, 266 Anomalocalyx, 174 Bertya, 179
Adenarake, 267 Anomostachys, 199 Bertyinae, 177, 178
Adenochlaena, 137 Anopyxis, 290 Beyeria, 179
Adenocline, 158 Anthodiscus, 14, 15 Bhesa, 18
Adenoclineae, 157 Anthostema, 206 Bia, 149
Adenoclininae, 158 Anthosteminae, 205 Bischofia, 89
Adenogyne, 198 Antidesma, 81 Bischofiaceae, 88
Adenopeltis, 202 Antidesmateae, 80 Bischofieae, 88
Adenophaedra, 145 Antidesmatinae, 80, 81 Blachia, 176
Adriana, 116 Antidesmatoideae, 80 Blastemanthus, 256, 265
Aerisilvaea, 76 Antidesmodae, 80 Blepharistemma, 288
Afrolicania, 23 Aparisthmium, 113 Blotia, 74
Afrotrewia, 155 Apodandra, 148 Blumeodendrinae, 124
Agatea, 318 Apodiscus, 83 Blumeodendron, 125
Agrostistachydeae, 110 Aporosa, 85 Bocquillonia, 114
Agrostistachys, 110 Aporuseae, 84 Bonania, 202
Alchornea, 112, 113 Archileptopus, 66 Bonnetiaceae, 6
Alchorneeae, 112 Argomuellera, 132 Borneodendron, 178
Alchorneinae, 112 Argythamnia, 140 Bossera, 112
Alchorneopsis, 109 Aristogeitonia, 92 Botryophora, 126
Aleurites, 183 A. sect. Ditaxis, 140 Boutonia, 128
Aleuritideae, 182 A. sect. Haematostemon, 147 Brackenridgea, 263
Aleuritinae, 182 A. sect. Philyra, 139 Brasiliocroton, 169
Algernonia, 199 Ashtonia, 86 Breynia, 80
Allantospermum, 235, 236 Astraea, 168 Breyniopsis, 79
Allexis, 316 Astrocasia, 73 Bridelia, 71
Alphandia, 172 Astrocasiinae, 72 Bridelieae, 68
Alsodeia, 315 A. subg. Chiropetalum, 140 Bruguiera, 292, 292
Amanoa, 69 Atuna, 23 Bruxanellia, 176
Amanoinae, 69 Aubletiana, 114, 115 B. sect. Adenophaedra, 145
Amperea, 134 Austrobuxus, 99 B. sect. Bia, 150
Ampereeae, 133 Avellanita, 123 B. sect. Zuckertia, 150
Amphirrhox, 317 Avellanitinae, 123 Bureavia, 99
DOI 10.1007/978-3-642-39417-1, # Springer-Verlag Berlin Heidelberg 2014
328 Index
Caletieae, 96 Clutia, 102 Deutzianthus, 186

Calophyllaceae, 6 Clutieae, 102 Diasperus, 78
Calycopeplus, 207 Cnemidostachys, 196 Dichapetalaceae, 6, 33
Calyptrion, 320 Cnesmone, 152 Dichapetalum, 34, 36
Campylospermum, 264 Cnidoscolus, 162 Dichostemma, 206
Canaca, 100 Cocconerinae, 177 Dicoelia, 73
Caperonia, 141 Cocconerion, 178 Didymocistus, 81
Carallia, 291 Codiaeae, 170 Dimorphocalyx, 174
Caryocar, 14, 15 Codiaeinae, 175 Diplomorpha, 79
Caryocaraceae, 6, 13 Codiaeum, 176 Discocarpus, 70
Caryodendreae, 108 Coelebogyne, 113 Discoclaoxylon, 121
Caryodendron, 109 Coelodiscus, 127 Discocleidion, 146
Cassipourea, 289, 289 Colliguaja, 204 Discoglypremna, 109
Cathetus, 78 Colobocarpos, 169 Dissilariinae, 99
Cavacoa, 184 Comiphyton, 289 Dissiliaria, 100
Celaenodendron, 96 Commia, 204 Distichirhops, 85, 85
Celianella, 87 Conceveiba, 114, 114 Ditaxinae, 139
Cenesmon, 152 Conceveibastrum, 114 Ditaxis, 140
Centroplacaceae, 6, 17 Conceveibinae, 114 Ditrysinia, 197
Centroplacus, 18 Conosapium, 204 Dodecastigma, 174
Cephalocroton, 137 Cordemoya, 128 Domohinea, 185
Cephalocrotonopsis, 137 Corynostylis, 320 Doryxylinae, 141
Cephalomappa, 137 Couepia, 24 Doryxylon, 141
Cephalomappinae, 137 Cremophyllum, 154 Droceloncia, 133
Ceratogynum, 79 Croizatia, 69 Drypetes, 275, 275
Ceriops, 293 Crossostylis, 291 Duckesia, 227
Cespedesia, 265 Croton, 169, 169 Durandea, 242
Chaenotheca, 73 Crotoneae, 167 Duvigneauia, 199
Chaetocarpeae, 102 Crotonogyne, 186 Dysopsidinae, 123
Chaetocarpus, 103, 103 Crotonogyninae, 186 Dysopsidoideae, 106
Chailletiaceae, 33 Crotonogynopsis, 155 Dysopsis, 123
Chamaesyce, 207 Crotonoideae, 156
Chascotheca, 73 Crotonopsis, 169 Echinus, 127
Cheilosa, 105 C. sect. Corylocroton, 170 Elaeophora, 148
Cheiloseae, 105 C. sect. Discoclaoxylon, 121 Elaeophorbia, 208
Cheilosoideae, 105 C. sect. Luteobrunnea, 121 Elateriospermeae, 161
Chiropetalum, 140 C. sect. Micrococca, 121 Elateriospermum, 161, 162
Chondrostylis, 111 C. subg. Croton, 170 Elatinaceae, 6, 39
Chonocentrum, 89 C. subg. Moacroton, 170 Elatine, 40, 41
Choriceras, 101 Ctenolophon, 29, 31 Elatinideae, 39
Choriophyllum, 99 Ctenolophonaceae, 6, 29 Eleutherostigma, 148
Chorisandrachne, 74 Ctenomeria, 151 Elvasia, 255, 264
Chrozophora, 143 Cubacroton, 170 Emblica, 78
Chrozophoreae, 138 Cubanthus, 208 Endadenium, 208
Chrozophorinae, 141 Cunuria, 164 Endopleura, 227
Chrysobalanaceae, 6, 19 Cyathogyne, 81 Endosperminae, 160
Chrysobalanus, 24 Cyrillopsis, 236 Endospermum, 160
Cicca, 78 Cyrtogonone, 186 Enriquebeltrania, 155
Cladogelonium, 161 Cyttaranthus, 111 Epiprineae, 135
Cladogynos, 136 Epiprininae, 135
Claoxylinae, 119 Dactyladenia, 24 Epiprinus, 135
Claoxylon, 120 Dactylopetalum, 289 Epistylium, 78
Claoxylopsis, 120 Dalechampia, 154, 154 Eremocarpus, 169
Clavistylus, 153 Dalechampiinae, 154 Eriococcus, 78
Cleidiinae, 117 Dalembertia, 195 Erismantheae, 107
Cleidiocarpon, 136 Danguyodrypetes, 76 Erismanthus, 107
Cleidion, 118, 119 Decorsella, 316 Erythrococca, 121
Cleistanthopsis, 236 Dendrocousinsia, 198 Erythroxylaceae, 6, 43
Cleistanthus, 71 Dendrothrix, 194 Erythroxylum, 45, 48
Cliococca, 244 Desbordesia, 231 E. sect. Calycopeplus, 207
Clonostylis, 129 Desmostemon, 189 E. sect. Sclerocroton, 197
Clusiaceae, 6 Deuteromallotus, 128 Euphorbia, 207
Index 329
Euphorbiac, 6 Heterosavia, 76 Kostermanthus, 25

Euphorbiaceae, 51 Hevea, 164, 165 Krukoviella, 265
Euphorbieae, 204 Heveeae, 164 Kurziodendron, 171
Euphorbiiinae, 207 Heywoodia, 73
Euphorbioideae, 190 Hieronyma, 84 Lachnostylis, 72
Euphronia, 217, 218 Hieronyminae, 84 Lacunaria, 278, 279, 280
Euphroniaceae, 6, 217 Hippomane, 201 Lasiococca, 129
Euthemis, 266 Hippomaneae, 191 Lasiococcinae, 129
Excoecaria, 204 Hirtella, 25 Lasiocroton, 144
Exellodendron, 25 Holstia, 185 Lauradia, 267
Homalanthus, 196 Leeuwenbergia, 181, 181
Fahrenheitia, 189 Homonoia, 130, 130 Leidesia, 123
Falconeria, 203 Hugonia, 237, 242 Leitgebia, 267
Fleurydora, 265 Hugoniaceae, 237 Leonia, 314, 315
Flueggea, 77 Hugonioideae, 242 Leonioideae, 314
Flueggeinae, 75 Humbertiodendron, 300 Leptonema, 84
Foersteria, 80 Humiria, 227 Leptonematinae, 84
Fontainea, 172 Humiriaceae, 6, 223 Leptopus, 66
Froesia, 280 Humiriastrum, 227 Leucocroton, 144
Fusispermoideae, 314 Humirieae, 226 Licania, 20, 26
Fusispermum, 314 Hunga, 25 Linaceae, 6, 237
Hura, 199, 200 Lingelsheimia, 76
Galearia, 270, 271 Hureae, 191 Linoideae, 243
Garcia, 183 Hyaenanche, 95 Linum, 240, 243
Garciadelia, 144 Hyaenanchinae, 94 Lobanilia, 121
Garciinae, 183 Hyaenanchoideae, 90 Loerzingia, 186
Gavarretia, 114 Hybanthopsis, 318 Longetia, 101
Gelonieae, 160 Hybanthus, 318 Lophira, 255, 264
Gelonium, 160 Hyeronima, 84 Lophopyxidaceae, 6, 247
Gitara, 152 Hylandia, 172 Lophopyxidoideae, 247
Givotia, 180 Hylocarpa, 227 Lophopyxis, 248, 248
Glochidion, 79 Hymenanthera, 316 Luxemburgia, 256, 262
Gloeospermum, 316, 317 Hymenocardia, 82, 83 Luxemburgieae, 262
Glycydendron, 165 Hymenocardiinae, 81 Luxemburgioideae, 262
Glyphostylus, 204 Hypericaceae, 6
Godoya, 264 Mabea, 197, 198
Gomphia, 263 Idertia, 263 Macaranga, 127
Gonatogyne, 69 Indorouchera, 242 Macarisia, 290
Goupia, 220, 220 Indosinia, 267 Macarisieae, 288
Goupiaceae, 6, 219 Indovethia, 267 Macraea, 78
Grangeria, 25 Irvingia, 231 Maesobotrya, 86
Grimmeodendron, 202 Irvingiaceae, 6, 229 Magnistipula, 26
Grossera, 184 Isidodendron, 300 Malesherbiaceae, 6
Grosserinae, 184 Isodendrion, 317 Mallotus, 126, 127, 128
G. sect. Ditrysina, 197 Ixonanthaceae, 6, 233 Malpighiaceae, 6
G. subg. Galearia, 271 Ixonanthes, 235 Mancanilla, 201
G. subg. Orthopetalum, 271 Manihot, 162
Gymnanthes, 195 Jablonskia, 87 Manihoteae, 161
Gymnorinorea, 316 Jablonskieae, 87 Manihotoides, 162
Gymnostillingia, 203 Jatropha, 166 Manniophyton, 187
Gynotrocheae, 290 Jatropheae, 166 Mappa, 128
Gynotroches, 291 Joannesia, 166 Maprounea, 193
Julocroton, 169 Maranthes, 26
Haematostemon, 147, 148 Mareya, 120
Halliophytum, 92 Kairothamnus, 97 Mareyopsinae, 115
Hamilcoa, 191 Kandelia, 293 Mareyopsis, 115
Hancea, 128, 128 Keayodendrinae, 70 Margaritaria, 75
Hasskarlia, 158 Keayodendron, 70 Martretia, 83
Hebepetalum, 239, 243 Kirganelia, 78 Martretieae, 83
Hekkingia, 317 Klaineanthus, 159 Mayanaea, 319
Hemicicca, 78 Klainedoxa, 230, 231 Medusagynaceae, 6, 249
Hesperolinon, 244 Kleinodendron, 69 Medusagyne, 250, 251
Heterocalymnantha, 79 Koilodepas, 136 Medusandra, 323, 324
330 Index
Medusandraceae, 323 Ochthocosmus, 234, 236 Phyllanthus, 78, 78

Megalostylis, 154 Octospermum, 127 Phyllanthus sect. Diandri, 75
Megistostigma, 153 Oldfieldia, 94, 95 Phyllanthus subg. Phyllanthopsis, 66
Meineckia, 68 Oldfieldioideae, 90 Phyllocosmus, 236
Melanolepis, 142 Oligoceras, 185 Picrodendraceae, 51, 90, 91
Melanthesa, 80 Omphalea, 159, 159 Picrodendreae, 91
Melanthesopsis, 80 Ophellantha, 168 Picrodendrinae, 95
Melicytus, 316 Ophthalmoblapton, 200 Picrodendron, 95
Mercurialinae, 122 Oreoporanthera, 67 Pimelodendron, 191
Mercurialis, 122 Orfilea, 112 Pinacopodium, 48
Micrandra, 163 Orthion, 318 Piranhea, 96
Micrandreae, 163 Ostodeinae, 173 Pistillate, 94
Micrandropsis, 164 Ostodes, 173 Plagiocladus, 75, 76
Micrantheum, 98 Ouratea, 255, 263 Plagiostyles, 191
Micrococca, 121 Platygyna, 151
Microdesmis, 270, 271 Pachystemon, 128 Pleradenophora, 201
Microstachys, 196, 196 Pachystroma, 200, 201 Plukenetia, 148
Mildbraedia, 188, 189 Pachystromateae, 191 Plukenetieae, 146
Mischodon, 93 Pachystylidium, 152 Plukenetiinae, 147
Mischodontinae, 92 Paivaeusa, 94 Podadenia, 125
Moacroton, 170 Paivaeusinae, 94 Podocalyceae, 91
Molina, 123 Palissya, 131 Podocalyx, 91
Monadenium, 208 Panda, 270, 271, 272 Podostemaceae, 6
Monotaxis, 134, 134 Pandaceae, 6, 269 Poecilandra, 266
Morelodendron, 48 Pantadenia, 175 Pogonophora, 104
Moultonianthus, 107, 108 Paracroton, 188 Pogonophoreae, 104
M. sect. Anopyxis, 290 Paracrotoninae, 188 Poinsettia, 207
M. sect. Blumeodendron, 125 Paradrypetes, 91, 293 Polyandra, 118
M. sect. Cordemoya, 128 Paragelonium, 92 Polydragma, 129
M. sect. Hancea, 128 Paranecepsia, 131 Poranthera, 67, 67
Myricanthe, 178 Parapantadenia, 175 Poranthereae, 65
Parastemon, 26 Protomegabaria, 85
Nealchornea, 190 Parinari, 20, 27 P. sect. Pseudanthus, 98
Necepsia, 131 Parodiodendron, 96 P. sect. Wetriaria, 132
Necepsinae, 131 Passifloraceae, 6 Pseudagrostistachys, 110
Neckia, 267 Pausandra, 175 Pseudanthinae, 97
Nectaropetalaceae, 43 Paypayrola, 318 Pseudanthus, 98, 99
Nectaropetalum, 48 Pedilanthus, 207 Pseudolachnostylidinae, 70
Neoboutonia, 187 Peglera, 48 Pseudolachnostylis, 71
Neoboutoninae, 187 Pellacalyx, 292 Pseudophyllanthus, 67
Neocarya, 27 Pentabrachion, 71 Pseudosagotia, 69
Neoguillauminia, 206, 207 Pentaspatella, 267 Pseudosenefeldera, 194
Neoguillauminiinae, 206 Pera, 104, 104 Ptychopyxis, 125
Neoholstia, 185 Peraceae, 51 Putranjiva, 275
Neojatropha, 188 Pereae, 104 Putranjivaceae, 6, 273
Neomanniophyton, 186 Peridiscaceae, 323 Pycnocoma, 133
Neopalissya, 131 Perissocarpa, 264 Pycnocomeae, 130
Neopycnocoma, 132 Peroideae, 102 Pycnocomina, 132
Neoroepera, 97 Petalodactylis, 289
Neoscortechinia, 105 Petalodiscus, 74 Quiina, 280
Neoshirakia, 195 Petalostigma, 97 Quiinaceae, 6, 277
Neotrewia, 127 Petalostigmatinae, 97
Niruri, 78 Philacra, 256, 262 Radcliffea, 189
Noisettia, 320 Philbornea, 242 Radiola, 244
Nothobaccaurea, 87 Philyra, 139 Reinwardtia, 243
Notoleptopus, 67 Phyllanthaceae, 51, 64 Reutealis, 183
Nymphanthus, 78 Phyllantheae, 74 Reverchonia, 78
Phyllanthinae, 77 Rhabdophyllum, 263
Ochna, 255, 262 Phyllanthodae, 65 Rhizophora, 293
Ochnaceae, 6, 253 Phyllanthodendron, 78 Rhizophoraceae, 6, 283
Ochnidae, 262 Phyllanthoideae, 64 Rhizophoreae, 292
Ochnoideae, 262 Phyllanthopsis, 66 Rhodothyrsus, 194
Index 331
Rhopalostylis, 154 Sebastiania, 198 Tirpitzia, 240, 243

Rhytidanthera, 265 Securinega, 72 Tithymalus, 207
Richeria, 85 Securineginae, 72 Touroulia, 278, 280
Richeria sect. Podocalyx, 91 Seidelia, 122 Toxicodendrum, 95
Richeriella, 77 Senefeldera, 194 Traganthus, 145
Ricineae, 115 Senefelderopsis, 194 Tragia, 150, 151
Ricinocarpeae, 177 Shirakia, 195 Tragiella, 153
Ricinocarpinae, 178 Shirakiopsis, 195 Tragiinae, 149
Ricinocarpos, 178 Shonia, 179, 179 Trevia, 127
Ricinodendreae, 180 Sibangea, 275 Trewia, 127
Ricinodendron, 180 Sinia, 267 Triadica, 194
Ricinoideae, 106 Sinopimelodendron, 136 Tridesmis, 169
Ricinus, 116 Spathiostemon, 129 Trigonia, 300, 301
Rinorea, 315, 316 Spathularia, 317 Trigoniaceae, 6, 297
Rinoreeae, 315 Spegazziniophytum, 202 Trigoniastrum, 300
Rinoreocarpus, 315 Speranskia, 139 Trigoniodendron, 300
Rockinghamia, 138 Speranskiinae, 138 Trigonopleura, 103
Rockinghamiinae, 138 Sphaerostylis, 153 Trigonostemon, 171
Romanoa, 148 Sphyranthereae, 111 Trigonostemoninae, 171
Roraimanthus, 267 Spirostachys, 204 Trilocularia, 10
Rottlera, 127 Spondiantheae, 87 T. sect. Bia, 149
Rottlerinae, 126 Spondianthus, 88 T. sect. Ctenomeria, 151
Roucheria, 242 S. sect. Falconeria, 203 Turneraceae, 6
R. subtribe Hymenantherinae, 316 S. sect. Heterosavia, 76 Tyleria, 266
R. subtribe Isodendriinae, 317 S. sect. Microstachys, 196
R. subtribe Paypayrolinae, 317 S. sect. Pleurostachya, 199 Uapaca, 88, 89
R. subtribe Rinoreinae, 315 S. sect. Stillingiopsis, 197 Uapacaceae, 88
S. subg. Schistostemon, 227 Uapaceae, 88
Sacoglottis, 228 S. subg. Sclerocroton, 197
Sacoglottis subg. Humiriastrum, 227 Stachyandra, 94 Vantanea, 224, 226
Sagotia, 168 Stachystemon, 98 Vantaneeae, 226
Salicaceae, 6 Stephanopodium, 34, 36 Vaupesia, 167
Sampantaea, 118 Sterigmapetalum, 290 Vausagesia, 267
Sandwithia, 167 Stilago, 81 Veconcibea, 114
Sankowskya, 100, 101 Stillingia, 203 Vernicia, 183
Sapium, 203 Stomatocalyceae, 190 Victorinia, 162
Sarothrostachys, 199 Strophioblachia, 176 Vigia, 148
Sauropus, 79 Sumbaviopsis, 142 Viola, 316, 320
Sauvagesia, 256, 267 Suregada, 160 Violaceae, 6, 303
Sauvagesieae, 264 Symphyllia, 136 Violeae, 318
Sauvages ioideae, 264 Synadenium, 207 Violoideae, 315
Savia, 69 Syndyophyllum, 108 Voatamalo, 93
Savia sect. Gonatogyne, 69 Synostemon, 79 V. subtribe Hybanthinae, 318
Savia sect. Petalodiscus, 74 V. subtribe Violinae, 319
Savia Willd. sect. Wielandia, 74 Tacarcuna, 70
Savieae, 69 Taeniosapium, 204 Wallacea, 266
Saviinae, 69 Tandonia, 185 Weihea, 289
Scagea, 98 Tapoides, 185 Wetria, 117, 118
Scepa, 85 Tapura, 34, 36 Wetriaria, 133
Scepeae, 84 Testulea, 265 Whyanbeelia, 100
Schinziophyton, 181 Tetracoccinae, 92 Wielandia, 74
Schistostemon, 227 Tetracoccus, 92 Wielandieae, 72
Schuurmansia, 266 Tetraplandra, 200 Wielandiinae, 73
Schuurmansiella, 266 Tetrorchidium, 158
Schweiggeria, 320 Tetrorchiopsis, 158 Zimmermannia, 68
Sclerocroton, 197 Thecacoris, 81, 82 Zimmermanniopsis, 68
Sclerolinon, 244 Thyrsanthera, 142 Zuckertia, 149
Volumes published in this series
Volume I Pteridophytes and Gymnosperms

Edited by K.U. Kramer and P.S. Green (1990)
Date of publication: 28.9.1990
Volume II Flowering Plants. Dicotyledons. Magnoliid, Hamamelid

and Caryophyllid Families
Edited by K. Kubitzki, J.G. Rohwer, and V. Bittrich (1993)
Volume III Flowering Plants. Monocotyledons: Lilianae (except Orchidaceae)

Edited by K. Kubitzki (1998)
Volume IV Flowering Plants. Monocotyledons: Alismatanae and Commelinanae

(except Gramineae)
Volume V Flowering Plants. Dicotyledons: Malvales, Capparales

and Non-betalain Caryophyllales
Edited by K. Kubitzki and C. Bayer (2003)
Volume VI Flowering Plants. Dicotyledons: Celastrales, Oxalidales,

Rosales, Cornales, Ericales
Volume VII Flowering Plants. Dicotyledons: Lamiales (except Acanthaceae

including Avicenniaceae)
Edited by J.W. Kadereit (2004)
Volume VIII Flowering Plants. Eudicots: Asterales

Edited by J.W. Kadereit and C. Jeffrey (2007)
Volume IX Flowering Plants. Eudicots: Berberidopsidales, Buxales, Crossosomatales,

Fabales p.p., Geraniales, Gunnerales, Myrtales p.p., Proteales, Saxifragales,
Vitales, Zygophyllales, Clusiaceae Alliance, Passifloraceae Alliance,
Dilleniaceae, Huaceae, Picramniaceae, Sabiaceae
Volume X Flowering Plants. Eudicots: Sapindales, Cucurbitales, Myrtaceae

Dates of publication: 10.12.2010 (online); 10.1.2011 (print)
Volume XI Flowering Plants. Eudicots: Malpighiales


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Edited by K.

ISBN 978-3-642-39416-4 ISBN 978-3-642-39417-1 (eBook)

Library of Congress Control Number: 2013954369

# Springer-Verlag Berlin Heidelberg 2014

Printed on acid-free paper

Springer is part of Springer ScienceþBusiness Media (www.springer.com)

Hamburg Klaus Kubitzki

Amaral, Maria do Carmo E. do Depto de Biologı́a Vegetal, Instituto de Biologı́a,

Introduction to Malpighiales K. KUBITZKI . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1

Addition to Peridiscaceae C. BAYER and S. DRESSLER . . . . . . . . . . . . . . . . . . . . . . 323

and characterised by the possession of cristarque interesting association between Hypericaceae

A F F I N I T I E S . The affinities of Balanops were

EMBRYOLOGY. Unknown. Only one genus:

POLLEN MORPHOLOGY. The pollen is 3–5-colpate, Balanops Baill. Fig. 2

F R U I T A N D S E E D . The pericarp consists mostly Selected Bibliography

F R U I T A N D S E E D . The fruits are drupes with 1–4

1. Centroplacus Pierre to racemose. Flowers bisexual, 5-merous; calyx

Fig. 4. Chrysobalanaceae. Licania granvillei. A Flowering

calyx is imbricate in bud, 5-lobed, entire; stipitate

P O L L I N A T I O N . Chrysobalanaceae have a wide P H Y T O C H E M I S T R Y . The chemistry of Chrysoba-

– Flowers bisexual (South America and Malesia) 4 GENERA OF CHRYSOBALANACEAE

KEY TO THE GENERA Selected Bibliography

compressed inflorescence axis. Pedicels are

R EPRODUCTIVE S YSTEMS AND POLLINATION. Since

Monoecious or dioecious trees, shrubs, or herbs, deciduous in fruit; petals + or 0, greenish to

autochorous dispersal is followed or superseded P H Y T O C H E M I S T R Y . Euphorbiaceae are one of

P A L E O B O T A N Y . The Euphorbiaceae, despite DISTRIBUTION AND H A B I T A T S . Although

– Erect shrubs to trees 0.2–6 m tall (Leptopus clarkei 2. Actephila Blume

curved; cotyledons broad, thin, longer and wider

Monoecious (dioecious) annual or perennial

10. Savia Willd. Dioecious trees or shrubs. Leaves alternate to

Dioecious trees or shrubs. Stipules deciduous. 15. Keayodendron Leandri

14. Tacarcuna Huft 2d. SUBTRIBE PSEUDOLACHNOSTYLIDINAE Pax &

Monoecious (dioecious) shrubs or trees; indumen- 19. Pseudolachnostylis Pax

22. Heywoodia Sim tillode 3-fid, adnate to staminal column. Pistillate

Chascotheca Urb., Symb. Ant. 5: 14 (1904); Alain, Fl. Cuba

thyrsoid, pedunculate; glomerules often bisex- Morphologically, Chorisandrachne and Dicoelia

and connective not enlarged; ovary 2–6(15)- 4a. SUBTRIBE FLUEGGEINAE M€

straight or curved, cotyledons longer than the

30. Lingelsheimia Pax

This subtribe, with 5 genera and nearly 1,000

34. Phyllanthus L. Fig. 13

on persistent or deciduous branchlets, sometimes Monoecious (dioecious) trees or shrubs; branch-

Peru; seed dispersal probably hydrochorous

41. Hymenocardia Wall. ex Lindl. Fig. 15

42. Martretia Beille

ovary (3)4(5)-locular; stylodia unlobed, recurved. Dioecious (monoecious) shrubs; indumentum

54. Jablonskia G.L. Webster 8. TRIBE SPONDIANTHEAE G.L. Webster (1975).

at least the staminate, petaliferous, subsessile; 57. Uapaca Baill. Fig. 16

flowers pedicellate; sepals 5, imbricate, cucullate-

– Leaf blades undivided [pseudo-trifoliolate in 60a. Paradrypetes Kuhlm.

interspinal tectum psilate; seeds ecarunculate petiole or intrapetiolar-connate; pollen 5–8-

63. Mischodon Thwaites 65. Androstachys Prain Fig. 17

Four spp., endemic to Madagascar. Closely

2c. SUBTRIBE PAIVAEUSINAE Pax & K. Hoffm. (1922).

67. Oldfieldia Benth. & Hook. f. Fig. 18

anthers introrse to extrorse, dehiscing longitudi-

2e. SUBTRIBE PICRODENDRINAE (Small) G.L. Webster

KEY TO THE GENERA OF PICRODENDRINAE

Nine spp., endemic to southwestern Western

3c. SUBTRIBE DISSILARIINAE Pax & K. Hoffm. (1922).

KEY TO THE GENERA OF DISSILARIINAE

cymose, unisexual or bisexual. Staminate flowers

85. Longetia Baill.