Acanthaceae (PDF)

ACANTHACEAE爵床科 jue chuang keHu Jiaqi ( 胡嘉琪 Hu Chia-chi) 1 , Deng Yunfei ( 邓云飞 ) 2 ; John R. I. Wood 3 , Thomas F. Daniel 4Prostrate, erect, or rarely climbing herbs (annual or perennial), subshrubs, shrubs, or rarely small trees, usually with cystoliths(except in following Chinese genera: Acanthus, Blepharis, Nelsonia, Ophiorrhiziphyllon, Staurogyne, and Thunbergia), isophyllous(leaf pairs of equal size at each node) or anisophyllous (leaf pairs of unequal size at each node). Branches decussate, terete to angularin cross-section, nodes often swollen, sometimes spinose with spines derived from reduced leaves, bracts, and/or bracteoles. Stipulesabsent. Leaves opposite [rarely alternate or whorled]; leaf blade margin entire, sinuate, crenate, dentate, or rarely pinnatifid. Inflorescencesterminal or axillary spikes, racemes, panicles, or dense clusters, rarely of solitary flowers; bracts 1 per flower or dichasialcluster, large and brightly colored or minute and green, sometimes becoming spinose; bracteoles present or rarely absent, usually 2per flower. Flowers sessile or pedicellate, bisexual, zygomorphic to subactinomorphic. Calyx synsepalous (at least basally), usually4- or 5-lobed, rarely (Thunbergia) reduced to an entire cupular ring or 10–20-lobed. Corolla sympetalous, sometimes resupinate 180ºby twisting of corolla tube; tube cylindric or funnelform; limb subactinomorphic (i.e., subequally 5-lobed) or zygomorphic (either 2-lipped with upper lip subentire to 2-lobed and lower lip 3-lobed, or rarely 1-lipped with 3 lobes); lobes ascending or descendingcochlear, quincuncial, contorted, or open in bud. Stamens epipetalous, included in or exserted from corolla tube, 2 or 4 and didynamous;filaments distinct, connate in pairs, or monadelphous basally via a sheath (Strobilanthes); anthers with 1 or 2 thecae; thecaeparallel to perpendicular, equally inserted to superposed, spherical to linear, base muticous or spurred, usually longitudinally dehiscent;staminodes 0–3, consisting of minute projections or sterile filaments. Disk annular and nectar-producing at base of ovary. Ovarysuperior, 2-locular, placentation axile, with 2 to many ovules per locule; style filiform, simple; stigma funnel-shaped or 2-lobed (onelobe sometimes suppressed), one or both lobes sometimes recurved or recoiled. Fruit a loculicidal, stipitate or not, 2-valved, explosivelydehiscent capsule [rarely drupaceous], 2- to many seeded, apex with or without a beak; septa remaining attached to orseparating from inner wall of mature capsule. Seeds usually borne on hooklike retinacula (attached to septa of capsule), or retinaculalacking (Nelsonia, Ophiorrhiziphyllon, Staurogyne, Thunbergia), surface smooth or roughened, pubescent or lacking trichomes,sometimes with hygroscopic trichomes that expand when moistened.About 220 genera and ca. 4,000 species: pantropical and subtropical, with a few species in temperate regions; 35 genera (one endemic) and 304species (134 endemic, eight introduced) in China.Pollen characters have traditionally been important in delimiting genera and higher taxa in the family; they are used sparingly in the key below.Avicennia Linnaeus, usually placed in either Verbenaceae (and treated as such in Fl. China 17: 49. 1994) or Avicenniaceae, is now included inAcanthaceae on the basis of molecular phylogenetic studies (e.g., A. E. Schwarzbach & L. A. McDade, Syst. Bot. 27: 84–98. 2002).The major economic importance of the family is horticultural. Species of Acanthus, Aphelandra R. Brown, Barleria, Crossandra Salisbury,Eranthemum, Fittonia Coemans, Justicia, Odontonema Nees, Pachystachys Nees, Ruellia, Sanchezia Ruiz & Pavon, Thunbergia, and numerous othergenera are widely cultivated as ornamentals. Many species are cultivated in China but not treated in the present account. These include: Aphelandrasquarrosa Nees, Barleria lupulina Lindley, Crossandra infundibuliformis (Linnaeus) Nees, Eranthemum pulchellum Andrews, Fittonia albivenis(Veitch) Brummitt, Justicia betonica Linnaeus, J. brandegeeana Wasshausen & L. B. Smith, J. carnea Lindley, Pachystachys lutea Nees, Ruelliaelegans Poiret, R. simplex C. Wright, Sanchezia parvibracteata Sprague & Hutchinson, Thunbergia erecta (Bentham) T. Anderson, and T. laurifoliaLindley.Hu Chiachi & Tsui Hongpin. 2002. Acanthaceae. In: Hu Chiachi, ed., Fl. Reipubl. Popularis Sin. 70: 1–309.1a. Vines; calyx cupular with 10–20 lobes or reduced to an entire ring; capsule with a prominent apical sterilebeak; seeds not borne on hooklike retinacula ................................................................................................................ 4. Thunbergia1b. Prostrate or erect herbs, shrubs, or rarely small trees; calyx 4- or 5-lobed; capsule lacking a prominent apicalsterile beak; seeds borne on hooklike retinacula or not.2a. Cystoliths absent; retinacula absent (except Acanthus and Blepharis).3a. Seeds laterally compressed, borne on hooklike retinacula; corolla 1-lipped (upper lip absent), lobesnot with descending cochlear aestivation; anthers 1-thecous.4a. Leaves opposite; flowers not subtended by up to 5 pairs of bracts; seeds lacking hygroscopic trichomes;lower filaments without an apical appendage ............................................................................................... 5. Acanthus4b. Leaves usually in pseudo-whorl of 4; flowers subtended by up to 5 pairs of bracts; seeds withhygroscopic trichomes; lower filaments with an apical appendage ............................................................. 6. Blepharis3b. Seeds subspherical or blocky, not borne on hooklike retinacula; corolla 2-lipped, lobes with descendingcochlear aestivation; anthers 2-thecous.1 Department of Ecology and Evolution, School of Life Sciences, Fudan University, 220 Handan Road, Shanghai 200433, People’s Republic of China.2 Key Laboratory of Plant Resources Conservation and Sustainable Utilization, South China Botanical Garden, Chinese Academy of Sciences, 723 Xingke Lu, Tianhe Qu,Guangzhou, Guangdong 510650, People’s Republic of China.3 Department of Plant Sciences, University of Oxford, South Parks Road, Oxford OX1 3RB, United Kingdom.4 Department of Botany, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, California 94118-4503, U.S.A.369

370ACANTHACEAE5a. Prostrate herbs; bracteoles absent; calyx 4-lobed, anterior lobe 2-cleft at apex; stamens 2 ......................... 1. Nelsonia5b. Erect herbs; bracteoles present; calyx 5-lobed; stamens 2 or 4.6a. Stamens 4, included or slightly exserted from mouth of corolla; anther thecae of a pairparallel; staminodes 0 or 1 ................................................................................................................... 2. Staurogyne6b. Stamens 2, long exserted; anther thecae of a pair divergent toward apex; staminodes 2 ...... 3. Ophiorrhiziphyllon2b. Cystoliths present; retinacula present, hooklike.7a. Calyx lobes heteromorphic, posterior lobe usually largest, anterior lobes connate to a greaterdegree than other lobes, lateral lobes smallest and innermost; corolla lobe aestivation quincuncial.8a. Corolla large (mostly > 3 cm), upper lip usually appearing 4-lobed and lower lip appearing 1-lobed ....... 29. Barleria8b. Corolla small (< 3 cm), upper lip 2-lobed and lower lip 3-lobed.9a. Corolla to 1 cm; anther thecae usually lacking basal appendages; capsule 4–7 mm .................... 30. Lepidagathis9b. Corolla ca. 2.5 cm; anther thecae spurred at base; capsule 12–16 mm ............................................ 31. Chroesthes7b. Calyx lobes homomorphic (or nearly so; sometimes partially fused to form a bipartite or tripartitecalyx in Strobilanthes), not as described above; corolla lobe aestivation not quincuncial.10a. Corolla lobes contorted in bud; stamens (or stamens + staminodes) 4.11a. Style held in place by trichomes on one side of corolla tube; stamens monadelphous(by a sheath) at base .................................................................................................................... 7. Strobilanthes11b. Style not held in place by trichomes on one side of corolla tube; stamens not monadelphousat base.12a. Inflorescence secund strobilate spikes with orbicular to reniform bracts; ovules 2 perlocule; seeds to 4 per capsule; corolla ca. 0.6 cm; septa with attached retinaculaseparating from inner wall of mature capsule ...................................................................... 8. Phaulopsis12b. Inflorescence not as described above; ovules 4 to many per locule; seeds 4 or moreper capsule; corolla 0.6–5.5 cm; septa with attached retinacula not separating frominner wall of mature capsule (except in Ruellia blechum).13a. Corolla limb distinctly 2-lipped, lobes usually not equal; calyx tube oftenrupturing in weak hyaline regions between lobes; pollen 4-colporate,interapertural regions multistriate with pseudocolpi ................................................... 9. Hygrophila13b. Corolla limb nearly regular, lobes equal to subequal; mature calyx tube notrupturing in weak hyaline regions between lobes; pollen otherwise.14a. Fertile stamens 2, staminodes 2 ..................................................................... 10. Eranthemum14b. Fertile stamens 4.15a. Anther thecae spurred at base ........................................................... 11. Echinacanthus15b. Anther thecae muticous at base.16a. Stems short; leaves in a basal rosette; anther thecae usuallywidely separated by a broad connective ....................................... 12. Pararuellia16b. Stems elongate; leaves cauline; anther thecae not separatedby a broad connective ........................................................................... 13. Ruellia10b. Corolla lobes ascending cochlear in bud; stamens 2 (except 4 in Asystasia), staminodes 0 or 2.17a. Ovules 3 to many per locule; seeds 6 to many per capsule; pollen with apertural marginsconspicuously thickened.18a. Capsule compressed perpendicular to septum; seeds subglobose, glabrous .................. 33. Andrographis18b. Capsule linear and 4-angled, not compressed perpendicular to septum; seedsstrongly compressed, glabrous or pubescent.19a. Staminodes 0 ..................................................................................................... 32. Gymnostachyum19b. Staminodes 2.20a. Corolla tube cylindric and slightly curved ................................................. 34. Phlogacanthus20b. Corolla tube abruptly inflated near middle and usually bent ca. 90° .......... 35. Cystacanthus17b. Ovules 2 per locule; seeds 4 per capsule; pollen with apertural margins not thickened.21a. Stamens 4 ............................................................................................................................... 14. Asystasia21b. Stamens 2.22a. Anthers 1-thecous.23a. Flowers subtended by a single pair of bracteoles; corolla not resupinate180° by a twisting of corolla tube (i.e., upper lip entire to 2-lobed andlower lip 3-lobed), or if position of lips reversed then this by geniculation(without twisting) of corolla tube .................................................................. 18. Clinacanthus23b. Flowers subtended by an involucre of 2 or more pairs of floral bracteoles;corolla resupinate 180° by twisting of corolla tube (i.e., lip in upper position3-lobed and lip in lower position entire to 2-lobed) .......................................... 28. Hypoestes

ACANTHACEAE 37122b. Anthers 2-thecous.24a. Flowers subtended by involucre of 2 or more pairs of floral bracteoles;corolla resupinate 180° (i.e., lip in upper position 3-lobed and lip inlower position entire to 2-lobed).25a. Septa with attached retinacula separating from inner wall of maturecapsule ....................................................................................................... 26. Dicliptera25b. Septa with attached retinacula remaining attached to inner wall ofmature capsule ........................................................................................ 27. Peristrophe24b. Flowers subtended by a single pair of bracteoles (or bracteoles rarelylacking); corolla not resupinate 180° (i.e., upper lip entire to 2-lobedand lower lip 3-lobed).26a. Staminodes 2.27a. Corolla ± salverform (i.e., tube slender and cylindric,limb spreading and inconspicuously 2-lipped), tubeconspicuously longer than limb, not or only slightlywidened distally ................................................................. 15. Pseuderanthemum27b. Corolla ± campanulate to strongly 2-lipped (notsalverform), tube not longer than limb, graduallywidened distally.28a. Corolla 3–4 cm; anther thecae 4–5 mm .................................. 16. Mackaya28b. Corolla less than 2 cm; anther thecae less than 3 mm.29a. Corolla campanulate, 6.5–10 mm, externallyglabrous; pollen 4-aperturate ............................... 22. Codonacanthus29b. Corolla strongly 2-lipped, 10–15 mm, externallypubescent (to subglabrous); pollen 3-aperturate............................................................................. 23. Cosmianthemum26b. Staminodes absent.30a. Upper lip of corolla lacking a stylar furrow on internalsurface; anther thecae lacking basal appendages.31a. Corolla 1.7–4.5 cm; anther thecae subequally to unequallyinserted; pollen 2-aperturate .................................................... 17. Isoglossa31b. Corolla less than 10 mm; anther thecae equally to subequallyinserted; pollen 3–5-aperturate.32a. Corolla glabrous outside ........................................ 19. Kudoacanthus32b. Corolla pubescent outside ........................................ 21. Leptostachya30b. Upper lip of corolla rugulate (i.e., with a furrow on innersurface of lip in which style resides); anther thecae (atleast lower theca) usually with a basal appendage.33a. Septa with attached retinacula separating from inner wallof mature capsule; inflorescence often dense with imbricatebracts 2- or 4-ranked (but only 2 ranks fertile) ........................... 20. Rungia33b. Septa with attached retinacula remaining attached to innerwall of mature capsule; inflorescence not as above.34a. Corolla not with combination of characters notedbelow ................................................................................. 24. Justicia34b. Corolla white or greenish white, tube narrowlycylindric, conspicuously longer than limb .............. 25. Rhinacanthus1. NELSONIA R. Brown, Prodr. 480. 1810.瘤子草属 liu zi cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, pubescent, without cystoliths. Leaves opposite, shortly petiolate; leaf blade pinnately veined, margin entire. Inflorescencesaxillary or terminal spikes; bracts overlapping. Flowers ± spirally arranged, sessile, subtended by a bract; bracteoles absent.Calyx 4-lobed to base; lobes unequal, anterior lobe apically 2-cleft. Corolla tube slender, sometimes widening into a throat; limb 2-liped; lower lip 3-lobed, upper lip 2-lobed; lobes ± equal, descending cochlear in bud. Stamens 2, included or slightly exserted; filamentsbasally pubescent; anthers 2-thecous; thecae subspherical, muticous, opening at base by a pore that has a small flap; stami-

372ACANTHACEAEnodes absent. Ovary with 8–28 ovules in 2–4 rows per locule; stigma usually unequally 2-lobed. Capsule conical; retinacula absent.Seeds subspherical, small, flattened on 1 face, lacking trichomes.Five species or fewer: widely occurring in tropical Africa, Asia, Australia, and South America (introduced elsewhere); one species in China.A single species, Nelsonia canescens, is sometimes recognized (e.g., A. B. M. E. Hossain, Willdenowia 14: 397–403. 1984), but Vollesen (e.g.,K. Vollesen in J. H. Seyani & A. C. Chikuni, eds., Proceedings of the XIII Plenary Meeting of AETFAT, Zomba, Malawi, 2–11 Apr 1991, 315–325.1994) presented arguments for recognizing up to five species.1. Nelsonia canescens (Lamarck) Sprengel, Syst. Veg. 1: 42.1824.瘤子草 liu zi caoJusticia canescens Lamarck, Tabl. Encycl. 1: 41. 1791.Herbs 10–20 cm tall, annual, creeping, sprawling, prostrate,or decumbent. Stems subterete, villous, often rooting atnodes. Petiole 0.2–3(–4) cm, villous; leaf blade elliptic to ovate,1–2.2 × 0.4–1.2 cm but basal ones sometimes 6–12 × 3.5–5 cm,both surfaces villous, secondary veins 3–7 on each side of midvein,base cuneate, margin entire, apex acute. Spikes 1.5–4 cm;bracts elliptic, 6–7.5 × 3–4 mm, 5–7-veined. Calyx abaxial lobeca. 2 × 0.6 mm, apex 2-lobed; adaxial lobe ca. 3 × 1 mm; laterallobes ca. 2 × 0.5 mm. Corolla bluish purple or white, externallyglabrous; tube cylindric for ca. 1.5 mm, contracted near midpointthen expanded into throat; lower lip ca. 2.3 mm; upper lipca. 2 mm. Stamens inserted at base of throat; filaments ca. 0.5mm, glabrous. Ovary glabrous; ovules 4–8 per locule. Capsuleca. 5 × 2 mm, 8–16-seeded. Seeds broadly ellipsoid, granulate.Fl. Oct–Mar, fr. Mar–May. 2n = 34 + ß, 36.Wet places, open forests; 400–2000 m. Guangxi, Yunnan [Bhutan,Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Nepal, Philippines,Thailand, Vietnam; Africa, Madagascar].2. STAUROGYNE Wallich, Pl. Asiat. Rar. 2: 80. 1831.叉柱花属 cha zhu hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielEbermaiera Nees.Annual or perennial herbs or small shrubs, without cystoliths. Stems very short or elongate. Leaves in a basal rosette or cauline,opposite (or alternate, especially apically), petiolate or sessile; leaf blade margin entire or nearly so. Inflorescences terminal or axillary,racemes or spikes; bracts opposite or alternate, leaflike or usually well differentiated from leaves; bracteoles present. Calyx 5-lobed almost to base; lobes equal or unequal. Corolla tube cylindric or basally cylindric and apically expanding into a campanulatethroat; limb subactinomorphic to 2-lipped, 5-lobed; lobes subequal, patent, descending cochlear in bud. Stamens 4, didynamous,included or anterior pair slightly exserted; filaments usually hairy; anthers 2-thecous; thecae parallel, usually equal; staminode 1 ormore, sometimes absent. Ovary with usually 12–60 ovules arranged in 2 or 4 rows per locule; style glabrous; stigmas 2-lobed, sometimesrepeatedly 2-lobed. Capsule oblong, many seeded; retinacula absent. Seeds minute, subspherical or cuboid.About 140 species: tropical regions worldwide; 17 species (eight endemic) in China.1a. Bracts pinnately veined.2a. Plants to 20 cm tall; leaf blade 2.5–6 × 1–2 cm; spikes ca. 1 cm ..................................................................... 7. S. paotingensis2b. Plants to 1 m tall; leaf blade 13–17 × 4.5–7 cm; racemes to 15 cm ....................................................................... 8. S. rivularis1b. Bracts not pinnately veined.3a. Leaf blade base cordate ....................................................................................................................................... 5. S. chapaensis3b. Leaf blade base cuneate, rounded, or attenuate.4a. Stems very short; leaves forming a basal rosette.5a. Leaf blade oblanceolate, widest above middle .................................................................................... 10. S. concinnula5b. Leaf blade ovate-oblong, elliptic, oblong, ovate, or oblong-obovate, widest below middle.6a. Leaf blade adaxially villous; corolla ca. 2 cm .......................................................................................... 6. S. sinica6b. Leaf blade adaxially strigose; corolla ca. 5.5 mm .............................................................................. 12. S. strigosa4b. Stems elongate; leaves cauline.7a. Calyx 1.8–2.4 cm; corolla 3–4 cm ......................................................................................................... 4. S. sesamoides7b. Calyx less than 1.5 cm; corolla less than 2.5 cm (corollas unknown in S. sichuanica andS. yunnanensis).8a. Leaf blade linear, 5–9 mm wide, length/width greater than 5 ...................................................... 13. S. stenophylla8b. Leaf blade lanceolate, ovate-lanceolate, oblong, or elliptic, usually broader than 1 cm,length/width less than 3.9a. Calyx lobes spatulate ................................................................................................................. 15. S. petelotii9b. Calyx lobes linear, lanceolate, or triangular.10a. Longer pair of stamens slightly exserted ........................................................................ 1. S. hypoleuca10b. Stamens all included.

ACANTHACEAE 37311a. Inflorescences all borne on axillary brachyblasts ............................................ 2. S. longicuneata11b. Inflorescences not all borne on axillary brachyblasts.12a. Inflorescences terminal only; calyx lobes unequal.13a. Bracts adaxially densely pubescent .............................................. 14. S. yunnanensis13b. Bracts glabrous or subglabrous ............................................................... 16. S. vicina12b. Inflorescences terminal and axillary; calyx lobes subequal or equal.14a. Stems suberect or ascending from a decumbent base ............................ 17. S. debilis14b. Stems erect.15a. Filaments glabrous ................................................................ 11. S. hainanensis15b. Filaments pubescent.16a. Peduncles, rachises, and pedicels puberulent .................. 3. S. sichuanica16b. Peduncles, rachises, and pedicels glabrous ............... 9. S. brachystachya1. Staurogyne hypoleuca Benoist, Notul. Syst. (Paris) 2: 338.1911.灰背叉柱花 hui bei cha zhu huaOphiorrhiziphyllon hypoleucum (Benoist) Benoist.Herbs to 1 m tall. Stems glabrous, lenticellate. Petiole 3–6cm; leaf blade oblong, elliptic, or lanceolate, 13–17 × 4.5–7 cm,both surfaces glabrous, abaxially glaucous, adaxially green, secondaryveins 8–11 on each side of midvein, base cuneate andslightly decurrent onto petiole, margin entire or undulate, apexacuminate to sometimes acute. Racemes terminal or in axils ofapical leaves, to 15 cm, unbranched or branched at base; rachisgland-tipped pilose; bracts linear, 4–4.5 × ca. 0.8 mm, apexacuminate; bracteoles linear, ca. 2 mm. Flowers alternate. Pedicel2–3.5 mm. Calyx ca. 5 mm, outside pubescent with glandtippedtrichomes; lobes triangular-ovate, ca. 4 × 1 mm, unequalwith lateral 2 lobes smaller, margin ciliate with gland-tipped trichomes.Corolla white, funnel-shaped, ca. 7 mm; lobes suborbicular,ca. 1.5 × 1.3 mm, subequal. Stamens with longer pairca. 3 mm and shorter pair ca. 2 mm; filaments glabrous; antherthecae ca. 1 mm, those of longer pair of stamens slightly exserted.Ovary ellipsoid, ca. 2 mm; ovules in 2 rows per locule,12 in each row; style ca. 3 mm, glabrous; stigma 2-cleft. Capsulecylindric, 5–7 mm. Fl. Dec–Apr.Forests; 300–1800 m. Yunnan [Vietnam].2. Staurogyne longicuneata H. S. Lo, Bull. Bot. Res., Harbin8(1): 2. 1988.楔叶叉柱花 xie ye cha zhu huaHerbs to 30 cm tall. Stems stout, 4-angled, pubescent.Petiole 2–5 cm, brown pubescent, soon glabrescent; leaf bladeelliptic to elliptic-lanceolate, 10–20 × 5–7 cm, both surfacesbrown or white pubescent, abaxially pale green, adaxially green,secondary veins 11–17 on each side of midvein, base cuneateand usually decurrent onto petiole, margin entire, apex acute toobtuse. Racemes borne on axillary brachyblasts, to 16 cm; rachisbrownish pubescent with gland-tipped trichomes; bractslinear, 3–5 × ca. 1 mm; bracteoles linear, 3–4 × ca. 1 mm. Pedicels2–5 mm. Calyx ca. 1 cm, outside brown pubescent, insideglabrous; lobes linear, ca. 9 × 1 mm, subequal. Corolla ca. 1.5cm, outside pubescent; lobes oblong, ca. 2 × 1.5 mm, subequal.Staminal filaments pubescent; anther thecae ca. 1 mm. Styleglabrous; stigma 3-lobed. Capsule ellipsoid, 5–6 mm. Seedsblack, in 2 rows, pubescent. Fl. Mar–Apr.● Forests or thickets by streams; 100–400 m. Yunnan.3. Staurogyne sichuanica H. S. Lo, Bull. Bot. Res., Harbin8(1): 2. 1988.金长莲 jin chang lianHerbs 10–30 cm tall. Stems purplish black, terete, sulcate,pubescent. Petiole purple, 1–2 cm, puberulent; leaf blade ovatelanceolateto elliptic-lanceolate, 4–9 × 1.3–3 mm, both surfacessparsely strigillose, secondary veins 6–11 on each side of midveinand abaxially inconspicuous, base cuneate to broadly cuneateand sometimes oblique, margin entire, apex acuminate. Racemesterminal or axillary, 3.5–6 cm, 2–5-flowered; peduncleand rachis slightly puberulent; bracts linear, ca. 5.5 × 0.5mm, subglabrous, 1-veined, margin ciliate; bracteoles similar tobracts. Pedicel 4–7 mm. Calyx ca. 1.1 cm, subglabrous; lobeslinear, 9–11 × 0.6–1.2 mm, subequal, 3-veined, margin ciliate.Corolla purplish blue in bud. Staminal filaments glabrous.Ovary glabrous. Capsule ca. 5 mm. Fl. Feb, fr. Jun–Jul.● Forests; 500–600 m. Sichuan (Changning, Hejiang).4. Staurogyne sesamoides (Handel-Mazzetti) B. L. Burtt, NotesRoy. Bot. Gard. Edinburgh 22: 310. 1958.大花叉柱花 da hua cha zhu huaLoxostigma sesamoides Handel-Mazzetti, Oesterr. Bot. Z.85: 217. 1936; Staurogyne dolichocalyx E. Hossain.Herbs to 40 cm tall, unbranched. Stems striate, tomentose,soon glabrescent. Petiole 1.5–4 cm, tomentose; leaf blade elliptic-lanceolate,5–13 × 2–5.5 cm, abaxially pubescent alongveins, adaxially glabrous, secondary veins 5–11 on each side ofmidvein, base cuneate, margin entire or sometimes irregularlyundulate, apex acuminate to sometimes acute. Racemes terminalor axillary, densely flowered, elongated after flowering; peduncleand rachis pubescent; bracts lanceolate, 6–7 × 2–2.2mm, pilose, 3-veined; bracteoles lanceolate, 8–8.5 × ca. 2 mm,pilose. Pedicel 0.8–1 mm but elongated to 5 mm in fruit, tomentose.Calyx 1.8–2.4 mm, outside sparsely pilose or glabrous;lobes lanceolate, 1.6–2.2 × 0.1–0.2 cm, subequal. Corollawhite or pinkish on lobes, 3–4 cm; lobes suborbicular, ca. 1cm in diam., subequal, 3-veined. Stamens with longer pair ca.1.7 cm and shorter pair ca. 1.3 cm; filaments gland-tipped hairyand apically more densely hairy; anther thecae ca. 2.5 × 1.5mm, with appendage, margin hispid. Ovary glabrous; ovules ca.60 in 2 rows; style ca. 2 cm, sparsely pubescent; stigma triangular,2-cleft; stigma lobes fimbriate on margin, posterior one

374ACANTHACEAEelliptic and ca. 3 × 2 mm, anterior one ca. 4 × 4 mm. Capsulenarrowly ellipsoid. Seeds small, alveolate. Fl. Mar–May.Moist places in forests; below 800 m. Guangdong, Guangxi [Vietnam].5. Staurogyne chapaensis Benoist, Bull. Mus. Natl. Hist. Nat.,sér. 2, 5: 172. 1933.弯花叉柱花 wan hua cha zhu huaHerbs to 10 cm tall. Stems short. Leaves in a basal rosette;petiole to 11 cm, brownish tomentose; leaf blade ovate to oblong,2.5–14.5 × 2–6 cm, abaxially glaucous and densely hairy,adaxially green and sparsely hairy, secondary veins 7–9 on eachside of midvein, base cordate, margin entire or slightly undulate,apex obtuse. Racemes terminal or axillary; peduncle ca. 4cm, villous; rachis villous; bracts obovate to linear-spatulate,2.5–3.5 cm, abaxially villous, adaxially glabrous; bracteoleslinear-spatulate, ca. 5 × 1.6 mm, abaxially villous, adaxiallyglabrous, margin ciliate, apex obtuse. Pedicel 1.5–7 mm, villous.Calyx ca. 1 cm, outside villous, inside glabrous; lobesunequal, margins ciliate, posterior one spatulate and ca. 8 × 2mm, lateral 2 linear and ca. 5 × 0.5 mm. Corolla light bluishpurple, ca. 1.5 cm; lobes orbicular, subequal. Stamens withlonger pair ca. 2 mm and shorter pair ca. 1.1 mm; filaments glabrous;anther thecae subequal, base with a rectangular appendage.Ovary ellipsoid, ca. 2 mm, glabrous; style ca. 5.5 mm,glabrous. Capsule obovoid-ellipsoid, ca. 6 mm. Fl. Mar–May,fr. Jul–Aug.Forests; 1000–2000 m. Guangdong, Guangxi, Hunan, Yunnan[Vietnam].6. Staurogyne sinica C. Y. Wu & H. S. Lo, Fl. Hainan. 3: 590.1974.中华叉柱花 zhong hua cha zhu huaTitanotrichum parviflorum X. B. Ye & W. T. Lin.Herbs, annual. Stems very short, densely pubescent.Leaves in a basal rosette; petiole 0.5–3 cm; leaf blade ovate-oblong,4–9 × 1–4.5 cm, papery, abaxially pale green and villousalong veins, adaxially green and villous, secondary veins 5–7on each side of midvein, base rounded to broadly cuneate, marginentire, apex rounded. Racemes terminal or axillary, 3–12cm; peduncle hispid; bracts linear-spatulate, 3–4 mm, pubescenton both surfaces, 3-veined, apex rounded; bracteoles similar tobracts. Pedicel 4–5 mm, hispid. Calyx 6–9 mm, outside villous;lobes linear-lanceolate, unequal, anterior and posterior ones 6–8mm and lateral ones 5–7 mm, 3-veined. Corolla whitish purple,tubular-campanulate, ca. 2 cm; tube basally cylindric for ca. 8mm; lobes oblong, 1.8–2.5 × 1.5–2 mm, slightly unequal, apexrounded. Stamens with longer pair ca. 7 mm and shorter pair ca.6 mm; filaments glabrous; anther thecae ca. 1.8 mm; staminodeca. 1 mm. Ovary ellipsoid, ca. 1.8 mm, glabrous; style ca. 8mm, glabrous. Capsule oblong-ovoid, ca. 6 mm, glabrous. Fl.Mar–May, fr. Aug.● Dense forests; 500–1200 m. Guangdong, Hainan.7. Staurogyne paotingensis C. Y. Wu & H. S. Lo, Fl. Hainan.3: 589. 1974.保亭叉柱花 bao ting cha zhu huaHerbs 10–20 cm tall, annual. Stems erect, pubescent.Leaves cauline; petiole to 4 cm; leaf blade elliptic, ellipticlanceolate,or rarely subovate, 2.5–6 × 1–2 cm, papery, abaxiallygreen and glabrous, adaxially pale green and glabrous exceptslightly pubescent along midvein, secondary veins 6–9 oneach side of midvein, base cuneate, margin entire, apex obtuseto rounded. Spikes borne on axillary brachyblasts, ca. 1 cm;peduncle and rachis pubescent; bracts elliptic to spatulateelliptic,6–7 × ca. 3.5 mm, abaxially pubescent, pinnatelyveined, apex obtuse; bracteoles linear-oblong, ca. 6 × 1 mm,abaxially pubescent, 1-veined, apex acute. Pedicel short or absent.Calyx ca. 1 cm, outside pubescent; lobes linear, 8–9 mm,unequal, margin ciliate. Corolla purple, ca. 9 mm, outside pubescent,inside pubescent at base; tube basally cylindric for ca.2 mm; lobes suborbicular, 1.7–2 mm in diam., slightly unequal,apex rounded. Stamens with longer pair ca. 5 mm and shorterpair ca. 3 mm; filaments flattened, ciliate; anther thecae ca. 1mm, base appendaged; staminode ca. 1 mm. Ovary oblong, glabrous;ovules ca. 24 in 2 rows per locule; style ca. 3 mm; stigma2-lobed, lobes unequal with posterior one 2-dentate. Capsuleovoid-oblong, ca. 6 mm. Seeds orbicular, ca. 0.2 mm indiam. Fl. Dec.● Wet places, paddy fields. Hainan (Baoting).8. Staurogyne rivularis Merrill, Phillipp. J. Sci., C, 7: 248.1912.瘦叉柱花 shou cha zhu huaHerbs to 1 m tall, usually ascending from a decumbentbase and rooting at nodes on lower portion. Stems striate andlenticellate, glabrous. Petiole 3–6 cm; leaf blade elliptic, oblong,or lanceolate, 13–17 × 4.5–7 cm, both surfaces glabrous,secondary veins 8–11 on each side of midvein, base cuneateand slightly decurrent onto petiole, margin entire or undulate,apex acuminate to acute. Racemes terminal or axillary fromapical leaves, to 15 cm, unbranched or with 2 branches at base;rachis gland-tipped pubescent; bracts linear, 4–4.5 × ca. 0.8mm, pinnately veined, apex acuminate; bracteoles linear, ca. 2× 0.5 mm. Pedicel 2–3.5 mm. Calyx ca. 5 mm, outside pubescentwith gland-tipped trichomes. Corolla white, funnelshaped,ca. 7 mm; lobes suborbicular, ca. 1.5 × 1.3 mm, subequal,apex rounded. Stamens with longer pair ca. 3 mm andshorter pair ca. 2 mm; filaments glabrous; anther thecae ca. 1mm. Ovary ellipsoid, ca. 2 × 1 mm, glabrous; ovules 12 in 2rows per locule; style ca. 3 cm, glabrous; stigma 2-lobed. Capsulecylindric, 5–7 mm. Fl. Dec–May.Forests along streams. Hainan, Yunnan [Vietnam].9. Staurogyne brachystachya Benoist, Bull. Mus. Natl. Hist.Nat., sér. 2, 5: 174. 1933.短穗叉柱花 duan sui cha zhu huaHerbs to 80 cm tall. Stems subterete, sulcate, brown pubescent.Petiole 0.5–2 cm, brown pubescent; leaf blade lanceolateto ovate-lanceolate, 3–8 × 1–1.6 cm, both surfaces glabrous,abaxially pale green but brown when dry, adaxially green,secondary veins 8–10 on each side of midvein, base narrowlycuneate, margin irregularly undulate, apex acute. Racemes axil-

ACANTHACEAE 375lary or terminal, 2–4 cm, 3–5-flowered; peduncle and rachisglabrous; bracts linear, 2.3–5 mm, thickened, 1-veined, glabrous;bracteoles linear, 2.3–3 mm, glabrous, 1-veined. Pedicel1.2–1.6 cm, glabrous. Calyx 0.9–1.1 cm, glabrous except marginslightly ciliate with gland-tipped trichomes; lobes linear,7.5–9 × ca. 1 mm, subequal, apex acuminate. Corolla white, 1–1.4 cm, glabrous; lobes orbicular, ca. 2 mm in diam. Stamenswith longer pair 7–9 mm and shorter pair 6–8 mm; filamentspubescent (especially apically); anther thecae with a long tieshapedappendage, apex hirsute; staminode small. Ovary glabrous;style ca. 1 cm, glabrous; stigma 2-lobed. Capsule notseen. Fl. Mar–Apr.Forests; 800–1200 m. Guangxi, Yunnan [Vietnam].10. Staurogyne concinnula (Hance) Kuntze, Revis. Gen. Pl. 2:497. 1891.叉柱花 cha zhu huaEbermaiera concinnula Hance, J. Bot. 6: 300. 1868.Herbs to 15 cm tall, perennial. Stems generally short,partially subterranean, 0.5–6 cm, usually simple, rarely fewbranched, terete, densely shortly pubescent. Leaves in a basalrosette; petiole 0.3–3 cm, pubescent; leaf blade oblanceolate,1.5–7 × 0.5–1.5 cm, scarious, abaxially glaucous and pubescentalong veins, adaxially green and sometimes pubescent alongveins, secondary veins 5–8 on each side of midvein and abaxiallyprominent, base gradually attenuate, margin obscurelyundulate, apex obtuse to rounded. Racemes 1–4, axillary, sometimesloosely branched and forming a panicle, 5–15 cm;branches slender, with 3–10 interrupted flowers; bracts linear tolinear-oblanceolate, 2–4 mm, apex obtuse to subacute; bracteolesborne near middle of pedicel, linear, ca. 2 mm, apex obtuse.Pedicel 1–4 mm, pubescent. Calyx campanulate, 5-lobedalmost to base, glabrous except for margin minutely pilose;lobes linear to linear-lanceolate, 5–6 mm but accrescent to 7mm in fruit, unequal, apex acute. Corolla white, tubular-funnelform,ca. 1 cm; tube ampliate, ca. 6 mm, ca. 4 mm wide; lobesorbicular, ca. 3 mm, subequal. Stamens with longer pair ca. 5mm and shorter pair ca. 2.5 mm; filaments glabrous; antherthecae ca. 1.5 mm. Ovary ellipsoid, glabrous; style ca. 5 mm;stigma subulate. Capsule oblong, 4–5 × ca. 1.5 mm, glabrous,shorter than persistent calyx. Seeds suborbicular, ca. 0.2 mm.Fl. Mar–May, fr. Jul–Sep.Forests along streams. Fujian, Guangdong, Hainan, Taiwan[Japan].11. Staurogyne hainanensis C. Y. Wu & H. S. Lo, Fl. Hainan.3: 589. 1974.海南叉柱花 hai nan cha zhu huaHerbs to 30 cm tall, perennial. Stems terete, villous.Leaves cauline, crowded at apex of branches; petiole 1–3 cm,densely villous; leaf blade oblong to oblong-lanceolate, 3–10 ×1.3–3 cm, subleathery, abaxially adpressed villous, adaxiallyglabrous except midvein slightly villous, secondary veins 6–10on each side of midvein, base cuneate to broadly cuneate, marginsentire or irregularly undulate, apex rounded. Racemes terminalor axillary, to 16 cm; rachis pubescent; bracts linear-spatulateto linear, ca. 2.5 mm, 1-veined; bracteoles similar tobracts. Pedicel ca. 3 mm. Calyx 6–7 mm, subglabrous; lobeslinear, unequal, 3-veined, apex acuminate. Corolla purple, ca. 8mm; lobes ovate-oblong, 1.2–1.4 × ca. 1 mm, apex rounded.Stamens with longer pair ca. 5 mm and shorter pair ca. 3 mm;filaments glabrous; anther thecae of longer stamens unequaland of shorter stamens equal; staminode ca. 0.7 mm. Ovaryovoid, ca. 1 mm, glabrous; style ca. 6 mm. Capsule ovoid-oblong,ca. 5 mm, glabrous. Fl. Feb–May, fr. Jul–Aug.● Streamsides; 600–900 m. Hainan.12. Staurogyne strigosa C. Y. Wu & H. S. Lo, Fl. Hainan. 3:590. 1974.琼海叉柱花 qiong hai cha zhu huaHerbs, annual. Stems very short, villous. Leaves in a basalrosette; petiole 0.5–2.2 cm, strigose; leaf blade elliptic, oblong,ovate, or oblong-obovate, 1–1.5 × 0.8–2.5 cm, papery, abaxiallyvillous, adaxially strigose, secondary veins 5–7 on each side ofmidvein, base cuneate to rounded, margin entire, apex roundedto obtuse. Raceme terminal or axillary, to 6 cm, simple or rarelybranched; peduncle villous; bracts spatulate-linear, ca. 6 mm,abaxially villous, apex obtuse, rounded, or acute; bracteolessimilar to bracts. Pedicel 1–3 mm, villous. Calyx 6–8 mm,abaxially villous; lobes linear, 3-veined, apex acuminate. Corollaca. 5.5 mm; lobes orbicular, ca. 1 mm in diam. Staminal filamentsglabrous; anther thecae of longer stamens ca. 0.7 mm andof shorter stamens ca. 0.2 mm; staminode ca. 1 mm. Ovary glabrous;stigma 2-lobed. Capsule ovoid-oblong, ca. 6 mm, glabrous.Fl. Feb–Mar, fr. Jul–Aug.● Forests; below 200 m. Hainan (Qionghai).13. Staurogyne stenophylla Merrill & Chun, Sunyatsenia 2:322. 1935.狭叶叉柱花 xia ye cha zhu huaHerbs, perennial. Stems short, densely villous. Leaves usuallycrowded at apex of branches (but not forming a basalrosette); petiole ca. 1 cm, pubescent; leaf blade linear to lanceolate-linear,6–12 × 0.5–0.9 cm, papery, abaxially paler andvillous especially along veins, adaxially dark green, denselywhite papillose, and subglabrous except for margin slightly pubescent,secondary veins 8–13 on each side of midvein, baseattenuate, margin entire or undulate, apex obtuse. Racemes axillary,5–10 cm, unbranched or with 1 or 2 branches at base; peduncleand rachis villous; bracts linear, 3.8–4.5 mm, abaxiallypubescent, 1-veined; bracteoles linear, ca. 3.7 mm, abaxially pubescent.Pedicel 2–3.5 mm. Calyx ca. 6 mm, abaxially slightlyvillous; lobes linear-triangular, 4.5–5.1 × 0.5–0.6 mm, unequal,margin ciliate. Corolla pink, ca. 8 mm; tube basally cylindricfor ca. 1.5 mm then gradually widened, convex at one side;lobes orbicular to suborbicular, 2.2–2.5 × ca. 2 mm, unequal.Stamens 4 with longer pair ca. 4.5 mm and shorter pair ca. 3mm; filaments glabrous; staminode ca. 0.7 mm. Ovary ellipsoid,ca. 2 × 1 mm, glabrous; style unequally 3-lobed. Capsuleovoid-oblong, ca. 3 mm. Fl. Mar–May, fr. Aug–Sep.● Dense forests; 1000–2000 m. Hainan.

376ACANTHACEAE14. Staurogyne yunnanensis H. S. Lo, Bull. Bot. Res., Harbin8(1): 3. 1988.云南叉柱花 yun nan cha zhu huaHerbs to 0.7 m tall. Stems densely pubescent. Leaves cauline;petiole 1.5–2 cm; leaf blade ovate-lanceolate, ellipticlanceolate,or lanceolate, 4.5–10.5 × 2–3.5 cm, abaxially villousalong veins, adaxially glabrous, secondary veins 9–14 on eachside of midvein, abaxially conspicuous, and adaxially inconspicuous,base cuneate, margin undulate or inconspicuouslydentate, apex obtuse. Racemes terminal only, to 8–15 cm infruit; peduncle and rachis densely hirsute; bracts linear, 2.5–3 ×ca. 0.5 mm, adaxially densely pubescent, 1-veined, apex acuminateto acute; bracteoles linear, ca. 1.2 mm, glabrous, apex acuminate.Pedicel 2–3 mm in fruit. Calyx ca. 4 mm, glabrous;lobes extremely unequal, anterior one triangular, 2.5–3 × ca. 0.5mm, and apex acuminate, posterior and lateral ones subtriangularand ca. 1 mm. Corolla not seen. Capsule ± oblong, 3.5–4mm. Seeds light brown. Fr. Jul.● Forests. Yunnan (Hekou).15. Staurogyne petelotii Benoist, Arch. Bot. Bull. Mens. 4: 75.1930.中越叉柱花 zhong yue cha zhu huaHerbs 10–30 cm tall, ascending from a decumbent base.Stems erect, densely pubescent, unbranched or rarely branchedin apical part, base with 1.5–3 cm internodes. Leaves cauline;petiole 2–4 cm, pubescent; leaf blade ovate to oblong, 3–6 × 2–3.5 cm, membranous, abaxially glabrous, adaxially darker andsparsely pubescent, secondary veins 4–6 on each side of midvein,base broadly cuneate to rounded, margin entire, apex obtuse.Raceme terminal, ca. 5 cm; bracts obovate, ca. 8 × 4 mm,abaxially sparsely pubescent, base attenuate, margin distallyciliate, apex rounded; bracteoles oblong, ca. 6 × 1 mm, abaxiallypubescent especially apically, base gradually attenuate,apex rounded. Calyx ca. 6 mm; lobes spatulate with apexslightly enlarged and rounded, unequal, anterior one ca. 5 × 0.5mm and others ca. 3 × 0.2 mm. Corolla 1.5–1.7 cm, slightlycurved; lobes orbicular, ca. 5 × 5 mm. Staminal filaments glabrous.Ovary glabrous; style 1.8–2 mm, glabrous; stigma 2-cleft.Capsule ellipsoid, ca. 7 mm, glabrous. Fl. Jan–Apr.Forests along ravines or on slopes; 1400–1700 m. Yunnan [Vietnam].16. Staurogyne vicina Benoist, Bull. Mus. Natl. Hist. Nat., sér.2, 5: 171. 1933.密花叉柱花 mi hua cha zhu huaHerbs to 50 cm tall. Stems erect, 4-angled, sulcate, tuberculateon basal part, pubescent on apical part. Leaves cauline;petiole 2–4 cm, pubescent; leaf blade elliptic, 6–15 × 3–7 cm,both surfaces glabrous except for pubescence along veins, abaxiallyglaucous, adaxially green, secondary veins 10–12 on eachside of midvein, base acute, margin entire or serrate, apex acuteto shortly acuminate. Racemes terminal only, unbranched orsometimes branched, 10–16 cm; peduncle 2–5 cm, glabrous;bracts linear, 4–7 × ca. 1 mm, glabrous, apex acute; bracteoleslanceolate, 2–3 × ca. 1 mm, glabrous, apex acute. Pedicel 2–3mm, glabrous. Calyx 5–7 mm, glabrous; lobes linear, unequal,apex obtuse. Corolla white, 1.8–2.2 cm, outside glabrous; lobesoblong, ca. 4 × 3 mm, subequal, apex rounded. Staminal filamentsglabrous; anther thecae spherical, ca. 1 mm in diam.Ovary glabrous; style glabrous; stigma 2-cleft. Capsule oblong,ca. 8 mm, glabrous. Fl. Apr.Yunnan [Vietnam].17. Staurogyne debilis (T. Anderson) C. B. Clarke ex Merrill,Phillipp. J. Sci., C, 2: 302. 1907.菲律宾叉柱花 fei lü bin cha zhu huaEbermaiera debilis T. Anderson, J. Linn. Soc., Bot. 9: 452.1866.Herbs 10–15 cm tall, somewhat succulent. Stems terete,ascending, unbranched or rarely branched, pubescent. Leavescauline; petiole 5–8 mm, pubescent; leaf blade elliptic, oblong,or oblong-ovate, 2.5–4 × 0.8–1.7 cm, abaxially pale green andglabrous except pubescent along veins, adaxially green and glabrous,secondary veins 6 or 7 on each side of midvein, base cuneateto rounded, margin undulate, apex obtuse. Racemes terminalor axillary, to 7 cm, 4–7-flowered; rachis sometimesbranched at base, slightly pubescent; bracts linear, ca. 2 × 0.4mm; bracteoles subulate, ca. 1 × 0.3 mm. Pedicel ca. 2 mm. Calyx5–6 mm; lobes linear, subequal, margin sparsely hirsute.Corolla white, ca. 1 cm, glabrous; lobes oblong, ca. 2 × 2 mm,subequal. Stamens with longer pair ca. 3 mm and shorter pairca. 2 mm; filaments strigose; anther thecae ellipsoid, ca. 1 ×0.7 mm. Ovary ca. 1.8 mm, glabrous; style ca. 8 mm, glabrous;stigma 2-cleft. Capsule oblong, ca. 6 mm, compressed, glabrous.Fl. Feb–Mar, fr. Aug.Evergreen broad-leaved forests; 300–400 m. Taiwan [Philippines].3. OPHIORRHIZIPHYLLON Kurz, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 40: 76. 1871.蛇根叶属 she gen ye shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielPhyllophiorhiza Kuntze.Herbs, erect, without cystoliths. Leaves opposite, petiolate; leaf blade pinnately veined, margin entire. Inflorescence terminalspikes to racemes, unbranched or branched at base; bracts subopposite to alternate; bracteoles present. Calyx subequally 5-lobed. Corollatube expanded apically, annular hairy at throat or glabrous; limb 2-lipped; lower lip 3-lobed; upper lip 2-lobed; lobes descendingcochlear in bud. Stamens 2, exserted; anthers 2-thecous; thecae ellipsoid to oblong, divergent toward apex; staminodes 2. Ovaryovoid-spherical; ovules 13–18 per locule, arranged in 2 rows per locule; style exserted; stigma 2-lobed. Capsule ovoid, with numerousseeds; retinacula absent. Seeds subglobose.

ACANTHACEAE 377Two species: China, Laos, Myanmar, Thailand, Vietnam; one species in China.Other species previously treated in this genus pertain to other genera. Ophiorrhiziphyllon hypoleucum (Benoist) Benoist = Staurogyne hypoleucaBenoist and O. laxum Lindau = Justicia sp., according to Hossain (A. B. M. E. Hossain, Studies in the Classification and Affinities of Acanthaceae(Ph.D. Dissertation), University of Edinburgh, United Kingdom. 1971).1. Ophiorrhiziphyllon macrobotryum Kurz, J. Asiat. Soc.Bengal, Pt. 2, Nat. Hist. 40: 76. 1871.蛇根叶 she gen yeOphiorrhiziphyllon poilanei Benoist.Herbs to 1 m tall, erect. Young branches terete to 4-angled,brown pubescent. Petiole 3–8 cm, sulcate, pubescent; leaf bladeoblong-ovate, oblong-elliptic, or lanceolate, (8–)15–17 × (2–)5–7 cm, abaxially pale green and pubescent along veins, adaxiallygreen and glabrous, secondary veins 7–10 on each side of midvein,base cuneate to subrounded and sometimes decurrent ontopetiole, margin entire, apex acute. Rachises, peduncles, pedicels,bracts, and bracteoles brown pubescent with gland-tippedtrichomes. Spikes or racemes terminal, unbranched or branchedat base; peduncle 0–2 mm; bracts subulate to lanceolate, ca. 4 ×1 mm; bracteoles linear, ca. 1 mm. Pedicel 0–1 mm. Calyx ca. 5mm, outside sparsely pubescent with gland-tipped trichomes;lobes triangular, ca. 2 mm, margin ciliate with gland-tipped trichomes.Corolla yellowish white, ca. 7 mm; lobes suborbicular,ca. 2 × 2 mm. Staminal filaments ca. 8 mm, glabrous; staminodesincluded. Ovary glabrous; style ca. 8 mm, glabrous. Capsule3.5–5.5 mm. Fl. Oct–Feb, fr. Mar.Dense forests, along streams; 100–1300 m. S Yunnan [Laos,Myanmar, Thailand, Vietnam].4. THUNBERGIA Retzius, Physiogr. Sälsk. Handl. 1(3): 163. 1780, nom. cons.,not Montin (1773).山牵牛属 shan qian niu shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielVines [or shrubs], lacking cystoliths. Leaves petiolate; leaf blade margin entire to lobed to dentate. Flowers solitary or paired inleaf axils, or in terminal dichasiate thyrses, pedunculate; bracteoles leaflike, paired, enclosing calyx and most or all of corolla tube,distinct or fused along one side, persistent. Calyx much shorter than bracteoles, cupular with 10–20 subulate lobes or reduced to anentire ring. Corolla funnel-shaped, large; tube with short cylindric base then widened; limb ± equally 5-lobed. Stamens 4, oftendidynamous, inserted near base of corolla tube, included; anthers 2-thecous; thecae oblong or ovoid, parallel, spurred or not at base,sometimes bearded. Disk shortly annular or pulvinate. Ovary fleshy; ovules 2 per locule, collateral; style glabrous or pubescent;stigma 2-cleft, entire, or fringed. Capsule basally subglobose, apically prominently beaked, 2–4-seeded; retinacula absent. Seedscompressed to spherical, lacking trichomes.More than 100 species: Old World tropical regions; six species (one introduced) in China.Some species of Thunbergia are naturalized in the New World tropics.Thunbergia erecta (Bentham) T. Anderson and T. laurifolia Lindley, both cultivated in gardens, are not treated here.1a. Calyx reduced to an entire ring.2a. Leaves glabrous ................................................................................................................................................... 3. T. eberhardtii2b. Leaves pubescent.3a. Corolla bluish; capsule pubescent ................................................................................................................. 1. T. grandiflora3b. Corolla red; capsule glabrous ............................................................................................................................ 2. T. coccinea1b. Calyx bearing 10–20 slender subulate lobes.4a. Leaves pinnately veined ................................................................................................................................................ 4. T. lutea4b. Leaves palmately 3–5-veined.5a. Petiole conspicuously winged; corolla orange with throat dark purple internally ................................................. 5. T. alata5b. Petiole wingless; corolla entirely white ............................................................................................................ 6. T. fragrans1. Thunbergia grandiflora Roxburgh, Bot. Reg. 6: 495. 1820.山牵牛 shan qian niuThunbergia adenophora W. W. Smith; T. chinensis Merrill;T. lacei Gamble.Vines to 10 m or longer, woody. Stems 4-angled, sulcate,pubescent. Petiole 1–7 cm, grooved, pubescent; leaf blade ovateto triangular-ovate, 5–10 × 4–8 cm, papery, both surfaces pubescent,palmately 3–7-veined, base subcordate to truncate, marginundulate, irregularly angular on basal half, or rarely entire,apex acuminate to acute. Flowers solitary, paired in leaf axils,or arranged in terminal racemes with 2–4 flowers per node;peduncle 4–7 cm, sulcate, pubescent; rachis pubescent withlarge cyathiform glands; apical inflorescence bracts subulate tolinear-subulate, 2–6 × 1–1.5 mm, pubescent; bracteoles oblongto ovate, 2.5–4 × 1.5–2.2 cm, both surfaces pubescent, 5–7-veined, base truncate, margin entire or ciliate, apex acute with ashort mucro. Calyx ca. 2 mm, annular, unlobed, densely pubescent.Corolla bluish with a yellowish throat, 4–6 cm, outside

378ACANTHACEAEglabrous; tube basally cylindric and ca. 3 mm wide for ca. 7mm then gradually widened to ca. 5 cm at throat; limb subactinomorphic;lobes ovate, ca. 3 × 2.5 cm. Staminal filaments7–9 mm; anther thecae pubescent, basally appendaged. Styleglabrous; stigma with 2 subequal lobes. Capsule 1.2–1.5 cm,pubescent, basal part 1.3–1.8 cm in diam., beak ca. 2.5 cm.Seeds ovate in outline, compressed, verrucose. Fl. Aug–Jan, fr.Nov–Mar. 2n = 56.Thickets; 400–1500 m. Fujian, Guangdong, Guangxi, Hainan,Yunnan [India, Myanmar, Thailand, Vietnam].Thunbergia grandiflora is naturalized in tropical regions worldwide.2. Thunbergia coccinea Wallich, Tent. Fl. Napal. 1: 48. 1826.红花山牵牛 hong hua shan qian niuHexacentris coccinea (Wallich) Nees.Vines, woody. Stems 9-angled, subglabrous, pubescent atnodes. Petiole 2–7 cm, grooved; leaf blade broadly ovate, ovate,or lanceolate, 8–15 × 3.5–11 cm, both surfaces pubescent,palmately 3–5-veined, base rounded to cordate, margin undulateor remotely toothed, apex acuminate. Racemes axillary orterminal, to 35 cm, pendulous; peduncle 3–4 cm, pubescent;bracts lanceolate, abaxially pubescent, adaxially glabrous; bracteolesoblong, 2.2–2.6 × 1.1–1.5 cm, apex acute. Calyx ca. 2mm, reduced to a minute rim. Corolla red; tube basally cylindricfor 5–6 mm, throat 1.5–1.6 cm; lobes suborbicular, ca. 7mm in diam. Staminal filaments 1.2–1.5 cm, glabrous but witha tuft of trichomes at base; anther thecae parallel, unequal withlonger one ca. 5.5 mm and shorter one ca. 4.5 mm, spurred atbase. Ovary glabrous; style glabrous, exserted; stigma 2-cleft.Capsule glabrous, basal part 1–1.2 × 1.5–2 cm, beak 1.5–2.3cm. Seeds compressed, ovate in outline, verrucose. Fl. Sep–Jan,fr. Jan–May. 2n = 20.Forested montane slopes; 800–1000 m. SE Xizang, Yunnan [Laos,Myanmar, Thailand].3. Thunbergia eberhardtii Benoist, Bull. Mus. Natl. Hist. Nat.27: 543. 1921.二色山牵牛 er se shan qian niuVines to 12 m, woody. Stems 4-angled, sulcate, pubescent,soon glabrescent, pilose at nodes. Petiole 3–4 cm; leaf bladebroadly ovate to ovate-lanceolate, ca. 10 × 5 cm, both surfacesglabrous, palmately 5–7-veined, base cordate, margin remotelydentate or sometimes entire, apex acuminate to acute. Racemesaxillary or terminal, erect; peduncle pubescent; bracts lanceolate,pubescent, 1–3-veined, margin dentate, apex acute; bracteolesovate-lanceolate, 1–1.4 × 0.8–1 mm, abaxially tomentose,apex acuminate. Calyx annular, unlobed. Corolla to 2 cm;tube yellowish brown; lobes ovate-elliptic, ca. 1.1 cm, subequal,lower lobes red, upper lobes yellow. Anther thecae glabrous,those of lower pair of stamens long spurred at base, thoseof upper pair of stamens spurred at base of only one theca peranther. Ovary pubescent. Capsule basal part 1–1.5 cm in diam.,beak ca. 1.6 cm. Seeds hemispherical, verrucose. Fl. Aug–Nov,fr. Jan–Apr.Dense forests; 300–800 m. Hainan [Vietnam].4. Thunbergia lutea T. Anderson, J. Linn. Soc., Bot. 9: 448.1866.羽脉山牵牛 yu mai shan qian niuThunbergia salwenensis W. W. Smith.Vines to 5 m or longer, climbing. Stems glabrous exceptfor a circle of trichomes on nodes. Petiole 5–7.5 cm, glabrous;leaf blade ovate to lanceolate-ovate, 4–13 × 2–6 cm, abaxiallysubglabrous, adaxially pubescent or setose, base roundedto broadly cuneate and decurrent onto petiole, margins obscurelytoothed to sometimes irregularly denticulate, apex acuminate.Flowers axillary, solitary; bracteoles elliptic, 2.1–2.4 ×1.5–1.7 cm, abaxially glabrous, adaxially gland-tipped hairy,palmately 7-veined, apex mucronate. Calyx minute, 10-toothed.Corolla pinkish or white; tube basally cylindric for ca. 7 mm,throat ca. 3.5 cm; lobes orbicular, ca. 1.2 cm in diam., equal.Staminal filaments glabrous, ca. 1 cm; anther thecae ca. 7 mm,barbate, spurred at base, spurs hairlike. Ovary glabrous; styleca. 2.5 cm, glabrous; stigma funnel-shaped, 2-cleft. Capsuleglabrous, basal part ca. 1 × 1.8 cm, beak ca. 1.8 cm. Seeds reniform,ca. 8 × 5 mm. Fl. Aug–Dec, fr. Mar–May.Forests or thickets; 1000–2500 m. SE Xizang, Yunnan [Bhutan,India, Myanmar].5. Thunbergia alata Bojer ex Sims, Bot. Mag. 52: t. 2591.1825.翼叶山牵牛 yi ye shan qian niuVines, herbaceous. Stems ± 4-angled to flattened, bisulcate,pubescent. Petiole 1.5–3 cm, winged, sparsely pubescent;leaf blade sagittate to deltoid ovate, 2–7.5 × 2–6 cm, abaxiallyhirsute, adaxially sparsely strigose, palmately 5-veined, basehastate to cordate, margin entire or undulate, apex acute. Flowersaxillary, solitary; peduncle 2.5–3 cm, sparsely strigose; bracteolesovate, 1.5–1.8 × 1–1.4 cm, abaxially hirsute, 5–7-veined,apex acute, acuminate, or obtuse. Calyx annular, unequally 10–13-lobed. Corolla orange with dark purple glandular “eye” inthroat, 2.5–4.5 cm; tube basally cylindric for 2–4 mm, throat 1–1.5 cm; lobes obovate, apically truncate. Staminal filaments ca.4 mm, glabrous; anther thecae 3.5–4 mm, unequal, pubescent atmargin and base. Ovary glabrous; style ca. 8 mm, glabrous;stigma funnel-shaped, unequally 2-lobed, lower lobe spreading,upper lobe erect. Capsule pubescent, basal part ca. 7 × 10 mm,2-seeded; beak ca. 1.4 cm, ca. 3 mm wide at base. Seeds reticulateon dorsal surface. Fl. Oct–Mar, fr. Feb–May. 2n = 18.Cultivated in gardens and naturalized along roads. Guangdong,Yunnan [native to Africa].Thunbergia alata is widely cultivated and naturalized in tropicalregions.6. Thunbergia fragrans Roxburgh, Pl. Coromandel 1: 47.1795.碗花草 wan hua caoFlemingia grandiflora Rottler; Thunbergia bodinieri H.Léveillé; T. fragrans subsp. hainanensis (C. Y. Wu & H. S. Lo)H. P. Tsui; T. hainanensis C. Y. Wu & H. S. Lo.

ACANTHACEAE 379Vines, herbaceous. Stems almost 4-angled to flattened, sulcate,hirsute. Petiole 0.5–4.5 cm, hirsute; leaf blade oblongovateto ovate to broadly ovate to oblong-lanceolate to lanceolate,3–14 × 1.8–7 cm, both surfaces puberulent to subglabrousexcept for trichomes along veins or rarely glabrous, palmately3–5-veined, base rounded to sometimes cuneate or cordate,margin entire, irregularly sinuate, or shallowly coarsely dentate,apex acute to acuminate. Flowers axillary, solitary; peduncle1.5–5.5 cm; bracteoles ovate, 1.5–2.5 × 0.8–1.5 cm, puberulent,apex acute. Calyx 3–5 mm, unequally 10–17-dentate, glabrous.Corolla white, 3–5 cm, sparsely puberulent; tube basally cylindricfor 4–7 mm, throat 1.8–2.3 cm; lobes obovate, 1.3–2.5 ×1.5–2.3 cm. Stamens included; filaments 6–10 mm, glabrous;anther thecae ca. 3 mm, divergent, glabrous. Ovary glabrous;style 1.5–2 cm, exserted; stigma funnel-shaped, ca. 2 mm. Capsuleglabrous, basal part ca. 7 × 10–13 mm, beak 1.5–1.9 cm.Seeds 4–5 mm in diam., smooth or with scales. Fl. Aug–Jan, fr.Nov–Mar.Thickets, roadsides; 800–2300 m. Guangdong, Guangxi, Guizhou,Hainan, Sichuan, Taiwan, Yunnan [Cambodia, India, Indonesia, Laos,Philippines, Sri Lanka, Thailand, Vietnam].Variation in the shape, size, pubescence, and margin form of theleaves is extensive in Thunbergia fragrans, and taxa have been recognizedbased on these characters. Additional studies of the species fromthroughout its range are needed to determine whether these should beaccorded formal taxonomic status.5. ACANTHUS Linnaeus, Sp. Pl. 2: 639. 1753.老鼠簕属 lao shu le shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielAcanthodus Rafinesque.Shrubs or perennial herbs, erect or decumbent, without cystoliths. Leaves opposite [whorled or in a basal cluster], petiolate; leafblade margin entire, pinnatifid, or spinously toothed. Inflorescences in terminal [or axillary], uninterrupted spikes; bracts imbricate,ovate, growing in size as plant matures, margin spiny; bracteoles 2 or sometimes absent. Calyx 4-lobed, posterior and anterior lobeslarger, lateral pair of lobes smaller. Corolla tube very short, horny; limb of a single (lower) lip (upper lip sometimes rudimentary); lipelongate, obtusely 3-lobed. Stamens 4, paired, inserted at top of corolla tube, exserted; filaments stout, thickened, bony; anthers 1-thecous; theca linear-oblong, usually densely bearded, lacking basal appendages; staminodes absent. Ovary with 2 ovules per locule;style slender; stigma 2-cleft. Capsule not stipitate, ellipsoid, shiny, 4-seeded; retinacula present. Seeds ± compressed.About 20–25 species: tropical and subtropical regions of Old World; three species in China.Acanthus mollis Linnaeus is cultivated in Chinese gardens so is not treated here.1a. Bracteoles absent ....................................................................................................................................................... 1. A. ebracteatus1b. Bracteoles 2.2a. Bracts broadly ovate, margin entire ........................................................................................................................ 2. A. ilicifolius2b. Bracts ovate to obovate, margin apically spiny .............................................................................................. 3. A. leucostachyus1. Acanthus ebracteatus Vahl, Symb. Bot. 2: 75. 1791.小花老鼠勒 xiao hua lao shu leAcanthus ilicifolius Linnaeus var. ebracteatus (Vahl)Benoist; Dilivaria ebracteata (Vahl) Persoon.Shrubs 1–3 m tall. Stems terete, stout, glabrous, often withstipulelike spines at nodes. Petiole 1–4 cm, glabrous; leaf bladeoblong to obovate-oblong, 5–12 × 3–5 cm, rigid, subleathery,glabrous, secondary veins 3 or 4 on each side of midvein andspreading to spinose teeth, base cuneate, margin 3- or 4-toothedor irregularly pinnatifid, apex truncate to slightly mucronateand obtuse. Inflorescences terminal spikes to 4 cm; bractsbroadly ovate, 6–7 × 4–5 mm; bracteoles absent. Calyx posteriorand anterior lobes broadly ovate and 8–12 × 5–9 mm, laterallobes elliptic and ca. 10 × 4 mm. Corolla white [blue or purplish],ca. 2.5 cm; tube basally cylindric and ca. 2.5 mm widefor 5–8 mm; lip oblong, ca. 2.2 cm; lobes oblong, outside glabrous,inside appressed pubescent. Stamens subequal; filamentsca. 9 mm; anther theca oblong, ca. 8 mm, with a line of trichomeson both sides of suture. Capsule ca. 1.8 cm. Seeds yellow,ca. 10 × 7 mm, rugose. Fl. Apr–May, fr. Aug–Sep. 2n = 44.Mangrove swamps; near sea level. Guangdong, Hainan [Cambodia,India, Indonesia, Myanmar, Papua New Guinea, Thailand, Vietnam;Australia, Pacific islands].2. Acanthus ilicifolius Linnaeus, Sp. Pl. 2: 639. 1753.老鼠簕 lao shu leAcanthus ebracteatus Vahl var. xiamenensis (R. T. Zhang)C. Y. Wu & C. C. Hu; A. ilicifolius var. xiamenensis (R. T.Zhang) Y. F. Deng, N. H. Xia & Heng B. Chen; A. xiamenensisR. T. Zhang; Dilivaria ilicifolia (Linnaeus) Jussieu.Shrubs to 2 m tall, erect. Stems stout, scarcely branched,glabrous, often with stipulelike spines at nodes. Petiole 3–6mm, thick, glabrous; leaf blade oblong to oblong-lanceolate, 6–14 × 2–5 cm, glabrous, secondary veins 5–7 on each side ofmidvein, base cuneate, margin pinnatifid and with few largespines, serrate, or subentire, apex truncate. Spikes terminal, to16.5 cm; bracts broadly ovate, 7–8 mm, caducous; bracteolesovate, ca. 5 mm, leathery. Calyx posterior and anterior lobes 1–1.3 cm, margin scarious and sometimes crispate, apex emarginate;lateral lobes ovate, ca. 1 cm, margin entire. Corolla

380ACANTHACEAEwhite [bluish to pinkish], 3–4 cm; tube ca. 6 mm; lip obovate,2.2–3 cm, thinly leathery, outside with soft trichomes, apex 3-cleft. Stamens equal; filaments ca. 1.5 cm; anther theca with aline of trichomes on both sides of suture. Style ca. 2.2 cm. Capsule2.5–3 cm. Seeds light yellow, reniform. Fl. Feb–Mar, fr.Aug–Sep. 2n = 44, 48.Mangrove swamps, marshes; near sea level. Fujian, Guangdong,Guangxi, Hainan [Cambodia, India, Indonesia, Malaysia, Myanmar,Papua New Guinea, Philippines, Sri Lanka, Thailand, Vietnam; Australia,Pacific islands].This species is used medicinally.3. Acanthus leucostachyus Wallich ex Nees, Pl. Asiat. Rar. 3:98. 1832.刺苞老鼠簕 ci bao lao shu leHerbs to 1 m tall, erect, unbranched. Stem brown, ca. 4mm in diam., hairy. Petiole 0.8–2.5 cm; leaf blade elliptic,ovate-lanceolate, oblanceolate, or lanceolate, 9–23 × 2–4.5 cm,abaxially glabrescent or brown pubescent along veins, adaxiallyglabrous, secondary veins 5–8 on each side of midvein andnetted near margin, tertiary veins inconspicuous, base roundedto subcordate, margin slightly spinose-dentate but sometimesinconspicuously dentate or occasionally entire, apex acute.Inflorescences terminal spikes, to 16 cm, 4-ranked; rachisgrayish pubescent; bracts ovate to obovate, 1–1.5 × 0.5–0.8 cm,abaxially pubescent, palmately veined, base rounded, marginsapically spiny, apex acute to acuminate; bracteoles lanceolate tooblanceolate, 1.3–2 × 0.2–0.4 cm, abaxially pubescent, baserounded, margin entire and ciliate, apex rounded with a shorttip. Calyx ca. 2 cm, pubescent; lobes unequal, margin ciliate,anterior lobe ca. 1.7 × 0.4 cm, lateral lobes lanceolate and ca.15 × 2.5 mm, posterior lobe oblong-lanceolate and ca. 2 × 0.5cm. Corolla white, ca. 2.1 cm, villous; lobes orbicular, ca. 5 × 5mm, apex emarginate. Staminal filaments ca. 8 mm, glabrous,upper pair curved (S-shaped); anther theca ca. 4.5 mm, pilose.Ovary ca. 3 mm; style ca. 1.1 cm. Capsule ca. 10 × 3 mm, glabrous.Seeds reniform, ca. 2 × 2.5 mm, glabrous. Fl. Feb–May,fr. Aug–Sep.Dense forests, wet places; 600–1200 m. Yunnan [India, Laos,Myanmar, Thailand, Vietnam].6. BLEPHARIS Jussieu, Gen. Pl. 103. 1789.百簕花属 bai le hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, perennial [annual herbs or shrubs], without cystoliths, anisophyllous. Leaves usually in pseudo-whorls of 4 [or opposite],sessile to subsessile; leaf blade margin entire to dentate. Flowers axillary and solitary or many in terminal dense headlike spikes;bracts often clustered in up to 5 pairs, imbricate, leathery, margin often bristly dentate with bristles barbed; bracteoles absent [or 2].Calyx included in bracts or ± exserted, 4-lobed; lobes in 2 unequal opposite pairs, thickened-horny at base, posterior and anteriorlobes larger and 2–7-veined, lateral pair smaller and 1-veined. Corolla tube shorter than limb, expanded from base and constrictednear apex; limb a single (lower) lip; lip elongate, 3[–5]-lobed. Stamens 4, didynamous, inserted at or near apex of corolla tube; filamentsbony, with lower pair straight and upper pair curved, broad, flattened, usually bifurcate apically into a branch bearing an antherand a toothlike appendage; anthers 1-thecous; theca bearded. Ovary glabrous, with 2 ovules per locule; style linear; stigma 2-cleft.Capsule not stipitate, oblong to ellipsoid, 2(–4)-seeded; retinacula present. Seeds discoid, covered with long branched hygroscopictrichomes.About 130 species: Old World tropical and subtropical regions; one species in China.Blepharis was recently revised by K. Vollesen (Blepharis (Acanthaceae): A Taxonomic Revision, 1–342. 2000).1. Blepharis maderaspatensis (Linnaeus) B. Heyne ex Roth,Nov. Pl. Sp. 320. 1821.百勒花 bai le huaAcanthus maderaspatensis Linnaeus, Sp. Pl. 2: 639. 1753;Blepharis boerhaviifolia Persoon, nom. illeg. superfl.; B. boerhaviifoliavar. maderaspatensis (Linnaeus) Nees.Herbs to 30 cm tall, perennial. Stems trailing or scramblingto decumbent, sometimes elongate, scabrous or pubescent.Leaves usually 4 per pseudo-whorl of 2 unequal pairs;petiole ca. 2 mm, puberulent; leaf blade elliptic to obovatelanceolate,2.5–5 × 1–2 cm, puberulent, secondary veins 3–5 oneach side of midvein, base cuneate, margin undulate or dentate,apex acute. Flowers axillary and solitary or in clusters of fewfloweredspikes; bracts paired, broadly spatulate to obovateoblong,4–13 × 2–6 mm, unequal with inner pairs larger thanouter ones, strongly 3-veined, margin bristly ciliate. Calyx puberulent,unequally lobed; posterior and anterior lobes oblong,1.2–1.8 × 0.4–0.6 cm, 3-veined, apex of anterior lobe often 2-cleft; lateral lobes linear-lanceolate, 9–12 × 2–3 mm, marginciliate. Corolla whitish, often with pink or purple; lip obovate,1.6–1.7 × ca. 0.8 cm, spreading; lobes ovate-oblong. Staminalfilaments ca. 5 mm. Style ca. 1.2 cm. Capsule ellipsoid, ca. 8mm, glabrous. Seeds ca. 3 × 2 mm. Fl. Dec. 2n = 16, 22, 24, 26,28, 30.Limestone cliffs or rocks; ca. 800 m. Hainan [Bangladesh, Cambodia,India, Myanmar, Pakistan, Sri Lanka, Thailand, Vietnam; tropicaland subtropical Africa, SW Asia, Madagascar].

ACANTHACEAE 3817. STROBILANTHES Blume, Bijdr. 781, 796. 1826.马蓝属 ma lan shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); John R. I. WoodAdenacanthus Nees; Aechmanthera Nees; Apolepsis (Blume) Haasskarl; Baphicacanthus Bremekamp; Championella Bremekamp;Diflugossa Bremekamp; Goldfussia Nees; Gutzlaffia Hance; Hemigraphis Nees; Hymenochlaena Bremekamp; Lepidagathissect. Apolepsis Blume; Parachampionella Bremekamp; Paragutzlaffia H. P. Tsui; Perilepta Bremekamp; PseudaechmantheraBremekamp; Pseudostenosiphonium Lindau; Pseudostonium Kuntze; Pteracanthus (Nees) Bremekamp; Pteroptychia Bremekamp;Pyrrothrix Bremekamp; Semnostachya Bremekamp; Sericocalyx Bremekamp; Strobilanthes subg. Pteracanthus Nees; S. subg.Sympagis Nees; Sympagis (Nees) Bremekamp; Tarphochlamys Bremekamp; Tetraglochidium Bremekamp; Tetragoga Bremekamp.Herbs, subshrubs, shrubs, [or small trees], isophyllous or weakly to strongly anisophyllous, woody species commonly pliestesial(living for several years then dying after flowering and fruiting). Stems and branches usually 4-angled, often sulcate, basallybecoming woody and hollow with age. Leaves opposite, petiolate or sessile; leaf blade adaxially usually with prominent linear cystolithsand sometimes also abaxially, margin variously dentate, serrate, crenate, undulate, or entire. Inflorescences axillary and/or terminal,bracteate heads, headlike clusters, spikes (sometimes distinctly secund), or less commonly of pedicellate flowers forming anopen panicle; sterile bracts usually resembling reduced leaves and often present in compound inflorescences; floral bracts usuallydifferent from leaves, persistent or caducous as flowers open, very variable in size and shape, sometimes of two types with basalsterile bracts (outermost bracts in a capitate inflorescence) differing from inner or apical fertile ones; bracteoles 2 per pedicel, usuallysmall, sometimes absent. Calyx usually 5-lobed to base, commonly accrescent in fruit; lobes equal or with middle one distinctlylonger than others, sometimes partially fused to form a bipartite or tripartite calyx. Corolla nearly always bluish, rarely white, yellow,or pink, tubular or funnel-shaped, inside glabrous apart from trichomes retaining style except in Strobilanthes parvifolia, S. oresbia,and related species; tube either gradually widened from base or narrowly cylindric and then abruptly widened, campanulate or gibbous;limb 5-lobed; lobes usually ovate, equal or subequal, spreading, contorted in bud. Stamens usually 4 and didynamous (rarely 2,2 fertile with 2 staminodes, or 4 fertile with a central staminode), basally monadelphous; usually 2 filaments distinctly longer thanother 2; anthers included or exserted, 2-thecous; thecae oblong or subspherical, parallel, erect or incurved, glabrous, basally muticous,rarely with connective extended to a mucronate tip; pollen spherical or ellipsoid, echinulate and/or variously ribbed, usuallytricolporate. Ovary oblong to obovoid, 2-locular, with 2(–8) ovules per locule; style filiform, long, slender, simple, sometimes persistentafter corolla falls, retained in place by trichomes on one side of corolla tube; stigma 2-cleft with one branch longer. Capsulecharacteristically oblong to narrowly obovoid but sometimes fusiform to narrowly ellipsoid, (2–)4(–16)-seeded; retinacula strong,curved. Seeds usually ovate or orbicular in outline and lenticular by being flattened, usually pubescent with appressed mucilaginoustrichomes which become spreading when wetted, trichomes caducous in a very few species with seed glabrescent; areola usuallyvery small but occasionally extending as a glabrous area over much of seed surface.About 400 species: tropical Asia; 128 species (57 endemic) in China.Note on indumentum: in many species the young inflorescence, even when a few capsules are present, is glabrous or nearly so. As the inflorescencematures, sessile glands become stipitate and older inflorescences become gland-tipped pilose, a process parallel to the development of glandsand trichomes as the ovary matures into the capsule. The calyx is also accrescent in many species and the apex appears to continue growing, so calyxlobes, which are linear-oblong at anthesis, may become subspatulate in fruit. There is also a tendency for the corolla indumentum to fall as the corollaopens. Corolla indumentum is most easily observed on buds.Note on pollen: fig. 1 and fig. 2 show 18 pollen types found in Chinese Strobilanthes. It should be noted that the pollen of some 20% of Chinesespecies is unknown and the types shown here could be construed as an oversimplification. However, the images do show the range of variation foundin the genus. Two important caveats should be taken into account. Many publications illustrating Strobilanthes pollen from China are marred by theuse of misidentified material. This is the explanation for cases where a different pollen type is given to that in a published source. A second problemrelates to the degree of infraspecific variation in pollen morphology. This is largely unstudied but it is known that some of the more widespread andvariable species are somewhat diverse in their pollen morphology although all plants sampled have pollen of one general type. Species known to havesomewhat variable pollen include S. atropurpurea, S. echinata, S. speciosa, and S. tomentosa.Attempts to divide Strobilanthes (Strobilanthinae sensu Bremekamp) into segregate genera have proved unsatisfactory largely because the levelof homoplasy displayed by morphological characters has to date rendered subdivision impossible. Molecular studies have not yet greatly clarified thesituation because of insufficient sampling, and this too is likely to prove problematic because many species are known only from the type collection orfrom material inadequate for molecular study. Clusters of related species are clearly discernible, but insufficient information is currently available toproduce a satisfactory infrageneric classification. Species in the following account are grouped with morphologically similar and presumably relatedspecies, but the linear sequence and lack of adequate data implies that the order of species is at least partially arbitrary.Strobilanthes lactucifolia H. Léveillé is not treated in the present account because no material has been traced. It is probably not a species ofStrobilanthes. Strobilanthes straminea W. W. Smith (Goldfussia straminea (W. W. Smith) C. Y. Wu & C. C. Hu) from N Myanmar, included in FRPS(70: 169. 2002), is excluded from the present account because no Chinese specimen can be traced. Strobilanthes deutziifolia H. Léveillé is Abeliamacrotera (Graebner & Buchwald) Rehder in the Linnaeaceae (see p. 645). Strobilanthes gentiliana H. Léveillé is Sesamum indicum Linnaeus in thePedaliaceae (see Fl. China 18: 226. 1998). Strobilanthes hypericifolia H. Léveillé is Abelia uniflora R. Brown in the Linnaeaceae (see p. 645).

382ACANTHACEAEType 1. S. bantonensis from Balansa 4273. Type 2. S. anamitica from Maclure 8578. Type 3. S. hupehensis from Wilson 6627.Type 4. S. cusia from Kingdon Ward 21622. Type 5. S. pinetorum from Forrest 8385. Type 6. S. tomentosa from Polunin 3304.Type 7. S. reptans from Alston 16886. Type 8. S. cumingiana from Merrill 9510. Type 9. S. vallicola from Xuan Xujie 74.Figure 1. Strobilanthes pollen. SEM images of species with prolate (ellipsoid) pollen types.

ACANTHACEAE 383Type 10. S. tetrasperma from Li Zhiyu 1218. Type 11. S. labordei from Henry 7421. Type 12. S. truncata from Qin Dehai 79901.Type 13. S. dalzielii from Ying Hu-Shiu 12402. Type 14. S. affinis from Wilson 4302. Type 15. S. henryi from Hancock 57.Type 16. S. parvifolia from Kingdon Ward 8718. Type 17. S. cyclus from Maire 730. Type 18. S. serrata from Lace 6019.Figure 2. Strobilanthes pollen. SEM images of species with spheroidal (globose) pollen types.

384ACANTHACEAE1a. Low herbs, sometimes trailing, always isophyllous; corolla less than 2.5 cm and often less than 2 cm,not obviously curved.2a. Inflorescences of solitary flowers in leaf axils.3a. Calyx 2-lipped, upper 3 calyx lobes partially fused ......................................................................................... 43. S. rankanensis3b. Calyx not 2-lipped, subequally 5-lobed to base.4a. Leaf blade oblong to lanceolate, 7 mm wide or less; capsule glabrous ................................................................... 7. S. sinica4b. Leaf blade ovate to obovate, 8 mm wide or more; capsule apically pilose .................................................... 8. S. szechuanica2b. Inflorescences of terminal or sometimes axillary heads or spikes.5a. Corolla outside glabrous; capsule 6–16-seeded.6a. Leaves sessile; corolla yellow ............................................................................................................................. 4. S. fluviatilis6b. Leaves petiolate; corolla blue or white.7a. Leaf blade pubescent, base abruptly truncate to slightly cordate; corolla blue ................................................. 5. S. reptans7b. Leaf blade subglabrous, base cuneate to rounded; corolla white ................................................................ 6. S. cumingiana5b. Corolla outside pubescent; capsule 4-seeded.8a. Stems, leaves, and inflorescences white pilose ................................................................................................... 10. S. labordei8b. Stems, leaves, and inflorescences glabrous or very sparsely hirsute.9a. Corolla less than 1 cm ......................................................................................................................................... 9. S. wilsonii9b. Corolla 1.5–2 cm.10a. Leaf blade ovate to oblong-elliptic, 2–3 × as long as broad; ovary and capsule apicallypubescent .............................................................................................................................................. 11. S. tetrasperma10b. Leaf blade lanceolate, ca. 4 × as long as broad; ovary and capsule glabrous .......................................... 12. S. japonica1b. Herbs or subshrubs, never trailing, usually greater than 50 cm tall, anisophyllous or isophyllous; corollagreater than 2 cm, often strongly curved.11a. Inflorescences subcapitate or capitellate, with flowers densely clustered in groups of 4 or more, headsoccasionally becoming subspicate in fruit.12a. Anthers incurved but shorter ones sometimes reflexed, ± as broad as long so subspherical or ellipsoid;bracts and bracteoles if present caducous except in S. cuneata, S. kingdonii, S. glomerata, andsometimes S. capitata and S. speciosa.13a. Bracts greater than 1.5 cm, outermost commonly infertile, persistent, or if caducous then ovate.14a. Plants isophyllous; flowers in subsessile axillary clusters; leaf blade obovate .......................................... 100. S. cuneata14b. Plants anisophyllous; flowers in pedunculate heads; leaf blade ovate to oblong-elliptic.15a. Indumentum of somewhat rigid purplish trichomes ............................................................................. 101. S. glomerata15b. Indumentum of soft whitish trichomes.16a. Outermost bracts linear-oblong to lanceolate, margin often dentate, apex obtuse; isolatedflowers sometimes present below main head ..................................................................................... 104. S. kingdonii16b. Outermost bracts lanceolate or ovate, margin entire, apex acute or acuminate; inflorescencesstrictly capitate.17a. Bracts ovate, equaling or shorter than head; heads relatively stout, 1–3 cm broad; plantsweakly anisophyllous ........................................................................................................................ 102. S. capitata17b. Bracts lanceolate, commonly overtopping head; heads relatively slender, 1–1.5 cm broad;plants usually strongly anisophyllous .............................................................................................. 103. S. speciosa13b. Bracts less than 1 cm, all normally flower-bearing, often scarious and early caducous.18a. Corolla outside glabrous even on lobes and in bud.19a. Leaf blade abaxially whitish; bracts fugacious; calyx in fruit with lobes subequal ............................... 114. S. discolor19b. Leaf blade abaxially paler but not whitish; bracts usually present at anthesis; calyx in fruitwith one lobe longer than others.20a. Calyx without gland-tipped trichomes ....................................................................................... 107. S. ningmingensis20b. Calyx with gland-tipped trichomes.21a. Apex of bracts acute to acuminate ........................................................................................ 108. S. dimorphotricha21b. Apex of bracts rounded or obtuse.22a. Bracts glabrous; inflorescences strictly capitellate; plants weakly anisophyllous ........... 106. S. penstemonoides22b. Bracts gland-tipped pubescent; some flower pairs usually present below capitulum;plants strongly anisophyllous ....................................................................................................... 116. S. multidens18b. Corolla pubescent or gland-tipped pubescent at least on lobes or in bud.23a. Inflorescences subterminal, arising only from apical leaf axils; leaf blade abaxially whitish,falcate ............................................................................................................................................... 117. S. oligocephala23b. Inflorescences not subterminal; leaf blade if abaxially whitish then not falcate.24a. Bracts fugacious, usually absent at anthesis; calyx lobes subequal.25a. Plants isophyllous or weakly anisophyllous; leaves petiolate; leaf blade apex acuminate;corolla tube and lobes gland-tipped pubescent ........................................................................... 113. S. oxycalycina

ACANTHACEAE 38525b. Plants strongly anisophyllous; leaves sessile; leaf blade apex acute; corolla pilose onlyon lobes ........................................................................................................................................ 115. S. rhombifolia24b. Bracts usually present; calyx with one lobe longer than others.26a. Apex of bracts obtuse or rounded.27a. Peduncle longer so inflorescence exceeding subtending leaves ............................................... 110. S. formosana27b. Peduncle very short so inflorescence shorter than subtending leaves ................................. 111. S. ovatibracteata26b. Apex of bracts acute or acuminate.28a. Bracts gland-tipped pubescent .......................................................................................... 109. S. glandibracteata28b. Bracts not gland-tipped pubescent.29a. Plants anisophyllous ....................................................................................................... 108. S. dimorphotricha29b. Plants isophyllous .................................................................................................................. 112. S. hupehensis12b. Anthers erect, oblong, ca. 2 × as long as broad; bracts persistent at least until fruit is formed.30a. Flowers arranged in small clusters along an axis forming a spike or panicle of capitula.31a. Capsule 6–8-seeded; calyx lobes linear-lanceolate; anthers apiculate with an excurrent connective ........ 1. S. tomentosa31b. Capsule 4-seeded; calyx lobes ovate and abruptly narrowed to an apiculate apex; anthers muticous ....... 70. S. congesta30b. Flowers arranged in axillary or terminal capitula.32a. Capsule 6-seeded or more; calyx hirsute inside at least near apex.33a. Corolla blue, bracts obtuse to acute ........................................................................................................ 2. S. quadrifaria33b. Corolla yellow, bracts apically beaked ...................................................................................................... 3. S. chinensis32b. Capsule 2–4-seeded; calyx glabrous inside (except in S. cruciata, S. oligantha, and S. wangiana).34a. Calyx 2-lipped, upper 3 lobes partially fused.35a. Fertile stamens 2; leaf blade margin entire ................................................................................................ 38. S. aprica35b. Fertile stamens 4; leaf blade margin serrate.36a. Lower calyx lip with 2 lobes, lobes distinct to base; capsule 2-seeded ................................................ 30. S. cyclus36b. Lower calyx lip 2-dentate, lobes connate for most of length; capsule 4-seeded ............................... 95. S. bipartita34b. Calyx subequally 5-lobed to base.37a. Corolla completely glabrous even on lobes and tube and in bud.38a. Isophyllous herbs; corolla purple ................................................................................................... 16. S. sarcorrhiza38b. Anisophyllous subshrubs; corolla white or rarely blue.39a. Stems winged; leaves sessile; anthers muticous ................................................................................. 96. S. retusa39b. Stems unwinged; leaves petiolate; anthers apiculate.40a. Outer bracts sessile, ovate ............................................................................................................ 99. S. echinata40b. Outer bracts with a broad petiolar base, pandurate in outline.41a. Petiolar base of bracts forming a rigid involucre, lamina spreading and ovate ....................... 97. S. cruciata41b. Petiolar base not forming an involucre, bracts pandurate, apical portion narrowlylanceolate ................................................................................................................................... 98. S. simonsii37b. Corolla pubescent at least on lobes or tube or in bud.42a. Bracts oblanceolate, obovate, subrhomboidal, or spatulate, broadest near apex, pilosewith reddish or white trichomes.43a. Bracts with reddish or brownish trichomes.44a. Leaves and stem reddish brown setose ............................................................................................ 20. S. hossei44b. Leaves and stem without reddish brown trichomes.45a. Bracts subrhomboidal-obovate, tips erect; pollen type 3 ...................................................... 21. S. torrentium45b. Bracts obovate, tips commonly recurved; pollen type 2 ........................................................ 22. S. anamitica43b. Bracts with white trichomes.46a. Bracts distinct from leaves, obovate, with gland-tipped trichomes, apex rounded ................... 18. S. lihengiae46b. Bracts oblanceolate to obovate, with a few marginal teeth, densely white pilose,without gland-tipped trichomes, apically narrowed to an obtuse apex.47a. Corolla 2.2–2.8 cm; plants drying dark green; inflorescences often becomingspicate in fruit .................................................................................................................. 15. S. austrosinensis47b. Corolla 2.8–3.2 cm; plants not drying dark green; inflorescences not becomingspicate in fruit ...................................................................................................................... 17. S. procumbens42b. Bracts linear, lanceolate, or oblong-elliptic, widest below middle.48a. Bracts commonly yellowish, overtopping capitula and giving a characteristic earedappearance ...................................................................................................................................... 19. S. esquirolii48b. Bracts with white trichomes, not overtopping capitula which do not have an earedappearance.49a. Heads strictly terminal; calyx lobes with trichomes apically on inner surface.50a. Leaf blade abaxially glabrous except on veins; corolla 3.5–4 cm ......................................... 13. S. wangiana50b. Leaf blade usually pubescent or rarely glabrous; corolla less than 3.5 cm ............................ 14. S. oligantha

386ACANTHACEAE49b. Heads arising from apical leaf axils; calyx glabrous inside.51a. Leaf blade glabrous .................................................................................................................. 24. S. fengiana51b. Leaf blade pilose or pubescent on both surfaces.52a. Corolla straight or nearly so; heads clearly axillary; plants isophyllous ........................ 23. S. yunnanensis52b. Corolla strongly bent; heads from apical leaf axils only; plants anisophyllous.53a. Heads slightly elongate; floral bracts lanceolate to oblong, pilose but marginnot ciliate .............................................................................................................................. 25. S. lamium53b. Heads very compact; floral bracts elliptic, margin ciliate .............................................. 26. S. cyphantha11b. Inflorescences spicate or flowers in leaf axils or arranged singly or in opposite pairs in a panicle.54a. Leaf blade pinnatifid ........................................................................................................................................ 34. S. pinnatifida54b. Leaf blade margin dentate or serrate but never pinnatifid.55a. Bracts decurrent as a wing on peduncle ....................................................................................................... 48. S. pteroclada55b. Bracts not decurrent on peduncle.56a. Anthers incurved and shorter ones sometimes reflexed, subspherical or ellipsoid, ± as broad aslong; bracts and bracteoles if present caducous except in S. longgangensis and cultivated formsof S. hamiltoniana and sometimes in S. lamiifolia.57a. Inflorescences an open panicle; flowers not arranged in opposite pairs.58a. Corolla arcuate, less than 3 cm; inflorescence branches with non-glandular trichomes ..................... 126. S. euantha58b. Corolla straight, greater than 3 cm; inflorescence branches glabrous or with gland-tippedtrichomes.59a. Inflorescences branching at a narrower angle from axis, ultimate branches relatively stoutand similar to primary branches ................................................................................................ 124. S. hamiltoniana59b. Inflorescences branches at 90° from axis, ultimate branches very slender ................................... 125. S. tenuiflora57b. Inflorescences spicate; flowers in opposite pairs, sometimes aggregated into a panicle.60a. Inflorescences a terminal panicle of spikes.61a. Leaf blade ovate, base rounded to broadly cuneate; basal leaves shortly petiolate, apicalones sessile ....................................................................................................................................... 122. S. mastersii61b. Leaf blade narrowly oblong-elliptic, base narrowly cuneate; leaves all petiolate ............................... 123. S. tenax60b. Inflorescences terminal or axillary but spikes never paniculate.62a. Inflorescences terminal or from apical leaf axils only.63a. Inflorescences a short terminal spike; bracts mostly 1.5–2.5 cm, relatively persistent .............. 105. S. lamiifolia63b. Inflorescences of flower pairs scattered along axis but often crowded and subcapitellatenear apex; bracts mostly less than 1.5 cm, caducous.64a. Corolla glabrous; bracts gland-tipped pilose ............................................................................ 116. S. multidens64b. Corolla gland-tipped pubescent on lobes; bracts not gland-tipped pilose ......................... 117. S. oligocephala62b. Inflorescences axillary spikes.65a. Corolla outside pubescent.66a. Calyx less than 1.3 cm; bracts often present at least on young parts; leaf blade marginserrate ....................................................................................................................................... 119. S. thomsonii66b. Calyx 1.6–2 cm; bracts very caducous; leaf blade margin entire or sinuate ............................. 121. S. secunda65b. Corolla outside glabrous.67a. Leaf blade ovate, base rounded to cordate; anthers slightly exserted .................................. 118. S. medogensis67b. Leaf blade narrowly elliptic, base cuneate; anthers included .......................................... 120. S. longgangensis56b. Anthers erect, oblong, ca. 2 × as long as broad; bracts persistent at least until fruit is formed.68a. Corolla outside glabrous but pubescent inside; leaves subentire; inflorescences leafy or1–3-flowered axillary spikes.69a. Leaf blade abaxially whitish; inflorescences paniculate ................................................................... 127. S. parvifolia69b. Leaf blade abaxially reddish brown; inflorescences 1–3-flowered axillary spikes ......................... 128. S. ferruginea68b. Corolla pubescent, or if outside glabrous then also inside glabrous except for trichomesretaining style; inflorescences not as above.70a. Flowers in axillary spikes, with basal persistent expanded leaflike bracts; flowers not imbricate;spikes sometimes reduced so flowers are solitary in leaf axils, not aggregated into a terminalpanicle.71a. Calyx 2-lipped with upper three calyx lobes distinctly connate for half their length.72a. Corolla blue, not resupinate .......................................................................................................... 62. S. flexicaulis72b. Corolla white, resupinate ............................................................................................................. 63. S. lanyuensis71b. Calyx subequally 5-lobed to base.73a. Corolla outside glabrous.74a. Leaf blade large with some greater than 10 cm, drying blackish; corolla straight, 3.5–5 cm .......... 61. S. cusia74b. Leaf blade mostly less than 6 cm, not drying blackish; corolla bent at mouth, 2.5–3.5 cm.

ACANTHACEAE 38775a. Calyx lobes linear, 1–1.5 mm wide, apex acute or rarely obtuse ..................................... 64. S. atropurpurea75b. Calyx lobes ovate-elliptic, 3–5 mm wide, apex obtuse ......................................................... 65. S. latisepala73b. Corolla outside pubescent.76a. Rachis of inflorescences strongly winged.77a. Calyx less than 2 cm; capsule glabrous ..................................................................................... 66. S. tibetica77b. Calyx 2.5–3.6 cm; capsule comose .............................................................................. 67. S. pterygorrhachis76b. Rachis of inflorescences not winged.78a. Bracts margin entire, apex obtuse; plants subglabrous ........................................................... 68. S. urophylla78b. Bracts margin dentate, apex acuminate; plants usually densely hirsute or rarely(in S. inflata var. gongshanensis) subglabrous ............................................................................. 69. S. inflata70b. Flowers in axillary or terminal spikes but if axillary without leaflike bracts (except sometimesS. oresbia); flowers imbricate or not; spikes aggregated into a terminal panicle or not.79a. Flowers in axillary leafless spikes (sometimes from apicalmost axils); bracts distinct from leaves,often inconspicuous and merging into bracteoles and calyx lobes; spikes not aggregated into aleafless panicle except in S. nobilis and S. rostrata; corolla outside glabrous.80a. Calyx glabrous or with gland-tipped trichomes, margin never ciliate or ciliolate.81a. Inflorescences with flowers solitary on one side of spike; bracts linear, persistent.82a. Spikes strictly axillary; inflorescences glabrous ......................................................................... 54. S. helicta82b. Spikes aggregated into a terminal panicle; inflorescences with gland-tipped trichomes ......... 55. S. rostrata81b. Inflorescences with flowers paired and on two sides of spike; bracts ovate tooblong-elliptic, somewhat caducous.83a. Corolla bent; leaf blade abaxially whitish, margin entire to crenate ..................................... 52. S. rubescens83b. Corolla straight; leaf blade abaxially green, margin coarsely serrate.84a. Apex of bracts acute to obtuse; stems winged ................................................................ 41. S. pateriformis84b. Apex of bracts truncate; stems not winged ............................................................................ 42. S. truncata80b. Calyx lobes sometimes also with gland-tipped trichomes, margin ciliate or ciliolate.85a. Bracts with a recurved tip; inflorescences reddish brown .................................................... 92. S. brunnescens85b. Bracts with an erect tip; inflorescences greenish or rarely reddish brown.86a. Bracts ovate, obovate, or broadly elliptic, distinctly shorter than calyx.87a. Bracts abaxially with 2 raised eyelike structures ................................................................ 84. S. biocullata87b. Bracts abaxially without raised eyelike structures although sometimes with largesessile glands.88a. Spikes from apical axils; bracts obovate-spatulate ................................................... 80. S. pseudocollina88b. Spikes not restricted to apical axils; bracts ovate to elliptic.89a. Bracts not abruptly narrowed and without a mucronate apex.90a. Spikes commonly branched; flower pairs close along rachis; calyx 2-lippedwith 3 upper lobes partially fused ...................................................................... 81. S. longzhouensis90b. Spikes simple; flower pairs distant along rachis; calyx subequally 5-lobedto base .................................................................................................................. 82. S. cystolithigera89b. Bracts abruptly narrowed with a mucronate and sometimes caducous apex.91a. Plant drying olive-green; bracts abaxially with sessile glandular area, apexnot fimbriate, lacking reddish brown trichomes ...................................................... 83. S. abbreviata91b. Plant drying blackish green; bracts abaxially without glandular area, apexoften fimbriate with reddish brown trichomes .......................................................... 91. S. fimbriata86b. Bracts lanceolate, ovate, or oblong, usually equaling or exceeding calyx.92a. Leaf blade often pandurate, base rounded; apical leaves sessile; spikes elongatingbeyond leaves and inflorescence often of paniculate spikes in fruit ....................................... 93. S. nobilis92b. Leaf blade not pandurate, base cuneate; all leaves petiolate; inflorescence not paniculatein fruit.93a. Spikes only from terminal leaf axils.94a. Spikes 1–3 per axil; bracts abaxially with sessile black glands .................................. 89. S. heteroclita94b. Spikes 1 per axil; bracts abaxially lacking black glands ............................. 90. S. mucronatoproducta93b. Spikes not restricted to terminal axils.95a. Spikes 1–5 per leaf axil .............................................................................................. 94. S. polyneuros95b. Spike usually 1 per leaf axil.96a. Bracts 2–3 cm, much exceeding calyx ..................................................................... 88. S. compacta96b. Bracts less than 1.5 cm, ± equaling calyx.97a. Stems, leaves, and bracts pubescent ................................................................. 87. S. guangxiensis97b. Plant completely glabrous except for ciliolate margin of calyx lobes.

388ACANTHACEAE98a. Spikes often branched, robust; leaf blade oblong, some greater than15 cm .............................................................................................................. 85. S. longespicata98b. Spikes simple, slender; leaf blade elliptic, less than 12 cm ..................................... 86. S. myura79b. Inflorescences terminal or terminal and axillary, sometimes forming a panicle of spikes,branches bearing leaves below spikes, if apparently axillary then corolla outside pubescentor bracts expanded and clearly different from calyx.99a. Fertile stamens 2.100a. Spikes simple; capsule glabrous apart from a few apical glands ................................................. 29. S. henryi100b. Spikes branched forming a terminal panicle of spikes; capsule pubescent.101a. Bracts broadly oblong-ovate to oblong-obovate, not imbricate so flowers clearlyseparate ...................................................................................................................................... 31. S. serrata101b. Bracts obovate, imbricate ........................................................................................................ 32. S. tubiflos99b. Fertile stamens 4.102a. Bracts imbricate.103a. Bracts greater than 1.5 cm.104a. Bracts distinct from apical leaves, ca. 3 cm, margin without prominent white cilia.105a. Bracts not sticky glandular; capsule 12-seeded ............................................................ 2. S. quadrifaria105b. Bracts sticky glandular; capsule 4-seeded ....................................................................... 60. S. cognata104b. Bracts indistinct from apical leaves, less than 2 cm, margin with prominent whitecilia.106a. Bracts ovate to elliptic; corolla 3–3.5 cm ...................................................................... 14. S. oligantha106b. Bracts obovate; corolla 2.2–2.8 cm ....................................................................... 15. S. austrosinensis103b. Bracts less than 1.5 cm.107a. Leaves sessile; leaf blade oblong; spikes axillary as well as terminal ........................... 35. S. auriculata107b. Leaves petiolate; leaf blade ovate or elliptic; spikes all terminal on leafy branchesor main stem.108a. Anthers strongly exserted from corolla .............................................................................. 33. S. affinis108b. Anthers included in corolla.109a. Corolla outside glabrous.110a. Bracts linear-spatulate, broadest at apex .............................................................. 57. S. tonkinensis110b. Bracts oblong or elliptic, broadest in middle or base.111a. Bracts elliptic, 5–6 mm wide ................................................................................. 27. S. strigosa111b. Bracts oblong , 2–3 mm wide ............................................................................ 58. S. stolonifera109b. Corolla outside pubescent.112a. Leaves glabrous ...................................................................................................... 28. S. longiflora112b. Leaves pubescent.113a. Calyx 2-lipped, 3 upper lobes partially fused .................................................. 36. S. mogokensis113b. Calyx subequally 5-lobed to base ......................................................................... 37. S. argentea102b. Bracts not imbricate so flowers clearly separate apart from basalmost.114a. Inflorescences a 15–25 cm uninterrupted spike from apicalmost leaf axils ......................... 59. S. vallicola114b. Inflorescences a much shorter spike but if similar in length then basalmost flowerswidely spaced.115a. Corolla outside glabrous.116a. Calyx subequally 5-lobed to base.117a. Stems and leaves sparsely pubescent ............................................................................. 51. S. taoana117b. Stems and leaves completely glabrous.118a. Inflorescences branched; calyx glabrous; leaf blade broadly ovate tosuborbicular; floral bracts 7–11 mm ...................................................................... 53. S. pinetorum118b. Inflorescences of simple terminal spikes; calyx gland-tipped pilose; leafblade elliptic; floral bracts 5–6.5 mm ................................................................. 56. S. chrysodelta116b. Calyx 2-lipped, 3 upper lobes partially fused.119a. Bracts 2–3 cm, much longer than calyx ...................................................................... 46. S. adpressa119b. Bracts less than 1.4 cm, equaling or shorter than calyx.120a. Bracts ovate ................................................................................................................... 49. S. ovata120b. Bracts linear-lanceolate or oblong.121a. Spikes with 1–3 flower pairs; bracts greater than 10 mm ................................. 47. S. atroviridis121b. Spikes with 1–7 flower pairs; bracts less than 8 mm.122a. Bracts linear-lanceolate; rachis unwinged; plant drying blackish green ....... 44. S. longispica122b. Bracts oblong-ovate; rachis winged; plant not drying blackish green .............. 45. S. dryadum

ACANTHACEAE 389115b. Corolla outside pubescent at least in bud, on lobes, or tube.123a. Plants strongly anisophyllous; corolla densely white pubescent in bud; capsuleglabrous.124a. Bracts linear, attenuate; calyx lobes finely acute ......................................................... 39. S. dalzielii124b. Bracts oblanceolate; calyx lobes obtuse ................................................................. 40. S. bantonensis123b. Plants not strongly anisophyllous; corolla not densely white pubescent in bud;capsule gland-tipped pubescent at apex.125a. Leaves sessile.126a. Leaf blade ovate to suborbicular, ± as long as broad, abaxially whitish;bracts orbicular to obovate, 5–8 mm ................................................................... 50. S. tamburensis126b. Leaves narrowly ovate, ca. 3 × as long as broad, abaxially green; bractslanceolate, 7–10 mm .................................................................................................. 75. S. forrestii125b. Leaves petiolate.127a. Corolla ventricose and strongly bent just below mouth, pubescent over itswhole length.128a. Corolla inside pubescent especially on lobes.129a. Bracts oblong, margin entire or with a few apical teeth; spikes 2-sided ............. 71. S. larium129b. Bracts broadly to narrowly ovate to lanceolate, margin mostly serrate;spikes often secund .............................................................................................. 72. S. oresbia128b. Corolla inside glabrous even on lobes.130a. Bracts much reduced even at base of spike; leaf blade mostly greater than5 cm; petiole 2–5 cm ................................................................................................ 73. S. flexa130b. Bracts leaflike almost to spike apex; leaf blade mostly less than 5 cm;petiole (0–)1–3 cm ........................................................................................... 74. S. versicolor127b. Corolla gradually widened above base, curved but not very strongly so,pubescent on lips only.131a. Spikes extremely lax, with apical flower pairs commonly separated bygreater than 1 cm along rachis; bracts ovate; calyx lobes conspicuouslyspatulate in fruit ....................................................................................................... 78. S. extensa131b. Spikes apically relatively dense, with flower pairs becoming imbricatealong rachis; bracts linear to lanceolate; calyx lobes not usuallyconspicuously spatulate.132a. Inflorescences mostly of simple terminal spikes; leaf blade glabrous .......... 79. S. spiciformis132b. Inflorescences a panicle of spikes; leaf blade pubescent at least whenyoung.133a. Leaf blade ovate, obovate, or broadly elliptic, ca. 2 × as long as broad ... 76. S. lachenensis133b. Leaf blade narrowly oblong-elliptic, 3–4 × as long as broad ...................... 77. S. nemorosa1. Strobilanthes tomentosa (Nees) J. R. I. Wood, Kew Bull.64: 16. 2009.尖药花 jian yao huaAechmanthera tomentosa Nees in Wallich, Pl. Asiat. Rar.3: 87. 1832; A. gossypina (Wallich) Nees; A. tomentosa var.wallichii C. B. Clarke; A. wallichii Nees, nom. illeg. superfl.; A.wallichii var. gossypina (Wallich) Nees; A. wallichii var. tomentosa(Nees) Nees; Ruellia gossypina Wallich; Strobilanthesblinii H. Léveillé; S. bodinieri H. Léveillé; S. cavaleriei H.Léveillé.Subshrubs to 1 m tall, erect, isophyllous. Stems usuallygray tomentose but sometimes thinly pilose and greenish. Petiole1–3 cm, with yellow multicellular gland-tipped trichomes;leaf blade elliptic-ovate to elliptic-oblong, 3–13 × 2–6.5 cm,abaxially sericeous, adaxially densely covered with linear cystoliths,secondary veins 7–10 on each side of midvein, basesubrounded, cuneate, or cordate and sometimes shortly decurrentonto petiole, margin crenate, apex acute. Inflorescencesusually paniculate, to 30 cm, with 3–8-flowered clusters commonlyon 3–8 cm axillary branches in axils of leaflike sterilebracts, clusters sometimes aggregated to form axillary thyrsesor more distant and forming an open panicle; floral bracts linear-ligulate,0.8–1.2 cm, abaxially and marginally gland-tippedpubescent, apex obtuse; bracteoles narrowly ligulate, ca. 5 mm,similar to and smaller than or equaling calyx lobes. Calyx 0.8–1.2 cm, deeply 5-lobed to base; lobes linear, unequal with 2longer and 3 shorter, slightly gland-tipped pilose, apex obtuse.Corolla blue, funnel-shaped, 2.5–3.5 cm, straight, ventricose,outside glabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and narrow for 4–8 mm then graduallywidened to ca. 1 cm at mouth; lobes ca. 5 mm, subequal,apex obtuse. Stamens 4, included; shorter filament pair ca. 6mm, glabrous; longer filament pair 1.1–1.2 cm, pilose; antherthecae oblong, apex mucronate; connective minutely excurrent;pollen type 6. Ovary comose; style sparsely hairy. Capsule oblong,7–10 mm, pubescent, 6–8-seeded. Seeds discoid, denselyhairy, sometimes glabrescent. Fl. Jun–Dec, fr. Aug–Dec. 2n =30.Grassy mountain slopes, open forests in relatively dry areas; 500–2300 m. Guangxi, Guizhou, Yunnan [Bangladesh, Bhutan, India, Laos,Myanmar, Nepal, Pakistan].

390ACANTHACEAEThis is a distinctive and apparently isolated species on account ofthe excurrent anther connective combined with the 6–8-seeded capsuleand seeds with caducous trichomes.Strobilanthes tomentosa was incorrectly reported as S. glutinosaNees (Pseudaechmanthera glutinosa (Nees) Bremekamp) by Handel-Mazzetti (Symb. Sin. 7: 894. 1936).2. Strobilanthes quadrifaria (Wallich ex Nees) Y. F. Deng,Acta Phytotax. Sin. 45: 849. 2007.四列马蓝 si lie ma lanRuellia quadrifaria Wallich ex Nees in Wallich, Pl. Asiat.Rar. 3: 83. 1832; Hemigraphis quadrifaria (Wallich ex Nees) T.Anderson; Sericocalyx quadrifarius (Wallich ex Nees) Bremekamp.Subshrubs (0.2–)1–1.5 m tall, isophyllous. Stems hirsutewith large-celled trichomes. Petiole (0.3–)1.5–4 cm, hirsute;leaf blade elliptic to ovate, 5–18 × 2–7 cm, both surfaces scabrousand rather sparsely pilose with large pustulose white trichomesespecially on veins, adaxially darker and with cystoliths,base attenuate, margin crenate-dentate, apex acute. Inflorescencesaxillary or terminal, subcapitate to shortly spicate, 2–5 × ca. 2 cm, very compact, densely covered with long silkywhite large-celled trichomes; peduncle 0.5–1.5 cm, hirsute;bracts ovate, elliptic, or broadly lanceolate, 1.5–2.5 × 0.6–1cm, pilose with white large-celled trichomes, apically obtuse toacute; bracteoles linear-lanceolate, ca. 10 × 2 mm, pilose withwhite large-celled trichomes. Calyx 8–11 mm, outside pilose,inside sericeous toward apex, subequally 5-lobed; lobes linear,apex acuminate. Corolla pale purple with a whitish tube, 3–3.2cm, straight, subglabrous except for pubescent lobes; tube basallycylindric and narrow for 8–10 mm then gradually widenedto 8–9 mm at mouth; lobes ovate, ca. 4 × 4 mm, apex rounded.Stamens 4, included; shorter filament pair 2–3 mm, glabrous;longer filament pair 4–7 mm, pilose; anther thecae oblong, ca. 2× 0.8 mm; pollen type 6. Ovary oblong, comose; style 2–2.2cm, pilose. Capsule oblong, 1–1.2 cm, sparsely pubescent nearapex, 12-seeded. Seeds ca. 1 mm in diam., pubescent. Fl.Oct–Jan.Moist shaded places in forests; 500–900 m. Yunnan [Laos, Myanmar,Thailand].3. Strobilanthes chinensis (Nees) J. R. I. Wood & Y. F. Deng,Bot. J. Linn. Soc. 150: 388. 2006.黄球花 huang qiu huaRuellia chinensis Nees in A. Candolle, Prodr. 11: 147.1847; Hemigraphis chinensis (Nees) T. Anderson ex Hemsley;Sericocalyx chinensis (Nees) Bremekamp.Herbs or subshrubs, 30–50(–150) cm tall, isophyllous.Stems often procumbent and rooting at base, 4-angled whenyoung, hirsute, basally slightly woody. Petiole 4–10 mm; leafblade obovate, obovate-elliptic, or sometimes oblong-elliptic,1.5–11 × 1–4.5 cm, both surfaces sparsely hispid, adaxiallydensely covered with cystoliths, midvein impressed, secondaryveins 5 on each side of midvein, purplish, and scabrous, baseattenuate and decurrent onto petiole, margin crenate to slightlydentate, apex acute to shortly acuminate. Inflorescences axillaryor terminal, subcapitate; bracts imbricate, broadly to narrowlyovate, 1.5–2 cm, hirsute, usually 3-veined, apically beaked to asubobtuse point; bracteoles linear, ca. 9 mm, resembling calyxlobes. Calyx 5–11 mm, subequally 5-lobed to base; lobes linearlanceolate,ca. 10 × 0.5 mm, outside gland-tipped pilose, insideapically pubescent, apex attenuate. Corolla yellow, 1.6–2 cm,straight; tube basally cylindric and ca. 1 mm wide for ca. 5 mmthen gradually widened to ca. 1 cm at mouth, outside pubescent,inside villous; lobes orbicular, ca. 4 mm in diam. Stamens 4,included or longer pair slightly exserted; shorter filament pairca. 2 mm; longer filament pair ca. 5 mm; anther thecae oblong,ca. 2 mm. Ovary pilose; style ca. 2 cm, villous; stigma pilose.Capsule ca. 1 cm, pubescent, 8-seeded. Seeds yellowish, broadlyovate in outline, ca. 2 × 2 mm, subglabrous; areole large. Fl.Oct–Mar, fr. Jun–Jul.Moist hollows along streams; near sea level to 1300 m. Guangdong,Guangxi, Hainan [Cambodia, Laos, Vietnam].4. Strobilanthes fluviatilis (C. B. Clarke ex W. W. Smith)Moylan & Y. F. Deng, Bot. J. Linn. Soc. 150: 389. 2006.溪畔黄球花 xi pan huang qiu huaHemigraphis fluviatilis C. B. Clarke ex W. W. Smith,Notes Roy. Bot. Gard. Edinburgh 10: 182. 1918; Sericocalyxfluviatilis (C. B. Clarke ex W. W. Smith) Bremekamp.Herbs to 20 cm tall, isophyllous. Stems decumbent, hirsute,basally woody. Petiole 1–5 mm, coarsely pilose; leaf bladeoblong, narrowly oblong-elliptic, or elliptic, 1.5–5 × 1–1.5 cm,abaxially subglabrous to thinly but coarsely pilose with pustulosetrichomes, adaxially darker green with prominent linearcystoliths and glabrous or strigose-pilose, secondary veins 4 or5 on each side of midvein, base cuneate, margin crenulate, apexobtuse. Inflorescences terminal, spikes, ca. 2.5 cm; bracts oblong,1–1.5 cm, persistent, pilose, base cuneate, apex obtuse;bracteoles narrowly linear-elliptic, ca. 7 × 1 mm, pilose. Calyxca. 8 mm, pilose, subequally 5-lobed almost to base; lobes linear,apex acute. Corolla yellow, 0.9–1.1 cm, glabrous; tube basallycylindric and narrow for ca. 3 mm then gradually widenedto ca. 3 mm at mouth; lobes orbicular, ca. 2 mm in diam.,spreading. Stamens 4, included; filaments with a few scatteredtrichomes, shorter pair ca. 1 mm, longer pair ca. 2 mm; antherthecae oblong; pollen type 6. Ovary oblong, comose; style glabrousor with a few trichomes. Capsule oblong, ca. 8 mm, comose,8–14-seeded. Seeds ca. 0.8 mm, pubescent; areola large.Fl. Feb–May.Along streams on rocks; 200–800 m. Guangxi, Guizhou, Yunnan[Myanmar, Thailand].5. Strobilanthes reptans (G. Forster) Moylan ex Y. F. Deng &J. R. I. Wood, comb. nov.匍匐半插花 pu fu ban cha huaBasionym: Ruellia reptans G. Forster, Fl. Ins. Austr. 44.1786; Hemigraphis okamotoi Masamune; H. pacifica Hosokawa;H. primulifolia (Nees) Fernández-Villar; H. reptans (G. Forster)T. Anderson ex Hemsley; H. tawadana Ohwi; R. primulifoliaNees.Herbs to 50 cm tall, perennial, isophyllous. Stems some-

ACANTHACEAE 391times rosette-forming but usually prostrate and rooting at nodes,sparsely to densely covered with multicellular trichomes. Petiole0.5–8 cm, pubescent, glabrescent; leaf blade oblong, elliptic,oblong-ovate, or suborbicular, 1.5–8 × 1–4 cm, both surfacespubescent with multicellular trichomes, secondary veins 4or 5 on each side of midvein, base abruptly truncate to slightlycordate, margin crenate to subentire, apex acute to obtuse. Inflorescencesterminal, spikes, elongating to 6–8 cm at maturity;bracts oblanceolate to elliptic, 8–12 × 2–5 mm, persistent, pubescenton veins, margin entire and ciliate; bracteoles absent.Calyx 6.5–10 mm, 5-lobed to base; lobes linear-lanceolate, ciliateand pubescent along veins, apex finely attenuate. Corollawhite or pale violet with darker veins, 1.3–1.5 cm, glabrous;tube basally cylindric for 4–6 mm; lobes orbicular, 2–5 mm,glabrous. Stamens 4, included; shorter filament pair ca. 1 mm,glabrous; longer filament pair ca. 2 mm, base slightly villous;anther thecae ca. 1 mm; pollen type 7. Ovary apex pubescent;style sparsely villous. Capsule 7–9 mm, apically pubescent, 8–16-seeded. Seeds orbicular in outline, ca. 1 mm in diam., pubescent.Fl. Apr–Sep, Dec.Weed of roadsides, ponds, and streamsides; near sea level to 400m. Taiwan [Indonesia, Japan, Malaysia, Papua New Guinea, Philippines;Australia, Pacific islands (New Caledonia)].The apparent earlier homonym, “Strobilanthes reptans Bojer”(Nees in Candolle, Prodr. 11: 368. 1847) was merely cited as a synonymand was therefore not validly published (Vienna Code, Art. 34.1(c));hence it does not preclude valid publication of the new combinationmade here. The apparent new combination “Strobilanthes primulifolia”(Y. F. Deng & J. R. I. Wood, J. Trop. Subtrop. Bot. 18: 483. 2010),based on Ruellia primulifolia Nees, was not validly published because afull and direct reference to the place of valid publication of the basionymwas not provided (Art. 33.4; “in DC., Prodr. 11: 144. 1847” wascited, but that publication cited the actual place: Nov. Actorum Acad.Caes. Leop.-Carol. Nat. Cur. 19(Suppl. 1): 382. 1843).Strobilanthes reptans is an ornamental plant.6. Strobilanthes cumingiana (Nees) Y. F. Deng & J. R. I.Wood, J. Trop. Subtrop. Bot. 18: 483. 2010.直立半插花 zhi li ban cha huaRuellia cumingiana Nees in A. Candolle, Prodr. 11: 148.1847; Hemigraphis cumingiana (Nees) Fernández-Villar.Herbs to 50 cm tall, erect, isophyllous. Stems 4-angled,thinly pubescent with multicellular trichomes. Petiole 0.8–2.5cm, sulcate, pubescent; leaf blade ovate-lanceolate, ovate, orelliptic, 4–10 × 1.5–5 cm, both surfaces densely covered withcystoliths, abaxially strigose along veins, adaxially darker andglabrous, secondary veins 5 or 6 on each side of midvein, basecuneate to rounded and often oblique, margin shallowly crenate,apex shortly acuminate. Inflorescences axillary or terminal,spikes, to 4 cm; peduncle 0.3–2 cm, pubescent to puberulent;rachis glabrous; bracts ovate, persistent, margin slightly ciliate,apex obtuse; basalmost bracts sterile, leaflike, 1.5–2 × 0.6–1cm, margin undulate; apical bracts fertile, ca. 10 × 6 mm, marginentire; bracteoles absent. Calyx 6.5–9.5 mm, subequally 5-lobed to base; lobes lanceolate, margin ciliate, apically attenuate.Corolla white, 1.3–1.8 cm, glabrous; tube basally cylindricfor 4–6 mm; lobes orbicular, ca. 2.5 mm in diam. Stamens 4,included; shorter filament pair 1–1.5 mm, glabrous; longer filamentpair 2–3 mm, villous; anther thecae oblong, ca. 1.5 mm;pollen type 8. Ovary comose; style ca. 1 cm, hirsute. Capsuleca. 9 mm, apically pubescent, 6–8-seeded. Seeds ca. 2 × 1.2–2mm, pubescent. Fl. Feb–May.Forests; 100–300 m. Taiwan (Lan Yu) [Indonesia, Malaysia, Philippines].7. Strobilanthes sinica (H. S. Lo) Y. F. Deng, comb. nov.安龙马蓝 an long ma lanBasionym: Dyschoriste sinica H. S. Lo, Acta Phytotax.Sin. 17(4): 85. 1979.Herbs to 10 cm tall, isophyllous. Stems prostrate, 4-angled, glabrous, internodes usually less than 1 cm, rooting atbasal nodes. Leaves shortly petiolate or sessile; leaf blade oblong-lanceolate,0.7–3 × 0.3–0.7 cm, both surfaces glabrous,secondary veins ca. 3 on each side of midvein, base cuneate,margin entire or apically inconspicuously serrate, apex obtuse.Flowers axillary, solitary. Bracts similar to leaves but smaller;bracteoles linear-spatulate, 4.5–5 mm, apex obtuse. Calyx 5.5–6mm, 5-lobed almost to base; lobes lanceolate, margin sparselyciliate. Corolla purple, ca. 1.1 cm, straight; tube basally cylindricand narrow for ca. 4 mm then widened to ca. 4.5 mm atthroat, outside reported, perhaps erroneously, to be glabrous;lobes ?obcordate, 2–2.5 mm, subequal. Stamens 4, included;filaments glabrous, shorter pair ca. 1.2 mm, longer pair ca. 3.5mm; anther thecae parallel; pollen type 10. Ovary glabrous;style ca. 8 mm, slightly hairy. Capsule oblong, ca. 6.5 mm, glabrous,4-seeded. Seeds ovate in outline, ca. 2 × 2 mm, denselyappressed pubescent. Fl. Aug–Sep.● Grassy slopes; ca. 1300 m. Guizhou (Anlong).8. Strobilanthes szechuanica (Batalin) J. R. I. Wood & Y. F.Deng, Bot. J. Linn. Soc. 150: 375. 2006.四川马蓝 si chuan ma lanHemigraphis szechuanica Batalin, Trudy Imp. S.-Peterburgsk.Bot. Sada 13: 384. 1894.Herbs to 20 cm tall, isophyllous. Stems decumbent, 4-angled, subglabrous, rooting at basal nodes. Petiole 3–5 mm,puberulent; leaf blade ovate to obovate, becoming suborbiculartoward branch tips, 1–2 × 0.8–1 cm, glabrous or margin with afew scattered cilia, both surfaces densely covered with whitishcystoliths, secondary veins 3 or 4 on each side of midvein, basebroadly cuneate and decurrent onto petiole, margin entire, apexobtuse to rounded. Flowers axillary, solitary; bracteoles oblanceolate,5–7 × ca. 1 mm, pilose and with abundant cystoliths,margin thinly ciliate. Calyx 8–10 mm, with abundant cystoliths,5-lobed almost to base; lobes linear to linear-lanceolate, marginciliate, apex obtuse. Corolla pale lilac, funnel-shaped, 1.8–2.3cm, outside pubescent; tube basally cylindric and narrow for ca.5 mm then gradually widened to ca. 8 mm at mouth; lobessuborbicular, ca. 3 × 3 mm, margin ciliate. Stamens 4, included;shorter filament pair ca. 3 mm, glabrous; longer filament pairca. 5 mm, pilose; anther thecae oblong, ca. 2 × 1.2 mm; pollentype 10 (requires confirmation). Ovary oblong, comose; style

392ACANTHACEAE1.4–1.8 cm, pilose. Capsule ca. 1 cm, pilose at tip, 4-seeded.Seeds ca. 2 mm in diam., pilose. Fl. Jul–Sep.● Streamsides, on rocks; 700–1500 m. Sichuan.9. Strobilanthes wilsonii J. R. I. Wood & Y. F. Deng, Bot. J.Linn. Soc. 150: 371. 2006.乐山马蓝 le shan ma lanHerbs to 30 cm tall, perennial, isophyllous. Stems procumbent,sulcate, glabrous, rooting at nodes; flowering branchesarising in pairs from nodes, apparently erect, 3–20 cm, sparselybifariously pilose especially apically, with reflexed whitishlarge-celled trichomes. Petiole 0.5–2 cm, bifariously hirsutewith short reflexed whitish trichomes; leaf blade narrowlyelliptic, broadly elliptic, or obovate-spatulate, 1.5–3 × 1–1.5cm, glabrous except for a few marginal trichomes near petiole,adaxially with prominent cystoliths, secondary veins 3–5 oneach side of midvein, base attenuate, margin serrate, apex veryshortly acuminate to obtuse. Inflorescences axillary or terminal,headlike spikes, 1–1.5 cm, few flowered; peduncle 0.2–1.8 cm,pilose; bracts obovate-spatulate, 4–10 × 2–6 mm, apicallydiminishing in size, abaxially glabrous, adaxially with a fewtrichomes toward base, margin basally entire and ciliate withthick white spreading trichomes and apically toothed and notciliate, apex ± obtuse; bracteoles oblanceolate to obovate-spatulate,2–3 × ca. 0.5 mm, margin ciliate with a few large white trichomesto 2 mm, apex ± rounded. Calyx ca. 4 mm, subequally5-lobed to ca. 1 mm above base; lobes pale but with a greentip, linear-oblong, ca. 3.2 × 1 mm, pilose with large-celled butmostly marginal trichomes, apex obtuse. Corolla blue, 7–8 mm,funnel-shaped, outside pilose; tube basally cylindric and ca. 1mm wide for ca. 2 mm then gradually widened to ca. 5 mm atmouth; lobes ovate, ca. 1.5 × 1.5 mm, apex acute. Stamens 4;filaments glabrous, shorter pair ca. 2 mm, longer pair ca. 4 mmand slightly exserted; anther thecae oblong, ca. 0.5 × 0.2 mm;central staminode present, ca. 1 mm; pollen type 9. Ovary pilose;style ca. 8 mm. Capsule (immature) ca. 2.5 mm, pilosewith large-celled trichomes, 4-seeded. Seeds ovate in outline,lenticular, ca. 2 × 2 mm, densely covered with appressed trichomes;areola small, glabrous. Fl. Sep–Oct.● Moist shaded places; 800–1100 m. Sichuan.10. Strobilanthes labordei H. Léveillé, Repert. Spec. Nov.Regni Veg. 12: 20. 1913.薄叶马蓝 bao ye ma lanAcanthopale debilis C. B. Clarke ex S. Moore; A. labordei(H. Léveillé) Handel-Mazzetti; Championella debilis Bremekamp;C. labordei (H. Léveillé) E. Hossain; Strobilanthesdebilis Hemsley (1890), not C. B. Clarke (1885).Herbs, creeping, branched, isophyllous. Stems white villous,rooting at nodes. Petiole 5–10 mm; leaf blade ovate, 2–3 ×1.5–2 cm, both surfaces white hirsute, abaxially glaucous, adaxiallygreen and sometimes purplish dotted, secondary veins 4 or5 on each side of midvein, base broadly cuneate to rounded,margin sparsely crenulate, apex acuminate. Inflorescences terminal,abbreviated to headlike spikes, ca. 1 cm, densely coveredwith large-celled white trichomes; bracts leaflike, 1–2 cm;bracteoles linear, ca. 5 mm. Calyx 5–7 mm, 5-lobed almost tobase; lobes linear-lanceolate, densely white hirsute. Corolla purplishblue to violet, 1.5–1.8 cm, outside pubescent, inside glabrousexcept for trichomes retaining style; tube basally cylindricand narrow then gradually widened to ca. 1.2 cm at mouth;lobes oblong, ca. 3 × 3 mm, subequal, apex rounded. Stamens4; shorter filament pair ca. 3 mm; longer filament pair ca. 7mm, exserted; anther thecae ca. 1.8 × 1 mm, weakly exserted;pollen type 11. Ovary pilose at tip; style ca. 1.2 cm, pilose. Capsulenarrowly obovoid, 6–8 mm, 4-seeded, apex pilose. Seedsovate in outline, ca. 2 × 2 mm, pubescent. Fl. Aug–Dec.● Valleys, moist places; 400–1800 m. Chongqing, Guangdong,Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan.11. Strobilanthes tetrasperma (Champion ex Bentham) Druce,Rep. Bot. Soc. Exch. Club Brit. Isles 1916: 649. 1917.四子马蓝 si zi ma lanRuellia tetrasperma Champion ex Bentham, Hooker’s J.Bot. Kew Gard. Misc. 5: 132. 1853; Acanthopale radicans (T.Anderson ex Bentham) C. B. Clarke ex Benoist; A. tetrasperma(Champion ex Bentham) Handel-Mazzetti; Championella tetrasperma(Champion ex Bentham) Bremekamp; Strobilanthesradicans T. Anderson ex Bentham.Herbs 30–50 cm tall, isophyllous. Stems often procumbentand rooting at base, 4-angled, sulcate, retrorsely pubescentalong sulci, glabrescent, basally slightly woody. Petiole 0.5–2.5cm, sulcate, glabrous; leaf blade ovate, elliptic, or oblong, 1.5–11 × 1–4.5 cm, glabrous, adaxially darker and densely coveredwith cystoliths, secondary veins ca. 5 on each side of midvein,base attenuate or decurrent onto petiole, margin serrate to dentate,apex acuminate to acute. Inflorescences terminal or (sometimes)axillary, short often headlike spikes, few flowered, oftenreduced to a single flower pair; bracts leaflike, obovate to spatulate,1–1.5 × 0.4–0.5 cm, sparsely pilose, pinnately veined;bracteoles linear, 5–6 × ca. 1 mm, abaxially glabrous or sparselypilose and densely covered with cystoliths. Calyx 5-lobed almostto base; lobes linear, 5–7 × ca. 1 mm, accrescent to ca. 8mm in fruit, glabrous except for large-celled trichomes on margin,densely covered with cystoliths. Corolla purple to purplishblue, funnel-shaped, 1.5–2 cm, straight, outside pubescent, insidevillous; tube basally cylindric and ca. 2 mm wide for ca. 5mm then gradually widened to ca. 1.5 cm at mouth; lobes oblong,ca. 4 × 5 mm, apex obtuse. Stamens 4, included; shorterfilament pair ca. 2 mm, glabrous; longer filament pair ca. 5 mm,sparsely pubescent; anther thecae oblong, ca. 2 mm; pollen type10. Ovary ovoid, ca. 3 mm, hispid at tip, with 4 ovules perlocule; style ca. 1.5 cm, hirsute; stigma 2-lobed, lobes unequal.Capsule oblong, 7–10 mm, apically pubescent, 4-seeded. Seedsovate in outline, ca. 2 × 2 mm, pubescent. Fl. Jul–Dec.Forests, shaded grasslands, rocks by streams, roadsides; 100–1000m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei,Hunan, Jiangxi, Sichuan [Vietnam].12. Strobilanthes japonica (Thunberg) Miquel, Ann. Mus. Bot.Lugduno-Batavi 2: 124. 1866.日本马蓝 ri ben ma lanRuellia japonica Thunberg in Murray, Syst. Veg., ed. 14,

ACANTHACEAE 393576. 1784; Acanthopale japonica (Thunberg) C. B. Clarke ex S.Moore; Championella japonica (Thunberg) Bremekamp; Strobilanthesbonatiana H. Léveillé.Herbs 20–50 cm tall, isophyllous. Stems slender, herbaceous,much branched; branchlets purplish red, 4-angled thenterete, glabrous. Petiole 2–5 cm, white pubescent; leaf bladenarrowly elliptic to lanceolate, 2–5 × 0.5–1.8 cm, both surfacesglabrous and densely covered with prominent linear cystoliths,secondary veins 3–5 on each side of midvein, base cuneate tobroadly cuneate, margin crenulate, apex long acuminate. Inflorescencesterminal, spikes, 1–3(–5) cm, sometimes reduced to asingle flower pair; bracts imbricate, oblanceolate to oblonglanceolate,7–10 × 2–3 mm, as long as corolla tube, glabrousexcept for pubescence at base, apically densely covered withcystoliths, margin entire, apex obtuse; bracteoles linear, ca. 5mm. Calyx equally 5-lobed almost to base; lobes linear, 6–7mm, margin ciliate, apex rounded. Corolla purplish white orwhite, funnel-shaped, ca. 1.5 cm, straight, outside pubescent,inside glabrous except for trichomes retaining style; tube basallycylindric for 5–6 mm; lobes oblong-elliptic, 5–6 mm, equal.Stamens 4, included; shorter filament pair ca. 8 mm; longerfilament pair ca. 1.2 cm; anther thecae oblong, ca. 1.5 mm; pollentype 10. Ovary glabrous; style filiform, ca. 1.2 cm, pilose;stigma 2-cleft. Capsule oblong, ca. 7 mm, presumably glabrous,4-seeded. Seeds ovate in outline, ca. 3 mm, densely appressedpubescent. Fl. Aug–Sep, fr. Oct–Nov. 2n = 30.Often near temples and religious sites; 500–1100 m. Chongqing,Guizhou, Hubei, Hunan, Sichuan [Japan].Strobilanthes japonica differs from S. tetrasperma in its lanceolateleaves and glabrous ovary. We have seen two collections intermediatebetween these two species (Yu Xunlin 91690 from Hunan and Exped. ToFlora of Yunnan, Guizhou and Guangxi 50124 from Guizhou).“Dyschoriste grandiflora H. S. Lo” (C. C. Hu, FRPS 70: 79.2002) belongs here but is a nomen nudum and was therefore not validlypublished (Vienna Code, Art. 32.1(d)).13. Strobilanthes wangiana Y. F. Deng & J. R. I. Wood, J.Trop. Subtrop. Bot. 18: 473. 2010.启无马蓝 qi wu ma lanHerbs to 1 m tall, perennial, isophyllous. Stems sulcate,obscurely bifariously pubescent, glabrescent; nodes pustulose,sparsely pubescent. Petiole 4–5(–20) mm, almost glabrous;leaf blade elliptic to ovate-elliptic, 2.5–6 × 1.1–2.8 cm, bothsurfaces glabrous except for large-celled trichomes adaxiallyalong midvein, cystoliths prominent, secondary veins 5–7 oneach side of midvein, base cuneate and attenuate onto petiole,margin crenate, apex obtuse. Inflorescences terminal, subcapitatespikes, 2–4 cm, few flowered; bracts leaflike; basalbracts elliptic, 2–4 × 1–1.4 cm, apically diminishing in size,margin serrate; apical bracts oblong, 1.4–2 × 0.4–0.6 cm,abaxially sparsely pubescent, adaxially glabrous, margin ciliateand entire except for a few teeth near apex; bracteoles linearlanceolate,9–11 × ca. 1 mm, abaxially silky pubescent, adaxiallyglabrous, margin ciliate. Flowers imbricate on rachis, sessile.Calyx 1–1.3 cm, 5-lobed to ca. 3 mm above base; lobeslinear-oblanceolate, 10–14 × 0.6–1 mm, outside silky pubescent,inside silky pubescent from middle apically and glabrescenttoward base, margin ciliate. Corolla purplish, 3.5–4 cm,ventricose and bent to ca. 90° at mouth, outside pubescent, insideglabrous except for trichomes retaining style; tube basallycylindric and ca. 2 mm wide for ca. 1 cm then graduallywidened to ca. 1 cm at mouth; lobes ovate, 4–5 × 4–5 mm,apex obtuse. Stamens 4, included; filaments pubescent, shorterpair 1–2 mm, longer pair 3–4 mm; anther thecae oblong, ca.1.5 mm. Ovary ca. 2 mm, glabrous except for a few trichomesat tip. Capsule oblong, ca. 10 × 3 mm, glabrous, 4-seeded.Seeds ovate in outline, ca. 3 × 2 mm, compressed, pubescent;areola small. Fl. Aug–Sep, fr. Nov.● Forests; 1200–2300 m. Yunnan.14. Strobilanthes oligantha Miquel, Ann. Mus. Bot. Lugduno-Batavi 2: 124. 1866.少花马蓝 shao hua ma lanAcanthopale oligantha (Miquel) C. B. Clarke ex S. Moore;Championella oligantha (Miquel) Bremekamp.Herbs 40–50 cm tall, isophyllous. Stems 4-angled, sulcate,sparsely and retrorsely white hairy. Petiole 3.5–4 cm, thinlypilose; leaf blade broadly ovate to elliptic, 4–7(–10) × 2–4 cm,both surfaces with prominent white cystoliths, abaxially thinlypilose on veins, adaxially glabrous to thinly pubescent, secondaryveins 4–6 on each side of midvein, base broadly cuneate,margin remotely serrate, apex acuminate. Inflorescences shortspikes, elongating to ca. 4 cm in fruit, sometimes reduced to asingle flower pair; bracts leaflike; outer bracts ca. 1.5 cm; innerbracts smaller, densely covered with white large-celled trichomes;bracteoles linear-spatulate, ca. 1 cm, densely coveredwith white large-celled trichomes. Flowers imbricate on rachis.Calyx ca. 8 mm, subequally 5-lobed almost to base; lobes linear,subequaling bracteoles, inside apically with trichomes. Corollabluish purple, 3–3.5 cm, usually bent, outside sparsely pubescenton lobes, inside glabrous except for trichomes retainingstyle; tube basally cylindric and narrow for ca. 1.5 cm thengradually widened to ca. 1.3 cm at mouth, expanded part campanulateand ca. 2.5 cm; lobes ca. 5 mm, subequal. Stamens 4,included; shorter filament pair ca. 2 mm, glabrous; longer filamentpair ca. 5 mm, pilose; anther thecae oblong, ca. 2 × 1 mm;pollen type 10. Ovary pubescent at tip; style ca. 3.2 cm, pilose.Capsule narrowly obovoid, ca. 1 cm, pubescent at tip, 4-seeded.Seeds ovate in outline, ca. 2.5 × 2 mm, pubescent. Fl. Jul–Sep,fr. Oct–Nov. 2n = 60.Forests, shaded moist grasslands; 100–800 m. Anhui, Fujian,Jiangxi, Zhejiang [Japan, Korea].15. Strobilanthes austrosinensis Y. F. Deng & J. R. I. Wood, J.Trop. Subtrop. Bot. 18: 470. 2010.华南马蓝 hua nan ma lanHerbs to 50 cm, perennial, drying a distinct dark greencolor, isophyllous. Stems basally procumbent and rooting atnodes but eventually ascending, sulcate, bifariously pubescentespecially apically, with reflexed whitish large-celled trichomes.Petiole 0–2 cm, bifariously hirsute with reflexedwhitish large-celled trichomes; leaf blade elliptic to suborbicular,2–8 × 1–4.5 cm, abaxially pilose along veins, adaxially piloseand hirsute especially toward base and densely covered

394ACANTHACEAEwith white cystoliths, secondary veins 3–5 on each side ofmidvein, base attenuate, margin serrate but becoming subentiretoward base, apex acute to obtuse. Inflorescences terminal,headlike spikes, 1–2 cm, often elongating at maturity, leafy, fewflowered; rachis densely whitish pilose; bracts shortly petiolate,leaflike, broadly obovate-spatulate, 1.5–1.8 × 0.5–0.8 cm, abaxiallydensely covered with whitish multicellular trichomes towardbase, adaxially subglabrous, margin basally entire and apically1–3-dentate, apex rounded to obtuse; bracteoles spatulate,8–11 × 1–2.5 mm, abaxially toward base densely covered withwhitish multicellular large-celled trichomes, adaxially whitishpilose. Calyx 1–1.2 cm, 5-lobed almost to base; lobes oblonglanceolate,outside pilose, inside subglabrous, midvein prominent,margin ciliate with thick white apically spreading cilia,apex obtuse. Corolla bluish purple, funnel-shaped, 2.2–2.8 cm,ventricose, straight to slightly curved, outside pilose whenyoung but glabrescent, inside glabrous except for trichomesretaining style; tube basally cylindric and ca. 2 mm wide for1.2–1.5 cm then gradually widened to ca. 1.5 cm at mouth;lobes ovate-elliptic, ca. 5 mm. Stamens 4, included; filamentshirsute, shorter pair ca. 6 mm, longer pair ca. 8 mm; antherthecae oblong, ca. 1.2 × 0.4 mm; pollen type 10. Ovary ovoid,ca. 2 mm, glabrous except for comose tip; style ca. 1.5 cm, hirsute;stigma unequally 2-cleft. Capsule oblong, 9–10 × ca. 3mm, glabrous except for pilose tip and along suture, 4-seeded.Seeds ovate in outline, ca. 2.5 × 2 mm, densely pilose; areolasmall, glabrous. Fl. Jul–Sep, fr. Oct–Nov.● Thickets, forest margins; 100–1500 m. Guangdong, Guangxi,Hunan, Jiangxi.16. Strobilanthes sarcorrhiza (C. Ling) C. Z. Zheng ex Y. F.Deng & N. H. Xia, Novon 17: 154. 2007.菜头肾 cai tou shenChampionella sarcorrhiza C. Ling, Acta Phytotax. Sin.13(3): 93. 1975.Herbs 20–40 cm tall, unbranched, isophyllous. Rhizomesswollen, fleshy. Stems 4-angled, sulcate, densely pubescent.Petiole 0–3 mm; leaf blade oblong-lanceolate, 4–18 × 1.5–3cm, abaxially pubescent along veins, adaxially glabrous anddensely covered with acerose cystoliths, secondary veins 5–9on each side of midvein, base narrowly cuneate, margin dentateto undulate, apex acuminate. Inflorescences terminal, subcapitatespikes, 2–4(–5) cm, 4–10-flowered; rachis pubescent; bractsleaflike, obovate-elliptic, 1–2 × 0.3–1 cm, glabrous; bracteoleslinear-lanceolate, ca. 8 mm, glabrous except for large-celled trichomeson margin, abaxially densely covered with white clavatecystoliths. Calyx ca. 1 cm, 5-lobed almost to base; lobeslinear, unequal, glabrous except for large-celled trichomes onmargin, outside densely covered with white clavate cystoliths.Corolla purple, funnel-shaped, 3.5–4.5 cm, ventricose, outsideglabrous, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 3 mm wide for ca. 7 mm thengradually widened to ca. 1.5 cm at mouth; lobes oblong, ca. 4 ×3 mm, subequal, apex emarginate. Stamens 4, included; filamentspilose, shorter pair ca. 4 mm, longer pair ca. 6 mm; antherthecae oblong, ca. 1.5 mm; pollen type 3. Ovary glabrous;style filiform, ca. 3.2 cm, pubescent. Capsule glabrous, 4-seeded. Seeds ovate in outline, ca. 2.5 × 2 mm, densely pubescent.Fl. Aug–Sep, fr. Nov.● Forests in valleys; 200–600 m. Zhejiang.This species is used medicinally.17. Strobilanthes procumbens Y. F. Deng & J. R. I. Wood, J.Trop. Subtrop. Bot. 18: 476. 2010.金佛山马蓝 jin fo shan ma lanHerbs to 70 cm tall, perennial, isophyllous. Stems at firstprocumbent and rooting at nodes then erect, sulcate, denselypubescent with multicellular trichomes. Petiole 3–11 mm, sulcate,densely pubescent with multicellular trichomes; leaf bladeovate to rhomboidal, 0.8–3 × 0.3–2 cm, both surfaces pubescentwith multicellular trichomes, adaxially with numerous cystoliths,secondary veins 3–5 on each side of midvein, abaxiallyelevated, and adaxially impressed, base cuneate and slightlydecurrent onto petiole, margin crenate, apex acute. Inflorescencesterminal, subcapitate spikes, ca. 1 cm; bracts leaflike,oblanceolate, 4–12 × 2–4 mm, apically diminishing in size,abaxially pubescent, adaxially pubescent with white to slightlyyellowish multicellular trichomes, margin entire or with 1 or 2teeth and silky ciliate, apex obtuse; bracteoles linear-spatulate,ca. 5 × 1 mm, indumentum as for bracts. Calyx subequally 5-lobed to base, white pilose; lobes linear, apex obtuse. Corollawhite, funnel-shaped, 2.8–3.3 cm, straight to slightly curved,outside pubescent especially on lobes, inside glabrous exceptfor trichomes retaining style; tube basally cylindric and ca. 1mm wide for ca. 6 mm then gradually widened to ca. 1 cm atmouth; lobes oblong, ca. 2 × 2 mm. Stamens 4, included;shorter filament pair ca. 2 mm, glabrous; longer filament pairca. 4 mm, pubescent; anther thecae oblong, ca. 1.3 mm; pollentype ?4. Ovary glabrous except for comose tip; style ca. 1 cm,glabrous. Capsule oblong, ca. 0.8 cm, glabrous except for comosetip, 4-seeded. Seeds elliptic in outline, ca. 1 × 1 mm, pubescent;areola small. Fl. Aug–Sep, fr. Nov.● Streamsides; 800–1000 m. Chongqing (Nanchuan).18. Strobilanthes lihengiae Y. F. Deng & J. R. I. Wood, Bot. J.Linn. Soc. 150: 379. 2006.李恒马蓝 li heng ma lanHerbs to 40 cm tall, perennial, isophyllous. Stems roundedto slightly sulcate, apparently ascending, bifariously pubescentto subglabrous. Petiole 0.2–4 cm, pilose with large whitemulticellular trichomes; leaf blade ovate to ovate-elliptic, 1.5–11 × 0.5–6 cm, both surfaces pilose with large-celled trichomesespecially on veins but adaxially sparser, secondary veins 4 oneach side of midvein, base cuneate, margin dentate to crenate,apex shortly acuminate. Inflorescences terminal on main stemand on small axillary branchlets, headlike spikes, very compact;bracts obovate, 8–11 × 6–9 mm, shortly and densely greenishtomentose and also pilose with long large-celled white glandtippedand non-glandular trichomes, margin entire, apexrounded; bracteoles oblong-oblanceolate, ca. 6 × 1.5–2 mm,margin long ciliate. Calyx subequally 5-lobed to ca. 1 mm frombase; lobes green-tipped, lanceolate-oblong, 8–9 × ca. 1.3 cm,exceeding bracteoles, margin long ciliate, apex obtuse. Corolla

ACANTHACEAE 395with a whitish tube base and blue apical part of tube and lobes,3.2–4 cm, ventricose, sparsely pilose on lobes; tube basallycylindric and ca. 1 mm wide for 8–10 mm then bent and expandedgradually to ca. 1.5 cm wide at mouth; lobes ovate,ca. 5 × 5 mm, apex acute. Stamens 4, included; filamentscoarsely pilose, shorter pair ca. 3 mm, longer pair ca. 5 mm;anther thecae narrowly oblong, ca. 3 × 0.5 mm; pollen type 3.Ovary villous; style 2.5–3 cm, basally sparsely pilose. Capsulecylindric, ca. 12 × 3 mm, comose, 4-seeded. Seeds ovate-ellipticin outline, ca. 2.5 × 2 mm, pilose. Fl. Nov–Dec.● Disturbed evergreen broad-leaved forests; ca. 2000 m. Yunnan(Tengchong).19. Strobilanthes esquirolii H. Léveillé, Repert. Spec. Nov.Regni Veg. 12: 18. 1913.白头马蓝 bai tou ma lanPyrrothrix heterochroa (Handel-Mazzetti) C. Y. Wu & C.C. Hu; Strobilanthes heterochroa Handel-Mazzetti; S. leucocephalaCraib; Tetragoga angustiphylla Q. H. Chen; T. esquirolii(H. Léveillé) E. Hossain.Subshrubs 0.5–2 m tall, much branched, anisophyllous.Stems 4-angled, basally glabrescent, apically rust-colored retrorselysetose. Petiole 1–3 cm, hispid-pilose; leaf blade ovateellipticto oblong-elliptic, 4–17 × 2–8 cm, smaller of pair ca. 3/4size of larger one, roughly hirsute, abaxially paler, adaxiallywith prominent cystoliths, secondary veins 7–10 on each side ofmidvein, abaxially prominent, and adaxially impressed, basecuneate and attenuate, margin serrulate and ciliate, apex acuteto caudate. Inflorescences terminal, heads, often yellowish, 2–3× ca. 2 cm, 8–10-flowered; rachis densely white sericeous withgland-tipped trichomes; bracts oblong-lanceolate, 1–3 × 0.5–0.7cm, outer ones longer with caudate tips giving inflorescences aneared appearance, inner ones smaller, indumentum variable,white sericeous, with gland-tipped or non-glandular trichomes,and apically often becoming brownish; bracteoles oblanceolate,7–9 × ca. 2 mm, abaxially basally white sericeous, trichomesapically often slightly rust-colored. Flowers imbricate on rachis.Calyx 7–10 mm, 5-lobed almost to base; lobes linear-oblanceolate,densely white sericeous toward base but slightly rustcoloredtoward apex, apex acute. Corolla violet, 3–3.5 cm,curved, ventricose, outside densely white pilose, inside glabrousexcept for trichomes retaining style; tube basally cylindric andca. 4 mm wide for ca. 6 mm then gradually widened to ca. 1 cmat mouth; lobes ovate, ca. 4 × 5 mm, apex obtuse. Stamens 4,included; shorter filament pair ca. 4 mm, glabrous; longer filamentpair ca. 7 mm, white pilose; anther thecae oblong, ca. 3 ×1 mm; pollen type 3. Ovary densely white pilose; style ca. 2.8cm, white pilose. Capsule oblong, 7–10 mm, apically white pilose,4-seeded. Seeds ovate in outline, ca. 3.5 × 2 mm, denselypubescent; areola small. Fl. Sep–Dec.Mountain slopes; 200–800 m. Guizhou, Yunnan [Laos, Thailand,Vietnam].20. Strobilanthes hossei C. B. Clarke, Bot. Jahrb. Syst. 41: 67.1907.红毛马蓝 hong mao ma lanChampionella fulvihispida (D. Fang & H. S. Lo) C. C. Hu& C. Y. Wu; Pyrrothrix hossei (C. B. Clarke) C. Y. Wu & C. C.Hu; P. rufohirta (C. B. Clarke ex W. W. Smith) C. Y. Wu & C.C. Hu; Strobilanthes fulvihispida D. Fang & H. S. Lo; S.rufohirta C. B. Clarke ex W. W. Smith.Herbs or subshrubs 20–30(–100) cm tall, much branched,anisophyllous. Stems slender, erect, 4-angled, basally glabrescent,apically reddish brown setose. Petiole 1.5–2.5 cm, denselyreddish setose; leaf blade ovate to ovate-lanceolate, 8–9 × 3.5–4cm, both surfaces reddish brown setose on veins, abaxiallywhitish, adaxially dark green, cystoliths absent or obscure, secondaryveins ca. 5 on each side of midvein, base broadly cuneateto subrounded, margin crenate, apex acuminate. Inflorescencesaxillary, spikes, 2–4 cm; rachis densely reddish setose;bracts spatulate, 1.3–1.6 × ca. 0.3 cm, persistent, basal clawpale reddish setose, expanded apex green and leaflike; bracteoleslinear-spatulate, 0.8–1.2 cm, reddish setose. Calyx ca. 1.3cm, 5-lobed almost to base; lobes linear-spatulate, reddishsetose. Corolla pale blue, 4–5 cm, straight, outside puberulent,inside glabrous except for trichomes retaining style; tube basallycylindric and ca. 2 mm wide for ca. 8 mm then graduallywidened to ca. 1.2 cm at mouth; lobes orbicular, 6–7 mm indiam. Stamens 4, included; filaments white villous, shorter pairca. 4 mm, longer pair ca. 6 mm; anther thecae oblong, ca. 2 × 1mm; pollen type 3. Capsule light brown, fusiform, ca. 1 cm,gland-tipped pubescent, 4-seeded, apex apiculate. Seeds brown,ovate in outline, ca. 2 × 2 mm, compressed, pubescent, apexshortly apiculate; areola absent. Fl. Aug–Dec, fr. Nov–Feb.Thickets, mountain slopes; 1000–1800 m. Guangxi, Yunnan [Indonesia,Laos, Malaysia, Myanmar, Thailand, Vietnam].21. Strobilanthes torrentium Benoist, Bull. Mus. Natl. Hist.Nat. 28: 188. 1922.急流马蓝 ji liu ma lanSubshrubs to 1 m tall, isophyllous or slightly anisophyllous.Stems dark green, erect, 4-angled, sulcate, bifariouslypubescent. Petiole 0.5–3 cm, pubescent and slightly hispid; leafblade ovate to elliptic, 3–13 × 1.5–5.5 cm, abaxially denselyyellow pilose along veins, adaxially sparsely pubescent, hispid,and with numerous cystoliths, secondary veins 5–7 on each sideof midvein, base narrowly cuneate and decurrent onto petiole,margin serrate, apex acuminate and slightly falcate. Inflorescencesaxillary or terminal, headlike spikes, 1.5–3 cm, verycompact, densely covered with long silky rufous trichomes;bracts imbricate, somewhat variable in size and shape; outerbracts leaflike, obovate to subrhombic, 1.5–1.7 × 0.5–0.7 cm,distinctly angled, densely villous, apex acute; inner bracts obovate,ca. 10 × 5 mm, angled, villous, apex subacute; bracteolesoblanceolate, ca. 7 mm, margin ciliate, apex obtuse. Calyx 1–1.1 cm, villous, 5-lobed to ca. 3 cm from base; lobes linearoblong7–8 × ca. 1 mm, with 2 slightly shorter than others,margin ciliate, apex obtuse. Corolla with a white tube and blueto mauve lobes, 3–4.5 cm, curved, outside sparsely pubescenton tube and on lobes, inside glabrous except for trichomes retainingstyle; tube basally cylindric and narrow for 8–10 mmthen gradually widened to 1–1.2 cm at mouth; lobes ovate, ca. 3× 5 mm, apex obtuse. Stamens 4, included; filaments pilose,

396ACANTHACEAEshorter pair ca. 2 mm, longer pair ca. 5 mm; anther thecae oblong,ca. 2.5 mm; pollen type 3. Ovary villous; style ca. 3.3 cm,basally glabrous, apically pubescent. Capsule oblong, ca. 9 mm,pilose at tip, 4-seeded. Seeds unknown. Fl. Aug–Nov.Forests by streams; 1900–2300 m. Yunnan [India, Myanmar].22. Strobilanthes anamitica Kuntze, Revis. Gen. Pl. 2: 498.1891.海南马蓝 hai nan ma lanChampionella maclurei (Merrill) C. Y. Wu & H. S. Lo;Strobilanthes maclurei Merrill.Herbs 50–100 cm tall, isophyllous. Stems reddish, stout,erect or procumbent and rooting at nodes, hirsute when young,glabrescent. Petiole 1–2 cm; leaf blade oblong-ovate, 3–13 ×1.5–5.5 cm, both surfaces golden to brown hirsute, adaxiallydensely covered with cystoliths, secondary veins ca. 5 on eachside of midvein, base cuneate to subrounded, margin crenulate,apex acuminate to shortly acuminate. Inflorescences axillary orterminal, headlike spikes, 1–3 cm, very compact, elongatedafter flowering; bracts obovate, ca. 1 cm, brown hirsute, pinnatelyveined, apex reflexed; bracteoles spatulate-linear, ca. 10× 1 mm, abaxially hirsute on apical half, adaxially subglabrous,margin ciliate. Calyx 6–8 mm, hirsute, 5-lobed almost to base;lobes linear, ca. 1 mm wide, outside hirsute on apical half, marginciliate. Corolla light blue or white, ca. 3 cm, outside pubescenton tube, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 0.2 mm wide for ca. 1.2 cmthen gradually widened to ca. 1 cm at mouth; lobes oblong, ca.6 mm. Stamens 4, included; filaments pilose, shorter pair ca. 3mm, longer pair ca. 6 mm; anther thecae oblong, ca. 2 × 1 mm;pollen type 2. Ovary pubescent at tip; style ca. 2.5 cm, hirsute.Capsule light brown, ca. 7 mm, pubescent at tip, 4-seeded.Seeds ovate in outline, ca. 2.5 × 2 mm, pubescent. Fl. Sep–Jan,fr. Dec–Mar.Forests, on rocks by streams; 400–1800 m. Guangxi, Hainan, Yunnan[Vietnam].23. Strobilanthes yunnanensis Diels, Notes Roy. Bot. Gard.Edinburgh 5: 164. 1912.云南马蓝 yun nan ma lanPteracanthus hygrophiloides (C. B. Clarke ex W. W.Smith) H. W. Li; P. hygrophiloides var. brachytrichus (Handel-Mazzetti) C. Y. Wu; P. mekongensis (W. W. Smith) C. Y. Wu &C. C. Hu; P. yunnanensis (Diels) C. Y. Wu & C. C. Hu;Strobilanthes hygrophiloides C. B. Clarke ex W. W. Smith; S.hygrophiloides var. brachytricha Handel-Mazzetti; S. hygrophiloidesvar. subnuda Benoist; S. limprichtii Diels; S. mekongensisW. W. Smith.Subshrubs 0.5–2.5 m tall, much branched, isophyllous.Stems 4-angled, bifariously whitish pubescent, sometimes glabrescent.Petiole 2–4 cm, whitish pubescent; leaf blade ovate,elliptic, or elliptic-ovate, 3–11 × 2–5 cm, abaxially pale greenand sparsely pubescent, adaxially dark green and pubescent,cystoliths abundant on both surfaces, secondary veins 4–7 oneach side of midvein, base narrowly cuneate and decurrent ontopetiole, margin crenate-serrate, apex acuminate and sometimesfalcate. Inflorescences axillary or terminal, subcapitate spikes,rarely reduced to a single flower; peduncle 0–4 mm, denselywhite pubescent; bracts persistent, hirsute; outermost sterilebracts leaflike, petiolate, lanceolate to ovate-lanceolate, 1.5–2.5× 0.3–0.4 cm; inner floral bracts linear-oblong, 1.2–1.6 cm,with 1 reddish brown vein; bracteoles linear to oblanceolate, ca.10 × 2 mm, pilose, apex obtuse. Flowers usually only one sterileper node. Calyx 1.2–1.6 cm, 5-lobed almost to base; lobes linear-lanceolate,11–15 × 1–1.5 mm, subhyaline, pilose especiallyon midvein, margin ciliate, apex finely acuminate. Corolla paleblue to pinkish, 3.5–4.5 cm, straight, outside densely white pilose,inside glabrous except for trichomes retaining style; tubebasally cylindric and 8–9 mm wide for 1–1.5 cm then graduallywidened to ca. 2 cm at mouth; lobes orbicular, 6–8 mm in diam.Stamens 4, included; filaments pilose, shorter pair ca. 2 mm,longer pair 4–7 mm; anther thecae oblong, ca. 4 × 1 mm; pollentype 3. Ovary whitish comose; style 2.8–3.4 cm, sparsely pilose.Capsule oblong, 11–13 × ca. 2.5 mm, apically sparselygland-tipped pubescent, 4-seeded. Seeds ovate in outline, 2.5–3.5 × 2–2.5 mm, densely appressed pilose. Fl. Jul–Sep, fr. Oct–Nov.● Moist open shady places in thickets; 800–2800 m. Gansu (Wenxian),Sichuan, Xizang (Mêdog, Zayü), Yunnan.24. Strobilanthes fengiana Y. F. Deng & J. R. I. Wood, J. Trop.Subtrop. Bot. 18: 481. 2010.冯氏马蓝 feng shi ma lanSubshrubs to 50 cm tall, perennial, weakly anisophyllous.Stems 4-angled, sulcate, glabrescent. Petiole 1–2 cm, sulcate,bifariously pubescent; leaf blade ovate, smaller of pair 6–8 × 3–4 cm and larger one 10–14 × 4–6 cm, abaxially glabrous exceptpilose along midvein, adaxially glabrous and with prominentcystoliths, secondary veins 4–7 on each side of midvein, baserounded to broadly cuneate, margin undulate and ciliate, apexacuminate. Inflorescences terminal, subcapitate spikes, 1.5–3cm; rachis pubescent; bracts lanceolate, 10–15 × ca. 2 mm,abaxially glabrescent, adaxially sparsely pubescent; bracteoleslanceolate, 8–10 × ca. 2 mm, abaxially glabrous, adaxiallydensely covered with multicellular large-celled trichomes. Calyxca. 1.2 cm, 5-lobed almost to base; lobes linear-spatulate,outside apically covered with multicellular large-celled trichomes,margin ciliate with multicellular large-celled trichomes,apex acute. Corolla purple, 3–3.5 cm, outside pilose with largecelledtrichomes, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 2 mm wide for ca. 1.5 cmthen bent to ca. 90° above middle and gradually widened to ca.1.3 cm at mouth; lobes rounded. Stamens 4, included; filamentspilose, shorter pair ca. 6 mm, longer pair ca. 8 mm; antherthecae oblong, ca. 2 mm. Capsule oblong, 12–14 × 3–4 mm,glabrous, 4-seeded. Seeds ovate in outline, ca. 3 × 2 mm, pubescent;areola small. Fl. Sep–Dec, fr. Nov–Feb.● Forests; 1200–1800 m. Yunnan (Xichou).25. Strobilanthes lamium C. B. Clarke ex W. W. Smith, NotesRoy. Bot. Gard. Edinburgh 10: 195. 1918.野芝麻马蓝 ye zhi ma ma lanPteracanthus lamium (C. B. Clarke ex W. W. Smith) C. Y.Wu & C. C. Hu.

ACANTHACEAE 397Herbs 30–80 cm tall, unbranched, anisophyllous. Stems 4-angled, sulcate, basally glabrescent, apically whitish glandtippedpubescent. Leaves apically denser; petiole 3–10 mm, hirsute;leaf blade ovate to ovate-lanceolate, 4–5 × 2.5–3 cm, bothsurfaces with very small conspicuous cystoliths, abaxially paler,sparsely pilose, and setose along veins, adaxially dark green,sparsely hirsute, and yellowish brown setose along midvein,secondary veins 7 or 8 on each side of midvein, base broadlycuneate and decurrent onto petiole, margin coarsely serrate,apex acute. Inflorescences terminal, subcapitate spikes, 3–5 cm,4–8-flowered; bracts oblong to lanceolate, resembling reducedleaves, 1.4–1.6 × 0.6–0.7 cm, persistent, abaxially hirsute especiallyon veins; bracteoles narrowly oblong-elliptic, ca. 1.2cm, abaxially gland-tipped pilose. Flowers imbricate on rachis.Calyx ca. 1.1 cm, gland-tipped pilose, 5-lobed almost to base;lobes linear-lanceolate, subequal, pilose, apex acute. Corollapurple, ca. 3.5 cm, bent at mouth, outside sparsely pilose, insideglabrous except for trichomes retaining style; tube basally cylindricand ca. 2 mm wide for ca. 6 mm then gradually widenedto ca. 1 cm at mouth; lobes orbicular, ca. 6 mm in diam. Stamens4, included; filaments densely pilose, shorter pair ca. 5mm, longer pair ca. 7 mm; anther thecae oblong, ca. 2.3 × 1.3mm, erect. Ovary pilose at tip; style pilose. Capsule not seen.Fl. Jul–Aug.● Forests, slopes; 800–1500 m. Chongqing, Hubei, Hunan.26. Strobilanthes cyphantha Diels, Notes Roy. Bot. Gard.Edinburgh 5: 162. 1912.弯花马蓝 wan hua ma lanPteracanthus cyphanthus (Diels) C. Y. Wu & C. C. Hu.Subshrubs 45–60 cm tall. Stems 4-angled. Petiole 2–5 cm;leaf blade ovate, 3–7 × 1.5–4.5 cm, both surfaces pilose, adaxiallywith numerous obscure cystoliths, secondary veins 6–8 oneach side of midvein, base decurrent onto petiole, margin serrate,apex acute. Inflorescences terminal or in apical leaf axils,subcapitate, dense, sessile or with peduncle to 5 mm, white pilose;outer sterile bracts leaflike; inner floral bracts elliptic, ca.12 × 5 mm, white pilose, margin dentate and ciliate; bracteoleslinear, ca. 16 × 1 mm, thinly pilose. Calyx 1–1.3 cm, 5-lobedalmost to base; lobes linear, ca. 11 × 1 mm, subequal, thinly pilose.Corolla blue, 3–4 cm, strongly bent and ventricose, outsidethinly pilose, inside pubescent on lobes and glabrous intube except for trichomes retaining style; tube basally cylindricand ca. 3 mm wide for ca. 1 cm then bent to ca. 90° at middleand widened to ca. 1.2 cm at mouth; lobes oblong, ca. 5 × 4mm. Stamens 4, included; shorter filament pair ca. 2 mm, glabrous;longer filament pair ca. 3 mm, glabrous except for a fewtrichomes basally; anther thecae oblong, ca. 2 × 1–1.5 mm;pollen type 3. Ovary comose; style 2.7–3.2 cm, pubescent. Capsuleca. 1.4 cm, pilose at tip, 4-seeded. Seeds ovate in outline,ca. 2.5 × 2 mm, densely covered with appressed trichomes. Fl.Jul–Oct, fr. Nov–Dec.● Mixed forests; 1200–3200 m. Gansu (Wenxian), Sichuan, SEXizang, Yunnan.“Strobilanthes curviflorus [sic!] C. B. Clarke” (C. Y. Wu, Index Fl.Yunnan. 2: 1685. 1984) belongs here but was not validly published becauseit was merely cited as a synonym (Vienna Code, Art. 34.1(c)).27. Strobilanthes strigosa D. Fang & H. S. Lo, Guihaia 17: 36.1997.糙毛马蓝 cao mao ma lanSubshrubs 30–60 cm tall, isophyllous. Stems black whendry, 4-angled, sulcate, bifariously pubescent and hirsute. Petiole5–12 mm, pubescent and hirsute; leaf blade elliptic to broadlyovate, 3.5–8 × 2–5 cm, both surfaces sparsely hispid pilose andwith conspicuous cystoliths, secondary veins ca. 7 on each sideof midvein and prominent, base cuneate to attenuate, marginserrate, apex acute. Inflorescences terminal or in apical leafaxils, spikes, 2–3 cm; bracts elliptic, 9–10 × 5–6 mm, persistent,pinnately veined, margin gland-tipped ciliate, apex acute;bracteoles oblanceolate, ca. 9 mm, densely covered with linearcystoliths, margin gland-tipped ciliate. Flowers imbricate on rachis.Calyx ca. 1.2 cm, 5-lobed to base; lobes falcately spatulate,ca. 2 mm wide, unequal, margin gland-tipped ciliate. Corollapurplish, ca. 4.5 cm, slightly bent, outside glabrous, insideglabrous except for trichomes retaining style; tube basallycylindric for ca. 1.5 cm then gradually widened to ca. 1 cm atmouth; lobes ovate, ca. 7 mm, apex rounded. Stamens 4, included;shorter filament pair ca. 2.5 mm, glabrous; longer filamentpair ca. 5 mm, pilose; anther thecae oblong, ca. 2 × 1.3mm; pollen type 4. Ovary comose; style ca. 2.8 cm, sparselypilose. Capsule not seen. Fl. Dec.● Limestone slopes; 400–600 m. Guangxi (Jingxi).Strobilanthes strigosa is a poorly understood species whose placementis uncertain.28. Strobilanthes longiflora Benoist, Bull. Mus. Natl. Hist.Nat. 28: 94. 1922.长花马蓝 chang hua ma lanChampionella longiflora (Benoist) C. Y. Wu & C. C. Hu.Herbs to 50 cm tall, isophyllous. Stems sulcate, glabrous.Petiole 2–4 mm, pubescent; leaf blade broadly lanceolate toovate, to ca. 10 × 5 cm, glabrous, cystoliths very prominent onboth surfaces, secondary veins 3–5 on each side of midvein,base cuneate, margin minutely crenulate, apex acuminate. Inflorescencesterminal or from apical axils, spikes, 0–4 cm, 8–12-flowered but sometimes reduced to a single flower in leaf axils;bracts oblanceolate-spatulate, 10–13 × 1.5–3 mm, sparsely pilose,apex entire to weakly 3-lobed; bracteoles oblong, ca. 6 × 1mm, pilose with a comose apex, apex obtuse. Calyx ca. 1 cm,5-lobed almost to base; lobes linear, ca. 9 × 1.5 mm, comose.Corolla blue, funnel-shaped, ca. 5 cm, strongly bent, outside pubescent,inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for ca. 8 mm thengradually widened to ca. 9 mm at mouth; lobes ovate, ca. 4 × 3mm, apex obtuse. Stamens 4, included; filaments pilose, shorterpair ca. 4 mm, longer pair ca. 6 mm; anther thecae oblong, ca.2.5 × 1.2 mm; pollen type 15. Ovary glabrous but pubescent attip; style sparsely white pilose; stigma linear. Capsule oblongfusiform,ca. 8 × 2 mm, glabrous apart from a few apical trichomes,4-seeded. Fl. Aug–Oct, fr. Nov–Dec.● Thickets; 700–1500 m. Yunnan.

398ACANTHACEAE29. Strobilanthes henryi Hemsley, J. Linn. Soc., Bot. 26: 240.1890.南一笼鸡 nan yi long jiGutzlaffia anisandra (Benoist) Handel-Mazzetti; G. anisandravar. drosothyrsa Handel-Mazzetti; G. forrestii S. Moore;G. henryi (Hemsley) C. B. Clarke ex S. Moore; G. lyi (H.Léveillé) E. Hossain; G. multiramosa Handel-Mazzetti; Paragutzlaffiahenryi (Hemsley) H. P. Tsui; P. lyi (H. Léveillé) H. P.Tsui; Ruellia lyi H. Léveillé; Strobilanthes anisandra Benoist.Subshrubs to 70 cm tall, variable from hirsute to subglabrous,weakly anisophyllous. Stems slender, erect or ascending,4-angled, much branched, villous to subglabrous. Petiole 0.5–3cm; leaf blade ovate, ovate-elliptic, or subrhomboidal, 2–9 ×1.5–4.5 cm, both surfaces thinly to densely pilose, abaxiallypaler, adaxially with linear cystoliths, secondary veins 5 or 6 oneach side of midvein, base broadly cuneate, slightly oblique,and usually decurrent onto petiole, margin crenulate, apexacute. Inflorescences terminal, spikes, usually 3–8 cm; bracts atbase of spike resembling sessile reduced leaves; floral bractsoblong to spatulate, 6–7 × 1–2 mm, persistent, abaxially glandtippedpubescent to subglabrous; bracteoles linear, ca. 7 × 0.8mm, abaxially gland-tipped pubescent to subglabrous. Flowerspaired, varying considerably in density, basally distant on rachisbut apically becoming imbricate. Calyx 8–11 mm, subequally5-lobed almost to base; lobes linear, 6–7 mm, outside glandtippedpubescent to subglabrous. Corolla light purple, blue, orwhite, funnel-shaped, 1.7–2.3 cm, outside pubescent, inside glabrousexcept for trichomes retaining style; tube basally cylindricand narrow for 3.5–6 mm then bent and gradually widenedto ca. 1 cm at mouth; lobes orbicular, ca. 2 mm in diam.,spreading. Fertile stamens 2, staminodes 2; filaments ca. 3.5mm, pubescent; anther thecae ovoid-oblong, exserted; pollentype 15. Ovary oblong, glabrous but with a few apical glands;style ca. 2 cm, glabrous. Capsule oblong, 7–10 mm, glabrousapart from a few apical glands, 4-seeded. Seeds suborbicular inoutline, ca. 2 × 2 mm, densely villous; areola small. Fl. Jan,Apr, Jul–Oct.● Mountain slopes; 1000–2800 m. Guizhou, Hubei, Hunan, Sichuan,SE Xizang, Yunnan.30. Strobilanthes cyclus C. B. Clarke ex W. W. Smith, NotesRoy. Bot. Gard. Edinburgh 10: 192. 1918.环毛马蓝 huan mao ma lanSubshrubs 40–60 cm tall, isophyllous, whole plant withhispid trichomes. Stems erect or suberect, with whitish multicellulartrichomes. Petiole 2–5 mm, densely pilose; leaf bladebroadly ovate to suborbicular, 2.5–5.5 × 2–4.5 cm, abaxiallypale green and hirsute along veins, adaxially dark green,densely hirsute, and with prominent linear cystoliths, secondaryveins 4 or 5 on each side of midvein, base broadly cuneateto subrounded, margin crenate to sometimes subentire, apexrounded to acute. Inflorescences axillary or terminal, capitatespikes, 1–3 cm, shortly pedunculate; bracts narrowly oblongelliptic,10–12 × 5–6 mm, densely pilose; bracteoles oblong, ca.7 × 2 mm, densely pilose on apical half, apex obtuse. Calyx ca.7 mm, outside pilose, weakly 2-lipped; lobes linear-oblong;lower 2 lobes distinct to base; upper 3 lobes fused in their basalhalf, with central lobe slightly longer. Corolla blue, 2–2.5 cm,substraight, outside glabrous except for pubescence on lobes,inside glabrous except for trichomes retaining style; tube basallycylindric and ca. 2 mm wide for ca. 1.2 cm then graduallywidened to ca. 1 cm at mouth; lobes suborbicular, ca. 5 mm indiam. Stamens 4; filaments glabrous, shorter pair ca. 3 mm andincluded, longer pair ca. 5 mm and exserted; anther thecae oblong,ca. 2.2 × 1.3 mm; pollen type 17. Ovary comose; style ca.2 cm. Capsule ellipsoid, ca. 7 × 3.5 mm, pilose at tip, 2-seededwith basal retinacula sterile. Seeds ovate to suborbicular in outline,ca. 1 mm, flattened, pubescent; areola small but prominent.Fl. Aug.● Forests; 1800–2300 m. Yunnan.31. Strobilanthes serrata J. B. Imlay, Bull. Misc. Inform. Kew1939: 117. 1939.齿叶马蓝 chi ye ma lanGutzlaffia glandulosa Lace.Subshrubs 50–100 m tall, perennial, erect, slightly anisophyllous.Stems sulcate, glabrous or with a few crisped trichomes.Petiole 0.5–7 cm, diminishing in length apically onstems, glabrous or sparsely pubescent; leaf blade ovate to oblong-ovate,5–12 × 4–6 cm, abaxially uniformly violet to lightgreen and glabrous except for pubescence along veins, adaxiallygreen variegated with white patches and glabrous, secondaryveins 7–9 on each side of midvein, base gradually narrowedand then ± decurrent to 2 cm onto petiole, margin coarsely serrate,apex acuminate. Inflorescences from apical leaf axils, simpleor branched pedunculate spikes, commonly forming a terminalpanicle of spikes; spikes 2–7 cm, densely gland-tippedpilose; peduncle of panicles 2–10 cm, subglabrous to thinly pubescent;rachis and branches densely gland-tipped pilose; bractsoblong-ovate to oblong-obovate, 3–4 × 1.5–2.5 mm, persistent,densely gland-tipped pilose, apex obtuse; bracteoles oblong toobovate, 3–4 × 1–1.5 mm, abaxially densely gland-tipped pilose,adaxially glabrous but densely covered with white cystoliths,apex obtuse. Flowers in opposite pairs, basally 1–2 cmapart on rachis, clearly separate all along spike except at veryapex. Calyx 5–6 mm, subequally 5-lobed to base; lobes oblong,outside densely gland-tipped villous, inside glabrous anddensely covered with white cystoliths. Corolla pale blue with awhite tube, 2–2.6 cm, straight or slightly bent, outside sparselyto densely pilose with large-celled trichomes; tube basally cylindricand narrow for 6–8 mm then widened to ca. 1 cm atmouth; lobes ovate, ca. 3 × 3 mm, apex obtuse. Fertile stamens2; filaments ca. 4 mm, glabrous; anther thecae ca. 1.5 mm,included; pollen type 18. Ovary villous; style ca. 2 cm, glabrous.Capsule oblong, 5–7 × ca. 2 mm, silky pubescent, 4-seeded. Seeds ca. 2 × 2 mm, pubescent. Fl. Sep–Jan.Partly shaded moist places in evergreen broad-leaved forests;700–1600 m. Yunnan [Myanmar, Thailand].32. Strobilanthes tubiflos (C. B. Clarke) J. R. I. Wood, EdinburghJ. Bot. 51: 264. 1994.管花马蓝 guan hua ma lan

ACANTHACEAE 399Strobilanthes petiolaris Nees var. tubiflos C. B. Clarke inJ. D. Hooker, Fl. Brit. India 4: 458. 1884.Subshrubs of unknown habit. Stems 4-angled to subterete,sulcate, glabrous except for a few trichomes in grooves.Petiole 5–10 mm; leaf blade ovate-elliptic, 6.5–16 × 3.5–8 cm,abaxially paler, sometimes purplish, and glabrous except forappressed trichomes on midvein, adaxially glabrous and withprominent cystoliths, secondary veins 4–7 on each side of midvein,base cuneate and decurrent as a wing onto petiole, margindentate, apex acuminate. Inflorescences terminal, branchedspikes, 2–4 cm; rachis gland-tipped pilose; sterile bracts leaflike,ovate, 9–10 × ca. 6 mm, with prominent cystoliths; floralbracts obovate, 5–6 × 2.5–3.5 mm, persistent, densely yellowishgland-tipped pubescent, cystoliths abaxially prominent,apex usually emarginate; bracteoles oblanceolate, ca. 5 × 1 mm,abaxially gland-tipped pubescent especially apically, apex obtuse.Calyx 7–8 mm, subequally 5-lobed almost to base; lobesoblong, ca. 1 mm wide, outside ciliate and apically gland-tippedpilose, inside glabrous. Corolla blue, 2–3 cm, curved, outsidepilose, inside glabrous except for trichomes retaining style; tubebasally cylindric and ca. 1 mm wide for ca. 3 mm then graduallywidened to ca. 1 cm at mouth; lobes broadly ovate, ca. 2 ×3 mm, apex rounded. Fertile stamens 2, included; filaments glabrous,ca. 4 mm; anther thecae oblong, erect, ca. 2 × 0.8 mm;pollen type 3. Ovary oblong-ovate, ca. 1 mm, apically pilose;style ca. 2 cm, glabrous. Capsule oblong, ca. 6 × 2 mm, pubescent,4-seeded. Seeds suborbicular in outline, ca. 1 × 1 mm,pubescent; areole prominent. Fl. Aug.● Valleys, streamsides; 200–300 m. Xizang (Mêdog).33. Strobilanthes affinis (Griffith) Terao ex J. R. I. Wood & J.R. Bennett, Kew Bull. 58: 134. 2003.肖笼鸡 xiao long jiAdenosma affinis Griffith, Not. Pl. Asiat. 4: 133. 1854;Strobilanthes acrocephala T. Anderson; S. darrisii H. Léveillé;S. thirionnii H. Léveillé; Tarphochlamys affinis (Griffith)Bremekamp; T. darrisii (H. Léveillé) E. Hossain.Herbs to 60 cm tall, isophyllous. Stems 4-angled, sulcate,white pubescent with large-celled trichomes. Petiole 0.5–1.5cm, pubescent; leaf blade ovate to ovate-elliptic, 3–8 × 2–4.5cm, both surfaces sparsely strigose, abaxially yellowish green,adaxially dark green and densely covered with subulate cystoliths,secondary veins 5–7 on each side of midvein, base subroundedto cuneate and decurrent onto petiole, margin crenulate,apex acute. Inflorescences axillary or terminal, densespikes, 2–6 cm; peduncle 0.5–2 cm, densely gland-tippedpubescent; sterile bracts sometimes present, sessile, suborbicularto ovate, 5–10 × 5–10 mm, strigose; floral bracts imbricate,broadly obovate, rhomboidal, or elliptic, 6–7 × 3–4 mm, glandtippedpilose with large-celled trichomes, 5-veined, margin ciliate,apex sometimes reflexed; bracteoles oblong-linear, ca. 5 ×1.2 mm, slightly shorter than calyx, gland-tipped pubescent,margin ciliate. Calyx 5–7 mm, 5-lobed almost to base; lobeslinear, one slightly longer than others, gland-tipped pubescent,1-veined, margin ciliate, apex obtuse. Corolla purplish blue toviolet, ca. 1.5 cm, curved, outside pilose, inside glabrous exceptfor trichomes retaining style; tube basally cylindric and ca. 1mm wide for ca. 7 mm then gradually widened to ca. 7 mm atmouth; limb somewhat 2-lipped, lower lip 3-lobed, upper lip2-lobed; lobes orbicular, ca. 3 × 3 mm, unequal. Stamens 4;shorter filament pair ca. 4 mm, glabrous; longer filament pairca. 6 cm, hirsute; anther thecae oblong, ca. 2 × 1.2 mm,strongly exserted; pollen type 14. Ovary pilose; style pilose, ca.1.2 cm. Capsule oblong, 7–9 mm, pubescent, 4-seeded. Seedsbrown, ovate to suborbicular in outline, ca. 1.5 × 1.5 mm, pubescent,glabrescent; areola small, glabrous. Fl. Sep–Nov.Mountain slopes, thickets; 600–1300 m. Guangxi, Guizhou, Hunan,Yunnan [India, Myanmar, Vietnam].Strobilanthes affinis is a very distinct species. Not only is the pollenunique but also the species is distinctive for having 4 exserted stamensand trichomes caducous on the seeds.34. Strobilanthes pinnatifida C. Z. Zheng, J. Hangzhou Univ.,Nat. Sci. Ed. 8: 431. 1981.羽裂马蓝 yu lie ma lanHu.Pteracanthus pinnatifidus (C. Z. Zheng) C. Y. Wu & C. C.Herbs to 40 cm tall, unbranched, isophyllous. Stems erect,brown hirsute. Petiole 1.5–2 cm, hirsute; leaf blade elliptic toovate-elliptic, 3–13 × 1–5 cm, whitish appressed hirsute, secondaryveins 6 or 7 on each side of midvein, base cuneate toattenuate and often decurrent onto petiole, margin coarsely dentateto deeply pinnatifid or lobed, apex acuminate to caudate.Inflorescences terminal, spikes, 5–9 cm; bracts spatulate-obovate,to 1.2 cm, pubescent; bracteoles linear, 1–1.5 cm, pubescent.Flowers distant on rachis. Calyx 1–1.5 cm, pubescent withgland-tipped trichomes, 5-lobed almost to base; lobes linear,equal. Corolla purplish blue, ca. 3.5 cm, ventricose, outside pubescent,inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 3 mm wide for ca. 1.2 mm thengradually widened to ca. 1 cm at mouth; lobes oblong, ca. 4 ×3.5 mm, apex emarginate to obtuse. Stamens 4, included; filamentspilose, shorter pair ca. 4 mm, longer pair ca. 6 mm;anther thecae oblong, ca. 2 mm. Ovary oblong, glabrous; styleca. 3 mm, sparsely hirsute. Capsule oblong, sparsely pubescent,4-seeded. Seeds ovate in outline, ca. 2.5 × 2 mm, pubescent;areola small. Fl. Aug–Sep, fr. Nov.● Forests in moist areas; 600–700 m. Zhejiang (Jingning, Qingyuan).Strobilanthes pinnatifida is unique because of its pinnatifid orlobed leaves.35. Strobilanthes auriculata Nees in Wallich, Pl. Asiat. Rar. 3:86. 1832.耳叶马蓝 er ye ma lanPerilepta auriculata (Nees) Bremekamp; P. edgeworthiana(Nees) Bremekamp; P. siamensis (C. B. Clarke) Bremekamp;Strobilanthes auriculata var. edgworthiana (Nees) C. B.Clarke; S. auriculata var. siamensis C. B. Clarke; S. edgeworthianaNees; S. siamensis C. B. Clarke.Subshrubs 0.5–2 m tall, much branched, weakly tostrongly anisophyllous. Stems 4-angled, sometimes zigzag, gla-

400ACANTHACEAEbrous or sparsely hirsute. Leaves sessile; leaf blade narrowly tobroadly oblong-elliptic or oblong-oblanceolate, rarely ovate forsmall leaves, 2–20 × 1.5–6 cm, both surfaces sparsely pilose,abaxially pale green or reddish when young, adaxially green,secondary veins 12–15 on each side of midvein, base attenuate,cuneate, rounded, or auriculate, margin serrate to undulate, apexacuminate. Inflorescences axillary or terminal, spikes, 3–10cm, sometimes branched; peduncle 2–4 cm, 4-angled, sulcate,bracteate; sterile bracts ovate, 0.8–2 × ca. 1.5 cm; floral bractsimbricate, broadly ovate to obovate-spatulate, 4–9 mm, oftenbecoming recurved, persistent, densely villous and ciliate, apexusually apiculate but sometimes rounded or emarginate; bracteolesabsent. Calyx 6–12 mm, gland-tipped pilose, 5-lobed almostto base; lobes linear, unequal with 2 slightly shorter thanothers, margin usually ciliate, apex subacute. Corolla pale purpleto violet, funnel-shaped, 2.5–3 cm, curved, outside glandtippedpilose on lobes, inside glabrous; tube basally cylindricand narrow for ca. 4 mm then weakly to strongly bent, abruptlyinflated, and widened to ca. 2.5 cm at mouth; limb weakly 2-lipped, lower lip 3-lobed, upper lip 2-lobed; lobes ca. 3 mm.Stamens 4, included; filaments glabrous, shorter pair ca. 2 mm,longer pair ca. 4 mm; anther thecae oblong, ca. 1.5 × 1 mm;pollen type 3. Ovary glabrous; style ca. 3.2 cm. Capsule narrowlyoblong-obovoid, 8–10 × ca. 2 mm, glabrous, 4-seeded,apex apiculate. Seeds ovate-suborbicular in outline, ca. 2 × 1.8mm, pubescent; areola very small. Fl. Sep–Jan, fr. Jan–Feb.Valleys; 300–1500 m. Guangxi, Yunnan [Bangladesh, India, Malaysia(Langkawi Island), Myanmar, Nepal, Pakistan, Thailand].Strobilanthes auriculata var. dyeriana (Masters) J. R. I. Woodwith leaf blade abaxially reddish rather than green is from Myanmarand Vietnam and is grown as a cultivated plant in Guangdong andYunnan.Strobilanthes auriculata is one of the few species of Strobilanthesthat can be recognized in a sterile state with confidence on account of itssessile, sometimes auriculate, oblong to oblanceolate leaves. It is knownto be pliestesial in part of its range.36. Strobilanthes mogokensis Lace, Bull. Misc. Inform. Kew1915: 406. 1915.卵叶马蓝 luan ye ma lanSubshrubs to 60 cm tall. Stems 4-angled, sulcate, whitegland-tipped pubescent, glabrescent. Petiole 0.6–1.5 cm, sulcate,pubescent; leaf blade variable in size and shape, basal onesbroadly ovate and 4–9 × 2–5 cm, apicalmost ones cordate and3–6 × 2–4.5 cm, both surfaces pubescent, secondary veins 4–6on each side of midvein, base truncate to narrowly cuneate andslightly decurrent onto petiole, margin crenulate-serrate, apexacuminate to obtuse. Inflorescences terminal, spikes, 3–5 cm;rachis gland-tipped pubescent; bracts lanceolate-oblong, 11–12× ca. 4 mm, abaxially yellowish gland-tipped pubescent, adaxiallygland-tipped pubescent; bracteoles narrowly oblong-elliptic,7–8 × ca. 1 mm, abaxially glabrous, adaxially gland-tippedpubescent. Calyx ca. 1 cm, outside pubescent, inside glandtippedpubescent, 2-lipped, lower lip 2-lobed almost to base,upper lip 3-lobed to middle; lobes linear, ca. 1 mm wide. Corollacolor not recorded, 2.5–3 cm, ventricose, curved, outsidepubescent, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for ca. 8 mm thengradually widened to ca. 1 cm at mouth; lobes ovate, ca. 4 × 4mm. Stamens 4, included; filaments glabrous, shorter pair ca. 3mm, longer pair ca. 7 mm; anther thecae oblong, ca. 2 × 1.2mm, erect; pollen type 9. Ovary white pilose at tip; style ca. 2.5cm, apically hirsute. Capsule ca. 1 cm, pilose at tip, 4-seeded.Seeds ovate in outline, ca. 2.5 × 2 mm, compressed, denselypilose. Fl. Oct–Dec, fr. Jan–Feb.Streamsides; 600–800 m. Yunnan (Ruili) [Myanmar].37. Strobilanthes argentea J. B. Imlay, Bull. Misc. Inform.Kew 1939: 121. 1939.银毛马蓝 yin mao ma lanSubshrubs to 50 cm tall, anisophyllous. Stems somewhatsulcate, bifariously pilose with large-celled trichomes, eventuallyglabrescent, basally woody. Petiole 0.2–4.3 cm, pilose; leafblade ovate, 4–12 × 2–6 cm, smaller of pair ca. 1/2 size oflarger one, both surfaces thinly or rather densely pilose withlarge white trichomes especially along veins, abaxially palerand with prominent cystoliths, base attenuate and ± decurrentonto petiole, margin crenulate to dentate, apex acute to shortlyacuminate and commonly falcate. Inflorescences terminal,spikes, simple or branched, 1–4 cm, 2–10-flowered; peduncle1–2.5 cm, pilose; rachis pilose; bracts oblong-lanceolate butbasal ones spatulate, 7–10 × 1.5–3.5 mm, densely pilose withgland-tipped and non-glandular trichomes, apex acute; bracteolesoblong-lanceolate, 6–7 × 1.5–2 mm, densely pilose withgland-tipped trichomes, apex acute. Flowers paired, basallydistant on rachis but apically becoming confluent. Calyx subequally5-lobed almost to base; lobes lanceolate, abaxially pilosewith gland-tipped and non-glandular trichomes, apex acute;central upper lobe longer than laterals. Corolla blue, 2.3–3.5cm, ventricose and strongly bent, outside pilose, inside glabrousexcept for trichomes retaining style; tube basally cylindric andca. 2 mm wide for ca. 1 cm then gradually widened to 7–8 mmat mouth; lobes broadly ovate, ca. 4 × 4 mm, apex rounded. Stamens4, included; filaments glabrous, shorter pair 2–3 mm,longer pair 5–7 mm; anther thecae oblong, ca. 2 × 1 mm. Ovarypilose at tip; style ca. 2.5 cm, pilose. Capsule not seen. Fl. Oct–Nov.Partly shaded and moist places along streams; 1100–1700 m.Yunnan [Thailand].38. Strobilanthes aprica (Hance) T. Anderson in Bentham, Fl.Hongk. 262. 1861.山一笼鸡 shan yi long jiGutzlaffia aprica Hance, Hooker’s J. Bot. Kew Gard. Misc.1: 143. 1849; G. aprica var. glabra (J. B. Imlay) H. S. Lo; G.dielsiana (W. W. Smith) S. Moore; Phlebophyllum apricum(Hance) Bentham; Strobilanthes aprica var. glabra J. B. Imlay;S. cavaleriei H. Léveillé var. angustifolia H. Léveillé; S. dielsianaW. W. Smith; S. mairei H. Léveillé.Shrubs or perennial herbs, to 70 cm tall, isophyllous.Stems 4-angled, sulcate, retrorsely white hirsute. Petiole 5–10mm, shortly white pubescent; leaf blade elliptic to oblong-elliptic,2–7 × 1–3 cm, leathery, indumentum very variable, both surfacesusually scabrous-hispid but abaxially sometimes whitish

ACANTHACEAE 401silky tomentose and occasionally glabrous except for pubescentmidvein, secondary veins 5–7 on each side of midvein, basebroadly cuneate, margin entire and scabrid to thinly ciliate, apexacute. Inflorescences axillary or terminal, short dense subcapitatespikes, becoming clustered in apicalmost leaf axils; spikesusually subsessile but occasionally basal ones with a 0.5–3(–7)cm peduncle; bracts ovate-lanceolate, 13–15 × 3–5 mm, subleathery,abaxially and on margin with white multicellular trichomes,adaxially glabrous, apex long acuminate; bracteoleslinear, 10–11 × ca. 1 mm, both surfaces covered with whitecystoliths, abaxially and on margin sparsely pilose with multicellulartrichomes, adaxially glabrous. Calyx 1–1.3 cm, 5-lobedfor ca. 1/2 its length; lobes linear, lower 2 becoming distinct,upper 3 remaining connate, outside pubescent to subglabrous,inside glabrous, margin pilose. Corolla lilac with a white tube,funnel-shaped, 2.5–3.5 cm, outside glabrous, inside glabrousexcept for trichomes retaining style; tube basally cylindric andca. 1 mm wide for 1.3–1.5 cm then abruptly inflated to ca. 1 cmin diam.; lobes suborbicular, ca. 5 × 5 mm, subequal, apex obtuse.Stamens 2, exserted; filaments ca. 1 cm, glabrous; antherthecae ellipsoid, ca. 2.5 mm; pollen type 11. Ovary white, narrowlyoblong, pilose at tip; style ca. 3.5 cm, curved, pilose.Capsule oblong, 1–1.3 cm, glabrous except for a few trichomesat tip, 4-seeded. Seeds ovate in outline, ca. 2 mm, convex onboth surfaces, pubescent; areola large, glabrous. Fl. Aug–Jan, fr.Sep–Feb.Dry forests, montane thickets; below 2200 m. Guangdong, Guangxi,Guizhou, Jiangxi, Sichuan, Yunnan [Cambodia, Laos, Myanmar,Thailand, Vietnam].39. Strobilanthes dalzielii (W. W. Smith) Benoist in Lecomte,Fl. Indo-Chine 4: 679. 1935.曲枝马蓝 qu zhi ma lanAcanthopale dalzielii W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 11: 193. 1919; Championella dalzielii (W. W. Smith)Bremekamp; C. dalzielii var. glabra Benoist; Pteroptychiadalzielii (W. W. Smith) H. S. Lo; Strobilanthes dalzielii var.inaequalis Benoist.Subshrubs or perennial herbs, 40–100 cm tall, branched,strongly anisophyllous. Stems slender, erect, zigzag, slightlypubescent to subglabrous. Petiole 0–3 mm; leaf blade ovate toovate-lanceolate, smaller of pair 2–5 × 1–2 cm and larger one9–14 × 3–5 cm, abaxially glaucous and glabrous or sparsely pubescentalong veins, adaxially green, glabrous, and with prominentcystoliths, secondary veins 3–5 on each side of midvein,base rounded but cordate for apical leaves, margin serrate, apexacuminate to acute and sometimes falcate. Inflorescences axillaryor terminal, spikes, 2–5 cm, 2–5-flowered; rachis usuallyzigzag, sparsely white puberulent; bracts linear-lanceolate, ca. 3× 1 mm, glabrous, 1-veined; bracteoles linear, ca. 5 × 1 mm,glabrous or puberulent with gland-tipped trichomes. Flowersdistant on rachis, solitary, not paired. Calyx ca. 1 cm, denselywhite pilose at base and along midvein, 5-lobed almost to base;lobes linear, apex finely acuminate. Corolla purplish blue orwhite, 3.5–4.5 cm, curved, outside densely white pilose in bud,inside glabrous except for trichomes retaining style; tube basallycylindric and ca. 2 mm wide for ca. 1 cm then ventricoseand gradually widened to 1–1.5 cm at mouth; lobes suborbicular,ca. 8 mm in diam. Stamens 4, slightly exserted; filamentshirsute, shorter pair ca. 5 mm, longer pair ca. 8 mm; antherthecae oblong, ca. 2 mm; pollen type 13. Ovary glabrous; styleca. 3.2 cm, pilose. Capsule linear-oblong, 1.8–2.2 cm, compressed,glabrous, 4-seeded, apex apiculate. Seeds ovate in outline,ca. 2 × 2 mm, densely appressed tomentose; areola verysmall. Fl. Oct–Jan.By streams; 400–1200 m. Fujian, Guangdong, Guangxi, Guizhou,Hainan, Hubei, Hunan, Jiangxi, Taiwan, Yunnan [Laos, Thailand, Vietnam].The strongly anisophyllous subsessile leaves combined with thedensely pubescent buds make Strobilanthes dalzielii very distinct.Specimens misidentified as Diflugossa divaricata (Nees) Bremekamp(H. P. Tsui & C. C. Hu, FRPS 70: 172. 2002; C. C. Hu & X. P. Fu,Higher Pl. China 10: 377. 2004) belong here.40. Strobilanthes bantonensis Lindau, Bull. Herb. Boissier 5:650. 1897.桂越马蓝 gui yue ma lanShrubs of unknown size, strongly anisophyllous. Stemszigzag, glabrous. Petiole 2–5 mm; leaf blade ovate-elliptic,smaller of each pair deciduous, larger one 7–15 × 4–7 cm, glabrous,secondary veins 4 or 5 on each side of midvein, base cuneate,margin sinuate-dentate, apex acute. Inflorescences axillaryor terminal, spikes 3–13 cm, fertile apically only; bractsoblanceolate to obovate, ca. 2 × 1 mm, sparsely gland-tipped pilose,± persistent, apex obtuse to rounded; bracteoles linear-oblanceolate,ca. 5 mm, sparsely gland-tipped pilose. Flowers 2 or3 per spike, 1–1.4 cm apart, one fertile at each node alternately.Calyx 7–10 mm; lobes linear, 7–9 × ca. 1 mm, subequal, glabrousor becoming sparsely gland-tipped pilose, apex obtuse.Corolla purplish blue or white, ca. 3.2 cm, outside densely pubescentin bud, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 2 mm wide for ca. 8 mmthen ventricose and gradually widened to ca. 1.3 cm at mouth;lobes oblong, ca. 4 × 6 mm. Stamens 4, included; filaments ofshorter stamen pair ca. 2 mm and glabrous, longer pair ca. 5mm and sparsely hirsute; anther thecae oblong ca. 2 mm; pollentype 1. Ovary glabrous; style ca. 3 cm, sparsely hirsute. Capsulefusiform, 1.2–1.3 cm, glabrous, 4-seeded. Seeds ovate in outline,ca. 3 mm, appressed pubescent. Fr. Dec.Forests in limestone hills; 600–1500 m. Guangxi [Vietnam].Strobilanthes bantonensis is a little-known species with verydistinct pollen.41. Strobilanthes pateriformis Lindau, Bull. Herb. Boissier 5:653. 1897.翅枝马蓝 chi zhi ma lanPteracanthus alatiramosus (H. S. Lo & D. Fang) C. Y. Wu& C. C. Hu; Pteroptychia pateriformis (Lindau) Bremekamp;Strobilanthes alatiramosa H. S. Lo & D. Fang.Herbs to 50 cm tall, anisophyllous, drying blackish green.Stems somewhat zigzag, internodes winged, wing 1.5–4 mmwide. Petiole 1–3 cm, glabrous; leaf blade oblong-ovate to lanceolate,smaller of pair 2–8 × 1–3 cm and larger one 6–13(–17)

402ACANTHACEAE× 2.5–5.5 cm, glabrous, secondary veins 4–6 on each side ofmidvein, base cuneate and decurrent onto petiole, margin dentateto crenate, apex acute. Inflorescences axillary or terminal,spikes, 3–10 cm; bracts ovate to lanceolate, 2.5–5 mm, caducous,glabrous, apex acute to obtuse; bracteoles spatulate-linearto linear, 4–7 mm, caducous, glabrous. Flowers 5–10 mm aparton rachis, often one sterile per node. Calyx 6–12 mm, glabrous,subequally 5-lobed or with 2 lobes slightly longer; lobes lanceolate,ca. 10 × 1 mm but accrescent in fruit, apex acute. Corollapale purple, 3–4 cm, ventricose, outside glabrous, insideglabrous except for trichomes retaining style; tube basally cylindricand ca. 2 mm wide for ca. 5 mm then gradually widenedto ca. 1 cm at mouth; lobes ovate, 3–4 mm, subequal, apexemarginate. Stamens 4, included; shorter filament pair ca. 2mm, glabrous; longer filament pair ca. 5 mm, hirsute; antherthecae oblong, ca. 2.5 × 1 mm; pollen type 12. Ovary apex minutelypubescent; style ca. 2.5 cm. Capsule fusiform, ca. 1.4cm, glabrous ?except apically, 4-seeded. Seeds ovate in outline,ca. 3 × 2.5 mm, densely appressed pubescent. Fl. Sep–Nov, fr.Dec–Feb.Dense forests; 400–1700 m. Guangxi, Guizhou, Hainan, Sichuan,Yunnan [Indonesia, Laos, Thailand, Vietnam].42. Strobilanthes truncata D. Fang & H. S. Lo, Guihaia 17:31. 1997.截头马蓝 jie tou ma lanHerbs to 3 m tall, perennial, anisophyllous. Stems 4-angled, somewhat zigzag, sulcate, glabrous. Petiole 0.6–7 cm;leaf blade broadly ovate, ovate, ovate-elliptic, or rarely elliptic,smaller of pair 3.5–7 × 2–4 cm and larger one 10–19 × 4.5–11.5 cm, glabrous, densely covered with linear cystoliths especiallyalong abaxial veins, secondary veins 4–7 on each sideof midvein, reticulate veins conspicuous, base broadly cuneateand oblique, margin remotely dentate to rarely serrate, apexacute. Inflorescences axillary or terminal, spikes, 7–15(–26) cm,sometimes branched; peduncle 0.5–5.5 cm, glabrous; rachis 4-angled, strongly zigzag, sulcate, bifariously pubescent; bractselliptic, 5–7 × ca. 4 mm, caducous, glabrous, margin ciliate,apex truncate to retuse; bracteoles yellowish but apically black,linear-lanceolate, ca. 8 × 1.5 mm, persistent, abaxially sparselypuberulent, margin ciliate, apex truncate to rounded. Flowersdistant on rachis. Calyx yellowish with a black apex, 8–10 mm,sparsely pubescent, usually sparsely covered with cystoliths, 2-lipped, lower lip 2-lobed almost to base, upper lip 3-lobed; lobeslanceolate, 5–6 mm, margin apically ciliate. Corolla white, 3.5–4 cm, slightly curved, outside glabrous, inside glabrous exceptfor trichomes retaining style; tube basally cylindric and ca. 4mm wide for 1–1.5 cm then ± ventricose and gradually widenedto ca. 2 cm at mouth; lobes ovate, ca. 7 × 5 mm, apex obtuse.Stamens 4, included; shorter filament pair ca. 3 mm, glabrous;longer filament pair ca. 7 mm, sparsely hirsute; anther thecaeoblong, 3–4 × ca. 1.5 mm; pollen type 12. Ovary comose; styleca. 2.7 cm, sparsely hirsute. Capsule fusiform, ca. 1.7 cm, glabrousexcept at apex, 4-seeded. Seeds ovate in outline, ca. 3 ×2.5 mm, densely appressed pubescent. Fl. Oct–Dec, Apr.Forests on limestone hills; 900–1300 m. Guangxi (Napo) [Vietnam].Strobilanthes truncata is doubtfully distinct from S. pateriformis,differing only in its truncate bracts and unwinged stem.43. Strobilanthes rankanensis Hayata, Icon. Pl. Formosan. 9:84. 1920.兰嵌马蓝 lan qian ma lanParachampionella rankanensis (Hayata) Bremekamp.Herbs 15–30 cm tall, rarely branched, isophyllous. Stemsslender, 4-angled, procumbent, usually rooting at basal nodes,sulcate, bifariously pubescent. Petiole 0.5–1.5 cm, sulcate, bifariouslypubescent or subglabrous; leaf blade ovate to elliptic,(0.5–)1–3.5 × 0.5–1.8 cm, glabrous or sparsely hirsute alongveins, secondary veins 3 or 4 on each side of midvein, base cuneateto subrounded, margin serrate, apex acute. Flowers solitaryin leaf axils. Bracts absent; bracteoles linear-spatulate, 6–9× 0.7–0.9 mm, sparsely hirsute, apex obtuse. Calyx 6–9 mm,slightly hirsute, 2-lipped, lower lip 2-lobed almost to base, upperlip unequally 3-lobed; lobes linear, margin ciliate. Corollapurple with white stripes, funnel-shaped, ca. 2 cm, outside glabrous,inside glabrous except for a few trichomes at base offilaments and adaxially in throat; tube basally cylindric and ca.1.5 mm wide for ca. 2.5 mm then gradually widened to ca. 2 cmat mouth; lobes oblong, ca. 3 × 4 mm, subequal, apex emarginate.Stamens 4, longer pair slightly exserted; filaments glabrous,shorter pair ca. 3 mm, longer pair ca. 6 mm; antherthecae oblong, ca. 2 mm, basally divergent; pollen type 11.Ovary ellipsoid, glabrous; style ca. 2 cm, basally sparselygland-tipped pilose. Capsule fusiform, 1–1.2 cm, glabrous, 4-seeded. Seeds ovate in outline, ca. 2.5 × 1.5 mm, lenticular,densely covered with long appressed trichomes. Fl. May–Sep.● Moist places in ravines; 600–1700 m. Taiwan.Strobilanthes rankanensis is a distinctive, superficially isolatedspecies because of the unique combination of trailing habit, 2-lippedcalyx, and globose echinulate pollen.44. Strobilanthes longispica (H. P. Tsui) J. R. I. Wood & Y. F.Deng, Bot. J. Linn. Soc. 150: 383. 2006.长穗腺背蓝 chang sui xian bei lanAdenacanthus longispicus H. P. Tsui, Fl. Reipubl. PopularisSin. 70: 348. 2002; Strobilanthes tripartita J. R. I. Wood.Herbs to 1.2 m tall, isophyllous, drying blackish green.Stems 4-angled, sulcate, bifariously pubescent. Petiole (0.2–)1–2.8 cm, hirsute; leaf blade elliptic to long ovate, 4.5–13 × 1.4–5cm, both surfaces slightly strigose, adaxially densely coveredwith linear cystoliths, secondary veins 7 or 8 on each side ofmidvein, base attenuate, margin serrate, apex acuminate. Inflorescencesaxillary or terminal, spikes, 3–20 cm, with 2–7 flowerpairs; peduncle ca. 1 cm, pubescent; rachis pubescent withgland-tipped trichomes; bracts linear-lanceolate, 6–8 × 1–2 mm,persistent, abaxially pubescent with gland-tipped trichomes, 1-veined, apex acute; bracteoles linear, 4–6 × ca. 1 mm, pubescentwith gland-tipped trichomes, apex acute. Flowers 1.5–3.5cm apart on rachis, usually only one sterile per node. Calyx1.2–1.8 cm, outside pilose with gland-tipped trichomes, 2-lipped; lower lip 2-lobed almost to base, lobes lanceolate and10–16 × 1.2–1.5 mm; upper lip 3-lobed, lobes triangular and 4–

ACANTHACEAE 4036 × 1.2–1.5 mm. Corolla violet blue, ca. 3.5 cm, slightly curved,outside glabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and narrow for ca. 7 mm then graduallywidened to ca. 1 cm at mouth; lobes oblong-ovate, ca. 7 ×6 mm, apex emarginate. Stamens 4, included; filaments hirsute,shorter pair ca. 2.2 mm, longer pair ca. 5 mm; anther thecae oblong,ca. 2 mm; pollen type 3. Ovary ellipsoid, ca. 3 mm, glabrous;style ca. 1.8 cm, sparsely gland-tipped pubescent. Capsuleoblong, 1.5–1.8 cm, gland-tipped pubescent at tip, 4-seeded. Seeds ovate in outline, ca. 3 × 2 mm, convex, denselypilose; areola small, glabrous. Fl. Oct–Jan.Thickets on mountain slopes; 1300–1600 m. Yunnan (Gongshan)[Myanmar].Strobilanthes longispica is very close to S. dryadum, differingonly in its lanceolate rather than ovate-oblong bracts, unwinged inflorescencerachis, and less dense inflorescence indumentum.45. Strobilanthes dryadum Benoist, Bull. Mus. Natl. Hist.Nat. 28: 94. 1922.林马蓝 lin ma lanPteracanthus dryadum (Benoist) C. Y. Wu & C. C. Hu.Herbs to 1 m tall, much branched, nearly isophyllous.Stems 4-angled, sulcate, bifariously pubescent. Petiole 1–3 cm,sulcate, bifariously pubescent; leaf blade ovate to lanceolate, to25 × 8 cm, glabrous, abaxially pale, secondary veins 6 or 7 oneach side of midvein, base attenuate and decurrent onto petioleforming wings, margin crenate-dentate, apex acuminate. Inflorescencesterminal, spikes, 3–6 cm, with 2–6 flower pairs;rachis pubescent with gland-tipped trichomes, winged, wing ca.1 mm; bracts ovate-oblong, 4–7 × 1–2 mm, persistent, glandtippedpubescent, apex obtuse; bracteoles oblong, ca. 4 × 0.5mm, gland-tipped pubescent. Flowers distant on rachis, usuallyone sterile per node. Calyx 1–1.5 cm, accrescent in fruit, outsidegland-tipped pubescent, 2-lipped; lower lip 2-lobed almostto base; upper lip 3-lobed to below middle, lobes linear-oblongand 11–15 × 1–2 mm. Corolla blue, outside glabrous, insideglabrous except for trichomes retaining style; tube basally cylindricand ca. 2 mm wide for ca. 8 mm then gradually widenedto ca. 1 cm at mouth; lobes oblong, ca. 8 × 8 mm. Stamens 4,included; filaments hirsute, shorter pair ca. 4 mm, longer pairca. 6 mm; anther thecae oblong, ca. 2 mm; pollen type 4. Ovarypuberulent at tip; style ca. 2.2 cm, sparsely pilose; stigma linear,compressed. Capsule narrowly oblong-obovoid, 1–1.2 cm, apicallygland-tipped pubescent, 4-seeded. Seeds pilose; areolasmall. Fl. May, Sep–Nov, fr. Jan–Mar.● Moist places in forests; 1300–2500 m. Guangxi, Yunnan.46. Strobilanthes adpressa J. R. I. Wood, Kew Bull. 58: 110.2003.紧贴马蓝 jin tie ma lanSubshrubs to 2 m tall, anisophyllous. Stems somewhatsulcate, bifariously pilose with large-celled trichomes, eventuallyglabrescent, basally woody; nodes pustulose, pubescent.Leaves basally petiolate, apically sessile; petiole (0–)1–2.5 cm,sulcate, sparsely pilose; leaf blade ovate-elliptic, (4–)6–16 ×(1–)3.5–8.5 cm, smaller of pair ca. 2/3 size of larger one, thinlyleathery, both surfaces villous with large-celled trichomes especiallyon veins, secondary veins 9–13 on each side of midvein,curved, and netted near margin, base decurrent onto petioleor those apically on stem amplexicaul, margin shallowlyserrate, apex acuminate. Inflorescences terminal, spikes, 5–7cm; rachis 4-angled, gland-tipped pilose; bracts somewhat imbricate,oblong, 1.1–1.7 × 0.5–0.7 cm, gland-tipped pilose,tardily deciduous as flowers open; bracteoles linear-oblong, 9–15 × 1.2–2 mm, persistent, abaxially gland-tipped pilose, apexacute. Flowers arranged in pairs, erect, ± appressed to rachis.Calyx 1.2–1.6 cm, 2-lipped; lower lip 2-lobed almost to base;upper lip 3-lobed, lobes 12–14 × ca. 1.5 mm. Corolla yellow orblue, outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric, ca. 2.5 mm wide for ca.1.5 cm then gradually ventricose and widened to ca. 1.5 cm atmouth, strongly bent ca. 2.5 cm above base; lobes ovate, ca. 6 ×5 mm, apex obtuse. Stamens 4, included; filaments sparselypilose, shorter pair ca. 4 mm, longer pair ca. 8 mm; antherthecae oblong, ca. 4 × 1.5 mm. Ovary comose with shortlystipitate glands; style ca. 2.5 cm, pilose. Capsule not seen. Fl.Oct–Dec.Streamsides in forests; 1500–1800 m. Yunnan (Maguan) [Vietnam].47. Strobilanthes atroviridis Y. F. Deng & J. R. I. Wood, J.Trop. Subtrop. Bot. 18: 480. 2010.景东马蓝 jing dong ma lanSubshrubs to 3 m tall, isophyllous or slightly anisophyllous,blackish green when dry. Stems branched, 4-angled, sulcate,glabrous, densely covered with white prominent cystoliths;nodes pustulose, pubescent. Petiole 2–5 cm, bifariouslypubescent; leaf blade ovate-elliptic, 9–22 × 5.5–14 cm, abaxiallypaler green and glabrous, adaxially green, subglabrous withsparse large-celled trichomes, and sparsely covered with prominentcystoliths especially along veins, secondary veins 6–10 oneach side of midvein, prominent on both surfaces, curved, andnetted near margin, tertiary veins conspicuous, base decurrentonto petiole, margin crenulate, apex acute to long acuminate.Inflorescences axillary, shortly pedunculate spikes, 1–3(–6) cm,2(–8)-flowered but commonly reduced to a subcapitate flowerpair; rachis gland-tipped pubescent; bracts linear-lanceolate, 9–14 × 2–3 mm, abaxially gland-tipped pubescent, adaxially glabrous;bracteoles linear-lanceolate, 9–13 × ca. 2.5 mm, abaxiallygland-tipped pubescent, adaxially glabrous. Flowers paired.Calyx 1–1.5 cm, outside gland-tipped pubescent, inside glabrous,2-lipped; lower lip 2-lobed almost to base, lobes linearlanceolate,9–14 × 2–3 mm, and apex acute; upper lip with 3lanceolate 5–7 mm teeth connate for 4–7 mm. Corolla purplishblue, 3–4 cm, straight, outside glabrous, inside glabrous exceptfor trichomes retaining style; tube ca. 2 mm wide at base thengradually widened to ca. 2 cm at mouth; lobes ovate, ca. 1 × 1cm, apex emarginate. Stamens 4, included; filaments basallypubescent, shorter pair ca. 6 mm, longer pair ca. 8 mm; antherthecae oblong; pollen type 3. Ovary not examined; style ca. 2cm, glabrous. Capsule not seen. Fl. Apr, Jun, Nov.● Moist places in forests; 2100–2400 m. Yunnan (Jingdong).

404ACANTHACEAE48. Strobilanthes pteroclada Benoist, Bull. Mus. Natl. Hist.Nat. 28: 187. 1922.延苞马蓝 yan bao ma lanHu.Hymenochlaena pteroclada (Benoist) C. Y. Wu & C. C.Subshrubs or perennial herbs, to 1 m tall, slightly anisophyllous,becoming black when dry. Stems 4-angled, sulcate,glabrous or subglabrous. Petiole (0–)0.5–1.5(–4) cm; leaf bladelanceolate, 7–20 × 3–8 cm, glabrous, abaxially densely coveredwith white cystoliths along veins, adaxially cystoliths prominent,secondary veins 5–9 on each side of midvein, base cuneateand decurrent onto petiole, margin serrulate to crenate, apexacuminate. Inflorescences axillary or terminal, spikes, 5–12 cm;peduncle 2–4.5 cm, apically widened to ca. 1.6 cm to form baseof bract; bracts lanceolate to narrowly ovate, 1.5–2 × 0.8–1.6cm, glabrous or gland-tipped pubescent, pinnately veined, basegradually narrowed and decurrent onto peduncle, margin entireor slightly dentate, apex acute to acuminate; bracteoles narrowlylanceolate, 5–7 × ca. 0.8 mm, glabrous, apex acuminate.Flowers ca. 3 cm apart on rachis. Calyx 7–9 mm, glabrous,weakly 2-lipped; lower lip 2-lobed, lobes 6–8 mm; upper lip 3-lobed, lobes lanceolate, ca. 6 mm, and apex acuminate. Corollapurplish blue, 3–4 cm, outside glabrous, inside glabrous exceptfor trichomes retaining style; tube basally cylindric and ca. 2.5mm wide for ca. 1.4 cm and then gradually widened to ca. 1.8cm at mouth; lobes ovate, ca. 6 × 5 mm, subequal, apex obtuse.Stamens 4, included; filaments pilose at base, shorter pair ca. 4mm, longer pair ca. 7 mm; anther thecae oblong, ca. 2.5 × 1.5mm; pollen type 3. Ovary glabrous; style ca. 2.4 cm, sparselypubescent. Capsule fusiform, ca. 1.3 cm, glabrous, 4-seeded.Seeds white, ovate in outline, ca. 2 × 2 mm, densely appressedtomentose. Fl. Apr–May, fr. Jun–Jul.Thickets or forests by streams; 300–900 m. Guangxi, Guizhou[Vietnam].49. Strobilanthes ovata Y. F. Deng & J. R. I. Wood, Novon 20:143. 2010.滇西马蓝 dian xi ma lanHerbs 30–40 cm tall, perennial. Stems with a narrow wingto 0.8 mm wide on each side, densely covered with cystoliths,glabrous. Leaves basally petiolate and apically becoming sessile;petiole of basal leaves 3–5 mm, glabrous; leaf blade ovateelliptic,2–5 × 1–2.3 cm, both surfaces glabrous and withprominent cystoliths, abaxially paler and with prominent veinscovered by cystoliths, secondary veins 2–4 on each side of midvein,base decurrent onto petiole, margin crenate to crenate-serrate,apex acuminate. Inflorescences terminal on main stem oron branches arising from apicalmost leaf axils, spikes, 1–7 cm;rachis narrowly winged, pilose with multicellular trichomes andsome gland-tipped trichomes; bracts ovate, oblong-ovate, orrounded, 3–7 × 1–5 mm, pubescent; bracteoles oblanceolate, ca.5 × 1 mm, subglabrous to pilose with multicellular trichomesand some gland-tipped trichomes, apex rounded. Calyx subglabrousor gland-tipped pilose on margin, slightly 2-lipped; upperlip 8–10 mm, 3-lobed to 3 mm from base, lobes triangular and5–7 mm; lower lip 2-lobed almost to base, lobes linear-oblanceolateto subspatulate, 9–10 mm, and apex obtuse. Corollalight blue, 3–3.5 cm, gently curved, subventricose, outside glabrous,inside glabrous except for trichomes retaining style; tubebasally cylindric and ca. 2 mm wide for ca. 1 cm and thengradually widened to ca. 1.3 cm at mouth; lobes ovate, ca. 4 × 4mm, subequal. Stamens 4, included; shorter filament pair ca. 2mm, glabrous; longer filament pair ca. 5 mm, basally pilose;anther thecae oblong, ca. 2 × 1 mm; pollen type 3. Ovary oblong,ca. 2 mm, comose at tip; style ca. 2.2 cm, sparsely pilose.Capsule narrowly oblong-ellipsoid, ca. 12 × 2 mm, glabrousexcept for gland-tipped pubescent tip, 4-seeded. Seeds ovate tosuborbicular in outline, ca. 3 × 2 mm, pilose. Fl. Oct–Feb, fr.Dec–Mar.● Moist forests; 1600–2200 m. Yunnan.50. Strobilanthes tamburensis C. B. Clarke in J. D. Hooker,Fl. Brit. India 4: 454. 1884.毛冠马蓝 mao guan ma lanHerbs to 50 cm tall, perennial, slightly anisophyllous.Stems decumbent and basally rooting then ascending, 4-angled,sulcate, bifariously pilose. Leaves usually sessile or with apetiole to 3 mm; leaf blade ovate to suborbicular, 8–45 × 8–45cm, both surfaces glabrous or pilose, abaxially whitish, adaxiallygreen, secondary veins 5–7 on each side of midvein, baserounded to subcordate, margin entire or obscurely crenulateserrate,apex acute to obtuse. Inflorescences terminal, spikes, 3–6 cm; bracts orbicular to obovate, 5–8 × 5–8 mm, persistentuntil flowers fall, pilose; bracteoles obovate, ca. 3 × 3 mm,pilose. Flowers ca. 2.5 cm apart basally on rachis but imbricateapically. Calyx 6–10 mm, 5-lobed almost to base; lobes linearspatulate,margin ciliate. Corolla blue, 2.5–3 cm, outside pilose,inside glabrous except for trichomes retaining style; tube basallycylindric and ca. 3 mm wide for ca. 1 cm then bent to ca.90º and abruptly widened to 8–10 mm at mouth; lobes broadlyovate, ca. 5 × 6–7 mm. Stamens 4, included; filaments subglabrous,shorter pair ca. 4 mm, longer pair ca. 7 mm; antherthecae oblong, ca. 2 × 1 mm; pollen type 16. Ovary glandtipped;style ca. 2.5 cm, sparsely pilose, apically glabrous.Capsule ca. 1.2 cm, gland-tipped pilose, 4-seeded. Seeds orbicularin outline, ca. 3 × 3 mm, densely pilose; areola absent. Fl.Aug–Nov.Wet places in forests; 1200–2100 m. Xizang (Cona) [Bhutan,India, Nepal].51. Strobilanthes taoana Y. F. Deng & J. R. I. Wood, J. Trop.Subtrop. Bot. 18: 479. 2010.陶氏马蓝 tao shi ma lanSubshrubs, weakly anisophyllous, becoming black whendry. Stems somewhat sulcate, pilose with large-celled trichomes,eventually glabrescent, basally woody. Petiole 0–2mm, pilose; leaf blade ovate, 4–10 × 2–5 cm, smaller of pair ca.2/3 size of larger one, abaxially paler, glabrous except for a fewtrichomes on veins, and densely covered with prominent cystoliths,adaxially green with inconspicuous cystoliths and subglabrousexcept for a few scattered hispid trichomes and a strigoseor scurfy midvein, base slightly asymmetric, rounded to subcordate,and briefly decurrent onto petiole, margin serrate, apex

ACANTHACEAE 405acute to shortly acuminate and commonly falcate. Inflorescencesaxillary or terminal, spikes, 2–5 cm; rachis straight,pilose; sterile bracts leaflike, apically diminishing in size, subsessile,ovate, 1–4 × 0.7–2.2 cm, apex acuminate; floral bractsoblong-elliptic, 3–6 × 2–4 mm, glabrous, apex acuminate;bracteoles obovate to oblong-lanceolate, 3–5 × 1–2 mm, glabrous.Flowers in opposite pairs but often solitary. Calyx 9–16× 1–2 mm, subequally 5-lobed to ca. 1 mm from base; lobesalmost colorless, linear, glabrous, covered with cystoliths, apexobtuse. Corolla white, 2–2.5 cm, straight, outside glabrous, insideglabrous except for trichomes retaining style; tube basallyca. 2 mm wide then gradually widened to ca. 1.2 cm at mouth;lobes ovate, 4–6 mm, apex rounded. Stamens 4, included; filamentsglabrous, shorter pair ca. 2 mm, longer pair ca. 6 mm;anther thecae spherical, ca. 1 mm in diam., glabrous. Ovaryglabrous; style ca. 2 cm, glabrous. Capsule not seen. Fl. Jan.● Limestone hills; ca. 2200 m. Yunnan (Jinghong).52. Strobilanthes rubescens T. Anderson, J. Linn. Soc., Bot. 9:479. 1867.红色马蓝 hong se ma lanPteracanthus rubescens (T. Anderson) Bremekamp.Subshrubs 0.5–2 m tall, isophyllous. Stems somewhat sulcate,pilose with large-celled trichomes, eventually glabrescent,basally woody. Petiole to 2 mm but apically leaves becomingsessile, pilose; leaf blade ovate to suborbicular, 4–10 × 2–5 cm,smaller of pair ca. 2/3 size of larger one, abaxially paler andglabrous except for a few trichomes on veins and densely coveredwith prominent cystoliths, adaxially green with inconspicuouscystoliths and subglabrous except for a few scattered hispidtrichomes and a strigose or scurfy midvein, base rounded,cordate, or cuneate and decurrent onto petiole, margin crenulate-dentate,apex acute to shortly acuminate and commonly falcate.Inflorescences axillary, spikes, 2–5 cm, secund, sometimesreduced to solitary flowers in leaf axils; rachis straight or obscurelyzigzag, pilose; sterile bracts leaflike, sessile, ovate, 1–4× 0.7–2.2 cm, variable in development, glabrous, densely coveredwith cystoliths, apex acuminate; floral bracts ovate, 2–3 ×1–1.5 mm, persistent, glabrous, apex acuminate; bracteoles oblong-lanceolate,1.5–3 × 0.5–1 mm, caducous, glabrous. Flowersto 2 cm apart basally on rachis but internodes shorter apically.Calyx 0.9–1.6 cm, subequally 5-lobed to ca. 1 mm frombase; lobes almost colorless, linear, 1–2 mm wide, glabrous,covered with cystoliths, apex acute. Corolla white, 2–2.5 cm,straight, outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric and narrow for ca. 5 mmthen gradually widened to ca. 1.2 cm at mouth; lobes ovate, 4–6mm, apex rounded. Stamens 4, included; filaments glabrous,shorter pair ca. 2 mm, longer pair ca. 6 mm; anther thecaespherical, ca. 1 mm in diam., glabrous; pollen type 3, requiresconfirmation. Ovary glabrous; style ca. 2 cm, glabrous. Capsule1–1.2 cm, glabrous except hairy at tip, 4-seeded. Fl. Oct–May.Limestone hills; ca. 1900 m. Yunnan [Bhutan, India].53. Strobilanthes pinetorum W. W. Smith, Notes Roy. Bot.Gard. Edinburgh 10: 197. 1918.松林马蓝 song lin ma lanDiflugossa pinetorum (W. W. Smith) C. Y. Wu & C. C.Hu.Subshrubs to 1 m tall, much branched, anisophyllous.Stems erect, glabrous or subglabrous. Petiole (0–)1–2 cm, subglabrous,winged; leaf blade broadly ovate to suborbicular,smaller of pair 2–3 × 1–1.5 cm and larger one 6–10 × 3.5–5 cm,both surfaces with prominent cystoliths, abaxially olive-greenand glabrous, adaxially dark green and sparsely pilose but soonglabrescent, secondary veins 5 or 6 on each side of midvein,base subrounded to broadly cuneate and abruptly decurrent ontopetiole or cordate for apical leaves, margin serrate, apexlong acuminate. Inflorescences axillary or terminal, simple orbranched spikes, to 10 cm; sterile bracts leaflike, sessile, ovate,1–5 × 0.7–3.5 cm, base cordate, apex shortly acuminate; floralbracts ovate-oblong, 7–11 × 2–3 mm; bracteoles linear-lanceolate,ca. 6 × 1 mm, glabrous, apex obtuse. Calyx ca. 7 mm, glabrous,5-lobed almost to base; lobes linear-oblong, ca. 6 × 1mm, apex obtuse. Corolla whitish purple, ca. 4 cm, curved, ventricose,outside glabrous; tube basally cylindric and ca. 2 mmwide for ca. 1.3 cm then curved and gradually widened to ca.1.2 cm at mouth; lobes orbicular, ca. 4 × 4 mm. Stamens 4,included; filaments subglabrous, shorter pair ca. 4 mm, longerpair ca. 7 mm; anther thecae oblong, ca. 2 × 1.5 mm; pollentype 5. Ovary pilose at tip; style ca. 3.2 cm, apically sparselyhirsute. Capsule not seen. Fl. Jun.● Forests; ca. 2000 m. Yunnan (Tengchong).Strobilanthes pinetorum is only known from the type, which isdistinctive but very immature.54. Strobilanthes helicta T. Anderson, J. Linn. Soc., Bot. 9:479. 1867.曲序马蓝 qu xu ma lanAsystasia calycina Nees (1832), not Strobilanthes calycinaNees (1836); Echinacanthus calycinus (Nees) Nees; Pteracanthuscalycinus (Nees) Bremekamp.Subshrubs 50–100 cm tall, pliestesial, nearly isophyllous.Stems 4-angled, erect, sulcate, usually glabrous but sometimesapically brown pubescent. Petiole 0.2–5 cm, bifariously pubescent;leaf blade ovate-lanceolate to elliptic, 5–12 × 1–6 cm,both surfaces glabrous except for pubescence on margin nearbase, abaxially paler, adaxially dark green, secondary veins 6 or7 on each side of midvein and abaxially prominent, base attenuateand decurrent onto petiole, margin serrate to rarely subentire,apex acuminate. Inflorescences axillary, spikes, 2–12 cm,secund, sometimes few flowered and frequently with bract pairsfertile only apically; rachis usually strongly zigzag; bracts linear,5–13 mm, persistent, one of pair per node shorter than other,glabrous or gland-tipped pilose; bracteoles linear, 2–3 mm, glabrousor gland-tipped pilose. Flowers 6–10 mm apart on rachis,only one sterile per node. Calyx 1.3–2.5 cm, glabrous or glandtippedpubescent, 5-lobed almost to base; lobes linear with oneshorter than others, often keeled, outside with white cystoliths,apex acuminate. Corolla white or flushed pale purple, 3.5–4 cm,outside glabrous, inside glabrous except for trichomes retainingstyle; tube basally narrow and cylindric for ca. 5 mm then

406ACANTHACEAEabruptly widened to ca. 1.8 cm and finally narrowed slightlyand bent to ca. 90° near mouth; lobes oblong, ca. 5 × 4 mm,unequal, apex emarginate. Stamens 4, included; filaments glabrous,shorter pair ca. 4 mm, longer pair ca. 6 mm; antherthecae oblong, ca. 3 × 1 mm; pollen type 4. Ovary glabrous;style 2.8–3.2 cm, apically hirsute. Capsule oblong, 1.8–2 cm,glabrous, 4-seeded, apex apiculate. Seeds ovate to obovate inoutline, ca. 4 × 2.5 mm, appressed pubescent. Fl. Sep–Nov, fr.Nov–Jan.Evergreen broad-leaved forests; 1700–2200 m. SE Xizang, Yunnan[Bhutan, India, Myanmar, Nepal].55. Strobilanthes rostrata Y. F. Deng & J. R. I. Wood, J. Trop.Subtrop. Bot. 18: 476. 2010.西畴马蓝 xi chou ma lanHerbs to 1 m tall, perennial, slightly anisophyllous. Stems4-angled, sulcate, glabrous. Petiole (1.5–)4–8 cm, glabrous; leafblade elliptic, (4.5–)11–17 × (2.5–)7–10 cm, both surfaces glabrous,abaxially paler, adaxially green and with numerouscystoliths, secondary veins (6–)9–12 on each side of midvein,base subrounded and decurrent onto petiole, margin dentate,apex acute. Inflorescences axillary, branched spikes, bearing apair of reduced leaves at branching point and forming a laxterminal panicle of spikes, ultimate branches slightly zigzag;bracts spatulate-linear, 2–3 mm, persistent, glabrous, both surfaceswith abundant white cystoliths; bracteoles linear, ca. 3mm. Flowers only one fertile per pair. Calyx ca. 1 cm, glabrous,outside densely covered with white cystoliths, inside with cystolithsin 2 lines along lobe margin, 5-lobed to ca. 2 mm frombase; lobes linear, equal. Corolla blue, ca. 3.8 cm, ventricose,outside glabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 2 mm wide for ca. 1.2 cm,bent to ca. 45° at middle, and gradually widened to ca. 1 cm atmouth. Stamens 4; filaments included, shorter pair ca. 6 mm,longer pair ca. 8 mm; anther thecae oblong, ca. 2 mm. Ovaryglabrous; style ca. 2.7 cm, pilose. Capsule ca. 1.6 cm, glabrous,4-seeded, apex with a ca. 4 mm beak. Seeds ovate in outline, ca.2 × 2 mm, appressed pilose; areola small. Fl. Aug–Sep, fr. Nov.● Forests by streams in limestone areas; 1400–1600 m. Yunnan(Xichou).56. Strobilanthes chrysodelta J. R. I. Wood, Kew Bull. 64: 41.2009.金三角马蓝 jin san jiao ma lanHerbs 30–60 cm tall, perennial, anisophyllous. Stemsslightly 4-angled, sulcate, glabrous. Petiole 0.5–2.5 cm, glabrous;leaf blade elliptic, 3–9 × 1.7–4 cm, smaller of pair ca. 2/3size of larger one but similar in shape, both surfaces glabrous,abaxially with prominent cystoliths, adaxially much paler, secondaryveins 4 on each side of midvein, base attenuate andslightly decurrent onto petiole, margin slightly serrate, apexacute. Inflorescences terminal, spikes, 2–10 cm, 2–10-flowered;rachis gland-tipped pilose; basal sterile bracts leaflike but sessileor almost so, ovate, base rounded; apical floral bracts ovateelliptic,5–6.5 × ca. 2 mm, persistent, gland-tipped pilose, apexobtuse; bracteoles oblong-oblanceolate, ca. 8 × 1 mm, persistentuntil after anthesis, gland-tipped pilose, apex obtuse. Flowersopposite pairs, 1–2 cm apart on rachis. Calyx 1–1.1 cm, accrescentto ca. 1.5 cm in fruits; lobes linear, gland-tipped pilose,apex obtuse; lower 2 lobes distinct to base; upper 3 lobes connatefor ca. 3 mm from base but becoming distinct in fruit, centrallobe slightly longer than others and becoming subspatulate.Corolla pale violet with a white tube, 3.7–4 cm, straight, outsideglabrous; tube basally cylindric and ca. 2.5 mm wide for ca. 1.1cm then gradually widened to ca. 1.2 cm at mouth; lobes ovate,9–10 × ca. 7 mm, apex rounded. Stamens 4, all fertile, included;filaments erect, very sparsely pilose, shorter pair ca. 2 mm,longer pair ca. 4 mm; anther thecae oblong, ca. 2 × 0.5 mm,erect, muticous. Ovary comose; style pilose. Capsule ca. 1.5cm, glabrous except for a few gland-tipped trichomes at apex,4-seeded. Seeds ca. 3 × 2.5 mm, lenticular, pilose; areola almostlacking. Fl. and fr. Apr.Moist places; 2100–2400 m. Yunnan [Myanmar].57. Strobilanthes tonkinensis Lindau, Bull. Herb. Boissier 5:651. 1897.糯米香 nuo mi xiangHerbs 50–100 cm tall, weakly anisophyllous. Stems 4-angled,pubescent, glabrescent, fragrant when dry. Petiole to 2 cm,pubescent; leaf blade broadly oblong-elliptic, to 18.5 × 6 cm,both surfaces sparsely pubescent especially on veins, adaxiallywith prominent cystoliths, secondary veins 4–7 on each side ofmidvein, base cuneate and decurrent onto petiole, margin crenate,apex acute. Inflorescences terminal, simple spikes, 3–11cm, sometimes with branches arising in apical leaf axils; rachispubescent with gland-tipped trichomes; bracts linear-spatulate,10–12 × 2–2.5 mm, sparsely pubescent and gland-tipped pubescent,densely covered with white cystoliths, 1-veined; bracteoleslinear, ca. 5 × 1 mm, pubescent, abaxially densely coveredwith cystoliths. Calyx 5-lobed to base; lobes linear, 8–9 × ca.1 mm, subequal, pubescent, outside covered with prominentcystoliths. Corolla pink to pure white, ca. 3.2 cm, straight, outsideglabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 2 mm wide for ca. 1 cmthen gradually widened to ca. 1 cm at mouth; lobes orbicular,ca. 5 mm in diam. Stamens 4, included; shorter filament pair ca.4 mm, glabrous; longer filament pair ca. 7 mm, puberulent; antherthecae oblong, ca. 3 mm; pollen type 4. Ovary obovoid, ca.3 mm, gland-tipped pilose at tip; style ca. 1.5 cm, glabrous.Capsule narrowly oblong-obovoid, 8–10 × ca. 2.5 mm, glandtippedpilose, 4-seeded, apex apiculate. Seeds ovate in outline,ca. 2 × 2 mm, pubescent; areola small, glabrous. Fl. Apr–Jun,Dec, fr. Jun–Jul.Moist forests; 200–1500 m. Guangxi, Yunnan [Thailand, Vietnam].Strobilanthes tonkinensis is a medicinal and aromatic plant.The name “Semnostachya menglaensis” (H. P. Tsui, FRPS 70:349. 2002) belongs here but was not validly published because theherbarium where the type specimen is conserved was not specified(Vienna Code, Art. 37.7).58. Strobilanthes stolonifera Benoist, Bull. Mus. Natl. Hist.Nat. 28: 98. 1922.匍枝马蓝 pu zhi ma lan

ACANTHACEAE 407Subshrubs to 20 cm tall, isophyllous. Stems ascending,usually rooting at nodes, glabrous but with cystoliths. Petiole 3–15(–20) mm, glabrous; leaf blade elliptic, (3–)5–10 × (1–)2–2.5cm, both surfaces with prominent cystoliths, abaxially palegreen to whitish and sparsely pubescent, adaxially pubescentbut soon glabrescent, secondary veins 5–7 on each side of midvein,base decurrent onto petiole, margin crenate to crenateserrate,apex acuminate. Inflorescences terminal, spikes, 1–2cm; rachis pilose; sterile bracts at base of inflorescence leaflike;floral bracts oblong to elliptic, 9–13 × 2–3 mm, pubescent;bracteoles oblong, ca. 8 × 1.5 mm, pilose with large-celled trichomes.Calyx lobes linear, 11–14 × ca. 1 mm, pubescent withsome trichomes gland-tipped. Corolla violet, 3–3.5 cm, subventricose,outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric and ca. 2 mm wide for ca.1 cm then slightly curved and gradually widened to ca. 1 cm atmouth; lobes ovate, ca. 4 × 4 mm. Stamens 4, included; filamentspilose, shorter pair ca. 3 mm, longer pair ca. 6 mm; antherthecae oblong, 2.5–3 × ca. 0.8 mm; pollen type 4, requiresconfirmation. Ovary oblong, 3–4 mm, puberulent at tip withsome gland-tipped trichomes; style ca. 3 cm, villous. Capsulenarrowly oblong, ca. 13 × 3 mm, pilose, 4-seeded. Seeds orbicularin outline, ca. 3 × 3 mm, villous. Fl. Jun.● Dry places; 1800–2400 m. Yunnan.59. Strobilanthes vallicola Y. F. Deng & J. R. I. Wood, Bot. J.Linn. Soc. 150: 380. 2006.河口马蓝 he kou ma lanHerbs 0.8–1.5 m tall, perennial, isophyllous. Stems black,4-angled, apically sulcate, glabrous. Petiole 1.3–2.5 cm, glabrous;leaf blade oblong-elliptic, 15–25 × 6.5–11 cm, both surfacesglabrous and with prominent cystoliths, abaxially palerand with prominent veins covered with cystoliths, secondaryveins 6–9 on each side of midvein, base cuneate, margin serrate,apex acuminate. Inflorescences terminal, spikes, 15–25cm, branched or rarely unbranched; rachis densely pilose withlarge-celled trichomes; bracts ovate-oblong, 8–12 × ca. 4 mm,persistent, pilose with large-celled trichomes and some glandtippedtrichomes; bracteoles lanceolate, 9–13 × 1.1–1.3 mm,persistent, pilose with large-celled trichomes and some glandtippedtrichomes. Flowers opposite pairs, sessile, 0.8–2 cmapart on rachis. Calyx 14–18 × 1–1.5 mm, persistent, pilosewith large-celled trichomes, some trichomes gland-tipped. Corollawhite, 3–3.5 cm, outside glabrous except for lobe margin,inside glabrous except for trichomes retaining style; tube basallycylindric and narrow for 1–1.5 cm then gradually ventricoseand strongly bent 1.5–2.5 cm from base; lobes ovate, ca. 4× 5 mm, apex obtuse. Stamens 4, included; filaments glabrous,shorter pairs 2–3 mm, longer pairs 3–5 mm; anther thecae oblong,3–4 × ca. 1 mm, glabrous; pollen type 9. Ovary ca. 3 × 1mm, glabrous; style ca. 2.5 cm. Capsule oblong, 1.5–2 × 0.2–0.3 cm, narrowed at both ends, comose, 4-seeded. Seeds ovateto suborbicular in outline, ca. 3 × 3 mm, pilose. Fl. Apr–Jun.● Moist places, stream and riversides in moist forests; below 200m. Yunnan (Hekou).60. Strobilanthes cognata Benoist, Bull. Mus. Natl. Hist. Nat.28: 189. 1922.奇瓣马蓝 qi ban ma lanPteracanthus cognatus (Benoist) C. Y. Wu & C. C. Hu.Herbs to 50 cm tall, isophyllous, drying blackish. Stemsstout, 4-angled, lenticellate, glabrous. Petiole ca. 1 cm, glabrous;leaf blade oblong to lanceolate, 6–12 × 3–4.5 cm, bothsurfaces densely covered with small cystoliths, very sparsely pilosewith scattered large-celled trichomes, secondary veins 8–10 on each side of midvein, base cuneate, margin crenate, apexacuminate. Inflorescences terminal, spikes, 3–5 cm; bracts lanceolate,7–20 × ca. 6 mm, sparsely pilose with large-celledwhite trichomes, margin ciliate, apex acuminate; bracteoles oblong,ca. 11 × 2 mm, gland-tipped pilose, apex acute. Flowersimbricate on rachis. Calyx 1–1.2 cm, outside gland-tipped pilose,5-lobed almost to base; lobes linear, ca. 1 mm wide, apexacute. Corolla violet, 4–5 cm, ventricose, outside glabrous, insideglabrous except for trichomes retaining style; tube basallycylindric and ca. 3 mm wide for ca. 2.5 cm then graduallywidened to ca. 1.5 cm at mouth; lobes oblong, ca. 6 × 5 mm,apex obtuse. Stamens 4, included; filaments erect, shorter pairca. 6 mm, longer pair ca. 8 mm; anther thecae oblong, ca. 3 × 1mm; pollen type 4. Ovary glabrous except pilose at tip; style3.5–4.2 cm. Capsule ca. 1.8 cm, pilose at tip, 4-seeded. Seedsovate in outline, ca. 3 × 2.5 mm, densely appressed pubescent.Fl. Aug–Oct, fr. Nov–Dec.● Moist places by streams; 1200–1400 m. Guizhou, Hubei, Hunan.61. Strobilanthes cusia (Nees) Kuntze, Revis. Gen. Pl. 2: 499.1891.板蓝 ban lanGoldfussia cusia Nees in Wallich, Pl. Asiat. Rar. 3: 88.1832; Baphicacanthus cusia (Nees) Bremekamp; Dipteracanthuscalycinus Champion; Ruellia indigofera Griffith; R. indigoticaFortune; Strobilanthes balansae Lindau; S. championii T.Anderson; S. flaccidifolia Nees.Herbs 0.5–1.5 m tall, erect, branched, drying blackish, isophyllousto weakly anisophyllous. Stems glabrous or minutelybrown puberulent. Petiole 0.5–7 cm; leaf blade elliptic to ovate,4–20 × 2–9 cm, both surfaces glabrous or abaxially minutelypuberulent along veins, abaxially paler green, adaxially darkgreen, secondary veins 7–9 on each side of midvein, base attenuate,margin serrate, apex acute. Inflorescences terminal or axillary,bracteate spikes, 1–6 cm, often aggregated to form a leafybranched panicle; peduncle 1–12 cm; bracts leaflike, petiolate,oblanceolate, obovate, or spatulate, 1.2–2.5 cm, basally usuallysterile; bracteoles linear-oblanceolate, 2–3 mm, deciduous beforebracts. Calyx 0.8–1.5 mm in flower, accrescent to ca. 2.5cm in fruit, minutely puberulent, 5-lobed almost to base; 4 lobeslinear-lanceolate, apex acute to obtuse; 1 lobe oblanceolate andmuch longer. Corolla blue, 3.5–5 cm, straight to slightly bent,outside glabrous; tube basally cylindric and ca. 3 mm widefor 1–1.5 cm then slightly curved and gradually widened to ca.1.5 cm at mouth; lobes oblong, ca. 9 × 9 mm, subequal. Stamens4, included; filaments glabrous, shorter pair ca. 3 mm,longer pair ca. 7 mm; anther thecae oblong, ca. 3 mm; pollentype 4. Ovary oblong, apex puberulent with few gland-tipped

408ACANTHACEAEtrichomes; style ca. 3.2 cm, glabrous. Capsule 1.5–2.2 cm, glabrous,4-seeded. Seeds ovate in outline, ca. 3.5 mm, coveredwith appressed trichomes; areola small. Fl. Jul–Feb, fr. Dec–Feb. 2n = 16, 32.Usually in moist wooded places, sometimes cultivated; 100–2000m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hunan, Sichuan,Taiwan, SE Xizang, Yunnan, Zhejiang [Bangladesh, Bhutan, India,Laos, Myanmar, Thailand, Vietnam].Strobilanthes cusia is reported to flower irregularly. The species isperhaps pliestesial.In at least part of its stated range Strobilanthes cusia may becultivated rather than native.This is a medicinal and dye plant and is the source of “Assamindigo.”62. Strobilanthes flexicaulis Hayata, Icon. Pl. Formosan. 5:135. 1915.曲茎兰嵌马蓝 qu jing lan qian ma lanParachampionella flexicaulis (Hayata) C. F. Hsieh & T. C.Huang; Strobilanthes fauriei Benoist; S. glandulifera Hayata; S.prionophylla Hayata; Triaenacanthus flexicaulis (Hayata) C. F.Hsieh & T. C. Huang.Subshrubs to 1 m tall, much branched, anisophyllous.Stems 4-angled, winged, glabrous. Petiole 0.2–2.5 cm, glabrousor sparsely pubescent; leaf blade ovate, ovate-lanceolate, orovate-elliptic, 3.5–18 × 1.8–5 cm, abaxially glaucous, adaxiallysparsely pilose to subglabrous, secondary veins 4–6 on eachside of midvein, base narrowly cuneate and decurrent onto petiole,margin serrate to crenate, apex acuminate. Inflorescencesaxillary or terminal, spikes, ca. 3 cm, few flowered; rachis oftenzigzag, glabrous to gland-tipped pilose; bracts persistent, basallymerging into leaves, ovate to suborbicular, and shortly petiolate,apically becoming ovate to oblanceolate and sessile, 4–15× 2–10 mm, apically diminishing in size, glabrous, apex obtuse;bracteoles linear, ca. 4 × 0.5 mm at anthesis but accrescent toca. 14 × 1 mm, glabrous or sparsely gland-tipped pilose. Pedicelto 1 mm or flowers sessile. Calyx 0.9–1.1 cm at anthesis,strongly accrescent to ca. 3.2 cm in fruit, 2-lipped; lower lip 2-lobed almost to base, lobes linear; upper lip 3-lobed to middle,lobes linear, ca. 6 × 1 mm at anthesis, margin ciliate with glandtippedtrichomes but soon glabrescent, and apex obtuse. Corollablue to purple, campanulate, ca. 4 cm, outside glabrous or glabrescent,inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 4 mm wide for ca. 1.5 cm thengradually widened to ca. 1.5 cm at mouth; limb slightly 2-lipped; lower lip ca. 1 cm, 3-lobed, center lobe orbicular, ca. 6 ×7 mm, base contracted, and apex emarginate, lateral lobes orbicular,somewhat oblique, and apex emarginate; upper lip 2-lobed, lobes semiorbicular and ca. 5 × 8 mm. Stamens 4;shorter filament pair ca. 2 mm, glabrous; longer filament pairca. 7 mm, hirsute; anther thecae oblong-linear, ca. 3.5 × 1 mm;pollen type 3. Ovary obovoid, glabrous; style filiform, ca. 3 cm,pubescent with gland-tipped trichomes; stigma subulate. Capsulelinear-oblong, 1.6–2 cm, glabrous, 4-seeded. Seeds oblong-ovatein outline, ca. 3.5 × 2 mm, densely covered withappressed trichomes. Fl. Sep–Nov, fr. Feb. 2n = 30.Streamsides in forests; 200–2300 m. Taiwan [Japan (RyukyuIslands)].Strobilanthes tashiroi Hayata (S. flexicaulis var. tashiroi (Hayata)Yamazaki) is endemic to the Ryukyu Islands (K. Iwatsuki et al., eds.,Fl. Japan 3a: 391. 1993).63. Strobilanthes lanyuensis Seok, C. F. Hsieh & J. Murata, J.Jap. Bot. 79: 151. 2004.兰屿马蓝 lan yu ma lanSubshrubs to 1.5 m tall, anisophyllous. Stems 4-angled,retrorsely pubescent. Petiole 0.5–5 cm, appressed pubescent;leaf blade variable in shape and size, oblong-lanceolate, oblongovate,or ovate, 3–20 × 2–8 cm, abaxially glabrous or sometimespubescent along veins, adaxially glabrous and with numerouscystoliths, secondary veins 3–5 on each side of midveinand abaxially prominent, base cuneate to narrowly cuneate anddecurrent onto petiole, margin mucronate-serrate, apex acuteto acuminate. Inflorescences terminal or from apical leaf axils,spikes, 5–10 cm; rachis 4-angled, glabrous or gland-tipped pubescent;bracts obovate, narrowly oblong, or linear-oblanceolate,1–2.5 × 0.3–2 cm, abaxially glabrous or sparsely pubescentand with numerous cystoliths, apex obtuse; bracteoles linear,10–20 × 3–4 mm, abaxially glabrous or sparsely pubescent. Calyx1.3–2.4 cm, accrescent to ca. 4 cm in fruit, 2-lipped, lowerlip 2-lobed almost to base, upper lip 3-lobed to 2/3 of length;lobes linear, glabrous, pubescent, or rarely gland-tipped piloseon margin, cystoliths numerous, apex acute. Corolla white, funnel-shaped,ca. 4 cm, bent near mouth, outside glabrous, insideglabrous except for trichomes retaining style; tube basally cylindricand ca. 3 mm wide for ca. 8 mm then gradually widenedto ca. 1.5 cm at mouth; lobes oblong, ca. 5 × 4 mm. Stamens4, included; filaments densely pilose, shorter pairs ca. 2 mm,longer pair ca. 2.5 mm; anther thecae oblong, ca. 2 mm; pollentype 3. Ovary glabrous; style ca. 2.5 cm; stigma very unequally2-lobed. Capsule cylindric-fusiform, ca. 1.7 cm, glabrous, 4-seeded. Seeds ovate in outline, ca. 3 × 3 mm, densely appressedpubescent. Fl. Sep–Nov, fr. Jan.● Moist places in forests; 200–1000 m. Taiwan (Lan Yu).Strobilanthes lanyuensis is close to S. flexicaulis, itself a variablespecies, differing only in the white, resupinate corolla and a few otherminor characters. Further studies are needed to confirm its status.64. Strobilanthes atropurpurea Nees in Wallich, Pl. Asiat.Rar. 3: 86. 1832.翅柄马蓝 chi bing ma lanHerbs 30–50(–100) cm tall, perennial, much branched,isophyllous or slightly anisophyllous. Stems slender, 4-angled,rooting at nodes, glabrous or pubescent. Leaves basally petiolate,apically sessile; petiole 0–2 cm, winged, glabrous or pubescent;leaf blade ovate, elliptic, or narrowly lanceolate, 1–6(–16) × 0.5–3.5(–6.5) cm, puberulent or glabrous, adaxially withlinear cystoliths, secondary veins 5 or 6 on each side of midvein,base rounded to attenuate and shortly decurrent onto petiole,margin serrate to coarsely crenate, apex acute to acuminate.Inflorescences axillary, spikes, 2–7 cm, zigzag, secund, fewflowered, often much reduced to flowers solitary or paired inleaf axil on main stem; bracts leaflike, ovate to subcordate, 8–

ACANTHACEAE 40910 × ca. 3 mm, apically smaller, persistent, glabrous, 3-veinedor pinnately veined; bracteoles linear-oblong, ca. 5 × 2 mm. Calyx1–1.5 cm, accrescent to 2–2.5 cm in fruit, 5-lobed almost tobase; lobes linear, 11–20 × 1–1.5 mm, unequal with one slightlylarger than others, glabrous or pilose, rarely gland-tipped pubescent,abaxially with linear cystoliths. Corolla pale purple, bluishpurple, or rarely white, 2.5–3.5 cm, outside glabrous, insideglabrous except for trichomes retaining style; tube basally cylindricand ca. 2 mm wide for ca. 1 cm then inflated, widened to1.5–1.8 cm, and bent at mouth; lobes oblong, ca. 3 × 4 mm,apex obtuse. Stamens 4, included; filaments glabrous, shorterpair ca. 4 mm, longer pair ca. 7 mm; anther thecae oblong, ca. 2mm; pollen type 3. Ovary glabrous; style ca. 2 cm, glabrous;stigma acute. Capsule fusiform, 1.2–1.8 cm, glabrous, 4-seeded.Seeds ovate in outline, ca. 3 × 3 mm, pubescent with appressedtrichomes; areola 0.2–0.3 mm. Fl. Jun–Oct.Moist places on mountain slopes, forests, by rivers; 700–2900 m.Chongqing, Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan,Taiwan, Xizang, Yunnan, Zhejiang [Bhutan, India, Myanmar,Nepal, Pakistan, Vietnam].There are no reports of Strobilanthes atropurpurea being pliestesialflowering from China or the E Himalaya, although this is a wellknownphenomenon in this species in the W Himalaya.1a. Leaf blade ovate to ovate-elliptic,1.5–6(–16) × 1–3.5(–6.5) cm,glabrous or pubescent ..................... 64a. var. atropurpurea1b. Leaf blade lanceolate,1–3 × 0.5–1.2 cm, glabrous .............. 64b. var. stenophylla64a. Strobilanthes atropurpurea var. atropurpurea翅柄马蓝 ( 原变种 ) chi bing ma lan (yuan bian zhong)Hemigraphis cuneata S. Y. Hu; Pteracanthus alatus(Wallich ex Nees) Bremekamp; Ruellia alata Wallich ex Nees;Strobilanthes densa Benoist; S. wallichii Nees; S. wallichii var.microphylla Nees.Leaf blade ovate to ovate-elliptic, 1.5–6(–16) × 1–3.5(–6.5) cm, glabrous or pubescent. Fl. Jun–Sep. 2n = 32, 40.Moist places on mountain slopes, forests; 700–2900 m. Chongqing,Guangdong, Guangxi, Guizhou, Hubei, Hunan, Jiangxi, Sichuan,Taiwan, Xizang (Cona, Nyalam), Yunnan, Zhejiang [Bhutan, India,Myanmar, Nepal, Pakistan, Vietnam].The name “Strobilanthes triflora” (Y. C. Tang, Iconogr. Cormophyt.Sin. 4: 163. 1975) belongs to this variety but was not validly publishedbecause no Latin description or diagnosis was provided (ViennaCode, Art. 36.1).64b. Strobilanthes atropurpurea var. stenophylla (C. B.Clarke) Y. F. Deng & J. R. I. Wood, J. Trop. Subtrop. Bot. 18:482. 2010.镇宁马蓝 zhen ning ma lanStrobilanthes stenophylla C. B. Clarke in J. D. Hooker, Fl.Brit. India 4: 472. 1884 [“stenophyllus”]; S. martini H. Léveillé.Leaf blade lanceolate, 1–3 × 0.5–1.2 cm, glabrous. Fl. Oct.By rivers. Guizhou (Zhenning) [India].65. Strobilanthes latisepala Hemsley, J. Linn. Soc., Bot. 26:241. 1890.薄萼马蓝 bao e ma lanHerbs to 50 cm tall, perennial, isophyllous. Stems 4-angled,sulcate, glabrous. Petiole 1–2.5 cm, glabrous; leaf bladelanceolate to ovate-lanceolate, 2–6 × 1–2.5 cm, abaxially glabrousand paler, adaxially with a few scattered trichomes andabundant cystoliths, secondary veins 4–6 on each side of midvein,base cuneate and decurrent onto petiole, margin crenulate,apex acuminate. Inflorescences axillary, subcapitate, 1–2 cm,2–4-flowered; peduncle 0–2 cm; bracts leaflike, ovate, 6–9 × 3–5 mm, persistent, glabrous, margin subentire, apex acute; bracteoleslinear, 5–6 × ca. 2 mm, glabrous. Calyx 8–10 mm, 5-lobed almost to base; lobes ovate-elliptic, 6–10 × 3–5 mm,unequal with one larger than others, outside sparsely pilose,apex obtuse. Corolla pale blue, ca. 3.2 cm, slightly curved, outsideglabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 2 mm wide for ca. 8 mmthen gradually widened to ca. 1 cm at mouth; lobes ovate, ca. 4× 3 mm. Stamens 4; filaments glabrous, shorter pair ca. 4 mm,longer pair ca. 8 mm; anther thecae oblong, ca. 2 × 1.2 mm,erect; pollen type 3. Ovary glabrous; style ca. 2.7 cm, glabrous.Capsule not seen.● Forests. Hubei (Xingshan).Strobilanthes latisepala is only known from the type and is possiblyconspecific with the very variable S. atropurpurea, from which itcan be distinguished by its distinctive broad sepals.66. Strobilanthes tibetica J. R. I. Wood, Edinburgh J. Bot. 51:262. 1994.西藏马蓝 xi zang ma lanPteracanthus tibeticus (J. R. I. Wood) C. Y. Wu & C. C.Hu.Subshrubs to 1.5 m tall, gregarious, probably pliestesial,nearly isophyllous. Stems slightly 4-angled, glabrous or obscurelybifariously pubescent. Petiole 1–3 cm, sparsely pilose;leaf blade ovate to elliptic, 7–14 × 3–9 cm, abaxially paler andsubglabrous except for a few trichomes near base of midvein,adaxially green, subglabrous except for a few scattered shorthispid trichomes, strigose or scurfy along midvein, and withconspicuous cystoliths, secondary veins 5 or 6 on each side ofmidvein, base narrowly cuneate and decurrent nearly to petiolebase, margin strongly serrate, apex shortly acuminate andslightly falcate. Inflorescences axillary, spikes, 4–12 cm, secund,sometimes reduced to single axillary flowers; rachis finelypuberulent to gland-tipped pilose, straight or obscurely zigzag,winged, wings 0.2–1 mm wide; bracts glabrous; basal bractsleaflike, with a petiole to 4 mm, ovate, 1.5–2.5 cm, apex acuminate;apical bracts sessile, narrowly oblong-elliptic, 1–1.2 cm,apex acuminate; bracteoles oblong, 2.5–4 mm, apex obtuse.Flowers to 1.5 cm apart on rachis. Calyx 0.9–2 cm, apparentlynot accrescent, subequally 5-lobed to ca. 1 mm above base;lobes almost colorless, linear, glabrous or sparsely gland-tippedpilose, apex obtuse. Corolla violet, 3.2–4 cm, outside pubescentbut silky gray in bud, inside glabrous; tube strongly inflatedfrom near base then widened to ca. 1.5 cm at middle and finallybent to ca. 90° but not narrowed at mouth; lobes ovate, ca. 5 × 7mm, apex rounded to slightly emarginate. Stamens 4, included;filaments ca. 2.5 mm, all equal in length, glabrous; anther

410ACANTHACEAEthecae oblong, ca. 1.5 mm. Ovary glabrous; style ca. 2 cm, glabrousor with a few trichomes. Capsule oblong, 1.1–1.2 cm,glabrous, 4-seeded, apex apiculate. Seeds oblong in outline, ca.3 × 2 mm, covered with stiff white trichomes. Fl. Aug–Sep, fr.Oct–Nov.Tsuga forests; 2300–2700 m. Xizang (Zayü) [India].67. Strobilanthes pterygorrhachis C. B. Clarke, J. Linn. Soc.,Bot. 25: 54. 1889.翅轴马蓝 chi zhou ma lanSubshrubs to 3.8 m tall, vigorous, subglabrous. Stems andbranches weakly 4-angled. Petiole 2–9 cm, glabrous; leaf bladeovate, ca. 15 × 7.5 cm, glabrous, adaxially darker and withabundant cystoliths, base cuneate and shortly decurrent ontopetiole, margin serrate, apex acute to shortly acuminate. Inflorescencesterminal or axillary, spikes, secund; rachis 3–25 cm,slightly zigzag, strongly winged, glabrous; bracts at base ofspike leaflike, petiolate, lanceolate, to 2 × 0.7 cm but diminishingin size apically to 6 × 3 mm; bracteoles linear-oblanceolate,5–10 mm. Flowers mostly paired, to 4 cm apart basallyon rachis but somewhat confluent apically. Calyx 2.5–3.6 cm,subequally 5-lobed to near base; lobes narrowly lanceolate,2.2–3.2 × ca. 0.3 cm, margin minutely ciliolate, apex narrowedinto a finely acuminate or aristate point. Corolla yellow, ca.3.5 cm, outside pubescent, inside glabrous except for trichomesretaining style; tube short, basally cylindric then abruptlywidened and bent just below mouth. Filaments glabrous, shorterpair 3–4 mm, longer pair 5–6 mm; anther thecae oblong, 3–3.5× ca. 1 mm. Ovary comose; style ca. 3 cm, glabrous. Capsulecomose. Fl. Aug.Moist places in forests; ca. 1800 m. Xizang (Mêdog) [India].68. Strobilanthes urophylla Nees in A. Candolle, Prodr. 11:192. 1847.尾叶马蓝 wei ye ma lanPteracanthus urophyllus (Nees) Bremekamp; Strobilanthesurophylla var. sikkimensis C. B. Clarke.Subshrubs, much branched, glabrous, anisophyllous. Stems4-angled, glabrous. Petiole 0.8–1.6 cm; leaf blade narrowlyovate-elliptic, 5–14 × 1.5–5.5 cm, abaxially pale green, pubescentalong veins, and with inconspicuous cystoliths, adaxiallyyellowish green, secondary veins 6–9 on each side of midvein,base cuneate to broadly cuneate and decurrent onto petiole,margin remotely serrate, apex shortly acuminate to caudate. Inflorescencesaxillary, short spikes; bracts leaflike, petiolate,ovate-elliptic, 0.8–4.5 cm, very persistent, abaxially with conspicuouscystoliths, pinnately veined, margin subentire, apexobtuse; bracteoles petiolate, obovate-spatulate, ca. 4 mm, persistent,subglabrous, 3-veined. Flowers in opposite pairs, 1–1.5cm apart on rachis, in axils of leaflike bracts. Calyx 7–8 mm,pubescent, 5-lobed almost to base; lobes lanceolate with onelonger than others, accrescent in fruit, apex acuminate. Corollapale purple, 2–2.5 cm, ventricose, outside pubescent, insidesubglabrous except for trichomes retaining style; tube basallycylindric and ca. 6 mm wide for ca. 5 mm then slightly bent andregularly widened to ca. 1.2 cm at mouth; lobes oblong, ca. 5 ×4 mm, apex obtuse. Stamens 4, included; filaments hirsute,shorter pair ca. 4 mm, longer pair ca. 6 mm; anther thecae oblong,ca. 2.5 × 1.2 mm; pollen type 3. Ovary comose; style ca.1.7 cm, hirsute. Capsule ca. 1.4 cm, glabrous, 4-seeded. Seedsovate in outline, ca. 2.5 × 2 mm, pubescent; areola small. Fl.May–Jun, fr. Jul–Aug.Evergreen broad-leaved forests; 1200–2000 m. Xizang (Mêdog)[India].69. Strobilanthes inflata T. Anderson, J. Linn. Soc., Bot. 9:476. 1867.锡金马蓝 xi jin ma lanHerbs 50–100 cm tall, branched, isophyllous. Stemsstraight or somewhat zigzag, glabrous, slightly 4-angled, sulcate,pubescent or with dense brown large-celled trichomes.Petiole 3–3.5(–7.5) cm, subglabrous, pubescent, or densely yellowishbrown hirsute; leaf blade oblong-ovate, elliptic, or ovate,5–20 × 1–10 cm, abaxially hirsute, pubescent, or subglabrousand with conspicuous cystoliths, adaxially pubescent or glabrousexcept brownish and whitish tomentose along veins,sparsely setose, densely yellowish brown hirsute on midvein,and with numerous cystoliths, secondary veins 6–8 on each sideof midvein, base oblique and broadly rounded to cuneate anddecurrent onto petiole, margin serrate to crenate, apex acuminateto caudate. Inflorescences spikes, 2.5–12(–21) cm, secund,sometimes reduced to 1 or 2 flower pairs; peduncle 1.5–3 cm;rachis usually geniculate at first node then straight or somewhatzigzag; bracts variable in size and shape with basalmost onesleaflike, with a 3–4 mm petiole, ovate to narrowly ovate, 2–3 ×0.8–1.3 cm, and margin serrate; apical bracts petiolate, obovate,elliptic, or ovate-spatulate, 0.6–3 × 0.3–1.2 cm, persistent, pubescent,margin usually with 1 or 2 teeth; bracteoles oblong tolinear, 2–3.5 mm, apex rounded. Flowers 1–2 cm apart on rachis.Calyx 1.1–1.4 cm, white pilose with large-celled glandtippedtrichomes, 5-lobed to base; lobes linear, unequal, reflexed,glabrous in anthesis, sometimes gland-tipped pubescentin fruit, densely covered with white linear cystoliths, marginpaler, apex acute. Corolla pinkish blue to blue, 3.5–4 cm, outsidepubescent, inside glabrous except for trichomes retainingstyle; tube basally cylindric and narrow for ca. 5 mm thenabruptly widened to ca. 1.3 cm at mouth, just basal to mouthslightly narrowed and bent to ca. 90°; lobes oblong, ca. 5 × 6mm. Stamens 4, included; filaments glabrous, shorter pair 3–4mm and incurved, longer pair 5–6 mm; anther thecae oblong,ca. 2 × 1 mm; pollen type 3. Ovary comose; style ca. 1.8 cm,sparsely gland-tipped pubescent at base. Capsule brownish yellow,narrowly ellipsoid, 1.6–2 cm, glabrous, 4-seeded. Seeds orbicularin outline, ca. 3 mm in diam., villous; areola inconspicuous.Fl. Jun–Nov.Evergreen broad-leaved forests, streamsides in shady places, forests,forest margins; 1700–3200 m. Xizang, Yunnan [Bhutan, India, Indonesia,Myanmar, Nepal].1a. Stems, petioles, and inflorescences denselycovered with long yellowish browntrichomes; leaf blade broadly ovate,12–20 × 6–10 cm ................................ 69c. var. aenobarba

ACANTHACEAE 4111b. Stems, petioles, and inflorescencessubglabrous or white pubescent; leafblade ovate-elliptic, less than 8 × 6 cm.2a. Leaf blade white pubescent ............... 69a. var. inflata2b. Leaf blade subglabrous ........ 69b. var. gongshanensis69a. Strobilanthes inflata var. inflata锡金马蓝 ( 原变种 ) xi jin ma lan (yuan bian zhong)Pteracanthus inflatus (T. Anderson) Bremekamp; Strobilantheswardii W. W. Smith.Stems, petioles, and inflorescences white pubescent. Leafblade elliptic to ovate, ca. 5.5 × 3 cm. Fl. Jun–Nov.Forests; ca. 1700 m. Xizang (Mêdog, Zayü), Yunnan [Bhutan,India, Myanmar, Nepal].69b. Strobilanthes inflata var. gongshanensis (H. P. Tsui) J. R.I. Wood & Y. F. Deng, Bot. J. Linn. Soc. 150: 387. 2006.贡山马蓝 gong shan ma lanPteracanthus gongshanensis H. P. Tsui, Acta Bot. Yunnan.12: 277. 1990; Strobilanthes unilateralis J. R. I. Wood.Stems, petioles, and inflorescences subglabrous. Leaf bladeovate-elliptic, 5–12 × 1–4.5 cm. Fl. Aug–Oct, fr. Dec.Evergreen broad-leaved forests; 1900–2200 m. Yunnan (Gongshan)[Indonesia, Myanmar].69c. Strobilanthes inflata var. aenobarba (W. W. Smith) J. R.I. Wood & Y. F. Deng, Bot. J. Linn. Soc. 150: 386. 2006.铜毛马蓝 tong mao ma lanStrobilanthes aenobarba W. W. Smith, Notes Roy. Bot.Gard. Edinburgh 13: 185. 1921; Pteracanthus aenobarbus (W.W. Smith) C. Y. Wu & C. C. Hu.Stems, petioles, and inflorescences densely covered withlong yellowish brown trichomes. Leaf blade broadly ovate, 12–20 × 6–10 cm. Fl. Aug–Oct.Streamsides in shady places, forest margins; 2300–3200 m. Xizang(Zayü), Yunnan [Indonesia, Myanmar].70. Strobilanthes congesta Terao, Notes Roy. Bot. Gard. Edinburgh40: 153. 1982.密序马蓝 mi xu ma lanPteracanthus congestus (Terao) C. Y. Wu & C. C. Hu.Subshrubs 0.6–1.8 m tall, anisophyllous. Stems subterete,whitish to brownish yellow pubescent with large-celled trichomes.Petiole 1–7 mm, densely hirsute; leaf blade ovate-lanceolate,2–9 × 0.8–3.5 cm, smaller of pair ca. 2/3 size of largerone, both surfaces brownish to yellow hirsute, secondary veins4–7 on each side of midvein and inconspicuous, base attenuate,oblique, and shortly decurrent onto petiole, margin crenate andciliate, apex acuminate and often falcate. Inflorescences axillaryor terminal, spicate with 1–4 small capitula borne singly alongrachis; capitula 2–8-flowered, to 4 cm apart; rachis 3–7 cm,often basally with a pair of sterile leaflike bracts so leavessometimes appearing verticillate, densely white or yellow pilose;bracts linear-oblanceolate, 3–8 × 0.8–1.5 mm, persistent,pilose, apex obtuse; bracteoles ca. 6 × 1 mm, linear, white pilose.Calyx 5-lobed almost to base; lobes ovate to lanceolate,pilose on both surfaces except basally inside, abruptly narrowedat middle with an obtuse often apically recurved cauda 11–14 ×1.5–5 mm; one lobe ca. 2 mm longer than others. Corolla palewhitish purple, ca. 2.5 cm, curved, outside pubescent; tube basallynarrow and cylindric for ca. 6 mm then abruptly inflatedand widened to ca. 1 cm at mouth; lobes oblong, ca. 4 × 3.5mm. Stamens 4, included; shorter filament pair ca. 3 mm, glabrous;longer filament pair 5–6 mm, base pilose; anther thecaeoblong, ca. 2 × 0.5 mm; pollen type 3. Ovary glabrous; style ca.1.8 cm, pilose and gland-tipped pubescent. Capsule oblong, ca.10 × 3 mm, glabrous, 4-seeded; seeds ovate in outline, ca. 3 ×2.5 mm, pubescent. Fl. Jul–Aug, fr. Dec.Thickets, tropical forests; 1600–1800 m. Yunnan [Myanmar,?Nepal].Strobilanthes congesta is a very distinct species because of theform of its inflorescence and unique calyx.71. Strobilanthes larium Handel-Mazzetti, Symb. Sin. 7: 893.1936.闭花马蓝 bi hua ma lanHerbs to 1.5 m tall, isophyllous. Branches 4-angled, sulcate,silky gland-tipped pubescent. Petiole 2–5 cm, sulcate, subglabrous;leaf blade ovate-elliptic, (5–)7–12 × (2–)3–7 cm,abaxially glabrous except pilose along midvein, glaucous, andwith numerous cystoliths, adaxially green and subglabrousexcept for few large-celled multicellular trichomes along veins,secondary veins 6–8 on each side of midvein, base cuneate anddecurrent onto petiole, margin dentate and ciliate, apex acuminateand falcate. Inflorescences axillary, spikes, (1–)3–10 cm;bracts oblong, 1.5–4 × 0.2–0.5 cm, variable in size and apicallydiminishing in size, abaxially densely covered with white largecelledmulticellular trichomes especially toward base, marginsubentire; bracteoles linear, ca. 4 × 1 mm, puberulent. Calyx 1–1.2 cm, 5-lobed almost to base; lobes unequal with one largerthan others, outside puberulent, margin large-celled multicellularpubescent with gland-tipped trichomes. Corolla purplish,curved, 3.5–4 cm, outside densely whitish pubescent whenyoung but glabrescent, inside densely pubescent when youngbut glabrescent except for trichomes below lobe bases; tubebasally cylindric and ca. 3 mm wide for ca. 5 mm then ventricose,bent to ca. 90°, and finally abruptly widened to ca. 1.3 cmat mouth; lobes oblong, ca. 5 × 4 mm, apex emarginate. Stamens4, included; filaments glabrous, shorter pair 3–4 mm,longer pair 6–7 mm; anther thecae oblong, ca. 3 × 1 mm. Ovaryglabrous; style 2.8–3 cm, apical part pilose. Capsule clavate, ca.1.5 cm, glabrous except for pilose apex, 4-seeded. Seeds ca. 2 ×1.5 mm, lenticular, densely covered with scalelike trichomes;areola ca. 0.3 mm. Fl. May–Sep.● By streams; 600–2700 m. Chongqing, Hubei, Hunan, Sichuan.72. Strobilanthes oresbia W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 10: 196. 1918.山马蓝 shan ma lan

412ACANTHACEAEGoldfussia grandissima H. P. Tsui; Pteracanthus grandissimus(H. P. Tsui) C. Y. Wu & C. C. Hu; P. oresbius (W. W.Smith) C. Y. Wu & C. C. Hu; Strobilanthes grandissima (H. P.Tsui) J. R. I. Wood.Herbs to 2 m tall, erect, nearly isophyllous. Stems 4-angled,densely white pubescent with gland-tipped trichomes, glabrescent.Petiole 2–13 cm, gland-tipped pubescent, glabrescent,apically winged; leaf blade ovate to orbicular, 7–22 × 4–18 cm,pubescent especially along veins, cystoliths abundant on bothsurfaces, secondary veins 9–11 on each side of midvein, basecuneate, rounded, or sometimes subcordate and decurrent ontopetiole, margin deeply dentate or serrate, apex acute to acuminate.Inflorescences axillary, spikes, to 15 cm, secund, sometimesreduced to single flowers; peduncle 0.5–7 cm, denselygland-tipped pubescent; rachis somewhat zigzag; bracts at baseof spike leaflike, petiolate, ovate, to 4 × 2 cm, gland-tipped pubescent,base cuneate, margin serrate, apex obtuse; bracts towardspike apex lanceolate, diminishing apically to ca. 10 × 1mm, margin entire, apex obtuse; bracteoles linear-oblanceolate,ca. 7 × 1 mm, gland-tipped pubescent, apex obtuse. Flowersusually 1 per node, 0.8–2.5 cm apart on rachis. Calyx 1.2–2.1cm, densely gland-tipped pubescent, 5-lobed almost to base;lobes linear-oblanceolate, apex obtuse; one lobe to 2 mm longerthan others. Corolla purplish blue, 4–5 cm, ventricose, outsidesparsely white pubescent, inside pubescent; tube basally cylindricand narrow for ca. 5 mm then bent to ca. 90°, strongly ventricose,and widened to ca. 1.5 cm at mouth; lobes oblong, ca. 6× 5 mm, apex obtuse. Stamens 4; filaments glabrous, shorterpair ca. 2.5 mm, longer pair ca. 4 mm; anther thecae oblong, ca.2.5 × 1 mm; pollen type 4. Ovary glabrous; style ca. 2.2 cm,shortly pilose. Capsule oblong, ca. 15 × 2–3 mm, glabrousexcept for very few stalked glands near apex, 4-seeded. Seedsblackish brown, elliptic in outline, 3–3.5 × ca. 2.5 mm, denselypilose; areola very small. Fl. Jul–Sep, fr. Oct–Dec.Moist alpine meadows and forests; 1600–3300 m. Chongqing, Sichuan,Xizang (Zayü), Yunnan [India, Myanmar].73. Strobilanthes flexa Benoist, Bull. Mus. Natl. Hist. Nat. 28:186. 1922.城口马蓝 cheng kou ma lanPteracanthus flexus (Benoist) C. Y. Wu & C. C. Hu.Herbs to 60 cm tall, isophyllous. Stems 4-angled, glabrescent.Petiole 1–5 cm, thinly pubescent; leaf blade ovate toelliptic, 3.5–15 × 3–7 cm, both surfaces glabrous except basallyon abaxial midvein, cystoliths apparently absent, secondaryveins 6–8 on each side of midvein, base cuneate and decurrentonto petiole, margin coarsely dentate or serrate, apex acute. Inflorescencesaxillary or terminal, spikes, 3–6 cm, lax, sometimesreduced to a single flower in axil of apical leaves; rachisgland-tipped pilose; bracts leaflike, oblong-oblanceolate, 1.5–2× 0.2–0.4 cm, apically diminishing in size, gland-tipped pubescent,apex acute to acuminate; bracteoles oblong, 7–8 × ca. 1mm, gland-tipped pubescent, apex obtuse. Flowers basally to 1cm apart on rachis but apically becoming congested. Calyx 0.8–1.2 cm, accrescent in fruit, gland-tipped pubescent, 5-lobed almostto base; lobes linear-oblong, 7–10 × ca. 1 mm, subequal,apex obtuse. Corolla purplish blue, 3.5–4 cm, outside pilose,inside glabrous except for trichomes retaining style; tube basallycylindric and ca. 2 mm wide for ca. 1 cm then abruptlyinflated, bent to ca. 90° above middle, and finally widened toca. 1.4 cm at mouth; lobes ovate, ca. 5 × 5 mm, apex rounded.Stamens 4, included; filaments glabrous, shorter pair ca. 2 mm,longer pair ca. 4 mm; anther thecae oblong, ca. 2 × 1 mm, erect;pollen type 4. Ovary comose; style ca. 3.2 cm, sparsely pilose.Capsule oblong, 1.5–1.8 cm, apically gland-tipped pubescent,4-seeded. Fl. Aug–Sep, fr. Oct–Dec.● Forests; 1400–3600 m. Chongqing, Guizhou, Hubei, Sichuan,Yunnan.Strobilanthes flexa appears to intergrade with S. versicolor andmay just be a more robust form.74. Strobilanthes versicolor Diels, Notes Roy. Bot. Gard.Edinburgh 5: 163. 1912.变色马蓝 bian se ma lanPteracanthus versicolor (Diels) H. W. Li.Herbs to 50(–100) cm tall, with several annual stemsarising from a perennial base, isophyllous. Stems sparsely todensely pubescent, glabrescent. Petiole (0–)1–3 cm, pubescent;leaf blade obovate to suborbicular, 2.5–6 × 1–3.5 cm, both surfacesthinly pilose especially on veins, densely covered withcystoliths, secondary veins 4–6 on each side of midvein, basebroadly cuneate and decurrent onto petiole, margin coarselyserrate, apex acute. Inflorescences terminal or axillary, spikes,4–15 cm, simple or sparingly branched; bracts leaflike, 1.2–3.5× 0.3–1 cm, persistent, apically diminishing in size, thinly pilose;basal bracts petiolate, suborbicular to oblong-elliptic, margindentate; apical bracts sessile, narrowly oblong-elliptic, marginentire; bracteoles linear-oblanceolate, 4–8 × 1–2 mm, pilose.Flower pairs 1–1.5 cm apart on rachis even apically. Calyxca. 1.5 cm, 5-lobed almost to base; lobes linear, 10–11 × ca. 1mm, subequal, pilose with large-celled trichomes, apex obtuse.Corolla white or blue, 3.5–4 cm, outside pilose, inside glabrousexcept for trichomes retaining style; tube basally cylindric andca. 2.5 mm wide for 1–1.5 cm then abruptly bent to ca. 90°above middle and finally widened to ca. 1.5 cm at mouth; lobesovate, ca. 6 × 5 mm, apex rounded. Stamens 4, included; filamentspilose, shorter pair ca. 5 mm, longer pair ca. 7 mm;anther thecae oblong, 2–2.5 × ca. 0.8 mm; pollen type 6. Ovarycomose; style ca. 2.8 cm, pilose. Capsule ca. 1.4 cm, glandtippedpubescent, 4-seeded. Seeds ovate in outline, ca. 3 × 3mm, appressed pubescent. Fl. May–Aug.● Grasslands at forest margins; 2800–3300 m. Sichuan, S Xizang,Yunnan.75. Strobilanthes forrestii Diels, Notes Roy. Bot. Gard. Edinburgh5: 162. 1912.腺毛马蓝 xian mao ma lanPteracanthus forrestii (Diels) H. P. Tsui; P. rotundifolius(Benoist) Bremekamp; Strobilanthes panpienkaiensis H. Léveillé;S. rotundifolia Benoist.Herbs to 30–75 cm tall, isophyllous. Stems 4-angled,thinly pilose with gland-tipped trichomes, glabrescent, basally

ACANTHACEAE 413woody. Leaves subsessile; leaf blade ovate to rarely ovate-oblong,2–5 × 1.2–3 cm, both surfaces thinly pilose, adaxiallywith cystoliths, secondary veins ca. 4 on each side of midvein,base rounded, cordate, or rarely cuneate, margin crenate to subentire,apex acute to obtuse. Inflorescences terminal or veryrarely also from apical leaf axils, spikes, 5–15 cm; rachis pilosewith gland-tipped trichomes; bracts leaflike, pilose with glandtippedtrichomes; basal bracts ovate-elliptic, 1–3 × 1–2 cm,margin obscurely crenate; apical bracts lanceolate, 7–10 × ca. 3mm, margin entire; bracteoles linear to narrowly oblong, 6–11 ×ca. 1 mm, pilose with gland-tipped trichomes. Flowers 1–2.5cm apart on rachis. Calyx 0.8–1.2 cm, 5-lobed almost to base;lobes linear with one ca. 2 mm longer than others, pilose withgland-tipped trichomes. Corolla blue to purple, 3.2–4 cm,curved, outside with a few trichomes, inside glabrous except fortrichomes retaining style; tube basally cylindric and ca. 2 mmwide for ca. 5 mm then bent to ca. 90° and widened to ca. 1.5cm at mouth; lobes ovate, ca. 6 × 7 mm, apex obtuse. Stamens4, included; shorter filament pair ca. 2 mm; longer filament pairca. 8 mm, shortly pubescent; anther thecae oblong, ca. 2 × 0.8mm; pollen type 3. Ovary gland-tipped puberulent at tip; styleca. 2.8 cm. Capsule oblong, ca. 1.2 cm, apically gland-tippedpilose, 4-seeded. Seeds dark brown, ovate in outline, ca. 3 mm,pubescent; areola white, small. Fl. Jul–Aug.● Forests, grass slopes; ca. 3000 m. Yunnan.Strobilanthes forrestii was misidentified as S. extensa (Pteracanthusextensus) by Handel-Mazzetti (Symb. Sin. 7: 895. 1936), C. Y. Wu(Index Fl. Yunnan. 2: 1682. 1984), and H. P. Tsui and C. C. Hu (FRPS70: 136. 2002; Fl. Yunnan. 16: 696. 2006).76. Strobilanthes lachenensis C. B. Clarke in J. D. Hooker, Fl.Brit. India 4: 465. 1884.白毛马蓝 bai mao ma lanChampionella xanthantha (Diels) Bremekamp; Pteracanthuslachenensis (C. B. Clarke) Bremekamp; P. leucotrichus(Benoist) C. Y. Wu & C. C. Hu; Strobilanthes leucotrichaBenoist; S. xanthantha Diels.Herbs 30–100 cm tall, isophyllous. Stems erect, basallyglabrescent, apically pubescent. Petiole 0.5–3.5 cm, thinly pubescentto glabrescent; leaf blade ovate, obovate, or broadlyelliptic, 3–11 × 2–6.5 cm, both surfaces thinly white pubescentespecially along veins, secondary veins 5–9 on each side ofmidvein, base cuneate and often decurrent onto petiole, marginserrate to crenate, apex acute to shortly acuminate. Inflorescencesaxillary or terminal, spikes, 3–10 cm, often forming adiffuse panicle of spikes; rachis gland-tipped pilose; bractsvariable in size and shape; basal bracts leaflike, sessile, ovate,2–5 × 0.5–2 cm, persistent, margin often toothed; apical bractsoblong to sometimes elliptic, 5–12 × 1–2 mm, persistent untilflowers fall; bracteoles linear, 4–5 × ca. 1 mm, white glandtippedpilose. Flowers basally to 2.5 cm apart on rachis but apicallycongested. Calyx 0.8–1.6 cm, accrescent in fruit, glandtippedpilose, 5-lobed almost to base; lobes linear, 7–12 × ca. 1mm, one longer and becoming spatulate. Corolla blue, 3–3.6cm, curved, outside pubescent on lobes, inside glabrous exceptfor trichomes retaining style; tube basally cylindric and ca. 2mm wide for ca. 8 mm, slightly bent at mouth, and graduallywidened to ca. 1.2 cm at mouth; lobes ovate, ca. 4 × 4 mm.Stamens 4, included; shorter filament pair 2–3 mm, glabrous;longer filament pair 7–8 mm, sparsely pilose; anther thecae oblong,ca. 2 × 1.3 mm, erect; pollen type 3. Ovary comose; style2.7–3.2 cm, sparsely gland-tipped pilose. Capsule 1.1–1.4 cm,gland-tipped pilose at tip, 4-seeded. Seeds ovate in outline, ca. 3× 2 mm, densely appressed pubescent. Fl. Jun–Oct.Forests; 1800–3400 m. Sichuan, Xizang (Dinggyê), Yunnan [Bhutan,India, Nepal].Strobilanthes lachenensis was incorrectly reported as Pteracanthusurticifolius (Kuntze) Bremekamp by H. W. Li (Fl. Xizang. 4: 179.1985).77. Strobilanthes nemorosa Benoist, Bull. Mus. Natl. Hist.Nat. 28: 97. 1922.琴叶马蓝 qin ye ma lanDiflugossa muliensis H. P. Tsui; Pteracanthus nemorosus(Benoist) C. Y. Wu & C. C. Hu; P. panduratus (Handel-Mazzetti)C. Y. Wu & C. C. Hu; Strobilanthes pandurata Handel-Mazzetti.Herbs to 60 cm tall, much branched, isophyllous. Stems 4-angled, sulcate, pubescent, basally woody. Leaves sessile orwith petiole to 1.5 cm; leaf blade lanceolate to oblong-elliptic,occasionally basally pandurate, 5–15(–20) × 1.5–6 cm, bothsurfaces sparsely pilose but glabrescent, adaxially dark greenand densely covered with cystoliths, secondary veins 6–12 oneach side of midvein, base rounded to attenuate, margin remotelyserrate, apex acuminate. Inflorescences terminal or axillary,spikes, usually several spikes forming a panicle; rachisdensely pilose, with gland-tipped trichomes; basal bracts leaflike,ovate-lanceolate, to 6 × 2 cm; apical bracts linear to narrowlyoblong-elliptic, 7–12 × 1–3 mm, persistent, pilose withgland-tipped trichomes; bracteoles linear, ca. 5 × 1 mm, pilosewith gland-tipped trichomes. Calyx 1–1.5 cm, pilose with glandtippedtrichomes, 5-lobed almost to base; lobes linear with onelonger than others, apex obtuse. Corolla blue, 3–3.7 cm, ventricose,outside sparsely pubescent, inside glabrous except fortrichomes retaining style; tube basally cylindric and ca. 2 mmwide for ca. 5 mm then gradually widened to ca. 1 cm at mouth;lobes oblong, ca. 5 × 4 mm, apex rounded. Stamens 4, included;shorter filament pair ca. 4 mm; longer filament pair ca. 6 mm,base pilose; anther thecae oblong, ca. 2.5 mm; pollen type 3.Ovary comose; style 2.8–3.2 cm, glabrous. Capsule clavate, ca.1.6 cm, apically gland-tipped pilose, 4-seeded. Seeds ovate inoutline, ca. 2.5 × 2 mm, densely appressed pubescent. Fl. Aug.● Pinus-Quercus forests, thickets; 1900–3400 m. Sichuan, Yunnan.Strobilanthes nemorosa is similar to S. lachenensis, differing inlittle more than leaf shape. It may not be a distinct species.78. Strobilanthes extensa (Nees) Nees in A. Candolle, Prodr.11: 191. 1847.棒果马蓝 bang guo ma lanGoldfussia extensa Nees in Wallich, Pl. Asiat. Rar. 3: 88.1832; Pteracanthus claviculatus (C. B. Clarke ex W. W. Smith)C. Y. Wu ex C. C. Hu; P. duclouxii (C. B. Clarke ex Benoist) C.

414ACANTHACEAEY. Wu & C. C. Hu; P. extensus (Nees) Bremekamp; Strobilanthesclaviculata C. B. Clarke ex W. W. Smith; S. duclouxii C. B.Clarke ex Benoist.Subshrubs 0.5–2 m tall, much branched, anisophyllous.Stems erect, glabrous or pilose. Petiole (0–)5–10 mm, narrowlywinged, sparsely pubescent; leaf blade ovate, lanceolate, or oblong-lanceolate,4–15 × 1–5 cm, both surfaces with prominentcystoliths, abaxially subglabrous to pilose, adaxially glabrous orsparsely pilose, secondary veins ca. 5 on each side of midvein,base cuneate, margin remotely serrate, apex acuminate and falcate.Inflorescences axillary or terminal, spikes, to 5 cm, verylax, 2–6-flowered; peduncle and rachis densely hirsute; sterilebracts intergrading with leaves but sessile, ovate to lanceolate,to 6 × 2.5 cm, glabrous or pubescent, base cuneate to cordate;floral bracts ovate to linear, 3–6 × 1.5–4 mm, persistent, glandtippedpilose; bracteoles oblong to elliptic, ca. 4 × 1–3 mm,gland-tipped pilose. Flowers usually paired, basally to 4 cmapart on rachis but apically closer. Calyx 1–2 cm, stronglyaccrescent in fruit, gland-tipped pilose, 5-lobed almost to base;lobes oblong-oblanceolate, one lobe 2–3 mm longer than others,apex spatulate and darker green. Corolla pale whitish purple toviolet, 3.2–4 cm, ventricose, outside pubescent only on lobes,inside glabrous except for trichomes retaining style; tube basallycylindric and ca. 2 mm wide for ca. 1.2 mm then graduallywidened to ca. 1.5 cm and bent at mouth; lobes orbicular, ca. 5mm in diam. Stamens 4, included; filaments glabrous, shorterpair 3–4 mm, longer pair 7–8 mm; anther thecae oblong, ca. 2.5× 1 mm, erect; pollen type 3. Ovary glabrous, apex with numeroussessile glands; style ca. 3.2 cm, sparsely pilose. Capsuleclavate, 10–16 × ca. 2 mm, apically thinly gland-tipped pilose,4-seeded. Seeds ovate in outline, ca. 3 × 2.5 mm, pubescent;areola small. Fl. Aug–Oct, fr. Oct–Dec.Mixed forests; 1900–2200 m. Sichuan, Yunnan [Bhutan, India,?Nepal].Strobilanthes extensa is very variable in leaf blade, bract, andbracteole shape and in the density of the indumentum.79. Strobilanthes spiciformis Y. F. Deng & J. R. I. Wood, J.Trop. Subtrop. Bot. 18: 473. 2010.黄连山马蓝 huang lian shan ma lanSubshrubs to 50 cm tall, anisophyllous. Stems to 5 mm indiam., young part sulcate and angled, subglabrous apart frompubescent at nodes, basally woody. Leaves petiolate or apicallysubsessile; petiole (0.5–)1.3–2.3(–2.5) cm, glabrous; leaf bladeovate, smaller of pair 4–7 × 2–3.5 cm and larger one 9–15 × 3–6 cm, both surfaces glabrous, adaxially with prominent cystoliths,secondary veins 5 or 6 on each side of midvein, base cuneateand decurrent onto petiole, margin serrate, apex acuminate.Inflorescences terminal, spikes, 4–9 cm; rachis 4-angled,gland-tipped pilose; bracts ovate, ca. 8 × 5 mm, deciduous asflowers open, abaxially sparsely gland-tipped pilose, adaxiallyglabrous, apex rounded; bracteoles absent. Calyx 1–1.6 cm;lobes linear-oblong, 8–16 × 1–2 mm, unequal with one distinctlylonger than others, gland-tipped pilose, margin entireand ciliate, apex acute. Corolla color not noted, probably bluish,ca. 2.5 cm, straight, outside pubescent in bud but soon glabrescent,inside glabrous; tube basally cylindric and ca. 2 mmwide for ca. 1 cm then gradually widened to ca. 8 mm at mouth;lobes orbicular, ca. 3 × 3 mm, apex obtuse. Stamens 4, included;filaments pilose, shorter pair ca. 5 mm, longer pair ca. 2cm; anther thecae oblong, ca. 2 × 1 mm; pollen type 3. Ovaryca. 2.5 × 0.8 mm, glabrous except for a few gland-tipped pilosetrichomes at tip; style ca. 2 cm, glabrous. Capsule oblong, 1.5–2× 0.3–0.4 cm, glabrous except for a few gland-tipped pilose trichomesat tip and along ribs toward tip, 4-seeded. Seeds ovateto suborbicular in outline, ca. 3 × 3 mm, pilose. Fl. Aug–Oct.● Moist places in forests; 1200–2400 m. Yunnan.80. Strobilanthes pseudocollina K. J. He & D. H. Qin, ActaPhytotax. Sin. 45: 701. 2007.阳朔马蓝 yang shuo ma lanHerbs to 50 cm tall, ascending, weakly anisophyllous.Stems terete, lenticellate, glabrous. Petiole 0.5–2.5 cm, sulcate;leaf blade ovate to lanceolate-ovate, 2.5–13 × 1.5–6 cm, smallerof pair 1/2–2/3 size of larger one, glabrous, cystoliths prominent,secondary veins 4–8 on each side of midvein, base obtuseto cuneate, margin subentire to sparsely crenulate, apex acuminate.Inflorescences from terminal leaf axils, spikes, (1–)3–11cm, 4–16-flowered; bracts subspatulate, 6–10 × 1.5–3 mm,leathery, glabrous, margin ciliate, apex acute; bracteoles lanceolate,ca. 4 × 0.6 mm, margin ciliate. Flower pairs 0.5–1.5 cmapart on rachis. Calyx 0.9–1.1 cm, 5-lobed almost to base; lobeslinear, ca. 1 mm wide, unequal, margin ciliate. Corolla purple,ca. 4.5 cm, outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric and ca. 1.5 mmwide for ca. 1 cm then gradually widened to ca. 1.5 cm atmouth; lobes suborbicular, 7–8 × 8–9 mm, apex obtuse. Stamens4, included; shorter filament pair ca. 3 mm, glabrous;longer filament pair ca. 6 mm, sparsely hirsute; anther thecaeoblong, ca. 2.5 × 1 mm, apiculate; connective shortly extended;pollen type 3. Ovary glabrous; style ca. 3 cm, glabrous. Capsule1–1.5 cm, glabrous. Seeds ovate in outline, ca. 4 × 2 mm,densely pubescent. Fl. Sep–Oct, fr. Oct–Nov.● Forests on limestone hills; 100–300 m. Guangxi (Yangshuo).81. Strobilanthes longzhouensis H. S. Lo & D. Fang, Guihaia17: 34. 1997.龙州马蓝 long zhou ma lanPerilepta longzhouensis (H. S. Lo & D. Fang) C. Y. Wu &C. C. Hu.Shrubs to 2.5 m tall, slightly anisophyllous. Stems basallygray, apically purplish black, glabrous. Petiole 0.5–1.5 cm, glabrous;leaf blade lanceolate-ovate to oblong-elliptic, 3–15.5 ×2.5–6 cm, leathery, glabrous, cystoliths numerous, secondaryveins 5–7(–9) on each side of midvein and prominent on bothsurfaces, base cuneate, margin serrulate to undulate, apex acuminateto acute. Inflorescences axillary, branched spikes, 1.5–5cm, to 4 per leaf axil, sometimes becoming compound andpaniculate; bracts ovate, 3–4 × ca. 2 mm, persistent, abaxiallyglabrous with numerous cystoliths, margin ciliolate, apex acute;bracteoles ca. 6 × 1.2 mm. Flowers 1 sterile per node. Calyx 8–10 mm, with prominent cystoliths, 2-lipped; lower lip 2-lobed,upper lip 3-lobed; lobes lanceolate-triangular, ca. 6 mm, mar-

ACANTHACEAE 415gins white ciliolate. Corolla pale purple, 3–3.5 cm, ventricose,outside glabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 2 mm wide for ca. 1 cmthen gradually widened to ca. 1.5 cm at mouth; lobes broadlyobovate, ca. 5 mm, apex emarginate. Stamens 4, included;shorter filament pair ca. 2 mm, glabrous; longer filament pairca. 7 mm, villous; anther thecae oblong, ca. 2 mm; pollen type3. Ovary glabrous; style ca. 2 cm, basally villous. Capsule oblong,1.2–1.4 cm, glabrous, 4-seeded. Seeds ovate in outline, ca.2.5 × 2.5 mm, densely covered with appressed trichomes. Fl.Sep–Dec, fr. Dec–Feb.Hilly limestone areas; 200–500 m. Guangxi, Hainan [Vietnam].82. Strobilanthes cystolithigera Lindau, Bull. Herb. Boissier5: 651. 1897.串花马蓝 chuan hua ma lanPteracanthus botryanthus (D. Fang & H. S. Lo) C. Y. Wu& C. C. Hu; Strobilanthes botryantha D. Fang & H. S. Lo; S.myriostachya D. Fang & H. S. Lo.Subshrubs 1.5–3.5 m tall, isophyllous, drying green. Stems4-angled, sulcate, glabrous. Petiole 0.5–2 cm, sulcate; leaf bladenarrowly elliptic, narrowly ovate, or rarely ovate-elliptic, (2–)4–17 × 1.5–4.5 cm, glabrous, densely covered with linear cystoliths,secondary veins 4–6 on each side of midvein, base cuneateand decurrent onto petiole, margin subentire to irregularlycrenate, apex acuminate. Inflorescences axillary, spikes, 2–6(–12) cm, sometimes branched, (2–)8–14-flowered; peduncle0.5–1.5 cm; bracts elliptic, 4–6 × 3–4 mm, persistent, abaxiallywith prominent cystoliths and frequently a single black squamategland, margin ciliolate, apex obtuse to acute; bracteolesovate-lanceolate, ca. 5 × 1 mm, glabrous except for ciliolatemargin, densely covered with cystoliths, apex acute. Flowers inopposite pairs, ca. 0.5 mm apart on rachis. Calyx 8–10 mm,subequally 5-lobed to base; lobes linear-lanceolate, 5–6 × ca. 1mm, basally overlapping, glabrous, apex obtuse. Corolla purple,3–4.5 cm, slightly curved, outside glabrous, inside glabrous exceptfor trichomes retaining style; tube basally cylindric andnarrow for ca. 1 cm then gradually widened to ca. 1.8 cm atmouth; lobes broadly ovate, ca. 6 × 7 mm, apex rounded. Stamens4, included; filaments pilose, shorter pair ca. 2 mm,longer pair ca. 5 mm; anther thecae oblong, ca. 2 × 1 mm; pollentype 3. Ovary glabrous; style ca. 2.8 cm, hirsute; stigmaglabrous. Capsule oblong, ca. 1.7 cm, glabrous, 4-seeded. Seedsovate in outline, ca. 1 cm, pilose; areola small, glabrous. Fl.Sep–Nov, fr. Dec.Ravines or streamsides on limestone hills; 800–1200 m. Guangxi,Hainan, Yunnan [Vietnam].83. Strobilanthes abbreviata Y. F. Deng & J. R. I. Wood, Bot.J. Linn. Soc. 150: 377. 2006.短尖马蓝 duan jian ma lanShrubs to 3 m tall, isophyllous, drying green. Stems terete,ca. 9 mm in diam., sulcate, striate, glabrous; bark dull brown,smooth. Petiole 1–2.5 cm, glabrous; leaf blade oblong-elliptic,6–25 × 2.5–9 cm, both surfaces glabrous and with numeroussmall cystoliths, abaxially olive-green, adaxially dark green,base long attenuate and decurrent onto petiole, margin serrate,apex tapered, long acuminate, and sometimes falcate. Inflorescencesaxillary, spikes, 2–14 cm, basally rarely with a reducedsecond spike, usually some forming a 30–50 cm leafless panicleof spikes; peduncle 1–3 mm; rachis glabrous; bracts broadlyoblong-obovate, 5–7 × 3–4 mm, concave, rigid, persistent intofruit, abaxially densely covered with cystoliths and dotted withsmall sessile brownish glands and often with larger sticky glandtippedpatches along midvein, apex abruptly truncate and with ashort deciduous mucro; bracteoles lanceolate to oblong-lanceolate,10–13 × 1–2 mm, slightly shorter than calyx, concave,rather rigid, glabrous or (rarely) scurfy along margin, abaxiallydensely covered with cystoliths and with scattered sessile glandssometimes becoming stipitate in fruit, apex obtuse. Flowersarranged in opposite pairs, both usually fertile, basally to 1 cmapart on rachis, apically confluent. Calyx 1.2–1.5 cm, slightlyaccrescent in fruit, subequally 5-lobed to just above base; lobesnarrowly lanceolate, 10–14 × 1–2 mm, glabrous except for apicalglands that sometimes become stipitate, outside non-glandularexcept for a few sessile glands near apex, densely coveredwith cystoliths, apex acute to acuminate. Corolla bluish purple,4–4.5 cm, sometimes apically bent, outside glabrous; tube basallycylindric and ca. 1.5 mm wide for 7–8 mm then ventricoseand widened to ca. 1.5 cm at mouth; lobes ovate, ca. 5 × 5 mm.Stamens 4, included; filaments setose, shorter pair ca. 2 mm,longer pair ca. 4 mm; anther thecae narrowly oblong, ca. 3 ×0.5 mm; pollen type 3. Ovary comose; style ca. 2.5 cm, denselyvillous. Capsule oblong, 17–18 × ca. 3 mm, comose with a fewtrichomes, 4-seeded. Seeds ovate in outline, flattened, denselypilose; areola small, glabrous. Fl. Jun–Oct, fr. Dec–Mar.Forests by streams; 200–1500 m. Yunnan [Cambodia, India,Myanmar, Thailand, Vietnam].84. Strobilanthes biocullata Y. F. Deng & J. R. I. Wood, Novon20: 406. 2010.湖南马蓝 hu nan ma lanSubshrubs 0.8–2 m tall, gregarious, anisophyllous. Stemssubterete, bisulcate, glabrous. Petiole 1–5 cm, sulcate, glabrous;leaf blade oblong-elliptic to elliptic, smaller of pair 6–12 × 2–4cm and larger ones 13–26 × 4–8 cm, both surfaces glabrous anddensely covered with cystoliths, secondary veins 5–10 on eachside of midvein and prominent on both surfaces, tertiary veinsprominent, base narrowly cuneate, margin serrate, apex acuminate.Inflorescences axillary, simple spikes, (3–)7–10 cm, solitary,leafless; rachis glabrous; bracts ovate, 3.5–5 × 2.5–3 mm,glabrous, abaxially with 2 swollen bulges resembling eyes, apexlong acuminate to mucronate; bracteoles ovate-oblong, ca. 4 × 2mm, somewhat fragile and caducous, glabrous, apex acuminateto mucronate. Flowers in opposite pairs, 1–2 cm apart on rachis.Calyx 1–1.2 cm, sparsely pubescent when young, glabrescent,subequally 5-lobed almost to base; lobes yellowish brown, lanceolate,1-veined, apex long acuminate. Corolla purplish blue,3.5–4 cm, outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric and ca. 3 mmwide for 6–11 mm then bent to ca. 90° and gradually widenedto ca. 2 cm at mouth; lobes ovate, ca. 4 × 7 mm, apex emarginate.Stamens 4, included; filaments pilose, shorter pair ca. 3mm, longer pair ca. 1 cm; anther thecae oblong, ca. 3 × 1.2 mm;

416ACANTHACEAEpollen type 3. Ovary ca. 3 mm, glabrous; style ca. 2.8 cm. Capsuleclavate, 1.5–2 cm, glabrous, 4-seeded. Seeds ca. 3 × 3 mm,lenticular, densely appressed pubescent; areola small. Fl. Jul–Oct, fr. Nov.● On rocks by streams; 200–800 m. Guangdong, Guangxi, Hunan.85. Strobilanthes longespicata Hayata, Icon. Pl. Formosan. 9:83. 1920.长穗马蓝 chang sui ma lanSemnostachya longespicata (Hayata) C. F. Hsieh & T. C.Huang.Shrubs 1.2–2 m tall, much branched, drying green.Branches ± 4-angled, sulcate, glabrous. Petiole 1–3 cm, sulcate,glabrous; leaf blade lanceolate to oblong-oblanceolate, 7–20 ×2–6 cm, glabrous, both surfaces with abundant cystoliths, secondaryveins 6–10 on each side of midvein, base attenuate, marginremotely serrate, apex acute to shortly acuminate. Inflorescencesaxillary, spikes, (5–)10–15 cm, simple or less commonlybranched; peduncle and rachis pubescent; bracts narrowly ovateto lanceolate, (5–)10–13 × (3–)6–7 mm, glabrous, cystolithsprominent, base truncate, apex acuminate; bracteoles linearlanceolate,9–11 × ca. 2 mm, scurfy-pubescent, apex acuminate.Flowers in opposite pairs, basally ca. 1 cm apart on rachis butapically imbricate. Calyx 5-lobed almost to base, accrescent infruit; lobes linear-lanceolate, 9–15 × 2–3 mm, scurfy-pubescent,prominently 1-veined, margin ciliolate, apex acuminate. Corollapale purple, 4.5–5 cm, straight or slightly bent, outside glabrous,inside glabrous except for trichomes retaining style; tubebasally cylindric and ca. 3 mm wide for ca. 1 cm then graduallywidened to 1.5–2 cm at mouth; lobes oblong, ca. 8 × 7 mm,subequal, apex emarginate. Stamens 4, included; filaments glabrous,shorter pair ca. 2 mm, longer pair ca. 8 mm; antherthecae oblong, 2–2.5 × ca. 1 mm; pollen type 3. Ovary cylindric-oblong,apically slightly hispid; style filiform, 3–3.2 mm,hispid. Capsule linear, ca. 1.7 cm, glabrous, 4-seeded. Seedssuborbicular in outline, ca. 3 mm in diam., flattened. Fl. Sep–Dec, fr. Feb–May.● Forests; 200–1000 m. Taiwan.86. Strobilanthes myura Benoist, Bull. Mus. Natl. Hist. Nat.28: 95. 1922.鼠尾马蓝 shu wei ma lanShrubs, drying green. Stems subterete, glabrous. Petiole 1–2.5 cm, glabrous; leaf blade ovate to elliptic, to 7–16 × 3–6 cm,glabrous, adaxially with cystoliths, secondary veins 8–11 oneach side of midvein, base cuneate, margin crenate-dentate, apexacuminate. Inflorescences axillary, spikes, 5–8 cm; bracts ovatetriangular,6–9 × ca. 2 mm, glabrous, midvein abaxially linedby cystoliths, margin sparsely ciliate, apex acuminate; bracteolesnarrowly ovate, ca. 3.5 × 1 mm, margin sparsely ciliate,apex acuminate. Flowers basally ca. 1.5 cm apart on rachis butapically somewhat confluent. Calyx 5-lobed to base; lobes lanceolate,ca. 7 × 1 mm, equal, margin minutely ciliolate, apexacuminate and shortly apiculate. Corolla purple, 2.5–3.4 cm,outside glabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and narrow for ca. 5 mm then bentand abruptly widened to 1–1.2 cm at mouth; lobes oblongovate,ca. 5 × 4 mm. Stamens 4, included; filaments pilose,shorter pair ca. 4 mm, longer pair ca. 6 mm; anther thecae oblong,ca. 2 × 1 mm; pollen type 3. Ovary sparsely pilose; styleca. 2.8 cm, sparsely pilose. Capsule unknown. Fl. Jul.● Thickets; 600–800 m. Guizhou (Dushan).87. Strobilanthes guangxiensis S. Z. Huang, Guihaia 6: 179.1986.广西马蓝 guang xi ma lanPteracanthus guangxiensis (S. Z. Huang) C. Y. Wu & C.C. Hu.Herbs to 1 m tall, perennial, anisophyllous. Stems erect orascending, densely pubescent, soon glabrescent. Petiole 0.3–3.5cm, densely pubescent; leaf blade elliptic, smaller of pair 4–9 ×2.3–4.5 cm and larger one 9–14(–18) × 3.3–6(–8) cm, abaxiallypale green and slightly pubescent, adaxially green, glabrous,and densely covered with prominent ca. 0.5 mm cystoliths, secondaryveins 4–8 on each side of midvein, base cuneate, marginentire or slightly undulate, apex acuminate. Inflorescences axillaryor terminal, spikes, 4–8 cm; peduncle ca. 4 cm, denselypubescent; rachis densely pubescent, usually with 2–6 nodes;bracts oblong to lanceolate, 10–13 × 2–3 mm, persistent, abaxiallypubescent, adaxially glabrous, margin ciliate, apex acuminateto caudate; bracteoles oblong to lanceolate, 11–16 × 2–3mm, abaxially pubescent, adaxially glabrous, margin ciliate,apex acuminate to caudate. Flowers 0.8–1.2 cm apart on rachis.Calyx ca. 1 cm, 5-lobed almost to base; lobes linear, ca. 9 × 1mm, equal, outside pubescent, inside glabrous, margin ciliatefrom middle apically, apex acute. Corolla purple, 3.5–4 cm,ventricose, outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric and ca. 3 mmwide for ca. 1.2 cm then gradually widened to ca. 1.4 cm atmouth; lobes oblong, ca. 5 × 5 mm, unequal, apex emarginate.Stamens 4, included; filaments glabrous, shorter pair ca. 3mm, longer pair ca. 6 mm; anther thecae oblong, 2.5–3.5 × ca.2 mm. Ovary oblong, ca. 3 mm, pilose at tip; style 2.5–3 cm,villous; stigma linear, glabrous. Capsule not seen. Fl. Oct–Dec.● Shaded valleys; 200–400 m. Guangxi (Liujiang).This species is used medicinally.88. Strobilanthes compacta D. Fang & H. S. Lo, Guihaia 17:31. 1997.密苞马蓝 mi bao ma lanSubshrubs to 2 m tall, slightly anisophyllous. Stems 4-angled,glabrous. Petiole 1–3 cm, sulcate, glabrous; leaf bladeelliptic to ovate-elliptic, 8.5–15 × 3.5–7 cm, both surfaces glabrousand densely covered with cystoliths, adaxially blackishwhen dry, secondary veins 8 or 9 on each side of midvein andprominent, base cuneate to attenuate, margin serrate, apex acuminate.Inflorescences axillary, simple spikes, 1.5–8(–10) × ca.1.2 cm, very dense; basal sterile bracts ovate-oblong, 3–4 × ca.1 cm, leathery, abruptly narrowed at middle with a long caudateapex; floral bracts ovate, 2–3 × 0.4–0.6 cm, leathery, persistent,abaxially glabrous, margin minutely ciliolate or glabrous, apex

ACANTHACEAE 417long acuminate; bracteoles ovate, ca. 1 × 0.5 cm, glabrous,densely covered with cystoliths, margin minutely ciliolate, apexacuminate. Flowers imbricate on rachis. Calyx 5-lobed almostto base; lobes broadly lanceolate, 12–15 × 3–4 mm, subequal,basally overlapping, margin minutely ciliolate but otherwiseglabrous, apex acuminate. Corolla purple, 3.5–4.8 cm, outsideglabrous, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for ca. 1 cm thenslightly ventricose and gradually widened to ca. 1.5 cm atmouth; lobes ovate, ca. 5 mm, apex rounded. Stamens 4, included;filaments pubescent, shorter pair ca. 2.5 mm, longerpair ca. 6 mm; anther thecae oblong, ca. 3 × 1 mm, glabrous.Ovary sparsely hirsute at tip; style ca. 2.5 cm, slightly hirsute atbase and glabrous apically. Capsule fusiform, ca. 1.5 cm, somewhathirsute on apical part, 4-seeded. Seeds ovate in outline, ca.3 × 2 mm, densely appressed hairy. Fl. Aug–Sep.● Forests on limestone hills; 200–300 m. Guangdong, Guangxi.89. Strobilanthes heteroclita D. Fang & H. S. Lo, Guihaia 17:32. 1997.异序马蓝 yi xu ma lanHerbs or subshrubs, to 1.5 m tall, gregarious, isophyllous.Stems ± 4-angled, sulcate, glabrous. Petiole to 5 cm; leaf bladebroadly ovate, elliptic, broadly elliptic, or suborbicular, 5.5–19 × 2.5–10 cm, glabrous, both surfaces densely covered withlinear cystoliths, secondary veins 7–10 on each side of midveinand prominent, base rounded to broadly cuneate, margin serrate,apex shortly caudate. Inflorescences axillary from apicalaxils, spikes, 1.5–5 cm, compact, 1–3 per axil; peduncle 0–1.3cm; bracts broadly lanceolate, 11–12 × 3–4 mm, leathery, persistent,rust-colored scurfy pubescent, abaxially with black sessileglands, glabrescent, apex acuminate; bracteoles lanceolate,8–9 mm, leathery, rust-colored tomentose, apex acuminate. Calyx1.2–1.4 cm, 5-lobed almost to base; lobes narrowly lanceolate,with 2 lobes slightly longer than others, apex acuminate.Corolla purple to light blue, 3.5–4 cm, outside glabrous, insideglabrous except for trichomes retaining style; tube basallycylindric and ca. 3 mm wide for ca. 1 cm then ventricose andgradually widened to ca. 1.2 cm at mouth; lobes ovate, 5–6 × 5–6 mm, apex obtuse. Stamens 4, included; filaments pilose,shorter pair ca. 2 mm, longer pair ca. 7 mm; anther thecae oblong,2–2.5 mm. Ovary sparsely hirsute at tip; style ca. 1.7 cm;stigma hirsute. Capsule fusiform, 1.5–2 cm, pilose at tip, 4-seeded. Seeds ovate in outline, ca. 3 × 3 mm, appressed pubescent.Fl. Jul–Sep, fr. Nov.● Forests on limestone hills; 100–500 m. Guangxi (Long’an).90. Strobilanthes mucronatoproducta Lindau, Bull. Herb.Boissier 5: 650. 1897.尾苞马蓝 wei bao ma lanShrubs to 1.5 m tall, weakly anisophyllous, dryingblackish. Stems ± 4-angled, glabrescent. Petiole 0.5–3 cm,scurfy pubescent; leaf blade ovate, elliptic, or oblong-elliptic,8.5–15 × 3.5–7 cm, abaxially puberulent along veins, adaxiallyglabrous, both surfaces densely covered with cystoliths, secondaryveins 8 or 9 on each side of midvein, base cuneate to attenuate,margin dentate, serrulate, or subentire, apex acuminate. Inflorescencesterminal and from apical leaf axils, simple spikes,4–9 cm, solitary; peduncle 0.3–1 cm, rust-colored pubescent;bracts lanceolate, ca. 2.4 × 0.6 cm, persistent, rust-colored pilose,apex acuminate; bracteoles lanceolate, ca. 5 × 4 mm, rustcoloredpilose, apex long acuminate and aristate. Flowers imbricateon rachis. Calyx 1.7–1.8 cm, 5-lobed almost to base;lobes lanceolate, 16–17 × ca. 2 mm, subequal, rust-colored pilose,apex acuminate. Corolla purple, 3.5–4 cm, slightly curved,outside glabrous, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 3 mm wide for ca. 1 mmthen gradually widened to ca. 1.3 cm at mouth; lobes ovate, ca.7 × 7 mm, apex obtuse. Stamens 4, included; filaments sparselypubescent, shorter pair ca. 2 mm, longer pair ca. 4 mm; antherthecae oblong, ca. 2 × 1.3 mm; pollen type 3. Ovary pilose;style ca. 2.7 cm; stigma ca. 3 mm, pubescent. Capsule fusiform,2–2.2 cm, pilose, 4-seeded. Seeds ovate in outline, ca. 4 × 3.5mm, appressed pubescent. Fl. Sep–Oct, fr. Dec.Forests on mountain slopes; 400–700 m. Guangxi, Yunnan [Vietnam].91. Strobilanthes fimbriata Nees in Wallich, Pl. Asiat. Rar. 3:85. 1832.流苏马蓝 liu su ma lanEndopogon macrostegius Nees; Strobilanthes fimbriatavar. majuscula W. W. Smith; S. macrostegia (Nees) C. B.Clarke; S. neesii Kurz.Shrubs 0.6–1.5 m tall, drying blackish green. Branches terete,apically thinly brown pubescent, glabrescent. Petiole 0.5–2.5 cm; leaf blade oblong-elliptic to elliptic, to 18 × 7 cm, bothsurfaces with cystoliths, abaxially glabrous except for brownscurfy pubescence along veins, adaxially glabrous, secondaryveins 8–12 on each side of midvein, base cuneate, margin subentire,apex acuminate. Inflorescences axillary, spikes, 3–6cm, with 6–8 flower pairs; peduncle 0–0.3 cm, glabrous; bractsobovate, ca. 13 × 5 mm, both surfaces densely covered withcystoliths, apical margin ciliate with rigid brown trichomes,apex abruptly narrowed to a mucro; bracteoles oblanceolate, ca.1 × 3 mm, abaxially apically fulvous pubescent, margin with adistinct shoulder, apex cuspidate. Flowers imbricate on rachis.Calyx 1–1.3 cm at anthesis, accrescent to ca. 2 cm; lobes linearlanceolate,to 18 × 1 mm, apex long acuminate. Corolla violet,ca. 4.5 cm, outside and inside glabrous except for trichomes retainingstyle; tube basally cylindric for ca. 8 mm then abruptlywidened to 1.5–1.8 cm at mouth; lobes broadly ovate, ca. 6 × 7mm, apex rounded. Stamens 4, included; shorter filament pairca. 2 mm, glabrous; longer filament pair ca. 7 mm, pilose; antherthecae oblong, ca. 3 × 1.5 mm, erect; pollen type 3. Ovaryglabrous except for densely comose tip; style ca. 4 cm, pilose.Capsule oblong, 1–2 cm, comose, 4-seeded. Seeds ovate in outline,ca. 4 × 3 mm, densely appressed pilose. Fl. May.Moist places in forests; ca. 1000 m. Xizang (Mêdog) [Bangladesh,India, Myanmar].92. Strobilanthes brunnescens Benoist, Bull. Mus. Natl. Hist.Nat. 27: 544. 1921.折苞马蓝 zhe bao ma lan

418ACANTHACEAEPerilepta refracta (D. Fang, Y. G. Wei & J. Murata) C. Y.Wu & C. C. Hu; Strobilanthes refracta D. Fang, Y. G. Wei & J.Murata.Subshrubs 1–2 m tall, anisophyllous, drying blackish.Stems 4-angled, sulcate, brownish puberulent. Petiole 0.2–2.5cm, brownish puberulent; leaf blade ovate to rarely obovateelliptic,(2–)4.5–21 × (1–)2.5–9.5 cm, both surfaces denselycovered with cystoliths, abaxially puberulent along veins, adaxiallyglabrous, secondary veins 4–12 on each side of midvein,base cuneate, margin subentire to crenulate, apex shortly acuminate.Inflorescences axillary, spikes, 1.5–3 cm; bracts oblong-obovatewith a broad base, ca. 11 × 4–5 mm, concave,rigid, abaxially pubescent and with marginal cilia of rigidbrown trichomes, apex acute and becoming reflexed; bracteoleslinear, 10–12 × ca. 1 mm, apically brownish pubescent, apexoften reflexed. Flowers imbricate on rachis. Calyx 1.2–1.5 cm,subequally 5-lobed; lobes lanceolate, ca. 10 × 1 mm, pubescentwith rigid trichomes, apex acuminate. Corolla purple, ca. 4 cm,slightly curved, outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric and narrow for ca.1 cm then gradually widened to ca. 2 cm at mouth; lobes ovate,ca. 5 mm, apex emarginate. Stamens 4, included; shorter filamentpair ca. 3 mm, glabrous; longer filament pair ca. 7 mm,pilose; anther thecae oblong, 3–3.5 mm. Ovary hirsute at tip;style ca. 2.7 cm, pilose; stigma obconical. Capsule not seen. Fl.Dec.Forests in valleys; 300–500 m. Guangxi, Yunnan [Vietnam].93. Strobilanthes nobilis C. B. Clarke in J. D. Hooker, Fl. Brit.India 4: 471. 1884.沙坝马蓝 sha ba ma lanStrobilanthes petelotii Benoist.Herbs 50–100 cm tall, perennial, isophyllous, dryingblackish. Stems ± 4-angled, sulcate, reddish brown pubescent,apically with distinctive scattered wartlike excrescences. Petiole1–1.5 cm or leaves sessile; leaf blade ovate-lanceolate to oblong-elliptic,sometimes pandurate, 7–17(–25) × 3.5–8 cm, glabrousor thinly pilose especially on abaxial veins, margin, andadaxial surface, abaxially paler, adaxially with cystoliths, secondaryveins 6 or 7 on each side of midvein and somewhat parallel,base cuneate to rounded and auriculate, margin crenate,apex shortly acuminate. Inflorescences axillary or terminal,simple or basally branched spikes, 5–20 cm, often forming apanicle when mature; rachis reddish brown puberulent, apicallybecoming very slender; bracts lanceolate, 1.5–2 × 0.2–0.4 cm,persistent, glabrous except for ciliolate margin, occasionallydeveloping stipitate glands when mature, 1-veined, apex longacuminate; bracteoles similar to bracts but narrowly linear-lanceolate,ca. 18 × 1 mm. Flowers 1–1.5 cm apart on rachis andapically not very confluent. Calyx 1–2.2 cm, accrescent in fruit,5-lobed almost to base; lobes linear, 14–20 × ca. 1 mm, equal,occasionally developing stipitate glands when mature, marginciliolate, apex finely acuminate. Corolla blue, ca. 3 cm, outsideglabrous, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2.5 mm wide for ca. 7 mm thenstrongly bent and abruptly widened to 1–1.4 cm at mouth; lobesovate, ca. 4 × 4 mm, apex emarginate. Stamens 4, included;shorter filament pair 2–3 mm, glabrous; longer filament pair ca.7 mm, basally pilose; anther thecae oblong, ca. 3 × 1.2 mm;pollen type 3. Ovary glabrous, with 2 ovules per locule; styleca. 2.5 cm, pilose. Capsule 1.2–2.2 cm, glabrous, 4-seeded.Seeds ovate in outline, ca. 3 × 2.5 mm, densely appressed pubescent;areola small. Fl. Sep–Nov, fr. Mar.Streamsides, forests; 1100–1800 m. Yunnan [India, Myanmar,Vietnam].94. Strobilanthes polyneuros C. B. Clarke ex W. W. Smith,Notes Roy. Bot. Gard. Edinburgh 10: 198. 1918.多脉马蓝 duo mai ma lanShrubs to 2 m tall, isophyllous. Stems 4-angled, sulcate,glabrous. Petiole 1–5 cm, sulcate, subglabrous; leaf blade ellipticto broadly lanceolate, 13–30 × 5–11 cm, glabrous, abaxiallypaler, adaxially with numerous cystoliths, secondary veins 12–20 on each side of midvein and parallel, base cuneate, marginserrate, apex acuminate. Inflorescences axillary, spikes, 2–8 cm,composed of 1–5 simple dense spikes arising at apex of peduncleforming a dense cluster of spikes; peduncle ca. 1 cm, puberulentand sometimes with warty excrescences; sterile bractsovate at base of spike, ca. 1 × 0.5 cm; floral bracts lanceolate,1.5–3 × 0.3–0.4 cm, abaxially minutely puberulent, apex finelyacuminate and slightly incurved; bracteoles linear-lanceolate,ca. 2 cm, minutely puberulent, apex acuminate. Flowers imbricateon rachis. Calyx 1.7–2.3 cm, somewhat accrescent andsometimes becoming gland-tipped pilose at maturity, 5-lobedalmost to base; lobes linear-lanceolate, thinly puberulent, apexfinely acuminate. Corolla purple, 3.5–4 cm, straight, outsidesparsely pilose, inside glabrous except for trichomes retainingstyle; tube basally cylindric and ca. 3 mm wide for ca. 1.4 cmthen gradually widened to ca. 1.4 cm at mouth; lobes orbicular,ca. 6 mm in diam. Stamens 4, included; filaments glabrous,shorter pair ca. 2 mm, longer pair ca. 5 mm; anther thecaeoblong, ca. 3.5 × 1 mm; pollen type 3. Ovary sparsely glandtippedpilose at tip; style ca. 3.5 cm, pilose. Capsule ca. 20 × 3mm, glabrous or with a very few apical trichomes, 4-seeded.Seeds ovate in outline, 3–5 × 2–4 mm, densely appressed pubescent.Fl. Oct–Jan, fr. Mar.Forests; 200–1600 m. Yunnan [Myanmar, Thailand, Vietnam].95. Strobilanthes bipartita Terao ex J. R. I. Wood, Kew Bull.58: 122. 2003.双萼马蓝 shuang e ma lanSubshrubs 50–100 cm tall, isophyllous. Stems terete, glabrous.Petiole 0.7–1.5 cm, sparsely brown pilose; leaf bladeelliptic, 6–22 × 3.5–10 cm, hirsute especially along veins, abaxiallycystoliths prominent, secondary veins 12–16 on each sideof midvein, base cuneate, margin remotely serrate, apex acute.Inflorescences axillary or terminal, abbreviated to headlikespikes, ca. 2 cm; peduncle 2–4 mm, brown pubescent; bractsovate, 3–3.5 × 1.2–1.7 cm, glabrous except for ciliate margin,margin entire or apically serrate, apex acute; bracteoles oblonglanceolate,ca. 15 × 3 mm, glabrous. Calyx with appressed silkytrichomes on both surfaces, 2-lipped; lower lip ca. 1.2 cm, 2-lobed, lobes triangular and ca. 1.5 × 1.5 mm; upper lip ca. 1.3cm, 3-dentate to ca. 1 mm. Corolla purplish blue, ca. 4 cm, ventricose,outside glabrous, inside glabrous except for trichomes

ACANTHACEAE 419retaining style; tube basally cylindric and ca. 2 mm wide for ca.1.2 cm then gradually widened to ca. 1.4 cm at mouth; lobesovate, ca. 5 × 7 mm, apex obtuse. Stamens 4, included; shorterfilament pair ca. 2 mm, glabrous; longer filament pair ca. 4.5mm, sparsely pilose; anther thecae oblong, ca. 3 × 1 mm. Ovaryglabrous; style ca. 3.2 cm, pilose. Capsule not seen. Fl. Dec.Thickets on limestone hills; 300–900 m. Yunnan (Jinghong)[Laos].96. Strobilanthes retusa D. Fang, Acta Phytotax. Sin. 38: 187.2000.凹苞马蓝 ao bao ma lanPerilepta retusa (D. Fang) C. Y. Wu & C. C. Hu.Herbs to 80 cm tall, branched, anisophyllous. Stems blackwhen dry, 4-angled, glabrous, winged between nodes, wing0.5–2 mm wide. Leaves sessile; leaf blade elliptic, rhombicelliptic,obovate, or ovate, 6.5–20.5 × 4–8.5 cm, abaxially glabrousexcept sparsely pilose along veins, adaxially pubescentand densely covered with cystoliths, secondary veins 5–9 oneach side of midvein and prominent on both surfaces, basallyattenuate to rarely broadly cuneate, base auriculate, amplexicaul,and decurrent onto petiole, margin crenate, apex acuteto acuminate. Inflorescences axillary or terminal, subcapitate,2.5–3.5 × 2–2.5 cm, elongating after flowering; peduncle 1–4cm; bracts ovate, ca. 2.2 × 1.5 cm, abaxially puberulent, basecordate, margin deeply crenate, apex acute; bracteoles elliptic,1.4–1.6 × ca. 0.7 cm, margin undulate, apex obtuse to emarginate.Calyx ca. 1.2 cm, accrescent to ca. 1.8 cm in fruit, subequally5-lobed; lobes oblanceolate, puberulent, lower 2 ca. 9.5mm and connate at base, upper 3 ca. 1.2 cm and extremelyshortly connate. Corolla white with purplish stripes on upperlobes, ca. 4 cm, slightly curved, outside glabrous; tube basallycylindric and narrow for ca. 1 cm then gradually widened to ca.2 cm at mouth; lobes oblong, ca. 5 × 5 mm, apex subrounded.Stamens 4, included; filaments pilose, shorter pair ca. 2.5 mmand basally sparsely pilose, longer pair ca. 5 mm; anther thecaeoblong, 3–4 mm. Ovary glabrous; style ca. 2.5 cm, pubescent;stigma unequally 2-lobed. Capsule ellipsoid, ca. 1.8 cm, glabrous,4-seeded. Seeds ovate in outline, ca. 2.5 × 2.5 mm,densely appressed pilose. Fl. Jul.● Moist places by streams; 500–600 m. Guangxi (Huanjiang).97. Strobilanthes cruciata (Bremekamp) Terao, Acta Phytotax.Geobot. 31: 59. 1980.四苞马蓝 si bao ma lanTetragoga cruciata Bremekamp, Verh. Kon. Ned. Akad.Wetensch., Afd. Natuurk., Sect. 2, 41(1): 300. 1944; T. nagaensisBremekamp.Subshrubs 1–2 m tall, anisophyllous. Stems 4-angled, sulcate,rust-colored hirsute or glabrous. Petiole 1–1.5 cm, pubescent;leaf blade ovate to ovate-lanceolate, 4–13 × 1.5–5.5 cm,smaller of pair ca. 1/3 size of larger one, abaxially pale whitetomentose especially along midvein, adaxially sparsely tomentoseand brownish green when dry, secondary veins 7–9 oneach side of midvein, base cuneate, margin serrate and ciliate,apex acuminate. Inflorescences axillary or terminal, subcapitate;peduncle 0–2.5 cm, gland-tipped pilose; outer bracts leaflike,petiolar base forming a rigid involucre around capitulum,1–2 × 0.5–1 cm, outside densely brownish gland-tipped piloseto subglabrous, inside glabrous, apical laminar portion ovate, 2–4 × 2–3 cm, and erect; inner bracts linear-pandurate, 12–16 ×3–5 mm, abaxially pilose and apically gland-tipped pubescent,adaxially glabrous, margin entire, apex acute; bracteoles narrowlylinear, ca. 13 × 1 mm. Calyx ca. 1.1 cm, outside piloseexcept glabrous toward base, inside apically silky villous, marginapically ciliate. Corolla white, ca. 2.5 cm, curved, outsideglabrous, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 3 mm wide for ca. 1 mm thenventricose and gradually widened to ca. 1.2 cm at mouth; lobesovate, ca. 4 × 4 mm, apex obtuse. Stamens 4, included; shorterfilament pair ca. 3 mm, glabrous; longer filament pair ca. 6 mm,hirsute; anther thecae oblong, ca. 3 × 1 mm, erect, apiculate;connective shortly excurrent; pollen type 1. Ovary oblong, glabrous;style ca. 2.2 cm, gland-tipped pilose. Capsule fusiform,ca. 1.5 cm, glabrous, 4-seeded. Seeds brown, ovate in outline,ca. 4 × 3 mm, densely appressed pubescent; areola small. Fl.Dec–Jul.Dense moist mixed forests; 800–1500 m. Hainan, Xizang(Mêdog), Yunnan [India, Indonesia, Myanmar, Thailand].Strobilanthes cruciata is a very distinct species with unique inflorescencestructure and unusual pollen combined with the calyx sericeousinside.98. Strobilanthes simonsii T. Anderson, J. Linn. Soc., Bot. 9:474. 1867.西蒙马蓝 xi meng ma lanSubshrubs to 1 m tall, anisophyllous. Stems and branches4-angled, sulcate, tomentose. Petiole 2–10 mm, sulcate, tomentose;leaf blade ovate-elliptic, smaller one 1.2–4 × 0.6–1.4 cmand larger one 3.5–18 × 1.6–6.5 cm, both surfaces tomentose,secondary veins 5–7 on each side of midvein, base cuneate andoblique, margin crenulate-serrate, apex acuminate and falcate.Inflorescences axillary, capitate, few flowered; peduncle 0.2–1cm, pubescent; bracts variable in size and shape; outermostbracts leaflike, pandurate with a constricted middle and broadpetiolar base, 3–4 × ca. 0.5 cm; inner bracts lanceolate, 13–20 ×ca. 3 mm, abaxially tomentose, apex acute; bracteoles linear, ca.2 × 0.3 cm, abaxially apically pubescent. Calyx ca. 1.4 cm,subequally 5-lobed almost to base; lobes linear, outside apicallypilose, apex acute. Corolla white, ca. 4 cm, straight, outside glabrous,inside glabrous except for trichomes retaining style; tubebasally cylindric and ca. 2.5 mm wide for ca. 8 mm then graduallywidened to ca. 1.8 cm at mouth; lobes oblong, ca. 6 × 5mm, apex rounded. Stamens 4, included; shorter filament pairca. 3 mm, glabrous; longer filament pair ca. 1 cm, hirsute; antherthecae oblong, ca. 2 × 1 mm, base divaricate, apex apiculate;connective shortly excurrent. Ovary glabrous; style ca.3.2 cm, hirsute. Capsule clavate, 1.8–2 cm, glabrous, 4-seeded.Seeds ovate in outline, ca. 3 × 2 mm, densely pubescent. Fl.Mar–Aug.Forests; 1300–1700 m. Xizang (Mêdog) [Bhutan, India].

420ACANTHACEAE99. Strobilanthes echinata Nees in Wallich, Pl. Asiat. Rar. 3:85. 1832.长苞马蓝 chang bao ma lanGoldfussia echinata (Nees) N. P. Balakrishnan; Strobilanthesjugorum Benoist; Tetraglochidium jugorum (Benoist)Bremekamp.Herbs to 2 m tall, anisophyllous. Stems subterete to 4-angled, sulcate, pubescent with brownish trichomes or glabrescent.Leaves petiolate but apicalmost ones subsessile; petiole0.5–4 cm, flat, slightly winged, pubescent or glabrous; leafblade ovate, elliptic, or ovate-lanceolate, 4–20 × 1.5–7.5 cm,smaller of pair ca. 1/2 size of larger one, both surfaces pilose tosubglabrous and with cystoliths, abaxially pale green, adaxiallygreen, secondary veins 7–11 on each side of midvein, base cuneateand decurrent onto petiole, margin crenate-serrate, apexacuminate. Inflorescences capitate, 4–8-flowered, enveloped bybracts; peduncle short, usually simple; outer sterile bracts sessile,ovate, ovate-oblong, or oblong-ovate, 3–5 × 2–2.5 cm,equal or unequal, persistent, margin dentate, apex obtuse,rounded, or fimbriate; inner bracts fertile, obovate to spatulate,2–3 × 0.7–0.8 cm, margin ciliate; bracteoles linear to oblanceolate,13–23 × 1.5–3 mm, pubescent or glabrous, margin entireor sometimes dentate or fimbriate toward apex, apex acuminate.Calyx 1.5–2 cm, pubescent, 5-lobed almost to base; lobes linear,12–19 × 1–2 mm, subequal, outside apically pubescent. Corollaviolet, 4.5–5.5 cm, outside glabrous, inside glabrous exceptfor trichomes retaining style; tube basally cylindric and ca.2 mm wide for ca. 1.5 mm then ventricose and widened to ca.1.5 cm at mouth; lobes ovate, 8–9 × 8–10 mm, apex rounded.Fertile stamens 4, included; shorter filament pair 3–5 mm,glabrous; longer filament pair 5–7 mm, pilose; anther thecaeoblong, 2–3 × ca. 2 mm, apex apiculate; connective excurrent;staminode occasionally present, small or inconspicuous; pollentype 3. Ovary glabrous; style ca. 2.5 cm, hirsute. Capsule fusiform,1.2–1.8 cm, glabrous, 4-seeded. Seeds brown, ovate inoutline, 3–5 × 2.5–4 mm, pilose; areola small. Fl. May–Sep.Moist forests; 100–2200 m. Guangdong, Guangxi, Yunnan [Bhutan,Cambodia, India, Indonesia, Laos, Malaysia, Myanmar, Thailand,Vietnam].Strobilanthes echinata has been misidentified in the recent literatureas Tetraglochidium gigantodes (Lindau) C. Y. Wu & C. C. Hu(Strobilanthes gigantodes Lindau) by D. Fang et al. (Guihaia 17: 38.1997), C. C. Hu (FRPS 70: 157. 2002), and C. C. Hu and X. P. Fu(Higher Pl. China 10: 373. 2004).100. Strobilanthes cuneata (Shakya) J. R. I. Wood, EdinburghJ. Bot. 51: 218. 1994.楔叶马蓝 xie ye ma lanDossifluga cuneata Shakya, J. Jap. Bot. 50: 99. 1975.Herbs to 30 cm tall, isophyllous. Stems 4-angled, glabrous.Petiole ca. 5 mm; leaf blade obovate to broadly elliptic, 4.5–7 ×2–3.5 cm, abaxially sparsely pilose along veins, adaxially glabrousand densely covered with cystoliths, secondary veins 6 or7 on each side of midvein, base narrowly cuneate, margin serrateand ciliate, apex acuminate. Inflorescences axillary, subsessileclusters; bracts leaflike, obovate, 6–12 × 6–8 mm, cystolithsprominent, margin dentate; bracteoles linear-oblanceolate, 2–4mm. Calyx 1.2–1.5 cm, 5-lobed almost to base; lobes linearlanceolate,1–1.5 mm wide, subequal, margin ciliate. Corollablue, campanulate, 3–4 cm, outside glabrous except for ciliaalong veins, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for ca. 1.2 cm thengradually widened to ca. 1.4 cm at mouth; lobes orbicular, ca. 8mm in diam. Stamens 4, included; filaments glabrous; shorterfilament pair ca. 3 mm, equal, incurved; longer filament pair ca.8 mm, unequal, incurved; anther thecae spherical, ca. 1.2 mm indiam., nodding; pollen type 3. Ovary glabrous; style 2.8–3.2cm. Capsule obovoid, ca. 10 × 3 mm, glabrous, 4-seeded. Seedssuborbicular in outline, ca. 1.5 mm, long pilose; areole large. Fl.Aug.Forests; ca. 2500 m. Xizang (Nyalam) [Nepal].101. Strobilanthes glomerata (Nees) T. Anderson, J. Linn.Soc., Bot. 9: 475. 1867.球序马蓝 qiu xu ma lanGoldfussia glomerata Nees in Wallich, Pl. Asiat. Rar. 3:88. 1832.Subshrubs to 1 m tall, strongly anisophyllous. Stems andbranches 4-angled to subterete, setose. Petiole 0.6–3 cm, denselypubescent; leaf blade elliptic, broadly elliptic, or narrowly ovate,5.5–19 × 2.7–10 cm, smaller of pair ca. 1/5 size of larger one,both surfaces densely pubescent with purplish trichomes, secondaryveins 5–7 on each side of midvein, base oblique,rounded, and decurrent onto petiole, margin serrate with shallowwidely spaced teeth and ciliate with red multicellular trichomes,apex acuminate. Inflorescences capitate; bracts variablein size and shape; outer bracts sterile, lanceolate, ca. 3 ×0.6 cm, densely pubescent with purplish trichomes, margin serrate;inner bracts lanceolate, ca. 1.6 × 0.4 cm, margin entire andciliate, apex acuminate; bracteoles lanceolate, 8–12 × ca. 2 mm,pubescent. Calyx ca. 1.5 cm, 5-lobed almost to base; lobes 10–13 × ca. 1.5 mm, slightly unequal, outside pubescent with purplishtrichomes especially apically, inside glabrous. Corolla purple,4–6 cm, subventricose, outside pubescent with gland-tippedtrichomes, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for 0.6–1.6 cm thenbent to ca. 90º and gradually widened to ca. 1.7 cm at mouth;lobes ovate, ca. 4 × 7 mm, apex retuse. Stamens 4, included;filaments glabrous, shorter pair ca. 1.5 mm, longer pair 7–9mm and unequal; anther thecae obliquely ellipsoid, ca. 1.5 mm;pollen type 3. Ovary glabrous; style ca. 3 cm, glabrous. Capsulefusiform, ca. 12 × 3 mm, glabrous, 4-seeded. Seeds ovate inoutline, ca. 2 × 2 mm, densely pubescent; areola small. Fl.Aug–Oct.Forests; 1000–1500 m. Xizang (Cona) [India].Strobilanthes glomerata is introduced in Indonesia.102. Strobilanthes capitata (Nees) T. Anderson, J. Linn. Soc.,Bot. 9: 475. 1867.头花马蓝 tou hua ma lanGoldfussia capitata Nees in Wallich, Pl. Asiat. Rar. 3: 88.1832.

ACANTHACEAE 421Subshrubs or perennial herbs, to 1 m tall, much branched,anisophyllous. Stems decumbent to ascending, pilose, glabrescent.Petiole 1–3 cm; leaf blade lanceolate, elliptic-ovate, orelliptic, 4–14 × 1.5–9 cm, abaxially pubescent along veins butoften glabrescent, adaxially sparsely pilose and with numerouscystoliths, secondary veins 6 or 7 on each side of midvein, baseattenuate and oblique, margin serrate, apex acuminate. Inflorescencesaxillary, pedunculate heads, ovoid to ellipsoid, 1.5–2 ×1–3 cm, 1–3 per axil, sometimes forming a panicle of headswith reduced leaves; peduncle 1–6 cm, glabrous or pubescent,simple or branched, sometimes with a sterile bract atbranching point; outer floral bracts sterile, ovate, 1.5–2.3 cm,not overtopping capitula, concave, deciduous as flowers open,pubescent or glabrous, margin crenate, apex obtuse, acute, oracuminate; inner bracts oblong-elliptic, 1.3–1.6 cm, margin entireor with 1 or 2 teeth; bracteoles oblong-elliptic, 7–11 mm,caducous, pilose. Calyx pale green, ca. 1.2 cm, accrescent to ca.2 cm in fruit, 5-lobed almost to base; lobes linear-oblanceolate,equal, outside pubescent, margin ciliate. Corolla blue, 4.3–5.5cm, straight, outside thinly pilose, inside glabrous except for trichomesretaining style; tube basally cylindric and narrow for ca.1.5 cm then gradually widened to ca. 1.5 cm at mouth; lobesoblong, ca. 5 × 4 mm. Stamens 4, included; filaments glabrous,shorter pair ca. 2 mm and nodding, longer pair 7–8 mm andunequal; anther thecae spherical, ca. 1.2 mm in diam., incurved;pollen type 3. Ovary gland-tipped pilose; style ca. 3.5 cm,thinly pilose. Capsule oblong, 1.4–2 cm, apical half glandtippedpilose, 4-seeded. Seeds ovate in outline, ca. 3 × 2.5 mm,pubescent; areola small. Fl. Sep–Feb.Forests; 900–1700 m. Xizang (Mainling, Nyalam) [Bhutan, India,Myanmar, Nepal].103. Strobilanthes speciosa Blume, Bijdr. 799. 1826.美丽马蓝 mei li ma lanBaphicacanthus multibractealatus Hung T. Chang & H.Zhu; Goldfussia speciosa (Blume) Bremekamp.Subshrubs or herbs, to 1.2 m tall, perennial, anisophyllous.Stems ascending, apically pubescent. Petiole 0.5–2 cm, hirsute;leaf blade of smaller pair ovate to suborbicular and ca. 3 × 1.6cm and of larger pair ovate and 12.5–16.5 × 6–8 cm, both surfaceshirsute, adaxially with numerous prominent linear cystoliths,secondary veins 7–9 on each side of midvein, baseoblique and decurrent onto petiole, margin irregularly crenatedentate,apex acute to caudate. Inflorescences axillary, heads,narrowly ovoid, 1–2.5 × 1–1.5 cm, borne on simple or 3-furcatebranches; peduncle 1–3 cm, simple or branched and sometimeswith a sterile bract at branching point, hirsute; sterile bractsresembling small leaves; floral bracts lanceolate, 1.2–2.3 × 0.7–1 cm, overtopping capitula, abaxially brownish pubescent withgland-tipped trichomes, margin entire and ciliate, apex acuminateand falcate; bracteoles oblanceolate, 7–13 × ca. 2 mm,abaxially pilose, margin ciliate. Calyx outside hirsute, 5-lobedalmost to base; lobes oblanceolate, 7–15 × ca. 0.2 mm, subequal,margin ciliate, apex acute. Corolla purplish blue, 3.5–5cm, ventricose, outside gland-tipped pilose, inside glabrous;tube basally cylindric and ca. 2.5 mm wide for ca. 1 cm thengradually widened to ca. 1.5 cm at mouth; lobes ca. 5 × 4 mm,apex rounded. Stamens 4, included; filaments glabrous, shorterpair 1–2 mm and nodding, longer pair 4–6 mm; anther thecaespherical, ca. 1.2 mm in diam., incurved; pollen type 3. Ovarycomose; style ca. 3.7 cm; stigma unequally 2-lobed. Capsulefusiform, 1.5–1.8 × ca. 0.4 cm, apically with a few gland-tippedtrichomes, 4-seeded. Seeds brown, ovate in outline, ca. 3 × 2.5mm, appressed pilose. Fl. Sep–Dec, fr. Jan.Forests; 500–1800 m. Yunnan [Cambodia, Indonesia, Laos, Malaysia,Thailand, Vietnam].This species intergrades with Strobilanthes capitata, from which itdiffers in little more than the narrower heads overtopped by the floralbracts and may not be distinct.Strobilanthes speciosa has been misidentified as Goldfussia glomerataNees by C. C. Hu (FRPS 70: 164. 2002).104. Strobilanthes kingdonii J. R. I. Wood, Edinburgh J. Bot.51: 244. 1994.合页草 he ye caoSubshrubs 0.5–3 m tall, branched, slightly anisophyllous.Stems 4-angled, bifariously pilose, apically glabrescent. Petiole0.5–2 cm, pubescent; leaf blade ovate, 5–12 × 1.5–5 cm, bothsurfaces pubescent, abaxially slightly paler, secondary veins 5–8 on each side of midvein, base long attenuate, margin crenateto serrate, apex acute to acuminate and slightly falcate. Inflorescencesaxillary, dense headlike spikes, borne on a short usuallysimple or (2 or)3-furcate peduncle; peduncle 1–3 cm, pilose;rachis sparsely pilose; outer sterile bracts leaflike, lanceolateto narrowly oblong-elliptic, 1.3–2.7 × 0.3–0.8 cm, pubescent,margin dentate; floral bracts ovate, ovate-elliptic, or subrhomboidal,7–10 × 2–4 mm, persistent, abaxially ciliate withlong non-glandular white trichomes and scattered shorter whitetrichomes, apex obtuse; bracteoles linear-oblanceolate, 4–8 ×1.5–2 mm, persistent, margin ciliate with long white trichomesand sometimes stalked glands at apex, apex obtuse to acute.Calyx outside densely pilose with scattered long white trichomesmixed with numerous gland-tipped trichomes, insideglabrous, subequally 5-lobed to base, not accrescent; lobes narrowlyoblong to lanceolate, 7–9 × ca. 1.5 mm, apex acute. Corollapurple, blue, or rose, 3.5–4.2 cm, straight, outside pilose,inside glabrous except for trichomes retaining style; tube basallycylindric and 1.5–2 mm wide for ca. 8 mm then ventricoseand widened to ca. 1.2 cm at mouth; lobes broadly ovate, ca. 7× 6 mm, apex rounded. Stamens 4, included; shorter filamentpair 1.5–3.5 mm, glabrous; longer filament pair 6–8 mm, pilose;anther thecae shortly ellipsoid, ca. 1.2 mm, incurved;pollen type 3. Ovary apically gland-tipped pilose; style 1.7–1.9mm, sparsely pilose. Capsule oblong, ca. 1 cm, gland-tipped pilose,4-seeded. Seeds ca. 3 × 1.8 mm, pubescent. Fl. Mar, Jul,Nov.● Evergreen broad-leaved forests by streams; 1500–2800 m. Xizang(Mêdog, Zayü), Yunnan (Fugong).Strobilanthes kingdonii was mistakenly treated as Sympagis monadelpha(Nees) Bremekamp by C. C. Hu (FRPS 70: 176. 2002) and C.C. Hu and X. P. Fu (Higher Pl. China 10: 378. 2004) and also asSympagis petiolaris (Nees) Bremekamp by C. C. Hu (FRPS 70: 178.2002).

422ACANTHACEAE105. Strobilanthes lamiifolia (Nees) T. Anderson, J. Linn.Soc., Bot. 9: 476. 1867.蒙自马蓝 meng zi ma lanGoldfussia lamiifolia Nees in Wallich, Pl. Asiat. Rar. 3: 88.1832; G. austinii (C. B. Clarke ex W. W. Smith) Bremekamp; G.feddei (H. Léveillé) E. Hossain; G. hancockii (C. B. Clarke exW. W. Smith) Bremekamp; G. mahongensis (H. Léveillé) E.Hossain; G. rotundifolia (D. Don) Bremekamp; Ruellia rotundifoliaD. Don; Strobilanthes austinii C. B. Clarke ex W. W.Smith; S. feddei H. Léveillé; S. hancockii C. B. Clarke ex W. W.Smith; S. mahongensis H. Léveillé.Herbs 30–60 cm tall, erect, much branched, isophyllous.Stems slender, decumbent and often rooting at nodes, bifariouslypilose or glabrescent. Petiole 0–3 cm but apically leavessessile, densely yellowish brown pilose; leaf blade elliptic,ovate, obovate, or ovate-lanceolate, 1–7(–11) × 1–4(–6) cm,abaxially glabrous or pilose, adaxially dark green and glabrousor sparsely pilose, secondary veins 4–7 on each side of midvein,base cuneate and often decurrent onto petiole, margin serrulateto crenulate, apex acute. Inflorescences terminal or axillaryfrom apical leaf axils, spikes, 2–5 cm; peduncle 0–6 cm,pubescent; sterile bracts resembling reduced leaves; floral bractsimbricate, oblong to ovate-elliptic, 1.1–2.5 cm, concave, caducousor somewhat persistent after flowering, dull green, abaxiallypubescent and covered with cystoliths, margin usuallycrenulate; bracteoles narrowly oblong-elliptic, 6–7 × 1–2 mm,scarious and caducous, abaxially puberulent. Calyx 0.8–1.4 cm,outside gland-tipped ciliate, 5-lobed almost to base; lobes linear-oblongwith one distinctly longer than others, apex acute.Corolla purplish red to blue, 3.5–5 cm, ventricose, outside pubescent;tube basally cylindric and ca. 2 mm wide for ca. 1.2cm then slightly curved and gradually widened to ca. 1.2 cm atmouth; lobes orbicular, 5–6 mm in diam. Stamens 4, included;filaments subglabrous, shorter pair ca. 2 mm and strongly reflexed,longer pair ca. 7 mm; anther thecae subspherical, ca. 1.2mm; pollen type 3. Ovary comose; style 3.2–3.5 cm. Capsulefusiform, ca. 1.2 cm, glabrous except for few trichomes at tip,4-seeded. Seeds light brown, ovate in outline, ca. 2.5 × 2 mm,densely pubescent; areola small, glabrous. Fl. Aug–Nov.Dry grasslands and open Pinus forests; 1000–2600 m. Guizhou,Sichuan, SE Xizang, Yunnan [Bhutan, India, Nepal].Strobilanthes lamiifolia was misidentified as S. oligocephala byQ. H. Chen (Fl. Guizhou. 10: 257. 2004).106. Strobilanthes penstemonoides (Nees) T. Anderson, J.Linn. Soc., Bot. 9: 477. 1867 [“pentstemonoides”].圆苞马蓝 yuan bao ma lanGoldfussia penstemonoides Nees in Wallich, Pl. Asiat. Rar.3: 88. 1832 [“pentastemonoides”]; G. flexuosa Nees; Ruelliacapitata Buchanan-Hamilton ex D. Don.Herbs to 1 m tall, anisophyllous. Stems 4-angled, sulcate,subglabrous. Leaves subsessile; leaf blade elliptic to ellipticlanceolate,4–15 × 1–4 cm, abaxially glabrous but sometimespubescent along veins, adaxially glabrous, secondary veins 5–7on each side of midvein, base cuneate, margin serrulate, apexacute. Inflorescences terminal, abbreviated headlike spikes, simpleor 3-furcately branched, sometimes with 1 or a few distantflowers; peduncle gland-tipped pilose; bracts orbicular, 6–9 mmin diam., unequal in size, soon scarious and caducous, glabrous;bracteoles absent. Calyx 0.5–1.2 cm, strongly accrescent infruit, 5-lobed almost to base; lobes linear, unequal with oneslightly longer than others, outside gland-tipped pilose. Corollaviolet, ca. 3.5 cm, slightly curved, outside glabrous, inside glabrousexcept for trichomes retaining style; tube basally cylindricand ca. 2 mm wide for ca. 8 mm then gradually widened toca. 1.2 cm at mouth; lobes ovate, ca. 4 × 4 mm. Stamens 4,included; filaments subglabrous, shorter pair ca. 4 mm andcurved, longer pair 7–8 mm; anther thecae spherical, ca. 1.3mm in diam., incurved; pollen type 3. Ovary pilose at tip; styleca. 2.8 cm, sparsely hirsute. Capsule clavate, ca. 1.2 cm, glandtippedpubescent, 4-seeded. Seeds suborbicular in outline, ca.2.5 mm in diam., pubescent. Fl. Aug–Nov, fr. Sep–Dec.Mountain slopes; 2100–2300 m. SE Xizang, Yunnan [Bhutan, India,Nepal].107. Strobilanthes ningmingensis D. Fang & H. S. Lo, Guihaia17: 39. 1997.宁明马蓝 ning ming ma lanGoldfussia ningmingensis (D. Fang & H. S. Lo) C. Y. Wuex C. C. Hu.Herbs or subshrubs, to 40 cm tall, branched, anisophyllous.Stems 4-angled, sulcate, glabrous except for pubescentnodes. Petiole 3–7 cm; leaf blade ovate-elliptic, smaller of pair1–1.7 × 0.7–1.2 cm and larger one 3–5.5 × 2–3 cm, both surfacesglabrous, abaxially cystoliths prominent, whitish, anddense along veins, adaxially with prominent linear cystoliths,secondary veins 4–6 on each side of midvein, base subrounded,margin serrulate, apex acute to acuminate. Inflorescences axillaryor terminal, headlike simple spikes, 3- or 4-flowered; peduncleca. 3 cm, apically thickened; bracts elliptic, 6–12 mm,caducous, abaxially densely covered with cystoliths, marginciliate, apex acute; bracteoles linear, ca. 5 × 1 mm, caducous,abaxially sparsely hirsute. Calyx 5-lobed almost to base; lobesoblong-elliptic, 6–7 × ca. 1.5 mm, slightly unequal with onelonger than others, outside glabrous, margin ciliate. Corollapurple, ca. 4 cm, ventricose, outside glabrous, inside glabrousexcept for trichomes retaining style; tube basally cylindric andca. 2.5 mm wide for ca. 1.2 cm then gradually widened to ca.1.3 cm at mouth; lobes orbicular, ca. 6 mm in diam., apexemarginate. Stamens 4, included; filaments glabrous; shorter filamentpair ca. 3 mm, equal, incurved; longer filament pair6–8 mm, unequal, erect; anther thecae subspherical, ca. 1.3 mmin diam., nodding. Ovary sparsely gland-tipped pilose at tip;style ca. 3 cm, sparsely gland-tipped pilose at base. Capsulenot seen. Fl. Dec.● Limestone hills; 200–300 m. Guangxi (Ningming).Known only from the type, Strobilanthes ningmingensis is veryclose to, and may be conspecific with, the very variable S. dimorphotricha,differing only in the non-glandular calyx, which may be only adevelopment stage.

ACANTHACEAE 423108. Strobilanthes dimorphotricha Hance, J. Bot. 21: 355.1883.球花马蓝 qiu hua ma lanHerbs 40–150 cm tall, perennial, much branched, isophyllousor anisophyllous. Stems apically somewhat zigzag, glabrousor yellowish pubescent. Petiole 0.5–1.5 cm, sparsely pubescentor glabrous; leaf blade elliptic, elliptic-lanceolate,ovate, oblong, or oblong-elliptic, 1.5–15 × 0.5–4.5 cm, smallerof pair 1/3–2/3 size of larger one, adaxially dark green, whiteappressed pubescent, and with slightly prominent cystoliths,abaxially glabrous or whitish pubescent along veins and with orwithout multicellular trichomes, secondary veins 5–7 on eachside of midvein and sometimes forming a net near margin, basecuneate to attenuate, margin serrulate, apex acuminate, caudate,or acute. Inflorescences axillary or terminal, headlike spikes,subspherical, very compact, 2- or 3-flowered, usually 1–3 spikesforming a forked panicle; peduncle ca. 2.5 cm, glabrous or pubescent;bracts ovate-elliptic, 1–1.3 cm, caducous, glabrous,apex acute to acuminate; bracteoles lanceolate, ca. 5 mm, caducous.Calyx 7–9 mm, accrescent to 1.5–1.7 cm, 5-lobed almostto base; lobes lanceolate, gland-tipped pubescent, one lobeslightly longer than others. Corolla violet, 3–4 cm, slightlycurved, outside subglabrous to sparsely pilose, inside glabrousexcept for trichomes retaining style; tube basally cylindric andca. 2 mm wide for ca. 5 mm then ventricose and graduallywidened to ca. 1.5 cm at mouth; lobes orbicular, 5–6 mm indiam., subequal, apex emarginate. Stamens 4, included; filamentsglabrous; shorter filament pair ca. 2 mm, equal, incurved;longer filament pair 5–7 mm, unequal, erect; anther thecaespherical, ca. 1 mm in diam.; pollen type 3. Ovary pilose; styleca. 3.5 cm, pilose. Capsule oblong-clavate, 1.4–1.8 cm, glandtippedpubescent, 4-seeded. Seeds ca. 2.5 × 2.5 mm, lenticular,pubescent. Fl. Aug–Feb(–Apr).Thickets on limestone hills, streamsides, thickets by streams; 200–2200 m. Chongqing, Fujian, Guangdong, Guangxi, Guizhou, Hainan,Hubei, Hunan, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [India,Laos, Myanmar, Thailand, Vietnam].Strobilanthes dimorphotricha has mistakenly been considered tobe synonymous with Goldfussia (= Strobilanthes) penstemonoides bymany authors.1a. Leaf blade ovate, elliptic, or ellipticlanceolate,pairs at each node quiteunequal in size, apex acuminateto caudate .............................. 108a. subsp. dimorphotricha1b. Leaf blade oblong to oblongelliptic,pairs at each nodeequal to subequal in size,apex acute .................................................. 108b. subsp. rex108a. Strobilanthes dimorphotricha subsp. dimorphotricha球花马蓝 ( 原亚种 ) qiu hua ma lan (yuan ya zhong)Goldfussia chaffanjonii (H. Léveillé) E. Hossain; G. dimorphotricha(Hance) Bremekamp; G. equitans (H. Léveillé) E.Hossain; G. geniculata (C. B. Clarke) Bremekamp; G. psilostachys(C. B. Clarke ex W. W. Smith) Bremekamp; G. seguinii(H. Léveillé) C. Y. Wu & C. C. Hu; Strobilanthes burkilii Dunn;S. chaffanjonii H. Léveillé; S. equitans H. Léveillé; S. geniculataC. B. Clarke; S. geniculata var. integra C. B. Clarke; S.gracilis T. Anderson (1867), not Beddome (1864); S. laxicalyxHayata; S. marchandii H. Léveillé; S. psilostachys C. B. Clarkeex W. W. Smith; S. seguinii H. Léveillé.Leaf blade ovate, elliptic, or elliptic-lanceolate, pairs ateach node quite unequal in size, apex acuminate to caudate. Fl.Aug–Nov(–Apr).Thickets on limestone hills, streamsides; 200–2200 m. Chongqing,Fujian, Guangdong, Guangxi, Guizhou, Hainan, Hubei, Hunan,Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [India, Vietnam].108b. Strobilanthes dimorphotricha subsp. rex (C. B. Clarke)J. R. I. Wood, Kew Bull. 61: 19. 2006.泰国马蓝 tai guo ma lanStrobilanthes rex C. B. Clarke, Bot. Jahrb. Syst. 41: 68.1907; Goldfussia anfractuosa (C. B. Clarke) Bremekamp; G.rex (C. B. Clarke) Bremekamp; S. anfractuosa C. B. Clarke;S. penstemonoides (Nees) T. Anderson var. anfractuosa (C. B.Clarke) Benoist; S. penstemonoides var. rex (C. B. Clarke) Benoist.Leaf blade oblong to oblong-elliptic, pairs at each nodeequal to subequal in size, apex acute. Fl. Dec–Feb.Thickets by streams; 600–1600 m. Yunnan [Laos, Myanmar, Thailand,Vietnam].109. Strobilanthes glandibracteata D. Fang & H. S. Lo, Guihaia17: 38. 1997.腺苞马蓝 xian bao ma lanGoldfussia glandibracteata (D. Fang & H. S. Lo) C. Y.Wu ex C. C. Hu.Herbs to 50 cm tall, anisophyllous. Stems 4-angled, sulcate,bifariously pubescent. Petiole (0–)3–7 mm, sparsely pubescent;leaf blade ovate-elliptic to rarely narrowly ovate orlanceolate, smaller of pair 2–3.5 × 0.4–2.9 cm and larger one5.5–12.5 × 2–4.5 cm, glabrous or sparsely pubescent alongveins, cystoliths prominent, secondary veins 4–7 on each sideof midvein, base attenuate, margin serrulate, apex caudate toacuminate. Inflorescences axillary, pedunculate heads; peduncle0.6–3.5 cm, simple or 3-furcately branched, sulcate, bifariouslypubescent, apically thickened; bracts elliptic to ovateelliptic,6–13 mm, caducous, multicellular gland-tipped pubescent,abaxially densely covered with cystoliths, apex acute;bracteoles lanceolate, ca. 5 × 1 mm. Calyx 5-lobed almost tobase; lobes lanceolate to linear, 6–8 mm, unequal with onelonger than others, outside multicellular gland-tipped pubescent.Corolla purple, ca. 3.5 cm, outside sparsely gland-tippedpilose but subglabrous at anthesis, inside glabrous except fortrichomes retaining style; tube basally cylindric and ca. 2 mmwide for ca. 8 mm then gradually widened to ca. 1.2 cm atmouth; lobes ovate, ca. 3 mm, apex emarginate. Stamens 4,included; filaments pilose; short filament pair ca. 2 mm, equal,incurved; longer filament pair 7–8 mm, unequal; anther thecaesubspherical, ca. 1.3 mm in diam., nodding. Ovary brownishgland-tipped pilose; style ca. 2.8 cm, gland-tipped pilose. Capsulenot seen. Fl. Oct.

ACANTHACEAE 425tube basally cylindric and ca. 1 mm wide for ca. 1 cm thengradually widened to ca. 9 mm at mouth; lobes ovate, ca. 5 × 5mm, apex obtuse. Stamens 4; filaments finely pubescent, shorterpair ca. 4 mm, longer pair 7–8 mm, unequal, and slightly exserted;anther thecae spherical, ca. 1.3 mm in diam., incurved;pollen type 3. Ovary gland-tipped pilose at tip; style ca. 2.5 cm,pilose. Capsule oblong, 1.1–1.3 cm, gland-tipped pubescent onapical half especially near tip, 4-seeded. Seeds ovate in outline,ca. 3 × 3 mm, densely covered with appressed trichomes. Fl.Dec–Feb.● Forests; 600–1000 m. Xizang (Mêdog).114. Strobilanthes discolor (Nees) T. Anderson, J. Linn. Soc.,Bot. 9: 477. 1867.异色马蓝 yi se ma lanGoldfussia discolor Nees in A. Candolle, Prodr. 11: 172.1847; Diflugossa nagaensis Bremekamp; Strobilanthes discolorvar. nudicalyx C. B. Clarke.Subshrubs to 1.8 m tall, strongly anisophyllous. Stems andbranches 4-angled to subterete, sulcate, glabrous, apically somewhatzigzag. Petiole 0.5–2.5 cm, sulcate, glabrous; leaf bladeelliptic to ovate-elliptic, 3–14 × 2–6.5 cm, glabrous, both surfaceswith prominent cystoliths, abaxially whitish, adaxiallydark green, secondary veins 6–8 on each side of midvein, baseattenuate, shortly decurrent onto petiole, and often slightlyoblique, margin serrate, apex acuminate to caudate and oftenfalcate. Inflorescences axillary, pedunculate heads, often withsecondary branching and becoming paniculate; floral bracts oblanceolateto spatulate, 3–5 × 1–2 mm, rapidly caducous, glabrous;bracteoles oblanceolate but often apparently absent. Calyx5-lobed almost to base; lobes at anthesis linear-oblong, 5–7× ca. 1 mm, subequal, apically pilose but otherwise glabrous,apex obtuse; lobes in fruit subspatulate, accrescent to ca. 1 cm,often gland-tipped pilose. Corolla violet, 3.2–3.8 cm, outsideglabrous, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for 8–10 mm thengradually widened to ca. 1 cm at mouth; lobes ovate, 5–6 × 6–7mm, apex rounded. Stamens 4; shorter filament pair 1.5–2 mm,strongly reflexed, glabrous; longer filament pair 7–8 mm, glabrousor with a few trichomes; anther thecae obliquely ellipsoid,ca. 1.2 mm, incurved; pollen type 3. Ovary glabrous; style2.3–2.8 cm, hirsute. Capsule narrowly obovoid, ca. 10 × 2.5mm, glabrous except for a few apical glands, 4-seeded. Seedsovate in outline, ca. 3 × 2 mm, pilose. Fl. Nov–Feb. 2n = 22.Forests. Xizang (Cona) [India].115. Strobilanthes rhombifolia C. B. Clarke in J. D. Hooker,Fl. Brit. India 4: 461. 1884.短柄马蓝 duan bing ma lanGoldfussia sessilis Nees.Herbs or subshrubs, strongly anisophyllous. Stems 4-angled,erect, glabrous. Petiole 0–10 mm but apical leaves sessile;leaf blade rhomboidal-elliptic, 2–15 × 1.5–10 cm, both surfacesgreen and with abundant cystoliths, secondary veins ca. 6 oneach side of midvein, base cuneate and shortly decurrent ontopetiole, margin serrate, apex acute. Inflorescences axillary, capitulaborne on simple or 2(or 3)-furcately branched peduncles,2–10 cm; peduncle sulcate, bifariously pubescent with trichomesin sulci; sterile bracts leaflike, ovate, ca. 0.5 × 0.5 mm,relatively persistent; floral bracts obovate to oblong-obovate,ca. 6 × 1.5 mm, soon scarious and caducous; bracteoles not observed.Calyx subequally 5-lobed to base; lobes linear-elliptic,7–12 × ca. 1.2 mm, pubescent and with sessile glands. Corollablue, 4–4.5 cm, outside sparsely gland-tipped pilose but glabrescent,inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for ca. 1 cm thenwidened to 1.2–1.7 cm at mouth; lobes suborbicular, ca. 6 × 7mm. Stamens 4, included; filaments glabrous, shorter pair ca. 2mm, longer pair ca. 5 mm; anther thecae obliquely ovoid, ca.1.5 × 1.2 mm, incurved. Ovary narrowly ovoid, apicallygland-tipped; style ca. 2.7 cm, slender, glabrous apart from afew trichomes on basal part. Capsule not seen. Fl. Dec.Forests. Xizang (Cona) [India].116. Strobilanthes multidens C. B. Clarke in J. D. Hooker, Fl.Brit. India 4: 461. 1884.分枝马蓝 fen zhi ma lanGoldfussia multidens (C. B. Clarke) Bremekamp; Pteracanthusagrestis (C. B. Clarke) Bremekamp; Strobilanthesagrestis C. B. Clarke; S. agrestis var. hemiotis C. B. Clarke; S.penstemonoides (Nees) T. Anderson var. multidens (C. B.Clarke) Kuntze.Subshrubs to 2 m tall, usually pubescent, strongly anisophyllous.Leaf blade ovate-elliptic, 2–20 × 1–9.5 cm, usuallypubescent at least on veins, base attenuate and oblique, marginserrate, apex shortly acuminate and sometimes falcate. Inflorescencesaxillary, usually capitate, borne on 3-furcate axillarygland-tipped pilose peduncles, sometimes with scattered flowerpairs below heads, rarely of branched spikes with flower pairsonly; bracts ovate-suborbicular, 3–6 mm, concave, caducous,gland-tipped pilose, apex acute; bracteoles absent. Calyx 6–10mm at anthesis, gland-tipped pubescent; lobes linear, accrescentto ca. 1.6 cm with one lobe 2–3 mm longer than others, apexacute. Corolla blue, 3.5–4.8 cm, outside glabrous, inside glabrousexcept for trichomes retaining style; tube basally cylindricand narrow then widened to ca. 1.5 cm at mouth. Stamens 4;shorter filament pair ca. 1 mm, recurved; longer filament pair5–6 mm, slightly unequal, glabrous; anther thecae subspherical,ca. 1 mm, incurved; pollen type 3. Ovary gland-tipped; style ca.2.5 cm, glabrous except for a few trichomes basally. Capsuleoblong, 13–18 × ca. 3 mm, apically gland-tipped pilose, 4-seeded. Seeds ovate in outline, ca. 3 × 2 mm, appressed pubescent.Fl. Sep–Apr.Streamsides; 1500–2000 m. Xizang (Cona) [Bhutan, India, Nepal].117. Strobilanthes oligocephala T. Anderson ex C. B. Clarkein J. D. Hooker, Fl. Brit. India 4: 461. 1884.菱叶马蓝 ling ye ma lanDiflugossa paupera (C. B. Clarke) Bremekamp; Goldfussiaoligocephala (T. Anderson ex C. B. Clarke) Bremekamp; G.thomsonii Hooker; Strobilanthes oligocephala var. treutleri C.B. Clarke; S. paupera C. B. Clarke.

426ACANTHACEAESubshrubs 30–50 cm tall, much branched, anisophyllous.Stems decumbent and basally rooting at nodes but eventuallyerect, pubescent to subglabrous. Petiole 0.5–4 cm but apicalmostleaves sessile; leaf blade rhombic, elliptic, narrowly ovate,or apicalmost suborbicular, 4–12 × 1.5–6 cm, abaxially glaucousand glabrous or pilose along veins and margin, adaxiallygreen, glabrous, and densely covered with prominent cystoliths,secondary veins 5–7 on each side of midvein, base attenuateand decurrent onto petiole, margin serrate, apex abruptly acuminateand falcate. Inflorescences terminal or arising from apicalleaf axil, simple or branched, subcapitate with ca. 2 flower pairsto shortly spicate, 4–5 cm; peduncle 1–5 cm, glabrous or pubescent;bracts ovate to oblong, 0.8–1.5 × 0.7–1.3 cm, caducous,green when young but soon scarious, often becoming glandtippedpilose, apex acute; bracteoles ca. 2 × 1 mm, caducous sooften appearing absent. Calyx 1–1.8 cm, gland-tipped pilose,5-lobed almost to base; lobes narrowly lanceolate with oneslightly longer than others, apex acuminate. Corolla blue orwhite, 3.5–5 cm, ventricose, outside gland-tipped pubescent butsometimes only on lobes, inside glabrous except for trichomesretaining style; tube basally cylindric and ca. 2.5 mm wide for1–1.5 cm then bent and widened to ca. 1 cm at mouth; lobesoblong, ca. 6 × 5 mm, apex obtuse. Stamens 4, included; filamentssparsely hirsute, shorter pair ca. 4 mm, longer pair ca.7 mm; anther thecae oblong-ovoid, ca. 2 × 1.2 mm. Ovarycomose; style 2.8–3.2 cm. Capsule ca. 1.4 cm, apically glandtippedpilose, 4-seeded. Seeds ovate in outline, ca. 3 × 2.5 mm,densely pubescent. Fl. Jul–Nov.Thickets on limestone hills; 2600–2800 m. Xizang (Cona) [Bhutan,India, Nepal].118. Strobilanthes medogensis (H. W. Li) J. R. I. Wood & Y. F.Deng, Bot. J. Linn. Soc. 150: 383. 2006.墨脱马蓝 mo tuo ma lanGoldfussia medogensis H. W. Li, Fl. Xizang. 4: 413. 1985.Subshrubs 2–3 m tall, anisophyllous. Stems erect, slightlysulcate, bifariously puberulent. Petiole 0–5 mm with smallerleaves subsessile; leaf blade ovate, 3–13 × 2–5.5 cm, smaller ofpair ca. 2/3 size of larger one, both surfaces glabrous, abaxiallypale green, adaxially green and with numerous cystoliths, baserounded to subcordate, margin serrulate, apex acuminate andfalcate. Inflorescences axillary, spikes, 3–6 cm, simple or 2-furcate;peduncle 2–5 cm; rachis bifariously pubescent; bractsobovate-oblanceolate, 2.5–3 mm, herbaceous, caducous, glabrous;bracteoles similar to bracts but ca. 1.5 mm. Flowers 3–11mm apart on rachis. Calyx ca. 1.2 cm in flower, 5-lobed to base;lobes narrowly oblong-elliptic, subequal, glabrous or glandtippedpubescent, apex acuminate. Corolla yellowish white butdull purple on lobes, campanulate, ca. 2.5 cm, straight, outsideglabrous, inside glabrous except for trichomes retaining style;tube basally cylindric and ca. 2 mm wide for ca. 1 cm thengradually widened to 1–1.5 cm at mouth; lobes broadly ovate,ca. 3 × 8 mm, apex rounded. Stamens 4, ± exserted; filamentsbasally pilose, apically glabrous, shorter pair 7–8 mm, longerpair 9–10 mm; anther thecae ca. 1 mm, incurved; pollen type 3.Ovary glabrous; style 2.7–2.8 cm, glabrous. Capsule not seen.Fl. Jul–Aug.● Forests; 1900–2500 m. Xizang (Mêdog).119. Strobilanthes thomsonii T. Anderson, J. Linn. Soc., Bot.9: 478. 1867.汤氏马蓝 tang shi ma lanSubshrubs 1–2 m tall, pliestesial, anisophyllous. Stems andbranches 4-angled, sulcate, glabrous or thinly pubescent. Petiole2–14 mm or leaves sessile, glabrous; leaf blade ovate to narrowlyovate-elliptic, 12–20 × 4–7 cm, abaxially pale, adaxiallyglabrous with abundant cystoliths, secondary veins 5–7 on eachside of midvein, base cuneate and decurrent onto petiole, marginserrate, apex long acuminate and usually falcate. Inflorescencesaxillary, simple or branched spikes, 3–9 cm; floral bractsvariable in shape, ovate, elliptic, oblanceolate, or spatulate, 4–6 × 2–3 mm, caducous, glabrous or gland-tipped pubescent;bracteoles linear, ca. 2.5 × 0.8 mm, caducous. Flowers imbricateon rachis or to 1 cm apart. Calyx 5–9 mm, accrescent to ca.1.3 cm in fruit, subequally 5-lobed almost to base; lobes linear,glabrous or pilose, apex acute. Corolla purple, funnel-shaped,2.5–3.2 cm, curved, outside pubescent, inside glabrous exceptfor trichomes retaining style; tube basally cylindric and ca. 2mm wide for ca. 1 cm then curved and widened to 1–1.3 cm atmouth; lobes ovate, ca. 4 × 4 mm, apex rounded. Stamens 4;filaments glabrous, shorter pair ca. 2 mm and strongly incurved,longer pair 3.5–4 mm; anther thecae subspherical, ca. 1 mm,incurved; pollen type 3. Ovary pubescent; style 2.5–3 cm, basallythinly pilose, apically glabrous. Capsule narrowly obovoid,gland-tipped pubescent, 4-seeded. Seeds orbicular in outline,ca. 2 mm in diam., densely covered with appressed trichomes.Fl. Sep–Nov.Dry forests; ca. 2500 m. Xizang (Cona) [Bhutan, India].120. Strobilanthes longgangensis D. Fang & H. S. Lo, Guihaia17: 33. 1997.弄岗马蓝 nong gang ma lanPerilepta longgangensis (D. Fang & H. S. Lo) C. Y. Wu &C. C. Hu.Subshrubs to 40 cm tall, strongly anisophyllous. Petiole 2–14 mm; leaf blade narrowly elliptic to rarely ovate-elliptic,smaller of pair (1.5–)3–8.5 × 0.8–3 cm and larger one 6–11.5 ×2.5–4.5 cm, both surfaces glabrous with prominent cystoliths,abaxially paler, secondary veins 4–6 on each side of midvein,base narrowly cuneate and decurrent onto petiole, margin denticulate,apex acuminate. Inflorescences axillary, spikes, 2–4cm, 1–5-flowered; peduncle ca. 8 mm; bracts narrowly triangular,3–4 × 1–1.5 mm, persistent, apex acute; bracteoles oblong,5–6 × ca. 1 mm, glabrous. Flowers distant on rachis, usuallyonly 1 sterile per node. Calyx ca. 5 mm, 5-lobed to nearbase; lobes resembling bracteoles, glabrous. Corolla purple,campanulate, ca. 2.5 cm, outside glabrous, inside glabrous exceptfor trichomes retaining style; tube basally cylindric andnarrow for ca. 5 mm then gradually widened to ca. 1.5 cm atmouth; lobes oblong, ca. 4 mm, apex emarginate. Stamens 4,included; filaments sparsely pilose, shorter pair ca. 2 mm,longer pair 6–8 mm; anther thecae subspherical, ca. 1 mm, incurved.Ovary glabrous; style ca. 1.8 cm, hirsute. Capsule notseen. Fl. Nov.

ACANTHACEAE 427● Limestone slopes; 300–600 m. Guangxi (Longzhou).121. Strobilanthes secunda T. Anderson, J. Linn. Soc., Bot. 9:480. 1867.偏花马蓝 pian hua ma lanShrubs 0.6–1.5 m tall, strongly but variably anisophyllous.Stems and branches 4-angled, often zigzag, sulcate, glabrous onridges but rufous scurfy pubescent in channels. Petiole 3–8 cm,appressed pubescent; leaf blade oblong-elliptic, 4–18 × 1.5–7.5cm, abaxially glabrous except for pubescent midvein, adaxiallyglabrous, secondary veins 6–9 on each side of midvein, basenarrowly cuneate, margin entire to sinuate, apex acuminate. Inflorescencesaxillary, spikes, 3–18 cm, lax; sterile bracts resemblingreduced leaves; floral bracts and bracteoles caducous,presence only known from presence of scars. Flowers solitaryor in opposite pairs, 1.5–3 cm apart on rachis. Pedicel ca. 1 mm.Calyx 1.6–2 cm, subequally lobed to 3–4 mm above base; lobeslinear-lanceolate, 1.2–1.6 cm, glabrous, apex finely acuminate.Corolla yellow, 2.6–3 cm, curved and somewhat ventricose, outsidepubescent, inside glabrous except for trichomes retainingstyle; tube basally cylindric and 2–3 mm wide for ca. 6 mmthen gradually widened to ca. 1.5 cm at mouth; lobes broadlyovate, ca. 9 × 4 mm, apex rounded. Stamens 4, included; shorterfilament pair ca. 3 mm, glabrous; longer filament pair ca. 5 mm,basally villous; anther thecae oblong, ca. 2 × 1 mm, erect; pollentype 3. Ovary clavate, comose; style ca. 2.6 cm, pilose. Capsulenarrowly oblong, ca. 20 × 2–2.5 mm, very shortly pubescent,4-seeded, apex acute. Seeds ovate in outline, ca. 3.5 × 2.5mm, pilose; areola densely pilose. Fl. Dec–Jan, fr. Jan–Apr.Forests. Xizang (Mêdog) [Myanmar].122. Strobilanthes mastersii T. Anderson, J. Linn. Soc., Bot. 9:481. 1867.瑞丽叉花草 rui li cha hua caoDiflugossa scoriarum (W. W. Smith) E. Hossain; D.shweliensis (W. W. Smith) E. Hossain; Goldfussia scoriarum(W. W. Smith) Bremekamp; Strobilanthes aborensis Dunn; S.scoriarum W. W. Smith; S. shweliensis W. W. Smith.Herbs 1–1.5 m tall, anisophyllous. Stems 4-angled, erect,sulcate, glabrous to gland-tipped pilose. Petiole 0–2 cm withapical leaves sessile, subglabrous; leaf blade ovate to ovatelanceolate,10–15 × 4–7 cm, glabrous except for a few cilia inteeth sinus, secondary veins 5–10 on each side of midvein, basebroadly cuneate on basal leaves and rounded on apical leaves,margin serrate, apex acuminate. Inflorescences terminal, paniclesof spikes, narrowly ovoid, to 30 cm; peduncle and rachisglabrous when young, densely gland-tipped pilose at maturity;sterile bracts leaflike, lanceolate to ovate, 5–15 × 3–8 mm,tardily caducous, apex acuminate; floral bracts elliptic, 5–6 ×2–3 mm, caducous, apex obtuse; bracteoles oblong, ca. 3 × 1mm, caducous, abaxially glabrous. Flowers in opposite pairs orsolitary through abortion. Calyx 0.8–1.2 cm, subglabrous withsessile glands when young but becoming densely gland-tippedpilose and accrescent, 5-lobed almost to base; lobes lanceolate,with 3 lobes slightly longer than other 2, apex acuminate. Corollapurple, 2.5–3 cm, straight, ventricose, outside glabrous, insideglabrous except for trichomes retaining style; tube basallycylindric and ca. 2 mm wide for 1–1.3 cm then widened to ca. 1cm at mouth; lobes orbicular, ca. 5 mm in diam. Stamens 4,included; filaments glabrous, shorter pair ca. 3 mm, longer pairca. 6 mm; anther thecae obliquely ellipsoid, ca. 0.8 × 1 mm,incurved; pollen type 3. Ovary slightly pilose at tip; style ca. 2.1cm, slender, sparsely pilose on basal part. Capsule clavate, ca.1.5 cm, apically gland-tipped pilose, 4-seeded. Seeds brown,ovate in outline, ca. 2.5 × 2 mm; areola small, verrucose. Fl.Sep–Jan.Forests on mountain slopes; ca. 1500 m. Xizang (Mêdog), Yunnan[India].Strobilanthes mastersii, like S. hamiltoniana, commonly has a glabrousinflorescence when it is young but becomes gland-tipped pilose asit matures.123. Strobilanthes tenax Dunn, Bull. Misc. Inform. Kew 1920:209. 1920.结壮马蓝 jie zhuang ma lanSubshrubs to 1 m tall. Stems 4-angled, sulcate, glandtippedpubescent. Petiole 2–5 mm but apical leaves sessile; leafblade lanceolate, 8–10 × 2.5–4 cm, glabrous, cystoliths numerous,linear, and prominent, secondary veins 4–6 on each sideof midvein, base cuneate, margin serrate, apex acuminate. Inflorescencesterminal, a panicle of branched spikes, 3–12 cm; rachisgland-tipped pilose; sterile bracts sessile, leaflike, oblonglanceolateto ovate, 8–12 × 3–5 mm; floral bracts oblongelliptic,5–7 × ca. 2 mm, scarious and early caducous, sparselygland-tipped pilose, apex obtuse; bracteoles apparently absent.Flowers sessile, with opposite pairs 1–2.5 cm apart on rachis.Calyx ca. 1 cm, outside gland-tipped pubescent, 5-lobed almostto base; lobes linear, apex acute. Corolla color unknown, ca. 3cm, straight, outside glabrous, inside glabrous except for trichomesretaining style; tube basally cylindric and 1–2 mm widefor ca. 0.8 cm then abruptly widened to ca. 1 cm at mouth;lobes broadly ovate, 5–6 × ca. 3 mm, apex rounded. Stamens 4,included; filaments glabrous, shorter pair 1–2 mm, longer pair3–4 mm; anther thecae obliquely ellipsoid, ca. 1.5 mm, incurved;pollen type 3. Ovary obovoid, ca. 2.5 mm, gland-tippedwith a few apical trichomes; style ca. 1.9 cm, thinly pilose.Capsule narrowly obovoid, ca. 1.5 cm, gland-tipped pubescent,4-seeded. Seeds immature. Fl. Mar.● Forests; ca. 800 m. Xizang (Mêdog).Strobilanthes tenax is only known from the type, and there aremany errors in the protologue.124. Strobilanthes hamiltoniana (Steudel) Bosser & Heine,Bull. Mus. Natl. Hist. Nat., B, Adansonia, Sér. 4, 10: 148. 1988.叉花草 cha hua caoRuellia hamiltoniana Steudel, Nomencl. Bot., ed. 2, 2:481. 1841, based on Goldfussia colorata Nees in Wallich, Pl.Asiat. Rar. 3: 89. 1832, not Ruellia colorata Blume (1826), norVellozo (1829); Diflugossa colorata (Nees) Bremekamp; Strobilanthescolorata (Nees) T. Anderson (1867), not Nees (1837).Shrubs 0.5–1.5 m tall, much branched, anisophyllous.Stems 4-angled, sulcate, glabrous. Leaves petiolate or apicalones sessile or subsessile; petiole (0–)0.8–5(–8) cm, glabrous;

428ACANTHACEAEleaf blade lanceolate to ovate, smaller pair at least 2/3 size oflarger one, 5–19 × 2–8.5 cm, glabrous, densely covered withlinear cystoliths, secondary veins 4–7 on each side of midvein,base oblique to rarely slightly oblique and attenuate or on verylarge leaves subcordate, margin serrulate, apex acuminate. Inflorescencesaxillary or terminal, panicles, to 30 cm, muchbranched; rachis angled at ca. 60°, glabrous or rarely pubescent,gland-tipped in fruit; bracts green, broadly obovate, 3–5 × ca. 2mm, concave, caducous, glabrous, becoming gland-tipped infruit, margin entire, apex retuse; bracteoles green, oblanceolateto narrowly elliptic, 3–4 × ca. 1 mm, concave, caducous, glabrous,margin entire, apex retuse. Flowers solitary on eachnode, distant on rachis. Pedicel 1.5–5 mm, glabrous. Calyx 8–10 mm, accrescent to ca. 1.3 cm in fruit, glabrous, sometimesapically gland-tipped pubescent, becoming densely glandtippedin fruit, 5-lobed almost to base; lobes narrowly ellipticwith one longer than others, apex acute to sometimes retuse.Corolla white, lilac, or (in cultivated forms) deep pink with awhite tube, 3.5–4 cm, subventricose, outside glabrous, insideglabrous except for trichomes retaining style; tube basally cylindricand ca. 2 mm wide for ca. 1.3 cm then slightly ventricoseand gradually widened to 1.4–1.8 cm at mouth; lobesovate, 4–5 × ca. 6 mm, apex rounded to retuse. Stamens 4,included; filaments glabrous; shorter filament pair ca. 1 mm,equal; longer filament pair 7–9 mm, unequal, erect; antherthecae white, spherical, ca. 1.2 mm in diam., partially recurved;pollen type 3. Ovary ca. 2.5 mm, glabrous; style ca. 3.8 cm,sparsely pubescent. Capsule green then dark purple, fusiform,1–1.5 cm, glabrous or with gland-tipped trichomes, 4-seeded.Seeds ovate in outline, ca. 3 × 2 mm, with long trichomes;areola small. Fl. Dec–Jan.Forests on mountain slopes; 800–2000 m. Xizang (Cona) [Bhutan,India, Myanmar, Nepal].Strobilanthes hamiltoniana is an ornamental plant. It is widelycultivated in tropical countries and in conservatories in temperate climates.The species is sometimes naturalized, occasionally becoming aninvasive weed as on the island of Réunion. Cultivated plants and plantsof cultivated origin seem always to be glabrous, pink-flowered, andsterile with dispersal being by cuttings or shoots broken off by grazinganimals.“Goldfussia tengyuehensis” (C. Y. Wu, Index Fl. Yunnan. 2: 1675.1984) belongs here but was not validly published.125. Strobilanthes tenuiflora J. R. I. Wood, Kew Bull. 58:691. 2003.纤序马蓝 xian xu ma lanSubshrubs to 1 m tall, anisophyllous. Stems 4-angled, sulcate,glabrous. Petiole 0.1–3.4 cm, usually very short on smallerleaves, often sulcate, glabrous; leaf blade ovate to ovate-elliptic,1–13 × 0.5–8 cm, smaller of pair 1/4–1/3 size of larger one, glabrousexcept for multicellular trichomes in teeth sinuses, adaxiallycystoliths prominent, secondary veins 5–7 on each side ofmidvein and abaxially prominent, base oblique and broadly cuneate,rounded, or slightly cordate, margin slightly serrate tosubentire, apex acuminate. Inflorescences terminal, panicles, to10 cm; peduncle to 3 cm, glabrous; rachis slender, glabrous,repeatedly 3-forked and bent to ca. 90º; bracts subulate, ca. 1mm, caducous; bracteoles similar to bracts. Flowers 1 per node.Calyx 4–6 mm, accrescent to ca. 8 mm in fruit, glabrous, 5-lobed almost to base; lobes oblong to oblong-lanceolate withone slightly longer than others, margin pale, apex emarginate.Corolla white, 3.2–4 cm, straight, outside glabrous, inside glabrousexcept for trichomes retaining style; tube basally cylindricand ca. 1.5 mm wide for 6–10 mm then slightly ventricoseand gradually widened to ca. 1.5 cm at mouth; lobes ovate, ca.5 × 5 mm, apex acute. Stamens 4, included; filaments glabrous,shorter pair ca. 0.5 mm, longer pair 4–5 mm; anther thecaespherical, ca. 1 mm in diam., partially recurved. Ovary glabrous;style 2.5–3.2 mm, glabrous. Capsule oblong, ca. 1.5 cm,glabrous, 4-seeded, apex acuminate. Seeds ovate in outline, ca.2 × 1.5 mm, puberulent. Fl. Oct.Forests; ca. 900 m. Yunnan (Cangyuan) [Thailand].126. Strobilanthes euantha J. R. I. Wood, Kew Bull. 58: 697.2003.腾冲马蓝 teng chong ma lanHerbs 0.5–1.5 m tall, perennial, slightly anisophyllous.Stems dark green, deeply sulcate and ridged, glabrous. Petiole0–2.6 cm, glabrous; leaf blade ovate to ovate-elliptic, 4–20 × 3–12 cm, abaxially paler green and sparsely pilose along veins,adaxially dark green, pilose along margin, and with prominentcystoliths, secondary veins 5–8 on each side of midvein, basecordate to abruptly narrowed and often somewhat oblique, marginserrate, apex acuminate. Inflorescences terminal, open panicles,to 15 cm; peduncle ca. 3 cm, gland-tipped pilose; rachisgland-tipped pilose; bracts obovate, 1–2 mm, caducous beforeflowers open, apex rounded; bracteoles similar to bracts. Flowersdistant on rachis, usually 1 per node. Pedicel ca. 3 mm,gland-tipped pilose. Calyx ca. 4 mm, accrescent to ca. 7 mm infruit, glabrous, 5-lobed to ca. 1 mm from base; lobes oblong tooblanceolate, apex truncate to emarginate. Corolla pale lavenderto pale pink, 2.2–3.5 cm, arcuate, glabrous; tube basally cylindricand narrow for 0.6–1.9 cm then ventricose and widenedto 8–10 mm at mouth; lobes ovate, ca. 4 × 4 mm, apex acute.Stamens 4; filaments glabrous; shorter filament pair ca. 2 mm,incurved; longer filament pair 1–1.2 cm, erect, slightly exserted;anther thecae spherical, ca. 1 mm in diam. Ovary glabrous;style 2.2–2.7 cm, glabrous. Capsule narrowly oblongobovoid,ca. 1.4 cm, glabrous, 4-seeded, apex acuminate. Seedsovate-suborbicular in outline, ca. 2.5 × 2 mm, pubescent. Fl.Sep–Oct, fr. Dec.Forests; 1500–2100 m. Yunnan (Gaoligong Shan) [Myanmar].127. Strobilanthes parvifolia J. R. I. Wood, Kew Bull. 64: 25.2009.小叶马蓝 xiao ye ma lanHerbs, perennial, anisophyllous. Stems somewhat sulcatewith distinct small parallel ridges, glabrous, basally geniculate.Petiole 0–4 mm, glabrous; leaf blade lanceolate, ovate, orovate-elliptic, 1–3.5 × 1–1.8 cm, smaller of pair ca. 1/3 size oflarger one, both surfaces glabrous and with somewhat obscurecystoliths or rarely abaxially with a few gland-tipped trichomeson veins, base broadly to narrowly cuneate and somewhat decurrentonto petiole, margin undulate to slightly crenate, apexacute. Inflorescences leaflike with long few-flowered branches;branches strict, densely covered with short stalked glands,

ACANTHACEAE 429sparingly 3-forked with subsessile flowers in opposite pairs(rarely solitary) in axils of leaflike bracts at tips of ultimatebranches; basal sterile bracts variable in size and shape, sessileor nearly so, oblong-lanceolate to sometimes ovate, elliptic, oroblong, 0.5–3.5 × 0.3–1.5 cm, often very unequal at each pair,abaxially densely stalked gland-tipped pubescent toward base,base attenuate, margin undulate and sparsely covered withstalked glands, apex acute; floral bracts oblong, ca. 5 × 1 mm,caducous, gland-tipped pilose; bracteoles oblong, ca. 2 × 1 mm.Calyx ca. 1.1 cm at anthesis, accrescent to ca. 2 cm in fruit, 5-lobed almost to base; lobes narrowly oblong-lanceolate, 1–1.5mm wide, subequal, densely covered with relatively short glandtippedtrichomes, apex obtuse. Corolla white flushed violet,2.5–3 cm, outside glabrous, inside pubescent; tube basally cylindricand ca. 3 mm wide for ca. 1 cm then gradually widenedto ca. 1.2 cm and bent to ca. 90° and then and widened to 1.2–1.5 cm at mouth; lobes broadly ovate, ca. 6 × 6 mm, apex obtuse.Stamens 4, included; filaments glabrous, shorter pair 1.5–2mm, longer pair 5–6 mm; anther thecae oblong, ca. 2 × 0.5 mm;pollen type 16. Ovary glabrous; style ca. 1.5 cm, glabrous. Capsulenarrowly clavate, ca. 2.2 cm, glabrous, 4-seeded. Matureseeds not seen. Fl. Sep, fr. Sep–Oct.● Forest margins on hills; ca. 1600 m. Xizang (Mêdog).128. Strobilanthes ferruginea D. Fang & H. S. Lo, Guihaia17: 29. 1997.锈背马蓝 xiu bei ma lanPerilepta ferruginea (D. Fang & H. S. Lo) C. Y. Wu & C.C. Hu.Shrubs to 4 m tall, almost glabrous throughout, anisophyllous.Stems 4-angled, sulcate. Petiole to 3 cm; leaf bladebroadly elliptic, elliptic, or rarely ovate-elliptic, 4–18 × 2–8 cm,smaller of pair ca. 1/3 size of larger one, both surfaces denselycovered with cystoliths, abaxially ferruginous when dry, adaxiallydark green, secondary veins 7–11 on each side of midvein,base broadly cuneate, margin apically remotely crenate, apexacuminate to caudate. Inflorescences axillary or terminal, spikes,1–2.5 cm, few flowered; bracts elliptic to oblong, 3.5–4 × ca. 2mm, persistent, margin ciliate, apex truncate to rounded; bracteolesoblong to elliptic, ca. 7 × 2.5 mm. Flowers distant on rachis,usually only 1 fertile per node. Calyx ca. 9 mm; lobesoblong-oblanceolate, ca. 4 mm, margin ciliolate, apex obtuse.Corolla purple, 3–3.5 cm, ventricose, outside glabrous; tubebasally cylindric and ca. 2 mm wide for ca. 6 mm thengradually widened to ca. 1.4 cm at mouth; lobes ca. 5 mm, apexemarginate. Stamens 4, included; filaments sparsely hirsute,shorter pair ca. 2 mm, longer pair ca. 8 mm; anther thecae oblong,ca. 2 mm. Ovary glabrous; style ca. 1.8 cm, densely hirsute.Capsule oblong, 1.7–2 cm, glabrous, 4-seeded. Seeds ovatein outline, ca. 3 mm, densely covered with appressed trichomes.Fl. Oct–Dec.● Dense forests on rocky mountains; 400–800 m. Guangxi (Longzhou).8. PHAULOPSIS Willdenow, Sp. Pl. 3: 4, 342. 1800 [“Phaylopsis”], nom. cons.肾苞草属 shen bao cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielAetheilema R. Brown; Micranthus J. C. Wendland; Theileamea Baillon.Herbs, perennial [or annual], prostrate [to erect], with cystoliths, often anisophyllous. Leaves opposite, long petiolate; leaf blademargin entire or crenate. Inflorescences terminal [or axillary], secund strobilate spikes; bracts subleaflike, orbicular or reniform,imbricate; bracteoles [present or] absent. Flowers sessile. Calyx 5-lobed almost to base; lobes unequal, posterior one largest. Corollasmall; tube cylindric, slender; limb ± 2-lipped, lower lip 3-lobed, upper lip 2-lobed; lobes contorted in bud. Stamens 4, didynamous,included in corolla tube; filaments coherent at base; anthers 2-thecous; thecae parallel, base minutely mucronate [or not mucronate].Ovary with 2 ovules per locule; style hairy; stigma 2-lobed, lobes unequal. Capsule shortly stipitate, 4-seeded; retinacula present;septa with attached retinacula separating from inner wall of mature capsule. Seeds discoid, densely covered with hygroscopictrichomes.Twenty-two species: tropical Africa, E, SE, and S Asia; one species in China.1. Phaulopsis dorsiflora (Retzius) Santapau, Kew Bull. [3]:276. 1948.肾苞草 shen bao caoRuellia dorsiflora Retzius, Observ. Bot. 6: 31. 1791;Aetheilema anisophyllum (Jussieu) E. Meyer (1837), not R.Brown (1810); A. parviflorum Sprengel; A. reniforme Nees;Blechum anisophyllum Jussieu; Micranthus dorsiflorus(Retzius) C. E. C. Fischer; M. oppositifolius J. C. Wendland;Phaulopsis oppositifolia (J. C. Wendland) Lindau; P. parvifloraWilldenow, nom. illeg. superfl.Herbs 30–50 cm tall, slightly anisophyllous. Stems ascending,4-angled, brown retrorsely pubescent. Petiole 4–6 cm;leaf blade ovate to elliptic, 7.5–11 × 3.5–5 cm, papery, abaxiallypubescent along veins, adaxially hirsute, secondary veins ca. 6on each side of midvein, base cuneate to attenuate and slightlyoblique, margin entire, apex acuminate. Spikes terminal, 3–6cm; bracts orbicular to reniform, ca. 1 × 1.4 cm, gland-tippedpubescent. Calyx lobes outside gland-tipped pubescent; posteriorlobe ovate-elliptic, 8–10 mm, bractlike, other lobes linear tosubulate and 5–6 mm. Corolla white, ca. 6 mm; lower lip 3-lobed, lobes ovate-oblong and 1–2.4 × 0.6–1 mm; upper lip narrow,2-lobed. Staminal filaments glabrous. Ovary apex sparselygland-tipped pubescent; style pilose. Capsule ellipsoid, ca. 6

430ACANTHACEAEmm, apex gland-tipped pubescent. Seeds ca. 2 × 1.5 mm. Fl.Nov–May, fr. Feb–Jul. 2n = 56, ca. 64, 70.Thickets, roadsides, along streams; 300–800 m. S and W Yunnan[Bangladesh, Bhutan, India, Myanmar, Thailand, Vietnam; ?South Africa,Indian Ocean islands (Réunion), ?Madagascar].Collections identified as Phaulopsis dorsiflora from South Africaand Madagascar remain questionable as to their source.9. HYGROPHILA R. Brown, Prodr. 479. 1810.水蓑衣属 shui suo yi shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielAdenosma Nees (1847), not R. Brown (1810); Asteracantha Nees; Cardanthera Buchanan-Hamilton ex Bentham & J. D. Hooker;Hemiadelphis Nees; Kita A. Chevalier; Nomaphila Blume; Physichilus Nees; Polyechma Hochstetter; Santapaua N. P. Balakrishnan& K. Subramanyam; Synnema Bentham; Tenoria Dehnhardt & Giordano (1832), not Sprengel (1813).Herbs, annual or perennial, often aquatic or hygrophilous, sometimes with axillary spines, cystoliths present. Leaves sessile orshortly petiolate; leaf blade margin entire, crenulate, or sometimes undulate. Flowers sessile, in terminal spikes or several in axillaryclusters; bracteoles present or absent. Calyx 5-lobed. Corolla ventricose; limb 2-lipped, lower lip 3-lobed, upper lip apex shallowlydentate; lobes contorted in bud. Stamens 2 or 4, didynamous (posterior pair similar to anterior pair or smaller); anthers 2-thecous;thecae equally inserted; staminodes 0 or 2. Ovary with 4 to many ovules per locule; stigma entire or 2-dentate. Capsule linearellipsoidto narrowly oblong, many seeded; retinacula present. Seeds discoid, covered with long mucilaginous trichomes.About 100 species: widely distributed in tropical and subtropical regions; six species (one endemic) in China.1a. Stamens 2 .................................................................................................................................................................. 1. H. polysperma1b. Stamens 4.2a. Flowers in terminal spikes ...................................................................................................................................... 2. H. biplicata2b. Flowers solitary or clustered in leaf axils.3a. Leaf blade glabrous or scabrous-hirsute.4a. Leaf blade glabrous or slightly pubescent ................................................................................................... 3. H. ringens4b. Leaf blade scabrous-hirsute ................................................................................................................. 4. H. pogonocalyx3b. Leaf blade densely strigose or hirsute on both surfaces.5a. Calyx ca. 0.7 cm; corolla ca. 1.5 cm ............................................................................................................. 5. H. erecta5b. Calyx ca. 1.1 cm; corolla 1.8–2.2 cm .................................................................................................... 6. H. phlomoides1. Hygrophila polysperma (Roxburgh) T. Anderson, J. Linn.Soc., Bot. 9: 456. 1867.小狮子草 xiao shi zi caoJusticia polysperma Roxburgh, Fl. Ind. 1: 120. 1820;Adenosma polysperma (Roxburgh) Sprengel; Hemiadelphispolysperma (Roxburgh) Nees.Annuals [or perennials] 10–20 cm tall. Stems often prostrate,4-angled, slightly swollen above nodes, puberulent, soonglabrescent. Petiole to 5 mm; leaf blade oblong-lanceolate toovate, 2–3.5 × 0.6–1.3 cm, glabrous, base attenuate, margin entire,apex subobtuse. Inflorescences terminal spikes, 1.3–5 cm;bracts imbricate, ovate, elliptic, or obovate, 5–10 × 1.5–3.5 mm,both surfaces densely hirtellous, apex obtuse; bracteoles lanceolate,3.5–5 × 0.5–1 mm, margin ciliate. Calyx 3.5–6 mm; lobeslanceolate, 2.5–3 mm, unequal, pubescent, apex acuminate. Corollawhite, ca. 6 mm; tube ca. 3.5 mm; lower lip ca. 2 mm, 3-lobed, lobes orbicular and subequal; upper lip oblong-ovate, ca.1 mm, 2-lobed. Stamens 2, slightly exserted; filaments ca. 1mm, coherent into a membrane near base; anther thecae subequal;staminodes 2, bristlelike, short. Capsule linear-oblong,5.5–8 mm, pubescent with non-gland-tipped trichomes (alsoglabrous?), 20–30-seeded. Seeds ca. 1 × 0.5 mm. Fl. Sep–Nov,fr. Oct–Jan. 2n = 32.Streamsides; below 600 m. Guangdong, Guangxi, Taiwan, Yunnan[Bhutan, India, Myanmar, Sri Lanka, Vietnam].Hygrophila polysperma is a problematic (e.g., invasive) naturalizedweed in some tropical and temperate regions outside of its nativerange.2. Hygrophila biplicata (Nees) Sreemadhavan, Bull. Bot. Surv.India 10: 223. 1969.连丝草 lian si caoAdenosma biplicata Nees in Wallich, Pl. Asiat. Rar. 3: 79.1832; Cardanthera avana (Wallich ex Bentham) Bentham ex C.B. Clarke; Hygrophila avana (Wallich ex Bentham) E. Hossain;Pedicularis avana Wallich ex Bentham; Synnema avanum(Wallich ex Bentham) Bentham.Annuals to 30 cm tall, ascending. Stems 4-angled, hirsute.Leaves sessile; leaf blade oblong, 2.5–3.8 × 1–1.8 cm, whitepubescent, secondary veins 3–5 on each side of midvein, basetruncate to subcordate, margin crenulate, apex obtuse. Inflorescencesterminal spikes, 2.5–10 cm; bracts narrowly elliptic, 6–8× 3–4 mm, margin crenulate or dentate; bracteoles linear, ca. 8× 3 mm. Calyx 8–9 mm, inside white hirsute; lobes linear,margin ciliate. Corolla purple, 1.2–1.6 cm, outside pilose; lowerlip elliptic, ca. 8 × 5 mm, 3-dentate, lobes triangular and ca. 1 ×1 mm; upper lip oblong-ovate, ca. 6 × 3 mm, 2-dentate. Stamens4, slightly exserted; filaments with longer pair ca. 3 mmand shorter pair ca. 2 mm; anther thecae ca. 1 mm. Ovary ca. 3mm, comose; style ca. 1 cm, hirsute. Capsule ca. 8 mm, pubescent.Fl. Dec, fr. Mar.

ACANTHACEAE 431Paddy fields, wet places; 800–1000 m. Yunnan [Myanmar, Thailand].3. Hygrophila ringens (Linnaeus) R. Brown ex Sprengel, Syst.Veg. 2: 828. 1825.水蓑衣 shui suo yiPerennials to 80 cm tall, branched. Stems erect or decumbentat base, 4-angled, striate, slightly pubescent. Petiole 0.5–1cm, glabrous; leaf blade narrowly lanceolate to oblanceolate, 3–10 × 0.5–1.5 cm, both surfaces with numerous cystoliths andglabrous or slightly pubescent, secondary veins 8–11 on eachside of midvein, base attenuate and decurrent onto petiole, marginentire or slightly undulate, apex acute to obtuse. Flowers(solitary or)2–10 clustered in leaf axils, sessile; bracteoles narrowlyovate, 3–5 mm, margin pubescent, apex obtuse. Calyxnarrowly campanulate, ca. 6 cm, 5-lobed to middle; lobes linear-lanceolate,grayish pubescent or fulvous strigose, apex acuminate.Corolla pale purple, 1–2.5 cm; tube ca. 7 mm, ca. 2 mmwide, glabrous; limb 2-liped; lower lip obovate, ca. 3 mm, 3-lobed to middle, lobes ovate with an obtuse apex; upper lipelliptic, ca. 3 mm, shallowly 2-lobed, outside puberulent. Stamens4, included; filaments glabrous, longer pair ca. 5 mm,shorter pair ca. 3 mm; anther thecae 1–2 mm. Ovary glabrous;style filiform, ca. 8 mm, included; stigma unequally 2-lobed.Capsule narrowly oblong, 0.8–2.2 cm, ca. 1.5 mm wide, glabrous,12–18-seeded. Seeds ca. 1 mm, pubescent. Fl. Aug–Oct,fr. Dec–Feb.Along streams, wet places; below 1000 m. Anhui, Chongqing,Fujian, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan,Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Bhutan, Cambodia,India, Indonesia, Japan, Laos, Malaysia, Myanmar, Nepal, Pakistan,Philippines, Thailand, Vietnam].1a. Bracts and calyx grayish pubescent ........... 3a. var. ringens1b. Bracts and calyx fulvous strigose ...... 3b. var. longihirsuta3a. Hygrophila ringens var. ringens水蓑衣 ( 原变种 ) shui suo yi (yuan bian zhong)Ruellia ringens Linnaeus, Sp. Pl. 2: 635. 1753; Dipteracanthusringens (Linnaeus) Abeywickrama; Hygrophila angustifoliaR. Brown; H. lancea (Thunberg) Miquel; H. megalanthaMerrill; H. quadrivalvis (Buchanan-Hamilton) Nees; H. salicifolia(Vahl) Nees; H. salicifolia var. megalantha (Merrill) H. S.Lo & L. D. Chou; Justicia lancea Thunberg; R. quadrivalvisBuchanan-Hamilton; R. salicifolia Vahl.Calyx grayish pubescent. Corolla to 1.2 cm.Along streams, wet places; below 1000 m. Anhui, Chongqing,Fujian, Guangdong, Guangxi, Guizhou, Hainan, Henan, Hubei, Hunan,Jiangsu, Jiangxi, Sichuan, Taiwan, Yunnan, Zhejiang [Bhutan, Cambodia,India, Indonesia, Japan, Laos, Malaysia, Myanmar, Nepal, Pakistan,Philippines, Thailand, Vietnam].3b. Hygrophila ringens var. longihirsuta (H. S. Lo & D.Fang) Y. F. Deng, comb. nov.贵港水蓑衣 gui gang shui suo yiBasionym: Hygrophila salicifolia (Vahl) Nees var. longihirsutaH. S. Lo & D. Fang, Guihaia 17: 41. 1997.Flowers 2 to several in a cluster; bracts and calyx fulvousstrigose. Corolla 1.2–1.7 cm.● Along streams; 200–300 m. Guangxi.4. Hygrophila pogonocalyx Hayata, Icon. Pl. Formosan. 9: 81.1920.大安水蓑衣 da an shui suo yiAnnuals to 50 cm tall. Stems erect, 4-angled, internodessubglabrous, nodes densely hirsute. Leaves sessile; leaf bladelinear-lanceolate, 5–9 × 0.7–1.2 cm, papery, both surfacesscabrous-hirsute, secondary veins 5–7 on each side of midvein,base broadly attenuate, margins subentire, apex obtuse. Flowersseveral clustered in leaf axils, sessile; bracteoles ovate to lanceolate,ca. 1.5 × 0.4–0.6 cm, abaxially glabrous, adaxially strigose,apex acuminate. Calyx ca. 1.5 cm, 5-lobed to middle;lobes linear, subequal, apex acuminate. Corolla ca. 2.5 cm;lower lip ± spreading, 3-lobed; lobes of lower lip oblong, 2.5–3× ca. 2 mm, subequal, inside barbate, apex obtuse; upper liperect, ca. 1 cm, 2-lobed. Stamens 4; anther thecae ca. 3 mm.Ovary ca. 2.5 mm, glabrous; style glabrous. Fl. Aug–Oct, fr.Oct–Jan.● Wet places. Taiwan.5. Hygrophila erecta (N. L. Burman) Hochreutiner, Candollea5: 230. 1934.小叶水蓑衣 xiao ye shui suo yiRuellia erecta N. L. Burman, Fl. Indica, 135. 1768; Hygrophilaphlomoides Nees var. roxburghii C. B. Clarke.Perennials to 1 m tall. Stems erect, 4-angled, 2-sulcate.Petiole (0–)3–5 mm, sulcate, hispid; leaf blade elliptic, obovate,or oblong, 2–9 × 1–3 cm, with numerous cystoliths, both surfacesdensely strigose, secondary veins 8–15 on each side ofmidvein and abaxially prominent, base decurrent onto petiole,margin entire or slightly undulate, apex acute to obtuse. Flowersseveral clustered in leaf axils, sessile; bracteoles linearoblong,ca. 6 mm, densely hispid-ciliate. Calyx ca. 1.1 cm, 5-lobed to middle; lobes linear-lanceolate, densely white hispidciliate.Corolla purplish blue, 1.8–2.2 cm, pilose; tube basallycylindric for ca. 5 mm and ca. 2 mm wide then graduallywidened to ca. 6 mm at mouth; limb 2-lipped; lower lip oblong,3-lobed, lobes ovate and ca. 2 × 2 mm; upper lip triangular, 2-lobed. Stamens 4; filaments glabrous, longer pair ca. 6 mm,shorter pair ca. 3 mm; anther thecae ca. 2 mm. Ovary glabrous;style pilose. Capsule 1.2–1.5 cm. Seeds blackish brown, ca. 1.5× 1 mm, pubescent.Wet places; below 1000 m. Guangdong, Hainan, Yunnan [India,Laos, Myanmar, Thailand, Vietnam].6. Hygrophila phlomoides Nees in Wallich, Pl. Asiat. Rar. 3:80. 1832.毛水蓑衣 mao shui suo yiPerennials to 1 m tall, erect. Stems 4-angled, brown strigose.Petiole 0–3 mm, hirsute; leaf blade elliptic, obovate, oroblong, 2–9 × 1–3 cm, papery, cystoliths dense, both surfaceshirsute, secondary veins 8–15 on each side of midvein, base

432ACANTHACEAEusually attenuate and decurrent onto petiole, margin entire orundulate, apex acute to sometimes obtuse. Flowers axillary,several clustered or in whorls upward; bracteoles linear-oblong,ca. 5 × 2 mm, hirsute. Calyx ca. 1.1 cm, white hirsute, 5-lobedto middle; lobes linear. Corolla 1.8–2.2 cm, pubescent; lowerlip oblong, sparely pilose, 3-lobed; upper lip triangular, 2-lobed.Stamens 4; filaments glabrous, longer pair ca. 5 mm, shorterpair ca. 3 mm; anther thecae ca. 2 mm. Ovary glabrous; styleca. 1.8 cm, pubescent. Capsule not seen.Wet places; below 1200 m. Yunnan [Cambodia, India, Indonesia,Laos, Myanmar, Pakistan, Philippines, Thailand, Vietnam].10. ERANTHEMUM Linnaeus, Sp. Pl. 1: 9. 1753.可爱花属 ke ai hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielDaedalacanthus T. Anderson; Pigafetta Adanson; Upudalia Rafinesque.Shrubs or perennial erect herbs with cystoliths. Leaves petiolate; leaf blade margin entire or crenate. Inflorescences terminal orrarely axillary, spikes, lax, sometimes several forming a panicle; bracts sometimes colored, subleaflike, large; bracteoles small,narrow. Calyx 5-lobed; lobes narrow, subequal. Corolla subsalverform; tube basally cylindric, long, slender, throat sometimesinconspicuous; limb 5-lobed; lobes obovate, subequal, contorted in bud. Stamens 2, inserted below throat; anthers 2-thecous; thecaeparallel, muticous; staminodes 2, clavate or filiform. Ovary with 2 ovules per locule; style filiform, glabrous or pubescent; stigma 2-lobed, lobes unequal. Capsule with a solid stalk at base, clavate, 4-seeded; retinacula present. Seeds discoid, compressed, pubescentwith hygroscopic trichomes.About 15 species: tropical and subtropical Asia; two species (one endemic) in China.The Himalayan species Eranthemum splendens (T. Anderson) Bremekamp & Nannenga-Bremekamp reported from S Yunnan (FRPS 70: 60.2002) is not treated here because no material has been seen by the authors. Eranthemum pulchellum Andrews is widely cultivated in S and SW Chinaas an ornamental.1a. Petiole ca. 0.5 cm; bracts oblong to lanceolate; stamens included; capsule gland-tipped pubescent ...................... 1. E. tetragonum1b. Petiole 0.7–2.5 cm; bracts ovate to elliptic; stamens exserted; capsule glabrous ................................................. 2. E. austrosinense1. Eranthemum tetragonum A. Dietrich ex Nees in A. Candolle,Prodr. 11: 454. 1847.云南可爱花 yun nan ke ai huaHerbs to 1 m tall. Stems 4-angled, glabrous or pubescent.Petiole ca. 5 mm; leaf blade lanceolate to linear-lanceolate tooblong, 6–20 × 2–4 cm, glabrous, secondary veins ca. 5 on eachside of midvein, base attenuate and decurrent onto petiole, marginentire or crenulate, apex acuminate. Spikes 3–8 cm; peduncle2–37 mm, gland-tipped pubescent; bracts yellowish whitewith green along veins, oblong to lanceolate, 1–2.2 × 0.2–0.9cm, apex acuminate; bracteoles lanceolate, ca. 4 mm, glandtippedpubescent. Calyx ca. 6 mm, outside pilose, 5-lobed tomiddle; lobes lanceolate, ca. 1 mm wide. Corolla blue to lightpurple, ca. 4.5 cm, gland-tipped pubescent; tube basally cylindricfor ca. 3 cm; lobes obovate, ca. 6 × 5 mm. Stamens included;filaments ca. 3 mm, glabrous. Ovary gland-tipped pubescent;style hirsute. Capsule 1–1.4 cm, gland-tipped pubescent.Seeds gold to reddish brown to blackish, 3–3.5 × 2.2–2.7mm. Fl. Dec–Mar.Forests or thickets; 400–800 m. Yunnan [Cambodia, Laos, Myanmar,Thailand, Vietnam].Eranthemum tetragonum, at least under this name, was not includedby C. C. Hu (FRPS 70: 1–397. 2002).2. Eranthemum austrosinense H. S. Lo, Acta Phytotax. Sin.17(4): 85. 1979.华南可爱花 hua nan ke ai huaHerbs to 50 cm tall. Stems 4-angled, pubescent. Petiole0.7–2.5 cm, pilose; leaf blade elliptic, ovate, or elliptic-ovate,2–10 × 1–4.5 cm, abaxially pubescent along veins, adaxiallyglabrous and cystoliths obviously subulate, secondary veins 4–6on each side of midvein, base cuneate to subrounded and usuallydecurrent onto petiole, margin entire, apex acute to acuminate.Spikes 3–11 cm; peduncle 1.5–3 cm, densely villous;bracts yellowish white with green along veins, ovate to elliptic,1.2–2 × 0.4–0.8 cm, abaxially villous, adaxially glabrous, marginciliate; bracteoles triangular-ovate, 5–6 × ca. 1.5 mm, marginciliate, apex acute. Calyx 5–7 mm, outside pilose, 5-lobedto middle; lobes lanceolate, ca. 1 mm wide. Corolla bluish purple,glabrous or pubescent; tube basally cylindric for 2.5–3 cm,throat ca. 4 mm; lobes obovate, 6–8 × 5–8 mm, apex 2-lobed.Stamens exserted; filaments 3–4 mm. Ovary glabrous or pubescent;style pilose. Capsule 1–1.5 cm, glabrous or pubescent.Seeds black, ca. 4 × 3 mm.● Forests, thickets in ravines; 100–700 m. Guangdong, Guangxi,Guizhou, Yunnan.1a. Corolla glabrous; ovary pilose ......... 2a. var. austrosinense1b. Corolla pubescent; ovary glabrous .... 2b. var. pubipetalum2a. Eranthemum austrosinense var. austrosinense华南可爱花 ( 原变种 ) hua nan ke ai hua (yuan bian zhong)Corolla glabrous. Ovary pilose.● Guangdong, Guangxi, Guizhou, Yunnan.

ACANTHACEAE 4332b. Eranthemum austrosinense var. pubipetalum (S. Z.Huang ex H. P. Tsui) T. L. Li & Y. F. Deng, J. Trop. Subtrop.Bot. 15: 260. 2007.毛冠可爱花 mao guan ke ai huaEranthemum pubipetalum S. Z. Huang ex H. P. Tsui, Fl.Reipubl. Popularis Sin. 70: 347. 2002.Corolla pubescent. Ovary glabrous.● Forests, thickets in ravines; 100–700 m. Guangxi, Guizhou,Yunnan.11. ECHINACANTHUS Nees in Wallich, Pl. Asiat. Rar. 3: 75, 90. 1832.恋岩花属 lian yan hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielShrubs or perennial herbs. Leaves opposite; leaf blade margin entire. Inflorescence axillary or terminal, branched cymes; bractsnarrow to subleaflike; bracteoles inconspicuous or absent. Calyx often gland-tipped hairy, deeply 5-lobed; lobes equal or subequal.Corolla purple or yellow, funnel-shaped to campanulate; tube ventricose; limb 5-lobed; lobes subequal, contorted in bud. Stamens 4,didynamous; filaments connate at base into 2 pairs; anthers 2-thecous; thecae parallel, hirsute, spurred at base. Ovary with 4–8 ovulesper locule; stigma 2-lobed, one lobe reduced. Capsule linear-cylindric to linear-oblong, 8–16-seeded; retinacula present. Seedssubovate in outline, compressed, covered with hygroscopic trichomes.Four species: Bhutan, China, India, Nepal, Vietnam; three species (two endemic) in China.1a. Herbs to 20 cm tall ......................................................................................................................................................... 1. E. longipes1b. Subshrubs or shrubs more than 50 cm tall.2a. Corolla purple, 2–2.5 cm, outside glabrous; anther thecae each with 2 basal spurs ..................................... 2. E. longzhouensis2b. Corolla yellow, ca. 4 cm, outside pubescent; anther thecae each with 1 basal spur ............................................ 3. E. lofouensis1. Echinacanthus longipes H. S. Lo & D. Fang, Acta Bot.Yunnan. 7: 138. 1985.长柄恋岩花 chang bing lian yan huaHerbs 5–20 cm tall, perennial, caulescent or acaulescent.Petiole 1.8–4.5(–5.5) cm, densely hirsute; leaf blade oblong tolanceolate-oblong, rarely ovate or oblanceolate, 3–7(–11) ×1.5–3(–4) cm, papery, both surfaces densely covered withcystoliths, abaxially pilose or sometimes only hairy along veins,adaxially dark grayish brown when dry and subglabrous, midveinraised on both surfaces, secondary veins 5–7 on each sideof midvein and raised on both surfaces, base oblique androunded to broadly cuneate, margin entire or slightly crenulate,apex acuminate to obtuse. Cymes axillary, (1 or)2–4(or 5)-flowered; peduncle 2–10 cm, densely hirsute; bracts subleaflike,ovate to ovate-elliptic, 8–10 × 4–7 mm. Calyx lobes linear, 1–1.2 cm but to 1.7 cm in fruit, white pilose. Corolla purple, ca. 4cm, straight, outside subglabrous; tube cylindric and ca. 2.5 mmwide for 1.3–1.5 cm then gradually widened to ca. 1 cm atmouth; lobes oblong, 6–8 × 5–6 mm. Staminal filaments 5–6mm; anther thecae ovoid, ca. 3 mm, subequal, ciliate, each with1 spur at base. Ovary densely pilose; style ca. 3.2 cm, pilose;stigma 2-lobed, with one lobe longer. Capsule cylindric, ca. 1.1cm, densely pilose, 12–16-seeded. Seeds elliptic in outline, ca.1.5 mm, margin white. Fl. Apr–Jun, fr. Aug–Sep.Limestone hills in forests; 500–2000 m. Guangxi, Yunnan [Vietnam].2. Echinacanthus longzhouensis H. S. Lo, Acta Bot. Yunnan.7: 140. 1985.龙州恋岩花 long zhou lian yan huaShrubs to 3 m tall. Older stems grayish brown, subterete,lenticellate and tuberculate; young stems 4-angled, pilose. Petiole1–2(–3.5) cm, pubescent; leaf blade elliptic to ovate, 4.5–8.5 × 1.5–4.5 cm, leathery, both surfaces glabrous, secondaryveins 6 or 7 on each side of midvein and slender, base broadlycuneate, margin entire, apex acute. Cymes axillary, 1–3-flowered;peduncle 1.5–3 cm, pubescent; bracts narrowly lanceolate,10–15 × ca. 1 mm, pubescent. Calyx 1–1.2 cm, abaxially glandtippedhirsute, adaxially hirsute; lobes linear, apex acuminate.Corolla purple, 2–2.5 cm, both surfaces glabrous; tube basallycylindric and ca. 1.5 mm wide for 5–8 mm then graduallywidened to ca. 8 mm at mouth; lobes oblong, ca. 4 × 3 mm,apex retuse. Staminal filament longer pair ca. 1 cm and shorterpair ca. 0.6 cm, pilose; anther thecae with 2 spurs at base.Ovary pubescent; style ca. 1. 4 cm, pubescent. Capsule linearoblong,1.3–1.4 cm, densely pubescent, usually 8-seeded. Seedsglobose, ca. 3 × 3 mm. Fl. Aug.● Limestone hills in forests; 300–400 m. Guangxi.3. Echinacanthus lofouensis (H. Léveillé) J. R. I. Wood,Edinburgh J. Bot. 51: 186. 1994.黄花恋岩花 huang hua lian yan huaStrobilanthes lofouensis H. Léveillé, Repert. Spec. Nov.Regni Veg. 12: 99. 1913; Echinacanthus flaviflorus H. S. Lo &D. Fang.Shrubs 1–3 m tall. Stems 4-angled, lenticellate, subglabrous.Petiole 0.5–2 cm, shortly hirsute; leaf blade ovate, lanceolate,or narrowly lanceolate, (2–)6–12 × (1–)1.5–3 cm,abaxially pubescent in vein axils, adaxially glabrous, midveinusually barbate, secondary veins 5 or 6 on each side of midvein,base broadly cuneate to subrounded, margin entire, apex acuminate.Cymes axillary, usually (1–)3-flowered; peduncle 1–2 cm,pubescent; bracts subleaflike, 1–1.5 × 0.5–1 cm, caducous. Calyx1.2–1.4 cm but to 1.8 cm in fruit, both surfaces denselywhite pubescent; lobes linear, apex acuminate. Corolla yellow,

434ACANTHACEAEca. 4 cm, outside pubescent; tube basally ca. 3 mm wide for ca.1 cm then expanding to mouth; lobes semicircular, ca. 5 mm.Staminal filaments 1–1.3 cm, densely villous; anther thecaeovoid, ca. 3 mm, each with 1 spur at base. Ovary densely villous;style pilose. Capsule linear-oblong, ca. 1.5 cm, denselypubescent, 8–12-seeded. Fl. May–Oct.● Limestone hills in forests; 500–1000 m. Guangxi, Guizhou.12. PARARUELLIA Bremekamp & Nannega-Bremekamp, Verh. Kon. Ned. Akad. Wetensch.,Afd. Natuurk., Sect. 2, 45(1): 25. 1948.地皮消属 di pi xiao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, perennial. Stems short. Leaves opposite, rosulate, petiolate; leaf blade margin undulate, crenulate, or rarely subentire.Inflorescences terminal, spikes or thyrses; bracts usually subleaflike; bracteoles present. Flowers sessile. Calyx subequally to equally5-lobed. Corolla white, bluish, or pink; tube basally cylindric and slender, apically expanded into a throat; limb 5-lobed; lobessubequal, contorted in bud, apex rounded or obtuse. Stamens 4, didynamous, inserted at base of throat; filaments glabrous; anthers 2-thecous; thecae usually widely separated by a broad connective, ± parallel; staminodes absent. Ovary glabrous, with 4–8 ovules perlocule; style pubescent; stigma pubescent, 2-cleft, upper lobe very short or reduced. Capsule cylindric, not stipitate, 8–16-seeded;retinacula present. Seeds lenticular, pubescent with hygroscopic hairs.About ten species: SE Asia, China; five species (all endemic) in China.Pararuellia is purported to differ from Ruellia by its pollen, which has “minute prickles, papillae or granules (not a honey-combed pattern)” (C.B. Clarke, J. Asiat. Soc. Bengal, Pt. 2, Nat. Hist. 74: 649. 1908) or at least papillae on the muri of reticulations (C. C. Hu et al., Acta Phytotax. Sin. 43:123–150. 2005). Because such structures are also known from some species of Ruellia (C. A. Furness & M. C. Grant, Grana 35: 231–239. 1996), thegeneric status of Pararuellia is questionable and deserves additional study.1a. Bracts longer than rachis internodes, imbricate, homomorphic; spikes cylindric; flowers imbricate on rachis ........ 1. P. glomerata1b. Bracts shorter than rachis internodes, distant, usually heteromorphic; spikes not cylindric; flowers distanton rachis.2a. Spikes usually with 1 or 2(or 3) nodes ................................................................................................................ 5. P. delavayana2b. Spikes with 4 to many nodes.3a. Rachis usually zigzag, winged below nodes .......................................................................................................... 2. P. alata3b. Rachis straight, wingless.4a. Bracts oblong, elliptic, or ovate, usually subtending many flowers ........................................................ 3. P. cavaleriei4b. Bracts cordate to suborbicular, usually subtending only 1 flower ........................................................ 4. P. hainanensis1. Pararuellia glomerata Y. M. Shui & W. H. Chen, Bot. Stud.(Taipei) 50: 261. 2009.云南地皮消 yun nan di pi xiaoHerbs to 20 cm tall, often rooting at nodes. Stems 1–2 cm,with 2–4 internodes; internodes constricted, 0.5–1.1 cm. Petiole1.3–3.4 cm, densely hispid; leaf blade oblanceolate, obovateoblong,or spatulate, 4–6.2 × 1.8–3.2 cm, abaxially densely hispid,adaxially slightly hispid, secondary veins 6–9 on each sideof midvein, base narrowly cuneate and decurrent onto petiole,margin erose to crenate, apex rounded to obtuse. Spikes cylindric,(0.6–)3.2–5.4 cm, several flowered; peduncle 0.7–5 cmand elongating to 13 cm in fruit, densely hispid; rachis (0.6–)3–5.5 cm and elongating in fruit, densely hispid, with (3 or)4–10nodes; bracts imbricate, ovate to spatulate, 8–10 × 6–9 mm,densely hispid, apex acute; bracteoles linear-lanceolate, 3–4 ×ca. 1 mm, abaxially hispid especially apically, apex acuminate.Calyx ca. 5 mm; lobes linear, ca. 4 × 0.7 mm and elongating toca. 6 mm in fruit, subequal, outside hispid. Corolla light purple,bluish, or pink, 1.4–1.7 cm, outside slightly hispid, inside glabrous;lobes suborbicular, 7–8 × 5–7 mm, equal, apex roundedto truncate. Stamen longer pair 1.5–3 mm, shorter pair ca. 0.5mm; anther thecae ca. 1.1 mm. Ovary 2–2.5 mm, with 5 or 6ovules per locule; style 0.8–1.4 cm. Capsule 1–1.5 cm, pubescent,10–12-seeded. Seeds broadly ovate to subcircular inoutline, 1–1.5 × 1–1.5 mm, covered with dense long whiteappressed trichomes. Fl. Apr–Jul, fr. Jun–Aug.● Thickets on limestone hills in hot dry valleys; 200–500 m. SEYunnan (Hong He valley).2. Pararuellia alata H. P. Tsui, Novon 18: 33. 2008.节翅地皮消 jie chi di pi xiaoHerbs to 27 cm tall. Stems hispid. Petiole 2.5–3.5 cm; leafblade oblong-elliptic to oblanceolate-elliptic, 5–8.5 × 1.5–3.5cm, glabrous, secondary veins 7–10 on each side of midvein,base cuneate, margin entire, apex rounded to acute. Spikes 3–5cm, with 4–6 nodes; rachis usually zigzag, winged for ca. 2 mmbelow nodes; bracts ovate to elliptic, 1.2–6 × 0.8–3 cm, reducedin size apically on rachis, glabrous except for ciliate margin,base rounded, apex acute; bracteoles lanceolate, ca. 5 × 2.5 mm,glabrous except for ciliate margin, margin serrate. Calyx ca. 7mm, glabrous; lobes linear-triangular, ca. 6 × 1.5 mm, equal.Corolla pink, ca. 1 cm, glabrous; tube basally cylindric and ca.2 mm wide for ca. 5 mm; lobes oblong, ca. 2 × 2 mm, equal,apex emarginate. Staminal filaments ca. 1 mm; anther thecaeca. 1 mm. Ovary ca. 2 mm; style ca. 7 mm. Capsule ca. 15 × 2.7mm, glabrous. Seeds broadly ovate to subcircular in outline, ca.

ACANTHACEAE 4351.3 × 1.3 mm, covered with dense long white appressed trichomes.Fl. Aug–Oct, fr. Nov–Jan.● Streamsides in forests; 700–800 m. Chongqing, Hubei, Yunnan.3. Pararuellia cavaleriei (H. Léveillé) E. Hossain, Notes Roy.Bot. Gard. Edinburgh 32: 409. 1973.罗甸地皮消 luo dian di pi xiaoRuellia cavaleriei H. Léveillé, Repert. Spec. Nov. RegniVeg. 12: 21. 1913.Herbs to 15 cm tall. Stems ca. 1.5 cm. Petiole 0.7–2.7 cm;leaf blade oblanceolate to spatulate, 4–12 × 1–3 cm, both surfacessparsely strigose, secondary veins 4–6 on each side ofmidvein, base attenuate and decurrent onto petiole, marginundulate, apex rounded to acute. Spikes ca. 3 cm, with 4–6nodes; rachis sparsely strigose; bracts ovate, 1–4 × 0.5–1.5cm, reduced in size apically on rachis, glabrous, 4–6-veined,base broadly cuneate, apex acute; bracteoles triangular-lanceolate,ca. 3.5 × 1 mm, glabrous. Calyx ca. 5 mm, outsideapically glandular pubescent; lobes triangular-lanceolate, ca. 3× 0.7 mm, equal. Corolla purple, light blue, yellow, or white,ca. 1.5 cm, outside pubescent but soon glabrescent on limb andthroat; tube basally cylindric and narrow for ca. 6 mm, throatca. 4 mm; lobes oblong, ca. 6 × 6 mm, equal, apex emarginate.Stamen longer pair ca. 3 mm, shorter pair ca. 1.5 mm;anther thecae glabrous. Ovary ca. 3 mm; style ca. 1 cm. Capsuleca. 15 × 3 mm, glabrous, 8–12-seeded. Seeds broadlyovate in outline, ca. 1.4 × 1 mm, covered with dense appressedtrichomes. Fl. Aug–Oct, fr. Nov–Jan.● Thickets, forests; 100–1400 m. Guangxi, Guizhou, Yunnan.4. Pararuellia hainanensis C. Y. Wu & H. S. Lo, Fl. Hainan. 3:593. 1974.海南地皮消 hai nan di pi xiaoHerbs. Petiole 0.5–1.5 cm; leaf blade obovate to oblanceolate,ca. 4.5 × 2.5–4.3 cm, abaxially pubescent, adaxiallysparsely strigose, secondary veins ca. 6 on each side of midvein,base cuneate, margin entire, apex rounded, sometimes obtuse,or rarely acute. Spikes with 2–6 nodes; rachis pubescent;bracts cordate-ovate to orbicular, 0.5–3(–6.5) × 0.5–1.8(–6.5)cm, abaxially strigose, adaxially glabrous; bracteoles lanceolate,2.5–3 × ca. 0.6 mm. Calyx 5–6 mm, inside sparsely strigose;lobes linear-lanceolate, ca. 3 × 1 mm. Corolla white orblue, ca. 1.2 cm, outside pubescent, inside glabrous; tube basallycylindric and ca. 1 mm wide for ca. 7 mm, throat ca. 2.3 ×4 mm; lobes oblong, ca. 2.5 × 2 mm, subequal. Stamen longerpair ca. 1.9 mm, shorter pair ca. 1 mm; anther thecae ca. 1 mm,glabrous. Ovary ca. 3 mm; style ca. 1.6 cm, white comose. Capsuleca. 1.4 cm, glabrous. Seeds obovate in outline, ca. 1 × 1mm, covered with dense appressed trichomes. Fl. Jun–Sep, fr.Dec–Mar.● Rocks or wet places along streams, forests; 100–600 m. Guangxi,Hainan.5. Pararuellia delavayana (Baillon) E. Hossain, Notes Roy.Bot. Gard. Edinburgh 32: 409. 1973.地皮消 di pi xiaoRuellia delavayana Baillon, Hist. Pl. 10: 408. 1891;Hemigraphis drymophila Diels; Pararuellia drymophila (Diels)C. Y. Wu & H. S. Lo; R. arcuata Lingelsheim & Borza; R.drymophila (Diels) Handel-Mazzetti; R. esquirolii H. Léveillé.Herbs. Stems 1–2 cm. Petiole 0.5–2 cm; leaf blade oblongto oblong-elliptic, sometimes obovate, elliptic, or lanceolate, 4–12 × 2.5–4 cm, abaxially pubescent, adaxially strigose, secondaryveins 7–9 on each side of midvein, base cuneate and decurrentonto petiole, margin undulate to crenate, apex acute, obtuse,or rounded. Spikes ca. 4 cm, with 1–3 nodes; bracts ellipticto ovate, 1–4.7 × 0.5–2.5 cm, reduced in size apically on rachis,both surfaces sparsely strigose, base rounded, margin ciliate,apex acute to obtuse; bracteoles linear, ca. 7 × 1 mm. Calyxca. 8 mm; lobes triangular-lanceolate, ca. 7 × 1 mm, marginapically ciliate. Corolla white, light blue, or pink, ca. 1.3 cm;tube basally cylindric and narrow for ca. 4 mm, throat ca. 5mm; lobes orbicular, ca. 4 × 4 mm, subequal, apex emarginate.Stamen longer pair ca. 3.5 mm, shorter pair ca. 1.5 mm; antherthecae oblong, ca. 1 mm. Ovary ellipsoid; style ca. 1.4 cm,white hirsute. Capsule to 2 cm, 16-seeded. Seeds ovate to subcircularin outline, 1.1–1.4 × 1.1–1.4 mm, covered with denseappressed trichomes. Fl. Jul–Sep, fr. Nov–Feb.● Forests, slopes; 700–3000 m. Guizhou, S Sichuan, Yunnan.13. RUELLIA Linnaeus, Sp. Pl. 2: 634. 1753.芦莉草属 lu li cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielBlechum P. Browne; Cryphiacanthus Nees; Dipteracanthus Nees.Herbs [or shrubs], perennial, prostrate to erect, with cystoliths. Leaves sessile or petiolate; leaf blade margin entire, crenulate, ordentate. Inflorescences axillary or terminal, reduced or expanded dichasia, sometimes forming dichasiate spikes, thyrses, or panicles,sometimes reduced to a solitary flower; bracts opposite, usually green, margin entire; bracteoles 2 [or absent]. Flowers [sessile or]subsessile to pedicellate. Calyx deeply 5-lobed; lobes equal or subequal. Corolla funnel-shaped; tube basally with a narrowly cylindricportion, usually expanded apically into a distinct throat; limb 5-lobed; lobes usually ovate to orbicular, equal to unequal in size,contorted in bud. Stamens 4, didynamous, usually included in corolla tube; filaments sometimes connate in pairs at base; anthers 2-thecous; thecae equal, parallel to subsagittate, muticous at base; staminode 1 or absent. Ovary with up to 10 ovules per locule; styleusually included in corolla tube or slightly exserted; stigma 2-lobed, lobes equal or unequal. Capsule stipitate or not stipitate, [4–]12–26-seeded; retinacula present; septa with attached retinacula remaining attached (or separating in Ruellia blechum) to inner wall ofmature capsule. Seeds discoid, usually pubescent with hygroscopic trichomes.

436ACANTHACEAEAbout 250 species: tropical to temperate regions worldwide; four species (one endemic, two introduced) in China.Blechum and Dipteracanthus are included in Ruellia based on morphological and molecular data (C. Ezcurra, Ann. Missouri Bot. Gard. 80:787–845. 1993; E. Tripp, Syst. Bot. 32: 628–649. 2007; E. Tripp et al., Taxon 58: 893–906. 2009).1a. Flowers in terminal dense 4-sided spikes; septa with retinacula separating from inner wall of mature capsule ......... 1. R. blechum1b. Flowers otherwise; septa with retinacula remaining attached to inner wall of mature capsule.2a. Creeping herbs; corolla white, 1.2–2 cm .................................................................................................................... 2. R. repens2b. Erect herbs; corolla purplish, 2.2–5.5 cm.3a. Roots lacking tuberlike swellings; flowers sessile ............................................................................................ 3. R. venusta3b. Roots with tuberlike swellings; flowers pedicellate ......................................................................................... 4. R. tuberosa1. Ruellia blechum Linnaeus, Syst. Nat., ed. 10, 2: 1120. 1759.赛山蓝 sai shan lanBarleria pyramidata Lamarck; Blechum brownei Jussieu;B. pyramidatum (Lamarck) Urban.Herbs to 50 cm tall. Roots lacking tuberlike swellings.Stems slightly 4-angled, often prostrate and rooting at nodes,sparsely pubescent to subglabrous. Petiole 0.5–2.5 cm, strigose;leaf blade ovate, 3–6 × 2–4 cm, abaxially paler and subglabrous,adaxially green, sparsely strigose and densely pilosealong veins, secondary veins 5–8 on each side of midvein, baseobtuse to rounded and slightly decurrent onto petiole, marginentire, apex acute. Inflorescences terminal, dense spikes, 2–6cm, subsessile to shortly pedunculate; rachis strigose; bractssubleaflike, ovate, ca. 1.5 × 1.3 cm, abaxially slightly pubescent,margin ciliate; bracteoles 2, lanceolate-ovate, 5–9 ×1.5–3 mm, margin ciliate. Calyx lobes linear to subulate, outsidepilose. Corolla white to pale blue, ca. 1.5 cm (cleistogamousflowers sometimes present, these much smaller andbudlike), outside puberulent on lobes; lobes ovate, ca. 3 × 2mm. Stamens with longer pair ca. 4 mm and shorter pair ca.3 mm; filaments glabrous; anther thecae ca. 1 mm. Ovarycomose; style 6–10 mm, slightly hirsute; stigma usually withonly 1 lobe evident. Capsule substipitate, ovoid to ellipsoid, ca.6 mm, puberulent, 8–12-seeded; septa with attached retinaculaseparating from inner wall of mature capsule. Seeds ca. 1.5 mmin diam., margin with a conspicuous band of appressed hygroscopictrichomes. Fl. Nov, fr. Nov. 2n = 34.Disturbed sites, drainage ditches; sea level to below 100 m. Naturalizedin Taiwan [native to tropical America].Ruellia blechum is widely naturalized in tropical and subtropicalareas.2. Ruellia repens Linnaeus, Syst. Nat., ed. 12, 2: 424; Mant. Pl.1: 89. 1767.楠草 nan caoDipteracanthus lanceolatus Nees; D. repens (Linnaeus)Hasskarl.Herbs 15–50 cm tall, perennial, creeping. Roots lackingtuberlike swellings. Stems almost 4-angled, finely pubescent orglabrescent. Petiole 3–5 mm; leaf blade ovate to linear-lanceolate,1.4–4 × 0.8–2 cm, thinly papery, both surfaces sparselystrigose, secondary veins 4 or 5 on each side of midvein, basebroadly cuneate to rounded, margin subentire, apex acuminateto acute. Flowers axillary, solitary [or clustered]; bracteolessubleaflike, ovate to ovate-oblong, 8–10 mm. Pedicel to 1 mm.Calyx lobes lanceolate, ca. 5 mm, margin sparingly ciliate. Corollawhite [or pink to light purple], 1.2–2 cm, outside pilose;tube basally ca. 2 mm wide for ca. 5 mm then ventricose andgradually widened to ca. 5 mm at mouth; lobes ovate, ca. 3 × 3mm, apex obtuse. Stamens with longer pair ca. 5 mm andshorter pair ca. 2 mm; filaments glabrous; anther thecae ca. 1mm. Ovary comose; style ca. 1 cm, hirsute; stigma lobesunequal. Capsule brownish yellow, stipitate, ca. 1.2 cm, slightlypilose, 12–16-seeded; septa with attached retinacula remainingattached to inner wall of mature capsule. Seeds discoid, ca. 3mm in diam., margin with a conspicuous band of appressedhygroscopic trichomes. Fl. May–Aug, fr. Jul–Aug. 2n = 24.Grasslands, roadsides; 100–900 m. Guangdong, Guangxi, Hainan,Taiwan, Yunnan [Indonesia, Malaysia, Myanmar, Papua New Guinea,Philippines, Thailand, Vietnam].3. Ruellia venusta Hance, J. Bot. 6: 92. 1868.飞来蓝 fei lai lanLeptosiphonium venustum (Hance) E. Hossain; Ruelliaseclusa S. Moore.Herbs to 70 cm tall, unbranched or rarely branched. Petiole3–8 mm; leaf blade oblong-lanceolate, lanceolate, or oblanceolate,4–14 × 1–3.5 cm, both surfaces glabrous, secondaryveins 5–8 on each side of midvein, base cuneate to attenuateand decurrent onto petiole, margin slightly undulate to crenate,apex acute to acuminate. Flowers in axils of leaves or leaflikebracts, solitary or in dichasial clusters, sessile; bracteoles subleaflike,lanceolate to elliptic, 7–30 × 1.5–6.5 mm, abaxiallyglabrous. Calyx 7–8 mm; lobes lanceolate, outside sparsely pubescentwith non-glandular trichomes, adaxially gland-tippedpuberulent. Corolla purplish white, 4–5.5 cm; tube basallycylindric and ca. 2 mm wide for 2.2–3.5 cm; lobes 7–17 × 4–6mm, subequal, apically undulate. Stamens with longer pair ca.7 mm and shorter pair ca. 4 mm; filaments glabrous; antherthecae ca. 2 mm. Ovary glabrous; style sparsely villous. Capsuleca. 1.3 cm. Fl. Aug–Sep.● Forests, along streams; 100–800 m. Anhui, Fujian, Guangdong,Guangxi, Hubei, Hunan, Jiangxi.Ruellia venusta was treated under Leptosiphonium F. Mueller byC. C. Hu (FRPS 70: 56–58. 2002). Leptosiphonium is often treated as asection of Ruellia. The generic distinctions between Leptosiphoniumand most Ruellia are based on floral form and color (white or yellowishcorollas that are hypocrateriform in Leptosiphonium) and likely reflectdifferent pollinators. It is doubtful that the genera are distinct, but theChinese plants (with their purplish and funnelform corollas) moreclosely resemble those of Ruellia than Leptosiphonium.

ACANTHACEAE 4374. Ruellia tuberosa Linnaeus, Sp. Pl. 2: 635. 1753.芦莉草 lu li caoHerbs to 45 cm tall, perennial, erect. Roots with elongatetuberlike swellings. Stems slightly swollen above nodes, almost4-angled, strigulose on angles. Petiole to 8 mm, glabrous; leafblade oblong-obovate, 4–8 × 1.5–4.2 cm, both surfaces glabrous,veins abaxially somewhat prominent, base cuneate andtapering onto petiole, margin undulate to subentire, apex acute.Inflorescences axillary, lax dichotomous cymes; peduncle to 2.3cm; bracts oblong-lanceolate, to 3–9 × 1–3 mm; bracteoleslinear-oblong, to 1.5 mm. Pedicel 6–10 mm, minutely pilose.Calyx tube to 6 mm, outside tomentose; lobes linear-lanceolate,0.9–2 cm. Corolla pale blue, 2.2–5.5 cm, outside hirtellous;lobes suborbicular, ca. 1.6 × 1.5 cm. Stamens with longer pairca. 8 mm and shorter pair ca. 4 mm; filaments glabrous; antherthecae white, hairy on dorsal side. Ovary glabrous except for afew gland-tipped trichomes at apex; style ca. 2.1 cm, sparselyhirsute; stigma 2-lamellate. Capsule not stipitate, linear-ellipsoid,1.8–2.5 × 0.3–0.4 cm, glabrous except for a few glandtippedtrichomes at apex, 20–26-seeded; septa with attachedretinacula remaining attached to inner wall of mature capsule.Seeds discoid, ca. 2 × 2 mm, covered with appressed hygroscopictrichomes. Fl. Dec–Jan, fr. May–Jul. 2n = 32, 34.Disturbed sites such as roadsides; below 100 m. Naturalized inTaiwan and Yunnan (Hekou) [native to tropical America].Ruellia tuberosa is widely naturalized in tropical areas.Intrusaria Rafinesque.14. ASYSTASIA Blume, Bijdr. 796. 1826.十万错属 shi wan cuo shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs or subshrubs, with cystoliths. Leaves petiolate; leaf blade margin entire to crenate. Inflorescences terminal or axillary,unilateral spikes or racemes, or panicles of these; bracts small, shorter than calyx; bracteoles similar to bracts or sometimes absent.Calyx 5-lobed; lobes equal to subequal. Corolla funnel-shaped; tube expanded distally into a throat; limb ± 2-lipped, often appearingsubactinomorphic with 5 ± equal lobes; lobes ascending cochlear in bud. Stamens 4, included in or partially exserted from corollatube; anthers 2-thecous; thecae parallel to sagittate to subperpendicular, equally to unequally inserted, base muticous or with a smallbasal appendage; staminodes 0. Ovary with 2 ovules per locule; stigma capitate, 2-lobed or 2-dentate. Capsule with a solid stalk atbase, clavate, up to 4-seeded; retinacula present. Seeds lenticular, lacking trichomes.About 40 species: tropical and subtropical regions of Old World; four species (one introduced) in China.1a. Corolla ± subsalverform with cylindric basal portion of tube at least 1.5 × as long as throat and limb combined .... 1. A. neesiana1b. Corolla ± campanulate with cylindric basal portion of tube less than 1.5 × as long as throat and limb combined.2a. Leaf blade ovate to elliptic, base truncate to rounded; corolla yellow or white ................................................... 2. A. gangetica2b. Leaf blade lanceolate to ovate-lanceolate or narrowly ovate, base attenuate; corolla red or purplish red.3a. Leaf blade (1.2–)1.8–2.8 cm wide, margin entire; corolla 3.5–4 cm, lobes ca. 5 mm wide ......................... 3. A. salicifolia3b. Leaf blade 3–5 cm wide, margin undulate-crenulate; corolla ca. 2.2 cm, lobes ca. 3.2 mm wide ................ 4. A. nemorum1. Asystasia neesiana (Wallich) Nees in Wallich, Pl. Asiat. Rar.3: 89. 1832.白接骨 bai jie guRuellia neesiana Wallich, Pl. Asiat. Rar. 1: 73. 1831;Asystasia chinensis S. Moore; Asystasiella chinensis (S. Moore)E. Hossain; A. neesiana (Wallich) Lindau.Herbs to 1 m tall. Stems 4-angled, sulcate, 2-fariously puberulentto subglabrous. Petioles 0.5–5 cm, glabrous or sparselypubescent; leaf blade lanceolate to ovate to elliptic to oblong oruppermost sometimes ovate-cordate, 4–25 × 2–11 cm, bothsurfaces glabrous or slightly pubescent, secondary veins 6–8,base attenuate and decurrent onto petiole (or cordate for uppermostleaves), margin entire to sinuate-crenate to subdentate,apex acuminate. Inflorescences terminal, spikes or racemes,unbranched or with one or more basal branches forming apanicle, 6–15 cm; rachis usually gland-tipped pubescent; bractsand bracteoles triangular, 1–2 × ca. 1 mm, minutely pubescent.Pedicel (0–)1–3 mm. Calyx 4–6.5 mm; lobes linear-lanceolate,usually gland-tipped pubescent. Corolla pink to purplish blue,4–6 cm, outside gland-tipped pubescent; tube with cylindricbasal portion narrow, elongate, 3–4 cm, at least 2 × as long asthroat and limb combined; lobes of lower lip ovate, ca. 6 × 8mm; lobes of upper lip suborbicular, ca. 7 × 7 mm. Stamenswith longer pair ca. 5 mm, shorter pair ca. 2 mm; anther thecaesubequally inserted. Style 3.6–5.2 cm, glabrous. Capsule 1.8–2.8 cm, gland-tipped pubescent, 4-seeded, basal stalk 1–1.5 cm.Seeds ovate in outline, ca. 4 × 3 mm, compressed. Fl. Jul–Sep,fr. Oct–Jan.Evergreen broad-leaved forests, moist banks, streamsides, disturbedsites, ditches, trails; 100–1800 m. Anhui, Fujian, Guangdong,Guangxi, Guizhou, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan,Yunnan, Zhejiang [India, Indonesia, Laos, Malaysia, Myanmar, Thailand,Vietnam].Asystasia neesiana is sometimes treated in Asystasiella Lindau, agenus of two species with spheroidal, 4- or 5-porate, and spinulate (vs.prolate-spheroidal to subprolate, 3-colporate, and reticulate) pollen (seeK. Ensermu et al., Kew Bull. 47: 669–675. 1992). The generic positionof Asystasia neesiana s.l. and some of its relatives has yet to bethoroughly investigated.

438ACANTHACEAEThis species is used medicinally.2. Asystasia gangetica (Linnaeus) T. Anderson, Enum. Pl. Zeyl.235. 1860.宽叶十万错 kuan ye shi wan cuoHerbs to 0.5 m tall, ascending. Stems 4-angled, pilose.Petiole 3–5 mm, pubescent; leaf blade ovate to elliptic, 3–12 ×1–5 cm, glabrous or sparsely pilose especially on veins, adaxiallywith numerous cystoliths, base truncate to rounded, marginentire or slightly crenulate, apex acuminate. Racemes axillaryor terminal, to 16 cm; bracts triangular, ca. 5 mm, pilose;bracteoles linear-lanceolate, 1–2.5 mm, pilose. Pedicel 2–3 mm,pilose. Calyx ca. 7 mm; lobes linear-lanceolate, 5–7 × 1–1.2mm, outside glandular pilose, margin ciliate. Corolla yellow orwhite, 1.2–3.5 cm, outside gland-tipped pilose, inside glabrous;tube basally cylindric and ca. 3 mm wide for ca. 8 mm thengradually widened to ca. 1 cm; lobes obovate to semicircular,0.7–1.2 × 0.8–1 cm; middle lobe of lower lip with violet ormaroon markings. Stamens included; filaments glabrous, longerpair ca. 5 mm, shorter pair ca. 3 mm; anther thecae ca. 3 × 1mm. Ovary ellipsoid, ca. 3.5 mm; style ca. 1.8 cm, velutinous;stigma slightly capitate, 2-lobed. Capsule ca. 1.3 × 2 cm, pubescent.Seeds irregularly obovate in outline, 3–5 × 0.5–3 mm,tuberculate-rugose. Fl. Sep–Dec, fr. Dec–Mar. 2n = 26, 52.Forest margins, roadsides. Naturalized in Guangdong, Guangxi,Taiwan, and Yunnan [paleotropical; SW Asia].Cultivated plants of Asystasia gangetica have corollas that varyfrom white to cream to lavender to purple.1a. Corolla 3–3.5 cm, ca. 1 cm wide atmouth, lobes erect .............................. 2a. subsp. gangetica1b. Corolla 1.2–1.5 cm, ca. 0.5 cm wideat mouth, middle lobe of lower lipslightly reflexed ................................ 2b. subsp. micrantha2a. Asystasia gangetica subsp. gangetica宽叶十万错 ( 原亚种 ) kuan ye shi wan cuo (yuan ya zhong)Justicia gangetica Linnaeus, Cent. Pl. 2: 3. 1756; Asystasiacoromandeliana Nees.Corolla 3–3.5 cm, ca. 1 cm wide at mouth, lobes erect.Forest margins. Naturalized in Guangdong, Guangxi, and Yunnan[tropical Asia; Pacific islands].Asystasia gangetica subsp. gangetica occurs from India and SriLanka through SE Asia and Malesia to several Pacific islands. Exactlywhere the subspecies is native in E Asia and eastward is not confirmed.2b. Asystasia gangetica subsp. micrantha (Nees) Ensermu inSeyani & Chikuni, Proc. XIII Plenary Meeting AETFAT 1: 343.1994.小花十万错 xiao hua shi wan cuoAsystasia coromandeliana var. micrantha Nees in A. Candolle,Prodr. 11: 165. 1847; A. micrantha (Nees) Y. F. Deng &N. H. Xia.Corolla 1.2–1.5 cm, ca. 0.5 cm wide at mouth, middle lobeof lower lip slightly reflexed.Forest margins, roadsides. Naturalized in Guangdong and Taiwan[Africa, SW Asia, Indian Ocean islands, Madagascar].3. Asystasia salicifolia Craib, Bull. Misc. Inform. Kew 1918:367. 1918.囊管花 nang guan huaHerbs to 1 m tall. Stems 4-angled, pilose. Petiole ca. 5mm, sparsely hirsute; leaf blade lanceolate, (5–)12–18 × (1.2–)1.8–2.8 cm, glabrous or hirsute along veins, secondary veins 6–9 on each side of midvein, base attenuate and decurrent ontopetiole, margin entire, apex acuminate. Racemes terminal orrarely axillary, 5–10 cm; bracts triangular, 2–3 × ca. 1 mm,pilose with gland-tipped trichomes; bracteoles similar to bracts.Pedicel 1–3 mm, pilose. Calyx 5–6 mm, outside pilose withgland-tipped trichomes; lobes lanceolate. Corolla red to purplishred, 3.5–4 cm, outside gland-tipped pilose, inside pubescentin throat; lobes ovate, 5–6 × ca. 5 mm. Stamens included;filaments glabrous, longer pair ca. 1 cm, shorter pair ca. 0.8 cm;anther thecae oblong, ca. 3 mm. Ovary pilose; style ca. 2.8 cm,base pilose; stigma apically 2-cleft. Capsule and seeds not seen.Fl. Aug–Oct, fr. Nov–Jan.Forests. Yunnan [India, Laos, Myanmar, Thailand].Specimens identified as “Asystasia henryi” (C. Y. Wu, Index Fl.Yunnan. 2: 1669. 1984), which is not a validly published name, belonghere.Outside of China variation in Asystasia salicifolia also includescorollas pinkish to purplish (or rarely entirely white) and leaf blademargins entire to subsinuate.4. Asystasia nemorum Nees in Wallich, Pl. Asiat. Rar. 3: 90.1832.十万错 shi wan cuoHerbs to 1 m tall, perennial. Stems pilose. Leaf blade narrowlyovate to ovate-lanceolate, 6–12 × 3–5 cm, abaxially pilosealong veins, adaxially glabrous or pilose along veins, secondaryveins 6–9 on each side of midvein, base attenuate, marginundulate-crenulate, apex acuminate. Racemes terminal oraxillary, 5–9 cm; bracts triangular, 2–3 × ca. 1 mm, pilose withgland-tipped trichomes; bracteoles similar to bracts. Pedicel1–2 mm. Calyx 5–6 mm, outside pilose with gland-tipped trichomes;lobes lanceolate. Corolla red to purplish red, ca. 2.2cm, outside pilose with gland-tipped trichomes, inside barbatein throat; lobes ovate, 4–5 × ca. 3.5 mm, unequal. Ovary andstyle pilose; stigma slightly 2-cleft. Capsule 1.8–2.2 cm. Seedsirregularly obovate in outline, 2–4 × ca. 2.5 mm, tuberculaterugose.Fl. Aug–Oct, fr. Nov–Feb.Paddy fields, streamsides, thickets. Guangdong, Guangxi, Yunnan[India, Laos, Myanmar, Thailand, Vietnam].The plants misidentified as Asystasia chelonoides Nees by C. C.Hu (FRPS 70: 215. 2002) belong here.Asystasia salicifolia can be difficult to distinguish from A. nemorum,and Chinese plants treated as these taxa might be conspecific.Indeed, according to R. Kiew and K. Vollesen (Kew Bull. 52: 965–971.1997), A. nemorum has a compact inflorescence with flowers clusteredtoward the apex, a long and slender corolla tube, blue corollas, and isknown only from Peninsular Malaysia (where cultivated) and Java(where native). It is possible that the plants treated here as A. nemorumare either not that species or represent cultivated plants of that species(which is used medicinally). Asystasia remains much in need of revisionarystudies.

ACANTHACEAE 43915. PSEUDERANTHEMUM Radlkofer, Sitzungsber. Math.-Phys. Cl. Königl. Bayer.Akad. Wiss. München 13: 282. 1883.山壳骨属 shan ke gu shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs or shrubs, with cystoliths. Leaves sessile or petiolate; leaf blade margin entire or subentire [or lobed]. Flowers in axillaryor terminal thyrses, racemes, or spikes; bracts and bracteoles present, mostly shorter than calyx. Calyx deeply 5-lobed; lobes equal tosubequal. Corolla salverform; tube slender, not or scarcely apically enlarged; limb 5-lobed or 2-liped; lobes ± equal or 3 of lower lipslightly larger than 2 of upper lip, ascending cochlear in bud. Stamens 2, included in corolla tube or exserted from corolla mouth(some species heterostylous); anthers 2-thecous; thecae equal to subequal, parallel, inserted at same height, muticous; staminodes 2or absent. Style included or exserted; stigma 2-lobed, lobes equal. Capsule clavate with a long solid stalk, 4-seeded; retinaculapresent. Seeds lenticular, ornamented or smooth, lacking trichomes.About 50 species: pantropical; seven species (two endemic) in China.1a. Flowers in thyrses, imbricate on rachis.2a. Calyx lobes pubescent with gland-tipped trichomes .......................................................................................... 1. P. polyanthum2b. Calyx lobes pubescent with non-glandular trichomes.3a. Bracts ca. 7 mm; ovary pubescent; leaf blade 5–15 cm ............................................................................... 2. P. crenulatum3b. Bracts ca. 3 mm; ovary glabrous; leaf blade 2.5–5.5 cm .............................................................................. 3. P. shweliense1b. Flowers in racemes (thyrses in P. coudercii), distant on rachis.4a. Racemes terminal or axillary ................................................................................................................................ 4. P. teysmannii4b. Racemes terminal only (or terminal and also rarely in apical leaf axils in P. haikangense).5a. Leaf blade lanceolate, length greater than 4 × width; filaments of fertile and sterile stamens connateat base ............................................................................................................................................................... 5. P. coudercii5b. Leaf blade elliptic-ovate or elliptic, length less than 3.5 × width; filaments of fertile and sterile stamensdistinct at base.6a. Stems terete, glabrous; bracts and bracteoles 1–3 mm; corolla ca. 4 cm ............................................ 6. P. haikangense6b. Stems 4-angled, pubescent; bracts and bracteoles 4–6 mm; corolla 2–3 mm .......................................... 7. P. latifolium1. Pseuderanthemum polyanthum (C. B. Clarke ex Oliver)Merrill, Brittonia 4: 175. 1941.多花山壳骨 duo hua shan ke guEranthemum polyanthum C. B. Clarke ex Oliver, Hooker’sIcon. Pl. 20: t. 2000. 1891.Herbs. Stems grayish, subterete, glabrescent. Petiole ca.2.5 cm; leaf blade broadly ovate to oblong, 7–17 × (3–)4–9 cm,both surfaces glabrous, secondary veins 7–9 on each side ofmidvein, base cuneate and decurrent onto petiole, margin entire,apex acute. Thyrses 5–12 cm; bracts triangular, 3.5–4 × ca. 1.5mm; bracteoles ca. 2 × 0.5 mm. Pedicel 1.5–2.5 mm. Calyx ca.1 cm; lobes lanceolate. Corolla bluish purple; tube 3–3.5 cm;lower lip 3-lobed, lobes oblong and ca. 1.5 × 0.6 cm; upper lip2-lobed, lobes ca. 11 × 3 mm. Staminal filaments short, distinct,inserted at throat. Ovary pilose; style basally pilose; stigmaspherical. Capsule clavate, ca. 2.5 cm, pubescent. Seeds ovatein outline, ca. 3 × 3 mm, verrucose, glabrous. Fl. Apr–Jun, fr.Aug.Forests, thickets; 300–1600 m. Guangxi, Yunnan [India, Malaysia,Myanmar, Thailand, Vietnam].Plants misidentified as Rhinacanthus calcaratus (Wallich) Neesby C. C. Hu (FRPS 70: 268. 2002) belong here.2. Pseuderanthemum crenulatum (Wallich ex Lindley) Radlkofer,Sitzungsber. Math.-Phys. Cl. Königl. Bayer. Akad. Wiss.München 13: 286. 1883.云南山壳骨 yun nan shan ke guEranthemum crenulatum Wallich ex Lindley, Bot. Reg. 11:t. 879. 1825; E. graciliflorum Nees; E. malaccense C. B.Clarke; Pseuderanthemum graciliflorum (Nees) Ridley; P.malaccense (C. B. Clarke) Lindau.Subshrubs or shrubs, to 3 m tall. Stems blackish brown,subterete, pubescent. Petiole 1–4 cm, puberulent; leaf bladeovate-elliptic to oblong-lanceolate, 5–15 × 3–5.5 cm, abaxiallylight green and pubescent along veins, adaxially slightly pubescent,secondary veins ca. 7 on each side of midvein, base cuneateto broadly cuneate, margin entire, apex acute to acuminate.Thyrses 3–10 cm, densely flowered; bracts triangular, ca. 7 × 3mm, brown tomentose; bracteoles subulate, 3–4 × ca. 1.5 mm,brown tomentose. Pedicel 1–3 mm, pubescent. Calyx 4–5 mm;lobes linear-lanceolate, densely pubescent with non-glandulartrichomes. Corolla white or light purplish, 3.5–4.5 cm, outsidefinely and sparingly pubescent with gland-tipped trichomes;tube 2.5–3.5 cm; lower lip 3-lobed, lobes oblong and ca. 1 cm;upper lip 2-lobed to middle. Stamens slightly exserted; filamentsca. 3 mm, glabrous; anther thecae ovoid. Ovary pilose;style basally white pilose; stigma spherical. Capsule clavate, ca.2.5 cm. Seeds subcircular in outline, ca. 3.5 mm in diam., verrucose,glabrous. Fl. Apr–Jun, fr. Aug.Forests, thickets; 200–1700 m. Guangxi, Guizhou, Yunnan [India,Laos, Malaysia, Thailand, Vietnam].3. Pseuderanthemum shweliense (W. W. Smith) C. Y. Wu &C. C. Hu, Fl. Reipubl. Popularis Sin. 70: 226. 2002.瑞丽山壳骨 rui li shan ke gu

440ACANTHACEAEEranthemum shweliense W. W. Smith, Notes Roy. Bot.Gard. Edinburgh 10: 176. 1918.Shrubs 1–2 m tall. Stems woody, grayish, hispid whenyoung. Petiole 3–5 mm, densely brownish hirsute; leaf bladelanceolate, ovate-lanceolate, or elliptic, 2.5–5.5 × 1–2 cm, abaxiallysparsely pilose or glabrous but densely brown hairy alongveins, adaxially sparsely pilose along veins, base cuneate anddecurrent onto petiole, margin entire, apex obtuse to acute. Inflorescencesterminal, 7–15 cm; rachis densely yellowish pilose,with flowers fascicled at each node; bracts linear to linearlanceolate,ca. 3 mm, pilose; bracteoles subulate, ca. 2 mm.Pedicel ca. 2 mm, pilose. Calyx ca. 5 mm; lobes linear-oblong,pilose, apex acute. Corolla white with violet spot on 2 lobes;tube ca. 2.2 cm, outside and inside pilose; lobes 5, ovate,1.8–1.9 cm. Stamens exserted. Ovary glabrous; style basallysparsely pilose; stigma slightly 2-cleft. Capsule clavate, ca. 2.5cm. Seeds ovate in outline, ca. 3 mm, verrucose, glabrous. Fl.Feb–May, fr. Jun–Aug.● Thickets, forest margins; 1200–1800 m. Yunnan.4. Pseuderanthemum teysmannii (Miquel) Ridley, Fl. Malay.Penin. 2: 592. 1923 [“teysmanni”].红河山壳骨 hong he shan ke guStrobilanthes teysmannii Miquel, Fl. Ned. Ind. 2: 799.1858 [“teysmanni”].Shrubs. Stems reddish brown, 4-angled, glabrous. Petioleto 2.5 cm; leaf blade ovate, 3.5–9.5 × 1.5–4.5 cm, abaxially pubescentalong veins, adaxially sparsely pubescent along midvein,secondary veins 8–10 on each side of midvein, basebroadly cuneate and decurrent onto petiole, margin subentire,apex acuminate. Racemes terminal or axillary; rachis with flowers0.5–1.5 cm apart, yellowish brown pubescent; bracts triangular,2–3 mm, margin ciliate; bracteoles similar to bracts. Pedicel0–4 mm, pilose. Calyx ca. 5 mm, pubescent; lobes linear,slightly unequal. Corolla pale purple, ca. 3.5 cm; tube ca. 2.5cm; limb 5-lobed, slightly 2-lipped. Stamens inserted at corollathroat; anthers as long as filaments. Ovary pilose; style ca. 2cm, slightly white villous; stigma 2-cleft. Capsule clavate, ca.2.3 cm, sparsely pubescent. Seeds ovate in outline, ca. 3 × 3mm, verrucose, glabrous. Fl. Apr–Jun, fr. Aug.Thickets; 100–300 m. Yunnan [Indonesia, S Thailand].5. Pseuderanthemum coudercii Benoist, Notul. Syst. (Paris)5: 111. 1936 [“couderci”].狭叶钩粉草 xia ye gou fen caoSubshrubs or perennial herbs, 30–50 cm tall. Stems subtereteto 4-angled, pubescent, glabrescent. Petiole ca. 5 mm; leafblade oblong, lanceolate, or linear, 3–11 × 0.2–2.5 mm, bothsurfaces glabrous, secondary veins 6–12 on each side of midvein,base cuneate, margin entire or slightly undulate, apex acuminateto rarely acute. Thyrses terminal, to 20 cm; rachis pubescent;dichasia 1–4-flowered; basal bracts 1–3, large, leaflike,longer than calyx; apical bracts linear, shorter than calyx, villous;bracteoles linear, villous. Calyx 5–6 mm; lobes linear, villous.Corolla light purplish red, slightly pubescent; tube ca. 3cm; lower lip ca. 1.2 cm, lobes ovate to ovate-elliptic; upperlip 2-lobed, lobes oblong. Stamens 2; filaments connate atbase; staminodes 2, smaller than stamens. Capsule ca. 1.8 cm,sparsely pubescent. Seeds obovate in outline, ca. 3.5 mm, tuberculate-punctate,glabrous. Fl. Jun–Aug, fr. Sep.Forests, streamsides; 100–400 m. Hainan [Cambodia].6. Pseuderanthemum haikangense C. Y. Wu & H. S. Lo, Fl.Hainan. 3: 595. 1974.海康钩粉草 hai kang gou fen caoShrubs 40–100 cm tall. Stems terete, glabrous; bark strawyellow.Petiole 0.5–1.5 cm; leaf blade elliptic-orbicular, oblong,ovate, or rarely lanceolate-elliptic, 5–11.5 × 2–3.5 cm,both surfaces glabrous, secondary veins 5–7 on each side ofmidvein, base broadly cuneate to rounded, margin entire or inconspicuouslyundulate, apex shortly acuminate to caudate. Racemesterminal or rarely in apical leaf axils, to 30 cm; bractsnarrowly triangular, 2–3 mm; bracteoles 1–2 mm. Pedicel 0–3mm. Calyx ca. 5 mm; lobes linear-lanceolate, apex acuminate.Corolla white or pink and with red spots on lobes of lower lips,ca. 4 cm, pilose; tube basally ca. 1.5 mm wide for ca. 3 cm;lower lip 3-lobed, lobes narrowly elliptic, ca. 10 × 4 mm, andapex obtuse; upper lip 2-lobed, lobes elliptic, ca. 10 × 6 mm,and apex obtuse. Stamens 2; filaments ca. 2 mm; anther thecaeoblong, ca. 1.5 mm; staminodes ca. 1.3 mm. Ovary pilose; styleca. 3 cm, slightly pubescent; stigma slightly 2-cleft. Capsuleclavate, 1.9–2 cm, pubescent. Seeds broadly ovate in outline,ca. 3.5 mm, tuberculate-punctate. Fl. Jun–Aug, fr. Sep.● Forests; 200–900 m. Guangdong, Hainan, Yunnan.7. Pseuderanthemum latifolium (Vahl) B. Hansen, Nordic J.Bot. 9: 213. 1989.山壳骨 shan ke guJusticia latifolia Vahl, Symb. Bot. 2: 4. 1791; Antheliacanthusmicranthus Ridley; Eranthemum palatiferum (Wallich)Nees; J. palatifera Wallich; Pseuderanthemum palatiferum(Wallich) Radlkofer.Herbs to 1 m tall, perennial. Stems 4-angled, pubescent.Petiole 1–2.5 mm; leaf blade elliptic, 11.5–12 × 3.5–6 cm,abaxially glabrous except villous on midvein, adaxially glabrous,secondary veins 5 or 6 on each side of midvein, basecuneate, margin subentire to slightly undulate-crenulate, apexacuminate. Racemes to 30 cm; rachis pubescent, usually with 3-flowered clusters at nodes; bracts triangular, 4–6 × ca. 2 mm,pubescent; bracteoles similar to bracts. Pedicel 0.5–2 mm. Calyx5–6 mm, outside pubescent; lobes linear, 4.5–5.5 × ca. 1mm. Corolla light purple with purple dots on lower lip, 2–3 cm;tube linear, 1.5–2.5 cm; lobes 3–4 × ca. 3 mm. Style basally pubescent.Capsule ca. 2.5 cm, pubescent. Seeds ovate in outline,ca. 4 × 4 mm, rugose. Fl. Apr–Jun, fr. Aug–Sep.Forests; 100–1600 m. Guangdong, Guangxi, Hainan, Yunnan[Cambodia, India, Laos, Malaysia, Myanmar, Thailand, Vietnam].

ACANTHACEAE 44116. MACKAYA Harvey, Thes. Cap. 1: 8. 1859, nom. cons., not Arnott (1838),nor Mackaia Gray (1821).太平爵床属 tai ping jue chuang shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, perennial, with cystoliths. Leaves petiolate. Inflorescences axillary or terminal, secund racemes; bracts linear-lanceolate[or narrowly deltoid]; bracteoles 2, borne at base of pedicel. Calyx much shorter than corolla tube, 5-lobed almost to base; lobeslinear-lanceolate. Corolla subactinomorphic; tube basally cylindric, campanulate toward apex; limb 2-lipped, lower lip 3-lobed,upper lip 2-lobed; lobes suborbicular [ovate or oblong], ascending cochlear in bud. Stamens 2, inserted at base of campanulate portionof corolla tube, subincluded; anthers 2-thecous; thecae oblong, dorsally pubescent; staminodes 2, lacking anthers. Disk obsolete.Ovary subellipsoid, with ovules 2 per locule; style subglabrous or pubescent near base; stigma 2-lobed. Capsule (not seen forMackaya tapingensis) stipitate, 2- or 4-seeded; retinacula present. Seeds (not seen for M. tapingensis) somewhat rugose.Three species: disjunctly distributed in Himalayan region and South Africa; one species in China.1. Mackaya tapingensis (W. W. Smith) Y. F. Deng & C. Y. Wu,Novon 19: 308. 2009.太平爵床 tai ping jue chuangEranthemum tapingense W. W. Smith, Notes Roy. Bot.Gard. Edinburgh 10: 177. 1918; Pseuderanthemum tapingense(W. W. Smith) C. Y. Hu & H. S. Lo.Herbs to 1 m tall, perennial. Stems subterete, glabrous orsubglabrous. Petiole 2–3.5 cm, sparsely puberulent or glabrescent;leaf blade lanceolate to oblong-lanceolate, 12–17 × 3–5(–7) cm, membranous, both surfaces glabrous or glabrescent,abaxially paler, adaxially dark green, secondary veins 8–12 oneach side of midvein, base cuneate, margin entire to undulate,apex acuminate. Inflorescences axillary or terminal, racemes,7–20 cm; peduncle sparsely pilose or glabrescent; bracts linearlanceolate,2–3 mm, pilose or glabrescent; bracteoles linear, ca.1 mm. Pedicel 2–3 mm. Calyx 6–8 mm; lobes linear-lanceolate,3–4 × 1.5–2 mm, glabrous, apex acuminate. Corolla rose, 3–4cm, outside glabrous; tube basally cylindric for ca. 1 cm; lobessuborbicular, 7–12 mm, subequal, margin dentate. Staminal filaments6–12 mm, pilose; anther thecae 4–5 mm, muticous atbase; staminodes 1–4 mm. Ovary glabrous; style ca. 1.8 cm, pubescent.Capsule not seen. Fl. Oct–Jan.Forests; 600–1800 m. Yunnan [Myanmar].17. ISOGLOSSA Oersted, Vidensk. Meddel. Dansk Naturhist. Foren. Kjøbenhavn1854: 155. 1854, nom. cons.叉序草属 cha xu cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielChingiacanthus Handel-Mazzetti; Ecteinanthus T. Anderson; Leda C. B. Clarke (1908), not Bory (1822) [Algae]; RhytiglossaNees; Strophacanthus Lindau.Shrubs or herbs, with cystoliths, isophyllous or slightly anisophyllous. Leaves petiolate. Inflorescences terminal and/or sometimesaxillary, open panicles or thyrses; bracts small, shorter than calyx. Calyx deeply 5-lobed, lobes similar. Corolla funnel-shaped,2-lipped; lower lip 3-lobed; upper lip shortly 2-lobed, lacking a stylar furrow (i.e., erugulate); lobes ascending cochlear in bud.Stamens 2; anthers 2-thecous; thecae equal or unequal, parallel (to perpendicular), subequally to unequally inserted, base muticous;staminodes absent; pollen 2-pororate [elsewhere also 3-colporate and 6-pseudocolpate]. Ovary with 2 ovules per locule. Capsuleclavate with a solid sterile basal stalk, 4-seeded; retinacula present; septa with attached retinacula remaining attached to inner wall ofmature capsule. Seeds subdiscoid.About 50 species: tropical Africa and Asia; two species (one endemic) in China.1a. Calyx lobes subulate, widest at base; inflorescence long branched, widely paniculate ................................................... 1. I. collina1b. Calyx lobes spatulate-oblanceolate, widest above middle; inflorescence shortly branched, narrowly paniculate orthyrsoid ............................................................................................................................................................................... 2. I. glabra1. Isoglossa collina (T. Anderson) B. Hansen, Nordic J. Bot. 5:12. 1985.叉序草 cha xu caoJusticia collina T. Anderson, J. Linn. Soc., Bot. 9: 515.1867; Chingiacanthus patulus Handel-Mazzetti; Dianthera collina(T. Anderson) C. B. Clarke; D. sinensis W. W. Smith.Herbs 40–100 cm tall. Stems prostrate and rooting at basebut apically erect, terete, slender, slightly puberulent whenyoung, later glabrescent. Petiole 1–3 cm, brownish pubescent;leaf blade ovate to ovate-elliptic, 3.5–11 × 2–4.8 cm, abaxiallysparsely puberulent especially along veins, adaxially glabrous,secondary veins 5–7 on each side of midvein, base cuneate oron uppermost leaves rounded, margin ± entire, apex acuminate.Inflorescences terminal or axillary from upper leaf axils, panicles,5–18 × 3–10 cm; rachis gland-tipped puberulent or subgla-

442ACANTHACEAEbrous with few sessile glands; branches slightly flattened; bractslanceolate, 1.5–2 × ca. 0.5 mm, glabrous or slightly glandtippedpubescent. Calyx 5–8 mm; lobes subulate, glabrous orgland-tipped puberulent. Corolla white with purplish dots, 1.7–4.5 cm, outside glabrous; tube basally cylindric for ca. 1/3 of itslength then gradually widening into a funnel-shaped throat; lips8–10 mm; lobes of lower lip oblong-elliptic, 3–6 mm, unequalwith lateral 2 narrower. Staminal filaments 4–10 mm, glabrous;anther thecae to 4.5 mm, equal, parallel, overlapping for ca. 1/2their length or more. Ovary glabrous; style glabrous. Capsule1–1.4 cm, glabrous. Seeds coarsely corrugated with irregularridges and often with minute acute protuberances. Fl. Aug–Nov.Evergreen broad-leaved forests, wetlands by streams; 300–2200m. Guangdong, Guangxi, Hunan, Jiangxi, Xizang, Yunnan [Bhutan, India,Thailand].2. Isoglossa glabra (Handel-Mazzetti) B. Hansen, Nordic J.Bot. 5: 12. 1985.光叉序草 guang cha xu caoChingiacanthus glaber Handel-Mazzetti, Sinensia 5: 12.1934.Herbs 40–100 cm tall, erect. Stems terete, glabrous. Petiole0.8–1.5 cm, minutely puberulent; leaf blade ovate-elliptic,5.8–7.5 × 3–3.5 cm, glabrous except puberulent on margin andmidvein, secondary veins 6 or 7 on each side of midvein, baserounded to constricted cuneate, apex attenuate. Inflorescencesterminal, interrupted panicles composed of cymes, ca. 10 × 2cm; rachis glabrous; bracts triangular, ca. 1.2 × 1 mm, glabrous.Calyx ca. 4.5 mm; lobes spatulate-oblanceolate, glabrous, apexacute. Corolla to 3.5 cm, outside glabrous, inside pubescent;tube basally cylindric for ca. 1/3 of its length then graduallywidening into a funnel-shaped throat; lobes of lower lip 3–5mm. Staminal filaments 4–6 mm, glabrous; anther thecae to2.5 mm, ± equal or with lower theca smaller, ± parallel, slightlyor not overlapping. Ovary glabrous; style glabrous. Capsule notseen. Fl. Aug–Sep.● Forests; ca. 1000 m. Guangxi.18. CLINACANTHUS Nees in A. Candolle, Prodr. 11: 511. 1847.鳄嘴花属 e zui hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielShrubs or perennial herbs, with cystoliths. Leaves petiolate; leaf blade margin entire or sinuate-crenate. Inflorescences axillaryor terminal, thyrsoid racemes; bracts and bracteoles narrow. Calyx deeply 5-lobed, lobes narrow. Corolla putatively resupinate(apparently by geniculation rather than torsion); tube long, geniculate, apically widened; limb 2-lipped; lower lip slightly recurved,much broader than upper lip, 3-lobed; upper lip shortly 2-lobed; lobes ascending cochlear in bud. Stamens 2; anthers 1-thecous, basemuticous; staminodes absent. Stigma shortly 2-cleft. Capsule oblong, with a short basal stalk, 4-seeded; retinacula present.Three species: tropical Asia; one species in China.1. Clinacanthus nutans (N. L. Burman) Lindau, Bot. Jahrb.Syst. 18: 63. 1893.鳄嘴花 e zui huaJusticia nutans N. L. Burman, Fl. Indica, 10. 1768; Clinacanthusburmanni Nees, nom. illeg. superfl.; C. nutans var.robinsonii Benoist.Shrubs to 1 m tall. Stems terete, striate, glabrescent. Petiole0.3–2 cm, sulcate, bifariously pubescent; leaf blade lanceolate-ovate,lanceolate, or linear-lanceolate, 7–12 × 1–4 cm, bothsurfaces pubescent when young then glabrescent except abaxiallypilose along veins, secondary veins 4–6 on each side ofmidvein and abaxially elevated, base cuneate to rounded, marginsubentire to sinuate-crenate, apex acuminate. Racemes 1 toseveral at or near shoot apex, usually 3 to several forming aterminal panicle; rachis gland-tipped pubescent; bracts andbracteoles linear, ca. 8 × 2 mm, abaxially gland-tipped pilose,apex acute. Calyx 0.9–1.3 cm; lobes linear, outside gland-tippedpilose, apex acute. Corolla dull red with a green base, 3–4.2 cm,outside gland-tipped pilose, inside glabrous except for a ring oftrichomes ca. 5 mm above base; tube basally cylindric for ca. 5mm and ca. 3 mm wide then gradually widened to ca. 6 mm atmouth; lower lip 1–2 × ca. 1 cm, with yellow streaks; upper liptriangular, 1–2 × ca. 0.5 cm, erect, apex emarginate; lobesovate, 3–5 × 1–2 mm. Stamens exserted from mouth of corolla;filaments 1–1.5 cm, glabrous; anthers ovate-linear, 4–5 × ca. 1mm, muticous. Style 3–3.7 cm, base sparsely pubescent. Capsuleca. 2 cm, pilose. Seeds ca. 2 mm in diam. Fl. Oct–Jan, fr.Mar.Thickets; below 700 m. Guangdong, Guangxi, Hainan, Yunnan[Indonesia, Malaysia, Thailand, Vietnam].This species is used medicinally.19. KUDOACANTHUS Hosokawa, Trans. Nat. Hist. Soc. Formosa 23: 94. 1933.银脉爵床属 yin mai jue chuang shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, decumbent, often rooting at basal nodes. Leaves petiolate. Inflorescences terminal, spikes or sometimes in panicles withspicate branches, bracteate; bracteoles 2. Calyx 5-lobed. Corolla tube short, distally slightly enlarged into a throat; limb 2-lipped;lower lip 3-lobed, lobes subequal; upper lip 2-cleft, lacking a stylar furrow (so far as known); lobes ascending cochlear in bud.

ACANTHACEAE 443Stamens 2, inserted slightly above throat; anthers 2-thecous; thecae equally to subequally inserted, parallel, subequal, base muticous;staminodes absent. Disk without an annulus. Capsule unknown.● One species: China (Taiwan).Kudoacanthus was recently shown to pertain to the “Tetramerium lineage” of Justicieae (L. McDade et al., Amer. J. Bot. 95: 1136–1152. 2008).Whether K. albonervosa is generically distinct remains to be determined; it appears very similar to Leptostachya wallichii.1. Kudoacanthus albonervosa Hosokawa, Trans. Nat. Hist.Soc. Formosa 23: 95. 1933.银脉爵床 yin mai jue chuangCodonacanthus albonervosa (Hosokawa) Yuen P. Yang.Herbs to 40 cm tall. Stems hirsute. Petiole 3–15 mm; leafblade ovate to orbicular-ovate, 0.7–4.5 × 0.7–2.5 cm, membranous,both surfaces pilose, abaxially slightly paler, adaxiallygreen, secondary veins 3–5 pairs on each side of midvein,prominent, and white, base broadly cuneate, margin remotelyundulate to entire, apex obtuse to acuminate. Spikes to 4 cm,simple or branched into a panicle, few flowered; rachis glandtippedpilose; bracts linear to oblanceolate, 2–3 × ca. 1 mm,abaxially gland-tipped pilose, adaxially glandular punctate.Flowers sessile; bracteoles linear, ca. 2 × 0.8 mm, abaxiallygland-tipped pilose, adaxially glandular punctate. Calyx ca. 4mm, outside gland-tipped pilose, inside glandular punctate, 5-lobed almost to base; lobes linear, outside pubescent, marginciliate. Corolla white, ca. 5 mm, outside glabrous, insideslightly puberulent on throat; tube basally cylindric for 2.5–3mm then slightly enlarged into a throat; lobes of lower lipobovate-elliptic, spreading, apex obtuse; lobes of upper liperect, concave. Staminal filaments ca. 1 mm, glabrous; antherthecae ca. 0.8 mm. Ovary oblong-ellipsoid, ca. 1.5 mm, glabrous,with 2 ovules per locule; style filiform, ca. 4 mm, glabrous;stigma minutely 2-cleft. Capsule and seeds unknown.Fl. Apr, Aug.● Evergreen broad-leaved forests; 600–700 m. Taiwan.Kudoacanthus albonervosa is known from few collections (butcollected as recently as 2002) and remains poorly characterized. Its pollenis described as 3-porate (C. F. Hsieh & T. C. Huang, Fl. Taiwan, ed.2, 4: 669. 1998).20. RUNGIA Nees in Wallich, Pl. Asiat. Rar. 3: 77, 109. 1832.孩儿草属 hai er cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, decumbent to erect, with cystoliths. Leaves petiolate; leaf blade margin entire. Inflorescences axillary or terminal spikes,secund or 4-sided, sometimes reduced to a single flower; bracts 2- or 4-ranked but usually only 2 ranks fertile, homomorphic orheteromorphic, usually imbricate, margin usually scarious or hyaline; bracteoles similar to bracts or narrower. Calyx deeply 5-lobed;lobes equal or subequal. Corolla small; tube basally cylindric, distally ± expanded; limb 2-lipped; lower lip spreading, 3-lobed; upperlip erect, rugula (stylar furrow) present, margin entire to emarginate; lobes ascending cochlear in bud. Stamens 2, inserted in corollathroat; anthers 2-thecous; thecae subequal in size, superposed, lower one white spurred at base. Ovary with 2 ovules per locule; stigmaentire or slightly 2-cleft. Capsule shortly stalked, ovoid or oblong, 4-seeded; retinacula present; septa (with attached retinacula)separating from inner wall of mature capsule. Seeds compressed, verrucose, lacking trichomes.About 50 species: Old World tropics and subtropics; 16 species (12 endemic) in China.1a. Flowers solitary in apical leaf axils or in 2-sided spikes; bracts 2- or 4-ranked, all fertile.2a. Flowers solitary in apical leaf axils ....................................................................................................................... 1. R. axilliflora2b. Flowers in axillary or terminal spikes.3a. Plants 6–10 cm tall; spikes globose or ovoid, 0.5–1 cm ........................................................................................ 2. R. mina3b. Plants more than 20 cm tall; spikes cylindric, 1–7 cm.4a. Bracts without hyaline margin ................................................................................................................ 3. R. densiflora4b. Bracts with hyaline margin.5a. Bracteoles elliptic, ca. 1/2 as long as bracts ....................................................................................... 4. R. chinensis5b. Bracteoles ovate-lanceolate, as long as bracts ................................................................................. 5. R. stolonifera1b. Spikes 1-sided; bracts 4-ranked, each 2 fertile or sterile.6a. Bract margins not ciliate.7a. Sterile and fertile bracts different in shape; fertile bracts oblong, ca. 0.7 cm, apex rounded ................... 6. R. guangxiensis7b. Sterile and fertile bracts similar in shape; fertile bracts oblong to elliptic-oblong, ca. 1.5 cm, apexacute ............................................................................................................................................................. 7. R. pinpienensis6b. Bract margins ciliate.8a. Fertile bracts circular, obovate, elliptic, obovate-elliptic, or sphenoid, apex rounded, obtuse, oremarginate.9a. Fertile bracts circular to obovate, apex rounded to obtuse to emarginate and mucronate ....................... 8. R. pectinata9b. Fertile bracts sphenoid, elliptic, obovate, or obovate-elliptic, apex rounded and not mucronate.10a. Fertile bracts sphenoid ................................................................................................................... 9. R. monetaria

444ACANTHACEAE10b. Fertile bracts elliptic, obovate-elliptic, or obovate.11a. Fertile bracts elliptic to obovate-elliptic ................................................................................ 10. R. longipes11b. Fertile bracts obovate ........................................................................................................... 11. R. bisaccata8b. Fertile bracts lanceolate-linear, oblong-lanceolate, lanceolate, ovate, or elliptic, apex acute, acuminate,or cuspidate.12a. Bracts abaxially gland-tipped pubescent ............................................................................................... 12. R. pungens12b. Bracts glabrous or non-glandular pubescent.13a. Fertile bracts elliptic, broader at middle.14a. Leaf blade pubescent abaxially ......................................................................................... 13. R. napoensis14b. Leaf blade glabrous on both surfaces ........................................................................... 14. R. yunnanensis13b. Fertile bracts ovate to ovate-lanceolate to lanceolate-linear, broader at base.15a. Leaf blade glabrous ........................................................................................................ 15. R. taiwanensis15b. Leaf blade pubescent along veins and ciliate on margin ....................................................... 16. R. hirpex1. Rungia axilliflora H. S. Lo, Acta Phytotax. Sin. 16(4): 92.1978.腋花孩儿草 ye hua hai er caoHerbs 10–20 cm tall. Branches slender, bifariously retrorselypubescent. Petiole 2–7 mm, pubescent; leaf blade ovateto broadly ovate, 5–20 × 4–14 mm, secondary veins ca. 4 oneach side of midvein, base cuneate to rounded, margin entire orslightly undulate, apex obtuse to rounded. Flowers solitary inapical leaf axils, sessile; bracteoles 2, broadly ovate, 4–4.5 × ca.2.5 mm, margin yellow, 0.1–0.2 mm broad, membranous, andciliate. Calyx 3–3.5 mm; lobes lanceolate, glabrous, apex acuminate.Corolla light bluish, 1.1–1.2 cm, outside glabrous; tubeca. 6 mm, straight; lower lip ca. 6 mm, patent, apex 3-lobed,lobes orbicular and ca. 1.5 mm in diam.; upper lip triangular,erect, apex emarginate. Staminal filaments ca. 5 mm; antherthecae ovoid. Ovary glabrous; style 8.5–9 mm; stigma 2-lobed.Capsule not seen. Fl. Sep–Nov.● Streamsides; 400–700 m. Guangxi, Guizhou.2. Rungia mina H. S. Lo, Acta Phytotax. Sin. 16(4): 93. 1978.矮孩儿草 ai hai er caoHerbs 6–10 cm tall. Stems ca. 1 mm in diam., erect, usuallyunbranched, white retrorsely gland-tipped pubescent. Petiole2–5 mm, densely pubescent; leaf blade ovate or broadlyovate, 5–20 × 4–10 mm, both surfaces sparsely pubescent,secondary veins 2 or 3 on each side of midvein, base broadlycuneate, margin entire, apex obtuse. Spikes terminal or axillary,sessile, globose to ovoid, 5–10 mm, 2-sided, contracted; bractsall fertile, obovate to spatulate-obovate, ca. 4 × 3 mm, abaxiallypilose, margin hyaline and ciliolate, apex obtuse to sometimesacute; bracteoles boat-shaped, ca. 3 × 2 mm, concave, marginciliolate. Calyx 5-lobed almost to base; lobes linear-lanceolate,1.5–1.7 mm, apex acuminate. Corolla purple, ca. 3.8 mm; tubeca. 2 mm; lower lip ca. 1.8 mm, patent, 3-lobed, lobes suborbicular,ca. 0.5 mm in diam., and outside slightly pubescent oncentral one; upper lip broadly triangular, ca. 1.5 mm, erect, apexemarginate. Staminal filaments ca. 1.5 mm; anther thecae ovoid.Ovary glabrous; style ca. 4 mm, glabrous. Capsule ca. 2.7 mm,glabrous. Fl. Sep.● Wet places in forests; ca. 1400 m. Yunnan.3. Rungia densiflora H. S. Lo, Acta Phytotax. Sin. 16(4): 94.1978.密花孩儿草 mi hua hai er caoHerbs to 50 cm tall. Stems stout, bifariously retrorselypubescent. Petiole 0.5–2 cm, pubescent; leaf blade ellipticovate,ovate, or ovate-lanceolate, 2–8.5 × 1–3 cm, papery, bothsurfaces glabrous or minutely hirsute, secondary veins 6–8 oneach side of midvein, base cuneate and slightly decurrent ontopetiole, margin entire, apex acuminate to slightly obtuse.Spikes terminal or axillary, sessile, to 3 cm; bracts 4-ranked, allfertile, homomorphic, usually spatulate to sometimes obovate,7–11 × 1.5–3 mm, 3-veined, margin not hyaline but ciliate anddensely so apically, apex rounded to obtuse; bracteoles 2, obovate,ca. 6 × 2–3 mm, margin hyaline. Calyx ca. 4 cm; lobes linear-lanceolate.Corolla 1.1–1.7 cm, outside pilose; tube basallycylindric for 6–9 mm; lower lip oblong, 5–8 mm, apex 3-lobed,central lobe smaller; upper lip triangular-ovate, erect, shortly 2-lobed. Staminal filaments 5–7 mm, glabrous; anther thecaeovoid. Capsule ca. 6 mm. Fl. Sep–Oct, fr. Nov–Dec.● Forests, streamsides; 400–800 m. Anhui, Guangdong, Jiangxi,Zhejiang.4. Rungia chinensis Bentham, Fl. Hongk. 166. 1861.中华孩儿草 zhong hua hai er caoHerbs to 70 cm tall, spreading to erect. Stems slender, 4-angled, sulcate, slightly puberulent, usually rooting at basalnodes. Petiole 0.5–3 cm, pubescent; leaf blade ovate to oblonglanceolate,2.5–9 × 1.8–3 cm, membranous, glabrous exceptsparsely hirsute along veins, secondary veins 5 or 6 on eachside of midvein, base cuneate to attenuate, margin entire orslightly undulate, apex acute to obtuse. Spikes terminal or axillary,1–7 cm, 2-sided; bracts 4-ranked, all fertile, elliptic, 7–8 ×2–4 mm, margin sparsely ciliate, apex acute; bracteoles elliptic,4–5 × ca. 2.5 mm, margin membranous and sparsely ciliate,apex mucronate. Calyx deeply 5-lobed; lobes linear-lanceolate,ca. 5 mm. Corolla bluish, 6–8 mm, outside slightly pilose andglandular punctate; lower lip 3-lobed, lobes ca. 7 mm; upper lipshallowly dentate. Ovary subglabrous; style ca. 1 cm, glabrous;stigma obtuse. Capsule ovoid, ca. 6 mm, 4-seeded. Seeds lenticular,ca. 1.5 mm wide, minutely tuberculate. Fl. Sep–Oct, fr.Nov–Dec.Grasslands; 300–1200 m. Anhui, Fujian, Guangdong, Guangxi,Jiangxi, Taiwan, Zhejiang [Vietnam].This species is used medicinally.

ACANTHACEAE 4455. Rungia stolonifera C. B. Clarke in J. D. Hooker, Fl. Brit.India 4: 547. 1885.匍匐鼠尾黄 pu fu shu wei huangJusticia stolonifera (C. B. Clarke) B. Hansen.Herbs 30–60 cm tall, erect, branched, rooting at basalnodes. Stems 4-angled, sulcate, pubescent along groove. Petiole5–10 mm; leaf blade ovate, 2.5–7.5 × 1.8–3.8 cm, subglabrousexcept for some pubescence on margin, secondary veins ca. 5on each side of midvein, base broadly cuneate, margin entire orundulate, apex acute. Spikes terminal or axillary; bracts obovateto spatulate, ca. 4 × 3 mm, minutely pubescent, margin hyaline,apex rounded; bracteoles ovate-lanceolate, similar to bracts.Calyx ca. 1.3 cm, glabrous; lobes lanceolate, margin hyaline.Corolla 1.2–1.6 cm; tube basally cylindric, 2–3 mm in diam.,and straight for ca. 8 mm; limb ca. 4 mm; lower lip 3-lobed,pendent, lobes oblong and ca. 1 × 1 mm; upper lip triangular,erect, subentire. Staminal filaments white, glabrous; antherthecae purplish black. Ovary sparsely hirsute; style 1.5–1.8 cm,exserted; stigma orbicular. Capsule 3–4 mm, subglabrous.Seeds ovate in outline, ca. 2 × 1.5 mm, minutely gland-tippedpapillose. Fl. Sep–Oct, fr. Nov–Dec.Forests, trailsides; 200–2300 m. Yunnan [Bangladesh, India].6. Rungia guangxiensis H. S. Lo & D. Fang, Guihaia 17: 46.1997.广西孩儿草 guang xi hai er caoHerbs to 70 cm tall, erect. Stems subterete, subglabrous orpubescent when young. Petiole 0.5–2.5 cm, bifariously pubescent;leaf blade elliptic to ovate, 9–17 × 2.5–4 cm, abaxially puberulent,adaxially glabrous, secondary veins ca. 7 on each sideof midvein, base cuneate, margin entire, apex shortly acuminate.Spikes terminal, 1-sided; peduncle absent or less than 3mm; bracts 4-ranked; fertile bracts oblong, ca. 7 × 3.5 mm;sterile bracts broadly obovate, 7–8 × 4–5 mm, glabrous, apexrounded. Calyx ca. 2.5 mm, 5-lobed. Corolla red, ca. 1.7 cm,outside glabrous; tube basally cylindric and ca. 1.5 mm indiam.; lower lip 3-lobed, patent; upper lip ovate-triangular, apexobtuse. Stamens slightly exserted; anther thecae ovoid, lowerthecae spurred. Capsule not seen. Fl. Jan–Apr.● Dense forests; 500–600 m. Guangxi.7. Rungia pinpienensis H. S. Lo, Acta Phytotax. Sin. 16(4): 91.1978.屏边孩儿草 ping bian hai er caoHerbs to 1 m tall, erect. Stems 4-angled when young thensubterete, subglabrous. Petiole 2–5 cm, subglabrous; leaf bladeelliptic to broadly elliptic, 13–15 × 6–8 cm, cystoliths white,abaxially glabrous or puberulent along veins, adaxially glabrous,secondary veins 9 or 10 on each side of midvein andnetted near margin, base broadly cuneate, margin slightly undulate,apex cuspidate. Spikes terminal, ca. 9 cm, 1-sided; peduncleca. 1 cm; bracts 4-ranked, each 2 sterile or fertile, oblong toelliptic-oblong, ca. 15 × 4–5 mm, homomorphic, glabrous, marginnarrowly hyaline, apex acute. Calyx ca. 5 mm; lobes linear.Corolla not seen. Capsule ca. 1.2 cm, glabrous. Seeds blackishbrown, ovate in outline, densely verrucose. Fr. Apr–Jun.● Streamsides; 900–1900 m. Yunnan.8. Rungia pectinata (Linnaeus) Nees in A. Candolle, Prodr. 11:470. 1847.孩儿草 hai er caoJusticia pectinata Linnaeus, Amoen. Acad. 4: 299. 1760;Rungia parviflora Nees subsp. pectinata (Linnaeus) L. H.Cramer; R. parviflora var. pectinata (Linnaeus) C. B. Clarke; R.pectinata var. clarkeana Handel-Mazzetti.Herbs 20–50 cm tall, annual or perennial. Stem basallyprostrate and rooting at nodes then erect, minutely pubescent.Petiole 0–7 mm; leaf blade oblong-elliptic, 1–4 × 0.4–1.4 cm,glabrous except for a few trichomes along veins, secondaryveins ca. 4 on each side of midvein, base cuneate and decurrentonto petiole, margin entire, apex acute. Spikes axillary or terminal,0.5–2 cm, 1-sided, solitary or sometimes 2 or 3 compound;bracts dimorphic; sterile bracts green, elliptic, ca. 4 × 0.7 mm,glabrous, apex acute; fertile bracts circular to obovate, 4–5 ×ca. 2 mm, pubescent, margin broadly hyaline, apex obtuse torounded to emarginate and mucronulate; bracteoles elliptic, 2–3mm, margin ciliate, apex 2-cleft and subacute. Calyx colorless,pubescent; lobes linear-lanceolate, ca. 3 × 0.5 mm, margin narrowlyhyaline, apex mucronulate. Corolla blue or white, ca. 5mm, outside pubescent; lower lip 3-lobed, lobes triangular; upperlip ovate, 1–2 mm, apex emarginate. Staminal filaments glabrous.Ovary glabrous. Capsule ellipsoid, ca. 2.5 mm, glabrous,2–4-seeded, apex apiculate. Seeds orbicular in outline, ca. 1mm in diam., minutely verrucose. Fl. Nov–Jan, fr. Jan–Apr. 2n= 26, 52.Wastelands. Guangdong, Guangxi, Hainan, Yunnan [Bangladesh,Bhutan, India, Laos, Myanmar, Nepal, Sri Lanka, Thailand, Vietnam].Rungia pectinata is a common weed in disturbed habitats.This species is used medicinally.9. Rungia monetaria (Benoist) B. Hansen, Nordic J. Bot. 9:211. 1989.中越孩儿草 zhong yue hai er caoJusticia monetaria Benoist, Bull. Soc. Bot. France 81: 605.1934.Herbs to 1.5 tall, perennial. Stems sulcate, glabrescent.Petiole 1–2.5 cm; leaf blade elliptic to ovate-elliptic, 6–17 × 3–8.5 cm, both surfaces densely covered with prominent cystolithsand glabrous, secondary veins 4–7 on each side of midvein,base cuneate and to decurrent and asymmetric or symmetric,margin entire, apex acute to acuminate. Spikes terminalor borne from leafless lateral branches, (2–)5–8(–19) cm; peduncleca. 1 cm, glabrous; rachis glabrous; bracts 4-ranked;sterile bracts yellowish green, circular, rhombic, or obovate,1.2–1.6 × 1.1–1.6 cm, glabrous, base rounded to cuneate, marginentire or apically serrate, apex rounded to acute; fertilebracts yellowish brown, sphenoid, 1.2–1.4 × 0.8–1.1 cm, glabrous,base cuneate, margin entire, apex rounded; bracteolesyellowish brown, narrowly lanceolate, 9–11 × ca. 2 mm, mem-

446ACANTHACEAEbranous, abaxially glabrous, margin ciliate. Calyx yellowishbrown, 6–7 mm, glabrous, 5-lobed almost to base; lobes 5–6 ×ca. 1 mm. Corolla white with purple dotted stripe in throat, ca.1.5 cm; tube ca. 6 × 3 mm; lower lip oblong, ca. 7 × 6 mm, 3-lobed, middle lobe ca. 3 × 2 mm, lateral lobes ca. 2 × 1 mm;upper lip triangular, ca. 5 × 6 mm, apex emarginate. Stamens 2,inserted at base of throat; filaments ca. 3 mm, glabrous; antherthecae ovoid. Ovary ca. 1.5 × 1 mm, glabrous; style ca. 8 mm,glabrous. Capsule obovoid-ellipsoid, ca. 10 × 3 mm, glabrous,2-seeded. Seeds orbicular in outline, ca. 3 mm in diam., glabrous.Fl. Sep–Oct, fr. Nov–Jan.Streamsides; 300–900 m. Yunnan [Vietnam].10. Rungia longipes D. Fang & H. S. Lo, Guihaia 17: 49.1997.长柄孩儿草 chang bing hai er caoHerbs to 50 cm tall. Stems subterete, bifariously pubescent.Petiole (1–)4–4.5 cm, pubescent; leaf blade ovate to elliptic,10–17 × 4–8.5 cm, thinly papery to papery, abaxially pubescentalong veins, adaxially densely covered with white linearcystoliths and glabrous, secondary veins 6 or 7 on each side ofmidvein, base broadly cuneate and slightly oblique, marginundulate-denticulate, apex acuminate. Spikes axillary, 2.5–11cm, 1-sided; bracts 4-ranked, heteromorphic; sterile bracts ellipticto ovate-elliptic, (6–)10–12 × 3–4 mm, abaxially subglabrous,hyaline margin ca. 0.5 mm and ciliate, apex obtuse; fertilebracts elliptic to obovate-elliptic, ca. 12 × 5 mm, abaxiallypuberulent, margin slightly ciliate, apex rounded; bracteolesobovate-elliptic, ca. 7 × 3 mm, apex obtuse. Calyx 6–10 mm;lobes subulate, margin ciliolate. Corolla not seen. Capsule ca. 1cm, pubescent. Seeds ovate in outline, ca. 2 × 2 mm, verrucose.Fr. Apr–Jun.● Forests in valleys. Guangxi.11. Rungia bisaccata D. Fang & H. S. Lo, Guihaia 17: 48.1997.囊花孩儿草 nang hua hai er caoHerbs 20–40 cm tall. Stems subterete when young then 4-angled, 2-sulcate, pubescent along furrow. Petiole 0.8–2.5 cm,pubescent; leaf blade ovate-elliptic, 5–13 × 2.3–5.7 cm, papery,abaxially subglabrous or puberulent along veins, adaxiallyglabrous, secondary veins 5–7 on each side of midvein, basebroadly cuneate to attenuate, margin slightly undulate, apexacuminate to obtuse. Spikes axillary, 1-sided; bracts 4-ranked,sterile and fertile ones similar, obovate, 7–11 × 4–8 mm, abaxiallyglabrous, margin slightly ciliate, apex rounded; bracteolesoblong-elliptic, ca. 10 × 4 mm, margin hyaline and ciliolate.Calyx ca. 7 mm; lobes subulate, margin ciliolate. Corolla whitewith rose dots, ca. 1.8 cm, outside and inside slightly glandtippedpuberulent; tube cylindric and ca. 2.5 mm in diam. forca. 9 mm, inside with a saccate papilla; lower lip ca. 6 × 3 mm,patent, 3-lobed, central lobe larger; upper lip ca. 5 mm, erect, 2-lobed. Staminal filaments ca. 5 mm, glabrous; anther thecae ca.2 mm. Ovary glabrous; style ca. 1.3 cm, glabrous. Capsule notseen. Fl. Jan–Feb.● Rocky mountains. Guangxi.12. Rungia pungens D. Fang & H. S. Lo, Guihaia 17: 47.1997.尖苞孩儿草 jian bao hai er caoHerbs 0.5–1.5 m tall, isophyllous. Stems erect or basalportion prostrate and rooting at nodes, 4-angled, glabrous orbifariously pubescent. Petiole 0.4–5.5 cm, glabrous; leaf bladeelliptic to ovate-elliptic, 2.5–16 × 0.9–5.5 cm, densely coveredwith linear cystoliths, glabrous, secondary veins 4–9 on eachside of midvein, base broadly cuneate, margin slightly undulate,apex acuminate. Spikes terminal or axillary, 2–5 cm, 1-sided;peduncle 1–9 cm; rachis pubescent or gland-tipped pubescent;bracts 4-ranked, heteromorphic; sterile bracts elliptic, 7.5–8 ×4–6 mm, abaxially gland-tipped puberulent, hyaline margin0.8–1.5 mm broad and ciliate, apex cuspidate; fertile bractslanceolate, 6.5–7.5 × ca. 4 mm, abaxially gland-tipped puberulent,hyaline margin ca. 1 mm wide and ciliate, apex cuspidate;bracteoles elliptic, ca. 6 × 4 mm, abaxially scabrid, adaxiallyglabrous. Calyx 6–7 mm; lobes linear, unequal, scabrid. Corollalight yellowish green outside and white with purple dots inside,ca. 1.5 cm, gland-tipped pubescent and pubescent; tube ca. 7mm; lower lip 3-lobed, middle lobe larger; upper lip ca. 6 mm,apex emarginate. Staminal filaments ca. 5 mm, glabrous; antherthecae ca. 1.5 mm. Ovary hirsute; style ca. 1 cm, basal portionhirsute. Capsule ca. 8 mm, gland-tipped puberulent or puberulent.Seeds ovate in outline, ca. 2 × 1.5 mm, verrucose. Fl. Dec–Feb, fr. Apr–May.● Limestone hills, forests; 600–1000 m. Guangxi, Yunnan.13. Rungia napoensis D. Fang & H. S. Lo, Guihaia 17: 46.1997.那坡孩儿草 na po hai er caoHerbs 70–90 cm tall. Stems 4-angled when young thensubterete, sulcate, bifariously pubescent. Petiole 0.5–2.5 cm,bifariously pubescent; leaf blade narrowly elliptic to lanceolate,3–10 × 1–4 cm, papery, both surfaces densely covered withlinear cystoliths, abaxially pubescent along veins, adaxially glabrous,secondary veins 4–7 on each side of midvein, base cuneate,margin subentire, apex acuminate. Spikes axillary or terminal,1–5 cm, 1-sided; peduncle 3–5 mm, pubescent; bracts 4-ranked; sterile bracts lanceolate-elliptic, ca. 4 × 1–2 mm, abaxiallyglabrous, margin hyaline and ciliate, apex acute; fertilebracts purplish, elliptic, ca. 9 × 4 mm, covered with cystoliths,abaxially glabrous, hyaline margin ca. 0.5 mm broad and ciliate,apex acute; bracteoles purple at tip, elliptic, ca. 9 × 5 mm,abaxially puberulent. Calyx ca. 6 mm; lobes linear. Corollawhite, ca. 1.6 cm, outside sparsely pubescent; tube ca. 7 × 2mm; lower lip ca. 7 mm, 3-lobed, central lobe larger; upper liptriangular, ca. 6 mm, apex emarginate. Staminal filaments ca. 3mm, glabrous; anther thecae ca. 1.5 mm. Ovary glabrous; styleca. 1.2 cm, subglabrous. Capsule not seen. Fl. Dec–Mar.● Limestone hills, streamsides; 200–500 m. Guangxi.14. Rungia yunnanensis H. S. Lo, Acta Phytotax. Sin. 16(4):92. 1978.云南孩儿草 yun nan hai er caoHerbs. Stems stout, subterete, sulcate, bifariously re-

ACANTHACEAE 447trorsely pubescent. Petiole 1.5–2 cm, pubescent; leaf bladeovate, 6–10 × 2.5–3.5 cm, papery, both surfaces glabrous,secondary veins 10–12 on each side of midvein and netted nearmargin, base attenuate and slightly decurrent onto petiole, marginentire, apex acuminate. Spikes terminal or axillary, to 8 cm,1-sided; bracts 4-ranked; sterile bracts ovate, 10–12 × 5–6 mm,hyaline margin broad, apex acuminate; bracts fertile, elliptic,ca. 9 × 3 mm, margin hyaline; bracteoles elliptic, ca. 6.5 × 3mm. Calyx ca. 7 mm; lobes linear, slightly scabrid, apex acuminate.Corolla ca. 1 cm, outside pilose. Staminal filaments glabrous;anther thecae ovoid. Ovary hirsute; style glabrous. Capsuleca. 9 mm, tomentose. Fl. Nov–Dec, fr. Jan–Mar.● Thickets, valleys, streamsides; 400–1000 m. Yunnan.15. Rungia taiwanensis T. Yamazaki, J. Jap. Bot. 43: 61. 1968.台湾明萼草 tai wan ming e caoHerbs 30–50 cm tall. Stems prostrate and rooting at nodeson basal portion, bifariously hirsute. Petiole 1–3 cm, pubescent;leaf blade ovate-oblong to oblong-lanceolate, 2.5–8 × 2.1–3.5cm, membranous, glabrous, secondary veins 4–7 on each sideof midvein, base cuneate to attenuate, margin entire or slightlyundulate, apex acuminate. Spikes terminal or axillary, to 4 cm,often solitary; peduncle 1–2 cm; bracts 4-ranked; sterile bractsoblong-lanceolate, 8–9 × 2–3 mm, hyaline margin narrow andciliate; fertile bracts ovate to ovate-lanceolate, hyaline marginbroad and ciliate; bracteoles ovate, ca. 6 × 3.5 mm. Calyx ca. 5mm, 5-lobed almost to base; lobes lanceolate. Corolla blue, ca.1.5 cm, outside slightly gland-tipped pubescent; throat ca. 3mm; lower lip 3-lobed; upper lip broad. Stamens exserted; filamentsca. 5 mm, glabrous. Ovary glabrous; style ca. 1 cm, glabrous.Capsule ovoid, ca. 6 mm. Seeds suborbicular, ca. 1.5 ×1.5 mm, minutely tuberculate.● Thickets, streamsides; 1000–1500 m. Taiwan.16. Rungia hirpex Benoist, Bull. Mus. Natl. Hist. Nat., sér. 2,2: 149. 1930.金沙鼠尾黄 jin sha shu wei huangHerbs to 60 cm tall. Stems 4-angled, sulcate, bifariouslypubescent along sulcae. Petiole 1–3 cm, pubescent; leaf bladeovate to oblong, 2.5–9 × 1.5–4.5 cm, slightly pubescent alongveins, secondary veins 3–5 on each side of midvein and inconspicuous,base cuneate and decurrent onto petiole, margin entireand ciliate, apex acuminate. Spikes terminal, 3–5 cm; bractshomomorphic but sterile ones slightly larger than fertile ones,lanceolate-linear, 10–14 × 2–2.5 mm, abaxially pubescent, marginciliate, apex acute; bracteoles oblong-lanceolate, ca. 6 × 2mm, margin ciliate, apex acuminate. Calyx 5-lobed almost tobase; lobes linear, ca. 5 × 2 mm, glabrous, apex acute. Corollaca. 1.4 cm; tube cylindric; lower lip ca. 2 × 3 mm, 3-lobed,lobes ovate and apex rounded; upper lip 2-lobed. Stamenswhite, ca. 4 mm, glabrous. Ovary glabrous. Capsule 7–8 mm,glabrous. Seeds ovate in outline, ca. 1 mm in diam. Fl. Aug–Oct, fr. Oct–Dec.● Streamsides; 500–600 m. Yunnan.21. LEPTOSTACHYA Nees in Wallich, Pl. Asiat. Rar. 3: 76, 105. 1832, nom. cons. prop.,not Leptostachia Adanson (1763).纤穗爵床属 xian sui jue chuang shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, perennial with cystoliths. Leaves petiolate. Inflorescences a panicle of spikes, usually reduced to a single terminal spike;bracts mostly shorter than calyx; bracteoles linear, minute. Calyx deeply 5-lobed, lobes similar (or posterior lobe shorter). Corollatube cylindric, short; limb 2-lipped; lower lip 3-lobed; upper lip emarginate and galeate, lacking a stylar furrow; lobes ascendingcochlear in bud. Stamens 2; anthers 2-thecous; thecae equally to subequally inserted, parallel, muticous at base; staminodes 0. Pollen3–5-aperturate. Ovary with 2 ovules per locule. Capsule clavate with a solid sterile stalk at base, 4-seeded; retinacula present, notrising from base. Seeds lenticular.One species: tropical Asia.The genus is similar to Justicia but differs from it by the combination of its muticous anthers, galeate upper lip of the corolla, and lack of a stylarfurrow (rugula) on the upper lip. B. Hansen (Nordic J. Bot. 5: 469–473. 1985) discussed typification, taxonomy, and morphological/geographicvariation of this genus.1. Leptostachya wallichii Nees in Wallich, Pl. Asiat. Rar. 3:105. 1832.纤穗爵床 xian sui jue chuangCodonacanthus spicatus Handel-Mazzetti; Dianthera leptostachyaC. B. Clarke; Justicia wallichii (Nees) T. Anderson;Odontonemella leptostachya (C. B. Clarke) Lindau; Ptyssiglottisleptostachya (C. B. Clarke) S. Moore.Herbs 30–100 cm tall. Stems prostrate at base then ascending.Branches subterete, striate, bifariously pubescent thenglabrescent. Petiole 1.5–4 cm, puberulent; leaf blade ovate, lanceolate,or elliptic, 2–11(–17) × 1–5 cm, abaxially subglabrousexcept sparsely pubescent on margin and veins, adaxiallyglabrous or with few scattered trichomes, secondary veins 4–7on each side of midvein, base rounded to broadly cuneate, marginentire, apex acute. Inflorescences with 1–3 spikes, 4–12 cm;rachis gland-tipped pubescent; bracts oblong-lanceolate, oblanceolate,or obovate, 1.2–2.5 mm; bracteoles lanceolate, 1–3mm, apex acute. Calyx 2.5–3 mm; lobes subulate, outsidegland-tipped pubescent. Corolla white, 5–8 mm, outside pilose;tube basal portion narrowly cylindric, 4–5 × ca. 0.5 mm; lobesof lower lip triangular-ovate, subequal. Stamens inserted in co-

448ACANTHACEAErolla throat; filaments 1.5–4 mm, glabrous or puberulent; antherthecae 1–1.5 mm. Ovary glabrous or apically hirsute; style glabrousor hirsute at basal portion; stigma 2-cleft. Capsule 1–1.5cm, finely pubescent. Seeds ca. 2 × 2 mm, with concentricridges of short linear warts. Fl. Jun–Sep.Moist forests; 900–1600 m. Guangdong, Guangxi, Hainan [Bhutan,India, Indonesia, Laos, Malaysia, Myanmar, Thailand, Vietnam].22. CODONACANTHUS Nees in A. Candolle, Prodr. 11: 103. 1847.钟花草属 zhong hua cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, small, erect, perennial, with cystoliths. Leaves petiolate; leaf blade margin entire to subsinuate. Inflorescences of axillaryor terminal racemes, loose, when terminal sometimes branched and forming a panicle, pedicellate; bracts and bracteoles minute.Flowers 1(–3 or more) in only 1 axil per node. Calyx deeply 5-lobed; lobes subequal. Corolla white with colored markings on lowerlip, broadly campanulate; tube short; limb slightly 2-liped; lower lip 3-lobed; upper lip 2-lobed; lobes ascending cochlear in bud.Stamens 2, included in to slightly emergent from corolla tube; anthers 2-thecous; thecae unequal, subparallel, ± equally to subequallyinserted; staminodes 2, short. Ovary with 2 ovules per locule; stigma capitate to shallowly 2-lobed. Capsule with a solid stalk at base,fertile region ellipsoid, usually with a medial constriction, up to 4-seeded; retinacula present. Seeds ± discoid, surfaces and marginsmooth to ± rugose, lacking trichomes.Two species: E and S Asia; one species in China.1. Codonacanthus pauciflorus (Nees) Nees in A. Candolle,Prodr. 11: 103. 1847.钟花草 zhong hua caoAsystasia pauciflora Nees in Wallich, Pl. Asiat. Rar. 3: 90.1832; Codonacanthus acuminatus Nees; Leptostachya repandaQ. H. Chen.Herbs 20–60 cm tall, much branched or unbranched. Stems4-angled, sulcate, 2-fariously pubescent. Petiole 3–15 mm,puberulent or glabrous; leaf blade elliptic, ovate, or lanceolate,(2–)4–12 × (0.5–)1.5–4.5 cm, thin, abaxially pale green andglabrous or sparsely pubescent on major veins, adaxially green,glabrous or with few large-celled hairs, and with numerouscystoliths, secondary veins 5–7 on each side of midvein, baserounded, cuneate, or attenuate and decurrent onto petiole, apexacute or acuminate. Racemes simple or terminal ones usuallyforming a panicle up to 15 cm; rachis puberulent; bracts lanceolateto linear, 1–2.5 mm, puberulent; bracteoles lanceolate,1–2 mm, puberulent. Pedicel 1–7 mm, puberulent. Calyx 2.5–4mm; lobes lanceolate, puberulent. Corolla white with purplespots at base of lower lip, 6.5–10 mm, glabrous; tube basallycylindric for 1–2 mm and 1–1.5 mm wide then abruptlywidened to 3.5–6.5 mm at mouth; lobes of lower lip 3.5–5.5 ×2–3.5 mm. Staminal filaments ca. 2.5 mm, glabrous; antherthecae 1.2–1.4 mm. Ovary glabrous; style 4–6.5 mm. Capsule1–1.9 cm, glabrous. Seeds 2–3 mm. Fl. and fr. Aug–Apr.Evergreen broad-leaved forests, wet places in ravines; below 100–1500 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Jiangxi, Taiwan,Yunnan [Bhutan, Cambodia, India, Japan, Myanmar, Thailand,Vietnam].23. COSMIANTHEMUM Bremekamp, Blumea 10: 166. 1960.秋英爵床属 qiu ying jue chuang shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, often prostrate then ascending, rarely erect, with cystoliths, isophyllous. Leaves petiolate or subsessile; leaf blade marginentire. Inflorescences terminal thyrsoid racemes or thyrses, sometimes with many-flowered cymes proximally and 1–3-floweredcymes distally; bracts small; bracteoles smaller than bracts. Calyx usually deeply 5-lobed; lobes subequal or slightly unequal, posteriorone sometimes shorter. Corolla white, greenish, or yellowish; tube straight or curved, sometimes with a dorsal pouch; throat shortor absent; limb 2-lipped; lower lip 3-lobed; upper lip emarginate or 2-lobed [or entire]; lobes ascending cochlear in bud. Stamens 2,inserted at throat; anthers 2-thecous; thecae equal or subequal, parallel, muticous at base; staminodes 2, inserted at base of upper lip.Stigma 2-cleft. Capsule stalked, 4-seeded; retinacula present.About ten species: SE Asia, especially Borneo; three species (two endemic) in China.Cosmianthemum resembles Pseuderanthemum in fruit and androecium but differs by the generally smaller corolla being distinctly 2-lipped (theupper lip only shallowly lobed) and lacking an elongate, narrow, cylindric tube; see B. Hansen (Nordic J. Bot. 5: 195–197. 1985) and B. L. Burtt andR. M. Smith (Notes Roy. Bot. Gard. Edinburgh 26: 365–381. 1965).1a. Leaf blade ovate to broadly ovate, 4–9 cm wide .................................................................................................. 1. C. viriduliflorum1b. Leaf blade elliptic-lanceolate, ovate-elliptic, or oblong-lanceolate, 1.4–7.3 cm wide.2a. Bracts abaxially gland-tipped pubescent; calyx outside gland-tipped pubescent ............................................ 2. C. knoxiifolium2b. Bracts abaxially non-gland-tipped pubescent; calyx outside non-glandular pubescent .................................. 3. C. guangxiense

ACANTHACEAE 4491. Cosmianthemum viriduliflorum (C. Y. Wu & H. S. Lo) H.S. Lo, Guihaia 17: 42. 1997.海南秋英爵床 hai nan qiu ying jue chuangGraptophyllum viriduliflorum C. Y. Wu & H. S. Lo, Fl.Hainan. 3: 594. 1974.Herbs to 60 cm tall, erect, perennial. Stems terete, glabrous;branches yellowish brown, pubescent. Petiole 1–2.5 cm,pubescent; leaf blade ovate to broadly ovate, 7–15 × 4–9 cm,both surfaces glabrous, abaxially light green, adaxially darkgreen and lustrous, secondary veins 5–7 on each side of midveinand netted near margin, base broadly cuneate to subrounded,margin entire or inconspicuously undulate, apex acuminate,caudate, or sometimes acute. Thyrses terminal, 4–10cm; peduncle 1–4 cm; rachis pubescent; bracts linear, 1–1.5 ×ca. 0.5 mm; bracteoles linear, ca. 1 × 0.5 mm. Pedicel 2–2.5mm. Calyx ca. 5 mm, gland-tipped pubescent; lobes linear-lanceolate,apex acuminate. Corolla light greenish yellow, ca. 1.2cm, outside and inside pubescent; lobes of lower lip oblonglanceolate,ca. 3 × 0.6 mm, subacute at apex; upper lip erect,apex 2-cleft to emarginate. Staminal filaments ca. 4 mm, pubescent;anther thecae ca. 2 mm; staminodes ca. 2 mm, sparselypubescent. Ovary glabrous; style ca. 6 mm, sparsely pubescent.Capsule ca. 2 cm. Seeds not seen. Fl. Jul–Oct.● Forests; 700–1000 m. Hainan.2. Cosmianthemum knoxiifolium (C. B. Clarke) B. Hansen,Nordic J. Bot. 5: 195. 1985.秋英爵床 qiu ying jue chuangGymnostachyum knoxiifolium C. B. Clarke, J. Asiat. Soc.Bengal, Pt. 2, Nat. Hist. 74: 663. 1908; Pseuderanthemumparviflorum Ridley; Ptyssiglottis parviflora (Ridley) Ridley;Sphinctacanthus malayanus Ridley.Shrubs to 50 cm tall or more. Stems terete, puberulent.Petiole 1–1.5 cm, glabrous; leaf blade elliptic-lanceolate, 8–20× 1.8–7.3 cm, leathery, abaxially puberulent along veins andmargin, adaxially glabrous, secondary veins 8–10 on each sideof midvein, base cuneate, margin remotely serrate or subentire,apex acuminate to attenuate. Thyrses terminal, (5.5–)10–30(–60) cm; rachis brownish pubescent with few gland-tippedtrichomes; bracts lanceolate, 2–3 × 0.5–1 mm, gland-tipped pubescent;bracteoles subulate, 1–2 × ca. 0.5 mm. Pedicels 1–3mm, pubescent. Calyx 2.5–5 mm, outside gland-tipped pubescent;lobes lanceolate, outside scaly hairy. Corolla white to paleyellow with purple spots on lower lip, ca. 1.5 cm, outside pubescent;tube ca. 5 mm; upper lip erect, apically 2-lobed toemarginate. Staminal filaments 6–7 mm, glabrous; staminodesca. 1 mm, pubescent. Ovary gland-tipped pubescent; style pubescent.Capsule 2–2.7 cm, puberulent. Seeds ovate in outline,ca. 2 × 2 mm, flattened, irregularly ridged or warty, lackingtrichomes. Fl. May–Sep, fr. Sep–Dec.Dense montane forests; 400–500 m. Hainan [Malaysia, Thailand,Vietnam].3. Cosmianthemum guangxiense H. S. Lo & D. Fang, Guihaia17: 42. 1997.广西秋英爵床 guang xi qiu ying jue chuangCosmianthemum longiflorum D. Fang & H. S. Lo.Herbs to 70 cm tall, branched or unbranched. Stems 4-angled to subterete, sulcate, sparsely pubescent. Petiole 0.5–1cm, grooved, pilose; leaf blade ovate-elliptic to oblong-lanceolate,2–13.5 × 1.4–4 cm, abaxially glabrous, adaxially pilosetoward base or along veins, secondary veins 5–7 on each side ofmidvein, base rounded to cuneate, margin entire, apex acuminate.Thyrses terminal, 3–10 cm; rachis pilose; peduncle(0.5–)1–3 cm, pubescent; cymes 1–6-flowered; bracts triangular-subulate,ca. 1.5 × 0.5 mm, pilose; bracteoles similar tobracts. Pedicel 1–3 mm. Calyx 3–4 mm, outside pilose; lobeslinear, apex acute. Corolla white or yellowish green, 1–1.5 cm,outside sparsely pubescent to subglabrous, inside glabrous; tubeca. 1.5 mm wide for 4–5 mm; lower lip 6–7 × 3.5–5 mm, lobesovate and ca. 1 × 1 mm; upper lip ovate, 5–6 × ca. 5 mm.Staminal filaments 4–5 mm, glabrous; thecae ca. 2 mm. Ovaryglabrous; style 0.9–1.1 cm, sparsely hirsute to subglabrous.Capsule and seeds not seen. Fl. Oct.● Forests; ca. 400 m. Guangxi.24. JUSTICIA Linnaeus, Sp. Pl. 1: 15. 1753.爵床属 jue chuang shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielAdatoda Adanson; Adhatoda Miller; Calophanoides Ridley; Ecbolium Kuntze (1891), not Kurz (1871); Gendarussa Nees;Mananthes Bremekamp; Rhaphidospora Nees; Rostellaria Nees (1832), not C. F. Gaertner (1807); Rostellularia Reichenbach.Herbs, subshrubs, or shrubs, cystoliths present. Leaves sessile or petiolate; leaf blade margin usually entire but sometimessinuate or slightly serrate. Inflorescences dichasia (sometimes reduced to a single flower) in leaf axils or axillary or terminal spikes,racemes, or thyrses, sometimes branched and becoming panicles; bracts variable in shape, sometimes prominent and/or brightlycolored; bracteoles 2, similar to or smaller than bracts. Calyx deeply 4- or 5-lobed; lobes equal or subequal. Corolla tubular orfunnel-shaped; limb strongly 2-lipped; lower lip 3-lobed; upper lip entire to 2-lobed, internally rugulate (i.e., with a stylar furrow);lobes ascending cochlear in bud. Stamens 2; anthers 2-thecous; thecae equally or unequally inserted, parallel to perpendicular,one or both spurred at base or sometimes lacking basal appendages; staminodes absent. Ovary with 2 ovules per locule;stigma slightly 2-lobed. Capsule with a sterile basal stalk and a fertile head (i.e., ± clavate), 2–4-seeded; retinacula present. Seedscompressed [to spheric].About 700 species: tropical and temperate regions worldwide; 43 species (26 endemic, three introduced) in China.

450ACANTHACEAEJusticia betonica Linnaeus, J. brandegeeana Wasshausen & L. B. Smith, and J. carnea Lindley are only cultivated in China so are not treatedhere.1a. Calyx equally 4-lobed or unequally 5-lobed (i.e., with one lobe conspicuously smaller than others).2a. Leaves sessile.3a. Plants diffuse, to 8 cm tall; leaf blade lanceolate to narrowly lanceolate, glabrous ................................................ 1. J. demissa3b. Plants usually erect, to 100 cm tall; leaf blade oblong to broadly ovate, pubescent ........................................... 2. J. mollissima2b. Leaves petiolate.4a. Leaf blade linear, to 5 mm wide ..................................................................................................................... 3. J. neolinearifolia4b. Leaf blade ovate to elliptic to subcircular, usually wider than 5 mm.5a. Bracts obovate-elliptic, broadest at or above middle, 2–3 mm wide .................................................................... 6. J. hayatae5b. Bracts ovate to elliptic-lanceolate to oblong-lanceolate, mostly broadest near base, 0.6–1.3 mm wide.6a. Spikes dense and cylindric, ± densely pubescent ........................................................................................ 4. J. procumbens6b. Spikes neither dense nor cylindric, sparsely pubescent ........................................................................................ 5. J. diffusa1b. Calyx equally 5-lobed.7a. Flowers in axillary abbreviated spikes, spikes ca. 1 cm, usually solitary or several clustered.8a. Leaf blade glabrous.9a. Leaf blade linear to linear-lanceolate ................................................................................................................... 7. J. neesiana9b. Leaf blade ovate to broadly elliptic (rarely lanceolate).10a. Bracts subcircular .................................................................................................................................... 8. J. kwangsiensis10b. Bracts lanceolate .......................................................................................................................................... 9. J. alboviridis8b. Leaf blade ± pubescent (sometimes glabrous or only pubescent along veins in J. quadrifaria).11a. Corolla ca. 2 cm ................................................................................................................................................... 10. J. wardii11b. Corolla less than 1.5 cm.12a. Bracts linear, lanceolate, or ovate, broadest at base, 1-veined.13a. Bracts 2–3 mm .......................................................................................................................................... 20. J. siccanea13b. Bracts 5–6 mm ..................................................................................................................................... 21. J. xantholeuca12b. Bracts ovate to circular to elliptic to obovate to spatulate-obovate, broadest above base, pinnatelyveined.14a. Leaf blade (2–)5.5–11.5 cm.15a. Bracts ovate to obovate, with 3–4 mm petiole, apex obtuse; calyx 5–6 mm ................................... 17. J. quadrifaria15b. Bracts obovate to circular, with 1–2 mm petiole, apex retuse or rounded; calyx ca. 1 cm .......... 18. J. kouytcheensis14b. Leaf blade 0.8–7(–10.5) cm.16a. Leaf blade adaxially glabrous ................................................................................................................ 13. J. xylopoda16b. Leaf blade adaxially pubescent at least when young.17a. Leaf blade sparsely pubescent, soon glabrescent or abaxially pilose along midvein ................... 11. J. championii17b. Leaf blade hispid, or densely grayish or white or yellowish pubescent.18a. Bracts circular to subcircular ..................................................................................................... 16. J. hainanensis18b. Bracts not circular (sometimes subcircular in J. xerobatica and J. xerophila).19a. Corolla white.20a. Stems yellow pilose; leaf blade 3–5 mm wide ........................................................................ 12. J. xerophila20b. Stems minutely white sericeous; leaf blade 5–8 mm wide ................................................... 19. J. xerobatica19b. Corolla whitish yellow, yellow, or yellowish green.21a. Stems villous .......................................................................................................................... 14. J. albovelata21b. Stems sericeous ................................................................................................................... 15. J. yunnanensis7b. Flowers in terminal or axillary elongated spikes (or racemes or panicles), or in axillary cymes, or inthyrses of 3-flowered cymes.22a. Flowers in terminal thyrses of 3-flowered cymes; both anther thecae spurred at base ......................................... 22. J. grossa22b. Flowers in spikes or axillary cymes; only lower anther theca spurred at base.23a. Corolla 2.5–3 cm ............................................................................................................................................. 23. J. adhatoda23b. Corolla 2.2 cm or less.24a. Flowers in axillary cymes ......................................................................................................................... 24. J. vagabunda24b. Flowers in terminal or axillary spikes (or panicles of spikes), or sometimes in racemes.25a. Bracts (at least lower ones) longer than calyx.26a. Bracts 1–1.5 cm ................................................................................................................................... 25. J. ventricosa26b. Bracts to 1 cm.27a. Leaf blade narrowly lanceolate, to 1.5 cm wide; basal bracts longer than calyx then graduallysmaller with apicalmost ones shorter than calyx ........................................................................... 26. J. gendarussa27b. Leaf blade lanceolate to ovate to elliptic to subcircular, often wider than 1.5 cm; all bractslonger than calyx.

ACANTHACEAE 45128a. Spikes interrupted; flowers distant on rachis; corolla yellow with purplish spots ................ 27. J. pseudospicata28b. Spikes continuous; flowers dense on rachis; corolla pale white or yellowish greenwith purplish red dots or stripes.29a. Leaf blade margin entire; bracts ovate to elliptic, basally pale green and purplish redabove middle, apex acuminate ..................................................................................................... 28. J. latiflora29b. Leaf blade margin entire or crenate; bracts broadly obovate or flabellate, green, apexacuminate to obtuse and mucronate with 1–3 teeth ............................................................ 29. J. austrosinensis25b. Bracts equaling or shorter than calyx.30a. Leaves sessile.31a. Leaf blade oblanceolate to rarely lanceolate, 8–17 × 2.5–6.5 cm, abaxially rust-coloredwhen dry .......................................................................................................................................... 30. J. ferruginea31b. Leaf blade obovate, 15–26 × 7.5–15 cm, abaxially dark green when dry ............................... 31. J. panduriformis30b. Leaves petiolate.32a. Leaf blade base cordate to truncate.33a. Flowers usually 1 per rachis node; stems 4-angled; calyx ca. 4 mm; calyx and corollapubescent with gland-tipped trichomes outside ......................................................................... 32. J. lianshanica33b. Flowers usually 2–5 per rachis node; stems terete; calyx ca. 2 mm; calyx and corollapubescent with non-glandular trichomes outside ..................................................................... 33. J. cardiophylla32b. Leaf blade base (rounded to) cuneate to attenuate.34a. Stems short; leaves in a basal rosette ............................................................................. 34. J. austroguangxiensis34b. Stems elongated; leaves cauline.35a. Flowers more than 1 per rachis node.36a. Shrubs to 2 m tall; leaf blade (2–)3–5.5 × 1.5–2.5 cm; inflorescences unbranchedspikes .................................................................................................................................... 36. J. microdonta36b. Herbs to 1 m tall; leaf blade 4–12.5(–18) × 2.5–6.5(–8) cm; inflorescencesunbranched or branched.37a. Leaf blade 10.5–12.5(–18) × 5–6.5(–8) cm; flowers in spikes ...................................... 35. J. leptostachya37b. Leaf blade 4–9 × 2.5–4.5 cm; flowers in spiciform racemes (pedicel ca. 1 mm) .......... 37. J. damingensis35b. Flowers 1 per rachis node.38a. Calyx lobes elliptic-lanceolate, apex obtuse or rounded ................................................. 38. J. amblyosepala38b. Calyx lobes linear to lanceolate or narrowly triangular, apex acute.39a. Stems sharply 4-angled, slightly winged along angles ..................................................... 39. J. acutangula39b. Stems subterete to 4-angled, wingless.40a. Corolla tube 2–3 × as long as limb ............................................................................... 40. J. patentiflora40b. Corolla tube less than 2 × as long as limb.41a. Calyx puberulent on both surfaces .................................................................................. 41. J. poilanei41b. Calyx glabrous adaxially.42a. Leaf blade lanceolate, broadest below middle, 4.3–5 cm wide; bractsca. 1.5 mm .............................................................................................................. 42. J. kampotiana42b. Leaf blade elliptic to obovate-elliptic, broadest at or above middle,1.2–4.2 cm wide; bracts 3–4 mm ........................................................................... 43. J. caudatifolia1. Justicia demissa N. H. Xia & Y. F. Deng, J. Trop. Subtrop.Bot. 13: 534. 2005.矮爵床 ai jue chuangRostellularia humilis H. S. Lo, Fl. Hainan. 3: 598. 1974,not Justicia humilis Michaux (1803).Herbs 6–8 cm tall, diffuse, annual. Stems 4-angled, glabrous.Leaves sessile; leaf blade lanceolate to narrowly lanceolate,1–2 × ca. 0.3 cm, glabrous, cystoliths inconspicuous, basecuneate, margin entire, apex obtuse. Spikes terminal, 2–4.5 cm,glabrous; bracts linear-lanceolate, ca. 3 mm, glabrous, marginmembranous; bracteoles similar to bracts. Calyx ca. 5 mm, glabrous,4-lobed to base; lobes linear, margin membranous, apexobtuse. Corolla not seen. Capsule ca. 6 mm, glabrous.● Hainan.2. Justicia mollissima (Nees) Y. F. Deng & T. F. Daniel, comb.nov.喀西爵床 ka xi jue chuangBasionym: Rostellaria mollissima Nees in Wallich, Pl.Asiat. Rar. 3: 101. 1832; Justicia khasiana C. B. Clarke; J.procumbens Linnaeus var. latispica C. B. Clarke; Rostellulariakhasiana (C. B. Clarke) J. L. Ellis; R. khasiana var. latispica(C. B. Clarke) C. Y. Wu ex C. C. Hu; R. mollissima (Nees)Nees.Herbs 60–100 cm tall. Stems 4-angled, erect, hispid.Leaves sessile; leaf blade oblong to broadly ovate, 3–4.5 × 0.3–0.5 cm, both surfaces hispid, secondary veins 4–6 on each sideof midvein, base broadly cuneate, margin entire, apex acute toobtuse. Spikes 2.5–4 cm, hispid; bracts linear-oblong, ca. 8 × 2–3 mm, hispid; bracteoles linear, ca. 6 mm. Calyx ca. 8 mm, out-

452ACANTHACEAEside hispid, 4-lobed almost to base; lobes linear, margin ciliate.Corolla purplish red, 0.8–1.2 cm; lower lip spreading, 3-lobed;upper lip erect, apex emarginate. Staminal filaments ca. 6 mm,sparsely hirsute; anther thecae superposed, lower one whitespurred at base. Ovary hirsute at tip; style ca. 8 mm, basal portionhirsute. Capsule ellipsoid, ca. 8 mm, pubescent. Seeds minutelyreticulate. 2n = 18.Trailsides; 2100–2700 m. Yunnan [India].“Justicia mollissima” was published as a nomen nudum (Wallich,Numer. List, no. 2445. 1830). The earliest name for this species isRostellaria mollissima, dating from 1832. Nees’ name has not beentransferred previously to Justicia.3. Justicia neolinearifolia N. H. Xia & Y. F. Deng, J. Trop.Subtrop. Bot. 13: 534. 2005.线叶爵床 xian ye jue chuangRostellularia linearifolia Bremekamp, Proc. Kon. Ned.Akad. Wetensch. C 60: 5. 1957; Justicia linearifolia (Bremekamp)H. S. Lo (1997), not Lamarck (1785); J. linearifoliasubsp. liangkwangensis (H. S. Lo) H. S. Lo; J. neolinearifoliasubsp. liangkwangensis (H. S. Lo) N. H. Xia & Y. F. Deng; R.linearifolia subsp. liangkwangensis H. S. Lo.Herbs to 50 cm tall. Stems 4-angled, sulcate, pubescent.Petiole 2–4 mm, pubescent; leaf blade linear, 2–4 × 0.2–0.5 cm,sparsely pubescent, secondary veins 3–5 on each side of midvein,base rounded, margin entire, apex acute. Spikes terminal,2–3.5 cm; rachis sparsely hirsute to subglabrous; bracts linear,ca. 6 × 1 mm, subglabrous, apex acute; bracteoles linear, ca. 5 ×1 mm, sparsely hirsute, apex acute. Calyx ca. 6 mm, outsidesparsely hirsute, 4-lobed almost to base; lobes linear, apex acuminate.Corolla purplish red, ca. 8 mm; lower lip 3-lobed; upperlip 2-lobed. Staminal filaments ca. 5 mm, sparsely hirsute;anther thecae superposed, lower one spurred at base. Ovary hirsute;style ca. 8 mm, basal portion hirsute. Capsule ca. 6 mm,pubescent. Seeds ovate in outline, ca. 1 × 1 mm, verrucose.Wetlands, trailsides. Guangdong, Guangxi, Yunnan [Thailand].4. Justicia procumbens Linnaeus, Sp. Pl. 1: 15. 1753.爵床 jue chuangEcbolium procumbens (Linnaeus) Kuntze; Justicia hayataeYamamoto var. decumbens Yamamoto; J. procumbens var.hirsuta Yamamoto; J. procumbens var. linearifolia Yamamoto;Rostellularia procumbens (Linnaeus) Nees; R. procumbens var.hirsuta (Yamamoto) S. S. Ying; R. procumbens var. linearifolia(Yamamoto) S. S. Ying; R. trichochila Miquel.Herbs 20–50 cm tall, procumbent. Stems 4-angled, sulcate,pubescent. Petiole 3–8 mm, pubescent; leaf blade elliptic,ovate-elliptic, or elliptic-oblong, 1.5–4 × 0.8–1.5 cm, subglabrousto sparsely hispid, cystoliths numerous, secondary veins3–6 on each side of midvein and prominent, base broadly cuneateto subrounded and slightly decurrent onto petiole, marginentire or slightly undulate, apex acute to obtuse. Spikes terminalor axillary in upper leaf axils, cylindric, 1–6 cm, dense; peduncle0.5–7 cm, densely pilose; bracts ovate to elliptic-lanceolate,2.5–8 × 0.6–1.3 mm, abaxially pilose, margin ciliate; bracteoleslanceolate, 2.5–5 mm, abaxially pilose, margin ciliate. Calyx 4–6 mm, 4-lobed to base; lobes linear, outside pilose along veins,1-veined, margin yellowish white and ciliate. Corolla pink orwhite and red-spotted on lower lip, 5–8.5 mm; lower lip ca. 3 ×3.5 mm, 3-lobed, lobes ovate, middle lobe slightly larger andca. 1 × 1–1.5 mm; upper lip ca. 3 mm, apex emarginate. Stamensexserted; filaments ca. 4 mm, glabrous; anther thecaesuperposed, lower one spurred at base, upper one muticous.Ovary pubescent; style ca. 5 mm, pubescent. Capsule 4–6 mm,4-seeded. Seeds ovate in outline, ca. 1 × 1 mm, rugose. Fl. andfr. all year. 2n = 18, 36.Wastelands, roadsides, lawns, open fields, littoral forests; near sealevel to 1500 m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou,Hainan, Hebei, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Shaanxi,Sichuan, Taiwan, Xizang, Yunnan, Zhejiang [Bangladesh, Bhutan, Cambodia,India, Indonesia, Japan, Laos, Malaysia, Myanmar, Nepal, Philippines,Sri Lanka, Thailand, Vietnam].5. Justicia diffusa Willdenow, Sp. Pl. 1: 87. 1797.小叶散爵床 xiao ye san jue chuangJusticia diffusa var. prostrata Roxburgh ex C. B. Clarke;Rostellaria hedyotidifolia Nees; Rostellularia diffusa (Willdenow)Nees; R. diffusa var. hedyotidifolia (Nees) Kumari; R.diffusa var. prostrata (Roxburgh ex C. B. Clarke) H. S. Lo; R.hedyotidifolia (Nees) Nees.Herbs 10–30 cm tall. Stems 4-angled, procumbent and diffuse,sulcate, sparsely pubescent. Petiole ca. 2 mm, pubescent;leaf blade lanceolate-elliptic, subcircular, or ovate, (0.7–)2–5 ×0.7–1.5 cm, glabrous or minutely pubescent, secondary veins2–4 on each side of midvein, base cuneate, margin entire, apexacute. Spikes (1.5–)2.5–5 cm, neither dense nor cylindric,sparsely hirsute; bracts oblong-lanceolate, 2–3 × ca. 1 mm,sparsely pubescent; bracteoles linear, ca. 2.5 × 1 mm, sparselypubescent. Calyx 3.5–5 mm, outside sparsely pubescent, 4- or5-lobed; lobes lanceolate, margin ciliate, apex acute. Corollapurplish red, ca. 6 mm; lower lip, 3-lobed, sparsely puberulent;upper lip emarginate. Staminal filaments ca. 4 mm, hirsute; antherthecae superposed, lower one spurred at base. Ovary pubescent;style ca. 5 mm, pubescent. Capsule oblong, ca. 5 mm,subglabrous to pubescent. Seeds subsmooth. 2n = 18.Grasslands, streamsides, sandy places. Fujian, Guangdong,Guangxi, Hainan, Taiwan, Yunnan [Bangladesh, India, Myanmar, SriLanka, Thailand, Vietnam].6. Justicia hayatae Yamamoto, Icon. Pl. Formosan. Suppl. 2:34. 1926 [“hayatai”].早田氏爵床 zao tian shi jue chuangJusticia ciliata (Yamamoto) C. F. Hsieh & T. C. Huang(1974), not Jacquin (1762); J. hayatae var. ciliata Yamamoto;J. procumbens Linnaeus var. hayatae (Yamamoto) Ohwi; Rostellulariahayatae (Yamamoto) S. S. Ying; R. procumbens (Linnaeus)Nees var. ciliata (Yamamoto) S. S. Ying.Herbs 30–50 cm tall. Stems 4-angled, sulcate, bifariouslypubescent along sulcae. Petiole 1–9 mm, glabrous; leaf bladeovate, 1.5–3.5 × 0.8–2 cm, glabrous, abaxially pale green andwithout cystoliths, adaxially green and with numerous cysto-

ACANTHACEAE 453liths, secondary veins 3 or 4 on each side of midvein, baseattenuate and decurrent onto petiole, margin entire, apex acute.Spikes mostly terminal or axillary at upper leaf axils, 1–4 cm,elongate to 7 cm in fruit; peduncle (0–)0.5–2 cm, glabrous;bracts obovate-elliptic, 4–5 × 2–3 mm, glabrous, margin white,apex acute; bracteoles lanceolate-elliptic, ca. 5 × 1.2 mm. Calyx4-lobed almost to base; lobes lanceolate, ca. 4 × 1 mm, glabrous,margin white. Corolla ca. 6 mm, glabrous, 2-lipped tohalf its length; lower lip oblong, ca. 3 × 3 mm, minutely 3-lobed; upper lip ovate, ca. 3 × 2 mm, apex emarginate. Stamensexserted; filaments ca. 2.5 mm, glabrous except for trichomesat base; anther thecae ovoid, ca. 1.3 mm, superposed, lowerone spurred at base, upper one muticous. Ovary glabrous; styleca. 6 mm, glabrous. Capsule with a brownish yellow short stipe,oblong, ca. 4 mm, glabrous, 4-seeded. Seeds ovate in outline,ca. 1 mm, rugose, glabrous.● Seashores, sandy places. Hong Kong, Taiwan.7. Justicia neesiana (Nees) T. Anderson, J. Linn. Soc., Bot. 9:513. 1867.狭叶爵床 xia ye jue chuangGendarussa neesiana Nees in Wallich, Pl. Asiat. Rar. 3:105. 1832; Adhatoda neesiana (Nees) Nees; Calophanoidesmultinodis (Benoist) C. Y. Wu & H. S. Lo; Justicia multinodisBenoist.Subshrubs to 30 cm tall. Stems basally woody, decumbent,and often rooting at nodes, then erect, apical potion muchbranched; branchlets blackish brown when dry, terete or ± 4-angled, pubescent at nodes. Leaves subsessile; leaf blade linearto linear-lanceolate, 1–3.5 × 0.2–0.8 cm, papery, glabrous, secondaryveins 4 on each side of midvein, base attenuate, marginentire, apex obtuse to rounded. Spikes axillary, ca. 1 cm, 1–3-flowered; bracts lanceolate to spatulate-lanceolate, 4–6 mm;bracteoles subulate, ca. 1 mm, glabrous. Calyx ca. 5 mm, 5-lobed almost to base; lobes lanceolate-linear. Corolla white,7–8 mm, pubescent; lower lip 3-lobed, lobes triangular and withan acute apex; upper lip triangular, 2-lobed. Staminal filamentspubescent on basal potion; anther thecae superposed, lower onewith a white spur at base. Capsule ca. 5 mm, glabrous. Fl. Jun–Aug, fr. Aug–Oct.On rocks, streamsides; 200–800 m. Hainan, Yunnan [Laos, Thailand,Vietnam].Specimens misidentified as Calophanoides loheri (C. B. Clarke)Bremekamp by C. C. Hu (FRPS 70: 284. 2002) belong here.8. Justicia kwangsiensis (H. S. Lo) H. S. Lo, Guihaia 17: 50.1997.广西爵床 guang xi jue chuangCalophanoides kwangsiensis H. S. Lo, Acta Phytotax. Sin.17(4): 86. 1979; C. buxifolia (H. S. Lo & D. Fang) C. Y. Wu &T. Y. Ding ex C. C. Hu; Justicia buxifolia H. S. Lo & D. Fang.Shrubs to 50 cm tall, erect. Branches stout, 4-angled, sulcate;branchlets glabrous, densely punctate with grayish whitecystoliths. Petiole 4–10 mm; leaf blade ovate to broadly elliptic,(0.5–)2–9 × 1–4 cm, leathery, glabrous, densely covered withcystoliths, secondary veins ca. 5 on each side of midvein, basebroadly cuneate, margin entire, apex acuminate and sometimesshortly caudate. Spikes axillary, ca. 1 cm, 1–3-flowered; bractssubcircular, (1.5–)4–5 mm in diam., glabrous; bracteoles subulate,ca. 2 × 1 mm, glabrous. Calyx 5–6 mm, glabrous, 5-lobedalmost to base; lobes narrowly lanceolate, apex acuminate. Corollawhite, 7–9 mm; lower lip 3-lobed; upper lip triangular, 2-toothed. Staminal filaments glabrous; anther thecae superposed,lower one spurred at base. Capsule fusiform, ca. 8 mm, glabrous,4-seeded. Seeds yellowish brown, tuberculate. Fl. Sep–Nov, fr. Dec–Feb.● Limestone hills; below 700 m. Guangdong, Guangxi, Hainan.9. Justicia alboviridis Benoist, Notul. Syst. (Paris) 5: 115.1936.大叶杜根藤 da ye du gen tengCalophanoides alboviridis (Benoist) C. Y. Wu & H. S. Lo.Herbs to 50 cm tall, perennial. Stems basally slightly decumbentthen erect; young branches 4-angled, sulcate and striate,bifariously pubescent, glabrescent. Petiole 3–8 mm; leafblade ovate to rarely lanceolate, (1–)3–9 × (0.5–)1.5–4 cm, papery,both surfaces glabrous, secondary veins 4 or 5 on eachside of midvein, base slightly oblique and cuneate to subrounded,margin subentire, apex shortly acuminate. Spikes axillary,ca. 1 cm, (1–)3–5-flowered; bracts lanceolate, 1–3 mm,glabrous; bracteoles linear, ca. 1.5 mm. Calyx ca. 4 mm, 5-lobed; lobes lanceolate, outside glabrous except pubescentalong midvein. Corolla white or greenish white, 8–9 mm, outsidepubescent; lower lip elliptic, 3-lobed, lobes ovate with anobtuse apex; upper lip narrowly triangular, 2-lobed. Ovaryslightly hirsute at tip; style basally hirsute. Capsule ca. 7 cm,glabrous. Seed light yellow, papillate. Fl. Aug–Sep, fr. Nov–Jan.Wet places, streamsides; 200–600 m. Hainan [Vietnam].10. Justicia wardii W. W. Smith, Notes Roy. Bot. Gard. Edinburgh10: 184. 1918.高山杜根藤 gao shan du gen tengHu.Calophanoides wardii (W. W. Smith) C. Y. Wu ex C. C.Shrubs to 60 cm tall, diffuse. Stems straw-colored, muchbranched, flexuous, densely grayish tomentose, soon glabrescent.Petiole ca. 5 mm, gray pilose; leaf blade elliptic, ellipticlanceolate,oblanceolate, or sometimes obovate, to 4.5 × 2 cm,papery, both surfaces first white tomentose then sparsely pilose,base cuneate, margin slightly undulate, apex shortly acuminate,acute, or rarely retuse. Spikes axillary, ca. 1 cm, (1–)3–5-flowered;peduncles 0–3 mm; bracts linear-lanceolate, ca. 5 mm,densely white villous; bracteoles linear, ca. 2 mm. Calyx to8 mm, 5-lobed almost to base; lobes oblanceolate, outsidedensely villous, apex acute. Corolla ca. 2 cm, outside white villous;tube longer than limb; lower lip 3-lobed to middle, lobesoblong; upper lip 7–8 mm, shallowly 2-lobed. Staminal filamentsglabrous, basally whitish; anther thecae superposed,lower one spurred at base. Ovary glabrous; style white villous.Capsule not seen. Fl. May.

454ACANTHACEAE● On rocks, thickets; ca. 1000 m. Yunnan.11. Justicia championii T. Anderson in Bentham, Fl. Hongk.264. 1861.圆苞杜根藤 yuan bao du gen tengAdhatoda chinensis Bentham, Hooker’s J. Bot. Kew Gard.Misc. 5: 134. 1853; Calophanoides chinensis (Bentham) C. Y.Wu & H. S. Lo; Dicliptera cyclostegia Handel-Mazzetti; Justiciachinensis (Bentham) Druce (1917), not Linnaeus (1753).Herbs to 50 cm tall. Stems 4-angled, sulcate, retrorsely pubescent.Petiole (0–)0.5–1.5 cm, sulcate, pubescent; leaf bladeelliptic, oblong-elliptic, ovate, or lanceolate, 1–7(–10.5) × 0.5–2(–3.5) cm, sparsely pubescent when young, soon glabrescentor abaxially pilose along midvein, cystoliths adaxially numerous,secondary veins 3–6 on each side of midvein, base cuneateto attenuate, margin entire, apex acute to obtuse. Spikes axillary,usually 3-flowered, several clustered; peduncle 2–4 mm,sparsely pubescent; bracts petiolate, subleaflike, circular to obovate-spatulateor sometimes elliptic, 5–11 × 2–6 mm, glabrousor adaxially sparsely pilose, veins 1–3 on each side of midvein,apex rounded to obtuse; bracteoles subulate to triangular, ca. 1mm, glabrous or sparsely yellow puberulent, apex acute. Calyx5-lobed almost to base; lobes yellowish white except for greenalong veins, linear-lanceolate, 6–7 mm, 1- or 2-veined, marginciliate. Corolla white with 2 purplish red streaks, 7–10 mm,outside puberulent; lower lip ovate, ca. 4 × 3 mm, 3-lobed, middlelobe semicircular and ca. 0.8 × 1.3 mm, lateral lobes ovateand ca. 1.2 × 1 mm; upper lip lanceolate, ca. 3.5 × 1.5 mm,apex emarginate. Stamens exserted; filaments ca. 4 mm, glabrous;anther thecae superposed, lower one spurred at base, upperone muticous. Ovary glabrous; style ca. 6 mm, sparsely pubescenttoward base; stigma capitate. Capsule 8–9 mm, glabrous,4-seeded. Seeds oblong, ca. 2 × 1.5 mm, verrucose. Fl.and fr. Aug–Oct.● Thickets, forests; 400–2000 m. Anhui, Fujian, Guangdong,Guangxi, Guizhou, Hainan, Hubei, Hunan, Jiangxi, Yunnan, Zhejiang.Inflorescences of Justicia championii appear to be axillary clustersof flowers but are much-reduced spikes or racemes with a very shortrachis.12. Justicia xerophila W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 11: 214. 1919.干地杜根藤 gan di du gen tengCalophanoides xerophila (W. W. Smith) C. Y. Wu ex C. C.Hu.Herbs to 10 cm tall. Stems basally semi-woody, slenderand flexuous, yellow pilose. Petiole 1–3 mm, pilose; leaf bladelanceolate to oblanceolate, 0.8–1.5 × 0.3–0.5 cm, papery, bothsurfaces slightly pilose along veins, secondary veins ca. 3 oneach side of midvein, base cuneate, margin ciliate and entire orapically 2 or 3 repandly toothed, apex acute to obtuse. Spikesaxillary, ca. 1 cm, 1–3-flowered; bracts leaflike, broadly obovateto subcircular, 3–5 mm; bracteoles linear, ca. 2 mm. Calyxgreen, ca. 5 mm, outside slightly pilose to subglabrous, 5-lobedalmost to base; lobes linear, apex acuminate. Corolla white, 7–8mm, outside sparsely pilose; lower lip 3-lobed; upper lip emarginate.Staminal filaments ca. 4 mm, pilose; anther thecae superposed,lower one spurred at base. Ovary glabrous; style ca. 6mm, subglabrous. Capsule ca. 5 mm, glabrous. Seeds ovate inoutline, ca. 1 × 1 mm, tuberculate.● Thickets on dry slopes. Yunnan.13. Justicia xylopoda W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 11: 214. 1919.木柄杜根藤 mu bing du gen tengCalophanoides xylopoda (W. W. Smith) C. Y. Wu ex H. P.Tsui.Shrubs 10–13 cm tall. Stems very short, much branched;branches basally often cylindric, erect or decumbent, apicallyascending, grayish or whitish pubescent, internodes sulcate.Petiole 1–3 cm, white pubescent; leaf blade ovate to narrowlyovate, 1–2.4 × 0.7–1.2 cm, papery, abaxially minutely andsparsely pilose on veins, adaxially glabrous, secondary veins 3on each side of midvein, base cuneate and decurrent onto petiole,margin entire, apex ± obtuse. Spikes axillary, ca. 1 cm, usually3-flowered; bracts ovate, ca. 1 cm, pubescent like leaves;bracteoles sublinear, ca. 1.5 mm. Calyx green, ca. 6 mm, 5-lobed to base; lobes linear-lanceolate, membranous, subequal,margin white, apex acuminate. Corolla yellow with purplishspots, ca. 1 cm, outside white pilose; lower lip broad, 3-lobedwith center one larger; upper lip triangular, 2-lobed. Stamensinserted at apical part of corolla tube; filaments pilose aroundinsertion; anther thecae superposed, lower one with a whitespur at base. Ovary glabrous. Capsule not seen.● Thickets on mountain slopes; 2300–2400 m. Sichuan, Yunnan.14. Justicia albovelata W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 10: 182. 1918.绵毛杜根藤 mian mao du gen tengCalophanoides albovelata (W. W. Smith) C. Y. Wu ex H.P. Tsui.Herbs 20–50 cm tall, perennial, much branched. Stemsbasally decumbent then erect and zigzag; young branchletsdensely white villous. Petiole 3–5 mm, pubescent; leaf bladelanceolate, 3.5–4.5 × 0.8–1.5 cm, membranous, abaxially villous,adaxially densely white villous, secondary veins 4 or 5 oneach side of midvein, base cuneate, margin entire, apex obtuse.Spikes axillary, ca. 1 cm, usually 3(or 4)-flowered; bracts spatulate-circularto obovate, 5–10 × 3–7 mm; bracteoles oblanceolateto linear. Pedicels ca. 2 mm. Calyx grayish green, 5–6 mm,5-lobed almost to base; lobes linear-lanceolate, with densewhite trichomes along veins, apex acuminate. Corolla whitishyellow to yellow, ca. 9 mm; tube slightly longer than limb,outside pilose; lower lip 3-lobed; upper lip short, 2-lobed.Staminal filaments basally hirsute; anther thecae superposed,lower one larger and with a white spur at base. Ovary glabrous.Capsule clavate, ca. 8 mm, glabrous, 4-seeded. Seeds ovate inoutline, ca. 1.5 × 1.5 mm, verrucose. Fl. Sep.● Thickets, streamsides; ca. 2700 m. Yunnan.

ACANTHACEAE 45515. Justicia yunnanensis W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 11: 215. 1919.滇杜根藤 dian du gen tengCalophanoides yunnanensis (W. W. Smith) C. Y. Wu ex H.P. Tsui.Shrubs to 45 cm tall. Stems basally unbranched, suberect,apically ascending, whitish sericeous. Petiole 2–5(–7) mm, pilose;leaf blade lanceolate to oblanceolate, ca. 2.5 × 1 cm, papery,both surfaces pilose, base narrowly cuneate and decurrentonto petiole, margin entire, apex ± obtuse. Spikes axillary, ca. 1cm, usually 3–5-flowered; bracts surrounded by leaves, circular-spatulateto obovate, 5–10 mm; bracteoles linear to oblanceolate.Pedicel short. Calyx light grayish green, ca. 5 mm, 5-lobed to base; lobes linear-lanceolate, midvein pale whitish yellow,apex acuminate. Corolla light whitish yellow, ca. 8 mm,outside minutely pilose; lower lip broad, 3-lobed with centerone larger; upper lip triangular, 2-lobed. Stamens inserted atapical part of corolla tube, pilose around insertion; antherthecae superposed, lower one with a white spur at base. Capsuleglabrous, 4-seeded. Seeds verrucose. Fl. Sep.● Limestone slopes; ca. 1600 m. Yunnan.Based on the known characters, Justicia yunnanensis appearsremarkably similar to J. albovelata, if it is indeed distinct. We haverefrained from making taxonomic alterations because no specimens ofJ. yunnanensis were studied.16. Justicia hainanensis (C. Y. Wu & H. S. Lo) N. H. Xia & Y.F. Deng, J. Trop. Subtrop. Bot. 13: 533. 2005.海南爵床 hai nan jue chuangCalophanoides hainanensis C. Y. Wu & H. S. Lo, Fl. Hainan.3: 597. 1974.Herbs to 1 m tall. Stems 4-angled, sulcate, pubescent.Petiole 2–5 mm; leaf blade ovate to lanceolate-ovate, (0.5–)1.5–3 × (0.3–)0.5–1.8 cm, thickly papery, both surfaces hispid, secondaryveins 3–5 on each side of midvein, base broadly cuneateto subrounded, margin entire, apex acute. Spikes axillary, ca. 1cm, (1–)3–5-flowered; bracts shortly petiolate, circular to subcircular,2–5 mm in diam.; bracteoles small or absent. Calyx ca.4.5 mm, hispid, 5-lobed almost to base; lobes linear-lanceolate.Corolla white, ca. 6 mm, outside densely pubescent; tubeslightly enlarged; lower lip spreading, apex 3-lobed; upper liperect, triangular and concave, apex emarginate. Staminal filamentsca. 2.5 mm; anther thecae superposed, lower one spurredat base. Ovary glabrous; style ca. 3 mm, glabrous. Capsule obovoid,ca. 6 mm, glabrous, 4-seeded. Seeds broadly ovate in outline,ca. 0.6 × 0.6 mm, compressed, verrucose. Fl. Jul–Nov, fr.Oct–Jan.● Forests; 200–1100 m. Guangdong, Hainan.17. Justicia quadrifaria (Nees) T. Anderson, J. Linn. Soc.,Bot. 9: 514. 1867.杜根藤 du gen tengGendarussa quadrifaria Nees in Wallich, Pl. Asiat. Rar. 3:105. 1832; Adhatoda quadrifaria (Nees) Nees; A. zollingerianaNees; Calophanoides quadrifaria (Nees) Ridley.Herbs. Stems 4-angled, basally decumbent and usuallyrooting at nodes then erect, sulcate, bifariously pubescent, soonglabrescent. Petiole 0.4–2.5 cm, sparsely pubescent; leaf bladeoblong, lanceolate, to rarely orbicular, (2–)6–8 × (1–)2–3.5 cm,both surfaces glabrous or only pubescent along veins, secondaryveins 5–7 on each side of midvein, base narrowly cuneateand decurrent onto petiole, margin entire or slightly serrate,apex acute. Spikes axillary, ca. 1 cm, usually 3-flowered, solitaryor several in a cluster; bracts with a 3–4 mm petiole, ovateto obovate, 3–8 × 3–5 mm, sparsely pubescent, pinnatelyveined, apex obtuse; bracteoles linear, ca. 1 mm, glabrous. Calyx5–6 mm, outside puberulent, 5-lobed almost to base; lobeslinear-lanceolate. Corolla white with purple spots on lower lip,ca. 8 mm, outside pubescent; lower lip ca. 4 × 3 mm, patent,apex 3-lobed; upper lip oblong, erect, apex 2-lobed. Antherswith thecae superposed, lower ones spurred at base. Ovary glabrous;style ca. 6 mm, sparsely hirsute. Capsule ca. 8 mm, glabrous.Seeds ovate in outline, ca. 1.5 × 1 mm, verruculose, glabrous.On rock in forested valleys; 800–1600 m. Chongqing, Guangdong,Guangxi, Guizhou, Hainan, Hubei, Hunan, Sichuan, Yunnan [India,Indonesia, Laos, Myanmar, Thailand, Vietnam].18. Justicia kouytcheensis (H. Léveillé) E. Hossain, NotesRoy. Bot. Gard. Edinburgh 32: 407. 1973.贵州杜根藤 gui zhou du gen tengRuellia repens Linnaeus var. kouytcheensis H. Léveillé,Repert. Spec. Nov. Regni Veg. 13: 175. 1914; Calophanoideskouytcheensis (H. Léveillé) H. S. Lo.Herbs to 40 cm tall. Stems basally decumbent and rootingat nodes. Petiole 1–2.5 cm; leaf blade oblong to rhombic, 5.5–11.5 × 3.5–5.5 cm, membranous, both surfaces pubescentespecially along veins, secondary veins 6 or 7 on each side ofmidvein, base cuneate and decurrent onto petiole, margin entireor sometimes apically undulate-serrate, apex acuminate tosubcaudate. Spikes axillary, ca. 1 cm, 1–3(–5)-flowered; bractswith a 1–2 mm petiole, obovate to circular, 7–8 × 4–5 mm,veins ca. 3 on each side of midvein, base broadly cuneate, apexretuse or rounded; bracteoles subulate, 1–2 mm. Calyx ca. 1cm, 5-lobed almost to base; lobes lanceolate, subequal, outsidewhite laterally, green along midvein, and pubescent. Corollawhite, ca. 1.4 cm; lower lip patently 3-lobed; upper lip erect, 2-lobed. Staminal filaments ca. 5 mm, glabrous; anther thecae oblong,superposed, lower one smaller and with a white spur atbase. Ovary glabrous; style ca. 9 mm, glabrous; stigma capitate,slightly enlarged. Capsule and seeds not seen. Fl. Aug–Oct.● Limestone hills, thickets; 800–1300 m. Guizhou, Yunnan.Justicia kouytcheensis is very similar to J. quadrifaria, and theputative distinctions between them require additional studies.19. Justicia xerobatica W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 11: 213. 1919.滇东杜根藤 dian dong du gen tengCalophanoides xerobatica (W. W. Smith) H. P. Tsui.Subshrubs to 30 cm tall, prostrate. Stems basally woody,slender, flexuous, and often rooting at nodes, apically as-

456ACANTHACEAEcending, minutely white sericeous. Petiole 2–3 cm, pilose; leafblade subcircular, broadly obovate, or rarely narrowly lanceolate,1–2 × 0.5–0.8 cm, softly membranous, both surfaces piloseon veins, base cuneate, margin entire, apex rounded to obtuse.Spikes axillary, ca. 1 cm, 1–3-flowered; bracts surrounded byleaves, leaflike, broadly obovate to subcircular, 3–4 mm; bracteolesovate to nearly subulate-lanceolate, small. Calyx green, ca.5 mm, 5-lobed almost to base; lobes linear-lanceolate, midribpilose, margins scarious, apex acuminate. Corolla white (reportedly),ca. 9 mm, outside sparsely pilose. Mature capsule notseen. Fl. Sep.● Limestone areas; 1200–1400 m. Sichuan, Yunnan.Justicia xerobatica is very similar to J. xerophila. Further studiesof both will be necessary to determine whether they should be treated asdistinct from one another.20. Justicia siccanea W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 10: 43. 1917.旱杜根藤 han du gen tengCalophanoides siccanea (W. W. Smith) C. Y. Wu ex C. C.Hu.Shrubs to 60 cm tall, prostrate. Stems woody, flexuous,grayish tomentose, soon glabrescent. Petiole 3–8 cm, villous;leaf blade ovate to lanceolate, 1.5–4 × 1–2 cm, both surfacessparsely pilose, secondary veins ca. 4 on each side of midvein,base cuneate, margin entire, apex rounded to acute. Spikes axillary,ca. 1 cm, 1–3-flowered; peduncle ca. 3 mm, grayish pubescent;bracts linear, 2–3 mm, abaxially grayish pubescent;bracteoles similar to bracts. Calyx ca. 7 mm, outside pubescent,5-lobed to base; lobes linear. Corolla white tinged with red, ca.1.3 cm, outside pubescent; lower lip very short, 3-lobed, lobescircular; upper lip shorter, apex emarginate. Staminal filamentsca. 4 mm, basal portion whitish villous; anther thecae superposed,lower one spurred at base. Ovary glabrous; style ca. 8mm, white villous. Capsule ca. 7 mm, glabrous. Seeds ovate inoutline, ca. 1 × 1 mm. Fl. Sep.● On rocks by trailsides, open slopes; 2000–2200 m. Sichuan,Yunnan.21. Justicia xantholeuca W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 11: 212. 1919.黄白杜根藤 huang bai du gen tengCalophanoides xantholeuca (W. W. Smith) C. Y. Wu ex C.C. Hu.Shrubs to 30 cm tall. Stems 4-angled, much branched, sulcate,pilose then glabrescent. Petiole 2–5 mm, pilose; leaf bladelanceolate, 2–3 × 0.7–1 cm, papery, both surfaces pilose alongveins, secondary veins ca. 3 on each side of midvein, base cuneate,margin entire, apex acute. Spikes axillary, 3–5-flowered;bracts lanceolate or ovate, 5–6 mm; bracteoles subulate, narrowlylanceolate, or ovate, smaller than bracts. Calyx grayishwhite, ca. 5 mm, white pilose along midvein, 5-lobed almost tobase; lobes linear-lanceolate. Corolla white, ca. 8 mm, outsidesparsely pilose; lower lip 3-lobed; upper lip emarginate. Staminalfilaments ca. 5 mm, sparsely hirsute; anther thecae superposed,lower one spurred at base. Ovary glabrous; style ca. 6mm, subglabrous. Capsule ca. 5 mm, glabrous. Seeds ovate inoutline, ca. 1 × 1 mm, tuberculate-verrucose. Fl. Nov–Jan.● Thickets on dry slopes; 400–800 m. Yunnan.22. Justicia grossa C. B. Clarke in J. D. Hooker, Fl. Brit. India4: 535. 1885.大爵床 da jue chuangShrubs 50–100 cm tall. Stems subterete, puberulent whenyoung then glabrescent. Petiole 1.3–4 cm, glabrous or puberulent;leaf blade elliptic to lanceolate, 16–30 × 5–9.5 cm, abaxiallyglabrous or puberulent along veins, adaxially glabrous,secondary veins 5–8 on each side of midvein, base broadlyrounded to cuneate, margin crenulate, apex acuminate to attenuateand with an obtuse tip. Inflorescences terminal, thyrsoid of3-flowered cymes, usually 3 or more branched, 5–15 × 2–3 cm;peduncle 1–3.5 cm; rachis hirsute; bracts ovate, elliptic, orlanceolate, 2–8 × 1–5 mm, abaxially puberulent; bracteoleslanceolate to rarely triangular, 1.5–3 × 0.5–1 mm, abaxially puberulent.Pedicel 1–5 mm, hirsute. Calyx ca. 6 mm, both surfacesgland-tipped pubescent, 5-lobed to base; lobes linear.Corolla greenish white with pale purple dots on lower lip, 1.2–1.5 cm, outside pubescent; tube 6–7 mm; limb 6–7 mm; lowerlip 3-lobed, lobes ovate and ca. 2 × 1.5–2 mm; upper lip emarginate.Staminal filaments 4–6 mm, basal portion densely pubescent;anther thecae 1.5–2 mm, both spurred at base. Ovarypubescent; style pubescent. Capsule ca. 1.6 cm, densely pubescent.Seeds elliptic to circular in outline, ca. 3.5 × 3.5 mm,wrinkly verrucose.Forests; 400–800 m. Hainan [Laos, Malaysia, Myanmar, Thailand,Vietnam].23. Justicia adhatoda Linnaeus, Sp. Pl. 1: 15. 1753.鸭嘴花 ya zui huaAdhatoda vasica Nees.Shrubs to 4 m tall. Branches thickened, ± 4-angled, lenticellate,pubescent when young but soon glabrescent. Petiole0.8–3 cm, puberulent; leaf blade ovate to elliptic-ovate, 7–18 ×2–7 cm, abaxially puberulent, adaxially densely tomentosewhen young but glabrescent except tomentose along veins, secondaryveins 9–12 on each side of midvein and meeting nearmargin, base broadly cuneate, margin entire, apex acuminateand sometimes slightly falcate. Spikes terminal or axillary,ovoid to broadly ovoid, 3–7 cm; peduncle 3–7 cm; bractsimbricate, ovate-oblong, 1–2 × 0.5–1.5 cm, puberulent, 3–7-veined, margin ciliate, apex acute; bracteoles linear-lanceolate,1–1.4 × ca. 0.4 cm, puberulent, 3–5-veined, margin ciliate, apexacute. Calyx 5-lobed; lobes linear-oblong, ca. 10 × 3 mm, marginnarrowly scarious and ciliate. Corolla white or pink withpurplish or pinkish stripes outside, broadly tubular, 2.5–3 cm,outside pilose; tube basally cylindric and ca. 5 mm wide for ca.5 mm and then slightly inflated and bent upward; upper lipovate-oblong, ca. 1.8 cm, erect, shallowly 2-lobed; lower lipoblong-circular, spreading, 3-lobed, middle lobe subcircular andca. 9 × 5 mm, lateral lobes ovate and ca. 8 × 4.5 mm. Stamensexserted; filaments ca. 1.5 cm, declinate, glabrous except basallysericeous; anther thecae ellipsoid, ca. 3.5 mm, equal,

ACANTHACEAE 457superposed, lower one spurred at base. Ovary pubescent especiallyat tip; style ca. 2.5 cm, recurved, basal part pubescent;stigma simple. Capsule obovoid, 2.5–4 × ca. 0.5 cm. Seeds circularin outline. Fl. Jan–Mar, fr. Jun–Jul. 2n = 34.Roadsides, thickets. Naturalized or cultivated in Guangdong,Guangxi, Hainan, and Yunnan [probably native to India, Indonesia, Malaysia,Nepal, Pakistan, and Sri Lanka, but widely cultivated and naturalizedin tropics, obscuring native distribution].Justicia adhatoda is widely cultivated beyond its native range.This species is used medicinally.24. Justicia vagabunda Benoist, Notul. Syst. (Paris) 5: 114.1936.针子草 zhen zi caoRhaphidospora vagabunda (Benoist) C. Y. Wu ex C. C.Hu.Shrubs to 1.3 m tall. Stems terete, zigzag, glabrous. Petiole7–12 mm; leaf blade oblong-lanceolate, 6–14 × 2–5 cm, glabrousor sometimes pilose along midvein, base obliquely cuneate,margin entire or slightly undulate, apex acute to falcate.Cymes axillary, ca. 2 cm; peduncle 4–6 mm, pubescent; bractssubulate, 1–2 mm, pilose; bracteoles 2–5 mm, pilose. Calyx ca.3.5 mm, outside pubescent; lobes 5, oblong-lanceolate. Corollawhite with blue lines, 1–1.1 cm, outside pubescent; tube as longas limb; lower lip 3-lobed; upper lip triangular, apex emarginate.Staminal filaments ca. 4 mm, glabrous; anther thecaesuperposed, lower one spurred at base. Ovary pubescent at tip;style basal portion hirsute. Capsule not seen.Forests, thickets, streamsides; 500–800 m. Yunnan [Cambodia,Vietnam].25. Justicia ventricosa Wallich ex Hooker, Bot. Mag. 54: t.2766. 1827.黑叶小驳骨 hei ye xiao bo guAdhatoda ventricosa (Wallich ex Hooker) Nees; Gendarussaventricosa (Wallich ex Hooker) Nees.Subshrubs or perennial herbs, to 1 m tall. Stems subterete,glabrous. Winter buds ovoid, ca. 2 mm, pubescent. Petiole 0.5–1.5 cm, sulcate, glabrous; leaf blade elliptic to obovate, 6–17 ×2–6 cm, glabrous, midvein abaxially prominent and adaxiallyimpressed, secondary veins 6–8 on each side of midvein,prominent on both surfaces, and meeting near margin, baseattenuate, margin entire or slightly undulate, apex shortly acuminateto acute. Spikes terminal, 5–10 cm, densely flowered,usually with 1–3 flowers on each inflorescence node; peduncleca. 1 cm; bracts leaflike, green to maroon, imbricate, broadlyovate to subcircular, 1–1.5 × 0.8–1 cm, abaxially pilose to subglabrous,pinnately veined, margin ciliate, apex acute; bracteoleslinear, 3–5 × 1–2 mm, margin ciliate. Calyx 5-lobed; lobeslinear-lanceolate, ca. 3 mm, margin ciliate. Corolla creamywhite with purplish pink stripes on lips, 1.5–1.8 cm, outsidepilose, inside pubescent; tube basally cylindric and ca. 1.7 mmwide for ca. 1 cm; lower lip 3-lobed, lobes ovate, ca. 2 × 1mm, and apex rounded; upper lip oblong-ovate, 2-lobed. Stamensexserted; filaments ca. 6 mm, glabrous; anther thecaeovoid, subequal, superposed, lower one with a white appendageat base. Ovary comose; style ca. 1.6 cm, base pilose. Capsuleca. 8 mm, pubescent, 4-seeded. Fl. Nov–Jan, fr. Apr–May.2n = 30.Sparse forests, thickets. Naturalized or cultivated in Guangdong,Guangxi, Hainan, and Yunnan [native to Cambodia, Laos, Myanmar, NThailand, and Vietnam].This species is used medicinally.26. Justicia gendarussa N. L. Burman, Fl. Indica, 10. 1768.小驳骨 xiao bo guGendarussa vulgaris Nees.Subshrubs 0.7–1.5 cm tall, much branched. Stems subterete,swollen at nodes, glabrous. Petiole 3–10 mm; leaf bladenarrowly lanceolate, 6–10 × 1–1.5 cm, glabrous, secondaryveins 5–8 on each side of midvein, base cuneate to attenuate,margin subsinuate, apex acute to shortly acuminate. Spikes terminalor axillary, 3–12 cm, interrupted, usually in a leafy panicle;peduncle 0.5–1.5 cm; bracts triangular, 2–6 × 1–2.5 mm,basal ones longer than calyx then gradually smaller with apicalmostones shorter than calyx, margin ciliate, apex acute;bracteoles elliptic to linear-lanceolate, ca. 3 × 1 mm, marginciliate, apex acute. Calyx ca. 5 mm, 5-lobed; lobes linear-lanceolate,3–4 × ca. 0.5 mm, subequal, apex acuminate. Corollacreamy white, 1.2–1.5 cm; tube basally cylindric and ca. 2 mmwide for 8–9 mm; lower lip violet dotted basally, cuneate-obovate,6–10 mm broad, 3-lobed, lobes oblanceolate and 3–5 × ca.3.5 mm; upper lip violet blotched, triangular, ca. 7 × 3.5 mm, 2-cleft. Stamens exserted; filaments 3–6 mm, glabrous; antherthecae oblong, ca. 1.2 mm, superposed, lower one spurred atbase, upper one muticous. Ovary glabrous; style ca. 1 cm, glabrous;stigma capitate, shortly 2-lobed. Capsule clavate, ca. 1.2cm. Fl. Jan–Apr. 2n = 28, 30.Roadsides, thickets. Naturalized or cultivated in Guangdong,Guangxi, Hainan, Taiwan, and Yunnan [native to or naturalized in Cambodia,India, Indonesia, Laos, Malaysia, Myanmar, Papua New Guinea,Philippines, Sri Lanka, Thailand, and Vietnam; widely cultivated].Justicia gendarussa is native to S and SE Asia but is widelycultivated and naturalized. Its actual native distribution is not certainbecause it is often not clear whether particular collections are native.27. Justicia pseudospicata H. S. Lo & D. Fang, Guihaia 17:52. 1997.黄花爵床 huang hua jue chuangMananthes pseudospicata (H. S. Lo & D. Fang) C. Y. Wu& C. C. Hu.Herbs to 30 cm tall, perennial. Stems 4-angled, basally usuallyrooting at nodes, sulcate, bifariously pubescent. Petiole 3–13 mm, pubescent; leaf blade ovate, (0.8–)1.5–6.5 × (0.4–)1–4cm, thickly papery, abaxially glabrous, adaxially glabrous or piloseon veins, secondary veins 4 or 5 on each side of midvein,base cuspidate to rounded, margin subundulate, apex acute.Spikes often terminal or rarely axillary in apicalmost leaf axil,1.5–3 cm, interrupted; peduncle 1–2.5 cm; rachis bifariouslypubescent; bracts broadly obovate to sometimes circular orovate, 3–5 × 2–4 mm, abaxially sparsely gland-tipped puberulent,apex cuspidate; bracteoles narrowly triangular, ca. 2 × 1mm, abaxially pubescent, margin ciliate, apex acute. Calyx 2–3

458ACANTHACEAEmm, 5-lobed almost to base; lobes linear. Corolla yellow withpurplish spots, 1.2–1.4 cm, outside pubescent; lower lip palate3-lobed, lobes triangular and 1–2 mm; upper lip 4–5.5 mm,apex emarginate. Staminal filaments 2.5–3 mm, glabrous; antherthecae ca. 1.5 mm, lower one spurred at base. Ovary glabrous;style 5–8 mm, subglabrous. Capsule not seen. Fl. Sep–Dec.● On rocks in forests; 1300–1700 m. Guangxi.28. Justicia latiflora Hemsley, J. Linn. Soc., Bot. 26: 245.1890.紫苞爵床 zi bao jue chuangMananthes latiflora (Hemsley) C. Y. Wu & C. C. Hu.Herbs or shrubs 50–100 cm tall, erect. Stems 4-angled,sulcate, sparsely pubescent. Petiole 1–3.5 cm, pubescent; leafblade lanceolate, ovate, or subcircular, 6–10 × 2–3.5 cm, bothsurfaces hispid along veins, secondary veins 8–10 on each sideof midvein, base cuneate to sometimes attenuate, margin entire,apex acuminate. Spikes terminal, 3.5–5.5 cm, densely flowered;bracts basally pale green and purplish red above middle, ovateto elliptic, 7–10 × 3–4 mm, abaxially puberulent, apex acuminate;bracteoles linear, 5–7 × ca. 1.5 mm, abaxially puberulent.Calyx 5–8 mm, outside puberulent, 5-lobed almost to base;lobes linear, apex acute. Corolla pale white with purplish reddots or stripes on lobes, ca. 1.5 cm, outside pubescent; tube ca.1 cm; lower lip patent, broadly 3-lobed, lobes circular and ca. 1mm in diam.; upper lip ca. 0.5 mm, broadly rounded andconcave, apex emarginate. Staminal filaments glabrous; antherthecae superposed, lower one minutely spurred at base. Ovaryglabrous; style glabrous. Capsule obovoid-clavate, 1–1.2 cm,glabrous. Seeds ovate in outline, ca. 4 × 3.5 mm, rugose. Fl.Feb–Apr, fr. May–Jun.● Forests, streamsides; 600–1800 m. Chongqing, Guizhou, Hubei,Hunan.29. Justicia austrosinensis H. S. Lo, Guihaia 17: 52. 1997.华南爵床 hua nan jue chuangHu.Mananthes austrosinensis (H. S. Lo) C. Y. Wu & C. C.Herbs 40–70 cm tall. Stems 4-angled, sulcate, glabrous.Petiole 0.5–3 cm, bifariously pubescent or glabrescent; leafblade broadly ovate to ovate-elliptic, 7–15 × 2.5–7 cm, glabrous,abaxially pale green, adaxially green and with numerouscystoliths especially on veins, secondary veins 5–8 on each sideof midvein, base cuneate and decurrent onto petiole, margincrenate or entire, apex shortly acuminate to acute. Spikesmostly terminal or rarely axillary, 5–7 cm; peduncle 1–3 cm,with numerous cystoliths, glabrous; bracts green, usually subtending1 or 2 flowers, flabellate or broadly obovate, 5–8 × 7–9mm, margin ciliate, apex acuminate to obtuse and mucronate,with 1–3 teeth; bracteoles subulate, ca. 3 mm, margin ciliate,apex acuminate. Calyx 5-lobed almost to base; lobes lanceolate,ca. 4 mm, margin ciliate, apex acuminate. Corolla yellowishgreen with purplish red dots on lobes, ca. 1.3 cm, outside pilose,inside glabrous except for trichomes retaining style; lip ca. 1/2as long as corolla; lower lip 3-lobed, lobes ovate, ca. 2 × 2 mm,and apex rounded; upper lip ovate-triangular, ca. 6 × 3 mm,erect, apex emarginate. Stamens exserted; filaments ca. 4 mm,glabrous; anther thecae ca. 2 mm, unequal, superposed, lowerone spurred at base, upper one muticous. Ovary glabrous; styleca. 1 mm, pilose. Capsule clavate, ca. 1.2 cm, pubescent, 4-seeded. Seeds ovate in outline, ca. 2 mm, rugose. Fl. Jul–Oct.● Thickets in ravines. Guangdong, Guangxi, Guizhou, Jiangxi,Yunnan.30. Justicia ferruginea H. S. Lo & D. Fang, Guihaia 17: 58.1997.锈背爵床 xiu bei jue chuangMananthes ferruginea (H. S. Lo & D. Fang) C. Y. Wu &C. C. Hu.Herbs 20–60 cm tall, anisophyllous. Stems 4-angled, sulcate,glabrous. Leaves sessile; leaf blade oblanceolate to rarelylanceolate, 8–17 × 2.5–6.5 cm, papery, glabrous, abaxially rustcoloredwhen dry, secondary veins 10–13 on each side of midvein,base auriculate and somewhat amplexicaul, margin entire,apex obtuse. Spikes terminal or axillary, 1.5–5 cm, interrupted,with 2–5 flowers per node; peduncle 1.5–3.5 cm, glabrous;rachis puberulent; bracts triangular, ca. 1 mm, subglabrous;bracteoles similar to bracts. Calyx ca. 2 mm, subglabrous, 5-lobed almost to base; lobes narrowly triangular. Corolla white,6.5–7 mm, outside puberulent; lower lip 3-lobed, lobes ca. 1mm and apex rounded; upper lip ovate-triangular, ca. 3 mm,erect, apex emarginate. Staminal filaments ca. 2 mm, glabrous;anther thecae ca. 1 mm, unequal, superposed, lower one spurredat base. Ovary glabrous; style subglabrous. Capsule not seen.Fl. May–Aug.● Forests on limestone hills. Guangxi.31. Justicia panduriformis Benoist, Notul. Syst. (Paris) 5: 116.1936.琴叶爵床 qin ye jue chuangMananthes panduriformis (Benoist) C. Y. Wu & C. C. Hu.Shrubs to 2 m tall. Stems 4-angled, sulcate, glabrous.Leaves sessile; leaf blade obovate, 15–26 × 7.5–15 cm, glabrous,base attenuate to rounded, margin entire or slightlyundulate, apex shortly acuminate. Spikes terminal, elongated;rachis densely pubescent, with 2 or 3 flowers clustered pernode; bracts lanceolate-linear, 2–3 mm, apex acute; bracteolessimilar to bracts. Calyx ca. 3 mm, outside puberulent, 5-lobedalmost to base; lobes linear, apex acute. Corolla white, ca. 8mm; tube as long as limb; lower lip 3-lobed; upper lip emarginate.Staminal filaments ca. 4 mm, glabrous; anther thecaeunequal, superposed, lower one spurred at base. Ovary pubescent;style ca. 6 mm, basal portion slightly pubescent. Capsulenot seen. Fl. May–Sep.Forests on limestone hills. Guangxi, Yunnan [Vietnam].32. Justicia lianshanica (H. S. Lo) H. S. Lo, Guihaia 17: 50.1997.广东爵床 guang dong jue chuang

ACANTHACEAE 459Mananthes lianshanica H. S. Lo, Bull. Bot. Res., Harbin1(4): 105. 1981.Herbs to 30 cm tall. Stems 4-angled, basally prostrate androoting at nodes then erect, sulcate, bifariously pubescent toglabrescent. Petiole 1.5–3.5 cm, pubescent; leaf blade ovate,3.5–14 × 2–6 cm, thinly papery, abaxially glabrous or sparselypuberulent along veins, adaxially glabrous, secondary veins 6–8on each side of midvein and meeting near margin, base shallowlycordate to subtruncate, margin slightly undulate or subentire,apex obtuse. Spikes terminal; rachis densely pubescent,usually 1-flowered per node; bracts ovate-lanceolate, 2–3 mm,abaxially pubescent; bracteoles similar to bracts. Calyx ca. 4mm, outside gland-tipped pubescent, 5-lobed almost to base;lobes lanceolate, ca. 3.5 × 0.7 mm. Corolla yellow with purplishspots on lower lip, ca. 8 mm, outside gland-tipped pubescent;tube slightly widened apically; lower lip ca. 3.5 mm, broaderthan upper lip, patent, 3-lobed, lobes ovate and apex rounded;upper lip triangular, ca. 2.2 mm, erect, apex emarginate. Staminalfilaments ca. 3 mm, glabrous; anther thecae ovoid, superposed,lower one spurred at base. Ovary hirsute; style ca. 9 mm,subglabrous. Capsule (immature) clavate, hirsute. Fl. May–Jul.● Forests on rocks; 300–800 m. Guangdong, Guangxi.33. Justicia cardiophylla D. Fang & H. S. Lo, Guihaia 17: 57.1997.心叶爵床 xin ye jue chuangMananthes cardiophylla (D. Fang & H. S. Lo) C. Y. Wu &C. C. Hu.Herbs 20–50 cm tall. Stems terete, glabrous. Petiole 3.5–11 mm, glabrous or somewhat gland-tipped puberulent; leafblade cordate to rarely broadly ovate, 3.5–12 × 2–9 cm, papery,glabrous, secondary veins 5–7 on each side of midvein, basecordate to ± truncate, margin subentire, apex acute to rarelyshortly acuminate. Spikes terminal, 3.5–11 cm, unbranched orsometimes 1-branched; peduncle 7.5–20.5 cm, somewhat bifariouslypubescent; rachis densely pubescent, (1 or)2–5-floweredper node; bracts triangular, 1.5–2 × ca. 1 mm, abaxiallypuberulent, margin ciliate; bracteoles subtriangular, ca. 1.5 mm,abaxially puberulent. Calyx ca. 2 mm, outside puberulent, 5-lobed almost to base; lobes linear-triangular, ca. 1.5 × 0.6 mm.Corolla greenish yellow, ca. 7 mm, outside pubescent; lower lippatent, 3-lobed, lobes ca. 1 mm and apex rounded; upper lip triangular,ca. 3.5 mm, erect, apex emarginate. Staminal filamentsca. 2 mm, glabrous; anther thecae linear, ca. 1 mm, unequal,superposed, lower one spurred at base. Ovary pubescent; styleca. 3.5 mm, basal portion sparsely puberulent. Capsule clavate,ca. 1.5 cm, pubescent, 4-seeded. Seeds ovate in outline, ca. 1.3× 1.3 mm, verrucose. Fl. Mar–Jun.Rocky limestone hills; 400–600 m. Guangxi [Vietnam].34. Justicia austroguangxiensis H. S. Lo & D. Fang, Guihaia17: 54. 1997.桂南爵床 gui nan jue chuangJusticia austroguangxiensis f. albinervia D. Fang & H. S.Lo; Mananthes austroguangxiensis (H. S. Lo & D. Fang) C. Y.Wu & C. C. Hu; M. austroguangxiensis f. albinervia (D. Fang& H. S. Lo) C. Y. Wu & C. C. Hu.Herbs 15–30 cm tall. Stems 0.5–5 cm, sometimes woody.Leaves in a basal rosette; petiole 1–3 cm, glabrous; leaf bladeobovate, obovate-elliptic, or rarely elliptic, (6–)10–15 × (2.5–)4–7(–9) cm, papery, glabrous, secondary veins 7 or 8 on eachside of midvein, base attenuate to narrowly cuneate and decurrentonto petiole, margin entire, apex acute to rarely obtuse.Spikes terminal or axillary in apical leaf axils, 5–35 cm, interrupted,usually branched; rachis puberulent, usually (1 or)2–9-flowered per node; bracts triangular to rarely narrowly triangular,1–2 mm, abaxially puberulent; bracteoles similar tobracts. Calyx 1.5–2 mm, outside puberulent, 5-lobed almost tobase; lobes subulate. Corolla light yellowish green, 8–9 mm,outside puberulent; lower lip 3-lobed, lobes 4–4.5 mm; upperlip triangular, 3.5–4 mm, erect, apex emarginate. Staminal filamentsca. 3 mm, glabrous; anther thecae superposed, lower onespurred at base. Ovary puberulent; style 4.5–5 mm, basal portionpuberulent. Capsule clavate, 1.3–1.5 cm. Seeds ovate inoutline, ca. 0.8 × 0.8 mm, tuberculate. Fl. Mar–Jul, fr. Aug.● Dense forests on rocks; 300–500 m. Guangxi.35. Justicia leptostachya Hemsley, J. Linn. Soc., Bot. 26: 245.1890.南岭爵床 nan ling jue chuangMananthes leptostachya (Hemsley) H. S. Lo.Herbs 40–60 cm tall. Stems 4-angled, sulcate, bifariouslypubescent. Petiole 1–2 cm; leaf blade ovate-lanceolate, 10.5–12.5(–18) × 5–6.5(–8) cm, papery, abaxially strigose alongveins, adaxially sparsely strigose, secondary veins 5–9 on eachside of midvein, base broadly cuneate, margin entire or slightlyundulate, apex acuminate. Spikes terminal, 3–6 cm, branched orrarely unbranched; rachis hispid, with several flowers per node;bracts linear, ca. 2 × 0.8 mm, abaxially puberulent; bracteolessimilar to bracts. Calyx ca. 3 mm, outside puberulent, 5-lobedalmost to base; lobes narrowly lanceolate to linear, apex acute.Corolla ca. 6 mm, outside pubescent; lower lip patent, 3-lobed,lobes ovate and apex rounded; upper lip oblong, apex emarginate.Staminal filaments ca. 3 mm, glabrous; anther thecaesuperposed, lower one spurred at base. Ovary glabrous; stylefiliform, glabrous. Capsule clavate, ca. 1.2 cm, puberulent,4-seeded. Seeds dark brown, ovate in outline, ca. 1 × 1 mm,slightly muricate. Fl. Mar–Jun, fr. Jul–Aug.● On rocks. Guangdong, Guangxi, Hunan.Specimens misidentified as Mananthes vasculosa (Nees) Bremekampby C. C. Hu (FRPS 70: 298. 2002) belong here.36. Justicia microdonta W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 10: 183. 1918.小齿爵床 xiao chi jue chuangHu.Mananthes microdonta (W. W. Smith) C. Y. Wu & C. C.Shrubs to 2 m tall, much branched. Young braches reddishwhen dry, 4-angled, sparsely pilose. Petiole 1–2 cm; leaf blade

460ACANTHACEAEovate, (2–)3–5.5 × 1.5–2.5 cm, variable in size, papery, abaxiallylight green and pilose along midvein, adaxially green andpilose especially along veins, base broadly cuneate and decurrentonto petiole, margin entire, apex shortly acuminate to acute.Spikes terminal, 5–7(–10) cm, unbranched, nodes distant, with2–4 flowers per node; peduncle short; rachis densely pilose;bracts linear-lanceolate, 3–5 cm; bracteoles similar to bracts.Calyx ca. 7 mm, 5-lobed almost to base; lobes linear-lanceolate,5–6 mm, pilose along midvein. Corolla white, 1–1.3 cm, outsidesomewhat pubescent to glabrescent; tube short; limb longerthan tube; lower lip 4–5 mm, 3-lobed, lobes pilose inside andciliate on margin; upper lip 2-lobed. Staminal filaments pilose;anther thecae parallel, superposed, lower one with a white spurat base. Ovary glabrous; style densely pilose. Capsule ca. 2 cm,4-seeded. Seeds reddish brown, circular in outline, 3–4 mm indiam., rugose-verruculose. Fl. Jun–Aug.● Thickets; 800–1200 m. Sichuan, Yunnan.37. Justicia damingensis (H. S. Lo) H. S. Lo, Guihaia 17: 51.1997.大明爵床 da ming jue chuangMananthes damingensis H. S. Lo, Bull. Bot. Res., Harbin1(4): 106. 1981.Herbs 15–30 cm tall. Stems subterete, basally prostrateand rooting at nodes then erect, sulcate, bifariously pubescent.Petiole 0.5–1.5 cm, pubescent; leaf blade ovate, 4–9 × 2.5–4.5cm, thinly papery, abaxially pubescent along veins, adaxiallyglabrous, secondary veins ca. 5 on each side of midvein, basenarrowly cuneate, margin shallowly undulate, apex acute. Racemesterminal, spiciform, 17–18 cm, branchless or trifurcatebranched, with 3 flowers per node; peduncle 5.5–10 cm, pubescent;bracts subulate, 1.5–2 mm, abaxially pubescent; bracteolessimilar to bracts. Pedicel ca. 1 mm, pubescent. Calyx ca. 6mm, 5-lobed almost to base; lobes narrowly triangular, ca. 5 ×0.6 mm, margin ciliate, apex acuminate. Corolla yellow, ca. 1cm, outside puberulent; lower lip ca. 3.2 mm, patent, 3-lobed,lobes ovate and apex rounded; upper lip subdeltoid, ca. 2.5 mm.Staminal filaments ca. 3 mm, glabrous; anther thecae ellipsoid,superposed, lower one spurred at base. Ovary glabrous; styleca. 6 mm, glabrous. Capsule not seen. Fl. May–Jun.● Thickets; 300–600 m. Guangxi.38. Justicia amblyosepala D. Fang & H. S. Lo, Guihaia 17:55. 1997.钝萼爵床 dun e jue chuangMananthes amblyosepala (D. Fang & H. S. Lo) C. Y. Wu& C. C. Hu.Shrubs 30–50 cm tall. Stems terete, glabrous. Petiole 0.8–3cm, glabrous; leaf blade ovate to elliptic, 9–17 × 4.5–8 cm, papery,glabrous, secondary veins 5–7 on each side of midvein,base attenuate to rounded, margin subentire, apex acute. Spikesterminal or axillary, 2–23 cm, interrupted, simple or branched;peduncle 1.5–5 cm; rachis 4-angled, sulcate, bifariously puberulent,with 1 flower per node; bracts triangular to rarely narrowlytriangular, 1.5–3 mm, subglabrous; bracteoles narrowlytriangular, ca. 2 mm, margin ciliolate. Calyx ca. 5 mm, 5-lobed;lobes elliptic-lanceolate, both surfaces usually puberulent tosubglabrous, apex obtuse to rounded. Corolla light yellow, ca.1.3 cm, outside retrorsely puberulent; lower lip 3-lobed, lobesca. 1.5 mm and apex rounded; upper lip ovate-triangular, ca. 5mm, erect, apex emarginate. Staminal filaments ca. 3 mm, glabrous;anther thecae ellipsoid, ca. 1.5 mm, unequal, superposed,lower one spurred at base. Ovary pubescent at tip; style 8–11mm, sparsely puberulent. Capsule clavate, ca. 1.5 cm, pubescent.Seeds ovate in outline, ca. 1 × 1 mm, verrucose. Fl. Mar–May, fr. Aug.● Forests on limestone slopes. Guangxi.39. Justicia acutangula H. S. Lo & D. Fang, Guihaia 17: 56.1997.棱茎爵床 leng jing jue chuangMananthes acutangula (H. S. Lo & D. Fang) C. Y. Wu &C. C. Hu.Herbs 50–100 cm tall. Stems sharply 4-angled and slightlywinged, sulcate, glabrous. Petiole ca. 1.3 mm, pubescent; leafblade elliptic to rarely broadly elliptic, 8.5–28 × 5–12.5 cm,glabrous, secondary veins 13–15 on each side of midvein, baserounded to attenuate and decurrent onto petiole, margin subentire,apex acute. Spikes terminal or axillary in upper leaf axils,3.5–14 cm, interrupted, unbranched or branched, with 1 flowerper node; bracts triangular, 2–3 mm, glabrous, margin ciliate;bracteoles ca. 2.5 mm, glabrous, margin ciliate. Calyx ca. 5mm, outside densely puberulent, 5-lobed almost to base; lobeslanceolate, apex acute. Corolla light greenish yellow with purplishdots on lower lip, 2–2.2 cm, outside pubescent; limb aslong as tube; lower lip as long as upper lip, 3-lobed, lobesovate, ca. 2 mm, and apex rounded; upper lip ca. 9 mm, apexemarginate. Staminal filaments ca. 7.5 mm, glabrous; antherthecae ca. 2.5 mm, superposed, lower one minutely spurred atbase. Ovary glabrous; style ca. 2 cm, basal portion pubescent.Capsule clavate, ca. 2 cm. Seeds ovate in outline, ca. 1.2 × 1.3mm. Fl. Nov–Jan, fr. Apr–May.● Forests on limestone hills; 500–700 m. Guangxi, Guizhou.40. Justicia patentiflora Hemsley, Hooker’s Icon. Pl. 28: t.2792. 1905.野靛棵 ye dian keMananthes patentiflora (Hemsley) Bremekamp.Herbs to 2 m tall, perennial. Stems subterete, glabrous.Petiole 2–6 cm; leaf blade ovate to oblong-lanceolate, 16–26 ×7.5–9.5 cm, membranous, glabrous, secondary veins 8 or 9 eachside, blade basally attenuate to a narrowly cuneate base, apexacuminate. Spikes terminal or axillary in upper leaf axils, 5–12cm, unbranched or bifurcately branched; rachis 4-angled, bifariouslyyellowish pubescent, with 1 flower per node; bracts scalelike,triangular-lanceolate, ca. 3 mm, abaxially puberulent;bracteoles similar to bracts. Calyx ca. 1.2 cm, puberulent, 5-lobed almost to base; lobes linear, apex acute. Corolla purplishred with purplish spots, 1.5–2 cm; tube 1–1.5 cm, abruptlyrecurved above base, outside puberulent; lower lip 3-lobed,lobes circular; upper lip subcircular, shorter than lower lip, apexemarginate. Staminal filaments ca. 5 mm, glabrous; anther

ACANTHACEAE 461thecae superposed, lower one spurred at base. Ovary glabrous;style ca. 7 mm, basal portion sparsely puberulent. Capsulenarrowly obovoid, ca. 1.5 cm. Seeds light yellow, circular inoutline, ca. 1.2 × 1.2 mm, verrucose. Fl. Nov–Jan, fr. Apr–May.Forests, streamsides, limestone hills; 500–1800(–2400) m. Yunnan[Vietnam].“Mananthes tubiflora” (Y. M. Shui & W. H. Chen, Seed Pl. KarstReg. China 1: 182. 2006) belongs here but was not validly published.41. Justicia poilanei Benoist, Notul. Syst. (Paris) 5: 125. 1936.毛萼爵床 mao e jue chuangShrubs to 2 m tall. Stems ± 4-angled, sulcate, glabrous.Petiole 1–6 cm, flat, sulcate, margin ciliate; leaf blade ovate toovate-lanceolate, 10–20 × 4–8 cm, glabrous, secondary veins6–10 on each side of midvein, base attenuate and decurrent ontopetiole, margin entire or undulate, apex acute. Spikes terminalor rarely axillary, 7–15 cm, usually trifurcately branched orrarely unbranched; peduncle 1–3 cm; rachis 4-angled, glabrous,with 1 flower per node; bracts triangular, 2–2.5 × ca. 2 mm,both surfaces glabrous, margin ciliate; bracteoles similar tobracts. Calyx 3–4 mm, both surfaces puberulent especially distallyon inner surface, 5-lobed almost to base; lobes lanceolate,margin ciliate, apex acute. Corolla ca. 1 cm, outside pubescent;tube as long as limb; lower lip 3-lobed, lobes subequal, ovate,and apex rounded; upper lip subtriangular, erect, apex emarginate.Staminal filaments ca. 4 mm, glabrous; anther thecaeunequal, superposed, lower one spurred at base. Ovary glabrous;style ca. 8 mm, glabrous. Capsule not seen. Fl. Dec–Feb.Limestone hills; 1000–2400 m. Yunnan [Vietnam].42. Justicia kampotiana Benoist, Notul. Syst. (Paris) 5: 118.1936.那坡爵床 na po jue chuangMananthes kampotiana (Benoist) C. Y. Wu & C. C. Hu.Shrubs to 1 m tall. Stems ± 4-angled, sulcate, glabrous.Petiole 3–6 mm, glabrous; leaf blade lanceolate, 9–16 × 4.3–5cm, glabrous, secondary veins ca. 8 on each side of midvein,base cuneate, margin entire, apex shortly acuminate. Spikesaxillary or terminal; rachis 4-angled, unbranched or 2-furcatelybranched, subglabrous, 1-flowered per node; bracts narrowlytriangular, ca. 1.5 mm, abaxially pubescent; bracteolessimilar to bracts. Calyx ca. 3 mm, outside pubescent, inside glabrous,5-lobed almost to base; lobes linear, subequal, apexacute. Corolla ca. 1 cm, outside pubescent; tube slightly longerthan limb; lower lip 3-lobed, lobes subequal, ovate, and apexrounded; upper lip subtriangular, erect, apex emarginate. Staminalfilaments ca. 4 mm, glabrous; anther thecae unequal, superposed,lower one spurred at base. Ovary and style glabrous.Capsule glabrous. Fl. Apr–May.On rocks in forests; 500–600 m. Guangxi [Cambodia].43. Justicia caudatifolia (H. S. Lo & D. Fang) Z. P. Hao, Y. F.Deng & T. F. Daniel, J. Trop. Subtrop. Bot. 18: 486. 2010.尾叶爵床 wei ye jue chuangLeptostachya caudatifolia H. S. Lo & D. Fang, Guihaia17: 45. 1997.Herbs to 1 m tall. Stems 4-angled, basally prostrate androoting at nodes, sulcate, glabrous. Petiole 2–15 mm, glabrous;leaf blade elliptic to obovate-elliptic, 3–10 × 1.2–4.2 cm, apicalhalf broadest, papery, glabrous, secondary veins 5 or 6 on eachside of midvein, base cuneate, margin subentire, apex caudate.Spikes axillary, 1–3 cm, interrupted; peduncle 0.5–1.5 cm; rachissulcate, puberulent, 1-flowered per node; bracts linear-triangular,3–4 mm, abaxially puberulent, margin ciliate; bracteolesovate, 2.5–3 mm, margin ciliate. Calyx ca. 4 mm, 5-lobedalmost to base; lobes narrowly triangular, outside glabrescent,inside glabrous. Corolla white, ca. 1.1 cm, outside slightly retrorselypubescent; tube as long as limb; lower lip 3-lobed,lobes ca. 2.5 mm and apex rounded; upper lip ca. 5 mm, apexemarginate. Staminal filaments ca. 3 mm, glabrous; antherthecae superposed, lower one minutely spurred at base. Ovaryglabrous; style ca. 5 mm, glabrous. Capsule not seen. Fl. Oct.● On rocks in forests on limestone hills. Guangxi.25. RHINACANTHUS Nees in Wallich, Pl. Asiat. Rar. 3: 76, 108. 1832.灵枝草属 ling zhi cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielShrubs, subshrubs, or herbs, with cystoliths. Leaves opposite, petiolate or subsessile; leaf blade margin entire or slightly undulate.Inflorescences axillary or terminal, spikes or racemes, sometimes branched into panicles; bracts and bracteoles shorter thancalyx. Calyx deeply 5-lobed; lobes subequal. Corolla white or greenish [or purple]; tube cylindric, elongate, narrow; limb 2-lipped;lower lip 3-lobed, lobes ascending cochlear in bud; upper lip entire or 2-cleft, inside rugulose. Stamens 2, exserted from mouth ofcorolla; anthers 2-thecous; thecae unequally inserted to superposed, lower theca with or without a basal appendage; staminodesabsent. Capsule with a solid stalk, clavate, 4-seeded; retinacula present. Seeds discoid, variously ornamented, lacking trichomes.About 25 species: tropical and subtropical regions of Africa and Asia; two species (one endemic) in China.Rhinacanthus calcaratus (Wallich) Nees was reported from Yunnan by C. C. Hu (FRPS 70: 268. 2002) based on a misidentification of Pseuderanthemumpolyanthum.1a. Leaf blade narrowly obovate to oblanceolate, 20–24 × 6–8 cm, abaxially glabrous, secondary veins 8–10 oneach side of midvein; corolla tube ca. 4 cm, lobes of lower lip 2–2.5 cm .................................................................. 1. R. beesianus1b. Leaf blade elliptic, ovate-elliptic, or rarely lanceolate, 2–7(–11) × 0.8–3 cm, abaxially pubescent, secondaryveins 5 or 6 on each side of midvein; corolla tube 1.5–1.8 cm, lobes of lower lip 2–4 mm ......................................... 2. R. nasutus

462ACANTHACEAE1. Rhinacanthus beesianus Diels, Notes Roy. Bot. Gard. Edinburgh5: 164. 1912.滇灵枝草 dian ling zhi caoShrubs to 1.5 m tall. Stems 4-angled, subglabrous. Petiole0–4 mm; leaf blade narrowly obovate to oblanceolate, 20–24 ×6–8 cm, abaxially glabrous, adaxially slightly pubescent, secondaryveins 8–10 on each side of midvein, base attenuate anddecurrent onto petiole, margin slightly undulate to subentire,apex acute. Panicles terminal, 12–15 × 8–9 cm; bracts leaflike,lanceolate, 0.8–1.2 × 0.3–0.4 cm. Flowers pedicellate. Calyxlobes linear, ca. 10 × 1.5 mm, pubescent, 1-veined, apex acute.Corolla white, fragrant, outside gland-tipped pubescent; tube ca.4 cm; lower lip lobes elliptic, 2–2.5 cm; upper lip ca. 2 cm,apex 2-cleft. Stamens ca. 1 cm. Fl. Aug–Sep.● Open dry situations on mountain slopes; 2100–2400 m. Yunnan.2. Rhinacanthus nasutus (Linnaeus) Kurz, J. Asiat. Soc. Bengal,Pt. 2, Nat. Hist. 39: 79. 1870.灵枝草 ling zhi caoJusticia nasuta Linnaeus, Sp. Pl. 1: 16. 1753; Rhinacanthuscommunis Nees.Subshrubs or perennial herbs, to 1.5 m tall. Stems stout, ±4-angled, faintly striate, densely pubescent when young thenglabrescent. Petiole 0.5–1.5 cm; leaf blade elliptic, ovate-elliptic,or rarely lanceolate, 2–7(–11) × 0.8–3 cm, abaxially denselypubescent, adaxially sparsely pubescent to subglabrous, secondaryveins 5 or 6 on each side of midvein, base cuneate, marginentire or slightly undulate, apex shortly acuminate to acute.Panicles terminal or axillary, to 50 cm; rachis densely pubescent;bracts lanceolate, to 2 × 0.5 mm; bracteoles ca. 1 mm.Flowers sessile to subsessile. Calyx ca. 5 mm, both surfacespubescent; lobes lanceolate, ca. 4 × 0.7 mm. Corolla greenishwhite, 2.1–2.7 cm, outside pubescent with gland-tipped andnon-glandular trichomes; tube 1.5–1.8 cm; lower lip 0.75–1.2cm, lobes 2–4 mm and subequal; upper lip linear-lanceolate, 6–7.5 mm, erect. Staminal filaments glabrous. Style sparsely pubescent.Capsule ca. 2 × 0.3 cm, pubescent with gland-tippedtrichomes. Seeds ca. 2.5 × 2.2 mm, papillose. Fl. Oct–Dec. 2n =30.Thickets, forests, cultivated around villages; below 700 m. Guangdong,Hainan, Yunnan [Cambodia, India, Indonesia, Laos, Malaysia,Myanmar, Philippines, Sri Lanka, Thailand, Vietnam; Madagascar].26. DICLIPTERA Jussieu, Ann. Mus. Natl. Hist. Nat. 9: 267. 1807, nom. cons.狗肝菜属 gou gan cai shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielDiapedium K. D. Koenig.Herbs, diffuse, annual, biennial, or perennial [or sometimes shrubs], with cystoliths; young stems ± 6-angled in cross-section.Leaves petiolate; leaf blade margin entire or subsinuate. Inflorescences of cymes in leaf axils or in terminal spikelike thyrses orpanicles of thyrses. Flowers subtended by an involucre of several pairs of bracteoles (= cymule); outermost pair equal or unequal insize, usually concealing inner pair(s), calyx, and base of corolla. Calyx deeply 5-lobed; lobes narrow, equal, apex acute. Corolla pink,resupinate 180° by torsion of tube; tube slender, widened apically, limb 2-lipped; lower lip (in upper position) 3-lobed, lobesascending cochlear in bud; upper lip (in lower position) entire or emarginate. Stamens 2, exserted from mouth of corolla; anthers 2-thecous; thecae unequally inserted to superposed, base muticous; staminodes absent. Stigma 2-cleft. Capsule substipitate or stipitate,2–4-seeded; retinacula present; placenta with retinacula separating elastically and rising from inner capsule wall at dehiscence. Seedslenticular, papillate, lacking trichomes.About 100 species: tropical and temperate regions worldwide; four species (two endemic) in China.1a. Outer bracteoles oblong-lanceolate to oblanceolate, greater than 2 × as long as wide.2a. Outer bracteoles oblong-lanceolate, 5–7 mm, sparsely covered with white or grayish trichomes orglabrescent; corolla ca. 0.5 cm ........................................................................................................................ 1. D. bupleuroides2b. Outer bracteoles oblanceolate, 10–14 mm, densely covered with yellow trichomes; corolla ca. 1.5 cm ................. 2. D. induta1b. Outer bracteoles elliptic, subcircular, or obovate, less than 2 × as long as wide.3a. Corolla pale purple, 1–1.2 cm; outer bracteoles 0.5–1.3 × 0.3–0.8 cm ................................................................ 3. D. chinensis3b. Corolla pinkish red, ca. 1.8 cm; outer bracteoles 1.5–2 × 1–1.5 cm ....................................................................... 4. D. elegans1. Dicliptera bupleuroides Nees in Wallich, Pl. Asiat. Rar. 3:111. 1832.印度狗肝菜 yin du gou gan caiDicliptera roxburghiana Nees var. bupleuroides (Nees) C.B. Clarke.Herbs to 50 cm tall. Stems sulcate, pubescent. Petiole 5–10mm; leaf blade ovate, 3.5–7 × 1.8–2.8 cm, glabrous or with 2-celled trichomes, secondary veins 4 or 5 on each side of midvein,base cuneate, margin subentire, apex acuminate. Inflorescencesaxillary and subsessile or sometimes terminal and shortlypedunculate, cymes, many flowered; peduncle 3–5 mm; bractslanceolate, 2–3 mm; outer bracteoles green, oblong-lanceolate,5–7 × ca. 2 mm, abaxially pubescent, base cuneate, marginentire and ciliate, apex mucronate; inner bracteoles green withyellowish margin, lanceolate, ca. 4 × 1.5 mm. Calyx ca. 3 mm;lobes subulate, pubescent. Corolla ca. 5 mm, outside pubescent;lip in lower position orbicular, ca. 1.5 × 1.5 mm; lip in upperposition oblong, ca. 2 × 1 mm, 3-lobed, lobes ovate and ca. 0.3× 0.3 mm. Staminal filaments ca. 2 mm; anther thecae spheri-

ACANTHACEAE 463cal. Ovary pilose at tip; style ca. 3 mm. Capsule ca. 3 mm, piloseat tip. Seeds ovate in outline, ca. 1 mm, papillate. Fl. Nov–May, fr. Dec–Jun. 2n = 26.Roadsides, forests, shady places; 800–1900 m. Guizhou, Sichuan,Yunnan [Cambodia, India, Laos, Myanmar, Thailand, Vietnam].2. Dicliptera induta W. W. Smith, Notes Roy. Bot. Gard. Edinburgh10: 175. 1918.毛狗肝菜 mao gou gan caiHerbs to 60 cm tall. Stems sulcate, densely yellow hirsuteespecially along sulcae. Petiole 5–12 mm, yellow tomentose;leaf blade ovate, 3–4 × 1.5–2.5 cm, both surfaces yellow tomentoseespecially along veins, secondary veins 3–5 on eachside of midvein, base broadly cuneate to subrounded, marginentire, apex acute. Inflorescences axillary, each of 1 or 2 cymes;peduncle 2–3 mm, yellow hirsute; bracts linear-subulate, ca. 5mm, tomentose; outer bracteoles dark green, oblanceolate, 10–14 × 2–4 mm, unequal, rigid, both surfaces tomentose especiallytoward base, base cuneate, apex acute and shortly mucronate;inner bracteoles linear-lanceolate, 8–10 mm, pilose, apexacuminate. Pedicel ca. 1 mm. Calyx ca. 5 mm; lobes linear,pilose, apex acute. Corolla pink, ca. 1.5 cm, pilose; tube basallyslender, white pilose; lip in lower position ca. 4 mm, 2-lobed;lip in upper position ca. 3 mm, 3-lobed, lobes ovate. Staminalfilaments ca. 6 mm, glabrous; anther thecae spherical, ca. 1mm. Ovary minutely pilose; style ca. 1.2 cm, minutely pilose.Capsule ca. 4 mm, pilose. Seeds ovate in outline, ca. 2 mm,papillate. Fl. Aug–Sep, fr. Oct–Nov.● Wastelands; 400–700 m. Yunnan.3. Dicliptera chinensis (Linnaeus) Jussieu, Ann. Mus. Natl.Hist. Nat. 9: 268. 1807.狗肝菜 gou gan caiJusticia chinensis Linnaeus, Sp. Pl. 1: 16. 1753; Diapediumchinense (Linnaeus) K. D. Koenig & Sims; Dicliptera burmanniNees; D. roxburghiana Nees.Herbs 30–80 cm tall, annual or biennial. Stems ridged,basally diffuse, apically erect, glabrous except for pubescenceon young branches and nodes. Petiole to 0.5–3 cm, puberulent;leaf blade ovate-elliptic, 2–8 × 1.5–4 cm, abaxially sparsely puberulentalong veins, adaxially glabrous, secondary veins 4 or 5on each side of midvein and abaxially slightly prominent, basecuneate to attenuate and decurrent onto petiole, margin entire orslightly sinuate, apex shortly acuminate to acute. Inflorescenceseach of (1–)3 or 4 pedunculate cymes from upper leaf axils, 1–4cm; inflorescence bracts subulate to linear to oblanceolate toobovate, 3–7.5 × 0.3–1 mm; peduncle 2–5 mm; cymules usually3 or 4 per cyme; outer bracteoles elliptic to obovate, 5–13 ×3–8 mm, unequal, abaxially puberulent, 3-veined, margin ciliate,apex acute and mucronate; inner bracteoles linear, ca. 5mm, abaxially hirsute, apex acuminate. Pedicel 0.5–3 mm,puberulent. Calyx ca. 5 mm; lobes linear, equal, margin hirsute,apex acuminate. Corolla pale purple, 1–1.2 cm, outsidepuberulent; tube slender, ca. 6 mm, ca. 1.5 mm wide; limb 2-lipped; lip in lower position ovate, ca. 5 × 3 mm, apex entire;lip in upper position oblong, ca. 4 × 1.5 mm, shallowly 3-lobed.Stamens inserted near apex of corolla tube; filaments ca. 4 mm,glabrous; thecae spherical, ca. 1.2 mm in diam., parallel. Ovarypilose at tip; style ca. 1.4 cm. Capsule broadly ellipsoid, ca. 6mm, puberulent, 4-seeded, apex apiculate. Seeds circular in outline,ca. 2 mm in diam., covered with apically barbed papillae.Fl. Sep–Jan, fr. Nov–Feb. 2n = 26.Streamsides, trailsides; below 1800 m. Fujian, Guangdong, Guangxi,Guizhou, Hainan, Sichuan, Taiwan, Yunnan [Bangladesh, India,Vietnam].This species is used medicinally.4. Dicliptera elegans W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 10: 174. 1918.优雅狗肝菜 you ya gou gan caiDicliptera mairei Benoist.Herbs 60–120 cm tall. Stems erect, branched, slender, sulcate,sparsely pubescent. Petiole 0.5–1.5 cm, sulcate, pubescent;leaf blade ovate to ovate-lanceolate, 4–8 × 2–5 cm, abaxiallypale gray and sparsely pubescent, adaxially green, densely coveredwith cystoliths, and densely brownish pilose on midveinsbut otherwise sparsely pubescent with few large-celled whitetrichomes, secondary veins 4–6 on each side of midvein, basebroadly cuneate and decurrent onto petiole, margin entire orsinuate, apex acuminate. Inflorescences axillary, each of 2 or3 cymes; peduncle to 7 cm, white tomentose; inflorescencebracts linear, 7–10 × 1–2 mm, puberulent; cymules 3–6 percyme; outer bracteoles subcircular to obovate, 1.5–2 × 1–1.5cm, unequal, abaxially with cystoliths and pilose, palmately3- or 5-veined, base rounded, apex mucronate; inner bracteoleslinear-lanceolate, 5–7 × ca. 1 mm, abaxially puberulent, apexacuminate. Pedicel 0.5–2 mm. Calyx 6–7 mm; lobes linear-lanceolate,outside puberulent, apex acuminate. Corolla pinkishred, ca. 1.8 cm, outside pubescent; tube slender; limb 2-lipped.Staminal filaments ca. 5 mm, glabrous; anther thecae spherical,ca. 1 mm, inserted at slightly different levels. Ovary slightly pilose.Capsule ca. 6 mm, light yellow pilose. Seeds light brown,circular in outline, ca. 1.5 × 1.5 mm, compressed, yellowishpapillate. Fl. Aug–Oct, fr. Oct–Dec.● Forest margins, thickets, roadsides; 600–2000 m. Sichuan, Yunnan.Dicliptera mairei was incorrectly reduced to D. induta by C. C.Hu (FRPS 70: 238. 2002).27. PERISTROPHE Nees in Wallich, Pl. Asiat. Rar. 3: 77, 112. 1832.观音草属 guan yin cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs or shrubs, with cystoliths. Leaf blade margin entire or slightly toothed. Inflorescences in axillary or terminal cymes, these

464ACANTHACEAEsometimes aggregated into a leafy panicle-like complex; cymes consisting of (1 or)2–4(or more) involucres; involucres of usually 1–3 flowers enclosed by 2(or more) pairs of bracteoles that conceal calyx and corolla tube. Outer pair of bracteoles equal to unequal insize and larger than inner pair(s). Calyx small, 5-lobed; lobes equal. Corolla white to pink or purple, usually large, resupinate 180°;tube slender, twisted, apically ± widened into throat; limb 2-lipped; upper lip (in lower position) entire to 2-cleft; lower lip (in upperposition) 3-lobed; lobes ascending cochlear in bud. Stamens 2, inserted at corolla throat, exserted from corolla tube; anthers 2-thecous; thecae unequally inserted or superposed (contiguous or separated by gap), base muticous; staminodes absent. Ovary with 2ovules per locule; stigma slightly enlarged or 2-cleft. Capsule with a solid stalk, 4-seeded; retinacula present; placenta not risingelastically from base of valves. Seeds discoid, smooth to tuberculate, lacking trichomes.About 40 species: tropical and subtropical Africa, Asia, and Madagascar, with one species possibly extending to Australia (see R. M. Barker, J.Adelaide Bot. Gard. 9: 192. 1986); ten species (three endemic) in China.1a. Outer involucral bracteoles abaxially pubescent with gland-tipped trichomes.2a. Corolla ca. 4.5 cm, pubescent with gland-tipped and non-glandular trichomes ................................................ 1. P. lanceolaria2b. Corolla ca. 1 cm, outside pubescent with non-glandular trichomes .................................................................... 2. P. paniculata1b. Outer involucral bracteoles abaxially glabrous or pubescent with non-glandular trichomes.3a. Anther thecae superposed (i.e., one entirely above other and with a gap between them).4a. Corolla ca. 1 cm; outer involucral bracteoles subulate to lanceolate to linear, less than 3 mm wide;capsule pubescent with non-glandular trichomes .......................................................................................... 2. P. paniculata4b. Corolla 3–3.4 cm; outer involucral bracteoles ovate to elliptic, more than 5 mm wide; capsulepubescent with gland-tipped and non-glandular trichomes ............................................................................ 10. P. japonica3b. Anther thecae inserted at same level or at different levels but overlapping.5a. Anther thecae inserted at same level ........................................................................................................... 3. P. yunnanensis5b. Anther thecae inserted at different levels.6a. Outer involucral bracteoles less than 2 × as long as calyx.7a. Leaf blade subglabrous ................................................................................................................... 4. P. tianmuensis7b. Leaf blade densely pubescent ............................................................................................................... 5. P. strigosa6b. Outer involucral bracteoles more than 2 × as long as calyx.8a. Corolla and capsule glabrous ............................................................................................................... 6. P. montana8b. Corolla and capsule pubescent.9a. Leaf blade adaxially puberulent (especially along midvein) or becoming subglabrescent;corolla 3–5.7 cm .............................................................................................................................. 7. P. bivalvis9b. Leaf blade adaxially sparsely pubescent; corolla 2–4 cm.10a. Outer involucral bracteoles narrowly ovate to lanceolate, more than 4 × as longas wide ............................................................................................................................... 8. P. floribunda10b. Outer involucral bracteoles ovate to oblong to elliptic, less than 3 × as long as wide.11a. Outer involucral bracteoles ovate to oblong, abaxially slightly villous, basesubcordate ........................................................................................................................... 9. P. fera11b. Outer involucral bracteoles ovate to elliptic, abaxially subglabrous topuberulent, base cuneate to truncate ........................................................................ 10. P. japonica1. Peristrophe lanceolaria (Roxburgh) Nees in Wallich, Pl.Asiat. Rar. 3: 114. 1832.五指山蓝 wu zhi shan lanJusticia lanceolaria Roxburgh, Fl. Ind. 1: 122. 1820.Herbs to 1 m tall, erect. Branches stout, densely coveredwith white retrorse trichomes or sometimes glabrescent. Petiole(0.5–)1.5–3.5 cm; leaf blade lanceolate to ovate-lanceolate,(6–)10–17 × (1–)3–6 cm, abaxially sparsely pubescent or glabrescent,adaxially glabrescent except for a few trichomes alongmidvein toward base, secondary veins 5–7 on each side of midvein,base narrowly cuneate and slightly oblique, margin entire,apex acuminate. Inflorescences of axillary pedunculate cymesof (1–)3 involucres with cymes sometimes in axils of distalbracts and collectively forming a terminal panicle-like complex5–7 cm; outer involucral bracteoles linear-lanceolate, 10–18 ×1.5–3 mm, gland-tipped pubescent; inner involucral bracteolessimilar to outer ones except smaller. Calyx ca. 9 mm; lobeslinear, ca. 1 mm broad. Corolla pink, ca. 4.5 cm, outside pubes-cent with gland-tipped and non-glandular trichomes; tube baseca. 1.5 mm wide for ca. 2.5 cm; lip in lower position obovate,ca. 2 × 0.8 cm, slightly 2-lobed; lip in upper position oblong,ca. 2 × 0.6 cm, 3-lobed, lobes ovate and ca. 1 × 1 mm. Staminalfilaments ca. 1.8 cm, glabrous; anther thecae oblong, ca. 3 ×0.6 mm, superposed. Capsule 1.4–1.8 cm, pubescent withgland-tipped trichomes. Seeds ca. 3 × 2 mm, smooth. Fl. Dec–Mar, fr. Mar–Jun.Wet places; 500–700 m. Hainan, Yunnan [India, Laos, Myanmar,Thailand, Vietnam].2. Peristrophe paniculata (Forsskål) Brummitt, Kew Bull. 38:451. 1983.双萼观音草 shuang e guan yin caoDianthera paniculata Forsskål, Fl. Aegypt.-Arab. 7. 1775;D. bicalyculata Retzius; Dicliptera paniculata (Forsskål) I.Darbyshire; Justicia bicalyculata (Retzius) Vahl; Peristrophebicalyculata (Retzius) Nees.

ACANTHACEAE 465Herbs to 1.2 m tall, erect, perennial. Stems hirsute, usually4- or 6-angled. Petiole 3–12 mm; leaf blade ovate to lanceolate,(1–)3–4.5 × (0.5–)1.5–2 cm, abaxially scabrid, adaxially slightlyscabrid, base attenuate to broadly cuneate, margin remotely serrulateand slightly ciliate, apex acuminate, acute, or sometimesrounded. Inflorescences of axillary pedunculate cymes with(1–)4(or more) involucres (cymes sometimes branching internally)in axils of leaves or bracts and collectively often forminga terminal leafy panicle-like complex; rachis and inflorescencebranches pubescent; outer involucral bracteoles unequal, abaxiallypubescent with gland-tipped (sometimes absent) and nonglandulartrichomes, larger outer involucral bracteole ± linear,ca. 10 × 2 mm, 1-veined, and apex acute and mucronate, shorterouter involucral bracteole subulate to lanceolate, ca. 6 × 0.1mm; inner involucral bracteoles narrowly linear to subulate,unequal in length. Calyx lobes lanceolate, ca. 3 mm, outsidesparsely pubescent with non-glandular trichomes, insidesparsely pubescent with sessile gland-tipped trichomes, marginmembranous at base, apex long acuminate. Corolla light to darkpurple, ca. 1 cm, outside pubescent with non-glandular trichomes;tube basally cylindric for ca. 5 mm; lip in lower positionnarrowly elliptic, 5–7 × 1–2 mm, apex emarginate; lip inupper position elliptic, 3–7 × 2–3 mm, 3-lobed. Staminal filamentsca. 5 mm, white hairy; anther thecae ovoid, superposed,not touching. Ovary ovoid, pubescent and glandular; style glabrous;stigma 2-lobed. Capsule 0.9–1.3 cm, pubescent withnon-glandular trichomes. Seeds 2.2–2.5 mm, rough and tuberculate.Fl. Aug–Dec, fr. Oct–Feb.Weedy areas, roadsides; 600–2200 m. Guangxi, Sichuan, Yunnan[Cambodia, India, Indonesia, Malaysia, Myanmar, Nepal, Pakistan,Philippines, Thailand, Vietnam; tropical Africa, Australia].3. Peristrophe yunnanensis W. W. Smith, Notes Roy. Bot.Gard. Edinburgh 10: 187. 1918.滇观音草 dian guan yin caoHerbs 1–2 m tall. Stems sulcate, densely grayish pubescentwhen young, later glabrescent. Petiole 1–10 mm; leaf bladelanceolate to ovate, 2.5–3 × ca. 1 cm, abaxially pubescent,adaxially sparsely puberulent especially on veins or glabrescent,base cuneate, margin entire, apex acuminate. Outer involucralbracteoles linear, ca. 3 mm; inner involucral bracteoles 2.5–6mm. Calyx 5–7 mm; lobes linear, glabrous or subglabrous, apexacute. Corolla pink to violet, ca. 1.8 cm, outside slightly pubescent.Staminal filaments basally pubescent; anther thecae oblong,ca. 2 mm, parallel, inserted at same level, touching eachother. Capsule not seen. Fl. Apr, Aug.● Thickets; 1900–2200 m. Sichuan, Yunnan.4. Peristrophe tianmuensis H. S. Lo, Bull. Bot. Res., Harbin8(1): 4. 1988.天目山蓝 tian mu shan lanHerbs 30–35 cm tall. Stems 6-angled, pubescent. Petiole3–10 mm; leaf blade broadly ovate, 1.5–6 × 1–4 cm, bothsurfaces subglabrous, secondary veins 3–6 on each side of midvein,base broadly cuneate to subrounded, margin entire, apexcuspidate to acute. Cymes terminal or pseudo-axillary; outerinvolucral bracteoles narrowly lanceolate, 3.5–6 mm, pubescent;inner involucral bracteoles similar to outer involucralbracteoles. Calyx ca. 3 mm; lobes linear, pubescent. Corollalight purple, ca. 2 cm; tube slender; lip in lower position oblong-elliptic,ca. 1 cm, apex emarginate; lip in upper positionoblong-spatulate, minutely 3-lobed. Staminal filaments 7.5–8cm, pubescent; anther thecae linear, parallel, inserted at differentlevels, touching each other for 2/3 of length. Ovary pubescent;style ca. 2 cm, glabrous. Capsule 8–9 mm, pubescent.Seeds ca. 1.2 × 1.2 mm. Fl. Aug, fr. Oct.● Thickets. Zhejiang (Tianmu Shan).5. Peristrophe strigosa C. Y. Wu & H. S. Lo, Fl. Hainan. 3:596. 1974.糙叶山蓝 cao ye shan lanHerbs 40–100 cm tall, annual. Stems 6-angled, pubescent.Petiole 1.5–2.5 cm; leaf blade ovate to lanceolate-ovate, 7–13 ×3–5.5 cm, both surfaces densely pubescent, secondary veins 6–8 on each side of midvein, base broadly cuneate, margin entire,apex acuminate to shortly caudate. Cymes terminal or axillary,3(or 5)-branched, with 3–5 involucres; peduncle ca. 4 mm;outer involucral bracteoles ovate, ca. 5 × 2 mm, concave, apexaristate; inner involucral bracteoles subulate, ca. 2.5 mm. Calyxca. 5 mm; lobes linear-lanceolate, apex acuminate. Corolla purple,ca. 4.2 cm, outside pubescent; tube basally cylindric for ca.2.4 cm; lip in lower position elliptic-obovate, ca. 1.6 × 0.6 cm,apex obtuse to retuse; lip in upper position oblong, ca. 1.6 × 0.5cm, slightly narrowed at tip, 3-lobed, lobes suboblong and ca. 1× 1 mm. Staminal filaments ca. 1 cm, pubescent; anther thecaelinear, parallel, inserted at different levels. Ovary oblong, ca. 2mm, pubescent; style ca. 3.5 cm, glabrous. Capsule clavate, ca.1.7 cm, densely strigose-pubescent. Seeds ca. 2.5 × 2 mm,tuberculate-punctate. Fl. Nov–Jan, fr. Jan–Mar.● Dense forests. Hainan.6. Peristrophe montana Nees in Wallich, Pl. Asiat. Rar. 3: 113.1832.岩观音草 yan guan yin caoHerbs to 1 m tall, erect, glabrous. Petiole 1.5–4.5 cm; leafblade narrowly oblong, lanceolate, or rarely ovate, 4–9(–20) ×3–5 cm, both surfaces glabrous, secondary veins 6–8 oneach side of midvein, base rounded, margin entire, apex acute.Cymes terminal or axillary; peduncle 1.5–1.7 cm; outer involucralbracteoles various, broadly ovate to linear-lanceolate, 1–1.8× 0.2–1 cm, pinnately veined, base rounded to truncate, marginentire, apex obtuse; inner cymule bracteoles linear, 3–4 mm.Calyx ca. 5 mm; lobes linear. Corolla pink, 4.5–6 cm, outsideglabrous; lip in lower position elliptic, apex emarginate; lip inupper position oblong, 3-lobed. Filaments pubescent; antherthecae linear. Ovary glabrous; style glabrous. Capsule ca. 2 cm,shiny, glabrous. Seeds ca. 3 × 2.6 mm. Fl. Oct–Dec, fr. Dec–Feb.Wet shady cliffs. Hainan [India, Sri Lanka].7. Peristrophe bivalvis (Linnaeus) Merrill, Interpr. Herb.Amboin. 476. 1917.观音草 guan yin cao

466ACANTHACEAEJusticia bivalvis Linnaeus, Syst. Nat., ed. 10, 2: 850.1759; Hypoestes bodinieri H. Léveillé; J. baphica Sprengel;J. roxburghiana Roemer & Schultes; J. tinctoria Roxburgh;Peristrophe baphica (Sprengel) Bremekamp; P. roxburghiana(Roemer & Schultes) Bremekamp; P. tinctoria (Roxburgh)Nees.Herbs to 1 m tall, perennial. Stems 5- or 6-angled, muchbranched, sulcate, pubescent. Petiole to 3.5 cm; leaf blade ovateto lanceolate-ovate, 3–14.5 × 1.5–5.3 cm, both surfaces puberulent(especially along midvein) or becoming subglabrescent,secondary veins 5 or 6 on each side of midvein, base broadlycuneate to subrounded, margin entire to sinuate, apex acuminateto acute. Cymes axillary or terminal, pedunculate, often consistingof 1–3(or more) pedunculate involucres terminatingbranches; peduncle 0.3–2.3 cm; outer involucral bracteolesbroadly ovate, ovate, or elliptic, unequal in size, puberulent tosubglabrous, base rounded to cuneate, apex acute; larger outerinvolucral bracteole (1.8–)2.3–3.3 × 0.8–1.5 cm; smaller outerinvolucral bracteole 0.71–0.85 × as long as larger one; innercymule bracteoles triangular-subulate, 1–2 mm. Calyx 4.5–5mm, pubescent; lobes lanceolate. Corolla pink, 3.5–5.7 cm, outsidepubescent with non-glandular trichomes; tube ca. 1.5 mmin diam.; lip in lower position broadly ovate-elliptic, 2.2–2.6 ×ca. 1.8 cm, apex emarginate; lip in upper position oblong, 2–2.5× ca. 0.9 cm, 3-lobed. Staminal filaments ca. 1.3 cm, puberulent;anther thecae linear, subequally to unequally inserted(overlapping for most of their length), 3.2–4 mm. Ovary pubescent;style glabrous; stigma 2-cleft. Capsule ca. 1.5 cm, pubescent.Seeds ca. 1.5 × 1.5 mm. Fl. Aug–Mar.Forests; 500–1000 m. Fujian, Guangdong, Guangxi, Guizhou,Hainan, Hunan, Jiangxi, Taiwan, Yunnan [Cambodia, India, Indonesia,Laos, Malaysia, Thailand, Vietnam].8. Peristrophe floribunda (Hemsley) C. Y. Wu & H. S. Lo, Fl.Hainan. 3: 561. 1974.海南山蓝 hai nan shan lanDicliptera crinita (Thunberg) Nees var. floribunda Hemsley,J. Linn. Soc., Bot. 26: 248. 1890.Herbs, erect, perennial. Stems stout, 4-angled, sulcate, pubescent.Petiole 0.5–2 cm; leaf blade ovate to lanceolate, 2–9 ×1.5–3.5 cm, both surfaces pubescent along veins, secondaryveins 4–6 on each side of midvein, base broadly cuneate to subrounded,margin entire, apex acuminate to acute-acuminate.Cymes terminal or axillary; outer involucral bracteoles narrowlyovate to lanceolate, unequal in size, both surfaces pubescent,base cuneate, rounded, or sometimes cordate, margin entireand ciliate, apex acuminate; larger outer involucral bracteoles1.5–2 × 0.3–0.5 cm; smaller outer involucral bracteoles4/5–5/6 size of larger ones; inner involucral bracteoles shorterthan calyx. Calyx ca. 4.5 mm, pubescent; lobes ca. 3.5 mm. Corollalight purple to pink, outside pubescent; tube slender, ca. 1mm in diam.; lip in lower position ovate-elliptic, apex slightlyemarginate; lip in upper position oblong, 3-lobed. Staminalfilaments ca. 1 cm, pubescent; anther thecae linear, parallel,slightly unequally inserted. Ovary hirsute; style basally hirsute.Capsule 1.2–1.3 mm, densely pubescent. Seeds ca. 2.5 mm indiam. Fl. Jun–Sep.Valleys, forests. Fujian, Guangdong, Guizhou, Hainan, Hunan, Sichuan,Yunnan [India].9. Peristrophe fera C. B. Clarke in J. D. Hooker, Fl. Brit. India4: 556. 1885.野山蓝 ye shan lanPeristrophe fera var. intermedia C. B. Clarke; P. jalappifoliaNees.Herbs to 1 m tall, erect. Stems 4-angled, sulcate, pubescent.Petiole 5–10(–20) mm; leaf blade ovate to ovate-lanceolate,5–10 × 2–4.3 cm, both surfaces pubescent, secondary veins3–5 on each side of midvein, base broadly cuneate, margin subentireor apically obscurely dentate, apex acuminate. Cymesterminal or axillary in apical leaf axils, densely pubescent;outer involucral bracteoles ovate to oblong, unequal in size,slightly villous, base subcordate, margin ciliate, apex acuminate;larger outer involucral bracteoles ca. 1.5 × 0.8–1 cm;smaller outer involucral bracteoles 4/5–5/6 size of larger ones;inner involucral bracteoles 2.2–2.5 mm. Calyx 4–5 mm, pubescent;lobes linear, apex acuminate. Corolla 3–4 cm, outsidepubescent; tube ca. 1.5 mm in diam.; lip in lower positionbroadly elliptic, apex emarginate; lip in upper position narrowlyoblong, apex 3-lobed. Staminal filaments ca. 1.5 cm, villous;anther thecae linear, parallel, slightly unequally inserted. Ovarypubescent; style basally hirsute. Capsule ca. 1.4 cm, pubescent.Seeds ca. 1.5 × 1.5 mm. Fl. Jul–Aug.Dense forests. Guizhou, Hainan, Yunnan [India].10. Peristrophe japonica (Thunberg) Bremekamp, Boissiera 7:194. 1943.九头狮子草 jiu tou shi zi caoDianthera japonica Thunberg in Murray, Syst. Veg., ed.14, 64. 1784; Dicliptera buergeriana Miquel; D. crinita (Thunberg)Nees; D. japonica (Thunberg) Makino; D. uraiensisHayata; Justicia crinita Thunberg; Peristrophe chinensis Nees;P. guangxiensis H. S. Lo & D. Fang.Herbs 20–80 cm, perennial. Stems 4-angled, basally prostrateand rooting at nodes, bifariously pubescent. Petiole 0.5–1.5 cm; leaf blade lanceolate to ovate-oblong, (3–)5–13 × (1–)2.5–4.5 cm, both surfaces sparsely pubescent (especially alongmidvein), secondary veins 3–5 on each side of midvein, baseobtuse to cuneate, margin entire or slightly sinuate, apex acuminateto caudate. Inflorescences terminal or axillary pedunculatecymes from axils of distal leaves, cymes consisting of 1–4(ormore) pedunculate involucres; involucres bearing 1–3 flowers;peduncle 0.7–5.5 cm; outer involucral bracteoles ovate to elliptic,subequal to unequal in size, abaxially subglabrous to puberulent,base cuneate to truncate, margin entire, apex acute toobtuse; larger outer involucral bracteoles 1.1–2.5 × 0.5–1.8 cm;smaller outer involucral bracteoles 8–17 × 3–9 mm; inner involucralbracteoles linear, 2–3 mm, abaxially puberulent, apexacuminate. Calyx ca. 4 mm; lobes subulate, pubescent. Corollawhite to pale pink to light purple and with pink specks or darklines on limb, 2–3.4 cm, outside pubescent with non-glandulartrichomes; tube basally ca. 1 mm wide for ca. 1.5 cm; lip inlower position elliptic to subcircular, 8–15 × 3–11 mm, entire orslightly emarginate; lip in upper position oblong, 9–15 × 4–7

ACANTHACEAE 467mm, 3-lobed. Staminal filaments ca. 7 mm, pubescent; antherthecae linear, ca. 1.2 mm, parallel, unequally inserted to superposed.Ovary pilose; style ca. 2 cm; stigma 2-cleft. Capsule0.8–1.2 cm, pubescent with gland-tipped and non-glandular trichomes.Seeds ca. 2 mm in diam., tuberculate. Fl. Jul–Feb, fr.Jul–Oct. 2n = 48.Roadsides, grasslands, streamsides, montane slopes, trailsides,evergreen broad-leaved forests, Machilus-Castanopsis forests; below1500 m. Anhui, Chongqing, Fujian, Guangdong, Guangxi, Guizhou,Hainan, Henan, Hubei, Hunan, Jiangsu, Jiangxi, Sichuan, Taiwan,Yunnan, Zhejiang [Japan].This species is used medicinally.28. HYPOESTES Solander ex R. Brown, Prodr. 474. 1810.枪刀药属 qiang dao yao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielShrubs or perennial herbs, decumbent to erect, with cystoliths. Leaves opposite, petiolate; leaf blade margin entire or dentate.Inflorescences of axillary or terminal bracteate spikes to panicles of spikes or of variously disposed cymes (these sometimesclustered in leaf axils); bracteoles usually 4, in 2 to rarely more pairs, distinct or connate, inner pair sometimes partially adnate toouter pair, forming an involucre enclosing 1 or sometimes more flowers. Calyx small, usually obscured by bracteoles, scarious ormembranous, 5-lobed; lobes equal or subequal in length. Corolla rose-pink, purplish, or whitish, resupinate 180° by torsion of tube;tube subcylindric to expanded distally into a narrow throat; limb 2-lipped; lower lip (in upper position) shallowly 3-lobed; upper lip(in lower position) subentire or shallowly 2-lobed; lobes ascending cochlear in bud. Stamens 2, inserted near apex of corolla tube,exserted from mouth of corolla; anthers 1-thecous; theca muticous. Ovary with 2 ovules per locule; style exserted from mouth ofcorolla; stigma 2-lobed, lobes equal or unequal. Capsule stalked, 2–4-seeded; retinacula present. Seeds smooth or ornamented.About 150 species: Old World; three species in China.1a. Outer bracteoles connate for more than half their length, forming a conspicuous tube; capsule glabrous ................ 1. H. purpurea1b. Outer bracteoles inconspicuously connate only at base if at all, not forming a conspicuous tube; capsulepubescent.2a. Outer bracteoles 2.5–4 mm, lanceolate-subulate to linear ................................................................................ 2. H. cumingiana2b. Outer bracteoles 7–17 mm, elliptic to obovate to oblanceolate ................................................................................ 3. H. triflora1. Hypoestes purpurea (Linnaeus) R. Brown, Prodr. 474. 1810.红丝线 hong si xianJusticia purpurea Linnaeus, Sp. Pl. 1: 16. 1753; Hypoestessinica Miquel; Peristrophe purpurea (Linnaeus) Hochreutiner.Shrubs or perennial herbs, to 1.5 m tall, spreading to erect.Stems ± 4-angled when young, sulcate, slightly puberulent.Petiole 1–3.5 cm, sparsely pubescent; leaf blade ovate toovate-elliptic, 5–13 × 1.5–7 cm, abaxially pale green and glabrous,adaxially green and sparsely pubescent or glabrous exceptfor pubescent along veins and on margin, secondary veins5–8 on each side of midvein, base narrowed and cuneate, marginentire, subundulate, or sinuate-crenate, apex acuminate toacute. Inflorescences of terminal or axillary spikes, 1–3 cm,often arranged in large panicles forming a many-floweredthyrse to 7 cm or of dense cymose clusters in axils of leaves orsubleaflike bracts; outer bracteoles connate for more than halftheir length, forming a 5–8 mm obconic involucre, abaxiallyglabrous or pubescent; inner bracteoles smaller than and partiallyadnate to outer pair. Calyx ca. 5 mm, 5-lobed ± halfway tobase; lobes equal, margin ciliate. Corolla reddish purple to purple,2–2.7 cm, outside puberulent; lip in lower position lanceolate,subentire; lip in upper position triangularly obovate. Staminalfilaments to 1.2 cm, glabrous. Style glabrous; stigma equally2-lobed. Capsule oblong-ovoid, 0.7–1 cm, glabrous. Seeds elliptic-ovatein outline, tuberculate. Fl. Sep–Oct, fr. Oct.Thickets, along trails and roads, evergreen broad-leaved forests,rocky shores; near sea level to 1200 m. Guangdong, Guangxi, Hainan,Taiwan [Laos, Philippines].Hypoestes purpurea is a medicinal and ornamental plant.2. Hypoestes cumingiana (Nees) Bentham & J. D. Hooker,Gen. Pl. 2: 1122. 1876.枪刀菜 qiang dao caiPeristrophe cumingiana Nees in A. Candolle, Prodr. 11:498. 1847.Herbs to 1 m tall, suffrutescent. Stems subterete, ascendingto decumbent, distally minutely pubescent. Petiole to 2 cm; leafblade ovate-oblong to linear-lanceolate, 4–14 × 1–4 cm, bothsurfaces glabrous, secondary veins 3–6 on each side of midvein,base truncate to cuneate, margin slightly undulate, apex acuminateto acute. Panicles terminal or axillary, to 40 cm, lax;branches of panicle usually subtended by subleaflike inflorescencebracts; floral bracts lanceolate, ca. 4.5 × 1 mm; bracteoleslanceolate-subulate to linear, outer pair 2.5–4 mm, puberulent,inner pair smaller. Flowers single or paired, crowded attips of branches. Calyx 5-lobed nearly to base, lobes subulate,outside puberulent with non-glandular and gland-tipped trichomes.Corolla white to light purple with darker purplemarkings, 1.6–2 cm, outside puberulent; tube basally cylindricaland ca. 1 mm wide for 5–8 mm; lip in lower positionelliptic, 8–1.2 × 7–8 mm, apex rounded; lip in upper positionca. 10 × 4 mm. Staminal filaments ca. 8 mm. Style 1.4–2 cm,sparsely strigose; stigma 2-lobed. Capsule oblong-ovoid, ca. 1.3cm, densely puberulent. Seeds blackish brown, ovate in outline,ca. 3 × 3 mm, verrucose. Fl. Nov–Mar, fr. Mar–Jun.Trailside slopes, along streams; 100–500 m. Taiwan [Philippines].

468ACANTHACEAE3. Hypoestes triflora (Forsskål) Roemer & Schultes, Syst.Veg. 1: 141. 1817.三花枪刀药 san hua qiang dao yaoJusticia triflora Forsskål, Fl. Aegypt.-Arab. 4. 1775; Diclipterariparia Nees var. yunnanensis Handel-Mazzetti.Herbs to 1 m tall, decumbent, much branched. Stems 4-angled, sulcate, strigose. Petiole 1–3 cm, pubescent; leaf bladeovate, ovate-elliptic, or oblong, (1.3–)3–10 × (1–)2–4 cm, bothsurfaces sparsely pubescent, secondary veins 4–7 on each sideof midvein, base cuneate and decurrent onto petiole, margin minutelycrenate to denticulate, apex acuminate. Cymes pedunculatein leaf axils and/or in axils of subleaflike bracts distallyor in a terminal cluster, (1–)3(–5)-flowered; outer pair of bracteoleselliptic to obovate to oblanceolate, 0.7–1.7 × 0.4–0.6 cm,green toward apex, abaxially pilose, base cuneate, apex roundedto acute; inner pair of bracteoles linear-lanceolate to oblanceolate,smaller, basally connate, apex acute. Calyx ca. 6 mm, 5-lobed almost to base; lobes linear-lanceolate, ca. 5 mm. Corollawhite to pink with maroon markings, ca. 1.5 cm, outside pubescent;tube basally cylindric and ca. 1 mm wide for ca. 1 cm;lip in lower position lanceolate, ca. 5 × 1.5 mm, apex slightly 2-lobed; lip in upper position oblong, ca. 5 × 3 mm. Ovary pubescent;style ca. 1.4 cm, glabrous; stigma unequally 2-lobed.Capsule ca. 9 mm, pubescent (at least distally). Seeds black,ovate in outline, ca. 2 × 2 mm, verrucose. Fl. Jul–Sep, fr. Sep–Jan. 2n = 30.Trailsides, forests; 300–2400 m. Yunnan [Bhutan, India, Myanmar,Nepal; Africa].29. BARLERIA Linnaeus, Sp. Pl. 2: 636. 1753.假杜鹃属 jia du juan shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielShrubs, subshrubs, or perennial herbs, with cystoliths, usually spiny. Leaves opposite, petiolate. Inflorescences axillary cymes(sometimes secund or reduced to dense clusters), terminal spikes, or flowers solitary; bracts present or absent; bracteoles 2, sometimesbecoming spinose. Calyx deeply 4-lobed; outer 2 lobes larger [anterior lobe sometimes 2-cleft]; inner 2 lobes smaller. Corollafunnel-shaped, usually large; limb 5-lobed (usually with upper lip appearing 4-lobed); lobes subequal, quincuncial in bud. Stamens [2or]4 and didynamous, one pair usually greatly reduced and/or sterile; anthers 2-thecous; thecae ± equal in size, equally inserted,lacking basal appendages; staminodes 1 or 3, sometimes bearing very small anthers. Ovary with 2 ovules per locule; stigma 2-cleft orentire. Capsule substipitate or lacking a distinct basal stalk, subellipsoid to ovoid, 2–4-seeded; retinacula present. Seeds discoid, pubescentwith appressed hygroscopic trichomes.About 80–120 species: tropical regions of Africa and Asia, one species in tropical America; four species (one endemic) in China.Barleria procumbens Loureiro (Fl. Cochinch. 2: 377. 1790) is not treated here because no material was available for study. Barleria lupulinaLindley is cultivated only in China and not treated here.1a. Margin of calyx lobes (at least outer lobes) serrate or spiny.2a. Flowers in dense, shortly pedunculate cymes from leaf axils; bracteoles linear to linear-lanceolate,2.4–6.5 cm .................................................................................................................................................................. 1. B. cristata2b. Flowers in axillary and terminal dense spikes; bracteoles elliptic, ca. 1.2 cm ........................................................ 3. B. strigosa1b. Margin of calyx lobes entire (apex spine-tipped in B. prionitis).3a. Plants with spines in at least some leaf axils; calyx lobes apically spinose; corolla yellowish to orange ............. 2. B. prionitis3b. Plants without spines; calyx lobes not apically spinose; corolla not yellowish to orange.4a. Flowers in dense spikes; calyx purple; narrow basal portion of corolla tube ca. 3 × as long as throat;bracteoles ca. 1/2 as long as calyx; seeds 4 per capsule .................................................................................... 3. B. strigosa4b. Flowers solitary in leaf axils; calyx green; narrow basal portion of corolla tube shorter than throat;bracteoles equaling or longer than calyx; seeds 2 per capsule ................................................................. 4. B. integrisepala1. Barleria cristata Linnaeus, Sp. Pl. 2: 636. 1753.假杜鹃 jia du juanBarleria cavaleriei H. Léveillé; B. ciliata Roxburgh; B.cristata var. mairei H. Léveillé; B. dichotoma Roxburgh; B.laciniata Nees; B. napalensis Nees.Subshrubs to 2 m tall. Stems terete, branched, coveredwith soft trichomes. Petiole (0–)0.3–1 cm; leaf blade elliptic tooblong to ovate, 2–10 × 1–4 cm, both surfaces villous especiallyalong veins, secondary veins 4–7 on each side of midvein,base cuneate and decurrent onto petiole, apex acute tosometimes acuminate. Inflorescences axillary short and densecymes, shortly pedunculate; bracts absent; bracteoles variable,linear to linear-lanceolate, 2.4–6.5 × 0.5–1.5 cm, 3–7-veined,base cuneate, margin usually spiny but sometimes bristly piloseand becoming spinescent with age, apex acuminate. Outer calyxlobes ovate to narrowly elliptic to lanceolate, 1.2–2.5 × 0.5–1.3cm, pilose, reticulately veined, margin spiny, apex mucronate;inner calyx lobes linear to lanceolate, 6–12 mm, 1-veined, marginscarious. Corolla purplish blue, 4.5–6.5 cm, outside pilose;tube basally narrowly cylindric then gradually widened; lobesoblong-elliptic, ca. 1.5 cm, equal. Stamens 4, didynamous; filamentspilose especially toward base; staminode 1, filamentsparsely pilose. Ovary oblong-ellipsoid, glabrous; style linear,ca. 2.5 cm, glabrous; stigma slightly inflated. Capsule 1.2–1.8cm, glabrous, 4-seeded. Seeds subglobose to ovoid, 4–5 × ca. 4mm. Fl. May, Aug–Dec, fr. May, Oct. 2n = 40.

ACANTHACEAE 469Roadsides, slopes, along streams, in xeric vegetation; below 100–2600 m. Fujian, Guangdong, Guangxi, Guizhou, Hainan, Sichuan, Taiwan,?Xizang, Yunnan [Bhutan, Cambodia, India, Indonesia, Laos,Myanmar, Nepal, Pakistan, Philippines, Singapore, Sri Lanka, Thailand,Vietnam].Barleria cristata is a medicinal and ornamental plant.2. Barleria prionitis Linnaeus, Sp. Pl. 2: 636. 1753.黄花假杜鹃 huang hua jia du juanShrubs to 1.8 m tall with spines in lower leaf axils,branched. Stems and branches terete, smooth, lenticellate, glabrous.Petiole 1–2.5 cm; leaf blade elliptic to ovate, 4–10.5 ×1.8–5.5 cm, both surfaces pubescent when young but soon glabrescent,sparsely strigose along midvein, base attenuate anddecurrent onto petiole, margin entire, apex acute. Flowers clusteredin axils of upper leaves and/or bracts; bracts linear-oblong,1.2–2.2 × 0.2–0.8 cm, margin ciliate, apex abruptly acuminate;bracteoles linear-lanceolate, to 1.4 × 0.2 cm, spinetipped.Outer calyx lobes ovate-oblong, ca. 1.5 × 0.4 cm, apexmucronate; inner calyx lobes linear-lanceolate, ca. 1.3 × 0.2 cm,apex mucronate. Corolla yellow to orange, 2.4–4 cm, outsidepilose; tube with narrow basal portion slightly shorter thanthroat; lobes ovate-oblong to orbicular, 8–10 × 6–7 mm, recurved.Stamens 4, longer pair ca. 1.1 cm with anthers ca. 3.2mm, shorter pair ca. 1.5 mm with anthers ca. 1 mm. Ovaryovoid; stigma slightly enlarged, 2-cleft, exserted. Capsule ovoid,ca. 1.8 cm, contracted gradually at tip forming a beak, 2-seeded.Seeds ovate in outline, ca. 7 × 5 mm. Fl. Oct–Dec, fr. Dec–Feb.2n = 40.Roadsides, thickets, dry places in evergreen broad-leaved forests;ca. 600 m. Yunnan [India, Laos, Myanmar, Sri Lanka, Thailand, Vietnam;Africa, Madagascar].This species is used medicinally.The distribution of Barleria prionitis is commonly given asthroughout tropical Africa and Asia, where it is often introduced; its nativedistribution is not known with certainty.3. Barleria strigosa Willdenow, Sp. Pl. 3: 379. 1800.紫萼假杜鹃 zi e jia du juanBarleria purpureosepala H. P. Tsui.Subshrubs to 80 cm tall, branched. Stems subterete,coarsely fulvous strigose. Petiole 0.7–2.5 cm, brownish yellowishstrigose; leaf blade elliptic to ovate, 5.5–15 × 2.2–5.5cm, both surfaces fulvous strigose especially along veins, secondaryveins ca. 7 on each side of midvein, base cuneate anddecurrent onto petiole, apex acute. Flowers in axillary and terminaldense spikes to 5 cm; bracts oblong to elliptic-oblong,abaxially shortly strigose along veins, apex sharply acute;bracteoles elliptic, ca. 1.2 cm, 1-veined, margin ciliate, apexacute. Outer calyx lobes purple, ovate, ca. 3.2 × 1.8 cm, baserounded, margin subentire to dentate, apex acute; inner calyxlobes yellowish brown, lanceolate, ca. 1.5 × 0.2 cm, both surfacesstrigose, apex acute. Corolla purplish red, 5–6 cm; tubebasally cylindric for ca. 3.7 cm, throat ca. 1.2 cm; lobes obovate-oblong,ca. 2 × 1–1.5 cm, subequal. Stamens 4; filamentsbasally pilose, longer pair ca. 1.3 cm with anthers ca. 5 mm,shorter pair ca. 3 mm with anthers ca. 1.5 mm; staminode 1.Ovary ovoid, ca. 3 mm; style ca. 2.5 cm. Capsule ellipsoid, 1.4–1.8 × ca. 0.5 cm, glabrous, shining, 4-seeded. Fl. Nov, fr. Feb.2n = 40.Dense forests; ca. 900 m. Yunnan [Bhutan, Cambodia, India, Indonesia,Malaysia, Myanmar, Nepal, Sri Lanka, Thailand, Vietnam].4. Barleria integrisepala H. P. Tsui, Acta Bot. Yunnan. 12:269. 1990.全缘萼假杜鹃 quan yuan e jia du juanShrubs to 1 m tall, evergreen. Stems sparsely lenticellate,strigose, soon glabrescent. Leaves clustered at shoot apices;petiole 3–5 cm, strigose; leaf blade ovate, narrowly ovate, orlanceolate, 3–4.5 × 0.7–1.8 cm, both surfaces pilose and strigosealong veins, secondary veins 4–6 on each side of midvein,base cuneate and decurrent onto petiole, apex acute. Flowersaxillary, solitary; bracteoles linear, ca. 14 × 1 mm, sparsely pilose.Pedicel short. Outer calyx lobes elliptic, ca. 1.3 × 0.7 cm,outside sparsely strigose, inside pilose, base cuneate, marginciliate, apex acute; inner calyx lobes lanceolate, ca. 5 × 1 mm,margin ciliate, apex acuminate. Corolla bluish purple, funnelshaped,ca. 3.5 cm; tube basally cylindric for ca. 7 mm, throatca. 1.5 cm; lobes suborbicular, ca. 1.4 × 1.4 cm, subequal.Stamens 4; filaments basally pilose, longer pair ca. 1.7 cm withanthers ca. 4 mm, shorter pair ca. 4 mm with anthers ca. 1.5mm; staminode 1, filament ca. 4 mm. Ovary subellipsoid, ca. 4mm, glabrous; stigma enlarged. Capsule ellipsoid, ca. 1.7 cm,contracted at tip forming a beak, 4-seeded. Seeds ovate in outline,ca. 5 × 5 mm. Fl. Aug–Sep, fr. Nov–Dec.● Thickets along streams; 1900–2000 m. Sichuan (Muli).30. LEPIDAGATHIS Willdenow, Sp. Pl. 3: 400. 1800.鳞花草属 lin hua cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs [or shrubs], perennial, with cystoliths, often anisophyllous. Leaves opposite; leaf blade margin entire or crenate-serrate.Inflorescences dense spikes or cymes condensed into globose heads, sometimes clustered; bracts and bracteoles present. Calyx 5-lobed; lobes heteromorphic; posterior lobe usually largest; lateral 2 lobes smallest; anterior 2 lobes usually connate to a greaterdegree than other lobes. Corolla small; tube usually expanded into a distinct throat near middle; limb 2-lipped; lower lip 3-lobed,patent, usually spotted, lobes subequal or middle one slightly larger; upper lip erect or slightly patent, entire to 2-lobed; lobesquincuncial in bud. Stamens 4, didynamous [or with one pair reduced or absent], usually included in corolla tube; filaments short;anthers 2-thecous [or with one pair of anthers sometimes 1-thecous]; thecae oblong, parallel or one lower and somewhat divergent,

470ACANTHACEAEbase muticous or spurred. Ovary with 2 ovules per locule; stigma subcapitate, entire or slightly 2-cleft. Capsule subellipsoid tooblong to linear-conical, lacking a stalk, to 4-seeded; retinacula present. Seeds discoid, pubescent with hygroscopic trichomes.About 100 species: tropical to subtropical regions worldwide (mostly paleotropical); seven species (three endemic) in China.1a. Inflorescences globose or ovoid heads to 1 cm; leaves sessile to subsessile; leaf blade usually less than 2 cm,base truncate to subcordate .............................................................................................................................................. 6. L. secunda1b. Inflorescences ± elongate and secund spikes to 5.5 cm; leaves usually petiolate; leaf blade to 14.5 cm, basecuneate to attenuate and usually decurrent onto petiole (in L. inaequalis abruptly truncate to rounded anddecurrent onto petiole).2a. Leaf blade linear to linear-lanceolate, (4.6–)6.5–16 × as long as wide ............................................................. 7. L. stenophylla2b. Leaf blade ovate to elliptic to subcircular to obovate, 1.6–4.5(–5.6) × as long as wide (in L. fasciculata,sometimes only distalmost 1 or 2 nodes with leaf blade linear to linear-lanceolate and 6–9.5 × as longas wide and others ovate to elliptic and 1.7–3 × as long as wide).3a. Anterior calyx lobes connate at base for 10%–25% of their length; bracts (1 or)3(or 5)-veined.4a. Corolla pinkish; bracts broadly ovate to ovate-oblong, 2.5–4.4 mm wide; posterior calyx lobe3-veined; distinct segments of anterior calyx lobe oblanceolate to linear, 6–6.5 mm; capsule4.5–5 mm, pubescent distally ................................................................................................................. 1. L. fasciculata4b. Corolla white; bracts narrowly lanceolate, 1–1.5 mm wide; posterior calyx lobe 5-veined; distinctsegments of anterior calyx lobe narrowly lanceolate, 9–10 mm; capsule 6–6.5 mm, pubescentthroughout .............................................................................................................................................. 3. L. hainanensis3b. Anterior calyx lobes connate at base for 30%–60% of their length; bracts 1(or 3)-veined.5a. Leaf blade 1–4(–6) cm, 1.6–2 × as long as wide, base abruptly rounded to truncate and decurrentonto petiole; plants decumbent, not conspicuously woody at base ........................................................ 4. L. inaequalis5b. Leaf blade to 14.5 cm, usually more than 2 × as long as wide, base cuneate to long attenuate anddecurrent onto petiole; plants not decumbent, conspicuously woody at base.6a. Calyx 5–6 mm ................................................................................................................................. 2. L. formosensis6b. Calyx 8–10 mm ...................................................................................................................................... 5. L. incurva1. Lepidagathis fasciculata (Retzius) Nees in Wallich, Pl.Asiat. Rar. 3: 95. 1832.齿叶鳞花草 chi ye lin hua caoRuellia fasciculata Retzius, Observ. Bot. 4: 28. 1786.Herbs to 40 cm tall, erect to spreading, isophyllous toanisophyllous. Stems 4-angled to 4-angled and sulcate, pubescentor glabrate. Petiole 1.5–3.5 cm; leaf blade ovate to elliptic(distalmost 1 or 2 nodes with leaves sometimes linear to linearlanceolate),1.3–12.5 × 0.7–5(–7) cm, 1.7–3 (6–9.5 in distalmostlinear leaves when present) × as long as wide, both surfacespubescent but trichomes sometimes sparse or restricted tomidvein, secondary veins 4 or 8 or more on each side of midvein,base attenuate and decurrent onto petiole, margin entire tocrenate-serrate, apex acuminate. Spikes elongate, 2–4 cm;bracts broadly ovate to ovate-oblong, 7–10 × 2.5–4.4 mm,abaxially pubescent, 1- or 3-veined, veins prominent, marginciliate with gland-tipped and non-glandular trichomes, apexaristate; bracteoles ovate-elliptic, abaxially pubescent, marginciliate with glandular and non-glandular trichomes. Calyx ca. 8mm, pubescent with gland-tipped and non-glandular trichomes;posterior lobe narrowly elliptic to oblong-lanceolate, ca. 8 ×1.5–1.7 mm, 3-veined; lateral lobes lanceolate, ca. 6.5 mm; anteriorlobes connate at base for 17%–25% of their length, distinctsegments oblanceolate to linear and 6–6.5 mm. Corollapinkish, 0.8–1.5 cm, outside pubescent; lower lip 2.5–4 mm,lobes ca. 1 × 0.7–1 mm. Capsule 4.5–5 mm, distally pubescent.Seeds subcircular in outline, 0.8–1.2 mm in diam. Fl. Nov–Mar,fr. Jan–May. 2n = 20.Thickets, roadsides, streamsides. Hainan, Yunnan [Bangladesh,Laos, Malaysia, Myanmar, N Thailand].2. Lepidagathis formosensis C. B. Clarke ex Hayata, J. Coll.Sci. Imp. Univ. Tokyo 30(1): 213. 1911.台湾鳞花草 tai wan lin hua caoHerbs to 45(–90) cm tall, suffruticose, erect, isophyllous toconspicuously anisophyllous. Stems 4-angled to 4-angled andsulcate, glabrous or slightly pubescent. Petiole 0.5–2 cm; leafblade ovate to narrowly elliptic to broadly elliptic to obovate,2.5–14.5 × (0.8–)1–6 cm, 2–3.8(–5.6) × as long as wide, abaxiallysparsely pubescent along veins, adaxially glabrous, secondaryveins 4–7 on each side of midvein, base long attenuateand decurrent onto petiole, margin entire to sinuate, apex acuminate.Spikes elongate, 1–5.5 cm, secund; bracts ovate to lanceolate,3.5–5 × 1–2 mm, 1(or 3)-veined, abaxially and marginallypubescent with non-glandular and sometimes glandtippedtrichomes, apex aristate; bracteoles similar to bracts ornarrower. Calyx 5–6 mm, pubescent with non-glandular andsometimes gland-tipped trichomes; posterior lobe ovate to oblong-lanceolate,5–5.9 × 1.2–1.8 mm, 3-veined; lateral lobeslanceolate to oblong-lanceolate, 4.5–5.8 × 0.7–0.9 mm; anteriorlobes connate at base for 50%–60% of their length, distinct segmentslanceolate-subulate and 2.5–3 mm. Corolla white withpurple markings on limb, 6.5–9 mm, outside pubescent; lowerlip 2.3–3.1 mm, lobes 1–2 × 1–2 mm. Capsule 5–7 mm, distallypubescent. Seeds subcircular to broadly ovate in outline, 1.3–1.8 mm in diam. Fl. Sep–May, fr. Jan–Apr.● Secondary evergreen broad-leaved forests, along streams,ditches, slopes; 100–2300 m. Guangdong, Taiwan.Lepidagathis formosensis is similar to L. incurva, if it is indeeddistinct. On most plants from Taiwan the bracts and calyx are pubescentwith only non-glandular trichomes. On some, however (e.g., B. Bar-

ACANTHACEAE 471tholomew & D. E. Boufford 6193 at CAS), conspicuous gland-tippedtrichomes are present on these structures. In other respects, such plantsagree with non-glandular specimens of L. formosensis.3. Lepidagathis hainanensis H. S. Lo, Fl. Hainan. 3: 598.1974.海南鳞花草 hai nan lin hua caoHerbs to 30 cm tall, erect, anisophyllous. Stems 4-angled,subglabrous to slightly pubescent. Petiole (0–)0.5–1.5 cm; leafblade ovate to elliptic, 1–12 × 0.5–3.5 cm, 1.8–3.5 × as long aswide, abaxial surface sparsely and inconspicuously puberulentespecially along midvein, adaxial surface glabrous, secondaryveins 4–6 on each side of midvein, base narrowly cuneate anddecurrent onto petiole, margin entire to irregularly subsinuate,apex obtuse to rounded to acute. Spikes elongate, 2–6 cm,secund; bracts narrowly lanceolate, 8–10 × 1–1.5 mm, abaxiallyand marginally pubescent with gland-tipped and non-glandulartrichomes, 3(or 5)-veined, apex aristate; bracteoles similar tobracts. Calyx 0.9–1.2 cm, pubescent with gland-tipped and nonglandulartrichomes; posterior lobe lanceolate, 8.5–11 × 1.5–2mm, 5-veined; lateral lobes lanceolate-subulate, ca. 7 × 1 mm;anterior lobes connate at base for ca. 10% of their length,distinct segments narrowly lanceolate and 9–10 mm. Corollawhite, 9–10 mm, outside pubescent; lower lip not seen. Capsule6–6.5 mm, pubescent throughout. Seeds not seen. Fl. Feb–Apr, fr. Feb–Apr.● Forests. Guangxi, Hainan.4. Lepidagathis inaequalis C. B. Clarke ex Elmer, Leafl.Philipp. Bot. 5: 1695. 1913.卵叶鳞花草 luan ye lin hua caoHerbs 10–20 cm tall, decumbent, not conspicuouslywoody, isophyllous. Stems 4-angled, sulcate, glabrous or 2-fariouslypubescent. Petiole 3–15 mm; leaf blade broadly ovate tosubcircular, 1–4(–6) × 0.65–2.5(–3) cm, 1.6–2 × as long aswide, sparsely pubescent especially along midvein, secondaryveins 3–5 on each side of midvein, base abruptly rounded totruncate and decurrent onto petiole, margin entire or shallowlysinuate, apex acute to acuminate. Spikes elongate, 1.2–2 cm,secund; bracts lanceolate, 3–5 × 1–1.4 mm, abaxially and marginallypubescent with gland-tipped trichomes (glands sometimesrestricted to distal portion of margin), 1(or 3)-veined,apex aristate; bracteoles similar to bracts or smaller. Calyx ca. 6mm, pubescent with gland-tipped and non-glandular trichomes;posterior lobe oblong-lanceolate, ca. 6.2 × 1.1 mm, 3-veined;lateral lobes lanceolate, ca. 5.8 × 0.7 mm; anterior lobes connateat base for ca. 50% of their length, distinct segmentslanceolate and ca. 3.2 mm. Corolla white, ca. 8 mm, outsidepubescent; lower lip ca. 3 mm, lobes ca. 2 × 2 mm. Capsule ca.5 mm, distally pubescent. Seeds not seen. Fl. and fr. Sep.Evergreen broad-leaved forests, shaded roadsides; below 100 m.Taiwan [Japan, Philippines].Corollas of Lepidagathis inaequalis have been reported as 4–5mm and glabrous outside (e.g., C. F. Hsieh & T. C. Huang, Fl. Taiwan,ed. 2, 4: 671. 1998). Those of Kao 704 (CAS) are ca. 8 mm and pubescentoutside.Lepidagathis inaequalis is similar to L. formosensis, if they areindeed distinct. Some plants from Taiwan with glandular bracts andcalyces (e.g., Hsieh 787 at CAS) appear somewhat intermediate betweenthem in leaf shape. Plants treated as L. inaequalis have glandularbracts and calyces, decumbent stems, and usually smaller leaf bladesthat are broadly ovate to subcircular and abruptly constricted basally(truncate to rounded). In L. formosensis, bracts and calyces are oftennon-glandular, stems are erect, and leaf blades are usually larger,ovate to elliptic to obovate, and gradually attenuate-tapered at thebase. Among the limited specimens studied, those of L. inaequalis areisophyllous, whereas most of those of L. formosensis are anisophyllous.5. Lepidagathis incurva Buchanan-Hamilton ex D. Don,Prodr. Fl. Nepal. 119. 1825.鳞花草 lin hua caoLepidagathis hyalina Nees.Herbs 30–100 cm tall, base often conspicuously woody,often ± anisophyllous. Stems 4-angled, sulcate, pubescent orsoon glabrescent. Petiole 0.5–3.5 cm; leaf blade ovate to ellipticto narrowly elliptic, 2.5–11 × 1–4.5 cm, 1.7–4.5 × as long aswide, both surfaces pubescent but soon glabrescent with fewtrichomes or sparsely pubescent along veins, secondary veins(4–)7–9 on each side of midvein, base cuneate and conspicuouslydecurrent onto petiole, margin entire and slightly sinuate,apex acute to shortly acuminate. Spikes elongate, (0.5–)1–3.8cm, secund; bracts oblong-lanceolate, 6–10 × 1.5–2 mm, abaxiallyand marginally pubescent with gland-tipped (sometimesabsent) and non-glandular trichomes, 1-veined, apex long acuminate;bracteoles similar to bracts except usually smaller. Calyx8–10 mm, pubescent with gland-tipped (sometimes absent)and non-glandular trichomes or glabrescent; posterior lobe oblong-lanceolate,8–9 × 1.5–1.7 mm, 3-veined; lateral lobes lanceolate,6–7.5 × 0.7–0.8 mm; anterior lobes connate at base for30%–55% of their length, distinct segments lanceolate and 4.8–6 mm. Corolla white streaked with purple, 8–10 mm, outsidesparsely pubescent; lower lip 2.5–3 mm, lobes 1.5–3 × 1.3–1.8 mm. Stamens not seen. Ovary not seen; style not seen; stigmanot seen. Capsule 5–6 mm, distally pubescent. Seeds subcircularin outline, 1.5–1.8 mm in diam. Fl. and fr. Oct–Mar. 2n= 20, 44, 84.Grasslands, thickets, roadsides, hedgerows, streamsides; 100–2200 m. Guangdong, Guangxi, Hainan, Yunnan [Bangladesh, India,Myanmar, Thailand, Vietnam].This species is used medicinally.6. Lepidagathis secunda Nees in A. Candolle, Prodr. 11: 259.1847.小琉球鳞花草 xiao liu qiu lin hua caoRuellia secunda Blanco, Fl. Filip. 495. 1837, not Vahl(1794).Herbs to 30 cm tall, erect or spreading. Stems 4-angled,subglabrous. Leaves sessile to subsessile; leaf blade ovate toovate-lanceolate, 1.5–2 × 0.7–1 cm, both surfaces pubescent,base truncate to subcordate, margin entire, apex acute to shortlyacuminate. Spikes globose to ovoid, to 1 cm; bracts oblong-lanceolate.Corolla to 6 mm, outside pubescent. Stamens included

472ACANTHACEAEin corolla tube; thecae ca. 1 mm, pilose at tip. Ovary oblong;style sparingly hirsute. Capsule ca. 4 mm. Seeds subcircular inoutline.Taiwan [Philippines].No specimens of Lepidagathis secunda have been seen, and publisheddescriptions are inadequate to evaluate the status of either thisspecies or its occurrence in Taiwan. Information above is based entirelyon that provided by C. F. Hsieh and T. C. Huang (Fl. Taiwan, ed. 2, 4:673. 1998).7. Lepidagathis stenophylla C. B. Clarke ex Hayata, J. Coll.Sci. Imp. Univ. Tokyo 30(1): 214. 1911.柳叶鳞花草 liu ye lin hua caoHerbs to 40 cm tall, suffrutescent, often ± anisophyllous.Stems 4-angled to 4-angled and sulcate, subglabrous or evenlyto ± bifariously pubescent. Petiole (1–)5–12 mm; leaf bladelinear to linear-lanceolate, 3.5–11 × 0.3–1.9 cm, (4.6–)6.5–16 ×as long as wide, both surfaces subglabrous or pubescent,secondary veins 4 or 5 on each side of midvein, base narrowedand decurrent onto petiole, margin shallowly sinuate, apexacute to acuminate. Spikes elongate, 1.3–3.8 cm, secund; bractslanceolate to lanceolate-ovate, ca. 5 × 1–1.5 mm, abaxially andmarginally pubescent with non-glandular trichomes, 1-veined,apex aristate; bracteoles similar to bracts except usually smallerand narrower. Calyx 5.5–8.2 mm, pubescent with non-glandulartrichomes; posterior lobe ovate to oblong-lanceolate, 5–7.5 ×1.1–1.3 mm, 3-veined; lateral lobes lanceolate, 4.5–5.7 × 0.7–0.8 mm; anterior lobes connate at base for 20%–55% of theirlength, distinct segments narrowly lanceolate and 3–6 mm.Corolla white, ca. 5 mm, outside pubescent; lower lip 2–2.3mm, lobes 1.5–1.6 × 0.6–0.8 mm. Capsule 4.5–5.5 mm, pubescent± throughout or only distally. Seeds subcircular in outline,1–1.8 mm in diam. Fl. and fr. Dec.● Forest trails; 200–400 m. Hong Kong, Taiwan.31. CHROESTHES Benoist, Bull. Mus. Natl. Hist. Nat. 33: 107. 1927.色萼花属 se e hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielShrubs, subisophyllous or anisophyllous. Leaves opposite, petiolate; leaf blade margin entire. Inflorescences terminal thyrses;bracts and bracteoles greenish. Calyx unequally 5-lobed; posterior lobe largest, 2 lateral lobes smaller than 2 anterior lobes. Corollatube basally cylindric and expanded distally into a throat; limb 2-lipped, lower lip 3-lobed, upper lip 2-lobed; lobes quincuncial inbud. Stamens 4, not connate, posterior pair shorter than anterior pair, inserted at base of throat; anthers 2-thecous; thecae parallel,inserted at different heights, dorsally pubescent, base spurred. Ovary with 2 ovules per locule; style basally sparsely pubescent.Capsule stipe absent or barely present; retinacula present. Seeds compressed, brownish pubescent.Three species: China, Laos, Malaysia, Myanmar, Thailand, Vietnam; one species in China.1. Chroesthes lanceolata (T. Anderson) B. Hansen, Nordic J.Bot. 3: 209. 1983.色萼花 se e huaAsystasia lanceolata T. Anderson, J. Linn. Soc., Bot. 9:524. 1867; A. kerrii Craib; A. silvicola W. W. Smith; Chroesthespubiflora Benoist; C. racemiflora Bremekamp; C. racemifloravar. glabrior Bremekamp; C. silvicola (W. W. Smith) E.Hossain.Shrubs 0.5–3 m tall, anisophyllous. Stems terete, slender,rarely branched, glabrous. Petiole 1–2.5 cm; leaf blade ellipticto oblanceolate to lanceolate, 10–16 × 3–7 cm, both surfacesglabrous, secondary veins 6–9 on each side of midvein, basecuneate, margin entire or subsinuate, apex acuminate. Thyrses3–7 cm; cymes sessile, 1–3-flowered; bracts elliptic to broadlylanceolate, 3–9 × 1–3 mm, gland-tipped pubescent, apex acute;bracteoles narrowly elliptic to broadly lanceolate, 4–9 × 0.7–1.2mm. Pedicel 1–5 mm. Calyx 1–1.6 cm, outside gland-tippedpubescent; posterior lobe lanceolate, ovate, or subelliptic; laterallobes linear-lanceolate; anterior lobes connate to 2/3 theirlength. Corolla white with pink or purple spots, ca. 2.5 cm, outsidepubescent; tube basal portion ca. 9 mm, throat ca. 1.5 cm;lower lip 3-lobed; upper lip 2-lobed. Stamens included in throat;filaments 1–1.2 cm, glabrous; anther thecae 2.1–2.3 mm, pubescentat apex and along sides, basal spur pointed. Ovary apexpubescent; style ca. 2.5 cm. Capsule subellipsoid to obovoid,1.2–1.6 cm, glabrous or only at apex pubescent, 4-seeded.Seeds subcircular in outline. Fl. Feb–Mar, fr. May–Jul.Forests; (200–)800–1400 m. Guangxi, S Yunnan [Laos, Myanmar,Thailand, Vietnam].32. GYMNOSTACHYUM Nees in Wallich, Pl. Asiat. Rar. 3: 76, 106. 1832.裸柱草属 luo zhu cao shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielCryptophragmium Nees; Sarcanthera Rafinesque.Herbs, perennial [or shrubs], with cystoliths. Leaves cauline or subradical, opposite; leaf blade margin usually entire. Inflorescencesaxillary or terminal, spikes, racemes, or panicles, often secund; bracts and bracteoles shorter than calyx. Calyx 5-lobed nearlyto base; lobes subequal. Corolla tube longer than limb, basal portion narrowly cylindric, ± expanded distally into a throat; limb 2-lipped, lower lip 3-lobed, upper lip emarginate to 2-lobed; lobes ascending cochlear in bud. Stamens 2, included in or slightly

ACANTHACEAE 473exserted from corolla tube; anther thecae 2, equal or subequal in size, parallel, one or both mucronate at base; staminodes absent.Ovary with 3 to many ovules per locule; stigma 2-lobed, lobes compressed. Capsule without a stipe, linear, 4-angled, many seeded;retinacula present. Seeds compressed, covered with hygroscopic trichomes.About 30 species: tropical Asia; three species (two endemic) in China.1a. Stems very short; leaves in a basal rosette ............................................................................................................. 1. G. subrosulatum1b. Stems elongate; leaves cauline.2a. Inflorescences axillary .................................................................................................................................................. 2. G. listeri2b. Inflorescences terminal .............................................................................................................................................. 3. G. sinense1. Gymnostachyum subrosulatum H. S. Lo, Acta Phytotax.Sin. 17(4): 86. 1979.矮裸柱草 ai luo zhu caoGymnostachyum kwangsiense H. S. Lo.Herbs 20–40 cm tall. Stems short, hirsute. Leaves in abasal rosette; petiole 2–9(–14) cm; leaf blade suborbicular tobroadly ovate, 5–16 × 3.5–11 cm, papery, abaxially subglabrousbut midvein and secondary veins hirsute, adaxially slightlyscabrous, secondary veins 7–9 on each side of midvein, baseslightly cordate to rounded, margin entire or slightly sinuate,apex rounded to acute. Thyrses 10–35 cm, with 1–4 flowers percyme; peduncle 8–19 cm; bracts subulate, ca. 2.5 mm, pilose,apex acute; bracteoles similar to bracts. Calyx 2.5–3 mm; lobessubulate, pilose. Corolla ca. 1.5 cm; lower lip patent, 3-lobed,lobes oblong-ovate and ca. 3 × 2 mm; upper lip triangular, ca. 5mm, erect, 2-dentate. Stamens exserted; filaments 7–8 mm; antherthecae linear, parallel. Ovary 2-locular, with 4 ovules perlocule; style 1.3–1.9 cm. Capsule linear, 1.3–1.6 cm, 8-seeded.Seeds ovate in outline, ca. 1 × 0.8 mm. Fl. Apr–May, fr. Jun–Sep.● Limestone hills; 200–600 m. Guangxi.2. Gymnostachyum listeri Prain, J. Asiat. Soc. Bengal, Pt. 2,Nat. Hist. 69: 171. 1900.云南裸柱草 yun nan luo zhu caoCryptophragmium tonkinense Benoist; Parajusticia petelotiiBenoist.Herbs to 50 cm tall, caulescent, isophyllous. Stemsbranched, 4-angled, pubescent. Leaves cauline; petiole 5–12mm, pubescent; leaf blade elliptic to obovate-elliptic, (4.5–)6–13(–17) × 2.1–5.5 cm, abaxially white pubescent, adaxially glabrousexcept for trichomes along midvein, secondary veins 11–14 on each side of midvein and abaxially prominently curved,tertiary veins inconspicuous, base asymmetric or symmetric and± decurrent onto petiole, margin entire or obscurely sinuate,apex acute to acuminate. Spikes 2–6 cm, secund; peduncle 0.5–1.5 cm; rachis pubescent and gland-tipped pilose; bracts linear,ca. 3 × 0.3 mm, pubescent; bracteoles linear-triangular, ca. 1.5 ×0.5 mm, pubescent. Calyx 0.1–1.1 cm; lobes linear, 8–10 × ca.0.5 mm, outside pubescent, inside glabrous, margin glandtippedpilose. Corolla purple, 1.6–2 cm, pubescent; tube basallycylindric and ca. 2.5 cm wide for ca. 8 mm, throat abruptlybending and widening to 5 mm; lower lip 4–5 mm, 3-lobed,lobes oblong and ca. 3 × 1.5 mm; upper lip 3–4 mm, 2-lobed,lobes broadly ovate and ca. 2 × 2 mm. Staminal filaments ca.4 mm, glabrous; anther thecae oblong, ca. 5 × 1 mm, pilose,spurred at base. Ovary ca. 3 × 1 mm, glabrous; style ca. 1.5 cm,villous. Capsule not seen. Fl. Apr–May.Limestone hills; 200–600 m. Guangxi, Yunnan [Bangladesh, Vietnam].3. Gymnostachyum sinense (H. S. Lo) H. Chu, Bull. Bot. Res.,Harbin 11(1): 48. 1991.华裸柱草 hua luo zhu caoAndrographis sinensis H. S. Lo, Bull. Bot. Res., Harbin1(4): 103. 1981.Herbs to 1 m tall. Stems purplish black, terete, glabrous;young shoots 4-angled, pubescent at nodes. Petiole 2–4 mm;leaf blade ovate to broadly ovate, 1.5–4 × 1–1.8 cm, thicklypaper, both surfaces glabrous, secondary veins ca. 6 on eachside of midvein, base cuneate, margin entire or subentire, apexacute to sometimes caudate. Spikes terminal, few flowered; peduncleca. 1 cm, subglabrous; bracts linear to linear-lanceolate,1.5–2 mm; bracteoles similar to bracts. Calyx ca. 7 mm, glabrous;lobes lanceolate-linear, apex acuminate. Corolla white,ca. 1 cm; lower lip ca. 4 mm, 3-lobed, lobes ovate and ca. 0.5 ×0.5 mm; upper lip subtriangular, ca. 3.5 mm, 2-lobed. Staminalfilaments ca. 2.5 mm, glabrous; anther thecae linear, ca. 3 mm,spurred at base. Ovary ellipsoid, ca. 3 mm, glabrous, ovules 5or 6 per locule; style ca. 7 mm. Capsule narrowly cylindric, 1–1.5 cm, glabrous. Seeds subglobose, ca. 1 × 1 mm, pubescent.Fl. Aug.● Limestone hills. Guangxi.33. ANDROGRAPHIS Wallich ex Nees in Wallich, Pl. Asiat. Rar. 3: 77, 116. 1832.穿心莲属 chuan xin lian shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHaplanthoides H. W. Li; Haplanthus Nees.Herbs or subshrubs, with cystoliths. Leaf blade margin entire. Inflorescences terminal or axillary, panicles, racemes, or sometimesspikes or dense clusters; bracts present; bracteoles small [or absent]. Calyx deeply 5-lobed; lobes narrow, equal or subequal.Corolla tubular to funnelform at base; limb 2-liped; lower lip 3-lobed; upper lip entire or 2-lobed; lobes ascending cochlear in bud.

474ACANTHACEAEStamens 2, exserted from or included in corolla tube; filaments sometimes pubescent; anthers 2-thecous; thecae parallel, equal orsubequal, base muticous and conspicuously pubescent. Capsule linear-oblong or ellipsoid, compressed perpendicular to septum,[6–]12–20-seeded; retinacula present. Seeds subglobose, ± blocky, or ± compressed, surfaces smooth or ornamented, lackingtrichomes.About 20 species: tropical and subtropical Asia; two species (one introduced) in China.1a. Leaf blade lanceolate to oblong-lanceolate; corolla deeply and prominently 2-lipped, lips divergent (i.e., upperlip ± perpendicular or oblique to tube and lower lip); stamens exserted from corolla tube ...................................... 1. A. paniculata1b. Leaf blade ovate; corolla shallowly and inconspicuously 2-lipped, lips not divergent (i.e., lips ± parallel to oneanother); stamens included in corolla tube .................................................................................................................... 2. A. laxiflora1. Andrographis paniculata (N. L. Burman) Wallich ex Neesin Wallich, Pl. Asiat. Rar. 3: 116. 1832.穿心莲 chuan xin lianJusticia paniculata N. L. Burman, Fl. Indica, 9. 1768.Herbs to 50 cm tall, annual, much branched. Stems 4-angled,glabrous. Petiole 0.3–1 cm; leaf blade ovate-lanceolate,lanceolate, or narrowly elliptic, 1.5–7 × 1–2.5 cm, both surfacesglabrous, abaxially pale green, adaxially green, secondary veins3–5 on each side of midvein, base attenuate and decurrent ontopetiole, margin entire, apex acute to shortly acuminate. Inflorescencesterminal, leafy panicles of secund racemes; rachis glabrousto sparsely pubescent; bracts triangular to subulate, 1–1.5mm; bracteoles linear to subulate, 1–1.5 mm. Pedicel 2–9 mm,sparsely pubescent with gland-tipped and non-glandular trichomes(gland-tipped pubescent). Calyx 2.5–3 mm, outside glabrousor gland-tipped pubescent, lobes subulate. Corolla white,0.9–1.5 cm, outside gland-tipped pubescent; tube basally funnelformfor 4–8 mm; lower lip with purple dots, 5–7 mm, erect,lobes ca. 3 mm; upper lip 5–7 mm, reflexed, 2-lobed, lobes ca.1 mm. Stamens exserted from corolla tube. Style 6–10 mm,sparsely pilose toward base. Capsule ellipsoid-compressed, 1.5–2 × 0.3–0.4 cm, glabrous or sparsely pubescent with glandtippedtrichomes, ca. 12-seeded. Seeds ca. 2 × 1.5 mm, rugose.Fl. and fr. throughout year. 2n = 50.Cultivated or naturalized in Anhui, Fujian, Guangdong, Guangxi,Hainan, Hubei, Hunan, Jiangsu, Jiangxi, Yunnan, and Zhejiang [nativeto India and Sri Lanka; cultivated or naturalized in Cambodia, Indonesia,Laos, Malaysia, Myanmar, Thailand, Vietnam, and Caribbean].This species is used medicinally.2. Andrographis laxiflora (Blume) Lindau in Engler & Prantl,Nat. Pflanzenfam. 4(3b): 323. 1895.疏花穿心莲 shu hua chuan xin lianJusticia laxiflora Blume, Bijdr. 789. 1826; Andrographisglomeruliflora Bremekamp; A. laxiflora var. glomeruliflora(Bremekamp) H. Zhu; A. monglunensis Hung T. Chang & H.Zhu; A. tenera (Nees) Kuntze; A. tenuiflora T. Anderson, nom.illeg. superfl.; Haplanthoides yunnanensis H. W. Li; Haplanthustener Nees.Herbs to 50 cm tall, isophyllous or anisophyllous. Stemserect or procumbent, 4-angled, glabrous or pubescent. Petiole3–10 mm; leaf blade ovate, 1.5–9 × 1.5–3(–5.5) cm, both surfacesglabrous, secondary veins 5 or 6 pairs on each side ofmidvein, base slightly attenuate and decurrent onto petiole,margin entire, apex acuminate to shortly acuminate. Inflorescencesaxillary or terminal, racemes or panicles of racemes,these sometimes reduced and flowers appearing as clusters inleaf axils or at ends of long peduncles; rachis glabrous or pubescent(trichomes non-gland-tipped or gland-tipped); bractslinear, subulate, or lanceolate, 1–3 mm, glabrous or pubescent;bracteoles subulate, glabrous or pubescent. Pedicel 0.5–2 mm,glabrous or pubescent. Calyx 2–4 mm, outside glabrous or pubescent,lobes subulate. Corolla white with purplish or pinkishcoloration on lips, 0.8–1.2 cm, outside pubescent with glandtippedtrichomes; tube 6–9 mm, base curved, dorsally ± gibbous;lower lip 1.5–4 mm, lobes 0.5–2 mm; upper lip 1.5–4mm, erect (± parallel to lower lip), entire to 2-lobed. Stamensincluded in corolla tube. Capsule linear-cylindric, 10–20 × ca. 2mm, glabrous or pubescent with gland-tipped trichomes, 16–20-seeded. Seeds ca. 1 × 0.7 mm, rugose. Fl. Aug–Nov, fr.Nov–Mar.Thickets, bamboo forests; 200–1500 m. Guangxi, Guizhou, Hainan,Yunnan [Cambodia, India, Indonesia, Laos, Malaysia, Myanmar,Thailand, Vietnam].Specimens misidentified as Gymnostachyum sanguinolentum(Vahl) T. Anderson (C. Y. Wu, Index Fl. Yunnan. 2: 1676. 1984; C.C. Hu, FRPS 70: 75. 2002) belong here.Variation is evident in the pubescence of rachises, pedicels, bracts,bracteoles, and calyces among plants of Andrographis laxiflora. Thesestructures may be glabrous, have non-glandular trichomes only, havegland-tipped trichomes only, or have a mixture of gland-tipped and nonglandulartrichomes. Capsule may be glabrous or pubescent with glandtippedtrichomes only. Plants with gland-tipped trichomes on thecalyces and capsules have been treated as A. laxiflora var. glomeruliflora(C. C. Hu, FRPS 70: 205. 2002). Because varying degrees of pubescenceare evident on calyces and capsules among specimens fromwithin and outside of China, and because these same differences are nottaxonomically recognized for plants of A. paniculata that show the verysame variations, var. glomeruliflora is not recognized here (in keepingwith B. Hansen, Nordic J. Bot. 5: 353–356. 1985, who did not recognizeit as distinct).34. PHLOGACANTHUS Nees in Wallich, Pl. Asiat. Rar. 3: 76, 99. 1832.火焰花属 huo yan hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielHerbs, shrubs, or small trees, with cystoliths. Leaves opposite; leaf blade large, margin entire or obscurely crenate. Inflores-

ACANTHACEAE 475cences in terminal thyrses or axillary cymes, pedunculate; bracts small; bracteoles small or absent. Calyx deeply 5-lobed; lobes equalto unequal. Corolla tubular, slightly curved (arched); limb subequally 5-lobed or 2-lipped; lower lip 3-lobed, lobes ascending cochlearin bud; upper lip 2-cleft. Stamens 2, inserted at middle or basal portion of tube, included or sometimes slightly exserted beyondcorolla lip; anthers 2-thecous; thecae equal, base divaricate and muticous; staminodes 2, small. Ovary usually glabrous; stigma subentire.Capsule linear, 4-angled, basal portion solid but lacking a conspicuous basal stalk; retinacula present, 8–16-seeded. Seedslenticular, glabrous or pubescent.About 15 species: mainland Asia; two species in China.1a. Inflorescences terminal thyrses, 14–18 cm; corolla purplish red, 4.5–5 cm; anthers not extending beyondcorolla lips; capsule 3.5–5 cm ..................................................................................................................................... 1. P. curviflorus1b. Inflorescences axillary cymes, ca. 4 cm; corolla orange, ca. 1.8 cm; anthers extending beyond corolla lips;capsule 2.5–3 cm ....................................................................................................................................................... 2. P. pubinervius1. Phlogacanthus curviflorus (Wallich) Nees in Wallich, Pl.Asiat. Rar. 3: 99. 1832.火焰花 huo yan huaJusticia curviflora Wallich, Pl. Asiat. Rar. 2: 9. 1831.Shrubs 3–6 m tall. Petiole 1.5–5 cm; leaf blade elliptic tooblong, 12–35 × 9–18 cm, abaxially puberulent along veins,adaxially glabrous, secondary veins 12–17 on each side of midvein,base broadly cuneate and decurrent onto petiole, marginentire to subcrenate, apex acute to acuminate. Thyrses terminal,14–18 × 6–8 cm; cymes (1–)3–5-flowered. Pedicel 5–12 mm,densely tomentose; bracts subulate to lanceolate, 3–7 mm;bracteoles subulate 2–3 mm. Calyx 6–11 mm; lobes subulate tolanceolate, subequal, outside sparsely pubescent, inside denselybrown tomentose. Corolla purplish red, 4.5–5 cm, outsidedensely brownish pubescent and sometimes including glandtippedtrichomes; tube ca. 4.2 cm; lower lip 3-lobed, lobes oblongand ca. 2 × 1.5 mm; upper lip 2-cleft. Stamens slightlyexserted; filaments glabrous; anther thecae ellipsoid, 5–5.5 mm,not extending beyond corolla lips; staminodes 2, small. Ovaryand style glabrous. Capsule 3.5–5 cm, 8–10-seeded. Seeds 5.5–6.3 × 4.2–4.5 mm, pubescent. Fl. Oct–Feb, fr. Feb–May.Thickets, forest margins, ravines; 400–1600 m. Xizang, Yunnan[Bhutan, India, Laos, Myanmar, Thailand, Vietnam].2. Phlogacanthus pubinervius T. Anderson, J. Linn. Soc., Bot.9: 508. 1867.毛脉火焰花 mao mai huo yan huaAeschynanthus dunnii H. Léveillé; Lonicera menelii H.Léveillé.Shrubs or small trees, to 5 m tall, much branched. Petiole0.5–2 cm; leaf blade elliptic-oblong to oblong, (5–)8–18 ×(1–)3.5–5 cm, abaxially puberulent along veins, adaxially scabrous,secondary veins 5–7 on each side of midvein, base attenuate,margin undulate, apex acuminate to long acuminate.Cymes axillary, 1–4-flowered; peduncle 0.8–1.6 cm, 4-angled,pubescent; bracts small, caducous; bracteoles subulate, ca. 3 × 1mm, caducous. Pedicel 0.8–1.5 cm. Calyx 7–8 mm; lobes linear-lanceolate,unequal, outside pubescent, inside gray tomentose.Corolla orange, ca. 1.8 cm, outside pubescent; tube ca. 1.3cm, slightly curved; lower lip deeply 3-lobed, lobes ovate;upper lip 2-cleft. Stamens much exserted; filaments ca. 2.5 cm,glabrous; anther thecae oblong, parallel, extending beyond corollalips; staminodes 2, small. Ovary glabrous; style ca. 2 cm,glabrous. Capsule 2.5–3 cm, glabrous, 8-seeded.Forests, thickets; 700–1500 m. Guangxi, Guizhou, Yunnan [Bhutan,India, Myanmar].35. CYSTACANTHUS T. Anderson, J. Linn. Soc., Bot. 9: 457. 1867.鳔冠花属 biao guan hua shuHu Jiaqi ( 胡嘉琪 Hu Chia-chi), Deng Yunfei ( 邓云飞 ); Thomas F. DanielShrubs or perennial herbs, with cystoliths. Leaf blade margin entire or obscurely crenate. Inflorescences thyrses or rarelyaxillary cymes or racemes; bracteoles 2, small. Calyx deeply 5-lobed; lobes usually covered with gland-tipped trichomes. Corollaorange or pink, funnel-shaped, usually bent ca. 90º and abruptly inflated near middle; limb 2-lipped; lower lip 3-lobed, lobes orbicularor ovate and equal; upper lip 2-lobed; lobes ascending cochlear in bud. Stamens 2, included; anthers 2-thecous; thecae parallel,equal, hirsute, muticous at base; staminodes 2. Ovary ovoid, long hirsute; style filiform; stigma 2-cleft. Capsule cylindric, 8–12-seeded; retinacula present. Seeds ovate in outline, compressed, tomentose.About 15 species: mainland Asia; eight species (three endemic) in China.Five of the Chinese species treated here in Cystacanthus have also been treated in Phlogacanthus. In addition to the macromorphological characteristics(geniculate corolla with an inflated throat in Cystacanthus) distinguishing Cystacanthus and Phlogacanthus, R. W. Scotland (Rev.Palaeobot. Palynol. 72: 229–243. 1992) discussed palynological differences between them in ornamentation of the aperture margins (i.e., aperturemargins with spines ± restricted to a single row in Phlogacanthus and with the spines throughout the aperture margin in Cystacanthus). A thoroughreevaluation of taxa treated in these two genera will be necessary to determine whether both genera are worthy of recognition and if so, to whichgenus each species pertains.

476ACANTHACEAE1a. Corolla outside pubescent with gland-tipped trichomes.2a. Thyrses to 2.5 cm, 3- or 4-flowered ............................................................................................................................ 1. C. affinis2b. Thyrses longer than 5 cm, many flowered.3a. Corolla purple; leaf blade oblong to oblong-lanceolate .............................................................................. 2. C. paniculatus3b. Corolla blue, red, white, or purplish blue; leaf blade ovate to ovate-lanceolate.4a. Young stems pubescent but not densely whitish or yellowish brown villous ........................... 3. C. yangtsekiangensis4b. Young stems densely whitish to yellowish brown villous ................................................................... 4. C. yunnanensis1b. Corolla outside glabrous or non-glandular pubescent.5a. Bracts leaflike, much longer than calyx; corolla glabrous outside ................................................................... 5. C. pyramidalis5b. Bracts small and not leaflike, subequal to calyx; corolla pubescent outside.6a. Leaf blade secondary veins ca. 6 on each side of midvein .......................................................................... 6. C. abbreviatus6b. Leaf blade secondary veins 9–17 on each side of midvein.7a. Leaf blade secondary veins 9–11 on each side of midvein; inflorescences spikelike thyrses; rachiswith 1 flower per node; calyx ca. 6 mm; corolla densely pubescent outside ........................................... 7. C. vitellinus7b. Leaf blade secondary veins 12–17 on each side of midvein; inflorescences paniculate thyrses;rachis with several flowers per node; calyx ca. 3 mm; corolla sparsely pubescent outside ..................... 8. C. colaniae1. Cystacanthus affinis W. W. Smith, Notes Roy. Bot. Gard.Edinburgh 9: 103. 1916.丽江鳔冠花 li jiang biao guan huaShrubs to 2 m tall. Stems terete. Petiole 5–10 mm; leafblade oblong to ovate, 2.5–6 × 1.5–3 cm, both surfaces pubescent,secondary veins 5 or 6 on each side of midvein, base cuneate,margin entire, apex acuminate. Thyrses terminal, to 2.5cm, 3- or 4-flowered; bracts oblong-lanceolate, shorter thancalyx, gland-tipped pubescent, caducous; bracteoles similar tobracts. Pedicel ca. 3 mm, gland-tipped pubescent. Calyx 1.5–2cm, outside pubescent with gland-tipped trichomes; lobes linear,ca. 1 mm wide, apex acute. Corolla yellow outside and lightyellow inside with purplish striae, 3.5–4 cm; lobes ovate, to 1.5cm. Stamens slightly exserted; filaments barbate; anther thecaeto 1 mm, connective barbate; staminodes ca. 2 mm. Ovary ca. 5mm; style ca. 2 cm, glabrous; stigma slightly 2-cleft. Capsulelight yellow, ca. 2 cm, glabrous, 6-seeded. Seeds ovate in outline,ca. 3 × 3 mm.● Thickets, streamsides, trailsides; 1700–2200 m. Sichuan, Xizang,Yunnan.2. Cystacanthus paniculatus T. Anderson, J. Linn. Soc., Bot.9: 458. 1867.鳔冠花 biao guan huaPhlogacanthus paniculatus (T. Anderson) J. B. Imlay.Shrubs to 2 m tall. Stems 4-angled, glabrous; youngbranches winged. Petiole 1–1.5 cm; leaf blade oblong to oblong-lanceolate,10–12 × 2.5–4 cm, abaxially pubescent alongveins, adaxially glabrous, base attenuate and decurrent ontopetiole, margin undulate to crenulate, apex acuminate. Thyrsesterminal, to 18 cm, many flowered; rachis densely villous withgland-tipped trichomes; bracts oblong-lanceolate, shorter thancalyx, gland-tipped pubescent, caducous; bracteoles similar tobracts. Pedicel 2–4 mm, gland-tipped pubescent. Calyx 1.2–1.4cm, gland-tipped pubescent; lobes lanceolate. Corolla light purple,2–2.4 cm, ventricose, outside gland-tipped pubescent, insideglabrous; lower lip with lobes orbicular, subequal, patent.Stamens included; filaments barbate; anther thecae ca. 1 mm,connective barbate; staminodes very small. Ovary pubescent,with 4 ovules per locule; style 1.6–2 cm, glabrous; stigmaslightly 2-cleft. Capsule grayish brown, clavate, ca. 2.3 cm, pubescent,8-seeded. Seeds ovate in outline, ca. 3.5 × 3.5 mm.Forests; 300–2100 m. Yunnan [Myanmar].3. Cystacanthus yangtsekiangensis (H. Léveillé) Rehder, J.Arnold Arbor. 16: 315. 1935.金江鳔冠花 jin jiang biao guan huaStrobilanthes yangtsekiangensis H. Léveillé, Cat. Pl. Yun-Nan, 6. 1915.Shrubs to 1 m tall. Stems subterete, striate, pubescent.Petiole 3–7 mm, pubescent; leaf blade ovate, ca. 3.5 × 1.5 cm,both surfaces pubescent and glaucous, secondary veins 10–12on each side of midvein, base cuneate, margin entire, apexacute. Thyrses terminal or axillary, racemose, secund, to 7 cm;bracts linear-lanceolate, 5–8 mm, densely gland-tipped villous;bracteoles similar to bracts. Pedicel 2–4 mm, gland-tipped pubescent.Calyx 7–10 mm; lobes linear, densely yellowish pubescent.Corolla blue or red, campanulate, ca. 2.5 × 1.2–1.3 cm,ventricose and curved; tube basally ca. 5 mm wide and short;lobes broadly triangular, apex obtuse. Stamens included; filamentsbarbate; anther thecae ovoid, connective barbate withyellowish brown trichomes; staminodes ca. 2 mm. Ovary pubescent;style ca. 2 cm, glabrous; stigma slightly 2-cleft. Capsuleand seeds not seen.● 400–500 m. Yunnan.4. Cystacanthus yunnanensis W. W. Smith, Notes Roy. Bot.Gard. Edinburgh 9: 104. 1916.滇鳔冠花 dian biao guan huaShrubs 1–1.5 m tall, erect. Stems subterete or slightly 4-angled, densely whitish or yellowish brown villous. Petiole 0.5–1.5 cm; leaf blade ovate to ovate-lanceolate, 5–10 × 3–4 cm,abaxially white villous, adaxially densely yellowish villous,secondary veins ca. 6 on each side of midvein, base attenuate,margin entire, apex acuminate to acute. Thyrses paniculate, to15 cm; bracts linear-lanceolate, 5–10 mm, densely gland-tipped

ACANTHACEAE 477villous; bracteoles similar to bracts. Pedicel 1–2 mm, glandtippedvillous. Calyx ca. 1.5 cm, outside densely gland-tippedvillous, inside slightly villous; lobes linear, ca. 13 × 2 mm, subequal,apex acute. Corolla white or purplish blue and with awhitish base, 3–3.5 cm, ventricose, incurved, outside glandtippedpubescent; tube basally ca. 1.5 mm wide and very shortthen wide at middle; lobes orbicular, ca. 10 × 7 mm, unequalwith anterior one longer. Stamens 2; filaments ca. 1.5 cm, basallypilose; anther thecae oblong, ca. 6 mm, connective barbate;staminodes filiform, ca. 2 mm. Ovary ca. 4 mm, villous, with 6ovules per locule; style ca. 2 mm, sparsely villous; stigma minutely2-dentate. Capsule ca. 3.5 cm, ca. 5 mm wide, glandtippedvillous. Seeds disciform, 2–3 × 2–3 mm, puberulent.● Forests; 800–1600 m. Yunnan.5. Cystacanthus pyramidalis Benoist, Bull Soc. Bot. France74: 907. 1928.金塔火焰花 jin ta huo yan huaPhlogacanthus pyramidalis (Benoist) Benoist.Herbs to 1 m tall, perennial. Stems 4-angled, glabrous.Petiole ca. 2.5 cm; leaf blade elliptic to elliptic-ovate, 9–23 ×5.5–10 cm, abaxially glabrous or pubescent along veins, adaxiallyglabrous, secondary veins 8–10 on each side of midvein,base cuneate and decurrent onto petiole, margin entire, apexacute to shortly acuminate. Thyrses terminal, to 26 cm; cymes3-flowered; peduncle 1–3 mm, pubescent; bracts leaflike, ovateto lanceolate, 0.6–3 × 0.3–1.2 cm, diminished in size apicallyon inflorescence, pubescent; bracteoles subulate, ca. 1 mm.Pedicel 3–5 mm. Calyx ca. 3.5 mm, glabrous or sparsely pubescent;lobes equal, linear, apex acuminate to acute. Corolla lightpurplish red, ca. 1 cm, bent to 90º and abruptly inflated at middle,glabrous; lobes ovate, ca. 2 × 1.5 mm. Stamens 2; filamentsca. 0.8 cm, glabrous; anther thecae ovoid, ca. 1.5 mm, connectivehirsute; staminodes ca. 2 mm. Ovary glabrous; style ca. 1cm, glabrous; stigma slightly 2-cleft. Capsule clavate, ca. 2.2cm, 4-angled, 8-seeded. Seeds ca. 2 × 2 mm, compressed.Forests. Hainan [Vietnam].6. Cystacanthus abbreviatus Craib, Bull. Misc. Inform. Kew1911: 438. 1911.缩序火焰花 suo xu huo yan huaPhlogacanthus abbreviatus (Craib) Benoist.Shrubs to 1 m tall. Stems green, terete or 4-angled, slightlypubescent. Petiole 1.5–3.5 cm; leaf blade oblanceolate to obovate-lanceolate,ca. 15 × 5 cm, abaxially slightly pubescentalong veins, adaxially glabrous, secondary veins ca. 6 on eachside of midvein, base narrowly attenuate, margin entire, apexshortly acuminate. Thyrses terminal, 8–12 cm; bracts minute,caducous; bracteoles subulate. Pedicel ca. 3 mm. Calyx ca. 8mm; lobes linear-lanceolate, outside appressed strigose, marginsparsely gland-tipped ciliate, apex obtuse. Corolla purple, ca.1.5 cm, bent to 90º and abruptly inflated at middle. Stamens included;filaments ca. 1.2 cm, hirsute at base; anther thecae oblong,connective gland-tipped hirsute; staminodes 2. Ovary ca.2 mm, glabrous; style ca. 1 cm, glabrous; stigma slightly 2-cleft. Capsule and seeds not seen.Yunnan [Vietnam].7. Cystacanthus vitellinus (Roxburgh) Y. F. Deng, comb. nov.糙叶火焰花 cao ye huo yan huaBasionym: Justicia vitellina Roxburgh, Fl. Ind. 1: 117.1820; Endopogon vitellinus (Roxburgh) Nees; J. asperula(Nees) G. Loddiges; Phlogacanthus asperulus Nees; P. vitellinus(Roxburgh) T. Anderson; Sphinctacanthus siamensis C. B.Clarke.Shrubs to 1.5 m tall. Stems terete, glabrous. Petiole 1–5cm; leaf blade broadly ovate to oblong-lanceolate, 10–30 × 6–11 cm, abaxially slightly pubescent or glabrescent, adaxiallyglabrous or slightly pubescent, secondary veins 9–11 on eachside of midvein, base attenuate and decurrent onto petiole, marginslightly undulate, apex acute to acuminate. Thyrses 10–20cm; bracts minute; bracteoles linear, ca. 1.5 × 0.5 mm, pubescent.Pedicel 1–3 mm. Calyx ca. 6 mm, pubescent; lobeslinear-lanceolate. Corolla purplish red or white, 1.2–1.8 cm,bent to 90º and abruptly inflated at middle, outside pubescent.Stamens ca. 1 cm; filaments hirsute at base; anther thecae oblong,ca. 2 mm, connective glabrous; staminodes very small.Ovary glabrous; style 1–1.5 cm, glabrous; stigma slightly 2-cleft. Capsule clavate, ca. 2 cm, 8-seeded. Seeds ovate in outline,ca. 2.5 × 2 mm.Forests; 200–1100 m. Yunnan [Bhutan, India, Myanmar].8. Cystacanthus colaniae (Benoist) Y. F. Deng, comb. nov.广西火焰花 guang xi huo yan huaBasionym: Phlogacanthus colaniae Benoist, Notul. Syst.(Paris) 5: 109. 1936.Shrubs to 1 m tall. Stems terete, pubescent, soon glabrescent.Petiole 1–5 cm, puberulent; leaf blade ovate to ovate-lanceolate,(5–)12–30 × (2–)5–12 cm, abaxially slightly pubescent,adaxially glabrous or puberulent along veins, secondary veins8–17 on each side of midvein, base shortly attenuate, marginentire or slightly crenate, apex shortly acuminate. Thyrses 10–20 cm; cymes few flowered; bracts triangular, 1.5–2 × 0.7–1mm, pubescent; bracteoles linear, ca. 1 × 0.5 mm, pubescent.Pedicel 1–2 mm, pubescent. Calyx ca. 3 mm, pubescent; lobeslinear. Corolla ca. 1.2 cm, ventricose, bent to 90º and abruptlyinflated at middle, outside slightly grayish white pubescent,inside pubescent on lobes; lobes oblong-ovate, ca. 5 × 3 mm,unequal, apex acute. Stamens 2; filaments hirsute at base, ca. 6mm; anther thecae oblong, ca. 2 mm, connective black and glabrous;staminodes filiform, very short, hirsute. Ovary oblong,ca. 2 mm, glabrous; style ca. 8 mm, glabrous; stigma slightly 2-cleft. Capsule 1.8–2 cm, glabrous. Seeds ovate in outline, ca. 2.5× 2 mm.Limestone hills; 200–500 m. Guangxi, Hainan, Yunnan [Vietnam].

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