Synopsis of the Genera and Suprageneric Taxa of Euphorbiaceae

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SYNOPSIS OF THE GENERA GradyL. Webster2 AND SUPRAGENERICTAXA OF EUPHORBIACEAE1

ABSTRACT

This revisedsynopsis of the taxa of Euphorbiaceaerecognizes, and provideskeys to, 5 subfamilies,49 tribes,and 317 genera.Two new tribes,Croizatieae and Podocalyceae,are proposed,as wellas fournew subtribes,Leptopinae, Podocalycinae,Pycnocominae, and Tetracoccinae;five taxa are reducedto subtribalstatus. One genus(Ophellantha) is reducedto a section; 14 new binomialcombinations are proposed.Lectotypifications are providedfor almostall taxa not previouslytypified.

The synoptic classification given here provides Euphorbiaceae A. L. de Jussieu, Gen. P1. 384. keys to and descriptions of the taxa of Euphorbi- 1789. TYPE: Euphorbia L. aceae at the subfamilial,tribal, and subtriballevels. It represents an extension and, in places, a con- Monoecious or dioecious trees, shrubs, or herbs siderable revision of the synoptic classification in (some climbing or twining); stems sometimes suc- which the five subfamilies were first recognized culent and/or with latex. Leaves alternate or op- (Webster, 1975). Within the tribes and subtribes, posite (rarely whorled); stipules free (less commonly keys are provided to the genera, and these are connate or absent), deciduous or persistent; leaf enumerated with citation of important synonyms. blades pinnatelyor palmately veined, entire to den- Keys are intended to provide the salient diagnostic tate or palmately lobed or compound; indumentum characters of taxa, but do not necessarily allow for simple to stellate or lepidote (sometimes absent). exceptions; in some instances, they may be of lim- Inflorescences terminal or axillary, basically cy- ited use in identifyingunknown specimens to genus. mose; flowerssolitary or in glomerules, these often Citations of synonyms and relevant works have grouped into spiciformor capitate thyrses or pseu- been deliberately kept brief and are not intended danthia; bracts often glandular. Flowers unisexual, to be all-inclusive; by and large, only strictlysys- actinomorphic (but pseudanthia often zygomor- tematic papers are cited. Some obscure synonyms, phic); perianth segments free or connate, valvate including most of the names proposed by Rafines- or imbricate, sometimes reduced or absent; sepals que and Otto Kuntze, have been omitted. Typifi- and petals (1-)3-6(-8), sometimes distinctlycol- cations followIndex Nominum Genericorum (Farr ored; disk present or absent, intrastaminal or ex- et al., 1979); new typificationsor alterations of trastaminal, entire to dissected; stamens (1 -)3- entries in ING are explicitly noted. Generic con- 50(-400) [always 1 in Euphorbia], hypogynous; cepts are relatively conservative; i.e., traditional filamentsfree or connate; anthers mostly 2-locular usage is followedwhere possible. In borderlinecases, and dehiscing longitudinally,introrse or extrorse; genera are enumerated as distinct even though pollen grains tectate or semi-tectate, mostly their claim to generic status may be questionable; 3-colporate (inaperturateto polytreme); gynoecium listing of taxa here is thereforenot intended to be syncarpous, ovary (1 -)2-5(-20)-locular; placen- canonical. A list of new taxa and an index to the tation axile; ovules 1 or 2 per locule, anatropous genera and higher taxa are given in the appendices or hemitropous (orthotropous in Panda), inserted to this article. beneath an obturator, crassinucellate with 2 integ-

I I expressmy gratitudeto JohnHayden and GeoffreyLevin forreviewing the treatmentof the Oldfieldioideae, LynnGillespie for tribe Plukenetieae, and Daryl Koutnikfor tribe Euphorbieae. Their suggestionshave led to some importantmodifications in circumscriptionsand interpretationsof taxa. I am also particularlyindebted to Michael Huftand GeoffreyLevin fortheir critical reviews of the manuscript,to GeoffreyLevin forcompiling the index,and to RobertRhode forhelp in formattingthe text. 2 Sectionof Plant Biology,University of California,Davis, California95616, U.S.A. ANN. MISSOURI BOT. GARD. 81: 33-144. 1994.

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uments, nucellus often beaked, embryo sac mostly lb. Locules of ovaryeach with1 ovule (except in 8-nucleate; styles free or connate, entire to lobed Dicoelia);latex present or absent;indumentum or multifid.Fruit typically a capsular schizocarp various;pollen grains binucleate or trinucleate. 3a. Milkylatex absent; laticifers(if present) with mericarps elastically dehiscent from a persis- inarticulate;leaves unlobedor lobed; in- tent columella, but sometimes baccate or drupa- dumentumsimple or stellate;petals present ceous; seeds 1 or 2 per locule (rarely 1 per fruit); or absent;pollen grains binucleate, mostly seed coat thin to indurate, sometimes with a sar- tricolporateor triporate...... III. ACALYPHOIDEAE cotesta; endosperm present or absent; embryo 3b. Latex milkyor colored(rarely absent); la- straight to curved or folded; cotyledons usually ticifersarticulate or inarticulate;leaves un- broader than radicle. lobed to compound;pollen grains binucleate or trinucleate. The family Euphorbiaceae is here construed as 4a. Latex reddishor yellowishto milky having 317 genera associated into 49 tribes in 5 (sometimesscanty or absent); latici- subfamilies. A reflection of the diversity of the fersarticulate or inarticulate;leaves unlobedto lobed or compound;indu- familyis that over 20 segregate familieshave been mentumsimple, stellate, dendritic, or proposed (Webster, 1987). Hurusawa (1954) was lepidote;bracts usually not biglandu- the firstmodern author to propose a major dis- lar at base; sepalsimbricate to valvate, mantling of the family,with his recognition of An- usuallycompletely covering anthers in tidesmataceae, Euphorbiaceae (sensu stricto), Por- bud; petalsmostly present (at least in staminateflower); pollen grains tri- antheraceae, and Ricinocarpaceae. This system, colporateor moreoften porate or in- based on inflation of the subfamilies of Pax & aperturate,with "crotonoid" pattern Hoffmann (1931), has met with little acceptance. of exinousprocesses ...... More influentialhave been the effortsof Airy ...... IV. CROTONOIDEAE 4b. Latex whitish,often caustic or toxic; Shaw, who recognized seven segregate families: laticifersinarticulate; leaves usually Androstachydaceae, Bischofiaceae, Hymenocardi- unlobed; indumentumsimple or ab- aceae, Peraceae, Picrodendraceae, Stilaginaceae, sent, never stellateor lepidote(den- and Uapacaceae (Airy Shaw, 1965, 1966). Among driticin Mabea); bractsoften biglan- modern authors,the most extreme splitteris Meeuse dular at base; sepals imbricateor reduced,anthers mostly not covered (1990), who recognizes nine segregate families, in bud; petals absent; pollen grains plus the Pandaceae. Radcliffe-Smithin Carter & tricolporate,sexine mostlyperforate- Radcliffe-Smith(1988) partiallyfollowed Airy Shaw reticulate,never with "crotonoid" in recognizing the Hymenocardiaceae and Pan- pattern. . V. EUPHORBIOIDEAE daceae, but treats Antidesma, Bischofia, and Uapaca as anomalous genera of Euphorbiaceae. Subfamily I. PHYLLANTHOIDEAE Asch- Of all these segregate taxa, only the family Pan- erson, Fl. Prov. Brandenburg 1: 59. 1864. daceae is recognized as distinctby Cronquist (1981) TYPE: PhyllanthusL. and Takhtajan (1980). Euphorbiaceae subordo Dispermae Zoll., Natuur-Ge- KEY TO THE SUBFAMILIES OF EUPHORBIACEAE neesk. Arch. Ned.-Indie2: 17. 1845. PhyllanthaceaeJ. G. Agardh,Theor. Syst. P1. 249. 1858 1a. Locules of ovaryeach with2 ovules(except in (as Phyllantheae);Klotzsch & Garcke,Monatsber. Scagea); milkylatex, intraxylary phloem, and K6nigl.Preuss. Akad. Berlin1859: 246. 1859. stinginghairs absent; indumentumsimple or rarelylepidote or dendritic;embedded foliar Trees, shrubs, or herbs; indumentum simple glandsrare; pollengrains binucleate. (rarely branched; lepidote in Hyeronima and 2a. Leaves alternate(very rarely opposite), leaves stipulate,blade simpleand unlobed(trifo- Uapaca); alternate (very rarely opposite), liolatein Bischofia); petalspresent or ab- spiral to distichous, usually stipulate; leaf blade sent; pollen grainstricolporate to porate, simple, entire (except in Drypetes and Bischofia), sexinenot withconspicuous spines (irreg- withoutembedded foliar glands. Inflorescences ax- ularly spiny in Amanoa); seeds ecarun- illary (rarely terminal), mostly racemiform or spi- culate ...... I. PHYLLANTHOIDEAE 2b. Leaves alternate,opposite, or whorled, ciform,or reduced to glomerules or solitaryflowers; stipulateor exstipulate,blade simpleor tri- bracts eglandular, mostly inconspicuous. Sepals foliolate;petals absent(except in Croiza- mostly 4-6, imbricate, sometimes connate; petals tia); pollengrains colpoidorate to porate, and disk present or absent; stamens (2-)4-8(-50), sexinespiny; seeds carunculateor ecarun- filamentsfree or culate, endosperm copious (except in united; pollen grains mostly 3-4- Hyaenanche and Picrodendron).-... colporate (rarely porate; periporate in Phyllan- .------II. OLDFIELDIOIDEAE thus), exine semitectate, rarely echinate; male ga-

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metophyte binucleate; pistillode present or absent; This large "basal" subfamily of Euphorbiaceae carpels (1-)2-5(-20); styles mostly free except at is so diverse that Meeuse (1990) proposed dividing base, usually bifid; ovules 2 in each locule, anat- it into six separate families.The 60 genera included ropous or hemitropous, nucellar beak sometimes are grouped into 10 tribes, a reduction from Web- prominent. Fruit capsular or less commonly bac- ster (1975), where 13 tribes were recognized. The cate or drupaceous; seeds ecarunculate or with work of Levin (1986a, b, c) has shown that some rudimentary caruncle; endosperm present or ab- proposed tribesneed to be incorporated as subtribes sent; cotyledons plane or folded. Base chromosome within the Antidesmeae. numbers mostly 12 or 13.

KEY TO THE TRIBES AND SUBTRIBES OF SUBFAMILY PHYLLANTHOIDEAE 1a. Leaves simple,mostly entire; styles usually bifid. 2a. Flowersin axillaryclusters (inflorescence axes less than 1 cm longexcept in some Phyllantheae);seeds withor withoutendosperm; leaves mostlywith paracytic stomata, lacking tanniniferous epidermal cells. 3a. Staminatedisk extrastaminal or absent;leaves entire;ovary 2-3(-6)-locular; fruitmostly capsular or baccate, less commonlydrupaceous. 4a. Petals present,at least in staminateflowers; sepals imbricate;wood parenchyma conspicuous, fibersmostly thick-walled; vessel element perforations scalariform or simple;ovules anatropous; endospermpresent or absent. 5a. Pollenexine finely reticulate, not coarsely spinose; stipules deciduous-- 1. WIELANDIEAE 5b. Pollenexine coarsely and deeplyreticulate or spinose;stipules persistent - 2. AMANOEAE 4b. Petals absent,or if presentthen sepals valvate, or wood and ovule charactersotherwise; endospermusually present. 6a. Sepals valvate;petals present, usually shorter than sepals -3. BRIEDELIEAE 6b. Sepals imbricate;petals absent, or ifpresent then often as longas sepals- 4. PHYLLANTHEAE 3b. Staminatedisk intrastaminal;leaves oftendentate; wood fibersthick-walled, not septate; fruit drupaceous;ovary 1-2-locular -5. DRYPETEAE 2b. Flowersin axillaryor terminalspicate or racemoseinflorescences (axis usuallyover 1 cm long;capitate in Uapaca); petals absent (or if present,inflorescence terminal); leaves withstomata paracytic or anomocytic;seeds withendosperm. 7a. Fruitsnot winged;leaves withtanniniferous epidermal cells, not granulose-glandular;floral disk presentor absent;pollen 3-colporate; styles usually bifid .6. ANTIDESMEAE 7b. Fruitswinged, samaroid; leaves granulose-glandular,without tanniniferous epidermal cells; inflo- rescencesamentlike; floral disk absent; pollen zonoporate; styles unlobed - 7. HYMENOCARDIEAE lb. Leaves trifoliolate;inflorescence paniculate; flowers apetalous, floral disk absent; styles entire; fruits baccate ------8. BIS C H O F IE A E

Tribe 1. WIELANDIEAE Baillonex Hurusawa, ovules anatropous, usually with a single obturator. J. Fac. Sci. Univ.Tokyo, Sect. 3, Bot. 6: 339. Fruit capsular; seeds 1 or 2 per locule; endosperm 1954. WielandiideesBaillon, Etude Gen. Eu- present or absent; embryo with cotyledons plane phorb. 568. 1858. Phyllantheaesubtribe or folded, much longer than the radicle. & Pflanzenr.147. WielandiinaePax Hoffm., The taxa with the greatest numbers of unspe- 81): 180. 1922. TYPE: Wielandia XV (Heft cialized morphological features are gathered to- Baillon. gether in this tribe, which is unified on the basis Monoeciousor dioecioustrees or shrubs;indu- of pollen morphology (K6hler, 1965) and leaf ve- mentumsimple; leaves alternate,entire, penni- nation (Levin, 1986a, b, c). It is probably para- nerved, eglandular;stipules deciduous. Inflores- phyletic in cladistic terms, which is essentially in- cencesaxillary; flowers in glomerulesor thepistillate evitable for the basal tribe in a large family such solitary;bracts inconspicuous. Sepals mostly4-6, as the Euphorbiaceae. Relationships among the imbricate;petals usually 5; disk usually annular; nine genera appear reticulate, and it does not seem stamens5-10, freeor filamentsconnate at base; feasible to group them into subtribes. It is notable anthersdehiscing longitudinally, connective not en- that all of the genera occur in South America, larged; pollengrains 3-colporate, semitectate, re- Africa, and the Indian Ocean islands, and only ticulate;pistillode present; carpels 3-5; stylesbifid; Savia extends into the Northern Hemisphere.

KEY TO THE GENERA OF TRIBE WIELANDIEAE la. Seeds withendosperm. 2a. Stamens8-10; sepalsand petalsmostly 4, verysimilar; ovary 4-5-locular -1. Heywoodia

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2b. Stamensmostly 5; sepals and petals mostly5; ovarymostly 3-locular. 3a. Pistillatedisk entire, not glandular --2. Savia 3b. Pistillatedisk glandular-lobed --3. Gonatogyne lb. Seeds withlittle or no endosperm. 4a. Staminatedisk dissected - - -4. Petalodiscus 4b. Staminatedisk annular, sometimes lobed 5a. Ovary 3- or 4-locular. 6a. Monoecious. 7a. Leaves coriaceous;petals longer than sepals; stamens free except at base - - 5. Blotia 7b. Leaves chartaceous;petals shorter than sepals; stamens connate into a column 6. Actephila 6b. Dioecious;leaves coriaceous. 8a. Petals present;staminate flowers pedicellate. 9a. Stylesdilated; staminate disk glabrous; stamens free from pistillode - 7. Discocarpus 9b. Stylesslender; staminate disk pubescent; stamens adnate to pistillode 8. Lachnostylis 8b. Petalsabsent; staminate flowers subsessile -9. Chonocentrum 5b. Ovary5-locular --10. Wielandia

1. Heywoodia Sim, For. Fl. Cape Col. 326, pl. West Indies and Madagascar; and the recently 140. 1907; Pax & Hoffm.,Pflanzenr. 147. describedsect. Afrosavia Radcl.-Sm. of Kenya XV. (Heft81): 280. 1922; Hutchinson,Kew (whichas notedbelow actually belongs to Petalo- Bull. 1922: 114. 1922; Milne-Redhead,Bull. discus). A numberof othergenera of tribeWie- Jard.Bot. Brux. 27: 327, pl. X. 1957; Rad- landieae have been includedin Savia by Mueller cliffe-Smith,Fl. Trop. E. Afr.,Euphorb. 1: and otherworkers. 85, fig. 12. 1987. TYPE: Heywoodia lucens Sim. 3. Gonatogyne Muell.Arg., Fl. Bras. 11(2): 13. 1873; Pax & Hoffm.,Repert. Sp. Nov. 31: Monotypic;Uganda and Kenya southto Natal 190. 1933. Savia sect. Gonatogyne and Transkei.On the basis of foliarvenation char- ("Klotzsch") Pax & Hoffm.,Pflanzenr. 147. acters,Levin (1986b) suggesteda relationshipbe- XV (Heft81): 187. 1922. TYPE: Gonatogyne tweenHeywoodia and Astrocasia, and regarded brasiliensis(Baillon) Muell. Arg. [Amanoa ? both genera as closer to the tribe Phyllantheae. brasiliensisBaillon]. Thereare indeedsome resemblances between Hey- woodia and Astrocasia,but the former genus seems Monotypic;Brazil. the moreprimitive, and appears to me as perhaps nearestin morphologicalcharacters to the hypo- 4. Petalodiscus (Baillon)Pax, Natiirl.Pflanzen- thetical ancestor of the family.This would not fam.I. 3(5): 15. 1890. Savia sect. Petalodis- precludean affinitywith Astrocasia, whichmay cus Baillon,Etude Gen. Euphorb.571. 1858. connectthe tribesWielandieae and Phyllantheae. TYPE: Petalodiscusplatyrhachis (Baillon) Pax (lectotype,selected here; thiswas the species illustratedby Pax, loc. cit.). 2. Savia Willdenow,Sp. P1.4: 771. 1805; Muell. Arg. in DC. Prodr. 15(2): 228. 1866; Ben- A genus of fiveor six species endemicto Mad- tham,Gen. PI. 3: 270. 1880; Fawc. & Rend., agascar; combinedwith Savia by Pax & Hoffmann Fl. Jam.4: 271. 1920; Pax & Hoffm.,Pflan- (1922, 1931) and Leandri (1958), but differing zenr. 147. XV (Heft81): 181. 1922; Urban, fromSavia in being monoeciousand having ex- Repert.Sp. Nov. 28: 209. 1928; Alain, Fl. albuminousseeds. Cuba 3: 41. 1953; Webster,J. ArnoldArbor. 48: 325. 1967; Taxon 31: 535. 1982; Rad- 5. Blotia Leandri,Mem. Inst. Sci. Madag. 88: cliffe-Smith,Kew Bull. 27: 508. 1972; Fl. 240. 1957; Fl. Madag. 111: 126. 1958. Sav- Trop. E. Afr.,Euphorb. 1: 83, fig.11. 1987. ia sect. Charidia Baillon, Etude Gen. Eu- TYPE: Savia sessilifiora(Sw.) Willd. phorb. 572. 1858; Muell. Arg., DC. Prodr. sect. Charid- KleinodendronSmith & Downs,Sellowia 16: 177. 15(2): 229. 1866. Petalodiscus 1964. TYPE: Kleinodendronriosulense Smith & ia (Baillon)Pax, Natuirl.Pflanzenfam. I. 3(5): Downs[= Savia dictyocarpaMuell. Arg.]. 15. 1890. TYPE: Blotia oblongifolia(Baillon) Leandri[Savia oblongifoliaBaill.]. About25 species of America,Africa, and Mad- agascar. Threesections have beenrecognized: sect. This genusof fiveor six Madagascan species is Savia, 2 spp., the type and S. dictyocarpa of closely related to Petalodiscus and may not be Brazil;sect. Heterosavia Urb., ca. 20 spp. of the distinct.

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6. Actephila Blume, Bijdr. Fl. Ned. Ind. 581. This genus, endemic to South Africa, includes 1825; Muell. Arg., DC. Prodr. 15(2): 221. a single variable species or, according to some 1866; Baillon, Adansonia I. 6: 231. 1866; workers (e.g., Dyer), two species. Although com- Bentham,Gen. P1. 3: 269. 1880; J. J. Smith, bined with Discocarpus by Pax & Hoffmann Add. Cogn. Fl. Arb. Jav. 12: 45. 1910; Pax (1922), it differsin a number of characters. & Hoffm.,Pflanzenr. 147. XV (Heft81): 191. 1922; AiryShaw, Kew Bull. 25: 496. 1971; 9. Chonocentrum Pierre ex Pax & Hoffmann, Kew Bull. Add. Ser. 8: 21. 1980. TYPE: Ac- Pflanzenr.147. XV (Heft 81): 205. 1922; tephilajavanica Miq. Jablonski, Mem. New York Bot. Gard. 17: 121. 1967. TYPE: Chonocentrumcyathopho- Dalzell,Hook. J. Bot. Kew Gard. Misc. Anomospermum rum (Muell. Arg.) Pax & Hoffm. [Drypetes 3: 228. 1851 (nonMiers, 1851). TYPE: Anomosper- mumexcelsum Dalz. [= Actephilaexcelsa (Dalz.) cyathophora Muell. Arg.]. Muell.Arg.]. A monotypic genus of Amazonian South Amer- An Asian/Australasiangenus of ca. 20 de- ica, regarded by Pax & Hoffmann as related to scribedspecies, stillpoorly understood taxonomi- Discocarpus; however, the pollen is stillunstudied, cally.The positionof the genushas been variously and its relationshipsmust be regarded as uncertain. interpreted.Pax & Hoffmann(1922) associatedit withAmanoa, whichhas similarflowers but very 10. Wielandia Baillon, Etude Gen. Euphorb. differentpollen. K8hler (1965) pointedout a pal- 568. 1858; Bentham, Gen. P1. 3: 270. 1880; ynologicalsimilarity to Andrachne,and Levin(1986 Hemsley, Hook. Ic. P1. 29: t. 2813. 1906; a, b, c) founda resemblancein leaf venationto Pax & Hoffm.,Pflanzenr. 147. XV (Heft 81): thatgenus. However, the exalbuminousseeds with 181. 1922; Leandri, Notul. Syst. (Paris) 7: plicateembryo of Actephila wouldappear to rule 190. 1939; Fl. Madag. 111(1): 135. 1958. out a close associationwith Andrachne. It is worth TYPE: Wielandia elegans Baillon. notingthat Mennega (1987) recommendedassign- Monotypic; restricted to Madagascar and the ment of Actephila to the Wielandieae. For the Seychelles. Although combined with Savia by present,it seems best to retainActephila in the Mueller, Wielandia appears closer to Petalodiscus tribeWielandieae. by virtue of its exalbuminous seeds and monoecious flowerproduction. 7. Discocarpus Klotzsch,Arch. Naturgesch.7: 201. 1841; Muell. Arg., DC. Prodr. 15(2): Tribe 2. AMANOEAE (Pax & Hoffmann)Web- 223. 1866; Fl. Bras. 11(1): 11, t. 2. 1873; ster, Taxon 24: 594. 1975. Phyllantheae sub- Benth.,Gen. P1. 3: 269.1880; Pax & Hoffm., tribe Amanoinae Pax & Hoffm., Pflanzenr. Pflanzenr.147. XV (Heft 81): 202. 1922; 147. XV (Heft 81): 190. 1922. TYPE: Ama- Mem. New York Bot. Gard. 17(1): Jablonski, noa Aublet. 84. 1967. TYPE: Discocarpus essequiboensis Klotzsch. Monoecious trees; indumentumsimple (extremely scarce to absent); leaves alternate, simple, entire, Accordingto the recenttreatment of Jablonski, penninerved, eglandular; stipules persistent,some- the fivespecies of Discocarpus are restrictedto times adnate to petiole. Flowers in glomerules,these the Amazonian/Guiananregion of SouthAmerica. sometimes forming spikes; bracts conspicuous or The phylogeneticposition of Discocarpus is some- inconspicuous. Sepals and petals 5, imbricate; sta- whatuncertain. Pax & Hoffmann(1922) assigned minate disk annular or dissected; stamens 5, free; it to a separatesubtribe (including Lachnostylis); anthers introrse; pollen grains tricolporate, very Ko1hler(1965) suggestedit was close to Amanoeae coarsely reticulate or irregularlyspinose; pistillode and Bridelieae;and Mennega (1987) regardedit prominent; pistillatedisk cupular; ovary 3-locular; as out of place in the Wielandieae. ovules anatropous; styles twice bifid or dilated, emarginate; fruita tardilydehiscent capsule; seeds 8. Lachnostylis Turczaninov,Bull. Soc. Imp. 1 per locule; endosperm present, scanty or absent; NaturalistesMoscou 19: 503. 1846; Muell. cotyledons massive but not plicate, much longer DC. Prodr.15(2): 224. 1866; Bentham, Arg., than radicle. Hook. Ic. P1. 13: 61, t. 1279. 1879; Gen. P1. 3: Dyer,Gen. S. Afr.Fl. P1. 1: 309. 1975. As here interpreted, the tribe Amanoeae in- TYPE: Lachnostyliscapensis Turcz.[= Lach- cludes the genera Amanoa and Pentabrachion. In nostylishirta (L.) Muell. Arg.]. the original circumscriptionof the tribe (Webster,

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1975), Actephila was also included. However, ev- leaves alternate, entire, pinnately veined, without idence from pollen morphology (K6hler, 1965), laminar glands; stipules usually deciduous. Flowers leaf venation (Levin, 1986a, b, c), and wood anat- in bracteate glomerules, these axillary or in spikes. omy (Mennega, 1987) argues against a close as- Sepals mostly 5, valvate; petals 5, imbricate or sociation of Amanoa and Actephila. valvate, usually smaller than the sepals, sometimes dentate or reduced; disk annular or dissected; sta- KEY TO THE GENERA OF TRIBE AMANOEAE mens mostly 5, filaments connate into a column; la. Stipulesnot intrapetiolar;floral bracts incon- anthers introrse; pollen grains tricolporate, retic- spicuous,not indurate; styles slender, twice bi- ulate or striate; pistillode present; pistillate sepals fid;fruit thin-walled, seeds 2 per locule, endo- 5, imbricate or valvate, deciduous; disk annular or 1 spermpresent ...... 1. Pentabrachion cupular; ovary 2- or 3-(rarely 4-)locular; ovules lb. Stipules? intrapetiolar;floral bracts conspicuous, indurate;styles dilated, emarginate; fruit anatropous; styles 2- or 4-fid; fruit capsular or thick-walled,seeds 1 per locule; endospermab- drupaceous, 1-3-celled; seeds with endosperm; sent ...... 12. Amanoa cotyledons much longer and broader than the radicle. 11. Pentabrachion Muell. Arg., Flora 47: 532. The has traditionally been considered to 1864; DC. Prodr. 15(2): 223. 1866 (as Pen- tribe comprise two closely related genera, Cleistanthus tabrachium); Pax & Hoffm.,Pflanzenr. 147. and Briedelia. However, the diagnostic characters XV (Heft 81): 188. 1922. TYPE: Pentabra- are not entirely satisfactory (as indicated, for ex- chion reticulatum Muell. Arg. ample, in the complicated distinctions in the key Monotypic; West Africa (Cameroon, Gabon). of Leonard, 1962a) and generic delimitation re- Levin (1986a, b, c) suggested a possible relation- quires furtherstudy. Baillon (1858) regarded Brie- ship to Dicoelia on the basis of foliar venation delia as being close to Amanoa, and K6hler (1965) characters, but this is not supported by pollen char- saw resemblances between pollen of Amanoa and acters (K6hler, 1965). Indeed, K6hler associated the African species of Cleistanthus; followingthe Pentabrachion with the Amanoeae and Bridelieae latter's suggestion, it would be possible to associate because of its coarsely reticulate pollen grains. It the Amanoeae and Bridelieae as subtribes of a appears that this palynological similaritymay be a single tribe. Nevertheless, neither the floral nor better guide to affinity,so Pentabrachion is here pollen characters of the Bridelieae appear suffi- placed in the same tribewith Amanoa, despitethe ciently close to Amanoa to associate them in the obviousdifferences between the genera. same tribe.

12. Amanoa Aublet,Fl. GuianeFr. 256, pl. 101. KEY TO THE GENERA OF TRIBE BRIEDELIEAE 1775; Muell. Arg., DC. Prodr. 15(2): 219. la. Ovarymainly of 3 carpels(rarely 2 or 4); fruits 1866; Bentham,Gen. P1. 3: 268. 1880; Bail- capsular,lobed, mainly3-locular with 2 seeds lon, in Grandidier,Atlas Hist. Madag. P1. tt. per locule; leaf ventationmainly reticulate. 210, 211. 1892; Pax & Hoffm.,Pflanzenr...... 13. Cleistanthus lb. Ovary mainlyof 2 carpels (rarely 3); fruits 147. XV (Heft 81): 195. 1922; Jablonski, drupaceousor rarelycapsular, unlobed, 1- or Mem. New York Bot. Gard. 17: 82. 1967; 2-locularwith 1 seed per locule; leaf venation Webster,Ann. MissouriBot. 54: 1968; Hay- mainlypercurrent .14. Briedelia den, Brittonia4: 260. 1990. TYPE: Amanoa guianensis Aublet. 13. Cleistanthus Hooker f. ex Planchon, Hook. A mainlyneotropical genus of 16 species; the Ic. PI. 8: t. 779. 1848; Muell. Arg., DC. three Africanspecies have been referredto an Prodr. 15(2): 503. 1866; Bentham, Gen. PI. endemicsection. 3(1): 268. 1880; Jablonski, Pflanzenr. 147. VIII (Heft 65): 8. 1915; Gagnepain, Fl. In- Tribe 3. BRIEDELIEAE Muell.Arg., Bot. Zeit. dochine 5: 482. 1926; Croizat, J. Arnold Ar- 22: 324. 1864 (as Bridelieae).Tribe Biovu- bor. 23: 38. 1942; Leandri, Fl. Madag. 111: latae sect.Bridelieae Thwaites, Enum. P1. Zeyl. 181. 1958; Leonard, Bull. Jard. Bot. Brux. 279. 1861. Antidesmataceaesubfam. Bridel- 30: 421. 1960; Fl. Congo 8(1): 5. 1962; ioideae Hurusawa,J. Fac. Sci. Univ. Tokyo, Radcliffe-Smith,Fl. Trop. E. Afr., Euphorb. Sect. 3, Bot. 6: 321. 1954. TYPE: Briedelia 130. 1987. TYPE: Cleistanthus polystachyus Willd. Hook. f.

Monoecious (rarely dioecious) trees or shrubs; NanopetalumHassk., Versl.Meded. Afd.Natuurk. Kon.

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Akad. Wetensch.Amsterdam 4: 140. 1856. TYPE: An Old World genus of more than 60 species, Nanopetalummyrianthum Hassk. [= Cleistanthus the majority Asian. I agree with Pfeiffer(1874) myrianthus(Hassk.) Kurz]. and Chapman (1991) in adopting Willdenow's orig- Leiopyxis Miquel,Fl. Ned. Ind., Erste Bijv. 445. 1861. TYPE: Leiopyxis sumatrana Miq. [= Cleistanthus inal spelling for the genus; Sprengel's correction sumatranus(Miq.) Muell. Arg.]. to match the name of the botanist honored (S. E. LebidieropsisMuell. Arg.,Linnaea 32: 80. 1863. TYPE: Bridel) is contrary to the current rules of nomen- Lebidieropsiscollina (Roxb.) Muell. Arg.[= Cleis- clature. tanthuscollinus (Roxb.) Benth.]. Schistostigma Lauterbach, Fl. Deutsches Schutzgeb. Nachtr.299. 1905. TYPE: Schistostigmapapuan- Anal. um Laut. [= Cleistanthuspapuanus (Laut.) Jabl.]. Tribe 4. PHYLLANTHEAE Dumortier, Paracleistus Gagnepain,Bull. Soc. Bot. France 70: 499. Fam. P1. 45. 1829. Euphorbiaceae sect. Phyl- 1923. TYPE: Paracleistus subgracilis Gagnep. [= lantheae Blume, Bijdr. 578. 1826 (nom. il- Cleistanthussaichikii Merr. & Chun; lectotype, leg.). Ordnung Phyllanthaceae Kl. & Gcke., designatedby Wheeler,1975.]. Monatsber. Kbnigl. Akad. Wiss. Berlin 1859: GodefroyaGagnepain, Bull. Soc. Bot. France 70: 435. 1923. TYPE: Godefroyarotundata (Jabl.) Gagnep. 246. 1859. TYPE: PhyllanthusL. [= Cleistanthusrotundatus Jabl.]. Monoecious or dioecious trees, shrubs, or herbs; leaves alternate, entire, pinnately veined, without A large and rather diverse genus of over 100 laminar glands; stipules persistent or deciduous. species, of which about 30 occur in Africa/Mad- Flowers in axillary racemes or glomerules, bracts agascar and the rest in tropical Asia, from India inconspicuous. Staminate sepals 4-6, free or con- to northernAustralia and Melanesia. Both Kohler nate, imbricate; petals present or absent; disk ex- (1965) and Levin (1986a) found Cleistanthus to trastaminal,dissected or entire (rarely absent); sta- be heterogeneous on the basis of pollen and leaf mens (2-)3-5(-10), filaments free or connate; characters, respectively; clearly, generic limits in anthers mostly extrorse; pollen grains prolate to the tribe need to be critically reevaluated. oblate, mostly 3-colporate (sometimes stephanocolporate or porate); pistillode present or absent; 14. Briedelia Willdenow, Sp. P1. 4(2): 978. pistillate sepals mostly 5, imbricate, persistent or 1805; Sprengel, Anleit. Kenntn. Gew. 2(2): deciduous; disk cupular to lobed (less commonly 887. 1818 (as Bridelia); Muell. Arg., DC. dissected or absent); ovary mostly 3-5-locular; Prodr. 15(2): 493. 1866; Pfeiffer,Nomencl. ovules anatropous or hemitropous; styles bifid or Bot. 1(1): 467. 1874; Bentham, Gen. P1. 3(1): entire(rarely multifid).Fruit usually capsular (rare- 267. 1880; Gehrmann, Bot. Jahrb. 41(Beibl. ly baccate or drupaceous); seeds 1 or 2 per locule; 95): 1. 1908; Hutchinson, Fl. Trop. Afr. 6(1): endosperm present; embryo with cotyledons broad- 611. 1912; Jablonski, Pflanzenr. 147. VIII er than and at least equal in length withthe radicle. (Heft 65): 54. 1915; Leandri, Fl. Madag. As here interpreted,tribe Phyllantheae is con- 111: 192. 1958; Leonard, Fl. Congo 8(1): siderably narrower in scope than the circumscrip- 27. 1962; Radcliffe-Smith,Fl. E. Trop. Afr., tion of Pax & Hoffmann(1922), but broader than Euphorb. 120. 1987; Chapman, Austral. P1. that of Webster (1975). The relationships of some Name Index 476. 1991. TYPE: Briedelia of the taxa (e.g., Astrocasia, Leptopus) point to- scandens (Roxb.) Willd. [Clutyia scandens ward the Wielandieae, but Phyllantheae are ana- Roxb.; lectotype, selected here; of the 3 spe- tomically more advanced in such characters as cies included in the original description of septate fibers and scarcity of scalariform perfora- Willdenow, it is the only one placed in subg. tions. The inclusion of petaliferous and apetalous Briedelia by Gehrmann (1908) and of which genera together seems justified by similarities in Willdenow saw herbarium material]. pollen (Punt, 1962; K6hler, 1965) and some of Candelabria Hochstetter,Flora 26: 79. 1843. TYPE: the leaf venation characters pointed out by Levin Candelabria micranthaHochst. [= Briedelia mi- (1986a). crantha (Hochst.) Baillon]. Subdivision of this heterogeneous tribe into sub- Neogoetzea Pax, Bot. Jahrb.28: 419. 1900. TYPE: Neo- goetzea brideliifoliaPax [= Briedelia brideliifolia tribes is stillcontroversial; the present classification (Pax) Fedde]. with six subtribes must be considered provisional. GentiliaBeille, Compte Rend. Acad. Sci. Paris 114: 1294. The distinction between anatropous and hemitro- 1907. TYPE: Gentilia hygrophilaBeille [= Brie- pous ovules, pointed out by Baillon (1858), appears delia ndellensisBeille]. systematicallysignificant even though not all gen- TzellemtiniaChiovenda, Ann. Bot. Roma 9: 55. 1911. TYPE: Tzellemtinia nervosa Chiov. [= Briedelia era have yet been carefullystudied, and there may scleroneuraMuell. Arg.]. prove to be transitional stages.

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KEY TO THE SUBTRIBESOF TRIBE PHYLLANTHEAE fromMexico and Cuba south to Bolivia and eastern 1a. Petalspresent; ovules anatropous; seeds smooth. Brazil. 2a. Pistillatesepals deciduous; staminalcolumn adnate to peltatepistillode .... Subtribe 4b. LEPTOPINAE Webster, subtribe ...... 4a. Astrocasiinae nov. TYPE: 2b. Pistillatesepals persistent;stamens free or Leptopus Dcne. connatebelow ...... 4b. Leptopinae Stipulaepersistentae; petala conspicua;stamina 5, fi- lb. Petals absent(except in some Andrachninae); lamentalibera vel connata;pistillodio trifidum; sepala pis- ovulesanatropous or hemitropous; seedssmooth tillatapersistentia; ovarium 3-loculare, ovulis anatropis; or sculptured. fructuscapsularis; semina laevia. 3a. Ovules anatropous;pistillode present. 4a. Staminatedisk annular; pollen grains Dioecious shrubs or herbs; stipules persistent; reticulate;seed coat dullor sculptured flowersin axillary glomerules; staminate sepals 5; ...... 4c. Pseudolachnostylidinae 4b. Staminatedisk dissected; pollen grains disk annular or dissected; stamens 5, filamentsfree verruculose;seed coat blackish, smooth or connate below; pollen grains 3-colporate, retic- ...... 4d. Securineginae ulate; pistillode 3-fid; pistillate sepals 5, persistent; 3b. Ovules hemitropous;staminate disk an- disk annular or dissected; ovary 3-locular; ovules nularor dissected;pistillode present or absent. anatropous; styles bifid;fruit capsular; seeds 2 per 5a. Petals present;stamens 5, free; sta- locule, testa smooth; cotyledons much longer than minatedisk dissected .4e. Andrachninae radicle. 5b. Petalsabsent; stamens 2-10 or more, free or connate; staminatedisk an- This subtribeincludes only the single genus Lep- nularor dissected...... 4f. Flueggeinae topus, which was submerged in Andrachne by Mueller (1866), Bentham (1880), and Pax & Hoff- Subtribe 4a. ASTROCASIINAE Webster,Syst. mann (1922). However, Leptopus differsfrom An- Bot. 17: 315. 1992. TYPE: Astrocasia Rob. drachne (sensu stricto) in its anatropous ovules, & Greenm. and appears to represent a connecting link between Astrocasia and Dioecious (or subdioecious) trees or shrubs; stip- Andrachne. It therefore appears best to assign it to a ules deciduous; flowersin axillary glomerules; sta- separate subtribe. minate sepals 5; disk annular; stamens 5, filaments 16. Leptopus Decaisne, in connate into a column; pollen grains 3-colporate, Jacquemont, Voy. Inde Bot. 4: 155. reticulate; pistillode peltate; pistillate sepals 5, de- 1836; Pojarkova, Not. Syst. Herb. Inst. Bot. ciduous; disk annular; ovary 3-locular; ovules anat- Acad. Sci. USSR 20: 269. 1960; Li ropous; styles bifid; fruits capsular; seeds 1 or 2 Ping-T'ao, Notes Roy. Bot. Gard. Edinburgh 40: TYPE: per locule, smooth; cotyledons much longer than 467. 1983. Leptopus radicle. cordifolius Dcne. Lepidanthus Nuttall,Trans. This monogeneric tribe includes only the type Amer. Phil. Soc. 5: 175. 1837 (non Lepidanthus Nees, 1830). TYPE: Lepi- genus Astrocasia, which was included in tribe Wie- danthus phyllanthoidesNutt. (loc. cit.) [= Lep- landieaeby Webster(1975), followingPax & Hoff- topus phyllanthoides (Nutt.) Webster, comb. mann (1922). However, Pax & Hoffmannreferred nov.]. Astrocasia to the Wielandieae on the basis of a ThelypetalumGagnepain, Bull. Soc. Bot.France 71: 876. 1924. TYPE: Thelypetalumpierrei Gagnep. [= Lep- mistaken interpretationof the floral morphology. topus australis (Zoll.) Pojark.]. Pollen studies of K6hler (1965) and leaf venation ChorisandrachneAiry Shaw, Kew Bull. 23: 40. 1969. studies of Levin (1986a, b, c) suggest, on the TYPE: Chorisandrachnediplosperma Airy Shaw [= contrary, that Astrocasia shows more similarities Leptopus diplospermus (Airy Shaw) Webster, comb. to the tribe Phyllantheae, particularlyto genera of nov.]. subtribes Leptopinae and Pseudolachnostylidinae. A genus of ca. 10 species of widely scattered distributionin the Old World (India to China, In- 15. Astrocasia Robinson & Greenman, Bot. donesia, and tropical Australia) and North America Jahrb.36 (Beibl. 80): 19. 1905; Pax & Hoffm., (Mexico, southern U.S., and Greater Antilles). Pflanzenr.147. XV (Heft 81): 189. 1922; Webster, Syst. Bot. 17: 311. 1992. TYPE: Subtribe 4c. PSEUDOLACHNOSTYLIDI- Astrocasia phyllanthoides Rob. & Greenm. NAE Pax & Hoffmann, Pflanzenr. 147. XV [= Astrocasia tremula (Griseb.) Webster]. (Heft 81): 206. 1922. TYPE: Pseudolachnos- A neotropical genus of five species distributed tylis Pax.

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Monoecious or dioecious trees, shrubs, or un- anatropous; styles bifid; fruitcapsular; seeds 1 or dershrubs; stipules persistent or deciduous; flowers 2 per locule, testa smooth or sculptured; cotyledons in axillary glomerules or cymes, bracts inconspic- much longer than radicle. uous; staminate sepals 5, petals absent; disk an- This subtribe of six genera is heterogeneous and nular; stamens usually 5, filamentsfree or connate; possibly unnatural, as it is quite possible that Pseu- pollen grains 3-colporate, reticulate; pistillodepres- dolachnostylisand Keayodendron may not be ent; pistillate sepals 5, persistent or deciduous; closely related to the other genera. petals absent; disk annular; ovary 3-locular; ovules

KEY TO THE GENERA OF SUBTRIBE PSEUDOLACHNOSTYLIDINAE la. Fruitcapsular, 3-locular, with 3-6 seeds; staminatedisk annular. 2a. Capsule thin-walled;cymes (glomerules) not pedunculate;seeds withsculptured testa; pistillatesepals persistent. 3a. Chalazadorsal on seed; staminateflowers subsessile - - - -17. Chascotheca 3b. Chalaza ventral;staminate flowers pedicellate. 4a. Filamentsfree from pistillode. 5a. Stipulesfoliaceous, persistent; sepals broad and imbricate - -18. Zimmermannia 5b. Stipulesnarrow, deciduous; sepals linear-lanceolate, not imbricate --- 19. Zimmermanniopsis 4b. Filamentsconnate to pistillode;stipules narrow - - -- 20. Meineckia 2b. Capsule thick-walled;pistillate cymes pedunculate; seeds withsmooth testa; pistillatesepals deciduous; filamentsconnate -- --- 21. Pseudolachnostylis lb. Fruitdrupaceous, 2-locular, with one seed; staminatedisk dissected -22. Keayodendron

17. Chascotheca Urban, Symb. Ant. 5: 14. t. 3698. 1971; Brunel & Roux, Bull. Mus. 1904; Alain,Fl. Cuba 3: 44. 1953. Chaeno- Hist. Nat. Paris, IV. 4: 79. 1982; Radcliffe- theca Urban,Symb. Ant. 3: 284. 1902 (non Smith,Fl. Trop. E. Afr.Euphorb. 76. 1987. T. M. Fries, 1860). TYPE: Chascotheca neo- TYPE: Meineckia phyllanthoides Baillon. peltandra (Griseb.) Urb. CluytiandraMuell. Arg., J. Bot. 2: 328. 1864. TYPE: Three specieshave been describedin thisGreat- Cluytiandratrichopoda Muell. Arg.[= Meineckia er Antilleangenus (Cuba and Hispaniola), but trichopoda(Muell. Arg.) Webster]. PeltandraWight, Icon. PI. Ind.Or. 5(2): 24. 1852 (non Chascotheca domingensis (Urb.) Urb. is probably Peltandra Raf., 1819). NeopeltandraGamble, Fl. a synonymof the typespecies. Madras 1285. 1925. TYPE: Peltandra parvifolia Wight[= Meineckia parvifolia (Wight)Webster; 18. Zimmermannia Pax, Bot. Jahrb.45: 235. lectotype]. 1910; Hutchinson,Fl. Trop. Afr.6(1): 739. A genusof 20 specieswith a disjunctdistribution 1912; Verdcourt,Kew Bull. 9: 38. 1954; in the New World and Old World, fromMexico Poole, Kew Bull. 36: 129. 1981; Radcliffe- to Colombiaand Brazil,and fromcentral Africa to Smith,Fl. E. Trop. Afr.Euphorb. 71. 1987. Madagascar, southernIndia, Sri Lanka, and As- TYPE: Zimmermannia capillipes Pax. sam. As interpretedby Radcliffe-Smith(1987), Zim- mermanniaincludes six east Africanspecies, plus 21. Pseudolachnostylis Pax, Bot. Jahrb.23: a dubiousMadagascan species transitionalto Mei- 19. 1899; Hutchinson,Hook. Ic. PI. 31: t. neckia. 3011. 1915; Pax & Hoffm.,Pflanzenr. 147. XV (Heft 81): 206. 1922; Phillips,Gen. S. 19. Zimmermanniopsis Radcliffe-Smith,Kew Afr. PI. ed. 2, 457. 1951; Radcliffe-Smith, Bull. 45: 152. 1990. TYPE: Zimmermanniop- Fl. E. Trop. Afr.Euphorb. 80. 1987. TYPE: sis uzungwaensis Radcl.-Sm. Pseudolachnostylis dekindtii Pax. A monotypicgenus from Tanzania (Iringa Distr.), An Africangenus of six describedspecies, re- very close to Zimmermannia,and questionably duced by Radcliffe-Smith(1987) to fourvarieties distinct. of P. maprouneifoliaPax. The phylogeneticposition of Pseudolachnostylis remains uncertain, 20. Meineckia Baillon, Etude Gen. Euphorb. althoughthe disposition of Pax & Hoffmann(1922) 586. 1858; Webster,Acta Bot. Neerl. 14: in locatingit adjacent to Meineckia is followed 323. 1965; Radcliffe-Smith,Hook. Ic. P1. 37: here. However,the pedunculateinflorescence and

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massive fruitsof Pseudolachnostylis are quite dif- onlyLeandri among modern writers has presented ferent from the preceding genera, and it is not the genus as accepted here. clear that it really belongs in the same subtribe. K6hler (1965) has proposed that Pseudolachnos- Subtribe 4e. ANDRACHNINAE Muell. Arg., tylis be classified with Amanoa because of its Linnaea 34: 64. 1865 (as Andrachneae);Pax, coarsely reticulate pollen grains and large fruits. Nat. Pflanzenfam.ed. 1, 3(5): 15. 1890; Pax However, the flowersdiffer from those of Amanoa & Hoffm.,Pflanzenr. 147. XV (Heft81): 169. in lacking petals or conspicuous bracts, and in 1922. TYPE: Andrachne L. having bifidstyles; furthermore,the plants are di- Monoeciousherbs or subshrubs;stipules persis- oecious, and the stipules are deciduous. Because tent; flowersin axillaryglomerules, bracts incon- of all these differences,it seems possible that the spicuous;staminate sepals and petals 5; disk dis- similarityin pollen ornamentation represents con- sected;stamens 5, filamentsfree or connate;pollen vergence rather than close affinity. grains3-colporate, sexine reticulate or striate;pistillodepresent; pistillate sepals 5, persistent;petals 22. Keayodendron Leandri, Soc. Bot. France 5; diskannular or dissected;ovary 3-locular; ovules 105: 517. 1959. TYPE: Keayodendron bri- hemitropous;styles bifid; fruit capsular; seeds 2 deijoides(Mildbr. ex Hutch.& Dalz.) Leandri. per locule, testa sculptured;cotyledons much lon- A monotypicgenus fromtropical west Africa, ger than radicle. still poorly known. As the specific epithet suggests, The subtribeincludes only the genus Andrachne the fruits and aspect suggest Briedelia, but the (sensustricto); Mueller (1866) and mostlater work- flowersare apetalous, with imbricate sepals. Keay- ers included Leptopus withinAndrachne. Except odendron is diffidentlyreferred to this subtribe for forthe presenceof petals, species of Andrachne lack of a better alternative. resembleherbaceous species of Phyllanthus;however, it is not clear whetherthis indicatesclose Subtribe 4d. SECURINEGINAE Muell. Arg., affinity.Earlier (Webster, 1975), I associatedthe DC. Prodr. 15(2): 446. 1866. TYPE: Securi- Australian genus Poranthera with Andrachne be- nega Comm. ex Juss. cause of similarityin the pollen pointedout by Dioecious shrubs or small trees; stipules decid- K6hler(1965). However,other characters of Por- uous; flowersin axillary glomerules, bracts incon- anthera now seem to me to pointtoward affinity spicuous; staminate sepals 5; petals absent; disk withtribe Antidesmeae. dissected; stamens 5(-10), free; pollen grains 3-colporate, sexine verruculose; pistillode present; 24. Andrachne L., Spec. P1. 1014. 1753; Gen. pistillatesepals 5, persistent;disk or angled absent; P1. ed. 5, 444. 1754; Muell.Arg., DC. Prodr. ovary 3-locular; ovules anatropous; styles bifid; 15(2): 232. 1866; Bentham,Gen. P1. 3: 270. fruitcapsular; seeds 1 or 2 per locule, testa dark 1880; Kossinki,Bot. Mater.Gerb. Glavn. Bot. and smooth;cotyledons about equaling the radicle. Sada RSFSR 2: 77. 1921; Pax & Hoffm., Although Mueller's name is applied to this sub- Pflanzenr.147. XV (Heft 81): 169. 1922; tribe, its present circumscriptionis very different, Croizat,J. Wash. Acad. Sci. 33: 11. 1942; since Mueller (1866) included Hymenocardia and Vindt,Trav. Inst. Sci. Cherifien6: 4. 1953; genera of subtribe Scepinae. The distinctivepollen Pojarkova,Not. Syst. Herb. Inst. Bot. Acad. and seeds of Securinega set it apart from other Sci. USSR 20: 256. 1960; Webster,J. Arnold taxa in tribe Phyllantheae and suggest a possible Arbor. 48: 327. 1967; Radcliffe-Smith,Fl. affinitywith subfamily Oldfieldioideae. Trop. E. Afr.,Euphorb. 7. 1987; Gilbert& Thulin,Nordic J. Bot. 8: 159. 1988. TYPE: 23. Securinega Commerson ex Jussieu,Gen. P1. Andrachne telephioides L. [lectotype, des- 388. 1789 (nom. cons.); Leandri, Fl. Madag. ignatedby Small in Britton& Brown,Ill. Fl. 111: 107. 1958. TYPE: Securinega durissima N. U.S. ed. 2, 2: 453. 1913]. J. F. Gmel. (typ. cons.). Eraclissa Forsskal,Fl. Aegypt.-Arab.208. 1775. TYPE: A Malagasian genus of five closely related spe- Eraclissa hexagyna Forssk.[= Andrachne telephioides L.]. cies, the type from Reunion and Mauritius, the TelephioidesGomez Ortega, Tabulae Bot. 15. 1773.TYPE: others from Madagascar. The majority of workers Telephioidesprocumbens Moench [= Andrachne have confounded the genus with Flueggea, and telephioidesL.].

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PhyllanthideaDidrichsen, Vidensk. Medd. Naturh.For- branchlets deciduous in some taxa; flowersin ax- en. Kj0benhavn1857: 450. 1857. TYPE: Phyllan- illary glomerules (except Richeriella); sepals most- mi- thidea microphylla(Lam.) Didr.[= Andrachne ly 4-6, free or connate; stamens 2-15, free or crophylla(Lam.) Baillon]. connate; disk extrastaminal and usually dissected In the restricted circumscriptionadopted here, (less commonly cupular or absent); anthers introrse Andrachne includes about 15 species with a pri- or extrorse; pistillode present or absent; pistillate marily Tethyan distributionfrom Persia through sepals mostly 5 or 6, persistentin fruit;disk usually the Mediterranean to the West Indies, with one cupular, often lobed (sometimes dissected); carpels species (A. microphylla) disjunct from Baja Cali- mostly 3 (rarely 2 or 4-10); styles free or connate, fornia to Pacific South America. The majority of bifid or entire; ovules hemitropous; fruit usually species listed under Andrachne by most authors capsular (sometimes baccate or drupaceous); seeds are better classified in a separate genus Leptopus. usually 2 per locule; testa smooth or sculptured; endosperm copious; cotyledons broader than and Subtribe 4f. FLUEGGEINAE Muell. Arg., Lin- 1-2 times longer than the radicle. naea 34: 64. 1865 (as Fliggeae). TYPE: Flueg- A variable subtribe of nine genera with over gea Willd. 1000 species, evidentlymonophyletic (possibly ex- Tribe Phyllantheaesubtribe Euphyllantheae Muell. Arg., cluding Aerisilvaea, however). The vast majority Linnaea 34: 64. 1865 (nom.illeg.). TYPE: Phyllan- of the species in genera lacking pistillodes (except thus L. for Margaritaria and some species of Phyllan- Tribe Phyllantheaesubtribe Sauropodeae Muell. Arg., share a distinctive vegetative specialization: Linnaea 34: 64. 1865. TYPE: Sauropus B1. thus) Tribe Phyllantheaesubtribe Phyllanthinae Pax, Natiirl. leaves are reduced to scales on main axes and are Pflanzenfam.ed. 1, 3(5): 17. 1890. TYPE: Phyl- developed only on lateral deciduous floriferousaxes. lanthusL. This has been designated as "phyllanthoid branch- Monoecious or dioecious trees, shrubs, or herbs; ing" (Webster, 1956).

KEY TO THE GENERA OF SUBTRIBE FLUEGGEINAE 1a. Pistillodepresent in staminateflower (sometimes minute). 2a. Dioecious; stamens3-7; pistillatecalyx not accrescent. 3a. Flowersin axillaryclusters; staminate flowers distinctly pedicellate; seeds mostly2 per locule ------25 . Flu e g g e a 3b. Flowersin axillaryor cauliflorousracemes or panicles;staminate flowers subsessile; seeds mostly 1 per locule-26. Richeriella 2b. Monoecious;stamens 17-21; pistillatecalyx accrescent -27. Aerisilvaea lb. Pistillodeabsent; monoecious or dioecious. 4a. Seed coat withbony endotesta(not ventrallyinvaginated) and fleshyexotesta; fruits dry, irregularly dehiscent;disk annular in bothstaminate and pistillateflowers; dioecious shrubs or trees-- 28. Margaritaria 4b. Seed coat lackingbony endotesta and fleshyexotesta (or else endotestausually ventrally invaginated); fruits,when capsular, usually regularly septicidal; disk usually dissected or absentin staminateflower; monoeciousor dioecioustrees, shrubs, or herbs. 5a. Floral disk usuallypresent; seeds dry,not ventrallyinvaginated. 6a. Staminatedisk extrastaminal;stamens various, but rarely2 and introrse;cotyledons much broaderthan radicle--- 29. Phyllanthus 6b. Staminatedisk lobed, intrastaminal; stamens 2, anthersintrorse; cotyledons scarcely broader thanradicle --- 30. Reverchonia 5b. Floral disk absentor completelyadnate to calyx; pollengrains stephanocolporate; styles bifid or entire;seeds withthickened dry or fleshyexotesta, and usuallyventrally invaginated. 7a. Stylesbifid or emarginate;anthers not apiculate;ovary 3-locular. 8a. Seed-coatdry; staminate calyx ? discoid,not turbinate -- 31. Sauropus 8b. Seed-coatfleshy; staminate calyx turbinate -- 32. Breynia 7b. Stylesusually entire; anthers apiculate; seed-coat usually fleshy; ovary 3-8-locular .....33. Glochidion

25. Flueggea Willdenow,Sp. PI. 4: 637. 1806; AcidotonP. Browne,Civ. Nat. Hist. Jamaica335. 1756 TYPE: Adelia Bentham,Gen. PI. 3: 276. 1880; Webster, (nom. rej.; non AcidotonSw., 1788). L. [= Flueggea acidoton (L.) Webster]. Brittonia acidoton Allertonia3: 273. 1984; Hayden, Bessera Sprengel,PI. MinusCogn. Pugillus2: 90. 1815. 39: 268. 1987. TYPE: Flueggea leucopyrus TYPE: Bessera inermisSprengel [prob. = Flueggea Willd. virosa (Roxb. ex Willd.) Voigt].

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Geblera Fischer& Meyer,Index Sem. Hort. Petropol. ProsorusDalz., J. Bot. Kew Misc. 4: 345. 1852. TYPE: 1: 28. 1835. TYPE: Geblera suffruticosa(Pallas) Prosorus indicus Dalz. [= Margaritaria indica Fischer& Meyer[= Flueggea suffruticosa(Pallas) (Dalz.) AiryShaw]. Baillon]. ZygospermumThw. ex Baillon, Etude G6n. Euphorb. Colmeiroa Reuter, Bibliot.Univers. Geneve 38: 215. 620. 1858. TYPE: Zygospermumzeylanicum Thw. 1842. TYPE: Colmeiroabuxifolia Reuter [= Flueg- ex Baillon[= Margaritaria cyanosperma(Gaert- gea tinctoria(L. in Loefl.)Webster]. ner) AiryShaw]. Pleiostemon Sonder, Linnaea 23: 135. 1850. TYPE: WurtziaBaillon, Adansonia I. 1: 186. 1861. TYPE: Wurt- Pleiostemonverrucosum (Thunb.) Sonder [= Flueg- zia tetracoccaBaillon [= Margaritaria tetracocca gea verrucosa(Thunb.) Webster]. (Baillon)Webster]. Neowawraea Rock, Indig. Trees Haw. I. 243. 1913. Calococcus Kurz ex Teijsm.& Binn.,Nat. Tijdschr.Ned. TYPE: Neowawraeaphyllanthoides Rock [= Flueg- Ind. 27: 48. 1864. TYPE: Calococcus sundaicus gea neowawraea Hayden]. Kurz ex Teisjm. & Binn. [= Margaritaria indica (Dalz.) AiryShaw]. A genus of 14 species, widespread in the tropics and temperate eastern Asia, but with a disjunct A genus of 14 species occurring in tropical relict distribution.The genus was merged with Se- America, Africa, Asia, and Australia. curinega by Mueller (1866), who was followed by many subsequent workers.However, Baillon (1858), 29. Phyllanthus L., Sp. P1. 981. 1753; Gen. Bentham (1880), and J. D. Hooker (1887) main- P1. ed. 5, 422. 1754; Baillon,Adansonia I. tained Flueggea as distinct; and indeed it is not 1: 24. 1860; Muell. Arg.,DC. Prodr. 15(2): closely related to Securinega, as indicated by both 274. 1866; Bentham,Gen. P1. 3(1): 272. pollen and seed characters. 1880; Hutchinson,Fl. Trop. Afr.6(1): 692. 1912; Beille, Fl. Indochine 5: 571. 1927; Pax & Hoffm., Natiirl. Pflanzenfam. ed. 2, 26. Richeriella Pax & Hoffmann, Pflanzenr. 19c: 60. 1931; Webster, J. Arnold Arbor. 147. XV (Heft 81): 26. 1922; Natiirl. Pflan- 48: 332. 1967; Brittonia22: 44. 1970; Web- zenfam. ed. 2, 19c: 48. 1931; Henderson, ster & Airy Shaw, Kew Bull. 26: 85. 1971; Garden Bull. Str. Settlem. 7: 122, t. 32. 1933; Brunel& Roux, Willdenowia11: 69. 1981; Airy Shaw, Kew Bull. 25: 489. 1971. TYPE: Bull. Mus. Hist. Nat. IV. 3B: 185. 1981; Richeriella gracilis (Merr.) Pax & Hoffm. NordicJ. Bot. 4: 469. 1984; Radcliffe-Smith, [Baccaurea gracilis Merr.]. Fl. Trop. E. Afr.Euphorb. 9. 1987; Rossignol A genus of two species native to southern China, et al., Amer. J. Bot. 74: 1853. 1987; San- the Philippines, Malaya, and Borneo. It is very tiago,Bradea 5(2): 44. 1988; Lebrun& Stork, close to Flueggea. Enum. P1. Afr.Trop. 1: 232. 1991; Schmid, Fl. Nouv.-Caled. 14(2): 31. 1991. TYPE: Phyl- lanthus niruri L. [lectotype, designated by 27. Aerisilvaea Radcliffe-Smith,Kew Bull. 45: Small in Britton & Brown, Ill. Fl. N. U.S. ed. 149. 1990; 47: 677. 1992. TYPE: Aerisilvaea 2, 2: 453. 1913]. sylvestris Radcliffe-Smith.

A genus of two species, one from Tanzania and NiruriAdanson, Fam. P1. 2: 356. 1763. TYPE: Phyllan- thusniruri L. [lectotype,selected here]. the other from Malawi. It is anomalous withinthis Cicca L., Syst.Nat. ed. 12, 2: 621. 1767. TYPE: Cicca subtribe because of its high stamen number, and disticha L. [= Phyllanthusacidus (L.) Skeels]. its placement must be regarded as provisional until XylophyllaL., Mant.Alt. 147, 221. 1771. TYPE: Xy- the pollen can be examined. Levin (pers. comm.) lophylla latifolia L. [= Phyllanthusepiphyllan- suggested that it may be referableto the Drypeteae. thusL; lectotype]. Conami Aublet,Hist. PI. Guiane Fr. 926. 1775. TYPE: Conami brasiliensisAubl. [= Phyllanthusbrasil- iensis (Aubl.) Muell. Arg.]. 28. Margaritaria L. f., Suppl. P1. 66. 1781; GenesiphyllaL'Herit., Sert. Angl. 29. 1778. TYPE: Ge- Airy Shaw, Kew Bull. 20: 386. 1966; Fos- nesiphyllaasplenifolia L'Herit. [= Phyllanthusla- berg, Kew Bull. 33: 185. 1978; Webster, J. tifoliusSw.]. Arnold Arbor. 60: 407. 1979; Airy Shaw, CathetusLoureiro, Fl. Cochinch.607. 1790. TYPE: Cath- Kew Bull. 35: 657. 1980; Radcliffe-Smith, etusfasciculata Lour. [= Phyllanthuscochinchi- nensisSpreng.]. Kew Bull. 36: 219. 1981; Fl. Trop. E. Afr. NymphanthusLoureiro, Fl. Cochinch.543. 1790. TYPE: Euphorb. 63. 1987. TYPE: Margaritaria no- Nymphanthusruber Lour. [= Phyllanthusruber bilis L. f. (Lour.)Spreng.].

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Emblica Gaertner,Fruct. Sem. P1. 2: 122. 1790. TYPE: WilliamiaBaillon, Etude G6n. Euphorb. 559. 1858. TYPE: Emblica officinalisGaertner [= Phyllanthusem- Williamiapruinosa Baill. [= Phyllanthusdiscolor blica L.]. Poepp. ex Spr.]. Kirganelia Juss.,Gen. PI. 387. 1789. TYPE: Kirganelia Orbicularia Baillon,Etude G6n. Euphorb. 616. 1858. virginea J. F. Gmelin[= Phyllanthus virgineus TYPE: Orbiculariaphyllanthoides Baillon [nom. il- (Gmel.) Pers.]. leg., = Phyllanthusorbicularis HBK]. TricariumLoureiro, Fl. Cochinch.557. 1790. TYPE: Tri- PhyllanthodendronHemsley, Hook. Ic. PI. 26: t. 2563, carium cochinchinenseLour. [= Phyllanthusaci- 2564. 1898. TYPE: Phyllanthodendronmirabilis dus (L.) Skeels]. (Muell.Arg.) Hemsl. [= Phyllanthusmirabilis Muell. EpistyliumSwartz, Fl. Ind. Occ. 2: 1095. 1800. TYPE: Arg.]. Epistyliumaxillare (Sw.) Sw. [= Phyllanthusax- Aporosella Chodat,Bull. Herb. Boiss. II. 5: 488. 1905. illaris (Sw.) Muell. Arg.; lectotype]. TYPE: Aporosella hassleriana Chod. [= Phyllan- GeminariaRafinesque, Western Minerva 42. 1821. TYPE: thus chacoensis Morong]. Geminaria obovata Raf. [= Phyllanthus caroli- Flueggeopsis (Muell. Arg.) K. Schumann,Fl. Deutsche niensisWalt.]. Schutzgeb.Nachtr. 289. 1905. TYPE: Phyllanthus Menarda Commersonex A. Jussieu,Euphorb. Tent. 23, glaucus Muell. Arg. 109. 1824. TYPE: Menarda cryptophilaComm. ex NymaniaK. Schumann,Fl. DeutscheSchutzgeb. Nachtr. A. Juss.[= Phyllanthuscryptophilus (Comm. ex 291. 1905. TYPE: Nymania insignisK. Schum. [ A. Juss.)Muell. Arg.]. Phyllanthusschumannianus L. S. Smith]. AnisonemaA. Jussieu,Euphorb. Tent. 19. 1824 (nom. UrantheraPax & Hoffm.,Pflanzenr. 147. III (Heft47): rej.). TYPE: Anisonemareticulatum (Poir.) A. Juss. 95. 1911. TYPE: Urantherasiamensis Pax & Hoffm. [= Phyllanthusreticulatus Poir.]. [= Phyllanthusroseus (Craib & Hutch.) Beille]. Scepasma Blume,Bijdr. Fl. Jav. 582. 1825. TYPE: Sce- DimorphocladiumBritton, Mem. TorreyBot. Club 16: pasma buxifoliaBi. [= Phyllanthusbuxifolius (Bi.) 74. 1920. TYPE: Dimorphocladiumformosum (Urb.) Muell. Arg.]. Britton[= Phyllanthusformosus Urb.]. SynexemiaRafinesque, Neogenyton 2. 1825. TYPE: Syn- Ramsdenia Britton,Mem. Torrey Bot. Club 16: 72. exemia caroliniana Raf. [= Phyllanthus caroli- 1920. TYPE: Ramsdenia incrustata(Urb.) Britton niensisWalt.; lectotype]. [= Phyllanthusincrustatus Urb.]. Hexadena Rafinesque,Sylva Tell. 92. 1838. TYPE: Hex- Roigia Britton,Mem. TorreyBot. Club 16: 73. 1920. adena angustifolia(Sw.) Raf. [= Phyllanthusan- TYPE: Roigia comosa(Urb.) Britton[= Phyllanthus gustifolius(Sw.) Sw.]. comosusUrb.]. Moeroris Rafinesque,Sylva Tell. 91. 1838. TYPE: Mo- DendrophyllanthusSpencer Moore, J. Linn. Soc. Bot. eroris stipulata Raf. [= Phyllanthus stipulatus 45: 395. 1921. TYPE: Dendrophyllanthuscomp- (Raf.) Webster]. tonii Sp. Moore [= Phyllanthusmoorei Schmid]. Nellica Rafinesque,Sylva Tell. 92. 1838. TYPE: Nellica PseudoglochidionGamble, Kew Bull. 1925: 329. 1925. maderaspatensis(L.) Raf. ["maderaspatana" = TYPE: Pseudoglochidion anamalayanum Gamble Phyllanthusmaderaspatensis L.]. [= Phyllanthus anamalayanus (Gamble) Web- AsterandraKlotzsch, Arch. Naturgesch. 7(1): 200. 1841. ster,comb. nov.]. TYPE: Asterandracornifolia (HBK) Kl. [= Phyl- HexaspermumDomin, Bibl. Bot. 89: 315. 1927. TYPE: lanthuscornifolius HBK]. Hexaspermumpaniculatum Domin [= Phyllanthus Eriococcus Hasskarl,Tijdschr. Natuurl. Gesch. Physiol. clamboides(F. Muell.) Diels]. 10: 143. 1843. TYPE: Eriococcus gracilis Hassk. ArachnodesGagnepain, Notul. Syst. (Paris) 14: 32. 1950; [= Phyllanthusgracilipes Muell. Arg.]. AiryShaw, Kew Bull. 14: 469. 1960. TYPE: Arach- CeramanthusHasskarl, Cat. P1. Hort. Bogor. Alt. 240. nodeschevalieri Gagnep., loc. cit.[= Phyllanthus 1844. TYPE: Ceramanthusgracilis Hassk. [= Phyl- chevalieri (Gagnep.) Webster,comb. nov.]. lanthusalbidiscus (Ridley)Airy Shaw]. Macraea Wight,Icon. P1. Ind. Or. 5(2): 27, t. 1901, A large and very diverse genus of 750-800 1902. 1852 (non Macraea Lindley,1828). TYPE: species, about 200 of which are American,100 Macraea oblongifoliaWight [= Phyllanthussim- African,70 Madagascan,and theremainder Asian plex Retz.; lectotype]. Reidia Wight,Icon. P1.Ind. Or. 5(2): 27, t. 1903, 1904. and Australasian.Many attemptshave been made 1852. TYPE: ReidiafioribundaWight [= Phyllan- to subdividethe genus,which very possibly is un- thuswightianus Muell. Arg.; lectotype chosen here]. natural.Here, thebroad concept of Mueller (1866) ChorisandraWight, Icon. P1. Ind. Or. 6: 13, t. 1994. has been modifiedby the recognition of Glochidion 1853. TYPE: Chorisandrapinnata Wight[= Phyl- and Margaritaria as distinct. lanthuspinnatus (Wight)Webster]. DichelactinaHance, Walp. Ann.Bot. Syst. 3: 375. 1852. TYPE: Dichelactina nodicaulis Hance [= Phyllan- 30. Reverchonia A. Gray, Proc. Amer. Acad. thus acidus (L.) Skeels]. ArtsSci. 16: 107. 1880; Webster& Miller, StaurothyraxGriffith, Notul. P1. Asiat. 4: 476. 1854. Rhodora65: 200. 1963. TYPE: Reverchonia TYPE: notdesignated [ex descr.,= Phyllanthusaci- dus (L.) Skeels]. arenaria A. Gray. Hemicicca Baillon, Etude Gen. Euphorb. 645. 1858. confinedto sand dunes in TYPE: Hemicicca japonica Baillon[= Phyllanthus A monotypicgenus fiexuosus(Sieb. & Zucc.) Muell. Arg.]. the southwesternUnited States and adjacentMex-

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ico. Although it is very close to Phyllanthus, it cies,found in tropicaleastern Asia, Indonesia, trop- would appear excessively foreign if placed in that ical Asia, and the Pacific islands. The genus is genus. barelyseparable from Sauropus on the one hand, and Glochidionon the other. 31. Sauropus Blume, Bijdr. 595. 1826; Muell. Agr., DC. Prodr. 15(2): 239. 1866; Bentham, 33. Glochidion J. R. & G. Forster,Char. Gen. Gen. P1. 3: 271. 1880; Pax & Hoffm., Pflan- P1. 113, t. 57. 1776 (nom.cons.); Muell. Arg., zenr. 147. XV (Heft 81): 215. 1922; Airy Flora 1865: 369. 1865; Hook. f., Fl. Brit. Shaw, Kew Bull. 23: 42. 1969; Kew Bull. Ind. 5: 305. 1887; Beille, Fl. Indochine5: 35: 669. 1980. TYPE: Sauropus albicans (L.) 608. 1927; Alston,Ann. Roy. Bot. Gard.Per- Blume [= Sauropus androgynus (L.) Merr.; adeniya 11: 1. 1928; Pax & Hoffm.,Natiirl. lectotype, selected here]. Pflanzenfam.ed. 2, 19c: 56. 1931; AiryShaw, Kew Bull. Add. Ser. 4: 115. 1975; Kew Bull. CeratogynumWight, Icon. P1. Or. 5(2): 26. 1852. TYPE: Add. Ser. 8: 92. 1980; Kew Bull. 35: 633. Ceratogynum rhamnoides Wight [= Sauropus 1980; 36: 298. 1981; A. C. Smith,Fl. Vi- quadrangularis (Willd.) Muell. Arg.]. Diplomorpha Griffith,Notul. P1. Asiat. 4: 479. 1854. tiensisNova 2: 467. 1981. TYPE: Glochidion TYPE: Diplomorphaherbacea Griffith[= Sauropus ramiflorumJ. R. & G. Forster. bacciformis(L.) AiryShaw]. SynostemonF. Muell., Fragm. Phyt. Austral. 1: 32. Agyneia L., MantissaP1. 2: 161. 1771. TYPE:Agyneja 1858. TYPE: Synostemonramosissimus F. Muell. pubera L. [= Glochidionpuberum (L.) Muell.Arg.; [= Sauropus ramosissimus(F. Muell.) AiryShaw; lectotype,selected here]. lectotype,designated by Wheeler,1975]. Bradleja Banks ex Gaertner,Fruct. Sem. P1. 2: 127. BreyniopsisBeille, Bull. Soc. Bot. France72: 157. 1925. 1790. TYPE:Bradleja sinica Gaertner[= Glochid- TYPE: Breyniopsis pierrei Beille [= Sauropus ion sinicum(Gaertner) H. & A.]. pierrei (Beille) Webster,comb. nov.]. GynoonA. Juss.,M6m. Mus. Hist. Nat. Paris 10: 335. HeterocalymnanthaDomin, Bibl. Bot. 89: 313. 1927. 1823. TYPE:Gynoon rigidum A. Juss.[= Glochid- TYPE: Heterocalymnanthaminutifolia Domin [ ion rigidum(A. Juss.)Muell. Arg.]. Sauropus rigens(F. Muell.) AiryShaw]. GlochidionopsisBlume, Bijdr. 588. 1826. TYPE: Glo- chidionopsis sericea Blume [= Glochidion seri- ceum (Blume) Hook. f.]. A genus of about 50 species distributedfrom Glochisandra Wight,Icon. PI. Ind. Or. 5: 26. 1852. India and Sri Lanka to southern China, the Phil- TYPE:Glochisandra acuminata Wight[= Glochid- ippines, Borneo, and Australia. Airy Shaw (1980) ion lanceolarium(Roxb.) Dalz.]. has transferreda large number of Australian spe- Zarcoa Llanos, Bot. Zeit. 15: 423. 1857. TYPE:Zarcoa philippica Llanos [= Glochidion album cies of Synostemon to Sauropus, and it appears (Blanco) Boerl.]. he may have good reason to do so, although the CoccoglochidionK. Schum.,Nachtr. Fl. Schutzgeb.292. demarcation of Sauropus from Breynia is almost 1905. TYPE:Coccoglochidion ethryococcus [= Glo- as problematical as its separation from Synoste- chidionphilippicum (Cav.) C. B. Robinson]. mon. Hemiglochidion(Muell. Arg.) K. Schum., Nachtr. Fl. Schutzgeb.289. 1905. TYPE: Glochidionramif1o- rumForst. [lectotype, designated by Wheeler, 1975; 32. Breynia J. R. & G. Forster,Char. Gen. P1. the choice of Hemiglochidion hylodendronK. 145. 1776 (nom. cons.); Muell. Arg., DC. Schum. as lectotypeby Farr et al. (1979) appears invalid,as thespecies was notincluded in Glochidion Prodr. 15(2): 438. 1866; Bentham,Gen. P1. sect. Hemiglochidionby Mueller]. 3: 277. 1880; Hook f., Fl. Brit.Ind. 5: 329. TetraglochidionK. Schum.,Nachtr. Fl. Schutzgeb.291. 1887; Beille, Fl. Indochine 5: 631. 1927; 1905. TYPE: Tetraglochidiongimi K. Schum. [ Pax & Hoffm.,Natiirl. Pflanzenfam. ed. 2, Glochidiongimi (K. Schum.) Pax & Hoffm.]. 19c: 59. 1931. Foersteria Scopoli, Intr. Hist. An Asian and Australasiangenus of morethan Nat. 98. 1777. TYPE: Breynia disticha J. R. 200 species, a few extendingto Polynesia. The & G. Forster. genuswas combinedwith Phyllanthus by Mueller (1866) and Bentham(1880), but mostsubsequent Melanthesa Blume,Bijdr. 590. 1826. TYPE: Melanthesa workershave treatedit as even it racemosa Blume[lectotype, selected here; = Brey- distinct, though nia racemosa (Blume) Muell. Arg.]. representsa specializedoffshoot from ancestral taxa MelanthesopsisMuell. Arg., Linnaea 32: 74. 1863. TYPE: withinPhyllanthus. Melanthesopsislucens (Poir.) Muell.Arg. [= Brey- niaftuticosa(L.) Hook. f.; lectotype,designated by Tribe 5. DRYPETEAE (Grisebach)Hurusawa, Wheeler, 1975]. J. Fac. Sci. Univ.Tokyo, Sect. 3, Bot. 6: 334. A difficultgenus of 10-15 highly variable spe- 1954 (as tribein Antidesmataceae).Phyllan-

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theae subtribe Drypeteae Griseb., Fl. Br. W. vensis Leandri [= Lingelsheimia manongari- Ind. 31. 1859. TYPE: Drypetes Vahl. vensis (Leandri)Webster, comb. nov.].

Antidesmeae* PutranjiveaeEndl., Gen. P1. 287. 1837. As interpretedby Leonard (1951, 1962b), Lin- Ordo PutranjiveaeEndlicher, Gen. P1. Ord. Nat. gelsheimia is an African genus of two species. 287. 1837. PutranjivaceaeEndl. ex Meeuse, Eu- However, the four Madagascan species referredto phorbiaceaeAuct. Plur. 30. 1990. TYPE: Putranjiva Danguyodrypetes by Leandri clearly appear con- Wallich. The appears anomalous in the Dry- Cyclostemon6esBaillon, Etude Gen. Euphorb.561. 1858. generic. genus Phyllantheaesubtribe Cyclostemoneae Baillon ex peteae, and it is possible that its correct place is Muell. Arg.,Linnaea 34: 64. 1865. TYPE: Cyclos- in tribe Phyllantheae. Levin (pers. comm.) suggests temonBlume [= DrypetesVahl]. a possible affinityto Aerisilvaea. Dioecious trees or shrubs; leaves entire or den- Amer. 3: 49. 1810; tate, stipulate; flowersin axillary clusters, or cau- 35. Drypetes Vahl, Eclog. 453. 1866; liflorous; sepals mostly 4 or 5, imbricate; petals Muell. Arg., DC. Prodr. 15(2): 1880; Pax & absent; staminate disk intrastaminal; stamens (2-) Bentham, Gen. P1. 3: 278. XV (Heft 81): 229. 3-20(-50), free, anthers introrse or extrorse; Hoffm., Pflanzenr. 147. J. Fac. Sci. Univ. Tokyo, pollen grains 3-colporate, reticulate; pistillodesmall 1922; Hurusawa, or obsolete; pistillate sepals deciduous (persistent Sect. 3, Bot. 6: 337. 1954; Leandri, Fl. Mad- Blumea in Sibangea); disk annular or lacking; ovary usu- ag. 111(1): 144. 1958; van Steenis, 10: 140. 1960; Leonard, Bull. Jard. Bot. Brux. ally 1- or 2-locular; stylesmostly entire and dilated; Smith & Ayensu, Brittonia ovules anatropous, with massive obturator; fruit 32: 513. 1962; 220. 1964; Airy Shaw, Kew Bull. 18: capsular or drupaceous; seeds 1 per locule or fruit; 16: 272. 1965; 27: 309. 1972; Add. Ser. 4: 97. endosperm copious; cotyledons plane, broader and Kew Bull. 36: 286. 1981; A. C. Smith, somewhat longer than radicle. 1975; Fl. Vitiensis Nova 2: 455. 1981. TYPE: Dry- As circumscribedhere, tribe Drypeteae includes petes glauca Vahl. four genera, with the vast majority of species in Koelera Willd.,Sp. P1.ed. 4, 750. 1806, nom.illeg. (non the large and variable genus Drypetes. The recent KoeleriaPers., 1805). Limacia Dietr.,Nachr. Vollst. study by Hayden has demonstrated that Neowa- Lex. Gartn.4: 383. 1818, nom.illeg. (non Limacia wraea, included in the Drypeteae by Webster Lour., 1790). TYPE: Koelera laurifolia Willd. [ Drypeteslateriflora (Sw.) Kr. & Urb.]. (1975), is really a species of Flueggea. Liparena Poit. ex Leman, Dict. Sci. Nat. 27: 6. 1823. TYPE: not designated. KEY TO THE GENERA OF TRIBE DRYPETEAE CyclostemonBlume, Bijdr. 597. 1826. TYPE: Cycloste- monmacrophyllum Blume [lectotype, selected here; 1a. Ovary3-locular; fruit capsular; styles bifid, style- = Drypetes macrophylla(Blume) Pax & Hoffm.]. branchesslender ...... 34. Lingelsheimia HemicycliaWight & Arn.,Edinb. New Phil.J. 14: 297. lb. Ovarymostly 1- or 2-locular;fruit drupaceous; 1833 (as Hemicyclia). TYPE: Hemicyclia sepiaria stylesnot bifid. Wight& Arn.[= Drypetessepiaria (Wight& Arn.) 2a. Disk present;stamens mostly 4 or more; Pax & Hoffm.]. stylesstigmatiform. Icon. P1. Ind. Orient.6: t. 1992. 1853. 3a. Pistillatesepals imbricatein bud, de- AstylisWight, TYPE: Astylisvenusta Wight [= Drypetesvenusta ciduousin fruit.35. Drypetes (Wight)Pax & Hoffm.]. 3b. Pistillatesepals open in bud,persistent Sphragidia Thwaites,Hook. J. Bot. Kew Misc. 7: 269. in fruit.36. Sibangea 1855. TYPE: Sphragidia zeylanica Thwaites [ 2b. Disk absent;stamens mostly 2 or 3; styles Drypeteslongifolia (Blume) Pax & Hoffm.]. petaloid-dilated.37. Putranjiva Dodecastemon Hasskarl, Versl. Meded. Afd. Natuurk. Kon. Akad. Wet. 4: 141. 1856. TYPE: Dodecas- temon teysmaniiHassk. [= Drypetes teysmanii 34. Lingelsheimia Pax, Bot. Jahrb. 43: 317. (Hassk.) Bakh. f. & van Steenis]. 1909; Mildbraed, Deutsche Zentral-Afr. PycnosandraBlume, Mus. Bot. Lugd.-Bat.2: 191. 1856. Exped. 2: t. 54, 55. 1912; Pax & Hoffm., TYPE: Pycnosandraserrata (Blume) Blume [= Dry- Pflanzenr. 147. XV (Heft 81): 279. 1922; petes serrata (Blume) Pax & Hoffm.,nom. illeg. (non D. serrata (Krug & Urb.) = Drypetes teys- Bot. 84: 49. Leonard, Bull. Soc. Roy. Belg. manii (Hassk.) Bakh. f. & van Steenis]. 1951; Brenan,Kew Bull. 1952: 445. 1953; Cometia Du Petit Thouars ex Baillon,Etude Gen. Eu- LeConard,Bull. Jard. Bot. Brux.32: 513. 1962. phorb.642. 1858. TYPE: Cometiathouarsii Baillon TYPE: Lingelsheimia frutescens Pax. [= Drypetesthouarsii (Baillon) Leandri]. Anaua Miquel, Fl. Ned. Ind., Eerste Bijv. 410. 1861. DanguyodrypetesLeandri, Bull. Soc. Bot. France 85: TYPE: Anaua sumatrana Miq. [= Drypetes suma- 524. 1938. TYPE: Danguyodrypetesmanongari- trana (Miq.) Pax & Hoffm.].

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Laneasagum Beddome,Madras J. Lit. Sci., ser. 2, 22: A small genus of three or four Asian species 71. 1861. TYPE: Laneasagum oblongifoliaBedd. (India, Sri Lanka, Taiwan, and the Ryukyu Is- [= Drypeteslongifolia (Bi.) Pax & Hoffm.]. lands), questionably separable from Drypetes. StelechanteriaThouars ex Baillon,Adansonia I. 4: 147. 1864. TYPE: Stelechanteriathouarsiana Baillon [ Drypetesthouarsiana (Baillon)Capuron]. Tribe 6. ANTIDESMEAE (Sweet) Hurusawa, FreireodendronMuell. Arg., DC. Prodr.15(2): 245. 1866. J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: 322. TYPE: Freireodendronsessiliforum (Fr. Allem.) 1954 [as tribe of Antidesmataceae]. Ordo Sti- Muell. Arg. [= Drypetessessilifiora Fr. Allem.]. HumblotiaBaillon, Bull. Mens. Soc. Linn. Paris 1: 593. lagineae Agardh, Aphor. 199, Class. P1. 9. 1886. TYPE: Humblotiacomorensis Baillon [= Dry- 1825. Ordo Antidesmeae Sweet, Hort. Brit. petes comorensis(Baillon) Pax & Hoffm.]. ed. 2, 460. 1830. Tribe Biovulatae sect. An- Guya Frappierex Cordemoy,Fl. Ile Reunion350. 1895. tidesmeae Thw., Enum. P1. Zeyl. 289. 1861. TYPE: Guya caustica Frapp. ex Cordem.[= Dry- petes caustica (Frapp. ex Cordemoy)Airy Shaw]. TYPE: AntidesmaL. Riseleya Hemsley,J. Bot. 55: 286. 1917. TYPE: Riseleya Dioecious (rarely monoecious) trees or shrubs griffithii(Hook. f.) Hemsl. [= Drypetes indica (Muell. Arg.) Pax & Hoffm.]. (rarely herbs); leaves alternate, entire, penni- CalyptosepalumSpencer Moore, J. Bot. 63 (suppl.):91. nerved, glandular or eglandular, stipulate; inflo- 1925. TYPE: Calyptosepalum sumatranum Sp. rescences usually axillary, amentlike; sepals 3-6, Moore [= Drypetescalyptosepala AiryShaw]. free or connate, imbricate; petals reduced or absent BrexiopsisH. Perrier,Notul. Syst. (Paris) 10: 192. 1942. (except Spondianthus); disk annular, lobed, or dis- TYPE: Brexiopsis aquifolia H. Perr. [= Drypetes bathiei Capuron& Leandri]. sected; stamens 2-8, free or nearly so; anthers extrorse or introrse; pollen grains mostly prolate, A large and variable genus of ca. 200 species; 3-colporate, reticulate; pistillodepresent or absent; poorly represented in the New World, with ca. 10 carpels (1) 2 or 3 (-5); ovules anatropous; styles species; ca. 60 species in Africa and Madagascar; bifidto bipartite (rarely entire, sometimes dilated); the remainder in Asia and Australasia. Mueller fruit capsular or drupaceous; seeds mostly 1 per (1866) and Bentham (1880) recognized Cyclos- locule (or 1 per fruit); seed-coat dry or fleshy; temon and Hemicyclia as distinctfrom Drypetes, endosperm thin to copious; cotyledons mostly but most 20th century students have followed Pax broader than radicle. & Hoffman (1922) in combining them. There is considerable anatomical and palynological varia- This mainly paleotropical tribe of 18 genera in tion withinDrypetes sensu lato, however, and the 5 subtribes is well characterized by its leaves with generic limitsrequire furtherstudy. I am indebted tanniniferousepidermis (Levin, 1986a) and prolate to Geoffrey Levin (pers. comm.) for critical sug- pollen grains with 3 narrow colpi (Khler, 1965). gestions on the generic synonymy. The tribe is here given a broader circumscription than that of Hurusawa (1954), who did not include 36. Sibangea Oliver, Hook. Ic. P1. 15: 9, t. the Spondianthinae, Uapacinae, or Porantherinae. 1411. 1883; Radcliffe-Smith,Kew Bull. 32: 480. 1978; Fl. Trop. E. Afr.,Euphorb. 101. KEY TO THE SUBTRIBES OF TRIBE ANTIDESMEAE 1987. TYPE: Sibangea arborescens Oliver. la. Stemswith resinous secretion; inflorescence paniculate,or capitulate. As interpretedby Radcliffe-Smith,Sibangea in- 2a. Petals present;inflorescence terminal, pa- cludes three Africanspecies. The genus is very niculate,bracts small; pollen sexine not close to Drypetes, and it remains to be seen wheth- spinulose;fruit capsular; stomata paracytic er it can be maintainedas distinct. 6a. Spondianthinae 2b. Petals absent;inflorescences axillary, capitate,bracts large and involucrate;pollen 37. Putranjiva Wallich, Tent. Fl. Nepal 61. sexinespinulose; fruit drupaceous; stomata 1826; Muell. Arg., DC. Prodr. 15(2): 443. anisocytic ...... 6b. Uapacinae lb. Stems lackingresin; inflorescence not panicu- 1866; Bentham, Gen. P1. 3: 277. 1880; Pax late or capitulate;petals mostlyabsent. & Hoffm.,Natiirl. Pflanzenfam. ed. 2, 19c: 3a. Inflorescencesaxillary, amentlike; trees or 59. 1931. TYPE: Putranjiva roxburghii Wal- shrubs(except Cyathogyne). lich [Nageia putranjiva Roxburgh]. 4a. Pollen grainsglobose or slightlypro- late; anther locules not pendulous; Palenga Thwaites,Hook. J. Bot. Kew Gard.Misc. 8: leaves sometimesglandular; stomata 270. 1856. TYPE: Palenga zeylanica Thwaites[ anisocytic..... 6c. Scepinae Putranjiva zeylanica (Thwaites)Muell. Arg.]. 4b. Pollengrains distinctly prolate; anther LiodendronKeng, J. WashingtonAcad. Sci. 41: 201. locules ? pendulous;leaves not glan- 1951. TYPE: Liodendron matsumurae(Koidzumi) dular; stomataparacytic..... Keng [= Putranjiva matsumuraeKoidz.]...... 6d. Antidesminae

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3b. Inflorescencesterminal, corymbiform; herbs phyllous; stamens 5, filaments free, anthers in- -...... 6e. Porantherinae trorse; pistillode apically dilated; pistillate calyx gamophyllous; ovary mostly 3-locular; styles free, Subtribe 6a. SPONDIANTHINAE (Webster) multifid;fruit drupaceous; seeds 1 per locule, al- Webster,stat. nov. Tribe Spondiantheae Web- buminous; cotyledons much longer than radicle. ster,Taxon 24: 594. 1975. TYPE: Spondian- thus Engler. This subtribe includes only the single genus Uapaca, which has always appeared isolated be- Dioecioustrees; leaves entire,penninerved, stip- cause of its remarkable involucrate inflorescences: ules caducous; inflorescencesterminal (at least in Airy Shaw (1965) and Meeuse (1990) have as- part),paniculate, bracts small; flowers (at least the signed it to a separate family, Uapacaceae. How- staminate)with petals; staminatedisk dissected; ever, Bentham (1880) juxtaposed it with Aporusa, stamens5, filamentsfree, anthers introrse; pistil- and Pax (1890) referred it to his subtribe Anti- lode present;pistillate disk lobed; stylesfree; fruit desminae. Those intuitions appear confirmed by loculicidallydehiscent; columella persistent; seeds the foliar anatomical studies of Levin (1986a, b, 1 per locule,testa dry; endosperm scanty; embryo c), who has found that the tanniniferousepidermal withcotyledons much broaderthan radicle. cells and anisocytic stomata indicate an affinity A monotypictribe, including only the African with subtribe Scepinae. On the other hand, the genus Spondianthus. Pax & Hoffmann(1922) distinctivepollen and resiniferousstems of Uapaca placed it adjacent to Thecacoris,a genus of sub- are divergent characters, and it appears best re- tribeAntidesminae. Levin (1986c), on the basis of ferred to a separate subtribe. foliarcharacters, suggested that it is the primitive sistergroup of the otherwoody tribes of Antides- 39. Uapaca Baillon, Etude Gen. Euphorb. 595. meae. The observationsof K6hler (1965) on pollen 1858; Muell. Arg., DC. Prodr. 15(2): 489. suggesta similarposition. 1866; Pax & Hoffm., Pflanzenr. 147. XV (Heft 81): 298. 1922; Leandri, Fl. Madag. 38. Spondianthus Engler,Bot. Jahrb.36: 215. 111(1): 163. 1958; Radcliffe-Smith,Fl. E. 1905; Hutch., Hook. Icon. P1. 30: t. 2986. Trop. Afr.,Euphorb. 567. 1988. TYPE: Uapa- 1911; Fl. Trop. Afr.6(1): 1044. 1913; Pax ca thouarsii Baillon [lectotype, selected by & Hoffm.,Pflanzenr. 147. XV (Heft81): 13. Airy-Shaw, Kew Bull. 18: 271. 1965]. 1922; Keay, Fl. W. Trop. Afr.ed. 2, 1: 372. A genus of ca. 60 species in Africa and Mad- 1958; Aubreville,Fl. Forest. Cote d'Ivoire, agascar. ed. 2, 2: 62. 1959; Radcliffe-Smith,Fl. E. Trop. Afr.Euphorb. 104. 1987; Leonard & Nkounkou,Bull. Jard. Bot. Nat. Belg. 59: Subtribe 6c. SCEPINAE (Lindley) Webster, stat. nov. Ordo Scepaceae Lindl., Nat. Syst. 133. 1989. TYPE: Spondianthuspreussii En- gler. Bot. ed. 2, 171. 1836. TYPE: Scepa Lindl. [ Aporusa Bl.]. Megabaria De Wild., Etud. Fl. Bas & Moy. Congo 2: 284. 1908. TYPE: Megabaria trillesiiDe Wild. [= Ordo AporuseaeLindl. ex Miq., Fl. Ind. Bat. 1(2): 430. Spondianthuspreussii Engler]. 1858. TribeAporuseae (Lindl. ex Miq.) AiryShaw, Hook. Icon. P1. 38: t. 3701. 1974; Webster,Taxon A monotypicgenus of a singlevariable species, 24: 594. 1975. TYPE: Aporusa Bi. widespreadin tropicalAfrica from Liberia to An- Dioecious or monoecious trees or shrubs, stems gola and Tanzania. without resinous secretion; leaves often with laminar glands, stomata mostly anisocytic (paracytic Subtribe 6b. UAPACINAE Muell. Arg., Lin- in Jablonskia), stipules deciduous or persistent; naea 34: 64. 1865 (as Phyllantheaesubtr. staminate bracts mostly each with 3 or more flow- Uapaceae). Tribe Uapaceae (Muell. Arg.) ers; sepals 4-8, free; petals absent; disk lobed, Hutch.,Amer. J. Bot. 56: 747. 1969. Family dissected, or absent; stamens 4-8, free; anthers Uapaceae (Muell. Arg.)Airy Shaw, Kew Bull. dehiscing longitudinally,connective not enlarged; 18: 270. 1965. TYPE: Uapaca Baillon. pollen grains subglobose to subprolate, 3-colporate, Dioecious trees or shrubs; leaves entire,pin- reticulate or tectate; pistillode present; ovary 2- natelyveined, stipules caducous; inflorescences ax- 5-locular; styles bifidor unlobed; fruitcapsular or illary,capitate, involucrate, pedunculate; flowers tardily dehiscent and subdrupaceous; seeds 1-6 apetalous and withoutfloral disk; calyx gamo- per fruit,seed-coat dry or fleshy; endosperm thin

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to copious,cotyledons mostly distinctly broader and Airy Shaw (1974) at the tribal rank. Since Lindley longerthan radicle. published the Scepaceae over 20 years before the ordo Aporuseae was proposed, it seems preferable Thistaxon, although early designated by Lindley to preserve his name. The seven genera of Scepinae (1836) as an "ordo," was confoundedby Mueller are scattered through the tropics in America, Af- (1866) and later authorswith the Antidesminae, rica, and Asia, with only Baccaurea attaining the and was firstrecognized in the 20th centuryby Pacific islands.

KEY TO THE GENERA OF SUBTRIBE SCEPINAE la. Pistillatedisk dissected; styles dilated, entire to slightlylobed 40. Protomegabaria lb. Pistillatedisk cupularor absent. 2a. Staminatedisk present. 3a. Pistillatedisk present; pistillate sepals persistentin fruit. 4a. Dioecious; flowersracemose; stipules deciduous. 5a. Capsule loculicidal;staminate flowers mostly one per bract 41. Maesobotrya 5b. Capsule septicidal;staminate flowers several per bract 42. Richeria 4b. Monoecious;flowers in axillaryglomerules; stipules persistent; capsule septicidal 43. Jablonskia 3b. Pistillatedisk absent; pistillate sepals deciduous. 6a. Dioecious;styles bifid -44. Baccaurea 6b. Monoecious;styles unlobed -45. Apodiscus 2b. Staminatedisk absent. 7a. Pistillodemassive, peltate; stamens 5 or 6; carpelsmostly 3 or 4; stylesstigmatiform --- 46. Ashtonia 7b. Pistillodesmall; stamens mostly 2 (rarely3-5); carpelsmostly 2 (rarely3 or 4); stylesbifid 7. Ap o rusa -...... 4

40. Protomegabaria Hutchinson,Hook. Icon. 1866; Fl. Bras. 11(2): 13. 1873; Bentham, P1. 30: t. 2929. 1911; Fl. Trop. Afr.6(1): Genera Plantarum 3: 286. 1880; Pax & 656. 1912; Pax & Hoffim.,Pflanzenr. 147. Hoffm.,Pflanzenr. 147. XV (Heft 81): 26. XV (Heft81): 43. 1922; Keay, Fl. W.' Trop. 1922; Jablonski,Mem. New York Bot. Gard. Afr.ed. 2, 1: 373. 1958; Aubreville,Fl. For. 17: 124. 1967; Philcox,Fl. Trin. Tob. 2(9): Cote, d'Ivoire 2: 68, t. 147. 1959. TYPE: 638. 1979; Webster,Ann. Missouri Bot. Gard. Protomegabaria stapfiana (Beille) Hutch. 75: 1093. 1989; Secco & Webster,Bol. Mus. (lectotype,selected here). Para. EmilioGoeldi, Bot. 6: 141-158. 1990. TYPE: Richeria grandis Vahl. Two species of West Africa. GuaraniaWeddell ex Baillon,Etude Gen. Euphorb. 598. 41. Maesobotrya Bentham,Hook. Icon. P1. 13: 1858. TYPE: Guarania gardneriana Baillon[= 75, t. 1296. 1879; Gen. P1. 3: 284. 1880; Richeria gardneriana (Baillon)Baillon; lectotype]. Hutchinson,Fl. Trop. Afr.6(1): 663. 1912; Pax & Hoffim.,Pflanzenr. 147. XV (Heft81): A genus of several poorly defined taxa, treated ex- 17. 1922; Leonard,Bull. Jard.Bot. Brux. 17: as two species by Secco & Webster(1990); rainforest 256. 1945; Keay, Fl. W. Trop. Afr.ed. 2, tendingin lowlandand montanetropical Peru 1: 373. 1958; Radcliffe-Smith,Fl. Trop. E. fromCosta Rica and the Lesser Antillesto Afr.Euphorb. 112. 1987. TYPE: Maesobotrya and Brazil. floribunda Bentham. 43. Jablonskia Webster, Syst. Bot. 9: 232. Pierre,Bull. Soc. Linn.Paris 1233. 1896; Staphysora congesta (Benth.ex Pax,Bot. Jahrb. 23: 521. 1897. TYPE: Staphysora 1984. TYPE: Jablonskia duseniiPax [= Maesobotryadusenii (Pax) Hutch.]. Muell. Arg.) Webster[Securinega congesta Benth.ex Muell. Arg.]. An Africangenus of ca. 20 species,all in West Africaand only1 speciesextending east to Uganda. Monotypic,the singlespecies confinedto Am- The genus is extremelyclose to the neotropical azonian South America.The genus is isolatedin Richeria. subtribeScepinae by its distinctivepollen type and paracyticstomata and shows some similaritiesto 42. Richeria Vahl, Eclog. Amer. 1: 30, t. 4. Celianella Jabl. in subtribe Antidesminae.Its 1797; Muell. Arg., DC. Prodr. 15(2): 466. placementrequires further evaluation.

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44. Baccaurea Loureiro, Fl. Cochinch. 661. 3: 282. 1880; Hook. f., Hook. Icon. P1. 16: 1790; Muell. Arg.,Prodr. 15(2): 456. 1866; 5. 1583. 1887; Fl. Brit. Ind. 5: 345. 1887; Bentham,Gen. P1. 3: 283. 1880; Pax & Pax & Hoffm.,Pflanzenr. 147. XV (Heft 81): Hoffim.,Pflanzenr. 147. XV (Heft 81): 45. 80. 1922; Backer & Bakh., Fl. Java 1: 455. 1922; AiryShaw, Kew Bull. 14: 353. 1960; 1963; AiryShaw, Kew Bull. 30: 380. 1966; Backer & Bakh., Fl. Java 1: 453. 1963; 23: 2. 1969; 25: 474. 1971; Thin, Cong Whitmore,Tree Fl. Malaya 2: 63. 1973; A. TrinhNghian Cu'u Khoa Hoc 10: 82. 1986. C. Smith,Fl. Vit. Nov. 2: 450. 1981; Thin, TYPE: Aporusaftutescens Blume. Cong TrinhNghien Cu'u Khoa Hoc 10: 78. Leiocarpus Blume,Bijdr. 581. 1826. TYPE: Leiocarpus 1986. TYPE: Baccaurea ramifiora Lour. [lec- fruticosusBlume [= Aporusa fruticosa (Blume) totype, selected by Merrill, 1935]. Muell.Arg.; lectotype, designated by Wheeler, 1975]. LepidostachysWallich ex Lindley,Nat. Syst.ed. 2, 441. PierardiaRoxb. ex Jack,Trans. Linn. Soc. London14: 1836. TYPE: Lepidostachysroxburghii Wallich ex 119. 1823. TYPE: Pierardia dulcis Jack [= Bac- Lindley[nom. illeg.; = Aporusadioica (Roxb.)Muell. caurea dulcis (Jack)Muell. Arg.]. Arg.]. AdenocrepisBlume, Bijdr. 579. 1825. TYPE: Adenocre- Scepa Lindley,Nat. Syst.ed. 2, 441. 1836. TYPE: Scepa pisjavanica Bl. [= Baccaurea javanica (Bi.) Muell. stipulaceaLindley [= Aporusadioica (Roxb.)Muell. Arg.]. Arg.]. CalyptroonMiquel, Fl. Ned. Ind., ErsteBijv. 471. 1861. TYPE: Calyptroonsumatranum Miq. [= Baccaurea A genus of 80 Old World species, from India sumatrana(Miq.) Muell. Arg.]. and Sri Lanka to Indonesia and Melanesia (Solomon Microsepala Miquel,Fl. Ned. Ind.,Erste Bijv. 444. 1861. Islands). TYPE: Microsepala acuminata Miq. [= Baccaurea javanica (Bi.) Muell.Arg.]. EverettiodendronMerrill, Phil. J. Sci. Bot. 4: 279. 1909. TYPE: Everettiodendronphilippinense Merr. [ Subtribe 6d. ANTIDESMINAE Muell. Arg., Baccaurea philippinensis(Merr.) Merr.]. Linnaea 34: 64. 1865 (as Antidesmeae); Pax, Gatnaia Gagnepain,Bull. Soc. Bot. France 71: 870. Natfirl. Pflanzenfam. ed. 1, 3(5): 26. 1890; 1924. TYPE: Gatnaia annamica Gagnep. [= Bac- Pax & Hoffm.,Pflanzenr. 147. XV (Heft 81): caurea oxycarpa Gagnep.]. 3. 1922. TYPE: Antidesma L. A diversifiedgenus of ca. 75 paleotropicalspe- Ordo StilagineaeC. A. Agardh,Aphor. Bot. 14: 199. from to the New cies, extending India Philippines, 1824. TYPE: Stilago L. [= AntidesmaL.]. Guinea,and the Pacificislands. SubtribeHieronymeae Muell. Arg., Linnaea 34: 64. 1865. TYPE: HyeronimaFr. Allem.

45. Apodiscus Hutchinson, Bull. Soc. Bot. Dioecious (rarely monoecious) trees or shrubs France 58, Mem. 8: 205. 1912; Fl. Trop. (rarely herbs); leaves eglandular, stomata para- Afr.6(1): 1045. 1913; Hook. Icon. P1. 31: cytic, stipules caducous; inflorescences axillary, ? t. 3032. 1915; Pax & Hoffim.,Pflanzenr. amentlike; staminate bracts mostly 1-flowered;se- 147. XV (Heft 81): 45. 1922. TYPE: Apodis- pals 3-6, free or connate; petals mostly absent or cus chevalieri Hutch. reduced; disk lobed or dissected; stamens 3-6, free A poorlyunderstood monotypic genus from the or nearly so, anthers with ? enlarged glandular Guinea region of west Africa; said by Pax and connectives, anther-sacs usually pendulous; pollen Hoffmannto be relatedto Maesobotrya. grains prolate, 3-colporate, reticulate; pistillode present; carpels 1-3(-5), ovules anatropous; styles bifid (rarely entire); fruit capsular or drupaceous; Bull. 357. 46. Ashtonia Airy Shaw, Kew 1968; seeds mostly 1 per locule or fruit; seed-coat dry; P1. t. Whitmore, Hook. Icon. 38: 3702. 1974; endosperm thinto copious, cotyledons broader than Tree Fl. Malaya 2: 62. 1972; Airy Shaw, but scarcely exceeding radicle. Kew Bull. Add. Ser. 4: 42. 1975. TYPE: Ash- tonia excelsa Airy Shaw. Subtribe Antidesminae corresponds to the family Stilaginaceae as recognized by Airy Shaw (1965, of two in Borneo An Old World genus species and many subsequent publications). However, pol- and the Malay peninsula. len and foliar anatomical evidence clearly supports membership of Antidesma in the Euphorbiaceae. 47. Aporusa Blume, Bijdr. 514. 1826 (as Apo- Two of the six genera (Phyllanoa and Leptonema) rosa); Fl. Jav. 1: viii. 1828; Muell. Arg., DC. are not well understood, and their inclusion here Prodr. 15(2): 469. 1866; Bentham, Gen. P1. must be regarded as provisional.

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KEY TO THE GENERA OF SUBTRIBE ANTIDESMINAE 1a. Indumentumabsent or of simpleor stellatehairs. 2a. Carpels 3-5; fruitdehiscent. 3a. Disk present;ovary 3-locular. 4a. Pistillodepresent; pistillate sepals not accrescent in fruit;pistillate flower sometimes with petals and staminodes. 5a. Leaves entire;pistillate flowers not bracteolate; fruits pedicellate 48. Thecacoris 5b. Leaves dentate;pistillate flowers bracteolate; fruits subsessile -49. Phyllanoa 4b. Pistillodeabsent; pistillate sepals accrescent; pistillate flowers lacking petals or staminodes ------.------50 . Ce lia n e lla 3b. Disk absent;ovary 4-5-locular- 51. Leptonema 2b. Carpelsolitary; fruit indehiscent -52. Antidesma lb. Indumentumlepidote; fruit indehiscent ..53. Hyeronima

48. Thecacoris A. Jussieu,Euphorb. Tent. 12. 51. Leptonema A. Jussieu,Euphorb. Tent. 19. 1824; Baillon, Etude Gen. Euphorb. 605. 1824; Bentham,Gen. P1. 3: 275. 1880; Pax 1858; Bentham,Gen. P1. 3: 286. 1880; Pax & Hoffim.,Naturl. Pflanzenfam. ed. 2, 19c: & Hoffrm.,Pflanzenr. 147. XV (Heft 81): 8. 59. 1931; Leandri,Notul. Syst. (Paris) 6: 22. 1922; Keay, Fl. W. Trop. Afr.ed. 2, 1: 371. 1937; Mem. Inst.Sci. Madag. 8B: 212. 1957; 1958; Leandri,Fl. Madag. 1: 4. 1958; Rad- Fl. Madag. 111(1): 12. 1958. TYPE: Lepto- cliffe-Smith,Fl. Trop. E. Afr.Euphorb. 107. nema venosum(Poir.) A. Juss. 1987. TYPE: Thecacorismadagascariensis A. A poorlyknown genus of two species endemic Juss. to Madagascar. CometiaDu PetitThouars ex Baillon,Etude, Gen. Eu- phorb.642. 1858. TYPE: Cometiathouarsii Baillon 52. Antidesma Burman ex L., Sp. P1. 1027. Thecacoriscometia [= Leandri]. 1753; Gen. P1. 451. 1754; Muell. Arg.,DC. CyathogyneMuell. Arg., Flora 47: 536. 1864. TYPE: Cyathogyneviridis Muell. Arg. [= Thecacorisvir- Prodr. 15(2): 247. 1866; Bentham,Gen. P1. idis (Muell.Arg.) Leandri]. 3: 284. 1880; Hooker,Fl. Brit.Ind. 5: 354. BaccaureopsisPax, Bot.Jahrb. 43: 319. 1909. TYPE: 1887; Pax & Hoffim.,Pflanzenr. 147. XV Baccaureopsislucida Pax [= Thecacorisludica (Heft 81): 107. 1922; AiryShaw, Kew Bull. (Pax) Hutch.]. 23: 277. 1969; 26: 457. 1972; 28: 269. As heredelimited, Thecacoris includes about 20 1973; 33: 15, 423. 1979; 36: 358, 635. species fromAfrica and Madagascar. The herba- 1981; Leonard, Bull. Jard. Bot. Nat. BeIg. ceous species have been segregatedas a separate 58: 4. 1988. Bestram Adanson, Fam. P1. genus Cyathogyneby Mueller(1866) and Pax & 354. 1763. TYPE: Antidesmaalexiteria L. Hoffmann(1922, 1931), butas notedby Radcliffe- StilagoL., Mant.P1. 16. 1767. TYPE: Stilagobunius L. Smith (1987), Thecacoris usambarensis Verdc. [= Antidesmabunius (L.) Spreng.]. bridgesthe gap betweenthe twogeneric concepts. RhytisLour., FH. Cochinchin. 660. 1790. TYPE: Rhytis fruticosaLour. [= Antidesmafruticosum (Lour.) 49. Phyllanoa Croizat,Caldasia 2: 123. 1943. Muell.Arg.]. TYPE: Phyllanoa colombiana Croizat. A large Old World genus of ca. 200 species, A monotypicgenus from the ColombianAndes, only 10 of whichoccur in Africaand Madagascar. knownonly from the imperfect type specimen. The The remainderof the species are Asian exceptfor floraldetails of the pistillateflower, especially the a fewspecies in Australiaand the Pacificislands. staminodes,suggest a possible affinitywith The- 53. P1. Bras. 1. cacoris, althoughthere is also a general resem- Hyeronima Allemao, Novas 1848; Muell. DC. Prodr. 268. blance to Richeria, in subtribeScepinae. Arg., 15(2): 1866 (as Hieronyma);Bentham, Gen. P1. 3: 284. 1880; Pax & Hoffm.,Pflanzenr. 147. 50. Celianella Mem. Jablonski, New York Bot. XV (Heft 81): 31. 1922; Jablonski,Mem. 12: 176. TYPE: Celianella Gard. 1965. mon- New York Bot. Gard. 17: 122. 1967; Web- tana Jabl. ster,Ann. Missouri Bot. Gard. 75: 1094. 1988; A monotypicgenus of the VenezuelanGuayana Franco, Bot. Jahrb.Syst. 111: 297. 1990. highlands.The plantis distinctivein the semisuc- TYPE: Hyeronimaalchorneoides Allemao. culentleaves and accrescentpistillate sepals. Levin StilaginellaTul., Ann.Sci. Nat. III. 15: 240. 1851. (1986c) allied it withHyeronima on the basis of TYPE: Stilaginellalaxifiora Tul. [= Hyeronimalax- foliarcharacters. ifiora(Tul.) Muell. Arg.].

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A neotropical genus of more than 20 described Latin description), even though he was obviously species, many of these difficultto distinguish.Fran- basing it on Gruning's section. co (1990) recognized 10 species for South Amer- ica, and there are a few additional species in the Tribe 7. HYMENOCARDIEAE (Muell. Arg.) West Indies. Hutchinson, Amer. J. Bot. 56: 746. 1969. Phyllantheae subtribeHymenocardieae Muell. Arg., Linnaea 34: 64. 1865. Hymenocardi- Subtribe 6e. PORANTHERINAE (Muell.Arg.) aceae Airy Shaw, Kew Bull. 18: 261. 1965; K6hler, Grana Palynologica 6: 99. 1965. Tribe Leonard & Mosango, Fl. Afr. Centrale, Hy- Poranthereae Muell. Arg., Bot. Zeitung 22: menocardiaceae 1. 1985. TYPE: Hymenocar- 324. 1864. TYPE: Poranthera Rudge. dia Wall. ex Lindley. Monoecious or dioecious herbs or subshrubs; Dioecious trees or shrubs; indumentumof simple leaves + ericoid; flowers in terminal corymbose hairs and sessile glands; leaves alternate, entire, racemes; sepals 5; petals present or absent; sta- penninerved, stipulate; inflorescences axillary, spi- mens 5; disk present; anthers pendulous; pistillode cate, racemose, or paniculate; staminate sepals 4- small or absent; pistillate disk annular; ovary 8, connate; petals absent; stamens 4-8, filaments 3-locular; ovules anatropous; styles bifid; fruits free or basally connate; anthers longitudinallyde- capsular; seeds 2 per locule; endosperm copious; hiscent; pollen porate; pistillode present; pistillate cotyledons about as broad as radicle. flowers pedicellate; sepals 4-9, free, caducous; Two closely related Australasian genera. Their ovary 2-locular; styles 2, elongated, entire or lac- systematic position remains controversial. Pax & erate, free; ovules anatropous, 2 per locule; fruit Hoffmann(1931) followed Mueller (1866) in plac- capsular or winged, samaroid; seeds 1 per locule; ing tribe Porantheae in the division "Stenolobae," endosperm scanty; cotyledons broader and longer because of the narrow cotyledons. In 1975, I as- than radicle. sociated Poranthera with Andrachne, partly be- A distinctive group originally including a single cause of the pollen data of Punt (1962). However, Old World genus, considered to be a distinctfamily K6hler (1965) showed that the pollen of Poranth- by a number of modern workers (e.g., Leonard & era was more compatible with an affinityto the Mosango, 1985; Meeuse, 1990). However, most Antidesma complex, and he demoted Porantheae floraland anatomical details (e.g., in Levin, 1 986a, to a subtribe of Antidesminae. Because he did not b, c) indicate that it is better retained within the cite the basionym, his combination was not validly Euphorbiaceae. A second genus, Didymocistus, is published, and it presumably is validated here. provisionallyincluded here, although it is very distinctfrom Hymenocardia in many ways, and could KEY TO THE GENERA OF SUBTRIBE PORANTHERINAE perhaps be placed in a separate tribe or subtribe. la. Monoecious;petals and pistillodepresent - 54. Poranthera KEY TO THE GENERA OF TRIBE HYMENOCARDIEAE lb. Dioecious; petalsand pistillodeabsent -____--__-_.._-_--__ _-__--._.. __-______-_-___-_-._____- __--___-__ _-_ 55. Oreoporanthera la. Fruitcapsular; inflorescencepaniculate; styles lacerate------56. Didymocistus lb. Fruitsamaroid; inflorescence racemose; styles 54. Poranthera Rudge, Trans. Linn. Soc. Lon- papillose.... 57. Hymenocardia don 10: 302. 1811; Bentham, Fl. Austral. 6: 54. 1873; Gen. P1. 3: 262. 1880; Grining, 56. Didymocistus Kuhlmann, An. Prim. Reun. Pflanzenr. 147. (Heft 58): 13. 1913. TYPE: Sud-Amer. Bot. 3: 82. 1940. TYPE: Didy- Poranthera Rudge. mocistus chrysadenius Kuhlm.

Ten species from Australia. A monotypic genus of Amazonian Brazil. Kuhl- mann related it to Aporusa, but the anatomical studies of Levin (1986a, b, c) suggest that it is 55. Oreoporanthera Hutchinson, Amer. J. Bot. possibly closer to Hymenocardia. It does appear 56: 747. 1969. Poranthera sect. Oreopor- that Didymocistus may possibly represent a link anthera Grining, Pflanzenr. 147. (Heft 58): between tribes Antidesmeae and Hymenocardieae. 21. 1913. TYPE: Oreoporanthera alpina (Cheeseman) Hutchinson. 57. Hymenocardia Wallich ex Lindley, Nat. A monotypic genus endemic to New Zealand. Syst. ed. 2, 441. 1836; Muell. Arg., DC. As pointed out by Huft (pers. comm.) Hutchinson Prodr. 15(2): 476. 1866; Oliver, Hook. Ic. described Oreoporanthera as a new genus (with P1. 12: 29, t. 1131. 1873; Bentham, Gen.

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P1. 3: 284. 1880; Pax & Hoffm., Pflanzenr. species from Gabon that was referred to a fla- 147. XV (Heft 81): 72. 1922; Leonard & courtiaceous affinityby Pierre, but to the Eu- Mosango, Fl. Afr. Centrale, Hymenocardi- phorbiaceae by Gilg (1908) and Hutchinson (1912). aceae 2. 1985; Radcliffe-Smith,Fl. E. Trop. Pax & Hoffmann(1931) placed it in the Antides- Afr., Euphorbiaceae 577. 1988. TYPE: Hy- minae next to Spondianthus, but its place within menocardia punctata Wallich ex Lindl. the Euphorbiaceae still appears uncertain.

Samaropyxis Miquel, Fl. Ned. Ind., Erste Bijv. 464. 60. Meborea Aublet,Hist. P1. Gulane 825.1775; 1861. TYPE: Samaropyxis elliptica Miq. [= Hy- menocardiawallichii Tul.]. Lanjouw, Euphorb. Suriname 9, t. 1. 1931. TYPE: Meborea guianensis Aublet. An Old World genus of eight species, of which seven are African and one southeast Asian; absent This monotypic genus from French Guiana was from Madagascar. referred by Mueller (1866) to Phyllanthus. Lan- jouw was unable to place the genus, but regarded Tribe 8. BISCHOFIEAE (Muell. Arg.) Huru- it as not synonymous with Phyllanthus, a view sawa, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. with which I agree. Judgingfrom Aublet's descrip- 6: 339. 1954. Phyllantheae subtribe Bis- tion and illustration, it seems probable that Me- choffieaeMuell. Arg., Linnaea 34: 64. 1865. borea belongs to tribe Phyllantheae, but it cannot Family Bischofiaceae (Muell. Arg.) Airy Shaw, be identifiedwith certainty. Kew Bull. 18: 252. 1965. TYPE: Bischofia Blume. Subfamily II. OLDFIELDIOIDEAE K6hler & Webster, J. Arnold Arbor. 48: 308. 1967; a Asian species, very A monotypic tribe of single Webster, Taxon 24: 595. 1975. TYPE: Old- distinct from other Phyllanthoideae by virtue of its fieldia Benth. trifoliolate leaves. Although Airy Shaw (1965) suggested a relationship with the Staphyleaceae, evi- Paivaeusaceae Meeuse, EuphorbiaceaeAuct. Plur. 30. 1990. TYPE: PaivaeusaWelw. ex Benth.[= Old- & 1974) dence from embryology (Bhatnagar Kapil, fieldiaBenth. & Hook.]. and leaf anatomy (Levin, 1986a, b, c) supports retention in the Euphorbiaceae. Monoecious or dioecious trees, shrubs, or subshrubs; leaves alternate or more commonly op- 58. Bischofia Blume, Bijdr. 1168. 1826; Muell. posite or verticellate, simple or palmately com- Arg., DC. Prodr. 15(2): 478. 1866; Bentham, pound, entireor dentate; stipulessmall and caducous Gen. P1. 3: 280. 1880; Pax & Hoffm., Pflan- or absent; inflorescences axillary, glomerulate, ra- zenr. 147. XV (Heft 81): 312. 1922; Airy cemose, capitulate, or paniculate; flowersapetalous Shaw, Kew Bull. 18: 253. 1965; 21: 327. (except in Croizatia); staminate sepals (3-)4- 1967; A. C. Smith, Fl. Vitiensis Nova 2: 494. 8(-12), imbricate, usually free; disk intrastaminal 1981. TYPE: Bischofia javanica Blume. or absent (extrastaminal in Croizatia); stamens (2)3-30, filaments free or connate; pollen grains Microelus Wight & Arnott,Edinb. New Philos. J. 14: spheroidal, tectate, exine mostly with conspicuous 298. 1833. TYPE:Microelus roeperianus Wight & Arnott[= Bischofiajavanica Blume]. spines, brevicolporate or porate; pistillode present Stylodiscus Bennett,P1. Jav. Rar. 133. 1840. TYPE: or absent; pistillate sepals (3-)4-8(-13), free; disk Stylodiscustrifoliatus (Roxb.) Bennett[Andrachne annular to lobed or dissected, or absent; ovary 2- = trifoliataRoxb. Bischofiajavanica Blume]. 4(-5)-locular; ovules 2 per locule (except in Sca- A genus of a single species distributed from India gea); styles entire or stigmatoid(rarely bifid);fruit and Sri Lanka to China, Indonesia, and the Pacific capsular (rarely drupaceous); seeds 1 or 2 per islands. locule, often carunculate, testa usually smooth and shiny; endosperm usually copious (rarely obsolete).

INCERTAE SEDIS This most recently recognized subfamily in- 59. Centroplacus Pierre, Bull. Mens. Soc. Linn. cludes 28 genera with only ca. 100 species, over- Paris II. 1: 114. 1899; Gilg, Bot. Jahrb. 40: whelminglyfound in the Southern Hemisphere. 516. 1908; Hutchinson, Fl. Trop. Afr. 6(1): 629. 1912; Pax & Hoffm., Natiirl. Pflanzen- KEY TO THE TRIBES OF SUBFAMILY OLDFIELDIOIDEAE fam. ed. 2, 19c: 46. 1931. TYPE: Centropla- cus glaucinus Pierre. 1a. Petals present;staminate disk extrastaminal, annular;styles twice bifid; ovules hemitropous This genus is based on a single poorly known ...... 9 . CR O IZ AT IE AE

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lb. Petals absent; staminatedisk dissected,intra- 4-brevicolporata,echinata; ovarium 3-loculare, stylis in- staminalor absent;styles entire or rarelyonce tegris,ovulis anatropis; semina albuminata. bifid;ovules anatropous. 2a. Pollen aperturesfour; vessel elementsat Dioecious trees; leaves alternate or opposite, least in part withscalariform perforation entire or serrate; stipules caducous. Inflorescences plates (except in Tetracoccus);staminate in thyrses or ebracteate cymes. Flowers apetalous, diskextrastaminal or absent;leaves simple disk absent; staminate sepals (3)4 or 5, free or 10. PODOCALYCEAE 2b. Pollenwith more than 4 apertures(rarely connate, imbricate; stamens 5-13, free; anthers inaperturate);vessel elementswith simple extrorse or introrse;pollen grains 4-brevicolporate, perforationplates; staminatedisk intra- echinate, tectum smooth; pistillode present or ab- staminalor absent;leaves simple,unifoli- sent; pistillate sepals 4 or 5, imbricate, persistent; olate, or palmatelycompound. or 3-locular; styles di- 3a. Leaves simple; monoeciousor dioe- disk present absent; ovary cious trees, shrubs,or herbs; seeds lated, stigmatiform;ovules anatropous. Fruit cap- mostlycarunculate 11. CALETIEAE sular or drupaceous; seeds carunculate or ecarun- 3b. Leaves compound,or if simplethen culate; endosperm present; cotyledons plane or laminastipellate or stipulesadnate to plicate, much broader than radicle. petiole;dioecious trees or shrubs;seeds carunculateor ecarunculate. This tribe includes three American genera that ...... 12. PICRODENDREAE are differentin many respects and are classified together on the basis of pollen characters (pollen Tribe 9. CROIZATIEAE Webster, trib. nov. grains with four shortened colpi). The differences TYPE: Croizatia Steyermark. between the genera are so strikingthat it seems necessary to assign them to differentsubtribes. Folia alternaintegra; discus annularis;petalae 5, im- bricatae; stamina 5, antherisextrorsis; pollinis grana 3-colporataspinulosa; ovarium 3-loculare, stylis bis bifidis, KEY TO THE SUBTRIBESOF TRIBE PODOCALYCEAE hemitropis;semina ecarunculata, exalbuminata. ovulis la. Pedunclefree from petiole; fruit capsular. Dioecious trees or shrubs; leaves alternate, en- 2a. Staminatesepals connate; pistillodepresent;staminate and pistillateflowers in elon- in tire; stipules deciduous or persistent; flowers gated spikes; seeds ecarunculate;leaves axillary clusters; staminate sepals and petals 5, alternate,entire ...... 1 Oa. Podocalycinae imbricate; disk annular; stamens 5, free, anthers 2b. Staminatesepals discrete; pistillode absent; introrse;pollen grains globose, 3-colporate, exinous pistillateflowers solitary or clustered;seeds carunculate;leaves alternate,opposite, or spines conspicuous; pistillode present; pistillate se- whorled,entire or dentate...... pals and petals 5, imbricate; disk annular, glabrous; ...... Tetracoccin ae ovary pubescent, 3-locular; styles free, twice bifid; lb. Peduncleadnate to thepetiole; staminate sepals ovules hemitropous; fruitcapsular; columella per- discrete;pistillode absent; fruit drupaceous .. sistent,dilated; seeds 1-2 per locule, ecarunculate; ...... P aradrypetin ae endosperm absent; cotyledons broader than and about equaling radicle. Subtribe lOa. PODOCALYCINAE Webster, subtr.nov. TYPE: Podocalyx K1. The tribe is represented by the single genus Croizatia. This taxon shows clear affinitieswith Folia alterna,integra; flores spicatis; antheris extrorsis; the tribe Amanoeae in subfamilyPhyllanthoideae, flos6 pistillodio;discus 9 sericeus;fructus capsularis; co- tyledonesplanis. and it is more or less arbitrarywhere it is placed. Trees with alternate leaves; stipules caducous; 61. Croizatia Steyermark, Fieldiana 28: 308, flowersin axillary spikes; staminate sepals 5, con- fig. 57. 1952; Webster, Gillespie & Steyer- nate below; disk obsolete; stamens 5 or 6, anthers mark, Syst. Bot. 12: 6. 1987. TYPE: Croizatia extrorse; pollen grains 4-colporate, echinate; pis- neotropica Steyermark. tillode present; pistillate flowerspedicellate; sepals 5, imbricate, persistent; disk cupular, sericeous; A neotropical genus of four species, distributed ovary sericeous; fruitcapsular; columella slender, from Panama to Venezuela and Ecuador. persistent; cotyledons plane. Tribe 10. PODOCALYCEAE Webster, trib. This subtribe includes only the single genus Po- nov. TYPE: Podocalyx Klotzsch. docalyx.

Folia alternavel opposita,integra vel serrata;stipulae deciduae; discus 6 nullus;petalae nullae; stamina5-13, 62. Podocalyx Klotzsch, Arch. Naturgesch. 7: antheris extrorsis vel introrsis; pollinis grana 202. 1841. Richeria sect. Podocalyx (Kl.)

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Muell. Arg., DC. Prodr. 15(2): 469. 1866. Subtribe lOc. PARADRYPETINAE G. Levin, TYPE: Podocalyx loranthoides K1. Syst. Bot. 17: 78. 1992. TYPE: Paradrypetes Kuhlmann. A monotypic genus restricted to Amazonian South America. Although Pax & Hoffmann(1922, Leaves opposite, margin ? spinose; stipules en- 1931) and recent workers (e.g., Jablonski, 1967) closing apical bud; staminate flowersin ebracteate have followed Mueller in treating Podocalyx as a cymes, pistillate solitary, the penduncle in both section of Richeria, its pollen is completely dif- sexes adnate to the subtending petiole; staminate ferentand characteristic of the subfamilyOldfield- sepals 3 or 4, free; disk obsolete; stamens 9-13, ioideae. anthers extrorse; pistillode absent; pistillate sepals 4; ovary 3-locular; fruitdrupaceous, 1-seeded; embryo with plicate cotyledons. Subtribe 1Ob. TETRACOCCINAE G. Levin, subtr. nov. TYPE: Tetracoccus Engelm. ex Par- A monogeneric subtribe, considered by Levin to ry. be closest to subtribe Podocalycinae. Fruticesdioicae; foliasimplicia, alterna, opposita, vel verticillata,dentata vel integra;stipulae 0; discus d in- 64. Paradrypetes Kuhlmann, Arq. Inst. Biol. extrorsis; trastaminalis,lobatus; stamina 4-10, antheris Veg. Rio Jan. 2: 84. 1935; Levin, Syst. Bot. pollinisgrana 4-porata,echinata; pistillodio 0; discus 9 1992. TYPE: Paradrypetes ilicifolia glabrus,lobatus; styli elongati; fructus capsularis; semina 17: 74. carunculata,albuminata, cotyledonibus plicatis. Kuhlmann.

Shrubs; leaves alternate, opposite, or whorled, A genus of two South American species, one in margins toothed or entire; stipules absent; stami- the Amazonian rainforests,the other in the Bra- nate flowers in axillary racemes or panicles, or zilian Atlantic rainforest. fasciculate; pistillateflowers solitary or fasciculate, axillary; staminate sepals 4-10, discrete; disk in- Tribe 11. CALETIEAE Muell. Arg., Bot. Zeit. trastaminal, lobed; stamens 4-10, anthers ex- 22: 34. 1865; DC. Prodr. 15(2): 190. 1866; trorse; pollen grains 4-porate, tectum verruculose Grining, Pflanzenr. 147 (Heft 58): 21. 1913. between spines; pistillode absent; pistillate sepals TYPE: Caletia Baillon [= Micrantheum Desfi. 5-13; disk lobed; ovary (2-)3-4(-5)-locular; styles trees or shrubs; leaves elongated; fruit capsular; seeds carunculate; en- Dioecious or monoecious or verticellate, simple, entire dosperm abundant; cotyledons plicate. alternate, opposite, or dentate, stipules obscure or absent; flowersapet- This monogeneric subtribe is very difficultto alous, in axillary clusters or panicles; staminal se- place and perhaps should constitutea distincttribe. pals 4-8(-12), free; disk present or absent; sta- Vegetatively, Tetracoccus is intermediatebetween mens 4-30, filamentsusually free; anthers extrorse Podocalyx and Paradrypetes in phyllotaxy and or latrorse; pollen grains spheroidal, with exinous leaf margins. In some characters, such as the in- spines, colporate or zonoporate; pistillode present trastaminal staminate disk and verruculose pollen or absent; pistillate sepals 3-8(- 13), free; disk sexine, there is a closer resemblance between Tet- present or absent; ovary 2-5-locular; styles un- Levin racoccus and other tribes of Oldfieldioideae. lobed, + elongated, often dilated; fruit capsular; & Simpson (1994) would refer the Tetracoccinae seeds usually 1 per coccus, carunculate or eca- to the tribe Picrodendreae. runculate; endosperm usually copious; cotyledons much broader or scarcely broader than the radicle. 63. Tetracoccus Engehnann ex Parry, W. Amer. with 15 genera Scientist 1: 13. 1885; Croizat, Bull. Torrey This variable tribe of 4 subtribes except for Bot. Club 69: 456. 1942; Dressler, Rhodora is entirely paleotropical, and indeed are nearly confined to 56: 49. 1954. TYPE: Tetracoccus dioicus Par- Hyaenanche, the genera species of Austrobuxus in ry. Australasia (with a few Indonesia). HalliophytumI. M. Johnston,Contr. Gray Herb. 68: 88. 1923. TYPE: Halliophytumfasciculatum (S. Wats. ) I. M. Johnston[= Tetracoccusfasciculatus KEY TO THE SUBTRIBES OF TRIBE CALETIEAE (S. Wats.) Croizat;lectotype]. 1a. Leaves whorled(at least in part); filaments shorterthan anthers;pollen tectum granulose, As interpreted by Dressler (1954), a genus of exinousspines long - 1 a. Hyaenanchinae four species endemic to southwesternNorth Amer- lb. Leaves alternateor opposite;filaments longer ica (southern California and adjacent Arizona, than anthers;pollen tectum smooth. southern Baja California, and northern Mexico). 2a. Pollen grainszonoaperturate; stipules de-

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ciduous or reduced; leaves oppositeor if Toxicodendrumcapense Thunb. [= Hyaenanche alternatethen capsule fleshy. globosa (Gaertner)Lambert]. 3a. Leaves opposite; stamens free; anthersnot apiculate;capsule dry;pis- A monotypic genus of the Cape region of South tillatesepals persistent. Africa. The seeds of Hyaenanche are unusual with- ...... Dissiliariinae in the Oldfieldioideae in the scanty endosperm, 3b. Leaves alternate;stamens connate; green cotyledons, and production of a unique toxin anthersapiculate; capsule fleshy; pistillatesepals deciduous (Hyaenanchin). .11...... I Ic. Petalostigmatinae 2b. Pollen grainspantoporate; stipules decid- Subtribe 1 lb. DISSILARIINAE Pax & Hoff- uous or persistent;leaves alternate;cap- mann, Pflanzenr. 147. XV (Heft 81): 288. sule dry.lId. Pseudanthinae 1922. TYPE: Dissiliaria F. Muell.

Monoecious or dioecious trees or shrubs; leaves Subtribe 1 la. HYAENANCHINAE Baillon ex opposite, simple, entire; stipules reduced or absent; Muell. Arg., Linnaea 34: 64. 1865 (as Hyae- flowers in axillary panicles or clusters; staminate nancheae); DC. Prodr. 15(2): 479. 1866. Tribe sepals 4-6, imbricate, free; disk absent; stamens Hyaenancheae (Baillon ex Muell. Arg.) Hutch., 8-50, free; pollen grains with long to short spines, Amer. J. Bot. 56: 746. 1969. TYPE: Hyaenan- tectum smooth; pistillode absent; pistillate sepals che Lambert. 4-6, free; disk cupulate or absent; ovary 2- or 3- (rarely 4-)locular, glabrous; styles short; fruitcap- ToxicodendrinaePax, Natiirl.Pflanzenfam. ed. 1, 3(5): 31. 1890; Pax & Hoffm.,Pflanzenr. 147. XV (Heft sular; seeds carunculate or ecarunculate. 81): 284. 1922. TYPE: ToxicodendrumThunb. (non This Australasian subtribeincludes 6 genera with P. Miller)[= Hyaenanche Lambert]. ca. 25 species. Generic limits require furthercrit- Dioecious trees or shrubs; leaves opposite or ical study. verticillate,simple, coriaceous, stipules obscure or absent; flowersin axillary clusters or panicles; sta- KEY TO THE GENERA OF SUBTRIBE DISSILARIINAE minate sepals 4-8(-12), ? connate; disk absent; la. Pollen exine with elongatedspines; pistillode stamens 8-30, filamentsfree, much shorter than mostlyabsent. anthers; anthers introrseor extrorse; pollen grains 2a. Dioecious;stamens 8-25; pistillatedisk cu- 6- or 7-zonoporate, spinose, tectum granulose; pis- pular or absent. 3a. Seeds withfimbriate caruncle; ovary tillode absent; pistillatesepals 3-8, free, deciduous 3(-4)-locular;styles ovate, erect. in fruit;disk absent; ovary 3-4-locular, sericeous; ...... A ustrobuxus styles elongated, ? dilated; fruit capsular; seeds 3b. Seeds ecarunculate. carunculate; endosperm sparse. 4a. Styleselongated, recurved; ovary 3(-4)-locular.-.. . 67. Dissiliaria This subtribe is here circumscribed to include 4b. Styles ovate, erect; ovary only the type genus Hyaenanche, as treated by 2-locular...-.. .. 68. Canaca 2b. Monoecious;stamens ca. 50; pistillatedisk Mueller and Pax & Hoffmann. Clearly Hyaenan- of subulatesegments ...-... 69. Whyanbeelia che is a very isolated genus whose relationships l b. Pollenexine with reduced spines; pistillode pres- are stilluncertain. It is isolated in the tribeCaletieae ent. because of the verruculose microsculpturingof the 5a. Stamens4-6; stylesseparated at base, re- pollen sexine, which recalls genera of tribe Picro- curved;seeds ecarunculate... 70. Choriceras 5b. Stamens8-17; stylesproximate, not .re- There is some resemblance to the Sri dendreae. curved;seeds carunculate.-.. 71. Longetia Lankan genus Mischodon in habit and pollen morphology, but it still is not clear whether the two 66. Austrobuxus Miquel, Fl. Ned. Ind. Suppl. be genera should grouped together. 444. 1861; Steenis, Blumea 12: 362. 1964; Airy Shaw, Kew Bull. 25: 506. 1971; Kew 65. Hyaenanche Lambert, Descr. Cinchona 52, Bull. 29: 303. 1974; Kew Bull. 35: 597. t. 10. 1797; Muell. Arg., DC. Prodr. 15(2): 1980; A. C. Smith, Fl. Vitiensis Nova 2: 495. 479. 1866; Dyer, Gen. S. Afr. Fl. P1. 1: 311. 1981; McPherson & Tirel, Fl. Nouv.-Cal6d. 1975. TYPE: Hyaenanche globosa (Gaertner) 14(1): 193. 1987. TYPE: Austrobuxus nitidus Lambert. Miquel.

ToxicodendrumThunberg, Kongl. Vetensk. Acad. Handl. Bureavia Baillon,Adansonia 11: 83. 1873. TYPE: Bur- 17: 188. 1796 (nonToxicodendron P. Miller,1754); eavia carunculata(Baillon) Baillon [= Austrobuxus Bentham,Gen. P1. 3: 280. 1880; Pax & Hoffm., carunculatus(Baillon) Airy Shaw]. Pflanzenr.147. XV (Heft 81): 284. 1922. TYPE: ChoriophyllumBentham, Hook. Icon. P1. 13: 62, t.

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1280. 1879. TYPE: Choriophyllummalayanum 7 1. Longetia Baillonex Muell. Arg.,DC. Prodr. Bentham[= Austrobuxusnitidus Miq.]. 15(2): 244. Aug. 1866; Baillon,Adansonia I. In the circumscriptionof AiryShaw (1974) and 6: 352, t. 9. Oct. 1866; Bentham,Gen. P1. McPherson & Tirel (1987), Austrobuxus is the 3: 279. 1880; Pax & Hoffm.,Pflanzenr. 147. largestgenus of subfamilyOldfieldioideae, with ca. XV (Heft81): 289. 1922; McPherson& Ti- 20 species. The distributionis disjunct,with areas rel,Fl. Nouv.-Cal6d.14(1): 188. 1987. TYPE: in Malaya and westernIndonesia, Queensland, New Longetia buxoides Baillonex Muell. Arg. Caledonia,and Fiji, but absentfrom New Guinea. As interpretedby McPherson& Tirel,Longetia is a monotypicgenus endemicto New Caledonia. 67. Dissiliaria F. Muellerex Baillon,Adansonia Pax & Hoffmann(1922) construedLongetia in a I. 7: 366. 1867; Bentham,Fl. Austral.6: 90. broad sense, includingspecies now placed in Aus- 1873; Pax & Hoffm.,Pflanzenr. 147. XV trobuxus.Airy Shaw (1971, 1974) held a similar (Heft 81): 291. 1922; AiryShaw, Muelleria broadview, except that Longetia specieswere now 4: 220. 1980; Kew Bull. 35: 625. 1980. transferredto Austrobuxus.In viewof the distinct TYPE: Dissiliaria baloghioides F. Muell. ex differencein pollen, McPherson's returnto the Baillon[lectotype, selected here; the generic classicalconcept of Baillon and Muellerseems fully name was associatedwith that species in Bail- justified. lon's originaldescription]. Accordingto AiryShaw (1980), Dissiliaria in- Subtribe lI c. PETALOSTIGMATINAE Pax cludesthree species endemic to Australia(Queens- & Hoffmann,Pflanzenr. 147. XV (Heft 81): land). 281. 1922. Tribe Petalostigmateae(Pax & Hoffm.)Webster, Taxon 24: 595. 1975. TYPE: 68. Canaca Guillaumin,Arch. Bot. Caen 1: 74. Petalostigma F. Muell. 1927; AiryShaw, Kew Bull. 25: 508. 1971. Dioecioustrees or shrubs;leaves alternate,sim- TYPE: Canaca vieillardii Guill. ple, entire;stipules caducous; flowersin axillary As noted by Airy Shaw (1971), Canaca was clusters;staminate sepals 4-6, imbricate;disk ab- originallymisplaced in the Monimiaceaeby Guil- sent;stamens 20-40, filamentsconnate below into lauminbut recognizedas relatedto Austrobuxus a column;anther connective typically pilose; pollen by McKee. McPherson & Tirel (1987) followed grainswith short spines; pistillodeobscure or ab- AiryShaw in relegatingCanaca to synonymyof sent; pistillatesepals 4-8, imbricate;disk absent; Austrobuxus. However, ifDissiliaria is maintained ovary 3-4-locular,pubescent; styles dilated; fruit as distinctfrom Austrobuxus, it appears that Can- a fleshycapsule; seeds carunculate. aca is equallydeserving of recognition. It is possible A monogenericsubtribe restricted to Australia thatall threegenera should be combinedunder the and New Guinea (Papua). name Austrobuxus.

72. Petalostigma F. Mueller, Hook. J. Bot. 69. Whyanbeelia AiryShaw & Hyland, Kew Kew Gard. Misc. 9: 16. 1857; Muell. Arg., Bull. 31: 375, fig. 2. 1976; Kew Bull. 35: DC. Prodr. 15(2): 273. 1866; Baillon,Adan- 691. 1980. TYPE: Whyanbeelia terra-reginae sonia I. 7: 352. 1867; Benth.,Fl. Austral.6: AiryShaw & Hyland. 92. 1873; Gen. P1. 3: 277. 1880; Pax & A monotypic genus of tropical Australia Hoffm.,Pflanzenr. 147. XV (Heft 81): 281. (Queensland). 1922; AiryShaw, Kew Bull. 31: 366. 1976; Kew Bull. 35: 661. 1980. TYPE: Petalostigma quadriloculare F. Muell. 70. Choriceras Baillon,Adansonia I. 11: 119. 1873; AiryShaw, Kew Bull. 14: 356. 1961; As treatedby AiryShaw (1980), Petalostigma Muelleria4: 220. 1980; Kew Bull. 35: 604. is a genus of six variableAustralian species, one 1980. TYPE: Choriceras australiana Baillon of whichextends to New Guinea (Papua). [= Choriceras tricorne(Bentham) Airy Shaw]. Subtribe lId. PSEUDANTHINAE Muell. A genus of two species restrictedto tropical Arg.,Linnaea 34: 55. 1865; DC. Prodr.15(2): Australiaand Papua. AlthoughPax & Hoffmann 195. 1866. TYPE: Pseudanthus Sprengel. (1922) reduced Choricerasto a synonymof Dis- siliaria, AiryShaw (1961) pointedout its distin- Monoecious(rarely dioecious) trees, shrubs, or guishingcharacters. herbs;leaves mostlyalternate, entire; stipules de-

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ciduous or persistent;flowers in axillaryclusters; familyCrotonoideae by McPhersonbecause of the staminatesepals 3-6, imbricate,free; disk intra- uniovulatecarpels, but the pollenclearly suggests staminalor absent; stamens3-20 (rarelymore), placementin the Oldfieldioideae.The genus ap- filamentsfree or connate;pollen grains with long pears very similarto Choriceras, but differsin or shortspines, tectum smooth; pistillode present ovule numberand pollenmorphology. or absent;pistillate sepals 4-6, free,usually persistent;disk present or absent;ovary 2-3-locular; 75. Neoroepera Muell. Arg. & F. Mueller,DC. stylesunlobed, dilated or elongated;fruit capsular; Prodr. 15(2): 488. 1866; Bentham,Fl. Aus- seeds carunculate. tral. 6: 116. 1873; Gen. P1. 3: 276. 1880; The subtribePseudanthinae is here circum- AiryShaw, Kew Bull. 35: 658. 1980. TYPE: scribedin concurrencewith suggestions by Levin Neoroeperabuxifolia Muell. Arg. & F. Muell. (pers. comm.) to includesix Australasiangenera. A genus to two species endemic to Australia The referralof Kairothamnusand Scagea to the (Queensland). Pseudanthinaeneeds confirmation,but appears compatiblewith the anatomicaland palynological evidence. 76. Micrantheum Desf., Mem. Mus. Hist. Nat. Paris 4: 253, t. 14. 1818; Muell. Arg., DC. KEY TO THE GENERA OF SUBTRIBE PSEUDANTHINAE Prodr. 15(2): 195. 1866; Bentham,Fl. Aus- la. Cotyledonsbroader than radicle; trees or shrubs. tral. 6: 58. 1873; Gen. P1. 3: 262. 1880; 2a. Staminatesepals 4; stamens10-12. Grining,Pflanzenr. 147. (Heft58): 23. 1913; -...... 73. Kairothamnus Pax & Hoffm.,Natiirl. Pflanzenfam. ed. 2, 2b. Staminatesepals (5)6; stamens4-6. 19c: 224. 1931. TYPE: Micrantheum eri- 3a. Ovules1 perlocule; pistillode present; coides staminateinflorescence racemose or Desf. umbellate;stipules conspicuous but CaletiaBaillon, Etude Gen. Euphorb. 553. 1858. TYPE: deciduous.74. Scagea Caletiamicrantheoides Baillon, nom. illeg. [= Mi- 3b. Ovules2 perlocule; pistillode absent; crantheumhexandrum Hook. f.]. staminateflowers in axillaryglomer- AlleniaEwart, Proc. Roy. Soc. Vict.II. 22: 7. 1909. ules;stipules minute or obsolete. TYPE: Alleniablackiana Ewart, nom. illeg. [= Mi- -...... 75. Neoroepera crantheumdemissum F. Muell.]. lb. Cotyledonsscarcely broader than radicle; herbs or undershrubs. A genus of threeAustralian species. 4a. Stipulesfoliose; fruits 3-seeded; pollen grains withlong exinous spines 76. Micrantheum 4b. Stipulesnot foliose; fruits 1-seeded; pollen 77. Pseudanthus Sieberex Sprengel,Syst. Veg. grainswith reduced spines. 4(2): 22, 25. 1827; Muell. Arg.,DC. Prodr. 5a. Stamensfree; pistillode present. 15(2): 196. 1866; Bentham,Fl. Austral.6: -...... 77. Pseudanthus 5b. Stamensconnate; pistillode absent. 58. 1873; Gen. P1. 3: 262. 1880; Grining, -...... 78. Stachystemon Pflanzenr.147. (Heft 58): 26. 1913. TYPE: Pseudanthuspimeleoides Sieber ex Sprengel. 73. Kairothamnus AiryShaw, Kew Bull. 34: ChrysostemonKlotzsch, PI. Preiss.2: 232. 1848. TYPE: 596. 1980; Kew Bull. Add. Ser. 8: 121, t. 1 Chrysostemonvirgatus Ki. [= Pseudanthusvir- fig. 1. 1980. TYPE: Kairothamnusphyllan- gatus(Ki.) Muell.Arg.]. thoides(Airy Shaw) AiryShaw. ChorizothecaMuell. Arg., Linnaea 32: 76. 1863. TYPE: Chorizothecamicrantheoides Muell. Arg. [= Pseu- A monotypicgenus of eastern New Guinea(Mo- danthusvirgatus (Ki.) Muell.Arg.]. robe District).The alternatecaudate-acuminate An Australiangenus of seven or eightspecies. leaves of this plant indeed give it an aspect sug- gestiveof Phyllanthus,as noted by Airy Shaw, but its pollenis typicalof the Oldfieldioideae,and 78. Stachystemon Planchon,Hook. LondonJ. the genus appears veryclose to Austrobuxus. Bot. 4: 471, t. 15. 1845; Muell. Arg., DC. Prodr. 15(2): 198. 1866; Bentham,Fl. Aus- 74. Scagea McPherson,Bull. Mus. Nat. Hist. tral. 6: 61. 1873; Gen. P1. 3: 263. 1880; Nat. Paris, IVB. 7: 247. 1985; Fl. Nouv.- Griining,Pflanzenr. 147 (Heft58): 32. 1913; Caled. 14(1): 90. 1987. TYPE: Scagea de- Pax & Hoffm.,Natiurl. Pflanzenfam. ed. 2, pauperata (Baillon)McPherson. 19c: 226. 1931. TYPE: Stachystemonver- micularePlanch. A genus of two species endemicto New Cale- donia. The genus was originallyreferred to sub- A genus of threespecies in westernAustralia.

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Tribe 12. PICRODENDREAE (Small) Web- The four American genera of tribe Picroden- ster, Taxon 24: 595. 1975. Family Picro- dreae are all referable to this subtribe. dendraceae Small, J. New York Bot. Gard. 18: 184. 1917. TYPE: PicrodendronPlan- KEY TO THE GENERAOF SUBTRIBE PICRODENDRINAE chon. la. Leaves mostly3-5-foliolate. 2a. Fruitsdehiscent; staminate sepals not re- Monoeciousor dioecioustrees or shrubs;leaves duced. alternateor opposite,3-9-parted (rarelyunifolio- 3a. Stamens6; pistillateflowers solitary late), entire;stipules caducous, fused to petiole,or -----..-79. Celaenodendron absent;flowers in axillarycymes, racemes, or glom- 3b. Stamens8-15; pistillateflowers race- erules; staminatesepals 4-8 (or obsolete),free; mose- 80. Piranhea 2b. Fruitsindehiscent, drupaceous; staminate se- disk intrastaminalor absent; stamens4-50, free; pals reduced- 81. Picrodendron anthersextrorse or introrse;pollen grains sphe- lb. Leavessimple (unifoliolate), stipellate; pistillate se- roidal,brevicolporate or porate,with exinous spines, pals foliose,persistent -- 82. Parodiodendron tectumgranulose; pistillode minute or absent;pistillate sepals 4-8, persistentor deciduous; disk 79. Celaenodendron Standley, Contr. Dudley presentor absent;ovary 2-3-locular; styleselon- Herb. 1: 76. 1927. TYPE: Celaenodendron mex- gated or stigmatiform;fruit capsular or drupa- icanum Standley. ceous; seeds carunculateor ecarunculate;endo- A monotypicgenus of westernMexico (Tres Ma- sperm usually copious; cotyledonsbroader and rias Islands, Jalisco). longerthan radicle. This tribeof ten generain threesubtribes con- 80. Piranhea Baillon, Adansonia I. 6: 235. 1866; trastsmarkedly with the Caletieae in its distribu- Muell. Arg., Fl. Brasil. 11(2): 79. 1873; Ben- tion:all thegenera are eitherAmerican or African/ tham, Gen. P1. 3: 281. 1880; Pax & Hoffm., Madagascan, withone genus (Mischodon) in Sri Pflanzenr. 147. XV (Heft 81): 295. 1922; Ja- Lanka. The tribeis generallywell characterized by blonski,Mem. New York Bot. Gard. 17: 121. its compoundleaves, althoughthe leaf blade ap- 1967. TYPE: Piranhea trifoliolata Baillon. pears unifoliolatein Mischodon, some species of A genus of two species endemic to South America. Androstachys,and Parodiodendron. 81. Picrodendron Planchon, Hook. London J. KEY TO THE SUBTRIBES OF TRIBE PICRODENDREAE Bot. 5: 579. 1846; nom. cons. prop.; Krug & 1a. Stipulescaducous or obsolete, free from petiole. Urban, Bot. Jahrb. 15: 308. 1893; Fawc. & 2a. Staminatesepals free; capsule septicidal Rend., J. Bot. 55: 268. 1917; Hayden et al., ...... 12a. Picrodendrinae 2b. Staminatesepals connate; capsule loculi- J. Arnold Arbor. 65: 110. 1984. TYPE (typ. cidal...... 12b. Paivaeusinae cons.): Picrodendron baccatam (L.) Krug & lb. Stipulespersistent, + adnateto petiole;seeds Urban. ecarunculate...... 12c. Mischodontinae A monotypicgenus endemic to the West Indies (Greater Antilles,Bahamas, Cayman Islands, Swan Subtribe 12a. PICRODENDRINAE (Small) Islands). Webster,stat. nov. Family Picrodendraceae Small,J. New YorkBot. Gard. 18: 184. 1917. 82. Parodiodendron Hunziker,Kurtziana 5: 331, TYPE: PicrodendronPlanchon. fig. 1. 1969. TYPE: Parodiodendron margini- Dioecioustrees or shrubs;leaves alternate; leaves villosam(Speg.) Hunziker. 3-foliolateor 1-foliolate,margins entire; stipules A monotypicgenus of northernArgentina. small and caducous or absent; inflorescencesaxillary,cymose, sometimes reduced to catkinsor Subtribe 12b. PAIVAEUSINAE Pax & Hoff- glomerules;staminate sepals 4-8, freeor obsolete; mann,Pflanzenr. 147. XV (Heft81): 294. 1922. disk intrastaminalor absent; stamens5-15, free; TYPE: Paivaeasa Welw. ex Bentham [= Old- pollengrains 5-8-porate or brevicolporate,sexine fieldia Benth. & Hook. f. withlong spines and verruculosetectum; pistillode minuteor absent;pistillate sepals 4-8, deciduous; Dioecious trees; leaves alternate or opposite, 5- diskcupular or absent;ovary 2-or 3-locular;styles 8-foliolate;stipules obsolete; staminateinflorescences elongatedor stigmatiform;fruit capsular or dru- in pedunculate ? congested cymes, the pistillateusu- paceous; seeds 1 or 2 per locule, carunculateor ally reduced to 1-3 flowers;staminate sepals 5-8, ecarunculate. basallyconnate; disk intrastaminal,lobed; stamens

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4-12, free, anthers extrorse; pistillode minute or lb. Leaves opposite;staminate disk or pistillode(or absent; pistillatesepals 5-8, persistent;disk annular; both)absent. 2a. Stamenswith filaments elongated, free; leaves ovary 2-3-locular; styles dilated, lobed; capsule tar- simple. dily loculicidallydehiscent; seeds carunculate. 3a. Staminatedisk absent; pistillode present; pistillatesepals deciduous 85. Mischodon As circumscribedhere, in agreementwith the stud- 3b. Staminatedisk present; pistillode absent; ies of Levin, subtribePaivaeusinae includes only the pistillatesepals accrescent in fruit single African genus Oldfieldia. Pax & Hoffmann ------.86. Voatomalo (1922) also included in the Paivaeusinae Piranhea, 2b. Stamenswith filaments united into a long column,anthers subsessile; leaves 1-7-fo- which is here referredto Picrodendrinae,and Aris- liolate. togeitonia, which appears better placed in Mischo- 4a. Leaves unifoliolate;fruit apically de- dontinae. pressed 87. Androstachys 4b. Leaves 3-7-foliolate;fruit beaked ------88. Stachyandra 83. Oldfieldia Bentham & Hooker f., Hook. J. Bot. Kew Gard. Misc. 2: 184. 1850; Bentham, 84. Aristogeitonia Prain, Kew Bull. Misc. Inf. Gen. P1. 3: 281. 1880; Pax & Hoffm.,Pflan- 1908: 439. 1908; Hook. Icon. P1. 30: t. 2926. zenr. 147. XV (Heft 81): 297. 1922; Milne- 1911; Fl. Trop. Afr.6(1): 625. 1912; Pax & Redhead, Kew Bull. 1948: 456. 1948; Leonard, Hoffm.,Pflanzenr. 147. XV (Heft 81): 296. Bull. Jard.Bot. Brux. 26: 338. 1956; Radcliffe- 1922; AiryShaw, Kew Bull. 26: 495. 1972; Smith, Fl. Trop. E. Afr.,Euphorb. 114. 1987. Radcliffe-Smith,Fl. E. Trop. Euphorb.1: 118. TYPE: Oldfieldia africana Benth. & Hook. 1987; Radcliffe-Smith,Kew Bull. 43: 627. 1988. Paivaeusa Weiw.ex Benth.,Gen. P1. 1: 993. 1867. TYPE: TYPE: Aristogeitonialimoniifolia Prami. Paivaeusa dactylophyllaWeiw. ex Oliver[= Old- fieldia dactylophylla(Weiw. ex Oliver)Leonard]. ParageloniumLeandri, Bull. Soc. Bot.France 85: 231. Cecchia Chiovenda,FR. Somala 2: 397, fig.227. 1932. 1939.TYPE: Parageloniumperrieri Leandri [= Aristo- geitoniaperrieri (Leandri) Radcl.-Sm.]. TYPE: Cecchia somalensisChiovenda [= Oldfieldia somalensis(Chiovenda) Milne-Redhead]. An Africangenus of three species (Angola, Kenya, As delimited by Leonard (1956) and Radcliffe- Tanzania,and Madagascar).Radcliffe-Smith (1988) Smith (1987), Oldfieldia includes four African spe- has recentlydemonstrated that Paragelonium from cies. The distinctionin phyllotaxy(alternate vs. op- Madagascarcannot be separatedfrom the mainland posite leaves) used by Pax & Hoffmannto distinguish AfricanAristogeitonia. Paivaeusa is clearly unworkable,because alternate and opposite leaves occur in both Paivaeusa and 85. Mischodon Thwaites,Hook. J. Bot. Kew Gard. Oldfieldia. Misc. 6: 299. 1854; Muell. Arg.,DC. Prodr. 15(2): 1124. 1866; Bentham,Gen. P1. 3: 280. Subtribe 12c. MISCHODONTINAE Muell.Arg., 1880; Hook. f.,Fl. Br. Ind. 5: 344. 1887; Pax Linnaea 34: 202. 1865; DC. Prodr. 15(2): 1124. & Hoffm.,Pflanzenr. 147. XV (Heft81): 292. 1866; Webster, Taxon 24: 595. 1975. TYPE: 1922; Raju, J. Econ. Tax. Bot. 5: 165. 1984. Mischodon Thwaites. TYPE: Mischodonzeylanicus Thwaites.

AndrostachydaceaeAiry Shaw, Kew Bull. 18: 250. 1965. A monotypicgenus of Sri Lanka; perhapsnative TYPE: AndrostachysPrain. (butvery rare) in southernIndia. Monoecious or dioecious trees or shrubs; leaves 86. Voatomalo Capuronex Bosser,Adansonia II. alternate or opposite, simple or 3-foliolate,entire; 15: 333. 1976. TYPE: Voatomaloeugenioidea stipulespersistent, adnate to petiole;flowers in axillary Capuronex Bosser. cymes or clusters; staminate sepals (2-)5-8, free; disk intrastaminalor absent; stamens 5-50, free; This genusof twospecies from Madagascar was pistillodepresent or absent; pistillatesepals 5 or 6, mentionedby Leandri(1958) butnot formallypub- deciduous; disk annular or absent; ovary 3-4(-5)- lisheduntil 1976. locular, glabrous or pubescent; styles slightlyto distinctlyconnate below; fruitcapsular; seeds ecarun- 87. Androstachys Prain, Kew Bull. Misc. Inf. culate. 1908. 438. 1908; Fl. Trop. Afr.6(1): 740. 1912; Pax & Hoffm.,Pflanzenr. 147. XV (Heft KEY TO THE GENERA OF SUBTRIBE MISCHODONTINAE 81): 287. 1922; Leandri,Fl. Madag. 111: 197. la. Leaves alternate;pistillode present 1958; AiryShaw, Kew Bull. 18: 251. 1965; ------84. Aristogeitonia AdansoniaII. 10: 519. 1970; Dyer, Gen. S.

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Afr.Fl. PI. ed. 3, 312. 1975; Leroy,C. R. leaves alternate (rarely opposite), usually stipulate; Acad. Sci. Paris283D: 147. 1976; Alvin,Ann. blade simple or palmately lobed (rarely compound), Bot. 59: 579. 1987. TYPE: Androstachysjohn- often with foliar glands. Inflorescences axillary or sonii Prain[= Androstachyssubpeltata (Sim) terminal,racemose or spicate, or reduced to glom- Phillips]. erules or solitaryflowers; bracts sometimesglandular. Staminate flower: sepals imbricate or often valvate If Stachyandrais recognizedas a distinctgenus, (and then often connate and rupturingat anthesis); Androstachysbecomes a monotypicgenus confined petals and disk presentor absent; stamens 2-100+, to easternAfrica. free or connate; pollen grains mostly 3-4-colporate, exine semitectate (very rarely echinate); male ga- 88. Stachyandra Leroyex Radcliffe-Smith,Kew metophyte binucleate; pistillode present or absent. Bull.45: 562. 1990. TYPE: Stachyandramera- Pistillate flower: sepals (2-)3-6(-12), imbricate or na (AiryShaw) Leroyex Radcl.-Sm. open at anthesis,sometimes connate; petals and disk This genus of fourMadagascan species has represent or absent; ovary mostly 2-4-locular; styles centlybeen segregated from Androstachys. The two entireto bifidor multifid;ovules solitaryin each locule taxa are closelyrelated, and it mightbe more ap- (except in Dicoelia), anatropous. Fruit capsular or propriateto treat Stachyandra as a subgenusof rarely baccate or drupaceous; seeds carunculate or Androstachys. ecarunculate,testa sometimesfleshy; endosperm usually copious; cotyledonslonger and broader than rad- Subfamily III. ACALYPHOIDEAE Ascherson, icle. Base chromosomenumbers mostly 9, 10, or 1 1. Fl. Prov. Brandenb.1: 58. 1864. FamilyAca- With 116 genera in 20 tribes, subfamilyAcaly- lyphaceaeKlotzsch & Garcke,Monatsber. K6n- phoideae is the largest and most complex of the igl.Preuss. Akad. Wiss. Berlin 1859: 246. 1859. subfamiliesof Euphorbiaceae. Through Clutia it ap- TYPE: Acalypha L. proaches the Phyllanthoideae, and through Om- Trees,shrubs, or herbs;milky latex absent (rarely phalea both the Crotonoideae and Euphorbioideae. present,or representedby coloredexudate); indu- These multipleconnections suggest that the Acaly- mentumsimple or hairsoften branched or lepidote; phoideae may not be a monophyleticgroup.

KEY TO THE TRIBES OF SUBFAMILY ACALYPHOIDEAE 1a. Staminateflowers with 5 imbricatesepals and petals,5 stamens,and perforate-tectatepollen grains; indumentum simpleor malpighiaceous;dioecious; fruit capsular. 2a. Petals not adaxiallybarbate; filaments connate; disk segments glandular-lobed; seeds carunculate;leaves usuallypellucid-punctate, indumentum simple -13. CLUTIEAE 2b. Petalsadaxially barbate; filaments free; disk segments not glandular-lobed;seeds ecarunculate;leaves not pellucid-punctate- 14. POGONOPHOREAE lb. Staminateflowers not with 5 imbricatesepals and petalsand 5 stamens(or ifso, thenpollen not perforate-tectate or fruitnot capsular). 3a. Staminatesepals distinctly imbricate, or else flowers enclosed in bibracteolate involucre; flowers apetalous; dioecioustrees or shrubs. 4a. Leaves alternate;pollen exine rugulose or micropunctate. 5a. Filamentsconnate; pistillode conspicuous; pollen tectum not spinulose; seeds carunculate, testa dry. 6a. Flowersin axillaryglomerules, not involucrate;stamens 8-15; styleselongated, bifid; capsuleechinate; leaves stipulate -15. CHAETOCARPEAE 6b. Flowersenclosed in bibracteolateinvolucres; stamens 2-6; stylesabbreviated, dilated; capsulenot echinate; leaves exstipulate -16. PEREAE 5b. Filamentsfree; pistillode small or absent;pollen tectum spinulose; seeds ecarunculate,testa fleshy------17.CHEILOSEAE 4b. Leaves opposite;pollen exine reticulate; seeds ecarunculate,testa dry 18. ERISMANTHEAE 3b. Staminatesepals valvate or slightlyimbricate; flowers not in bibracteolateinvolucres; petals present or absent;pollen grains various, not micropunctate; indumentum various. 7a. Staminatepetals with depressions opposite anthers, or else fruit drupaceous with channeled endocarp. 8a. Ovules2 per locule;fruit capsular; plants monoecious; styles unlobed - 19. DICOELIEAE 8b. Ovules 1 per locule;fruit drupaceous; plants dioecious; styles bifid 20. GALEARIEAE 7b. Staminatepetals lacking depressions opposite anthers; fruit not drupaceous with channeled endocarp. 9a. Cotyledonsscarcely broader than radicle; monoecious (rarely dioecious) herbs or subshrubs withericoid foliage; indumentum simple; pollen grains perforate-tectate, colpi distinctly marginate;seeds carunculate-21. AMPEREAE 9b. Cotyledonsdistinctly broader than radicle; habit otherwise.

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1Oa. Petalspresent, at leastin staminateflowers. 1 la. Dioecious;indumentum simple; anther connective enlarged, anther sacs pendulous; pollensexine coarsely reticulate, not heterobrochate - 22. AGROSTISTACHYEAE 1lb. Monoecious(rarely dioecious); indumentum (at least in part) malpighiaceous, stellate,or lepidote;anthers not with enlarged connective and pendulousanther- sacs; pollensexine often distinctly heterobrochate - 23. CHROZOPHOREAE 1Ob. Petalsabsent. 12a. Stylesfree or basally connate (if long-connate, then styles bifid or indumentum stellate);plants rarely scandent; stinging hairs absent. 13a. Staminatedisk present; pollen exine coarsely reticulate, colpi mostly inoperculate. 14a. Leaves usuallywith laminar glands; indumentum simple or stellate; inflorescenceaxillary or terminal. 15a. Pollencolpi not marginate;staminate disk massive, pubescent;anther connective enlarged ------24. CARYODENDREAE 15b. Pollencolpi marginate; staminate disk tenuous, glabrous; antherconnective not enlarged - 25. BERNARDIEAE 14b. Leaves eglandular;indumentum simple; stamens many; inflorescenceaxillary - 26. PYCNOCOMEAE 13b. Staminatedisk absent (or ifpresent, then pollen colpi operculate). 16a. Pollengrains coarsely reticulate or perforate-tectate;indumentum stellate;stamens inflexed in bud (exceptin Koilodepas); monoecious;pistillate calyx often accrescent; seeds ecarunculate ------27. EPIPRINEA E 16b. Pollengrains finely perforate-tectate to rugulose; indumentum simpleor stellate;stamens not inflexed in bud. 17a. Pollengrains with operculate colpi; staminate disk present or absent;seeds mostlyecarunculate, testa not fleshy; stipulesdeciduous or obsolete. 18a. Leaves eglandular;styles subentire to multifid;staminatedisk (if present) extrastaminal or interstaminal; pollensexine finely perforate-tectate (nanos- pinulesregularly arranged) - 28. ADELIEAE 18b. Leavesusually with embedded laminar glands; styles entireor if dividedthen inflorescencesterminal; staminatedisk (if present) intrastaminal; pollen sexine ruguloseto striate(nanospinules not regularly arranged) 29. ALCHORNEAE 17b. Pollencolpi inoperculate, often reduced; seeds carunculate or ecarunculate,testa dry or fleshy- 30. ACALYPHEAE 12b. Stylesunlobed, mostly connate into a distinctcolumn; often scandent or twining,sometimes with stinging hairs; inflorescences mostly terminal or pseudoaxillary,bisexual; seeds ecarunculate,testa dry. 19a. Staminatesepals valvate, splitting apart at anthesis;latex absent; in- florescencenot paniculate; bracts eglandular; filaments free or connate - 31. PLUKENETIEAE 19b. Staminatesepals imbricate;latex clear or reddish;inflorescence paniculate;bracts foliose and biglandular;stamens 2 or 3, filaments connate;pollen grains colporate, exine rugulose - 32. OMPHALEAE

Tribe 13. CLUTIEAE (Muell.Arg.) Pax, Natiirl. imbricate;petals 5, free; disk dissected, segments 5, Pflanzenfam.ed. 1, 3(5): 81. 1890 (as Cluy- ? palmatelylobed or glandular; stamens 5, filaments tieae). Hippomaneaesubtribe Cluytieae Muell. connate; anthersmuticous, introrse; pollen grains pro- Arg.,Linnaea 34: 202. 1865. Cluytieaesubtribe late, 3-colporate, colpi inoperculate and slightlyto CluytiinaePax, Pflanzenr.147. III (Heft47): distinctlymarginate, exine perforate-tectate;pistillode 49. 1911. TYPE: ClutiaL. present; ovary 3-(rarely 4-)locular, glabrous or pubescent; stylesfree or nearly so, bifid;fruit capsular; Dioecious(rarely monoecious) perennial herbs or seeds carunculate, smooth (or rarelypunctate), testa shrubs;indumentum simple or absent;leaves alter- dry; endosperm copious; cotyledons broader and nate,simple, entire, penninerved, often pellucid-punc- somewhat longer than the radicle. tate,eglandular; stipules small or obsolete;flowers in axillaryglomerules (pistillate often solitary); sepals 5, As here interpreted,the tribeClutieae is restricted

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to the type genus Clutia, in accordancewith the claimed that the membership of the taxon in the originalconcept of Mueller.None of the generaas- Euphorbiaceae is suspect. Airy Shaw (1966) sug- sociatedin subtribeClutiinae by Pax (1911) are at gested that it may be more closely allied to the all closelyrelated to Clutia.Jussieu (1824) hadplaced Icacinaceae. However, the pollen of Pogonophora Clutiain thetribe Phyllantheae, presumably because is similarto thatof Clutia, and the floralresemblances ofits striking similarity in habit.Baillon (1858), who to the Icacinaceae are probably superficial. clearlydemonstrated that Clutia has uniovulatecarpels, consideredthe resemblanceto Briedelia and 90. Pogonophora Miers ex Bentham, Hook. J. Phyllanthusto be superficial,and suggestedan af- Bot. Kew Gard. Misc. 6: 372. 1854; Muell. finitywith Chiropetalum. However, the pollen struc- Arg., DC. Prodr. 15(2): 1040. 1866; Fl. Bras. turedoes not supportBaillon's view, and in factis 11(2): 435. 1874; Bentham, Gen. P1. 3: 288. morecompatible with Jussieu's original suggestion of 1880; Pax, Pflanzenr. 147. III (Heft 47): 108. a relationshipwith the Phyllanthoideae. 1911; Letouzey, Adansonia II. 9: 275. 1969; Secco, Rev. Gen. Anomalocalyx . . . Amer. SWl, 89. Clutia L., Sp. PI. 1042. 1753; Gen. PI. ed. 88. 1990. TYPE: Pogonophora schomburg- 5, 464. 1754; Muell. Arg.,DC. Prodr.15(2): kiana Miers ex Bentham. 1043. 1866; Bentham,Gen. PI. 3; 302. 1880; A genus of two species, one neotropical (Amazo- Pax, Pflanzenr.147. III (Heft47): 50. 1911; nian and coastal Brazil), the other west African. Hutchinson,Fl. Trop. Afr.6(1): 801. 1912; Prain,Fl. Cap. 5(2): 427. 1920; Dyer,Gen. S. Tribe 15. CHAETOCARPEAE (Muell. Arg.) Afr.Fl. PI. 1: 321. 1975; Leonard,Fl. Congo Taxon 24: 595. 1975. Hippomaneae Belge 8(1): 93. 1962; Radcliffe-Smith,Fl. E. Webster, subtribeChaetocarpeae Muell. Arg., Linnaea 34: Trop.Afr., Euphorb. 1: 331.1987. TYPE: Clutia 202. 1865. TYPE: Chaetocarpus Thwaites. pulchella L. trees or shrubs; indumentumsimple or A largeAfrican genus of ca. 75 species,with two Dioecious alternate,simple, entire,penninerved, extendingto tropicalArabia; the greatest diversity is absent; leaves flowers in axillary in SouthAfrica. eglandular; stipules deciduous; bracteate glomerules; staminate sepals 4 or 5, imbricate, free or connate; petals present or absent; 14. Tribe POGONOPHOREAE (Muell. Arg.) disk dissected; stamens 5-15, filamentshirsute, con- Webster,Taxon 24: 595. 1975. Hippomaneae nate below; anthers dehiscing longitudinally;pollen subtribePogonophoreae Muell. Arg., Linnaea grains globose, 3-colporate (colpi withirregular mar- 34: 202. 1865; DC. Prodr.15(2): 1040. 1866. gins), exine rugulose and micropunctate; pistillode TYPE: PogonophoraMiers. prominent,hirsute; pistillatesepals 4-8, imbricate; Dioecioustrees; indumentum simple or (in inflo- disk urceolate; ovary 3-locular; styles bipartite,pa- rescence)malpighiaceous; leaves alternate,simple, pillose; fruitcapsular; seeds smooth,blackish, carun- entire,penninerved, eglandular; stipules very small culate; endosperm copious; cotyledons broader and orobsolete; inflorescences axillary, of thyrses or glom- longer than radicle. erules;staminate sepals 5, coriaceous,broadly im- This taxon, originallyincluding only the type ge- bricate(? biseriate);petals 5, imbricate,adaxially nus, was enlarged by Webster (1975) through the barbate;disk intrastaminal,urceolate, 5-lobed; sta- inclusion of Trigonopleura. The close similarityof mens5, filamentsfree; anthers basifixed, elongated; the pollen in the two genera seems decisive as an pollengrains prolate, 3-colporate, colpi inoperculate, indicationof affinity.The blackish carunculate seeds not marginate,sexine perforate-tectate;pistillode are reminiscentof those of Clutia. There may also present;pistillate perianth as in staminate;disk cu- be a more distantrelationship with Cheilosa, as sug- pulate,tenuous; ovary 3-locular; styles bibbed; fruit gestedby Pax & Hoffmann(1912). AiryShaw (1975) capsular,seeds ecarunculate, smooth, shining, hilum independentlysuggested a tribe Chaetocarpeae (not large; endospermcopious; cotyledons much longer formallypublished) including Chaetocarpus and Tri- and broaderthan radicle. gonopleura, and saw a resemblance to Casearia in The tribePogonophoreae is unigeneric,with the the Flacourtiaceae. same circumscriptionas that of Mueller (1866). The relationshipsof the genus Pogonophora have re- KEY TO THE GENERA OF TRIBE CHAETOCARPEAE mainedproblematical; affinities have been suggested la. Petalspresent; anthers nearly sessile on staminal withMicrandra, Microdesmis, and it has evenbeen column;ovary smooth -- 91. Trigonopleura

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lb. Petalsabsent; filaments well developed; ovary echi- ers apetalous, disk absent; staminate calyx 2-4-fid nate-92. Chaetocarpus or rudimentary;stamens 2-8, filamentsconnate; pollen grains 3-colporate, rugulose; pistillodes (repre- 91. Trigonopleura Hook. f., Fl. Brit.Ind. 5: 399. senting reduced pistillate flowers) surroundingsta- 1887; Hook. Icon. P1. 18: t. 1753. 1888; Pax minateflower. Pistillate flower without perianth; ovary & Hoffm., Pflanzenr. 147. III (Heft 47: 95). 3-locular; styles stigmatiform.Fruit capsular, valves 1911; Merrill, Phil. J. Sci. llC: 76. 1916; woody; seeds smooth,dark, carunculate; endosperm Ridley, Fl. Malay Pen. 3: 263. 1924; Whit- copious; cotyledons much longer and broader than more, Tree Flora Malaya 2: 134. 1973; Airy radicle. Shaw, Kew Bull. Add. Ser. 4: 201. 1975; Kew This tribe,by the unanimous consensus of workers Bull. 36: 350. 1981. TYPE: Trigonopleura ma- genus Pera. layana Hook. f. since Baillon, includes only the single The relationshipsof Pera have been in doubt, how- A monotypicgenus of southeast Asia (Malaya to ever, because of the reduced nature of the flowers the Philippinesand Borneo). and the peculiar inflorescencesthat have no coun- terpartelsewhere in the family.Mueller (1866) placed 92. Chaetocarpus Thwaites, Hook. J. Bot. Kew Pera at the end of the Acalypheae (Acalyphoideae), Gard. Misc. 6: 300, t. l0a. 1854 (nom. cons.); Bentham (1880) at the end of the Hippomaneae, and Muell. Arg., DC. Prodr. 15(2): 1121. 1866; Pax & Hoffmann(1922) followingDalechampieae. Bentham,Gen. P1. 3: 323. 1880; Pax & Hoffm., AiryShaw (Dict. Fl. PI. ed. 7, xxii. 1966) recognized Pflanzenr. 147. IV (Heft 52): 7. 1912; Fawc. Peraceae as a distinctfamily, but withoutdiscussion & Rend., Fl. Jam. 4: 318. 1920; Pax & Hoffm., of its affinities.I believe that the closest relationship Natiirl. Pflanzenfam.ed. 2, 19c: 179. 1931; of Pera may be withtribe Chaetocarpeae, as indicated Leonard, Fl. Congo 8(1): 127. 1962; Capuron, by similarityin the pollen and also the woody capsules Adansonia II. 12: 209. 1972; Whitmore,Tree with smooth carunculate seeds. Fl. Malaya 2: 76. 1973. TYPE: Chaetocarpus pungens Thwaites [= Chaetocarpus castani- 93. Pera Mutis,Kongl. Vetensk.Akad. Nya Handl. carpus (Roxb.) Thw.]. 5: 299. 1784; Muell. Arg., DC. Prodr. 15(2): MetteniaGrisebach, FH. Brit.W. Ind. 43. 1859. TYPE: 1025. 1866; Fl. Bras. 11(2): 421. 1874; Ben- Metteniaglobosa Griseb.[= Chaetocarpusglobosus tham, Gen. PI. 3: 340. 1880; Pax & Hoffm., (Griseb.)Fawc. & Rend.]. Pflanzenr. 147. XIII (Heft 68): 2. 1919; Ja- Regnaldia Baillon,Adansonia I. 1: 87. 1861. TYPE: Reg- blonski,Mem. New York Bot. Gard. 17: 147. naldia cluytioidesBaillon [= Chaetocarpuscastani- carpus(Roxb.) Thw.]. 1967. Perula Schreber, Gen. PI. 703. 1791. NeochevalieraBeille, Compt. Rend. Hebd. Acad. Sci. Paris TYPE: Pera arborea Mutis. 145: 1295. 1907. TYPE: Neochevalierabrazzavillen- sis Beille[= Chaetocarpusafricanus Pax]. Spixia Leandrode Sacrameto,Denkschr. K6nigl. Akad. Wiss. Mfinchen,Math. Phys. 7: 231. 1821 (non A genus of 12 species witha disjunctdistribution: Schrank,1821). TYPE: Spixia heterantheraSchrank West Indies, South America, west Africa,Madagas- [= Pera heteranthera(Schrank) I. M. Johnston]. SchismatoperaKlotzsch, Arch. Naturgesch. 7: 178. 1841. car, and tropical Asia. TYPE: Schismatoperamartiana Ki. [= Pera disticho- phylla (Mart.)Baillon]. Tribe 16. PEREAE (Kl. & Gcke.) Pax & Hoff- mann, Pflanzenr.147. XIII (Heft 68): 1. 1919. A neotropical genus of ca. 30 species, extending Peraceae K1. & Gcke., Monatsber. Konigl. fromCuba and Central America south to Brazil, with Preuss. Akad. Wiss. Berlin 1859: 246. 1859. the greatest concentrationof taxa in the Amazon Acalypheae subtribePerinae (Kl. & Gcke.) Pax, basin. Natuirl. Pflanzenfam. ed. 1, 3(5): 69. 1890. TYPE: Pera Mutis. Tribe 17. CHEILOSEAE (Muell. Arg.) AiryShaw & Webster, Taxon 24: 595. 1975. Hippoma- ProsopidoclineaeKlotzsch, Arch. Naturgesch. 7: 176. 1841. Muell. Arg., Linnaea TYPE: Pera Mutis. neae subtribe Cheiloseae 34: 202. 1865. Gelonieae subtribeChaetocar- Dioecious trees or shrubs; indumentum simple, pinae series Cheilosiformes Pax & Hoffm., stellate,or lepidote;leaves alternate(rarely opposite), Pflanzenr.147. XIV (Heft 68): 50. 1919. TYPE: entire, pinnatelyveined, exstipulate. Flowers in ax- Cheilosa Blume. illary involucrateglomerules; involucre usually of 2 valvate bracts entirelyenclosing flowers in bud. Flow- Dioecious trees; indumentumsimple, stellate, or

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nearlyabsent; leaves alternate,simple, entire or cren- A tropicalAsian genus of four species, distributed ulate, penninerved,glandular or eglandular; stipules fromBurma to the Philippinesand Solomon Islands. caducous; inflorescencesaxillary or sometimes terminal, paniculate, bracts eglandular; flowersapetal- Tribe 18. ERISMANTHEAE Webster,Taxon ous; sepals 4 or 5, imbricate;staminate disk dissected, 24: 595. 1975. TYPE: Erismanthus Wall. ex intrastaminalor extrastaminal;stamens 4-10, free; Muell. Arg. anthers muticous, introrse; pollen grains globose, 3-colporate, colpi inoperculate, exine echinate; pis- Monoecious trees or shrubs; indumentumsimple; tillodepresent; pistillate disk annular or absent; ovary leaves opposite,subsessile, simple, entire or obscurely 2-3-locular; styles bifid; fruitcapsular; seeds eca- dentate,basally oblique, penninerved,stipulate; inflo- runculate,testa fleshy;cotyledons longer and broader rescences axillary, racemose or spicate; staminate than radicle. sepals 4 or 5, imbricateor nearly valvate; petals 5; disk absent; stamens 5-15, filamentsfree; anthers This tribeincludes two genera, but has a different muticous, introrse;pollen grains 3-colporate, angu- circumscriptionfrom the "series" Cheilosiformesof laperturate,colpi inoperculate,not marginate,sexine Pax & Hoffmann (1919, 1931), since Elaterios- reticulate;pistillode present; pistillate sepals 5-6, im- permum is assigned to the subfamilyCrotonoideae, bricate; petals present or absent; disk absent; ovary and Neoscortechinia is added. Airy Shaw (1972) 3-locular; ovules anatropous; styles free to connate, appears to have been the firstto suggest that Neos- entire or bifid;fruit capsular; seeds smooth,ecarun- cortechinia should be placed in the same tribe as culate, testa dry. Cheilosa. This tribeincludes three tropical Asian genera that KEY TO THE GENERAOF TRIBE CHEILOSEAE share some common features with tribe Epiprineae. Erismanthus and Moultonianthus have a distinctive la. Fruitthick-walled, capsular, with 3 seeds; indumentumsimple; stamens 8-10 - 94. Cheilosa habit due to the opposite phyllotaxyand oblique leaf lb. Fruitthin-walled, with 1 seed; indumentumstel- bases; this is present, but less pronounced, in Syn- late; stamens5 or 6 .. 95. Neoscortechinia dyophyllum.

94. Cheilosa Blume,Bijdr. 613. 1826; Muell. Arg., KEY TO THE GENERA OF TRIBE ERISMANTHEAE DC. Prodr.15(2): 1122. 1866; Bentham,Gen. la. Pistillodein staminateflower greatly elongated, P1. 3: 322. 1880; J. J. Sm., Add. Cogn. Fl. clavate; youngstaminate inflorescence covered Arb. Jav. 12: 604. 1910; Pax & Hoffm.,Pflan- withclosely imbricate bracts; pistillate flowers apet- zenr. 147. IV (Heft 52): 12. 1912; Airy Shaw, alous;staminate petals shorter than calyx; stamens 12-15 -96. Erismanthus Kew Bull. 16: 364. 1963; Whitmore,Tree Fl. lb. Pistillodenot greatlyelongated; young staminate Malaya 2: 77. 1973; Airy Shaw, Kew Bull. inflorescencesnot covered with densely imbricate Add. Ser. 4: 68. 1975; Kew Bull. 36: 276. bracts;stamens 5-10. 1981. TYPE: Cheilosa montana Blume. 2a. Stylesdistinctly bifid; staminate petals longer thancalyx; pistillate flowers petaliferous; stip- A genus of two species (perhaps conspecific) in ules persistent,foliaceous, and clasping southeast Asia (Malaya to the Philippinesand Bor- - ...... ------97. Moultonianthus 2b. Stylesentire or nearlyso; staminatepetals neo). shorterthan sepals; pistillate flowers apetalous; stipulessmall, deciduous ------98. Syndyophyllum 95. Neoscortechinia Pax, Natiirl. Pflanzenfam. Nachtr. 1: 213. 1897; Pax & Hoffm., Pflan- zenr. 147. XIV (Heft 68): 52. 1919; AiryShaw, 96. Erismanthus Wallich ex Muell. Arg., DC. Kew Bull. 16: 368. 1963; Whitmore,Tree Fl. Prodr. 15(2): 1138. 1866; Bentham, Gen. P1. Malaya 2: 119. 1973; Airy Shaw, Kew Bull. 3: 325. 1880; Hook. f., Fl. Brit. Ind. 5: 405. 36: 333. 1981. Scortechinia Hook. f., Hook. 1887; Pax & Hoffm.,Pflanzenr. 147. III (Heft Ic. P1. 18: t. 1706. 1887 (non Scortechinia 47): 33. 1911; Gagnep., Fl. Indochine 5: 464. Saccardo, 1885). TYPE: Scortechinia kingii 1926; Airy Shaw, Kew Bull. 26: 260. 1972; Hook. f. [= Neoscortechinia kingii (Hook. f.) Whitmore,Tree Fl. Malaya 2: 95. 1973; Airy Pax & Hoffm.]. Shaw, Kew Bull. Add. Ser. 4: 110. 1975; Kew Bull. 36: 294. 1981. TYPE: 1?rismanthusobli- AlcinaeanthusMerrill, Phil. J. Sci. Bot.7: 379. 1912. TYPE: quus Muell. Arg. Alcinaeanthusphilippinensis Merr. [= Neoscor- techiniaforbesii Hook. f.) Pax ex S. Moore]. An Asian genus of two species, the type species

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occurringfrom Thailand to Sumatraand Borneo,the to be subdividedinto segregate families,this arrange- otherspecies (E. sinensisOliver) in Indochinaand ment still seems logical. However, in the present Hainan. synopsis,where a broad concept ofthe familyis taken, it appears necessary to include the Dicoelieae in the 97. Moultonianthus Merrill,Phil. J. Sci. Bot.11: Acalyphoideae adjacent to the Galearieae. It might 70. 1916; Pax & Hoffm.,Pflanzenr. 147. XIV be possible to recognize a separate subfamilyfor the (Heft68): 41. 1919; Natiirl.Pflanzenfam. ed. tribesDicoelieae and Galearieae, but this seems pre- 2, 19c: 170. 1931; Steenis,Bull. Bot. Gard. matureuntil phylogenetic relationships within the sub- Buitenz.III. 17: 404. 1948; AiryShaw, Kew familyAcalyphoideae are much betterunderstood. Bull. Add. Ser. 4: 176. 1975; Kew Bull. 36: 332. 1981. TYPE: Moultonianthusborneensis 99. Dicoelia Bentham, Hook. Ic. P1. 13: 70, t. Merr. [= Moultonianthusleembruggianus 1289. 1879; Gen.P1. 3: 286. 1880; J.J. Smith, (Boerl.& Koord.)Steenis]. Bull. Jard. Bot. Buitenz. III. 1: 392. 1920; Pax A monotypicgenus known only from Sumatra and & Hoffm.,Pflanzenr. 147. XII (Heft 81): 15. Borneo. 1922; Croizat, J. Arnold Arbor. 23: 38. 1942; Airy Shaw, Kew Bull. 27: 3. 1972; 36: 285. 98. Syndyophyllum Lauterbach& K. Schu- 1981. TYPE: Dicoelia beccariana Benth. mann,Fl. DeutscheSchutzg. 403. 1901; Pax A monotypicgenus of Malaya, Sumatra, and Bor- & Hoffm.,Pflanzenr. 147. III (Heft47): 104. neo according to Airy Shaw (1981), who does not 1911; AiryShaw, Kew Bull. 14: 392. 1960; accept D. affinisJ. J. Sm. as distinct. Hook Ic. P1.38: t. 3722. 1974; Kew Bull.Add. Ser. 4: 199. 1975; 8: 204. 1980. TYPE: Syn- Tribe 20. GALEARIEAE Bentham, Gen. P1. 3: dyophyllumexcelsum Lauterb. & K. Sch. [lec- 247, 287. 1880. Cluytieae subtribeGaleariinae totype]. Pax, Natiirl.Pflanzenfam. ed. 1, 3(5): 81. 1890. A monotypicgenus, the singlevariable species TYPE: Galearia Zoll. & Mor. foundin Sumatra,Borneo, and New Guinea. BennettiaceaeR. Brown,P1. Jav. Rar. 250. 1850 (nom. prov.).Bennettieae R. Brownex Schlizlein,Iconogr. Tribe 19. DICOELIEAE Hurusawa,J. Fac. Sci. Fam. Nat. RegniVegetab. 3: 172. 1860. TYPE: Ben- Univ.Tokyo Sect. 3, Bot. 6: 322. 1954; Web- nettiaR. Brown[non Bennettia S. F. Gray,1821; = ster, Bot. J. Linn. Soc. 94: 6. 1987. TYPE: Galearia Zoll. & Mor.]. Dicoelia L. Pandaceae Pierre, Bull. Soc. Linn. Paris 1255. 1896; Engler& Gilg,Syll. Pflanzenfam. ed. 7, 223. 1913; Monoecioustrees or shrubs;indumentum simple; Forman,Kew Bull. 20: 309. 1966; Webster,Bot. leaves simple,entire, pinnately veined, eglandular; J. Linn. Soc. 94: 6. 1987. TYPE: Panda Pierre. stipulescaducous. Inflorescences bisexual, racemose. Staminatesepals 5, valvateor nearlyso; petals 5, Dioecious trees or shrubs; indumentum simple; valvate,with 2 concavitiesopposite anthers; disk leaves alternate, simple, entire or dentate, pin- absent;stamens 5; filamentsslightly connate to base nately veined, eglandular, stipulate. Inflorescences of3-5-fid pistillode; pollen grains 3-colporate, tectate. terminal thyrses or axillary glomerules; bracts Pistillatesepals 5, valvate;petals 5, slightlyimbricate; sometimes persistent.Staminate sepals free or con- disk absent;ovary 3-locular;ovules 2 per locule, nate, slightlyimbricate or valvate; petals valvate anatropous;styles free or nearlyso, unlobed,slender. or somewhat imbricate, with ventral paired de- Fruitcapsular; seeds ecarunculate. pressions opposite anthers; disk absent; stamens 5- 15, free; anthers introrse;pistillode unlobed, some- A monogenerictribe, referred to subfamilyPhyl- times peltate; pollen grains subglobose, angulap- lanthoideaeby Pax & Hoffmann(1922) because of erturate, 3-colporate, sexine finelyreticulate. Pis- the biovulatecarpels. Although Airy Shaw (1981) tillate sepals free or connate; petals valvate; ovary suggestedan affinityto Trigonostemonin the Cro- mainly 3-(2-5-)locular; ovules 1 per locule, anat- tonoideae,Bentham's original suggestion of a rela- ropous or orthotropous; styles bifid or multifid. tionshipto theGalearieae seems perceptive and best Fruit drupaceous; exocarp fleshy or woody; en- supportedby the availabledata. docarp bony and corrugated or channeled; seeds In my last reviewof the placementof Dicoelia ecarunculate, endosperm copious. (Webster,1987), the tribeDicoelieae was referred to thefamily Pandaceae as one ofits two tribes, the The tribe Galearieae as treated here corresponds otherbeing the Galearieae. If the Euphorbiaceae were exactly, except forrank, withthe familyPandaceae

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as circumscribed by Forman (1966). The group 3: 287. 1880; Hook. f., Fl. Brit. Ind. 5: 377. has had a rather complicated taxonomic history, 1887; Pax & Hoffm., Pflanzenr. 147. III which Forman has discussed thoroughly as far as (Heft 47): 97. 1911; Gagnepain, Fl. Indochine Panda is concerned. Bentham (1878, 1880) per- 5: 456. 1926; Forman, Kew Bull. 26: 155. ceptively noted the resemblance of Dicoelia with 1971; Whitmore, Tree Fl. Malaya 2: 97. Galearia, even though he retained Dicoelia in the 1973. TYPE: Galearia sessilis Zoll. & Mor. Phyllantheae (no doubt because of its biovulate [= Galearia filiformis (B1.) Boerl.; lectotype carpels). His tribeGalearieae, proposed rather apol- selected here because the other species of Zol- ogetically as a transitionalgroup between Phyllan- linger & Moritzi, G. pedicellata, could be theae and Crotoneae, included Galearia and Mi- confused with Bennettia pedicellata R. Br.]. crodesmis juxtaposed. CremostachysTulasne, Ann. Sci. Nat. III. 15: 259. 1851. The Pandaceae have generally been accepted TYPE: Cremostachysfiliformis(Bi.) Tul. [= Galearia in the 20th century as a distinct family, usually filiformis(Bi.) Boerl.; lectotype]. with a position next to the Euphorbiaceae (e.g., BennettiaR. Brown,P1. Jav. Rar. 249. 1852 (non S. F. Cronquist, 1981; Radcliffe-Smith, 1987). In my Gray, 1821). TYPE: Bennettiajavanica R. Br. [ Galearia filiformis(Bi.) Boerl.]. previous review of the group (Webster, 1987), Pandaceae was expanded to include the tribe Di- As treated by Forman (1971), Galearia includes coelieae (previously situated in subfamilyPhyllan- six species of tropical southeast Asia (Burma to thoideae). Whatever their eventual disposition, it Solomon Islands). appears that Dicoelieae and Galearieae (as here defined) form a coherent and apparently mono- 102. Panda Pierre, Bull. Mens. Soc. Linn. Paris phyletic group. 1255. 1896; Aubreville, For. Fl. CUte Iv. 1: The key to the three genera follows that of 300. 1959; Keay, Fl. W. Trop. Afr., ed. 2, Forman. 1: 634. 1958; Forman, Kew Bull. 20: 309. 1966. TYPE: Panda oleosa Pierre. KEY TO THE GENERA OF TRIBE GALEARIEAE PorphyranthusEngler, Bot. Jahrb.Syst. 26: 367. 1899. la. Inflorescencesaxillary, flowers solitary or in TYPE: Porphyranthuszenkeri Engler [= Panda glomerules;staminate petals imbricate;leaves oleosa Pierre]. usuallypellucid-punctate ...... 100. Microdesmis lb. Inflorescencesterminal or cauliflorous,thyr- A monotypic genus of west tropical Africa. Al- soid;staminate petals valvate or nearlyso; leaves though Panda diverges from nearly all other Eu- not pellucid-punctate. phorbiaceae in its distinctive characters (e.g., or- 2a. Endocarpthin-walled; ovules usually anat- thotropous ovules), Forman appears correct in ropous;petals valvate; inflorescences mostly terminal...... 101. Galearia seeing it as the terminus of a sequence of mor- 2b. Endocarp thick-walled;ovules orthotro- phological changes within the Galearieae. pous; petals ? imbricate;inflorescences cauliflorous...... 102. Panda Tribe 21. AMPEREAE Muell. Arg., Bot. Zeit. 100. Microdesmis Planchon, Hook. Ic. P1. 8: 22: 324. 1864; DC. Prodr. 15(2): 211. 1866. t. 758. 1848; Muell. Arg., DC. Prodr. 15(2): TYPE: Amperea A. Juss. 1041. 1866; Bentham, Gen. P1. 3: 287. 1880; Ampereaesubtribe Monotaxideae Muell. Arg., Linnaea Pax & Hoffm., Plfanzenr. 147. III (Heft 47): 34: 63. 1865. TYPE: Monotaxis Brongn. 105. 1911; Gagnepain, Fl. Indochine 5: 458. Monoecious (rarely dioecious) herbs or sub- 1926; Leonard, Bull. Jard. Bot. Brux. 31: shrubs; indumentum simple; leaves alternate, eri- 159. 1961; Fl. Congo 8(1): 102. 1962; Whit- coid, stipulate; flowersin terminalor axillary glom- more, Tree Fl. Malaya 2: 118. 1973; Airy erules; staminate calyx 3-5-lobed, lobes valvate or Shaw, Kew Bull. Add. Ser. 4: 222. 1975. slightly imbricate; petals present or absent; disk TYPE: Microdesmis puberula Hook. f. ex entire or dissected; stamens 6-10, biseriate, fila- Planch. ments free, anthers with disjunct or pendulous loc- A paleotropical genus of ten species, eight Af- ules; pollen grains 3-colporate, colpi inoperculate, rican and two Asian. distinctlymarginate, sexine perforate-tectate;pistillode present or absent; pistillate sepals 4 or 5, 101. Galearia Zollinger & Moritzi, Syst. Verz. imbricate or valvate; petals small or absent; disk 19. 1846 (nom. cons. prop.); Muell. Arg., DC. annular, dissected, or absent; ovary 3-locular; styles Prodr. 15(2): 1036. 1866; Bentham, Gen. P1. bifid to multifid;fruit capsular; seeds smooth, ca-

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runculate; endosperm copious; embryo cylindric, to genera in tribe Epiprineae; this suggestion merits cotyledons scarcely broader than radicle. furtherinquiry.

of Pax (1890) and Grii- In the circumscription Tribe 22. AGROSTISTACHYDEAE (Muell. ning (1913), which is followed here, tribe Amper- Arg.) Webster, Taxon 24: 596. 1975. Aca- eae includes two genera confined to Australia. Al- lypheae subtribe Agrostistachydeae Muell. though the pollen grains are similar in exine Arg., Linnaea 34: 143. 1865. TYPE: Agros- sculpturing and marginate colpi, the genera are tistachys Dalz. dissimilar in many ways, and it is not certain that they should be associated in the same tribe. Dioecious (rarely monoecious) trees or shrubs; indumentum simple or absent; leaves alternate, KEY TO THE GENERA OF TRIBE AMPEREAE simple, entire or denticulate, lamina glandular or 1a. Staminatepetals present; sepals imbricate;an- eglandular; stipules persistent or deciduous; inflo- ther locules disjunct;pollen grains 3-angled, rescences axillary, racemose or spicate, sometimes sexinereticulate; style-branches fimbriate; cymes with scarious imbricate bracts; staminate calyx terminal. 103. Monotaxis closed in bud, splittinginto 2-5 valvate lobes; petals lb. Staminatepetals absent; sepals valvate;anther 3-8; disk receptacular or extrastaminal and dis- locules pendulousfrom glandular connective; pollen grains 3-lobed,sexine perforate-foveo- sected; stamens 10-50, filamentsfree (at least in late; style-branchesentire; flowers axillary. part), anthers introrse,with ? enlarged connective, 104. Amperea anther-sacs often pendulous; pollen grains subglobose, 3-colporate, colpi inoperculate, not distinctly pistillode 103. Monotaxis Brongniart,Voy. Coquille Bot. marginate, sexine coarsely reticulate; or 223. 1829; Muell. Arg., DC. Prodr. 15(2): present or absent; pistillate sepals 4 or 5, open 212. 1866; Bentham, Fl. Austral. 6: 78. 1873; valvate; petals 5, small and deciduous, or absent; Gen. P1. 3: 264. 1880; Grining, Pflanzenr. disk lobed, pitted, or dissected; ovary 3-locular, seeds 147. (Heft 58): 75. 1913; Pax & Hoffm., often tomentose; styles bifid; fruitcapsular; testa some- Natiirl. Pflanzenfam. ed. 2, 19c: 227. 1931; smooth or granulose, ecarunculate, Airy Shaw, Muelleria 4: 239. 1980. TYPE: times fleshy. Monotaxis linifolia Brongn. [lectotype]. This tribe is here considered to include four Asiatic. Hippocrepandra Muell. Arg., Linnaea 34: 61. 1865. genera, two African and two TYPE: Hippocrepandra gracilis Muell. Arg. [= Monotaxis lurida (Muell. Arg.) Benth.; lectotype, KEY TO THE GENERAOF TRIBE AGROSTISTACHYDEAE designatedby Wheeler,1975]. la. Petals present;flowers in spikesor racemes. An Australian genus of about ten species. Brong- 2a. Dioecious;staminate petals imbricate in bud; petals presentin pistillateflower. niart, in his original description, suggested an af- 3a. Staminatedisk dissected; stamens 8- finityto Chiropetalum and Ditaxis in tribe Chro- 10; stipulesfree; leaves eglandular zophoreae. Punt (1962) related pollen of Monotaxis 105. Agrostistachys to his "Sumbavia type," which includes genera of 3b. Staminatedisk receptacular; stamens connate,leaving an tribe Chrozophoreae. Perhaps further study will 20-55; stipules annularscar; leaves glandularat base demonstrate that Monotaxis is indeed an aberrant 106. Pseudagrostistachys genus of Chrozophoreae. 2b. Monoecious;staminate petals not imbricate in bud; pistillateflowers apetalous; leaves glandularat base 107. Cyttaranthus Tent. 35. 104. Amperea A. Jussieu, Euphorb. lb. Petals absent;flowers in panicles;monoecious, 1824; Muell. Arg., DC. Prodr. 15(2): 213. leaves glandularat base 108. Chondrostylis 1866; Bentham, Fl. Austral. 6: 81. 1873; 265. 1880; Grining, Pflanzenr. Gen. P1. 3: 105. Agrostistachys Dalzell,Hook. J.Bot. Kew 86. Pax & Hoffm., 147 (Heft 58): 1913; Gard. Misc. 2: 41. 1850; Muell. Arg., DC. Natiirl. Pflanzenfam. ed. 2, 19c: 228. 1931; Prodr. 15(2): 725. 1866; Bentham, Gen. P1. Henderson, Austral. Syst. Bot. 10 (in press). 3: 302. 1880; Hook. f., Fl. Brit. Ind. 5: 405. 1993. TYPE: Amperea ericoides A. Juss. 1887; Pax & Hoffm.,Pflanzenr. 147. VI (Heft According to Henderson (1993), Amperea is a 57): 98. 1912; Gagnep., Fl. Indochine 5: 463. genus of eight Australian species, all except two 1926; AiryShaw, Kew Bull. 14: 472. 1960; confined to Western Australia. On the basis of 26: 210. 1972; Whitmore,Tree Fl. Malaya pollen characters, Punt (1962) related Amperea 2: 52. 1973; Airy Shaw, Kew Bull. Add. Ser.

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4: 26. 1975; Kew Bull. 36: 248. 1981. TYPE: or dentate,pinnately or palmatelyveined, some- Agrostistachys indica Dalz. times withlaminar glands; stipulespersistent or deciduous,sometimes obsolete; inflorescences ax- HeterocalyxGagnepain, Notul. Syst. (Paris) 14: 33. 1950. illaryor terminal,mostly racemose or spicate.Sta- TYPE: Heterocalyx laoticus Gagnep. [= Agrostis- tachys indica Dalz.]. minateflower: sepals 3-5, valvate; petals 4 or 5 (rarely 10 or absent); disk dissected or absent; An Asiaticgenus of ten species, some difficult stamens 5-many, filamentsfree or connate; an- to distinguish,distributed from India and Sri Lanka thers introrse,muticous, locules not pendulous; to New Guinea. pollen grains subgloboseto oblate, mostly3-6- colporate,colpi sometimes operculate, not distinct- 106. Pseudagrostistachys Pax & Hoffmann, ly marginate,sexine reticulate, often distinctly het- Pflanzenr. 147. VI (Heft 57): 96. 1912; Le- erobrochate;pistillode present or absent.Pistillate brun, Bull. Soc. Roy. Belge 67: 97. 1934; flower:sepals 5 or 6, imbricateor valvate; petals Leonard, Fl. Congo 8(1): 183. 1962; Rad- 5 (sometimesreduced or absent); disk annularor cliffe-Smith,Fl. E. Trop. Afr. Euphorb. 1: dissected(rarely absent); ovary (2-)3-locular, smooth 166. 1987. TYPE: Pseudagrostistachys af or muricate;styles unlobed to bifidor twice-bifid, ricana (Muell. Arg.) Pax & Hoffm. sometimeslacerate. Fruit capsular; seeds dry or An Africangenus of two species distributed from fleshy,endotesta smooth or roughenedto pitted, Sao Tome and Nigeria to Zaire and Uganda. ecarunculate. As delimitedby Webster(1975) and here,tribe 107. Cyttaranthus Leonard, Bull. Jard. Bot. Chrozophoreaecorresponds almost exactly to the Brux. 25: 286. 1955; Fl. Congo 8(1): 180. subtribeRegulares as treatedby Pax & Hoffmann 1962. TYPE: Cyttaranthus congolensis Leon- (1931); their other subtribeIrregulares includes ard. genera belongingto tribe Agrostistachydeaeand A monotypicgenus confinedto west Africa (Con- subfamilyCrotonoideae. Pollen charactersin gen- go and Cabinda); extremely close to Chondrostylis era such as Doryxylonsuggest a possiblerelation- and perhaps not separable. shipbetween Chrozophoreae and tribeEpiprineae.

KEY TO THE SUBTRIBESOF TRIBE CHROZOPHOREAE 108. Chondrostylis Boerlage, Ic. Bogor. 1: t. 23. 1897; Koorders, Ann. Jard. Bot. Buitenz. 1a. Pollengrains not distinctly heterobrochate; indumentumsimple or malpighiaceous; leaves not 19: 45. 1904; Pax & Hoffm.,Pflanzenr. 147. withbasal laminar glands. VII (Heft 63): 15. 1914; Airy Shaw, Kew 2a. Inflorescencesterminal; indumentum en- Bull. 14: 358. 1960; Whitmore, Tree Fl. Ma- tirelysimple; stamens 10-15, free;pistil- laya 2: 77. 1973; Airy Shaw, Kew Bull. Add. latedisk annular; ovary muricate Ser. 4: 69. 1975; Kew Bull. 36: 276. 1981. .------. . . - 23a. Speranskiinae 2b. Inflorescencesaxillary; indumentum mal- TYPE: Chondrostylis bancana Boerl. pighiaceousor stellate(at leastin part); ifmore than connate;pistillate Fl. Penin.3: 283. 1924. stamens, 6, KunstlerodendronRidley, Malay diskdissected or obsolete...... 23b. Ditaxinae TYPE: nom. Kunstlerodendronsublanceolata Ridl., lb. Pollen indu- ex Hook. f.) grainsdistinctly heterobrochate; illeg.[= Chondrostyliskunstleri (King mentumstellate; leaves withbasal laminar AiryShaw]. glands. A monotypicgenus of southeastAsia (Thailand 3a. Shrubsor trees; pollen grains 3-colporate; stamensfree; seeds fleshy .... 23c. Doryxylinae to Sumatraand Borneo). 3b. Herbs;pollen grains stephanocolporate; stamensconnate; seeds not fleshy ...... Tribe 23. CHROZOPHOREAE (Muell. Arg.) ...... 23d. Chrozophorinae Pax & Hoffmann,Pflanzenr. 147. XIV (Heft 68): 5. 1919. Chrozophoreaesubtribe Regu- Subtribe 23a. SPERANSKIINAE Webster, lares (Pax & Hoffm.) Pax & Hoffm., Natiirl. Taxon 24: 596. 1975. TYPE: Speranskia Bail- Pflanzenfam. ed. 2, 19c: 89. 1931. TYPE: lon. Chrozophora Necker ex A. Juss. Monoeciousherbs; indumentumsimple; leaves Trees, shrubs, or herbs, monoecious or dioe- alternate,coarsely dentate or lobulate,with glan- cious; indumentum usually stellate, lepidote, or dular teeth,stipulate; inflorescences terminal, spi- malpighiaceous (at least in part; rarely entirely cate, bisexual,pistillate flowers proximal; staminate simple); leaves alternate, unlobed or lobed, entire sepalsand petals5; diskdissected; stamens 1 0(- 15),

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filaments free; pollen grains 3-colporate, evenly which resembles the Chinese genus in its muricate reticulate; pistillode absent; pistillate sepals 5; pet- ovary and laciniate style-branches, but differsin als 5, small or absent; disk annular; ovary 3-locular, its axillary racemes and stephanocolporate pollen. verrucose; styles free,bifid, branches lacerate; fruit Generic delimitationwithin the subtribe is still con- capsular; seeds dry, ecarunculate, testa foveolate- troversial, but pollen characters such as pointed asperate. out by Punt (1962) support the narrow circumscriptionof Pax & Hoffmannrather than the broad This monogeneric subtribe is morphologically delimitationof Mueller (1866) and Ingram (1979). similar to subtribe Ditaxinae, and indeed Mueller (1866) included Speranskia as a section of Ar- KEY TO THE GENERAOF SUBTRIBEDITAXINAE gythamnia. However, the Speranskiinae diverge 1a. Pistillodepresent; styles dissected; indumentum in number of characters, from the Ditaxinae a simpleor glandular(not malpighiaceous); leaves particularly the androecium of free stamens and finelyserrate with straight parallel lateral veins; the terminal inflorescence. pollengrains 6-colporate 110. Caperonia lb. Pistillodeabsent; stylesbifid to dissected;in- 109. Speranskia Baillon, Etude Gen. Euphorb. dumentum(at least in part) malpighiaceous; leavesotherwise; pollen grains 3- or 4-colporate. 388. 1858; Bentham,Gen. P1. 3: 305. 1880; 2a. Dioecious;tepals scarious; shrubs or trees, Pax & Hoffm., Pflanzenr. 147. VI (Heft 57): branchesoften spiny ...... 111. Philyra 14. 1912; Hurusawa, J. Fac. Sci. Univ. Tokyo 2b. Monoecious; tepals not scarious; under- III. Bot. 6: 310. 1954. TYPE: Speranskia shrubsor herbs,not spiny. 3a. Petalsentire; styles twice bifid to mul- tuberculata (Bunge) Baillon. tifid;stellate hairs absent. A genus restricted to China; there are three 4a. Stamens 8-10, anthers in 2 whorls; pollen grains 3- or Pax Hoffmann (1912) or species according to & 4-colporate,bilaterally symmet- two according to Hurusawa (1954). rical. .... 112. Ditaxis 4b. Stamens 4 or 5, anthersin 1 whorl;pollen grains stephanocol- Subtribe 23b. DITAXINAE Grisebach, Fl. Brit. porate 113. Argythamnia W. Ind. 43. 1859; Abh. Gesell. Wiss. Goet- 3b. Petals toothedor lacerate;styles once bifid;stellate hairs sometimes present; tingen 9: 15. 1861. TYPE: Ditaxis Vahl ex A. pollengrains 3-colporate Juss. 114. Chiropetaluin Acalypheae subtribeCaperonieae Muell. Arg., Linnaea 34: 143. 1865. TYPE: Caperonia St. Hil. 110. Caperonia St. Hilaire, P1. Remarq. Bresil 244. 1826; Muell. Arg., DC. Prodr. 15(2): Monoecious (rarely dioecious) trees, shrubs, or 751. 1866; Fl. Bras. 11(2): 315. 1874; Ben- herbs; indumentumpredominantly malpighiaceous, tham, Gen. P1. 3: 304. 1880; Pax & Hoffm., sometimes also stellate, simple, or glandular; leaves Pflanzenr. 147. VI (Heft 57): 27. 1912; Le- alternate, entire or serrate, withoutlaminar glands; onard, Bull. Jard. Bot. Brux. 26: 313. 1956; inflorescences axillary, racemose or glomerulate, Fl. Congo 8(1): 166. 1962; Webster, J. Ar- usually bisexual; staminate sepals mostly 4 or 5; nold Arbor. 48: 363. 1967; Dyer, Gen. S. petals 4 or 5; disk dissected or absent; stamens 4- Afr. Fl. P1. 1: 314. 1975; Radcliffe-Smith, 15, filamentsconnate (rarely almost free); anthers Fl. E. Trop. Afr. Euphorb. 1: 163. 1987. introrse; pollen grains subglobose to oblate, TYPE: Caperonia castaneifolia (L.) St. Hil. 3-colporate to stephanocolporate, sexine reticulate, [lectotype, designated by Britton & Wilson, not distinctlyheterobrochate; pistillode present or Bot. Porto Rico 6: 486. 1925]. absent; pistillate sepals 5 or more, imbricate to valvate; petals 5(6), sometimes obsolete; disk dis- A genus of ca. 40 species, mostly American, sected or absent; ovary sometimes muricate; styles less than 10 species in Africa. Caperonia is quite bifidto laciniate; fruitcapsular; seeds dry, smooth sharply distinct from the other genera in subtribe or ornamented (foveolate, asperate, or reticulate), Ditaxinae, and Mueller referred it to a separate ecarunculate. subtribe Caperonieae. On the basis of present re- ports (Hans, 1973), Caperonia appears to have a In the circumscriptionof Webster (1975), which differentchromosome number (x = 11) from the is continued here, subtribe Ditaxinae includes five other genera (x = 13). genera that are confined to the New World except for a few species of Caperonia. The subtribe ap- 1 1 1. Philyra Klotzsch, Arch. Naturgesch. 7(1): proaches subtribe Speranskiinae via Caperonia, 199. 1841; Baillon, Etude Gen. Euphorb. 297.

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1858; Pax & Hoffmann, Pflanzenr. 147. VI 147. VI (Heft 57): 86. 1912; Ingram, Gentes (Heft 57): 49. 1912; Naturl. Pflanzenfam. ed. Herb. 11(7): 437. 1979; L. Smith et al., Fl. 2, 19c: 92. 1931; O'Donell & Lourteig, Lilloa Ilustr. Catar. EUFO 154. 1988. TYPE: Chi- 8: 60. 1942. TYPE: Philyra brasiliensis KI. ropetalum tricuspidatum (Lam.) A. Juss.

A monotypic genus of Paraguay and southern DesfontaenaVellozo, Fl. Flum. 95. 1825. TYPE: DesJbn- Brazil. taena tricocca Vell. [= Chiropetalumtricoccum (Vell.) Chod. & HassI.]. Chlorocaulon Ki. ex Endl., Gen. P1. Suppl. 4(3): 89. 1 12. Ditaxis Vahl ex A. Jussieu, Euphorb. Tent. 1850. TYPE: Chiropetalumschiedeanum (Muell. 27. 1824; Pax, Natiirl. Pflanzenfam. ed. 1, Arg.) Pax & Hoffm. 3(5): 44. 1890; Pax & Hoffm., Pflanzenr. AonikenaSpegazzini, Anal. Mus. Nac. Hist. Nat. Buenos 147. VI (Heft 57): 51. 1912; Natiirl. Pflan- AiresII. 7: 162. 1902. TYPE: Aonikenapatagonica Speg. [= Chiropetalumpatagonicum (Speg.) O'Don. zenfam. ed. 2, 19c: 93. 1931. TYPE: Ditaxis & Lourt.]. fasciculata Vahl ex A. Juss. A genus of ca. 20 species, almost entirely in Nuttall,Trans. Amer. Phil. Soc. II. 5: 174. 1837. Aphora temperate South America except for a single Mex- TYPE: Aphora mercurialinaNutt. [= Ditaxis mer- curialina (Nutt.)Coulter]. ican species. Although reduced to a section of Ar- SerophytonBentham, Bot. Voy. Sulphur 52. 1844. TYPE: gythamnia by Ingram, Chiropetalum is distinctive Serophytonlanceolatum Benth. [= Ditaxis lan- in its lobed petals and differentpollen type (Punt, ceolata (Benth.)Pax & Hoffm.]. 1962). Stenonia Didrichsen,Vidensk. Medd. Dansk Naturhist. Foren.Kj0benhavn 1857f: 146. 1857 (nonEndlich- er, 1947). TYPE: Stenonia montevidensisDidr. [ Subtribe 23c. DORYXYLINAE Webster,Tax- Ditaxis montevidensis(Didr.) Pax]. on 24: 596. 1975. TYPE: Doryxylon Zoll. Paxiuscula Herter,Rev. Sudamer. Bot. 6: 92. 1941. TYPE: Paxiuscula acaulis (Herterex Arech.)Herter Monoecious shrubs or trees; indumentum stel- [= Ditaxis acaulis Herter]. late; leaves alternate, entire or dentate, with laminar glands; stipules small; inflorescences terminal An Americangenus of 40-50 speciesdistributed or axillary, usually bisexual; staminate sepals 3- in drier areas from the United States south to 5; petals 5-10 or absent; disk reduced or absent; Argentina. Ditaxis is very close to Argythamnia, stamens many (ca. 30-250), free or partially con- and it remains to be demonstratedwhether it should nate; pollen grains 3-colporate, angulaperturate, be combined with that genus, as many workers, coarsely reticulate, slightly heterobrochate along including Mueller (1866), Bentham (1880), and colpi; pistillode present or absent; pistillate sepals Ingram (1979), have proposed. However, evidence 5 or 6, imbricate or valvate; petals rudimentary from pollen morphology (Punt, 1962; Webster, or absent; disk annular (or obsolete); ovary 2- or ined.) tends to support its status as a distinctgenus. 3-locular, stellate-pubescent; styles unlobed or bifid; seeds with fleshyexotesta, foveolate endotesta. 113. Argythamnia P. Brown, Civ. Nat. Hist. Jamaica 338. 1756; Swartz,Fl. Ind. Occid. This subtribe of four Asian genera stands out 1: 335. 1797; Muell. Arg., DC. Prodr. 15(2): from other Chrozophoreae by the combination of 732. 1866; Bentham,Gen. P1. 3: 303. 1880; woody habit and stellate pubescence. Pax & Hoffm.,Pflanzenr. 147. VI (Heft57): 78. 1912; Johnston& Warnock,Southw. Nat. KEY TO THEGENERA OF SUBTRIBEDORYXYLINAE 7: 154. 1962; Ingram, Brittonia16: 271. La. Petals present;ovary 3-locular; stamens ca. 1964; GentesHerb. 10(1): 1-46. 1967; 11(7): 30-100. 426. 1979; Correll & Correll,Fl. Bahama 2a. Inflorescencesaxillary; styles unlobed, ba- stamensfree 115. Doryxylon Arch.777. 1982; Sneep & De Roon,Fl. Neth. sally connate; 2b. Inflorescencesterminal; styles bifid, + free. Ant. 3: 250. 1984. TYPE: Argythamnia can- 3a. Stamensfree; staminate disk absent; dicans Swartz. pistillatesepals imbricate ...... 116. Sumbaviopsis In the revisional study by Ingram (1967), 18 3b. Stamensconnate; staminate disk pres- species of Argythamnia (sensu stricto) are recent; pistillatesepals valvate . ognized. It is a Caribbean genus with over half the 1 17. Thyrsanthera lb. Petals absent;ovary usually 2-locular; stamens species in the West Indies, but a number also 200 or more; inflorescencesterminal; styles occurring in Mexico and Central America. entire... 118. Melanolepis

114. Chiropetalum A. Jussieu, Ann. Sci. Nat. 1 15. Doryxylon Zollinger, Tijdschr. Ned.-Indie I. 25: 21. 1832; Pax & Hoffm., Pflanzenr. 14: 172. 1857; Linnaea 29: 469. 1859; Ba-

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lakrishnan,Bull. Bot. Surv. India 9: 56. 1967. Subtribe 23d. CHROZOPHORINAE Muell. TYPE: Doryxylon spinosum Zoll. Arg., Linnaea 34: 143. 1865; DC. Prodr. 15(2): 726. 1866; Pax & Hoffm.,Pflanzenr. SumbaviaBaillon, Etude Gen. Euphorb. 390. 1858; Muell. Arg.,DC. Prodr.15(2): 727. 1866; Pax & Hoffm., 147. VI (Heft57): 1. 1912. TYPE: Chrozoph- Pflanzenr.147. VI (Heft57): 11. 1912. TYPE: Sum- ora tinctoria(L.) A. Juss. bavia rottleroidesBaillon [= Doryxylonspinosum Zoll.]. Monoecious herbs or subshrubs;indumentum stellate;leaves alternate,entire or lobed, biglan- This Asian genus, according to Balakrishnan, dularat base, stipulate;inflorescences axillary, ra- includes two species, but he treated Sumbaviopsis cemose, bisexual; staminatesepals and petals 5; as a synonym of Doryxylon. He may be correct diskinconspicuous, adnate to staminalcolumn; sta- in this, but until the relationships can be further mens 4-15, filamentsconnate into a column;an- investigated, also taking Thyrsanthera into ac- thers introrse;pollen grains oblate, stephanocol- count, it seems best to provisionallykeep the genera porate, colpi short and broad, sexine distinctly distinct. Earlier workers, from Mueller (1866) to heterobrochate;pistillode absent; pistillatesepals Pax & Hoffmann (1912, 1931), used Sumbavia 5; petals5, smallor absent;disk lobed or dissected; as the name for this genus because they thought ovary3-locular, stellate or lepidote,sometimes mu- Doryxylon was firstpublished in Linnaea in 1859, ricate; stylesbifid; fruit capsular; seeds caruncu- and hence did not have priority. late, somewhatfleshy, endotesta smooth or tuberculate. 1 16. Sumbaviopsis J. J. Smith,Add. Fl. Arb. Jav. 12 (Med. Dept. Landbouw 10): 356. In the presentcircumscription, subtribe Chro- 1910; Pax & Hoffm.,Pflanzenr. 147. VI (Heft zophorinaeis monogeneric.It is veryclose to sub- 57): 13. 1912; Gagnep.,Fl. Indochine5: 418. tribeDoryxylinae, as notedby AiryShaw (1972, 1926; AiryShaw, Kew Bull. 14: 357. 1960; 1975), and furtherstudy may show that Thyr- Whitmore,Tree Fl. Malaya 2: 132. 1973; santhera bridgesthe apparent gap betweenthe AiryShaw, Kew Bull.Add. Ser. 4: 197. 1975. two taxa. TYPE: Sumbaviopsis albicans (B1.) J. J. Sm. 119. Chrozophora Necker ex A. Jussieu,Eu- A monotypic tropical Asian genus (Assam to phorb.Tent. 27. 1824 (as Crozophora),nom. Borneo and Palawan). & orth.cons.; Muell. Arg.,DC. Prodr. 15(2): 746. 1866; Bentham,Gen. P1. 3: 305. 1880; 117. Thyrsanthera Pierre ex Gagnepain, Bull. Pax & Hoffm.,Pflanzenr. 147. VI (Heft57): Soc. Bot. France 71: 878. 1925; Gagnepain, 17. 1912; Prain,Kew Bull. Misc. Inf. 1918: Fl. Indochine5: 299. 1925; Pax & Hoffm., 49. 1918; Pojarkova, Fl. URSS 14: 288. Natiirl.Pflanzenfam. ed. 2, 19c: 90. 1931; 1949; Rech. f. & Schiman-Czeika,Fl. Iranica Backer & Backh.,Fl. Jav. 1: 477. 1963; Airy 6: 5. 1964; Berhaut,Fl. Ill. S6n6gal 3: 397. Shaw, Kew Bull. 19: 308. 1965; 26: 343. 1975; Radcliffe-Smith,Fl. E. Trop. Afr.Eu- 1972. TYPE: Thyrsanthera suborbicularis phorb. 1: 160. 1987. TYPE: Chrozophora Pierreex Gagnep. tinctoria(L.) A. Juss. A monotypic genus known from Thailand and An Old World genus of ca. ten species distrib- Cambodia. Airy Shaw (1972, in clavi) suggested uted fromsouthern Europe to east Africa,central that it may be more closely related to Chrozophora and southernAsia. than to Sumbaviopsis. Tribe 24. CARYODENDREAE Webster,Tax- 1 18. Melanolepis Reichb.f. & Zoll., Verh.Na- on 24: 596. 1975. TYPE: CaryodendronKar- tuurk.Ver. Ned. Ind. 1: 22. 1856; Linnaea sten. 28: 324. 1856; Pax & Hoffm.,Pflanzenr. 147. VII (Heft 63): 142. 1914; Gagnepain, Dioecious trees or shrubs;indumentum simple; Fl. Indochine 5: 347. 1925; Kanehira, Fl. leaves alternate,penninerved or triplinerved,with Micronesica180. 1933; AiryShaw, Kew Bull. basal glands;stipules persistent, reduced, or absent; Add. Ser. 8: 174. 1980. TYPE: Melanolepis inflorescencesterminal or axillary,spicate. Flowers multiglandulosa(Reinw.) Reichb. f. & Zoll. apetalous; staminatesepals 3-5, valvate; disk intrastaminal,pubescent; stamens 4-15, filaments An Asian genus of two species extendingfrom free;anthers introrse, connective enlarged or glan- Thailandand Cambodiato Taiwan, Marianas,and dular; pollen grainsoblate, 3-colporate,colpi not Bismarck Archipelago. marginate,sexine coarsely reticulate; pistillode

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presentor absent. Pistillatesepals 4-6, imbricate of Alchorneopsis; the two taxa could as easily be or valvate, deciduous; disk annular or dissected, regarded as subgenera of a single genus. pubescent;ovary 3-(rarely2- or 4-)locular;styles free,unlobed. Fruit capsular; seeds ecarunculate, 122. Alchorneopsis Muell. Arg., Linnaea 34: testa dryor fleshy. 156. 1865; DC. Prodr. 15(2): 764. 1866; Fl. Bras. 11(2): 327. 1874; Bentham, Gen. P1. This tribeof threegenera is representedby two 3: 315. 1880; Pax & Hoffm.,Pflanzenr. 147. generain the New Worldand one genusin Africa. VII (Heft 63): 267. 1914; Webster, Ann. The pollengrains are verysimilar to thosein tribe Missouri Bot. Gard. 54: 284. 1968. TYPE: Agrostistachydeae,which may be the mostclosely Alchorneopsisfioribunda (Benth.) Muell. Arg. related taxon; however,they clearly differfrom Caryodendreaein havingpetaliferous flowers, more A neotropical genus of three described species stamens,a glabrousfloral disk, and bifidstyles. that are probably conspecific, recorded from the West Indies (Puerto Rico), Panama, Brazil, and KEY TO THE GENERA OF TRIBE CARYODENDREAE the Guianas. la. Pistillodeabsent; inflorescence terminal. 2a. Staminatedisk massive, pulviniform; cap- Tribe 25. BERNARDIEAE Webster, Taxon sulethick-walled, seed coat dry; leaves pen- 24: 596. 1975. Acalypheae subtribe Mer- ninerved,stipules deciduous ...... curialinae series Bernardiiformes Pax & .------. . ... - 120. Caryodendron 2b. Staminatedisk dissected;capsule thin- Hoffm., Pflanzenr. 147. VII (Heft 63): 13. walled,seed coat fleshy;leaves tripli- 1914. TYPE: Bernardia Houstoun ex Miller. nerved,exstipulate . 121. Discoglypremna lb. Pistillodepresent; inflorescences axillary; sta- Monoecious or dioecious shrubs or herbs; in- minatedisk massive,annular; capsule thin- dumentum simple or stellate; leaves alternate, sim- walled;seed coat fleshy; leaves triplinerved, ex- ple, penninerved, often with laminar glands, stip- stipulate .. 122. Alchorneopsisulate; inflorescences terminal, pseudoterminal, or axillary, racemose or spicate; flowers apetalous. 120. Caryodendron Karsten,Fl. Colombiae1: Staminate calyx splittinginto 3 or 4 valvate seg- 91, t. 45. 1860; Muell. Arg.,Fl. Bras. 11(2): ments; disk intrastaminal, pulviniform,dissected, 706. 1874; Bentham,Gen. P1. 3: 314. 1880; or obsolete; stamens 3-60, filamentsfree; anthers Pax & Hoffm.,Pflanzenr. 147. VII (Heft63): muticous or apiculate; pollen grains 3-lobed, 263. 1914; Ducke, Trop. Woods 76: 18. 3-colporate, colpi inoperculate, conspicuously mar- 1943; Webster,Ann. Missouri Bot. Gard. 54: ginate, sexine perforate-tectate;pistillode rudimen- 287. 1968; Huft,Ann. MissouriBot. Gard. tary or absent. Pistillate sepals 4-6, imbricate; disk 76: 1077. 1989. TYPE: Caryodendronori- annular or dissected; ovary 3-locular; styles bifid, nocense Karst. branches sometimes lacerate. Fruit capsular; seeds ecarunculate, smooth, roundish, sometimes carinate. A neotropicalgenus of three species recorded fromPanama to Colombiaand Brazil.Huft (1989) This tribe includes only part of the genera as- has recentlyshown that Centrodiscus Muell. Arg., signed to the series Bernardiiformes by Pax & citedby Pax & Hoffmannand othersas a synonym Hoffmann(1914). The type genus, Bernardia, has of Caryodendron,was notvalidly published; hence oftenbeen associated withAdelia, and indeed Bail- it is not listedas a synonym. lon (1858) treated Bernardia as a synonym of Adelia. However, the pollen of Adelia differs 12 1. Discoglypremna Prain,Kew Bull. 1911: markedly in having operculate colpi and a finely 317. 1911; Fl. Trop. Afr.6(1): 931. 1912; punctate-tectate sexine; tribes Bernardieae and Hook. Ic. P1. 30: t. 2988. 1913; Pax & Adelieae do not appear to be closely related. Hutch- Hoffm.,Pflanzenr. 147. VII (Heft 63): 18. inson (1969) referredBernardia to tribe Macaran- 1914; Keay, Fl. W. Trop. Afr.,ed. 2, 1: 403. geae, but its pollen is very differentfrom Macar- 1958; Radcliffe-Smith,Fl. E. Trop. Afr.Eu- anga. phorb.1: 222. 1987. TYPE: Discoglypremna caloneura (Pax) Prain. KEY TO THE GENERA OF TRIBE BERNARDIEAE A African from monotypic genus, distributed la. Stylesbifid; stamens 4-50, antherconnective Guinea and Sao Tome to Cabinda and Uganda. not greatlyenlarged. Discoglypremna is clearlythe Africanvicariant 2a. Leaf blades not stipellate.

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3a. Stamens 30 or fewer,anthers emar- Fl. E. Trop. Afr. Euphorb. 1: 218. 1987. ginate;seeds carinate 123. Bernardia TYPE: Necepsia afzelii Prain. 3b. Stamensmore than 30, anthersapiculate;seeds notcarinate 124. Necepsia Neopalissya Pax, in Pax & Hoffm.,Pflanzenr. 147. VII 2b. Leaf blades stipellate. (Heft63): 16. 1914. Palissya Baillon,Etude G6n. 4a. Leaves penninerved,short-petioled; Euphorb.502. 1858 (nonPalissya Endlicher,1847). anthersapiculate -- 125. Paranecepsia TYPE: Neopalissya castaneifolia (Baillon) Pax [= 4b. Leaves palmatinerved,long-petioled; Necepsia castaneifolia(Baillon) Bouch. & Leon.]. anthersmuticous - 126. Discocleidion lb. Stylesunlobed, dilated, forming a cap over the As interpreted by Bouchat & Leonard (1986), top of the ovary;stamens 2 or 3, anthercon- Necepsia is an African genus of three species. nectivegreatly dilated - 127. Adenophaedra 125. Paranecepsia Radcliffe-Smith,Kew Bull. 123. Bernardia Houstounex Miller,Gard. Dict. 30: 684. 1976; Fl. E. Trop. Afr. Euphorb. abr. ed. 1754; Houstounex P. Browne,Civ. 1: 220. 1987. TYPE: Paranecepsia alchor- Nat. Hist. Jamaica 361. 1756; Muell. Arg., neifolia Radcl.-Sm. Prodr. 15(2): 915. 1866; Fl. Bras. 11(2): 389. 1874; Bentham,Gen. P1. 3: 308. 1880; A monotypic east African genus (Tanzania and Pax & Hoffm.,Pflanzenr. 147. VII (Heft63): Mozambique). 21. 1914; Fawc. & Rend., Fl. Jam. 4: 290. 1920; Pax & Hoffm.,Natiirl. Pflanzenfam. 126. Discocleidion (Muell. Arg.) Pax & Hoff- ed. 2, 19c: 105. 1931; Standley& Steyerm., mann, Pflanzenr. 147. VII (Heft 63): 45. Fieldiana Bot. 24(6): 52. 1949; Buchheim, 1914. TYPE: Discocleidion ulmifolium(Muell. Willdenowia2: 291. 1960; 3: 217. 1962; Arg.) Pax & Hoffm. Allem,Rev. Brasil.Biol. 39: 529. 1979; Lio- gier, Fl. Espafiola 4: 85. 1986. TYPE: Ber- A genus of two species found in southeast China nardia carpinifoliaGriseb. and the Ryukyus; reduced to Alchornea by Hu- rusawa (1954). Bivonia Spreng.,Neue Entd. Pflanzenk.2: 116. 1820. TYPE: Bivonia axillaris Spreng.[= Bernardia axillaris (Spreng.)Muell. Arg.]. 127. Adenophaedra (Muell. Arg.) Muell. Arg., TraganthusKlotzsch, Arch. Naturgesch. 7(1): 188. 1841. Fl. Bras. 11(2): 385. 1874; Bentham, Gen. TYPE: Traganthus sidoides Kl. [= Bernardia si- P1. 3: 314. 1880; Pax & Hoffmann, Pflan- doides (KI.) Muell. Arg.]. zenr. 147. VII (Heft 63): 261. 1914; Croizat, Phaedra Klotzschex Endlicher,Gen. P1. Suppl.4(3): 88. Trop. Woods 88: 30. 1946; Jablonski,Mem. 1850. TYPE: Bernardia jacquiniana Muell. Arg. [lectotype,selected here]. New York Bot. Gard. 17: 140. 1967; Huft, Polyboea Klotzschex Endlicher,Gen. P1. Suppl. 4(3): Ann. Missouri Bot. Gard. 75: 1099. 1988. 88. 1850. TYPE: Acalypha corensisJacq. [= Ber- TYPE: Adenophaedra grandifolia (Kl.) Muell. nardia corensis(Jacq.) Muell. Arg.]. Arg. [lectotype, designated by Jablonski, Tyria Klotzschex Endlicher,Gen. P1. Suppl. 4(3): 88. 1850. TYPE: Bernardia mexicana (Hook. & Arn.) 1967]. Muell. Arg. [lectotype,selected here]. A neotropical genus of three species extending Alevia Baillon,Etude Gen. Euphorb.508. 1858. TYPE: Alevia leptotaschiaBaillon, sphalm. [= Bernardia from Costa Rica south to Brazil. The relationships interrupta(Schlecht.) Muell. Arg.]. of the genus require furtherstudy, as it is somewhat Passaea Baillon,Etude Gen. Euphorb.507. 1858. TYPE: aberrant in the tribe Bernardieae. Although Muel- Passaea spartioides Baillon [= Bernardia spar- ler (1866) originally treated Adenophaedra as a tioides (Baill.) Muell. Arg.]. section of Bernardia, Bentham (1880) placed it A diverse Americangenus of ca. 50 species, next to Caryodendron, as did Pax & Hoffmann dividedinto seven sectionsby Pax & Hoffmann (1914). There is indeed some resemblance, but the (1914, 1931); some of these, especiallysection pollen studies of Punt (1962) appear to confirm Traganthus,may deserve generic status on further the intuitionof Mueller that the closer relationship study. of Adenophaedra is with Bernardia.

Tribe 26. PYCNOCOMEAE Hutchinson,Amer. 124. Necepsia Prain,Kew Bull.Misc. Inf. 1910: J. Bot. 56: 753. 1969; Webster, Taxon 24: 343. 1910; Hook. Ic. P1. 30: t. 2987. 1913; 596. 1975. TYPE: Pycnocoma Bentham. Pax & Hoffm.,Pflanzenr. 147. VII (Heft63): 16. 1914; Bouchat & Leonard, Bull. Jard. Dioecious trees; indumentumsimple, often glan- Bot. Brux. 56: 179. 1986; Radcliffe-Smith,dular, or absent; leaves alternate, opposite, or ver-

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ticillate, simple, penninerved, eglandular, some- flowersdistinctly pedicellate; filaments in- times resinous; stipules caducous or absent; flexedin bud .... 128. Pycnocoma 2b. Ovaryunappendaged, glabrous; staminate axillary, racemose or paniculate, or inflorescences flowerssubsessile, filaments not inflexedin contracted into capitula; flowers apetalous. Sta- bud.129. Droceloncia minate calyx closed in bud, splittinginto 3 or 4 lb. Pistillateflower not terminatinginflorescence; valvate segments; disk glabrous and receptacular, pistillatedisk annular. 130. Argomuellera or absent; stamens 15-many, free, anthers sometimes withenlarged connective; pollen grains oblate 128. Pycnocoma Bentham, Niger Fl. 508. or subglobose, 3-colporate, colpi short and narrow, 1849; Muell. Arg., DC. Prodr. 15(2): 951. inoperculate, sexine tectate-perforate,gemmate or 1866; Bentham,Gen. P1. 3: 326. 1880; Prain, spinulose; pistillode absent. Pistillate sepals 3-7, Fl. Trop. Afr.6(1): 955. 1913; Pax & Hoffm., imbricate; disk glabrous or absent; ovary (2-)3- Pflanzenr.147. VII (Heft 63): 52. 1914; locular; stylesbifid or unlobed. Fruit capsular, often Leandri, Notul. Syst. (Paris) 9: 161. 1941; thick-walled and subdrupaceous; seeds ecaruncu- Keay, Fl. W. Trop. Afr.ed. 2, 1: 405. 1958; late, testa dry or fleshy. Uonard, Bull. Soc. Roy. Bot. Belge 91: 273. 1959; Racliffe-Smith,Fl. E. Trop. Afr. Eu- As defined by Webster (1975) and here, only phorb.1: 228. 1987. TYPE: Pycnocomamac- the type genus Pycnocoma, of the genera included rophylla Benth. by Hutchinson, is retained in the tribe Pycnoco- meae; the other genera are relegated to tribes A tropicalAfrican genus of ca. 15 species. Epiprineae and Plukenetieae. However, the membership of the taxa here included in the Pycno- 129. Droceloncia Leonard,Bull. Soc. Roy.Bot. comeae is still a matter of doubt. The two subtribes Belge 91: 279. 1959. TYPE: Droceloncia rig- recognized here are divergentboth morphologically idifolia(Baillon) Leonard. and in geographical distribution,so that they may A monotypicgenus of the Comoresand Mad- have to be separated on furtherstudy. However, agascar. there is some resemblance in pollen characters between Pycnocoma and Blumeodendron, so the 130. Argomuellera Pax, Bot. Jahrb.19: 90. two subtribes are kept together provisionally. 1894; Prain,Fl. Trop. Afr.6(1): 925. 1912; Leandri,Notul. Syst. (Paris) 9: 161. 1941; KEY TO THE SUBTRIBES OF TRIBE PYCNOCOMEAE Leonard,Bull. Soc. Roy. Bot. Belge 91: 274. 1a. Monoecious;seed-coat not fleshy; pollen sexine 1959; Radcliffe-Smith,Fl. E. Trop. Afr.Eu- gemmate 26a. Pycnocominae phorb. 1: 225. 1987. Pycnocoma sect. We- fleshy;pollen sexine punc- lb. Dioecious;seed-coat DC. Prodr. 15(2): 952. tate-tectateor coarselyreticulate. triaria Muell. Arg., -...... 26b. Blumeodendrinae 1866. Wetriaria(Muell. Arg.) Pax, Pflanzenr. 147. VII (Heft 63): 49. 1914. TYPE: Argo- muellera macrophylla Pax. Subtribe 26a. PYCNOCOMINAE Webster, subtr. nov. TYPE: Pycnocoma Bentham. An Afro-Malagasiangenus of ca. ten species,as delimitedby Leonard (1959). Monoicae; stamina numerosa;antherarum connecti- vum incrassatum;pollinis grana 3-colpoidorata,sexino gemmato;semina non carnosa. Subtribe 26b. BLUMEODENDRINAE Web- ster, Taxon 24: 596. 1975. TYPE: Blumeo- Monoecious shrubs or small trees; leaves alter- dendron(Muell. Arg.) Kurz. nate or pseudo-verticillate; stamens 50 or more; pistillateflowers sessile; pistillatedisk absent; styles Dioecioustrees; leaves long-petiolate, entire, with elongated, connate below, barely lobed at tip; fruit scalariformveinlets; inflorescences racemose or pa- thin-walled,horned or not; seeds with dry testa. niculate;stamens 15-many; antherconnective enlarged; pollengrains perforate-tectate or coarsely This subtribeincludes three genera, two African reticulate;ovary (2-)3-locular; styles unlobed; fruit and one Malagasian. large, thick-walled;seeds withfleshy testa.

KEY TO THE GENERA OF SUBTRIBE PYCNOCOMINAE This subtribeincludes three Asian genera that differin aspectfrom the Pycnocominae by the long- la. Pistillateflower terminating the inflorescence; pistillatedisk absent. petiolateleaves and more or less paniculateinflo- 2a. Ovary 6-horned, pubescent; staminate rescence.

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KEY TO THE GENERA OF SUBTRIBE BLUMEODENDRINAE ClaroriviniaPax & Hoffm.,Pflanzenr. 147. VII (Heft 63): 17. 1914. TYPE: Claroriviniachrysantha 1a. Antherconnective no more than moderately (K. Schum.)Pax & Hoffm.[= Ptychopyxischrysantha enlarged,not umbraculiform; diskten- pistillate (K. Schum.) AiryShaw]. uous or absent;styles elongated. 2a. Anther-cells2; fruitlacking spines or ridg- A tropical Asian genus of 10-12 species dis- es. 3a. Anther-cellsadnate to connective;pis- tributed from Thailand through Indonesia to the tillatedisk annular; flowerswithout Philippines and New Guinea. glandularhairs 131. Blumeodendron 3b. Anther-cellspendulous; pistillate disk 134. Botryophora Hooker f., Fl. Brit.Ind. 5: absent;flowers with stalked glandular 476. 1888 (nom. cons.); Pax & Hoffm.,Na- hairs 132. Podadenia 2b. Anther-cells4; fruitwith spines or ridges tiirl.Pflanzenfam. ed. 2, 19c: 228. 1931; Airy -...... 133. Ptychopyxis Shaw, Kew Bull. 3: 484. 1949; 14: 374. lb. Antherconnective umbraculiform, with 4 pen- 1960; Hook. Ic. P1. 36: t. 3576. 1962; Kew dentlocules; pistillate disk massive, pulviniform; Bull. Add. Ser. 4: 61. 1975; Kew Bull. 36: stylesstigmatiform - 134. Botryophora 267. 1981. TYPE: Botryophora kingii Hook. f. [= Botryophora geniculata (Miq.) Beum~e 131. Blumeodendron (Muell. Arg.) Kurz, J. ex AiryShaw]. Asiatic Soc. Bengal 42: 245. 1873; J. J. Smith, Meded. Depart. Landbouw 10: 458. 1910; A monotypic genus of tropical Asia (Burma to Pax & Hoffm.,Pflanzenr. 147. VII (Heft 63): Sumatra and Borneo). 47. 1914; Airy Shaw, Kew Bull. 16: 348. 1963; Whitmore, Tree Fl. Malaya 2: 68. Tribe 27. EPIPRINEAE (Muell. Arg.) Huru- 1973; Airy Shaw, Kew Bull. Add. Ser. 4: 57. sawa, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 1975; Kew Bull. Add. Ser. 8: 37. 1980; Kew 6: 309. 1954. Acalypheae subtribe Epipri- Bull. 36: 267. 1981. TYPE: Blumeodendron neae Muell. Arg., Linnaea 34: 144. 1865; tokbrai (Bl.) Kurz [lectotype, designated by DC. Prodr. 15(2): 1024. 1866. TYPE: Epi- Wheeler, 1975]. prinus Griffith.

An Asian genus of six species distributedfrom Monoecious trees or shrubs; indumentum stel- the Andaman Islands and Burma throughIndonesia late; leaves alternate, simple, pinnately veined, to the Bismarck Archipelago. eglandular or with petiolar glands, stipulate; inflorescences terminal or axillary, paniculate, race- 132. Podadenia Thwaites, Enum. PI. Zey 4: mose, or spicate (or staminate capitular); flowers 273. 1861; Muell. Arg., DC. Prodr. 15(2): apetalous. Staminate calyx closed in bud, splitting 791. 1866; Bentham, Gen. PI. 3: 318. 1880; into 2-6 valvate segments; disk absent; stamens Hook. f., Fl. Brit. Ind. 5: 422. 1887; Trimen, 5-15, filaments free or connate, usually inflexed Handb. Fl. Ceylon 4: 62. 1898; Pax & Hoffm., in bud (except in Koilodepas); pollen grains subglo- Pflanzenr. 147. VII (Heft 63): 19. 1914. TYPE: bose to subprolate, 3-colporate, angulaperturate, Podadenia sapida Thwaites. colpi inoperculate, scarcely marginate, sexine per- A monotypic genus endemic to Sri Lanka. Al- forate-tectate or rather coarsely reticulate. Pistil- though combined by Croizat (1942) with Ptycho- late sepals 4-8, imbricate, sometimes foliaceous or pyxis, it is a very distinctive group, as noted by toothed; disk absent or rudimentary; ovary 3-(4-) Airy Shaw (1963). Until its status can be critically locular; styles free or connate, bifid or multifid. evaluated, it seems best to retain it as a distinct Fruit capsular (rarely drupaceous); seeds subglo- genus. bose, smooth, dry, ecarunculate.

133. Ptychopyxis Miquel, Fl. Ned. Ind. Suppl. This tribeincludes nine paleotropical genera that 402. 1861; Hooker f., Hook. Ic. PI. 18: t. have never all been associated in previous treat- 1703. 1887; Croizat, J. Arnold Arbor. 23: ments, although a number were brought together 47. 1942; Airy Shaw, Kew Bull. 14: 363. under subtribe Cephalocrotoninae by Mueller 1960; 16: 347. 1963; Whitmore, Tree Fl. (1866), and by Pax & Hoffmann(1914, 1931) in Malaya 2: 126. 1973; Airy Shaw, Kew Bull. their series Cladogyniformes.There is considerable Add. Ser. 4: 188. 1975; Add. Ser. 8: 197. resemblance between the Epiprineae and the 1980; Kew Bull. 36: 340. 1981. TYPE: Pty- Agrostistachydeae, and in some respects with the chopyxis costata Miq. Erismantheae.

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KEY TO THE SUBTRIBES OF TRIBE EPIPRINEAE Leaves entire or dentate, pinnately or palmately 1a. Staminatecalyx splitting into distinct segments; veined; staminate flowersin clusters on spicate or pollensexine not spinulose; pistillate sepals often racemose axes, or in capitula; pistillate flowers at persistent,accrescent 27a. Epiprininae same nodes as staminate, or proximal; stamens 4- lobesver- lb. Staminatecalyx turbinate, 2-5-lobed, 10, filamentsfree; pollen grains globose, with rel- rucose; pollensexine spinulose; pistillate sepals caducous.27b. Cephalomappinae atively large colpi, sexine not spinulose; pistillate sepals 4-6, ? persistent; capsule smooth.

Subtribe 27a. EPIPRININAE Muell. Arg., The eight genera of this subtribeoccur in Africa, Linnaea34: 144. 1865. TYPE: EpiprinusGrif- Madagascar, and tropical Asia. Although Pax & fith. Hoffmann (1919, 1931) and Hurusawa (1954) Acalypheaesubtribe Cephalocrotoneae Muell. Arg.,Lin- maintained Epiprinus in a monotypic tribe or sub- naea 34: 143. 1865. TYPE: CephalocrotonHochst. tribe, its relationship to the other genera of Epi- Acalypheae subtribe Mercurialinae series Cladogyni- prineae enumerated below seems clear from re- formesPax & Hoffm.,Pflanzenr. 147. VII (Heft semblances in pollen and floral structures (e.g., 63): 264. 1914. TYPE: Cladogynos Zipp. ex Span. staminate flowers with filaments inflexed in the Epiprineaesubtribe Cleidiocarpinae Thin, Tap Chi Sinh Hoc 10(2): 32. 1988. TYPE: Cleidocarpon Airy bud). Shaw.

KEY TO THE GENERA OF SUBTRIBE EPIPRINAE 1a. Staminateflowers in racemesor spikes;stipules (if present)glandular. 2a. Pistillatecalyx foliose,accrescent, involucrate with deciduous biglandular bracts; styles connate into a column,distally bifid or multifid;stamens 5-15; leaveswith petiolar glands -135. Epiprinus 2b. Pistillatecalyx neitheraccrescent nor involucrate;styles ? free,distally multifid; stamens 3-6. 3a. Filamentsinflexed in bud. 4a. Fruitscapsular; leaves eglandular. 5a. Pistillatesepals entire; staminate flowers not in heads - -136. Symphyllia 5b. Pistillatesepals glandular-dissected; staminate flowers in heads - -- 137. Adenochlaena 4b. Fruitindehiscent; petiole biglandular - - -- 138. Cleidiocarpon 3b. Filamentsnot inflexed in bud - -- -- 139. Koilodepas lb. Staminateflowers in pedunculatecapitula; stipules not glandular. 5a. Inflorescencesaxillary, inconspicuous; stamens 3-5; pistillatesepals accrescent,unlobed; leaves white- tomentosebeneath ------140. Cladogynos 5b. Inflorescencesterminal; leaves not white-tomentosebeneath. 6a. Pistillatesepals entire, connate; styles free; leaves coriaceous - - 141. Cephalocrotonopsis 6b. Pistillatesepals pinnatifid; styles connate; leaves thinner - 142. Cephalocroton

135. Epiprinus Griffith,Notul. P1. Asiat. 4: 487. 763. 1866; Gagnepain,Fl. Indochine5: 477. 1854; Muell. Arg., DC. Prodr. 15(2): 1024. 1926. TYPE: Symphylliasiletiana Baillon. 1866; Bentham,Gen. P1.3: 325. 1880; Hook. f., Fl. Brit.Ind. 5: 463. 1887; Pax & Hoff- A genus of threespecies distributedfrom India mann,Pflanzenr. 147. X (Heft68): 109. 1919; to Hainan and Malaya. Croizat(1942) combined Gagnepain,Bull. Soc. Bot. France 72: 465. Symphylliawith Epiprinus on the basis thatEpi- 1925; Fl. Indochine5: 474. 1926; Croizat, .prinus balansae (Pax & Hoffm.)Gagnep. repre- J. ArnoldArbor. 23: 52. 1942; AiryShaw, sentsa transitionallink between the two taxa. Airy Kew Bull. 16: 356. 1963; Kew Bull. 26: 259. Shaw (1963, 1972) and Thin(1988) followedhim 1972; Whitmore,Tree Fl. Malaya 2: 95. in this, but neitherdiscussed the morphologyof 1973; Thin, Tap Chi Sinh Hoc 10(2): 30. the pistillateflower or demonstratedthat the "in- 1988. TYPE: Epiprinus malayanus Griff. volucre"described by Mueller is withoutsystematic significance.It is possiblethat Croizat was correct, A rathervariable genus of ca. six species in but until the problemis given furtherstudy, it tropical Asia, from Assam to Malaya. Mueller seems preferableto maintainSymphyllia as dis- (1866), followedby Pax & Hoffmann(1919), placed tinct. Epiprinus by itselfin subtribeEpiprininae, on the basis thatthe pistillateflower is involucrate. 137. Adenochlaena Baillon, Etude GCn. Eu- 136. Symphyllia Baillon,Etude Gen. Euphorb. phorb. 472. 1858; Pax, Pflanzenr.147. II 473. 1858; Muell. Arg., DC. Prodr. 15(2): (Heft 44): 12. 1910; Pax & Hoffm.,Natiirl.

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Pflanzenfam.ed. 2, 19c: 124. 1931. TYPE: 1972. TYPE: Cladogynos orientalis Zipp. ex Adenochlaena leucocephala Baillon. Span.

CentrostylisBaillon, Etude Gen. Euphorb.469. 1858. AdenogynumReichenbach & Zollinger,Acta Soc. Regiae TYPE: Centrostyliszeylanica Baillon[= Adenoch- Sci. Indo-Neerl.1: 23. 1856. TYPE: Adenogynum laena zeylanica (Baillon)Thw.]. discolor Reich. & Zoll. [= Cladogynos orientalis Niedenzua Pax, Bot. Jahrb.19: 106. 1894. TYPE: Nie- Zipp. ex Span.]. denzua cordata Pax [= Adenochlaena leucocephala Baillon]. A monotypic genus of tropical southeast Asia, Thailand to the Philippines, Java, and Timor. A genus of two disjunctspecies (Madagascar/ Comores; Sri Lanka); treatedby Mueller (1866) 141. Cephalocrotonopsis Pax, Pflanzenr. 147. as a section of Cephalocroton. II (Heft 44): 15. 1910; Radcliffe-Smith,Kew Bull. 28: 131. 1973. TYPE: Cephalocroton- 138. Cleidiocarpon AiryShaw, Kew Bull. 19: opsis socotrana (Balf. f.) Pax. 313. 1965; Kew Bull. 32: 410. 1978; Thin, Tap Chi Sinh Hoc 10(2): 32. 1988. TYPE: A monotypic genus of Socotra; reduced by Rad- Cleidiocarpon laurinum Airy Shaw. cliffe-Smith(1973) to a section of Cephalocroton.

Bot. 132. SinopimelodendronTsiang, Acta Sinica 15: 142. Cephalocroton Hochstetter, Flora 24: 1973. TYPE: Sinopimelodendron kwangsiense Tsiang [= Cleidiocarpon cavaleriei (Levl.) Airy 370. 1841; Muell. Arg., DC. Prodr. 15(2): Shaw]. 760. 1866; Bentham, Gen. PI. 3: 307. 1880; Pax, Pflanzenr. 147. II (Heft 44): 7. 1910; A genusof twoclosely related species of Burma Prain, Fl. Trop. Afr. 6(1): 843. 1912; Rad- and China; Thin (1988) placed it in a separate cliffe-Smith,Kew Bull. 28: 123. 1973; Fl. E. subtribe,Cleidiocarpinae. Trop. Afr. Euphorb. 1: 282. 1987. TYPE: Cephalocroton cordofanus Hochst. 139. Koilodepas Hasskarl, Versl. Med. Afd. Natuurk. Kon. Akad. Wetensch. 4: 139. An Old World genus of five species recorded 1856; Flora 40: 531. 1857 (as Coelodepas); from east Africa, Madagascar and the Comores, Muell. Arg., DC. Prodr. 15(2): 759. 1866; and Sri Lanka. Bentham,Hook. Ic. P1. 13: t. 1288. 1879; Gen. P1. 3: 313. 1880; Hook. f.,Fl. Brit.Ind. Subtribe 27b. CEPHALOMAPPINAE Web- 5: 419. 1887; Pax, Pflanzenr.147. VII (Heft ster, Taxon 24: 597. 1975. TYPE: Cephalo- 63): 268. 1914; Croizat,J. ArnoldArbor. 23: mappa Baillon. 50. Kew Bull. 14: 382. 1942; Airy Shaw, Leaves entire, pinnately veined; staminate flow- 1960; 16: 354. 1963; 26: 284. 1972; Whit- ers in capitula on racemose axes, pistillate flowers more, Tree Fl. Malaya 2: 103. 1973; Airy at proximal nodes of inflorescence; stamens 2-4, Kew Bull. Add. Ser. 4: 137. 1975; Shaw, filaments connate; pollen grains globose, coarsely Airy Shaw, Kew Bull. 36: 310. 1981. TYPE: reticulate, reticulum spinulose; pistillate sepals 4- Koilodepas bantamense Hassk. 6, deciduous; capsule verrucose.

Calpigyne Blume,Ann. Mus. Bot. Lugd.-Batav.2: 193. This subtribe includes only the type genus. 1857. TYPE: Calpigyne frutescensBi. [= Koilo- depas (B1.) AiryShaw]. ftutescens 143. Cephalomappa Baillon, Adansonia I. 11: NephrostylusGagnepain, Bull. Soc. Bot. France72: 467. 1925. TYPE: Nephrostyluspoilanei Gagnep. [ 130. 1874; Bentham,Gen. PI. 3: 323. 1880; Koilodepas longifoliumHook. f.]. Pax, Pflanzenr.147. II (Heft44): 16. 1910; AiryShaw, Kew Bull. 14: 378. 1960; Kos- A genusof tenspecies of India and southeastern termans,Reinwardtia 5: 413. 1961; Airy Asia as far as Hainan and Borneo. Shaw, Kew Bull. Add. Ser. 4: 66. 1975; Kew Bull. 36: 274. 1981. TYPE: Cephalomappa 140. Cladogynos Zippeliusex Spanoghe,Lin- beccariana Baillon. naea 15: 349. 1841; Muell. Arg.,DC. Prodr. 15(2): 895. 1866; Bentham, Gen. P1. 3: 323. MuricococcumChun & How, Acta Phytotax.Sin. 5: 14. 1880; J. J. Smith,Meded. Dept. Landb. 10: 1956. TYPE: Muricococcumsinense Chun & How & Kos- 383. Pflanzenr.147. VII [= Cephalomappa sinensis (Chun How) 1910; Pax, (Heft term.]. 63): 264. 1914; Gagnepain,Fl. Indochine5: 478. 1926; Backer & Bakhuizen,Fl. Java 1: A genus of fivespecies in southeast Asia (Malaya 485. 1963; AiryShaw, Kew Bull. 26: 232. to Sumatra and Borneo, southern China).

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Tribe 28. ADELIEAE Webster, Taxon 24: 597. A neotropicalgenus of ca. 10- 12 species,Mex- 1975. Acalypheae subtribe Mercurialinae se- ico to Paraguay and Brazilbut best representedin ries Adeliiformes Pax & Hoffm., Pflanzenr. the West Indies. 147. VII (Heft 63): 59. 1919. TYPE: Adelia L. 145. Crotonogynopsis Pax, Bot. Jahrb. 26: 328. 1899; Prain, Fl. Trop. Afr.6(1): 924. Dioecious (rarely monoecious) trees or shrubs; 1912; Pax & Hoffmann,Pflanzenr. 14. XII indumentum simple or stellate; leaves alternate, (Heft63): 14. 1914; Keay, Fl. W. Trop. Afr. simple, entire or dentate, eglandular; stipules small; ed. 2, 1: 404. 1958; Radcliffe-Smith,Fl. Trop. inflorescences axillary, flowersapetalous, in glom- E. Afr. Euphorb. 1: 213. 1987. TYPE: Cro- erules or racemes. Staminate calyx splittinginto 4 tonogynopsisusambarica Pax. or 5 valvate segments; disk extrastaminal, annular or obsolete; stamens 8-18, + connate at base; A monotypicAfrican genus (Zaire to Uganda anthers versatile; pollen grains 3-(4-)colporate, col- and Tanzania). pi operculate, exine tectate-perforate with polyg- onally arranged nanospinules; pistillode present or 146. Enriquebeltrania Rzedowski,Bol. Soc. absent. Pistillate sepals 5 or 6, valvate or open at Bot. Mexico 38: 75. 1979. Beltrania Miran- anthesis, often reflexed; disk annular, glabrous or da, Bol. Soc. Bot. Mexico 21: 4. 1957 (non pubescent; seeds roundish, smooth, ecarunculate Beltrania Penzig, 1882). TYPE: Enriquebel- (rarely carunculate). trania crenatifolia (Miranda) Rzedowski [Beltrania crenatifoliaMiranda]. An American tribe of five genera. A monotypicgenus of the Yucatan peninsula, KEY TO THE GENERA OF TRIBE ADELIEAE verysimilar to Adelia and questionablydistinct. 1a. Indumentumsimple; stipules deciduous; pollen grainswith finely perforate tectum; styles lac- 147. Lasiocroton Grisebach,Fl. Brit.W. Ind. erate. 46. 1859; Abh. K6nigl.Ges. Wiss. Gottingen 2a. Ovary 3-locular;staminate disk present; 9: 20. 1861; Bentham,Gen. P1.3: 317. 1880; stamens8-20; seeds ecarunculate. 3a. Pistillodepresent; staminate disk an- Pax, Pflanzenr.147. VII (Heft63): 60. 1914; nularor segmented;flowers in axillary Fawc. & Rend.,Fl. Jam.4: 293. 1920; Alain, clusters.144. Adelia Fl. Cuba 3: 87. 1953. TYPE: Lasiocrotonmac- 3b. Pistillodeabsent; staminate disk of in- rophyllus(Sw.) Griseb. terstaminalprocesses; flowersin racemes 145. Crotonogynopsis A West Indian genusof fivespecies (Bahamas, 2b. Ovary 2-locular; staminatedisk absent; Cuba, Jamaica,and Hispaniola). stamensmore than 20; seeds carunculate -...... 146. Enriquebeltrania lb. Indumentumstellate or stellate-lepidote;stip- 148. Leucocroton Grisebach,Abh. Konigl. Ges. ules absent; pollengrains with coarsely perfo- Wiss. G6ttingen9: 20. 1861; Muell. Arg., rate tectum. DC. Prodr. 15(2): 757. 1866; Bentham.Gen. bifidor sub- 4a. Pistillatedisk entire;styles P1. 3: 312. 1880; Pax, Pflanzenr.147. VII entire;filaments free; pollen 3-colporate; leaves triplinerved,indumentum stellate (Heft 63): 62. 1914; Urban, Ber. Deutsch. .------147. Lasiocroton Bot. Ges. 36: 504. 1918; Alain,Fl. Cuba 3: 4b. Pistillatedisk lobed; styleslacerate; fila- 89. 1953; Borhidi,Acta Bot. Acad. Sci. Hun- ments? connate;pollen 4-colporate; leaves gar. 21: 222. 1975. TYPE: Leucocroton penninerved,indumentum stellate-lepidote -...... 148. Leucocroton wrightiiGriseb. A genus of 20 species of Cuba and Hispaniola. 144. Adelia L., Syst. Nat. ed. 10, 2: 1298. 1759 (nom. cons.); Bentham, Gen. P1. 3: 312. 29. ALCHORNEAE (Hurusawa)Hutch- 1880; Pax, Pflanzenr. 147. VII (Heft 63): Tribe Bot. 56: 752. 1969. Acaly- 64. 1914; Fawc. & Rend., Fl. Jam. 4: 291. inson,Amer. J. subtribeAlchorneinae Hurus., J. Fac. 1920; Webster, Ann. Missouri Bot. Gard. 54: pheae III. 6: 302. 1954. TYPE: 272. 1968; Sneep & De Roon, Fl. Neth. Ant. Sci. Univ. Tokyo Alchornea Sw. 3: 254. 1984. TYPE: Adelia ricinella L. (typ. cons.). Dioecious trees or shrubs;indumentum simple or stellate;leaves alternate,entire or dentate,pin- Ricinella Muell. Arg., Linnaea 34: 153. 1865. TYPE: Ricinella pedunculosa (A. Rich.) Muell. Arg. [= natelyor palmatelyveined, sometimes stipellate or Adelia ricinella L.]. with laminar glands; stipules mostlydeciduous,

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sometimes obsolete; inflorescences terminal or ax- 3b. Stylesstigmatiform ordilated; indumentum illary, spicate or paniculate (compounded spikes); simple;ovary 3-locular. 4a. Stamens usually 8, free; stigmas bracts glandular or eglandular; flowers apetalous. smooth;pistillate sepals glandularat Staminate calyx splitting into 2-5 valvate seg- base 152. Coelebogyne ments; disk intrastaminal or absent; stamens (2-) 4b. Stamens2-4, basallyconnate. 4-60, free; anthers introrse,muticous or apiculate; 5a. Stamens4; styleselongated, di- pollen grains 3-colporate, colpi operculate, sexine lated; seeds ecarunculate. 153. Aparisthmium rugulose to vermiculate; pistillode rudimentaryor 5b. Stamens2 or 3; stylesstigmati- absent. Pistillate sepals 3-8, imbricate,usually per- form;seeds minutely carunculate sistent,sometimes glandular; disk mostly rudimen- 154. Bocquillonia tary or absent; ovary 2- or 3- (rarely 4-)locular; styles entire to bifid or multifid. Fruit capsular; 149. Orfilea Baillon, Etude Gen. Euphorb. 452. seeds smooth or tuberculate, testa not fleshy. 1858. TYPE: Orfilea coriacea Baillon.

As delimited earlier (Webster, 1975) and here, Diderotia Baillon,Adansonia I. 1: 274. 1861. Laurem- tribe Alchorneae is somewhat more comprehensive bergia Baillon, Etude Gen. Euphorb. 451. 1858 (sphalm.Lautembergia; non LaurembergiaBergi- than the circumscription of Hutchinson (1969), us, 1767). TYPE: Diderotia multispicata(Baillon) since Conceveiba and its allies referred to the Baillon[= Orfilea multispicata (Baillon)Webster, Malloteae by Hutchinson are included. The dis- comb. nov.]. tinctive pollen grains that characterize this tribe A Malagasian genus of four species, closely re- indicate an affinityto Mallotus and related genera, lated to Alchornea but differingin the distinctly but are distinctive in the operculate colpi. bifidstyles. A species from Mauritius discussed by Coode (Kew Bull. 33: 111. 1978) also appears to KEY TO THE SUBTRIBES OF TRIBE ALCHORNEAE belong to this genus: Orfilea neraudiana (Bail- la. Styles entireor if distinctlybifid, then indu- lon) Webster (comb. nov.; based on Claoxylon mentumsimple; stamens 4-9; staminateinflo- neraudianum Baillon, Adansonia I. 1: 280. 1861). rescencesaxillary .. 29a. ALCHORNEINAE lb. Stylesbifid; indumentum stellate; stamens 15- II. 2: 216. 60; staminateinflorescences terminal or axillary 150. Bossera Leandri, Adansonia ...... 29b. CO NCEV EIBINA E 1962. TYPE: Bossera cristatocarpa Leandri. This monotypic genus described from Mada- ALCHORNEINAE Subtribe 29a. Hurusawa, gascar is very close to Alchornea and is weakly Sect. Bot. 6: 302. J. Fac. Sci. Univ. Tokyo, 3, distinguishedby its androecium of ten stamens and se- 1954. Acalypheae subtribe Mercurialinae the distinctivelycristate ovary. ries Alchorneiformes Pax & Hoffm., Pflan- zenr. 147. VII (Heft 63): 220. 1914. TYPE: 151. Alchornea Swartz, Prodr. 6, 98. 1788; Alchornea Sw. Fl. Ind. Occ. 2: 1153. 1800; Muell. Arg., DC. Prodr.15(2): 899. 1866; Fl. Bras. 11(2): Indumentum simple or stellate; staminate inflo- 374. 1874; Bentham,Gen. P1. 3: 314. 1880; rescences axillary; stamens 2-9; pistillate sepals Pax, Pflanzenr.147. VII (Heft63): 220. 1914; eglandular; styles mostlyentire, oftenstigmatiform Webster,Ann. MissouriBot. Gard. 54: 279. or dilated; ovary 2- or 3-(rarely 4-)locular. 1968; 75: 1100. 1989; Whitmore,Tree Fl. This subtribe of six genera with ca. 70 species Malaya 2: 53. 1973; Dyer, Gen. S. Afr.Fl. is well represented in both the Neotropics and Pa- P1. ed. 3, 317. 1975; AiryShaw, Kew Bull. leotropics. Most of the genera are only weakly Add. Ser. 4: 28. 1975; Thin, Tap Chi Sinh separated from Alchornea, and the number may Hoc 6(3): 26. 1984. TYPE: Alchornea lati- be reduced on furtherstudy. folia Sw.

Cladodes Loureiro,Fl. Cochinch.574. 1790. TYPE: Clad- TO THE GENERA OF SUBTRIBE ALCHORNEINAE KEY odes rugosa Lour. [= Alchornea rugosa (Lour.) la. Stylesbifid; indumentum simple. Muell. Arg.]. 2a. Dioecious;ovary smooth; pistillode absent; Hermesia Humb. & Bonpl. ex Willd., Sp. P1. 4: 809. seeds ecarunculate...... 149. Orfilea 1805. TYPE: Hermesiacastanifolia Humb. & Bonpl. 2b. Monoecious;ovary cristate; pistillode pres- ex Willd. [= Alchornea castanifolia (Humb. & ent; seeds carunculate...... 150. Bossera Bonpl. ex Willd.) A. Juss.]. lb. Stylesunlobed or bifidonly at tip. Schousboea Schumacher,Beskr. Guin. P1. 449. 1827 3a. Styleselongated and slender;indumentum (non Willd., 1799). TYPE: Schousboea cordifolia simpleor stellate;ovary 2- or 3-locular. Schum. [= Alchornea cordifolia(Schum.) Muell. ------151. Alchornea Arg.].

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Stipellaria Bentham,J. Bot. Kew Gard.Misc. 6: 2. 1854. 154. Boequillonia Baillon, Adansonia I. 2: 225. TYPE: Stipellaria trewioidesBenth. [= Alchornea 1861; Muell. Arg., DC. Prodr. 15(2): 894. trewioides(Benth.) Muell. Arg.; lectotype,designated by Thin, 1984]. 1866; Bentham, Gen. P1. 3: 313. 1880; Pax, LepidoturusBojer ex Baillon,Etude GCn. Euphorb. 448. Pflanzenr. 147. VII (Heft 63): 260. 1914; 1858. TYPE: Lepidoturusalnifjlius Boj. ex Baillon Airy Shaw, Kew Bull. 29: 321. 1974; Mc- [= Alchornea alnifolia (Baillon)Pax & Hoffm.]. Pherson & Tirel, Fl. Nouv.-Cal6donie 14(1): Bleekeria Miquel, Fl. Ind. Bat. 1(2): 407. 1859 (non 114. 1987. TYPE: Bocquillonia Hassk., 1855). TYPE: Bleekeria zollingeri(Hassk.) sessiliflora Miq. [= Alchornea villosa (Benth.)Muell. Arg.]. Baillon [designated by McPherson & Tirel, 1987]. A variable genus of ca. 50 species distributed in the tropics of both the New World and Old Ramelia Baillon, Adansonia I. 11: 132. 1874. TYPE: Ramelia codonostylisBaillon [= Bocquillonia co- World. Pax (1914) drew the generic boundaries donostylis(Baillon) Airy Shaw]. more narrowlythan Mueller (1866) by recognizing such segregates as Aparisthmium and Coelebo- As treated by McPherson & Tirel (1987), Boc- gyne. However, the Paxian genus still includes quillonia is a genus of 14 species endemic to New three sections, and the relationshipsbetween these Caledonia. and the segregate taxa badly need clarification. Subtribe 29b. CONCEVEIBINAE Webster, 152. Coelebogyne J. Smith, Proc. Linn. Soc. Taxon 24: 597. 1975. TYPE: Conceveiba Aubl. London 1: 41. 1839; Baillon, Etude GCn. Eu- phorb. 416. 1858; Pax, Pflanzenr. 147. VII Leaves long-petiolate, entire or subentire, not stipellate; indumentum stellate (at least in part); (Heft 63): 255. 1914. TYPE: Coelebogyne ilicifolia J. Smith. floral disk absent; stamens 15-60; pistillate sepals often glandular; ovary 2-3-locular; styles bifid. A monotypicgenus fromtropical Australia, very This subtribe of three weakly distinguishedfrom Alchornea and perhaps genera and about ten was better treated as a section of that genus. species thought to be entirely American until the recent discovery of Conceveiba in west Africa. Pax Hoffmann 153. Aparisthmium Endlicher, Gen. P1. 1112. & (1914) included Conceveiba and in 1840; nom. cons. prop.; Pax, Pflanzenr. 147. Gavarretia their series Trewiiformes (Mallo- teae), an VII (Heft 63): 257. 1914; Jablonski, Mem. entirely paleotropical group. New York Bot. Gard. 17: 135. 1967. Con- ceveibum A. Rich. ex A. Jussieu, Euphorb. KEY TO THE GENERA OF SUBTRIBE CONCEVEIBINAE Tent. 42. 1824. TYPE: Conceveibum corda- la. Inflorescencesterminal. tum A. Juss. [= Aparisthmium cordatum (A. 2a. Pistillatesepals separate,often glandular; anthersmuticous; ovary 3-locular;style- Juss.) Baillon). branchesblunt 155. Conceveiba 2b. Pistillatesepals eglandular,connate, lobes In the concept of Pax (1914), accepted by Ja- scarcelyevident; anthers apiculate; ovary blonski and other neotropical floristic authors, 2-locular;style-branches acute. Aparisthmium is a monotypic neotropical genus -...... 156. G avarretia widely distributed from Colombia to Bolivia and lb. Inflorescencesaxillary; anthers muticous. southern Brazil. Unfortunately,the genus is tech- ...... 157.Polyandra nically a synonymof Conceveiba, because Endlich- er cited A. Jussieu's figure42B, which in the legend 155. Coneeveiba Aublet, Hist. P1. Guiane Fr. is clearly indicated as Aublet's Conceveiba gui- 923. 1775; Muell. Arg., DC. Prodr. 15(2): anensis. Endlicher's description,on the other hand, 895. 1866; Bentham,Gen. P1. 3: 316. 1880; clearly seems to fitthe plant that Baillon and later Pax, Pflanzenr.147. VII (Heft63): 214. 1914; writers have accepted as Aparisthmium. If End- Jablonski,Mem. New York Bot. Gard. 17: licher's citation is regarded as a misprint, then 131. 1967; Thomas, Ann. Missouri Bot. Gard. Aparisthmium becomes a synonym of Concevei- 77: 856. 1990. TYPE: Conceveiba guianensis bum Rich. ex A. Juss., unless Conceveibum can Aubl. be regarded as an intentionalorthographic variant of Conceveiba. The easiest course out of this tangle Conceveibastrum(Muell. Arg.) Pax & Hoffm.,Pflanzenr. would be to propose the generic name Aparisth- 147. VII (Heft 63): 217. 1914. TYPE: Concevei- bastrummartianum (Baillon) Pax & Hoffm.[= Con- mium for which seems in conservation, warranted ceveiba martiana Baillon]. view of the abundance of the plant in neotropical Veconcibea(Muell. Arg.)Pax & Hoffm.,Pflanzenr. 147. vegetation. VII (Heft 63): 218. 1914. TYPE: Veconcibea lati-

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folia (Benth.)Pax & Hoffm.[= Conceveibalatifolia Tribe 30. ACALYPHEAE Dumortier,Anal. Benth.]. Fam. P1. 45. 1829. TYPE: AcalyphaL. A genus of seven or eight neotropical species Monoeciousor dioecioustrees, shrub, or herbs; extending fromCosta Rica to Amazonian Peru and indumentumsimple or stellate;leaves alternateor Brazil, and one recently discovered African species opposite,simple, entire or dentate,pinnately or from Gabon (Thomas, 1990). palmatelynerved, often punctate or withlaminar 156. Gavarretia Baillon, Adansonia I. 1: 185, glands, stipulate;inflorescences terminal or axil- t. 7. 1861; Bentham, Gen. PI. 3: 316. 1880; lary,unisexual or bisexual,bracts sometimes glan- Pax, Pflanzenr. 147. VII (Heft 63): 213. 1914; dular; flowersapetalous. Staminate calyx closedin Jablonski, Mem. New York Bot. Gard. 17: bud, valvatelysplitting into 2-5 segments;disk 130. 1967. TYPE: Gavarretia terminalis Bail- receptacular(intrastaminal), of interstaminalpro- Ion. cesses, or absent; stamens4-many, usuallyfree; anthers sometimesapiculate, anther-sacssome- A monotypic South American genus; the single times pendulous;pollen grains globose,brevicol- species occurs in Amazonian Venezuela, Guiana, porate or porate,apertures not or scarcelyoper- and Brazil. Mueller (1874) treated Gavarretia as culate; sexine coarsely foveolate to striate or a section of Conceveiba, and this may yet prove rugulose;pistillode usually absent. Pistillate sepals to be the best systematic disposition. mostly3-6, free;disk cupular or absent;ovary 2- basally connate,unlobed 157. Polyandra Leal, Arch. Jard. Bot. Rio de 4-locular; stylesfree or Fruit capsular or drupaceous; seeds Janeiro 11: 63. 1951. TYPE: Polyandra brac- to lacerate. testa dryor fleshy. teosa Leal. carunculateor ecarunculate, A poorly understood genus known only fromthe The largesttribe of Acalyphoideae,here divided type species in the Brazilian Amazon; pistillateflow- into 11 subtribeswith 30 genera and over 1000 ers and fruitsare still unknown. species.

KEY TO THE SUBTRIBES OF TRIBE ACALYPHEAE 1a. Seeds carunculate;inflorescences terminal; indumentum stellate or absent;styles bifid. 2a. Monoecious;filaments connate into fascicles; pistillate sepals deciduous; stipules connate - 30a. Ricininae 2b. Dioecious; filamentsfree; pistillate sepals persistent;stipules free ... 30b. Adrianinae lb. Seeds ecarunculate,or if carunculatethen inflorescences axillary, or stylesunlobed. 3a. Stamensnot connateinto fascicles. 4a. Anther-sacsnot vermiformand pendulous;pollen colporate; seeds usuallyecarunculate. 5a. Indumentumusually simple (except Lobaniliinae);leaves alternate,eglandular or withem- beddedlaminar glands, not granulose-glandular. 6a. Herbs,often with opposite leaves; indumentumsimple; styles undivided. 7a. Staminatesepals free; stamensfree; ovary 2-locular; pollen grains 3-colporate; anthersnot apiculate;cotyledons broader than radicle ..... 30c. Mercurialinae 7b. Staminatesepals connate;filaments connate; ovary 3-locular; pollen grains inaperturate;anthers apiculate; cotyledons scarcely broader than radicle - 30d. Dysopsidinae 6b. Shrubsor trees(or herbswith bifid styles). 8a. Anther-sacsnot partedto the connective;staminate disk absent; leaves mostlywith laminarglands. 9a. Stylesbifid; anthers 2-locular; pollen grains vermiculate-rugulose; seed-coat dry or fleshy...... 30e. Cleidiinae 9b. Stylesunlobed; anthers 3-4-locular; pollen grains spinulose-rugulose; seed-coat usuallyfleshy ...... 30f. Macaranginae 8b. Anther-sacsparted to theconnective, erect or pendulous;interstaminal disk present; stylesunlobed. 1lOa. Indumentumsimple ...... 30g. Claoxylinae lOb. Indumentumstellate ...... 30h. Lobaniliinae 5b. Indumentumusually stellate; leaves oftenopposite, often with embedded laminary glands and usuallyglandular-granulose as well;usually dioecious; inflorescences often terminal; staminate diskpresent or absent;pollen grains ? spinulose-rugulose... -..- ...... 30i. Rottlerinae 4b. Anthersvermiform and pendulose;staminate disk absent;pollen porate; styles lacerate; seeds + carunculate;pistillate bracts usually large and foliaceous...... 30j. Acalyphinae 3b. Stamensmany, connate into fascicles; shrubs or trees withalternate eglandular leaves, indumentum simple or lepidote...... -...... -...30k. Lasiococcinae

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Subtribe 30a. RICININAE Grisebach,Fl. Brit. stipules glandular; inflorescences spicate, terminal W. Ind. 37. 1859; Abh. Konigl. Ges. Wiss. or opposite the leaves, bracts glandular; stamens GCttingen9: 15. 1860; Muell.Arg., DC. Prodr. numerous, free; anthers apiculate, sacs adnate to 15(2): 143. 1866; Pax & Hoffm.,Pflanzenr. linear connective; ovary 3-locular, smooth; styles 147. XI (Heft68): 112. 1919; Natiirl.Pflan- bifid; fruitcapsular; seeds carunculate. zenfam. ed. 2, 19c: 149. 1931; Webster, In contrast to the concepts of Taxon 24: 597. 1975. TYPE: Ricinus L. Bentham (1880) and Pax (1910), subtribe Adrianinae is here re- Monoecioustrees or shrubs(herbs in temperate stricted to the type genus Adriana. Pax (1910) climates); indumentumabsent; leaves alternate, rather slavishlyfollowed Bentham in includingwith- long-petiolate,palmately lobed; stipules connate in his Adrianinae such unrelated genera as Manihot into a sheath,deciduous; inflorescences terminal, and Pachystroma. In his later work (Pax & Hoff- paniculate,proximal cymules staminate, distal pis- mann, 1931), he abandoned the tribe Adrianeae tillate;bracts glandular; stamens many, filaments and included Adriana in his subtribeMercurialinae united into branching fascicles; pollen grains adjacent to Conceveiba and Gavarretia, which is 3-colporate,colpi narrowand inoperculate;pistil- a better and not implausible suggestion of affinity. late flowerspedicellate; ovary muricate; styles bifid; capsule echinate;seeds carunculate. 159. Adriana Gaudichaud, Ann. Sci. Nat. Paris As here defined,subtribe Ricininae is restricted 5: 223. 1825; Muell. Arg., DC. Prodr. 15(2): to the single genus Ricinus. Pax & Hoffmann 889. 1866; Bentham, Fl. Austral. 6: 133. (1919) also includedHomonoia and Lasiococca 1873; Gen. P1. 3: 306. 1880; Pax, Pflanzenr. in the Ricininae,mainly because of theirrather 147. II (Heft 44): 17. 1910; Airy Shaw, Kew similarandroecium of stamens united into fascicles. Bull. 35: 589. 1980. TYPE: Adriana tomen- However,Ricinus differsfrom those genera (here tosa Gaud. [lectotype]. referredto subtribeLasiococcinae) in its terminal A genus endemic to Australia; treated by Airy inflorescence,distinctly carunculate seeds, pal- Shaw (1980) as having five species and several matelylobed leaves withglandular petiole, and bifid varieties. The resemblances between Ricinus and styles.I am inclinedto agreewith Airy Shaw (1974) Adriana appear too significant to be explained thatthe characterof fasciculatestamens is homo- away as due to convergence, despite the great plasious,and the Ricininaedo not appear closely geographical disjunction. However, Adriana also relatedto the Lasiococcinae. On the other hand, has a number of characters in common withgenera Adriana is much more similarto Ricinus and such as Cephalocroton and Cephalomappa that appears to be the mostclosely related genus. were included in the Adrianeae by Pax (1919) but are referredto the Epiprineae here. It is possible, 158. Ricinus L., Sp. P1. 1007. 1753; Gen. P1. therefore,that Adriana provides the phylogenetic ed. 5, 437. 1754; Muell. Arg., DC. Prodr. link between the tribe Epiprineae and subtribe Ri- 15(2): 1016. 1866; Bentham,Gen. P1. 3: cininae. It may be necessary to remove the Ad- 321. 1880; Pax & Hoffm.,Pflanzenr. 147. rianinae and Ricininae fromthe Acalypheae as we XI (Heft68): 119. 1919; Webster,J. Arnold accumulate furtherknowledge. Arbor.48: 379. 1967; Correll& Correll,Fl. Bahama Arch. 840. 1982; Radcliffe-Smith, Subtribe 30c. MERCURIALINAE Fl. E. Trop. Afr. Euphorb. 1: 322. 1987. Pax, Na- TYPE: Ricinus communisL. turl. Pflanzenfam. ed. 1, 3(5): 46. 1890. Aca- lypheae subtribe Mercurialinae series Mer- A monotypicgenus native to northeastAfrica curialiiformesPax & Hoffm., Pflanzenr. 147. (accordingto Radcliffe-Smith),now widelyculti- VII (Heft 63): 270. 1914. TYPE: Mercurial- vatedand ruderalin tropicaland subtropicalregions is L. worldwide. Monoecious or dioecious herbs; indumentum simple; leaves alternate or Subtribe 30b. ADRIANINAE Bentham,Gen. opposite, pinnately veined, eglandular; P1. 3: 250. 1880; Webster,Taxon 24: 597. inflorescences axillary, spicate or in glomerules; stamens 1975. Tribe Adrianeae (Benth.) Pax, Pflan- 4-20 (rarely fewer),free; anthers extrorse, muticous; pollen grains 3- zenr. 147. II (Heft 44): 1. 1910. TYPE: Ad- riana Gaud. colporate, colpi operculate; ovary 2-locular, smooth; styles unlobed; fruitcapsular, smooth or muricate; Dioecious;indumentum stellate or absent;leaves seeds carunculate; cotyledons much broader than alternateor opposite,unlobed or palmatelylobed, radicle.

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This subtribe includes three genera, one Eur- the name Leidesia procumbens (L.) Prain for the asian and the other two South African. two species of Mueller combined, this epithet belongs to the same taxon as Leidesia capensis Muell. KEY TO THE GENERA OF SUBTRIBE MERCURIALINAE Arg., which unfortunatelyis based on a confusion 1 a. Dioecious; stamens8-20; pistillateflower with with Urtica capensis L. f. (a name referable to a 2 subulatestaminodia (or disk-segments);seeds species of Acalypha). It thereforeseems better to carunculate. -... . 160. Mercurialis choose Mueller's other species as lectotype. lb. Monoecious;stamens 2-7; pistillatestaminodia absentor verysmall; seeds ecarunculate. 2a. Capsule smooth;pistillate sepals 3; leaves Subtribe 30d. DYSOPSIDINAE Hurusawa,J. entireor denticulate .. 161. Seidelia Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: 302. 2b. Capsule setose-muricate;pistillate sepals 1954. TYPE: Dysopsis Baillon. nearlyobsolete; leaves crenulate. .. . . 162. Leidesia Monoecious herbs; leaves alternate, simple, crenate, eglandular; stipules caducous; flowers axillary, mostly solitary; staminate calyx gamophyl- 160. Mercurialis L., Sp. PI. 2: 1035. 1753; Gen. PI. ed. 5, 437. 1754; Muell. Arg., DC. lous; stamens 3-6, interior connate; anthers apiculate, sacs adnate to connective; pollen grains Prodr. 15(2): 794. 1866; Bentham, Gen. PI. globose, porate; pistillatesepals 3; ovary 3-locular, 3: 309. 1880; Pax & Hoffm.,Pflanzenr. 147. fruit VII (Heft 63): 271. 1914; Zimmermann et pubescent; styles unlobed, lacerate; capsular; seeds with obsolete caruncle. al., in Hegi, Ill. Fl. Mitteleur. 5(1): 126. 1925; Webster, J. Arnold Arbor. 48: 366. 1967; A monogeneric tribe of South America; closely Tutin, Fl. Europaea 2: 212. 1968. TYPE: Mer- related to the Mercurialinae, and furtherstudy may curialis perennis L. [lectotype, designated by indicate that the two subtribes should be combined. Small, in Britton& Brown,Ill. Fl. N. U.S. ed. 2: 2, 460. 1913]. 163. Dysopsis Baillon, Etude GCn. Euphorb. A Eurasian genus of eight species, seven in 435. 1858; Muell. Arg., DC. Prodr. 15(2): Europe and Mediterranean Africa, and one in east 949. 1866; Bentham, Gen. P1. 3: 264. 1880; temperate Asia. Grining, Pflanzenr. 147. (Heft 58): 10. 1913; Pax & Hoffmann, Pflanzenr. 147. VII (Heft 63): 286. 1914. Molina Gay, Hist. Chile Bot. 161. Seidelia Baillon, Etude GCn. Euphorb. 465. 5: 345. 1851 (non Molina Cav., 1790). TYPE: 1858; Bentham, Gen. PI. 3: 310. 1880; Prain, Molina chilensis Gay = Dysopsis gayana Ann. Bot. 27: 398. 1913; Pax & Hoffm., Baillon, nom. illeg. [= Dysopsis glechomoides Pflanzenr. 147. VII (Heft 63): 282. 1914; (A. Rich.) Muell. Arg.]. Prain, Fl. Capensis 5(2): 464. 1920; Dyer, Gen. S. Afr. Fl. PI., ed. 3, 316. 1975. TYPE: A monotypic genus of the South American An- Seidelia mercurialis Baillon (nom. illeg.) [= des (Ecuador to Chile), Juan Fernandez, and Costa Seidelia triandra (E. Mey.) Pax; lectotype, Rica. designated by Pfeiffer,Nomencl. Bot. 2: 1128. 1874]. Subtribe 30e. CLEIDIINAE Webster,Taxon A South African genus of two species. 24: 598. 1975. TYPE: Cleidion Blume. Monoecious or dioecious trees or shrubs; indu- 162. Leidesia Muell. Arg., DC. Prodr. 15(2): mentum simple; leaves alternate, simple, pinnately 792. 1866; Bentham, Hook. Ic. PI. 13: 66, veined, with laminar glands, stipulate; inflores- t. 1284. 1879; Gen. PI. 3: 310. 1880; Pax cences axillary, spicate or racemose; bracts eglan- & Hoffm.,Pflanzenr. 147. VII (Heft 63): 284. dular; flowers apetalous, without disk; staminate 1914; Prain, Ann. Bot. 27: 399. 1913; Fl. sepals 3 or 4; stamens 25-80, free or basally Capensis5(2): 462. 1920; Dyer,Gen. S. Afr. connate; anthersmuticous or apiculate; pollen grains Fl. PI., ed. 3, 316. 1975. TYPE: Leidesia 3-colporate, colpi not operculate; sexine rugulose- obtusa (Thunb.) Muell. Arg. [loc. cit. 793; vermiculate, spinules vestigial; pistillode absent; lectotype, chosen here]. pistillatesepals 3-6, imbricate; ovary (2-)3-locular; styles bifid,branches elongate; fruitcapsular; seeds A South African genus of two or three species, with dry or fleshy testa. very closely related to Seidelia; the two genera could certainlybe combined withoutobscuring phy- As here construed, subtribe Cleidiinae includes logeny. Although Pax & Hoffmann (1914) used three genera, of which two are paleotropical and

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the third pantropical. The characteristic globose Lasiostyles Presl, Abh. Konigl. B6hm.Ges. Wiss. V. 3: pollen grains with vermiculate sexine ornamenta- 579. 1845. TYPE: Lasiostyles salicifolia Presl [ Cleidion Blume]. tion indicate an affinitybetween Wetria and Clei- Javanicum Tetraglossa Beddome, Madras J. Lit. Sci. II. 22: 70. dion not clearly expressed in previous classifica- 1861. TYPE: Tetraglossa indica Beddome[= Clei- tions. Pax & Hoffmann(1914) assigned Wetria to dion JavanicumBlume]. the series Trewiiformes adjacent to Alchorneae A well-markedgenus of ca. 25 species, of which subtr. Conceveibinae; however, in those genera the 5 are neotropical, 1 west African, 12 New Cale- indumentum is stellate and the pollen grains oper- donian, and the remainder tropical Asian. culate.

166. Sampantaea Airy Shaw, Kew Bull. 26: KEY TO THE GENERA OF SUBTRIBE CLEIDIINAE 328. 1972; Hook. Ic. P1. 38: t. 3717. 1974. la. Staminateflowers pedicellate, usually 2 or more TYPE: Sampantaea amentifiora (Airy Shaw) per bract; stamens25-80. 2a. Anthersmuticous, connective scarcely en- Airy Shaw. larged; stamens25-30; pedicel pistillate A monotypic genus restricted to Thailand and less than 1 cm long 164. Wetria 2b. Anthersapiculate, connectiveenlarged; Cambodia. It appears closely related to Wetria and stamensmostly 30-80 or more; pistillate may prove congeneric. pedicelover 1 cm long 165. Cleidion lb. Staminateflowers sessile, 1 per bract;stamens Subtribe 30f. MACARANGINAE (Hutchin- 15-25, anthersapiculate, connective not enlarged 166. Sampantaea son) Webster, Taxon 24: 598. 1975. Tribe Macarangeae Hutchinson, Amer. J. Bot. 56: 164. Wetria Baillon, Etude GCn. Euphorb. 409. 755. 1969. TYPE: Macaranga Du Petit 1858; J. J. Smith, Med. Dept. Landb. 10: Thouars. 470. 1910; Pax & Hoffm., Pflanzenr. VII Dioecious trees or shrubs; indumentum simple; (Heft 63): 219. 1914; Backer & Bakhuizen, leaves alternate, unlobed or palmately lobed, pin- Fl. Java 1: 485. 1963; Airy Shaw, Kew Bull. nately to palmately veined, glandular or eglandular; 26: 350. 1972; Whitmore, Tree Fl. Malaya stipules minute to enlarged; inflorescencesaxillary, 2: 136. 1973; Airy Shaw, Kew Bull. 36: 358. racemose or paniculate; bracts ? glandular; flow- 1981. TYPE: Wetria trewioides Baillon, nom. ers apetalous and lacking disk; staminate calyx of illeg. [= Wetria insignis (Steud.) Airy Shaw]. 2-5 segments; stamens (1)2-50, free; anthers mu- A monotypic genus, the single species occurring ticous, 2-4-valved; pollen grains 3- or 4-colporate, from Burma to Indonesia and New Guinea. Airy sexine finelyto coarsely rugulose-spinulose; pistil- Shaw has suggested that the genus is closely related late sepals 3-5, free or connate; ovary (1 -)2- or to Alchornea, but it differsin its simple indumen- 3-(-6)locular; styles unlobed; fruitcapsular; seeds tum, bifid styles, and larger stamen number. ecarunculate, testa fleshy. As definedby Webster (1975) and here, subtribe 165. Cleidion Blume, Bijdr. Fl. Ned. Ind. 612. Macaranginae is much more narrowlydefined than 1826; Muell. Arg., DC. Prodr. 15(2): 983. Hutchinson's tribe Macarangeae, which included 1866; Bentham, Gen. PI. 3: 320. 1880; Pax seven other genera from various tribes of subfam- & Hoffm.,Pflanzenr. 147. VII (Heft 63): 288. ilies Acalyphoideae and Crotonoideae; only the ge- 1914; Croizat, J. Arnold Arbor. 24: 166. nus Macaranga is included here. 1943; Leonard, Bull. Jard. Bot. Nat. Belge 42: 297. 1972; Leandri, Adansonia II. 12: 167. Macaranga Du Petit Thouars, Gen. Nova 193. 1972; A. C. Smith, Fl. Vitiensis Nova Madag. 26. 1806; Muell. Arg., DC. Prodr. 2: 514. 1981; McPherson & Tirel, Fl. Nouv.- 15(2): 987. 1866; Bentham, Gen. P1. 3: 320. Caled. 14: 143. 1987; Huft, Ann. Missouri 1880; Hook. f., Fl. Brit. Ind. 5: 445. 1887; Bot. Gard. 75: 1103. 1989. TYPE: Cleidion Pax & Hoffm.,Pflanzenr. 147. VII (Heft 63): javanicum Blume. 298. 1914; Gagnepain, Fl. Indochine 5: 434. Redia Casaretto,Nov. Stirp.Bras. 51. 1843. TYPE: Re- 1926; Perry, J. Arnold Arbor. 34: 191. 1953; dia tricoccaCasar. [= Cleidion tricoccum(Casar.) Whitmore, Tree Fl. Malaya 2: 105. 1973; Baillon]. PsilostachysTurcz., Bull. Soc. Imp. NaturalistesMoscou Kew Bull. Add. Ser. 4: 140. 1975; Kew Bull. 16: 58. 1843. TYPE: Psilostachysaxillaris Turcz. Add. Ser. 8: 123. 1980; A. C. Smith, Fl. [= Cleidion tricoccum(Casar.) Baillon]. Vitiensis Nova 2: 500. 1981; Coode, Taxon

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25: 184. 1976; McPherson & Tirel, Fl. Nouv.- Subtribe 30g. CLAOXYLINAE Hurusawa,J. Caled. 14: 172. 1987; Radcliffe-Smith,Fl. E. Fac. Sci. Univ. Tokyo,Sect. 3, Bot. 6: 301. Trop. Afr. Euphorb. 1: 239. 1987. TYPE: Ma- 1954. Tribe Acalypheaesubtribe Mercuriali- caranga mauritiana Bojer ex Muell. Arg. nae series ClaoxyliformesPax & Hoffm., [lectotype, selected by Coode, 1976, who re- Pflanzenr.147. VII (Heft63): 75. 1914. Tribe jected the proposed selection of M. roxburgii ClaoxyleaeHutchinson, Amer. J. Bot. 56: 752. Wight by Wheeler, 1975]. 1969. TYPE: Claoxylon A. Juss. TribeMareyeae Hutchinson, Amer. J. Bot. 56: 751. A. Jussieu,Euphorb. Tent. 44. 1824. TYPE: Map- Mappa 1969. TYPE: MareyaBaillon. pa glabra A. Juss.[= Macaranga glabra (A. Juss.) Pax & Hoffm.]. Monoeciousor dioecioustrees, shrubs, or herbs; 1826. PachystemonBlume, Bijdr. Fl. Ned. Ind. 626. indumentumsimple; leaves alternate, unlobed, TYPE: Pachystemontrilobum (Reinw. ex Bi.) Bl. [ Macaranga triloba(Reinw. ex Bl.) Muell. Arg.]. mostlydentate, pinnately veined, without laminar MecostylisKurz ex Teysmann& Binnendijk,Natuurk. glands;stipules small, persistent or deciduous;in- Tijdschr.Ned.-Indie 27: 44. 1864. TYPE: Mecostylis florescencesaxillary, racemose or paniculate;bracts acalyphoides Kurz ex Teysm. & Binn. [= Maca- eglandular;flowers apetalous; staminate sepals 2- ranga involucrata(Roxb.) Baillon]. mostlyof interstaminal processes or annular Phocea Seemann, J. Bot. 8: 69. 1870. TYPE: Phocea 5; disk andersoaniSeem. [= Macaranga sp.; notidentified (rarelyabsent); stamens 5-40(-200), free;anther by Pax &8 Hoffm.,1931, or McPherson& Tirel, sacs separatedto the connective,? erect; pollen 1987]. grainsrugulose-spinulose; pistillate sepals 2-4, open or imbricate;disk annular or dissected;ovary most- A and diverse paleotropical genus of very large ly 2- or 3-locular;styles ? elongated,unlobed but there are about 50 species in ca. 300 species; lacerate; fruitcapsular; seeds ecarunculate,testa over 200 in tropical Asia from Africa/Madagascar, usuallyfleshy. India to New Guinea, and a few in Fiji and other Pacific islands. The genus has acquired several A paleotropicalsubtribe of nine genera, some generic synonyms, but no plausible system of seg- hardlydistinct, and othersnot certainlybelonging regate genera has ever been proposed. here.

KEY TO THE GENERA OF SUBTRIBE CLAOXYLINAE la. Stylesundivided; pistillate disk unlobed to 5-lobed. 2a. Buds perulate;stipules mostly persistent . . . . 168. Erythrococca 2b. Buds not perulate;stipules mostly deciduous. 3a. Racemes uniformlyfloriferous; capsules coriaceous;leaves not stipellate. 4a. Staminatedisk of interstaminalsegments. 5a. Anthersacs erect; mostlydioecious. 6a. Stamensmostly 20 or more; stylesrecurved, papillose but hardlylacerate. .~~~~~~~~~~~~~~~~~1 69. Claoxylonz 6b. Stamensup to 15; styleserect, lacerate .170. Claoxylopsis 5b. Anthersacs pendent;styles lacerate; stamens 10-40; monoecious.. 171. Mareya 4b. Staminatedisk urceolate;stamens 6-12; stylesrecurved, lacerate .. 172. Discoclaoxylon 3b. Racemes interrupted;capsules crustaceous;leaves stipellate . . . 173. Micrococca lb. Stylesbifid; pistillate disk 8-10-lobed; staminatedisk of interstaminalsegments; stamens up to 25; seed- coat not fleshy.174..Amyrea

168. Erythrococca Bentham,Niger Fl. 506. Poggeophyton aculeatum Pax [= Erythrococca 1849; Muell. Arg., DC. Prodr. 15(2): 790. poggeophytonPrain]. ChloropataneEngler, Bot. Jahrb.26: 383. 1899. TYPE: 1866; Bentham,Gen. P1. 3: 308. 1880; Prami, Chloropatane africana (Baillon) Engler [= Ery- Ann. Bot. 25: 606. 1911; Fl. Trop. Afr.6(1): thrococcaafricana (Baillon)Prain]. 847. 1912; Pax & Hoffm.,Pflanzenr. 147. Athroandra(Hook. f.) Pax & Hoffm.,Pflanzenr. 147. VII (Heft63): 86. 1914; Keay, Fl. W. Trop. VII (Heft63): 76. 1914. TYPE: Claoxylon mannai f.) Prain]. Afr.ed. 2, 1: 400. 1954; Radcliffe-Smith,Fl. Hook. f. [= Erythrococcainannii (Hook. E. Trop. Afr.Euphorb. 1: 265. 1987. TYPE: A diverse African genus of ca. 50 species, whose Erythrococcaaculeata Benth.,nom. illeg. [= delimitation apparently requires further analysis. Erythrococcaanomala (Juss.ex Poir.)Prain]. Although Prain, Keay, and Radcliffe-Smithtreat PoggeophytonPax, Bot. Jahrb.19: 88. 1894. TYPE: Athroandra as part of Erythrococca, Airy Shaw

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(1966) suggestedthat Erythrococca is close to in the lectotype species appropriately reflects Claoxylon. the taxon name].

A mainly west African genus of three species, 169. Claoxylon A. Jussieu,Euphorb. Tent. 43. extending fromSierra Leone and Fernando P6o to 1824; Muell. Arg., DC. Prodr. 15(2): 775. Uganda. 1866; Bentham,Gen. P1. 3: 309. 1880; Pax & Hoffm.,Pflanzenr. 147. VII (Heft63): 100. 173. Micrococca Bentham, Niger Fl. 503. 1914; Merrill,Enum. Phil. P1. 2: 429. 1923; 1849; Gen. P1. 3: 309. 1880; Prain, Ann. Whitmore,Tree Fl. Malaya 2: 78. 1973; A. Bot. 25: 628. 1911; Fl. Trop. Afr. 6(1): 876. C. Smith,Fl. Vit. Nova 2: 516. 1981. TYPE: 1912; Airy Shaw, Kew Bull. 25: 524. 1971; Claoxylon parvifiorum A. Juss. Dyer, Gen. S. Afr. Fl. P1., ed. 3, 1: 315. ErytrochilusReinw. ex Blume,Bijdr. Fl. Ned.-Ind.614. 1975; Radcliffe-Smith,Fl. E. Trop. Afr. Eu- 1826. TYPE: Erytrochilusindicus Reinw. ex Bl. [= phorb. 1: 260. 1987. TYPE: Micrococca mer- Claoxylon indicum(Reinw. ex Bl.) Hassk.; desig- curialis (L.) Benth. nated by Wheeler,1975]. QuadrasiaElmer, Leafl. Phil. Bot. 7: 2656. 1915. TYPE: A paleotropical genus of 12 species, reported Quadrasia euphorbioidesElmer [= Claoxyloneu- from tropical Africa and Madagascar to Malaya. phorbioides(Elmer) Merr.].

A paleotropicalgenus of ca. 75 species,absent 174. Amyrea Leandri, Notul. Syst. (Paris) 9: fromAfrica but extendingfrom Madagascar to 168. 1940. TYPE: Amyrea sambiranensis Melanesia and Hawaii. Leandri[lectotype, selected here].

170. Claoxylopsis Leandri, Bull. Soc. Bot. A genus of two species endemic to Madagascar. France 85: 526. 1938; Radcliffe-Smith,Kew Amyrea diverges from all of the other genera of subtribe Claoxylinae in its bifid styles, and its po- Bull. 43: 642. 1988. TYPE: Claoxylopsis per- sition requires evaluation. rieri Leandri. A genus of threespecies endemicto Madagas- Subtribe 30h. LOBANILIINAE Radcliffe- car. Althoughapparently very close to Claoxylon, Smith, Kew Bull. 44: 339. 1989. TYPE: Lo- it is stillpoorly known and its positionand status banilia Radcl.-Sm. mustbe regardedas tentative. Dioecious trees or shrubs; indumentum stellate 171. Mareya Baillon,Adansonia I. 1: 73. 1860; (sometimes simple as well); leaves alternate, un- Muell. Arg., DC. Prodr. 15(2): 792. 1866; lobed, mostly pinnately veined, without embedded Bentham,Hook. Ic. P1. 13: 63, t. 128. 1879; or granular laminar glands; stipules minute and Gen. P1. 3: 312. 1880; Prain,Fl. Trop. Afr. deciduous; inflorescences axillary, racemose; flow- 6(1): 910. 1912; Pax & Hoffm.,Pflanzenr. ers apetalous; staminate sepals 3: disk of inter- 147. XIV (Heft68): 11. 1919; Leonard,Bull. staminal segments; stamens 17-30, free; anther- Jard. Bot. Brux. 25: 291. 1945; Radcliffe- sacs free to base; pistillate sepals 3; disk annular or ? Smith,Fl. E. Trop. Afr. Euphorb. 1: 216. dissected; ovary 3-4-locular; styles unlobed, papillose or plumose; fruitcapsular; seeds ecarun- 1987. TYPE: Mareya spicata Baillon. culate. MareyopsisPax & Hoffm.,Pflanzenr. 147. XIV (Heft 68): 13. 1919. TYPE: Mareyopsis longifolia(Pax) This monotypic subtribe, recently proposed by Pax & Hoffm.[= Mareya longifoliaPax]. Radcliffe-Smith,appears very similar to subtribe Claoxylinae except for the stellate indumentum. It An Africangenus of threespecies. is provisionallyrecognized here until detailed corf- parisons with taxa of Claoxylinae can be made to 172. Discoclaoxylon (Muell. Arg.) Pax & determine whether Lobanilia should be kept in a Hoffmann,Wiss. Ergebn. Deutsche Zentral- separate subtribe. Afr. Exped. 2: 452. 1912; Pflanzenr.147. VII (Heft 63): 137. 1914; Radcliffe-Smith, 175. Lobanilia Radcliffe-Smith,Kew Bull. 44: Fl. E. Trop. Afr. Euphorb. 1: 279. 1987. 334. 1989. TYPE: Lobanilia luteobrunnea Claoxylon sect. Discoclaoxylon Muell. Arg., (Baker) Radcl.-Sm. Flora 47: 137. 1864. TYPE: Discoclaoxylon hexandrum (Muell. Arg.) Pax & Hoffm.[lec- A genus of seven species confined to Madagas- totype, chosen here; the larger staminate disk car. Originally these were treated as a section of

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Claoxylon, but Radcliffe-Smithpresented persua- nate or opposite, unlobed or lobed, pinnately or sive arguments for recognitionof a separate genus. palmately veined, sometimes with embedded laminar glands; stipules present or obsolete; inflores- Subtribe 30i. ROTTLERINAE Meisner, P1. cences terminalor axillary, racemose or paniculate; Vasc. Gen. 1: 339. 1841; Webster, Taxon flowers apetalous; staminate sepals 3-5; disk in- 24: 598. 1975. TYPE: Rottlera Roxb. [ Mal- trastaminal, of interstaminal segments, or absent; lotus L.]. stamens 15-300, free; anthers muticous, anthersacs not pendulous; pollen grains 3(-4) colporate, FamilyTrewiaceae Lindley,Intr. Nat. Syst. ed. 2, 174. finelyto coarsely tectate-perforate and spinulose; 1835. Acalypheaesubtribe Mercurialinae series Tre- wiiformesPax & Hoffm.,Pflanzenr. 147. VII (Heft pistillate sepals 3-6(-10), free or connate; disk 63): 139. 1914. TYPE: Trewia L. absent; ovary 2-4(-8)-locular, sometimes echi- TribeAcalypheae subtribe Coelodisceae Muell. Arg.,Lin- nate; styles unlobed, + plumose or laciniate; fruit [ naea 34: 143. 1865. TYPE: CoelodiscusBaillon capsular or baccate; seeds ecarunculate (rarely ca- Mallotus Lour.]. testa often fleshy. TribeMalloteae Hutchinson, Amer. J. Bot. 56: 752. 1969. runculate), TYPE: Mallotus Lour. This Old World subtribe of eight closely related Dioecious (rarely monoecious) trees or shrubs; genera is most stronglyrepresented in tropical Asia. indumentum stellate (rarely simple); leaves alter-

KEY TO THE GENERA OF SUBTRIBE ROTTLERINAE la. Carpels 2-4 (rarely5); fruitcapsular or drupaceous. 2a. Stylesunlobed; leaves usuallyglandular. 3a. Fruitcapsular, dehiscent; pistillate sepals freeor connate. 4a. Ovarysmooth to echinate,not winged;inflorescences terminal or oppositeleaves (sometimes axillaryas well);bracts small; stamens20-200; pollensexine tectate-perforate to rugulose. 5a. Styleselongate; leaves usuallygranulose-glandular ...... 176. Mallotus 5b. Stylescontracted; leaves eglandular...... 177. Deuteromallotus 4b. Ovary ? winged;inflorescences axillary. 6a. Stamensmany (over 200); seeds ecarunculate...... 178. Cordemoya 6b. Stamens 15-20; seeds carunculate...... 179. Coccoceras 3b. Fruitindehiscent; pistillate calyx gamophyllous. 7a. Staminatedisk absent;ovary 2-4-locular ...... 180. Trewia 7b. Staminatedisk present;ovary 1(2)-locular...... 181. Neotrewia 2b. Stylesbifid; leaves eglandular . . . .182. Rockinghamia lb. Ovary 8-locular;fruit baccate; leaves alternate,glandular; stamens 15-20 .183. Octospermum

176. Mallotus Loureiro, Fl. Cochinch. 635. nus trisulcus Lour. [= Mallotus cochinchensis 1790; Muell. Arg., DC. Prodr. 15(2): 956. Lour.]. Rottlera Roxburgh,P1. Coromandel2: 36. 1798 (non 1873; 1866; Bentham,Fl. Austral.6: 138. Willd., 1797). TYPE: Rottlera tinctoriaRoxb. [ Gen. P1. 3: 319. 1880; Forbes & Hemsley, Mallotus philippensis(Lam.) Muell. Arg.]. J. Linn. Soc. Bot. 26: 439. 1894; Schum. & Adisca Blume, Bijdr. Fl. Ned. Ind. 609. 1826. TYPE: Lauterb.,Fl. DeutscheSchutzgeb. 394. 1901; AdiscafloribundaBi. [= Mallotusfloribundus(Bi.) Arg.; lectotype,chosen here]. Meded. Dept. Landb. 10: 394. Muell. J. J. Smith, Plagianthera Reichb.f. & Zollinger,Verh. Natuurk. Ver. 1910; Prami,Fl. Trop. Afr.6(1): 927. 1912; Ned. Ind. 1(4): 19. 1856. TYPE: Plagianthera op- Pax & Hoffm.,Pflanzenr. 147. VII (Heft63): positifolia(Bi.) Reichb.f. & Zoll. [= Mallotus blu- 145. 1914; Natiirl.Pflanzenfam. ed. 2, 19c: meanus Muell. Arg.]. 113. 1931; Merrill& van Steenis, Webbia Hancea Seemann, Bot. Voy. Herald 409. 1857. TYPE: Hancea hookeriana Seem. [= Mallotus hookeri- 8: 405. 1951; Hurusawa,J. Fac. Sci. Univ. anus (Seem.) Muell. Arg.]. Tokyo,Sect. 3, Bot. 6: 304. 1954; AiryShaw, Axenfeldia Baillon, Etude Gen. Euphorb. 419. 1858. Kew Bull. 20: 41. 1866; 21: 379. 1968; 25: TYPE: Axenfeldiaintermedia Baillon [= Mallotus 526. 1971; Whitmore,Tree Fl. Malaya 2: muricatus(Wight) Muell. Arg.]. Etude Gen. Euphorb.293. 1858. 113. 1973; AiryShaw, Kew Bull. Add. Ser. Coelodiscus Baillon, TYPE: Ricinus dioicus Wall. ex Baill. [= Mallotus 4: 160. 1975; 8: 162. 1980; Radcliffe-Smith, eriocarpus(Thw.) Muell. Arg.]. Fl. E. Trop. Afr. Euphorb. 1: 235. 1987. AconceveibumMiquel, Fl. Ind. Bat. 1(2): 389. 1859. TYPE: Mallotus cochinchensis Lour. TYPE: Aconceveibumtrinerve Miq. [= Mallotus philippensis(Lam.) Muell. Arg.]. EchinusLoureiro, Fl. Cochinch.633. 1790. TYPE: Echi- EchinocrotonF. Mueller,Fragm. Phytogeogr.Austral.

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1: 31. 1859. TYPE: Echinocrotonclaoxyloides F. 182. Rockinghamia Airy Shaw, Kew Bull. 20: Muell. [= Mallotus claoxyloides(F. Muell.) Muell. 29. 1966; 35: 667. 1980. TYPE: Rocking- Arg.]. hamia angustifolia (Benth.) Airy Shaw. DiplochlamysMuell. Arg.,Flora 47: 539. 1864. TYPE: Diplochlamysgriffithianus Muell. Arg. [= Mallo- A genus of two species endemic to tropical Aus- tus griffithianus(Muell. Arg.) Hook. f.]. tralia (Queensland). This large and complex paleotropical genus includes ca. 150 species, of which there are only 183. Octospermum Airy Shaw, Kew Bull. 19: two in Africa and Madagascar, the rest in Asia and 311. 1965; Hook. Ic. P1. 38: t. 3716. 1974; Australasia (to Fiji). Kew Bull. Add. Ser. 8: 176. 1980. TYPE: Octospermum pleiogynum (Pax & Hoffm.) 177. Deuteromallotus Pax & Hoffmann, Airy Shaw. Pflanzenr. 147. VII (Heft 63): 212. 1914; A single species of New Guinea. Leandri, Bull. Soc. Bot. France 103: 605. 1956. TYPE: Deuteromallotus acuminatus Subtribe 30j. ACALYPHINAE Grisebach,Fl. (Baillon) Pax & Hoffm. Br. W. Ind. 45. 1859; Pax & Hoffm., Pflan- A genus of two species endemic to Madagascar. zenr. 147. XVI (Heft 85): 1. 1924. TYPE: Acalypha L. 178. Cordemoya Baillon, Adansonia I. 1: 255. Monoecious or dioecious trees, shrubs, or herbs; 1861; Pax & Hoffm., Pflanzenr. 147. VII indumentum simple or stellate, often glandular; (Heft 63): 208. 1914. TYPE: Cordemoya in- leaves alternate, unlobed, pinnately or palmately tegrifolia Willd. ex Baill. [lectotype]. veined, stipulate; inflorescences spicate or the pis- A monotypic genus endemic to Malagasia. tillate (in a few species) paniculate, terminal or axillary, unisexual or bisexual; pistillatebracts usu- 179. Coccoceras Miquel,Fl. Ind. Batav. Suppl. ally enlarged in fruit;staminate sepals 4; disk ab- 455. 1860; Muell. Arg., DC. Prodr. 15(2): sent; stamens usually 8, filamentsfree, anther sacs 949. 1866; Bentham, Gen. PI. 3: 308. 1880; pendulous and vermiform; pollen grains porate, Pax & Hoffm.,Pflanzenr. 147. VII (Heft 63): rugulose; pistillate sepals 3-5, imbricate; disk ab- 209. 1914; Airy Shaw, Kew Bull. 16: 349. sent; ovary (2-)3-locular; styles mostly free, lacin- 1963. TYPE: Coccoceras sumatranus Miq. iate (rarely entire); fruit capsular; seeds with caruncle minute or -absent. A tropical Asian genus of three or four species, distributed from Burma to Borneo. Airy Shaw As here defined, subtribe Acalyphinae includes (1963) reduced Coccoceras to a section of Mal- only the genus Acalypha; it appears closely related lotus, perhaps correctly so. to subtribe Claoxylinae, but differsin its unique anthers, enlarged pistillate bracts, distinctive por- 180. Trewia L., Sp. PI. 1193. 1753; Klotzsch, ate pollen, and dissected styles. Arch. Naturgesch. 7: 255. 1841; Muell. Arg., DC. Prodr. 15(2): 953. 1866; Bentham, Gen. 184. Acalypha L., Sp. P1. 1003. 1753; Gen. PI. 3: 318. 1880; J. J. Smith, Meded. Dept. P1. ed. 5, 436. 1754; Muell. Arg., DC. Prodr. Landb. 10: 389. 1910; Pax & Hoffm.,Pflan- 15(2): 799. 1866; Bentham, Gen. P1. 3: 31 1. zenr. 147. VII (Heft 63): 140. 1914; Airy 1880; Hutchinson, Fl. Trop. Afr. 6(1): 880. Shaw, Kew Bull. 20: 406. 1966; Kew Bull. 1912; Pax & Hoffm., Pflanzenr. 147. XVI Add. Ser. 4: 200. 1975. TYPE: Trewia nu- (Heft 85): 12. 1924; Natiurl.Pflanzenfam. ed. difiora L. 2, 19c: 134. 1931; Hurusawa, J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: 295. 1954; A monotypic genus, the single species widely Webster, J. Arnold Arbor. 48: 370. 1967; A. distributed from India to southern China and In- C. Smith, Fl. Vit. Nova 2: 522. 1981; Seberg, donesia. Nordic J. Bot. 4: 159. 1984; Radcliffe-Smith, Fl. E. Trop. Afr. Euphorb. 1: 185. 1987. 181. Neotrewia Pax & Hoffmann, Pflanzenr. TYPE: Acalypha virginica L. [lectotype, des- 147. VII (Heft 63): 211. 1914; Merrill,Enum. ignated by Small, in Britton & Brown, Ill. Fl. Phil. P1. 2: 437. 1923. TYPE: Neotrewia cum- N. U.S. ed. 2, 2: 457. 1913]. ingii (Muell. Arg.) Pax & Hoffm. Caturus L., Syst. Nat. ed. 12, 2: 650. 1767; Mant. P1. A monotypic genus of the Philippines and Cele- 19, 127. 1767. TYPE: Caturus spiciflora L. [ bes, dubiously separable from Trewia. Acalypha hispida Burm.f.].

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LinostachysKlotzsch ex Schlect.,Linnaea 19: 235. 1846. 185. Lasiococca Hooker f., Hook. Ic. P1. 16: TYPE: Linostachyspadifolia Schlecht.[= Acalypha t. 1587. 1887; Fl. Brit. Ind. 5: 456. 1887; schlechtendalianaMuell. Arg.]. Inf. 70. OdonteilemaTurczaninov, Bull. Soc. Imp. Natur.Moscou Haines,Kew Bull.Misc. 1920; 1920; 21: 587. 1848. TYPE: Odonteilemaclaussenji Turcz. Airy Shaw, Kew Bull. 16: 358. 1963; 21: [= Acalypha claussenji (Turez.) Muell. Arg.]. 406. 1968; Whitmore,Tree Fl. Malaya 2: GymnalyphaGrisebach, Bonplandia 6: 2. 1858. TYPE: 104. 1973; Thin,J. Biol. Sinh Hoc 8(3): 36. Gymnalypha Griseb.,nom. illeg.[= Aca- jacquinii 1986. TYPE: Lasiococca symphyllifolia(Kurz) lypha villosa Jacq.]. CorytheaS. Watson, Proc. Amer.Acad. Sci. 22: 451. Hook. f. 1887. TYPE: S. Wats. [= Acalypha Corytheafihipes A tropicalAsian genus of threespecies, with a filipes(S. Watson) McVaugh]. AcalyphopsisPax & Hoffm.,Pflanzenr. 147. XVI (Heft disjunctdistribution in India, Malaya and Indo- 85): 178. 1924. TYPE: Acalyphopsis celebica Pax china, and Hainan. & Hoffm.[= Acalypha hoffmannianaHurus.].

With ca. 450 species, Acalypha is the fourth 186. Spathiostemon Blume,Bijdr. Fl. Ned. Ind. largest genus of Euphorbiaceae; in addition to the 621. 1826; AiryShaw, Kew Bull. 16: 357. small number of temperate species, species are 1963; 20: 408. 1966; 26: 341. 1972; Hook. widespread throughoutthe tropics except in Hawaii Ic. P1. 38: t. 3720. 1974; Kew Bull. Add. 4: TYPE: and a few Pacific archipelagos. Ser. 196. 1975; 8: 202. 1980. Spa- thiostemon javense Bl. Subtribe 30k. LASIOCOCCINAE Webster, Polydragma Hook. f., Hook. Ic. P1. 18: t. 1701. 1887; Taxon 24: 597. 1975. TYPE: Lasiococca Fl. Brit.Ind. 5: 456. 1887. TYPE: Polydragma Hook. f. mallotiformeHook. f. [= Spathiostemonjavense BI.]. Monoecious or dioecious trees or shrubs; indu- ClonostylisSp. Moore, J. Bot. 63 (suppl.): 101. 1925. mentum simple or lepidote; leaves alternate, un- TYPE: Clonostylisforbesii Sp. Moore [= Spathios- temonforbesii (Sp. Moore) AiryShaw]. lobed, pinnately veined; eglandular, stipulate; inflorescences axillary, unisexual, racemose or the A genusof two species distributed from Thailand pistillate flowers solitary, bracts eglandular; sta- to thePhilippines and NewGuinea. Although united minate sepals usually 3; disk absent; stamens many, with Homonoia by Mueller (1866), Bentham filamentsconnate, anthers on filamentsof branch- (1880), and Pax & Hoffmann(1919), it may be ing fascicles; pollen grains 3-colporate; not oper- distinct,as indicatedby AiryShaw. culate; pistillatesepals 5-8, imbricate; disk absent; ovary 3-locular; styles unlobed, smooth to papillose 187. Homonoia Loureiro,Fl. Cochinch.636. or plumose; fruitcapsular; seeds ecarunculate, tes- 1790; Muell. Arg., DC. Prodr. 15(2): 1022. ta usually fleshy. 1866; Bentham,Gen. P1. 3: 322. 1880; J. J. Smith,Med. Dept. Landb. 10: 542. 1910; This subtribe includes three genera that were Pax & Hoffm.,Pflanzenr. 147. XI (Heft68): treated by Pax & Hoffmann (1919) as part of 114. 1919; Gagnepain,Fl. Indochine5: 330. subtribe Ricininae, doubtlessly because of the an- 1925; AiryShaw, Kew Bull.Add. Ser. 4: 136. droecium of many stamens with anthers borne on 1975; Kew Bull. 36: 310. 1981. TYPE: Homo- branches of a dendritic column. However, this noia riparia Lour. character almost surely represents a convergence, since the Lasiococcinae diverge from Ricinus in Lumanaja Blanco,Fl. Filip.821. 1837. TYPE: Lumanaja many importantcharacters, includingecarunculate fluviatilisBlanco [= Homonoia riparia Lour.]. seeds, pinnatelyveined unlobed leaves, and axillary A genus of twoAsian species (India to Taiwan, inflorescences. Airy Shaw (1974) has suggested a the Philippines,and New Guinea). close relationshipof subtribe Lasiococcinae to Mal- lotus. Tribe 3 1. PLUKENETIEAE (Bentham) Hutchinson,Amer. J. Bot. 56: 753. 1969. OF KEY TO THE GENERA SUBTRTBE LASIOCOCCINAE Crotoneae subtribe Plukenetieae Bentham, la. Monoecious; pistillatesepals persistent;indu- Gen. P1. 3: 252. 1880. TYPE: Plukenetia L. mentumsimple; ovary muricate -- 185. Lasiococca lb. Dioecious;pistillate sepals deciduous;indumen- Monoecious(rarely dioecious) trees, shrubs, or tumsimple or lepidote. herbs,sometimes climbing or twining;indumentum 2a. Lepidotescales absent;ovary tuberculate; often sometimes leaves pollensexine not striate 186. Spathiostemon simple, glandular, urticant; 2b. Lepidotescales present;ovary smooth; pol- alternate,unlobed to palmatelyparted, pinnately len sexinestriate ...... 187. Homonoia to palmatelyveined, sometimesglandular or sti-

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pellate at base; stipules present. Inflorescences ter- mann (1919, 1931). In contrast to subtribe Tra- minal or axillary, usually bisexual, mostlyracemose giinae, the distributionof the Plukenetieae is almost or spicate; bracts eglandular; flowers apetalous. entirely neotropical (a few species of Plukenetia Staminate calyx splitting into 3-5 valvate seg- occur in Africa). ments; disk interstaminal,extrastaminal, or absent; stamens (2)3-100, usually free; anthers introrse KEY TO THE GENERA OF SUBTRIBE PLUKENETIINAE or extrorse, muticous or apiculate; pollen grains la. Ovary 3-locular;pistillate sepals 5 or 6; trees, 3-colporate, porate, or inaperturate,sexine varying shrubsor vines. fromtectate to intectate; pistillodeabsent. Pistillate 2a. Stamens4; staminatedisk segments 4 (or sepals 3-6, imbricate; disk absent; ovary 3- or if absentreplaced by 4-lobedpseudodisk); leaves pinnatelyveined; trees or shrubs. 4-locular; styles unlobed, slender or dilated, some-, 3a. Disk absent;base of stamensdilated, times papillose or lacerate, usually connate into a formingan intrastaminalpseudodisk; distinct column. Fruit capsular; columella persis- styleurceolate 188. Haematostemon tent; seeds ecarunculate, testa usually dry; endo- 3b. Disk extrastaminal;style ovoid. 189. sperm copious; cotyledons much longer and broad- -...... Astrococcus 2b. Stamens 10 or more; staminatedisk seg- er than radicle. mentsabsent or minute,numerous. 4a. Stamensca. 20; styleurceolate; flow- This distinctive tribe includes 16 genera with ers fasciculatein leafaxils; leaves pin- more than 250 species in the tropics of both the natelyveined; tree or shrub. Old World and New World; the majority are neo- -...... 190. Angostyles tropical. 4b. Stamens10; stylecylindrical, clavate, apically3-lobed; flowers in axillaryra- cemes; leaves palmatelyveined; twin- KEY TO THE SUBTRIBESOF TRIBE PLUKENETIEAE ing vine 191. Romanoa la. Flowersin racemes or spikes,not bibracteate lb. Ovary4-locular; pistillate sepals 4; twiningvines pseudanthia;pollen grains subglobose. or lianas. 2a. Stinginghairs absent; fruit winged, crested, 5a. Stylesmostly free, connate for less than carinate, or indehiscent;pollen grains one-halftheir length,tips bifid;stamens 3-colpate . 3 la. Plukenetiinae 40-50 192. Eleutherostigma 2b. Stinginghairs present; fruit usually unap- 5b. Stylesentirely connate into a massivecol- pendaged;pollen grains 3-colpate, 3-porate, umn;stamens 8-30(-40). or inaperturate,rugulose 31 b. Tragiinae 6a. Fruitdehiscent (rarely indehiscent, but lb. Flowersin bibracteatepseudanthia; pollen grains thenstaminate receptacle not globose) prolate,coarsely reticulate; stinging hairs pres- -...... 193. Plukenetia ent 31c. Dalechampiinae 6b. Fruitindehiscent; staminate receptacle globose 194. Vigia

Subtribe 3 la. PLUKENETIINAE Bentham, 188. Haematostemon (Muell. Arg.) Pax & Gen. P1. 3: 253. 1880; Pax, Naturl. Pflan- Hoffmann,Pflanzenr. 147. IX (Heft 68): 31. zenfam. ed. 1, 3(5): 62. 1890; Pax & Hoffm., 1919; Sandwith, Kew Bull. 1950: 133. 1951; Pflanzenr. 147. IX (Heft 68): 1. 1919; Na- Jablonski, Mem. New York Bot. Gard. 17: turl. Pflanzenfam. ed. 2, 19c: 141. 1931; 143. 1967. Astrococcus sect. Haematoste- Webster, Taxon 24: 598. 1975. TYPE: Plu- mon Muell. Arg., Linnaea 34: 157. 1865. kenetia L. TYPE: Haematostemon coriaceus (Baill.) Pax & Hoffm. Trees, shrubs, lianas, or twining vines; indumentum entirelyof simple hairs; leaf blades simple A genus of two species fromrainforests in south- and unlobed, entire or dentate, often with embed- ern Venezuela and Guiana. The gynoecium is very ded laminar glands, sometimes stipellate; inflores- similar to that of Angostyles, but the staminate cence axillary or terminal and appearing leaf-op- flowers are quite different. posed; stamens 4-50, filaments free or anthers subsessile; pollen grains suboblate to subglobose, 189. Astrococcus Bentham, Hook. J. Bot. Kew tricolpate, tectate-perforate to semitectate-reticu- Gard. Misc. 6: 327. 1854; Muell. Arg., DC. late; ovary 3- or 4-locular, often winged; styles Prodr. 15(2): 766. 1866; Fl. Bras. 11(2): massive, partly to entirely connate; fruitcapsular 330. 1874; Bentham, Gen. P1. 3: 326. 1880; or rarely indehiscent. Pax & Hoffm., Pflanzenr. 147. IX (Heft 68): 30. 1919. TYPE: Astrococcus cornutus Benth. As delimited by Webster (1975), subtribe Plu- kenetiinae is here treated in a narrower sense than A monotypic genus from Amazonian Brazil and the concepts of Bentham (1880) or Pax & Hoff- Venezuela.

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190. Angostyles Bentham,Hook. J. Bot. Kew (Muell. Arg.)Pax & Hoffm.[= Plukenetiaconoph- Gard. Misc. 6: 328. 1854; Muell. Arg., DC. ora Muell. Arg.]. Prodr. 15(2): 767. 1866; Fl. Bras. 11(2): Apodandra Pax & Hoffm.,Pflanzenr. 147. IX (Heft68): 20. 1919. TYPE: Apodandra loretensis(Ule) Pax 331, t. 50. 1874; Bentham,Gen. P1. 3: 327. & Hoffm.[= Plukenetia loretensisUle]. 1880; Pax & Hoffm.,Pflanzenr. 147. IX Elaeophora Ducke, Ann. Jard. Bot. Rio Jan. 4: 112. (Heft 68): 29. 1919. TYPE: Angostyles lon- 1925. TYPE: Elaeophora abutaefoliaDucke [= Plu- gifolia Benth. kenetia abutiJclia(Ducke) Pax & Hoffm.]. A monotypicgenus from the BrazilianAmazon. A genus of about 15 species, mostlyneotropical, but withthree species in Africa, one in Madagascar, 191. Romanoa Trevisan,Saggio Monogr. Alghe and one in southeast Asia. The genus is rather Cocc. 99. 1848 (nom.cons.); Radcliffe-Smith,variable, and a number of workers have accepted Kew Bull. 34: 589. 1980. Anabaena A. Juss., either Pterococcus or Tetracarpidium as distinct Euphorb.Tent. 46. 1824. Anabaenella Pax genera. However, the present conservative concept & Hoffm.,Pflanzenr. 147. IX (Heft68): 27. may not be broad enough, since some of the other 1919. TYPE: Anabaena tamnoides A. Juss. genera of subtribe Plukenetiinae listed here (e.g., [= Romanoa tamnoides (A. Juss.) Radcl.- Vigia and Romanoa) are only questionably distinct Sm.]. from Plukenetia. A monotypicgenus of Brazil,also veryclose to Plukenetia and questionablydistinct. 194. Vigia Vellozo, Fl. Flum. 9: t. 127. 1832. TYPE: Vigia serrata Vell. 192. Eleutherostigma Pax & Hoffmann, FragariopsisSt. Hilaire,Le~ons Morph.Veg. 426. 1840. Pflanzenr.147. IX (Heft68): 11. 1919. TYPE: TYPE: Fragariopsis scandens St. Hil. [= Vigia ser- Eleutherostigma lehmannianum Pax & rata Vell.]. Hoffm. Accia St. Hilaire,Le~ons Morph.Veg. 499. 1840. TYPE: Accia scandens St. Hil. [= Vigia serrata Vell.]. A poorlyknown monotypic genus of Colombia BotryantheKlotzsch, Arch. Naturgesch. 7(1): 190. 1841. and Ecuador, apparentlyclosely related to Plu- TYPE: Botryanthediscolor K. [= Vigia serrata kenetia,and combinedwith it by Gillespie(1993). Vell.]. A South American genus, probably monotypic, 193. Plukenetia L., Sp. P1. 1192. 1753; Gen. although Pax & Hoffmann (1931) recognized a P1. ed. 5, 438. 1754; Muell.Arg., DC. Prodr. second species, Fragariopsis paxii Pitt. Even 15(2): 768. 1866; Bentham,Gen. P1. 3: 327. though the name Fragariopsis has been univer- 1880; Pax & Hoffm.,Pflanzenr. 147. IX sally used for this segregate fromPlukenetia since (Heft 68): 12. 1919; Jablonski,Mem. New the treatment of Baillon (1858), the name Vigia York Bot. Gard. 17: 142. 1967; Webster, of Vellozo clearly has priority.Although the plate Ann. Missouri Bot. Gard. 54: 293. 1968; of Vellozo is crude, it represents publication as Huft, Ann. MissouriBot. Gard. 75: 1105. valid as the casual mention of Fragariopsis by St. 1989; Gillespie,Syst. Bot. 18: 575. 1993. Hilaire in his textbook. There seems no good reason TYPE: Plukenetia volubilis L. to propose the name Fragariopsis for conserva- PterococcusHasskarl, Flora 25(2), Beibl. 41. 1842(nom. tion, especially since it is only dubiously distinct cons.); Croizat,J. ArnoldArbor. 22: 423. 1941; from Plukenetia. Airy Shaw, Kew Bull. Add. Ser. 4: 187. 1975. CeratococcusMeissner, P1. Vasc. Gen. 2: 369. 1843. Sajorium Endl.,Gen. P1. Suppl. 3: 98. 1843. TYPE: Subtribe 31b. TRAGIINAE Webster,Taxon Pterococcusglaberrimus Hassk., nom. illeg. [= Plu- 24: 598. 1975. TYPE: Tragia L. kenetia corniculataJ. E. Smith]. HedraiostylusHasskarl, Tijdschr. Natuurl. Gesch. Phys- Shrubs or herbs, often twining; indumentum of iol. 10: 141. 1843. TYPE: Hedraiostylusglaber- simple and stinginghairs; leaves simple to distinctly rimus(Hassk.) Hassk. [= Plukenetia corniculata lobed, entire or dentate, withoutembedded laminar J. E. Smith]. TetracarpidiumPax, Bot. Jahrb.26: 329. 1899. TYPE: glands, not stipellate; inflorescencesaxillary or ter- Tetracarpidiumstaudtii Pax [= Plukenetia co- minal and opposite leaves; stamens (2)3-50, fila- nophora Muell. Arg.]. ments free or connate; pollen grains subglobose to Pseudotragia Pax, Bull. Herb. Boiss. II. 8: 635. 1908. suboblate, 3-colpate, 3-porate, or inaperturate, TYPE: Pseudotragia scandens Pax [= Plukenetia corniculataJ. E. Sm.]. sexine intectate-baculate, semitectate-reticulate,or AngostylidiumPax & Hoffm.,Pflanzenr. 147. IX (Heft tectate-perforate;ovary 3-locular; stylesnearly free 68): 17. 1919. TYPE: Angostylidiumconophorum to distinctlyconnate; fruitcapsular.

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As here delimited,the subtribeincludes 8 genera Nair, Gardener'sBull. 31: 49. 1978. TYPE: with ca. 150 species; besides the widespread Tra- Cnesmone javanica Bi. gia, there are 5 Old World and only 2 neotropical CenesmonGagnep., Bull. Soc. Bot.France 71: 866. 1924; genera. Fl. Gen.Indochine 5: 386. 1926. TYPE: Cenesmon tonkinense Gagnep. [= Cnesmone tonkinense KEY TO THE GENERA OF SUBTRIBE TRAGIINAE (Gagnep.)Croizat; lectotype, designated by Wheeler, Taxon24: 534. 1975]. la. Staminatesepals adaxiallyinflexed to forma pseudodisk;stamens 3 or 4, anthersintrorse; This Asian genus of ca. ten species is here de- stylesmassive. limitedfollowing Croizat (1941). 2a. Pistillatesepals entireor toothed;pistillode absentin staminateflower. 3a. Pistillatesepals entire; anther connec- 196. Megistostigma Hooker f., Hook. Ic. P1. tive enlarged,triangular, sometimes 16: t. 1592. 1887; Fl. Brit. Ind. 5: 466. caudate; stamensfree. 1888; Merrill,Phil. J. Sci. 16C: 563. 1920; 4a. Stylesfree or nearlyso, long-pap- illoseon adaxialsurface; inflores- Croizat,J. ArnoldArbor. 22: 425. 1941; Airy cence terminaland leaf-opposed; Shaw, Kew Bull. 23: 119. 1969. TYPE: leaves withstinging hairs Megistostigma malaccense Hook. f. 195. Cnesmone 4b. Stylesconnate into a massiveglo- ClavistylusJ. J. Smith,Med. Dept. Landb.10: 517. bose or clavatecolumn, not papil- 1910. TYPE: Clavistyluspeltatus J.J. Sm. [= Me- lose; inflorescenceaxillary; leaves gistostigmapeltatum (J. J. Sm.) Croizat]. subglabrous 196. Megistostigma 3b. Pistillatesepals toothed;anther con- A tropicalsoutheast Asian genusof fivespecies nectivenot enlargedor caudate; sta- distributedfrom Burma to China, the Philippines, mensconnate at base; stylesconnate, and Borneo; combinedwith Sphaerostylis by Pax columnglobose; inflorescencetermi- & Hoffmann(1919) but maintainedas distinctby nal and leaf-opposed;leaves subglabrous 197. Sphaerostylis Croizat(1941). 2b. Pistillatesepals distinctlypinnatifid; pistillode presentin staminateflower; leaves 197. Sphaerostylis Baillon, Etude Gen. Eu- withstinging hairs 198. Tragiella phorb. 466. 1858; Muell. Arg., DC. Prodr. lb. Staminatesepals not adaxially inflexed; stamens 15(2): 768. 1866; Bentham,Gen. P1. 3: 327. 2 to many,anthers introrse or extrorse;styles usuallyslender. 1880; Baillon,Hist. Phys. Nat. Madag. 4(29): 5a. Anthers3 or more,if 2 thennot subsessile; t. 196. 1891; Pax & Hoffm.,Pflanzenr. 147. staminatecalyx concave; styles free to IX (Heft 68): 106. 1919; Croizat,J. Arnold connate. partly Arbor.22: 430. 1941. TYPE: Sphaerostylis 6a. Anther connective withouttuft of stinginghairs; stamens (2)3-many; tulasneana Baill. subshrubs,herbs, or twiningvines, A genusof twospecies endemicto Madagascar. usuallymonoecious. 7a. Styles thick,papillose; staminal receptacle convex; stamens 4- 198. Tragiella Pax & Hoffmann,Pflanzenr. 18 199. Platygyna 147. IX (Heft 68): 104. 1919; Radcliffe- 7b. Stylesslender, not papillose;sta- Smith,Kew Bull. 35: 777. 1981; Fl. E. Trop. minalreceptacle plane; stamens 1: 318. 1987. TYPE: Tragiella 2-8(-50) 200. Tragia Afr.Euphorb. 6b. Antherconnective ending in tuftof natalensis (Sond.) Pax & Hoffm.[lectotype, stinginghairs; erect shrubs, leaves gla- chosen here]. brescent,dioecious 201. Acidoton 5b. Anthers2, subsessile;staminate calyx flat An Africangenus of five species, extremely close withlobes reflexed;styles connate into a to Sphaerostylisand doubtfullyseparable. thickcolumn 202. Pachystylidium 199. Platygyna Mercier,Bull. Bot. (Seringe)1: 195. Cnesmone Blume,Bijdr. Fl. Ned. Ind. 630. 168. 1830; Muell. Arg., DC. Prodr. 15(2): 1826 (as Cnesmosa);Fl. Javaevi. 1828; Muell. 913. 1866 (as Platygyne); Bentham,Gen. Arg.,DC. Prodr.15(2): 926. 1866; Bentham, P1. 3: 328. 1880; Pax & Hoffm.,Pflanzenr. Gen. P1. 3: 330. 1880; Hook. f.,Fl. Brit.Ind. 147. IX (Heft68): 26. 1919; Alain,Fl. Cuba 5: 466. 1888; J. J. Smith,Med. Dept. Land- 3: 100. 1953; Mem. New York Bot. Gard. bouw 10: 513. 1910; Pax & Hoffm.,Pflan- 21: 132. 1971; Borhidi,Ann. Hist.-Nat. Mus. zenr. 147. IX (Heft68): 102. 1919; Croizat, Nat. Hung. 64: 89. 1972. TYPE: Platygyna J. ArnoldArbor. 22: 427. 1941; AiryShaw, urens Merc. [= Platygyna hexandra (Jacq.) Kew Bull. 26: 240. 1972; Balakrisnan& Muell. Arg.].

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AcanthocaulonKlotzsch ex Endl., Gen. P1. Suppl. 4(3): Pflanzenr. 147. IX (Heft 68): 24. 1919; Fawc. 88. 1850. TYPE: Acanthocaulonpruriens KI. ex & Rend., Fl. Jam. 4: 303. 1920; Webster, Endl. [= Platygyna hexandra(Jacq.) Muell. Arg.]. Ann. Missouri Bot. Gard. 54: 191. 1967; 54: In the most recent revision of Borhidi (1972), 289. 1968; Liogier, Fl. Espaiiola 4: 71. 1986. Platygyna is a genus of seven species endemic to TYPE: Acidoton urens Sw. Cuba. Alain (1971) reduced Platygyna to a syn- Gitara Pax & Hoffm.,Pflanzenr. 147. XVII (Heft 85): onym of Tragia, a disposition that merits further 187. 1924. TYPE: Gitara venezolana Pax & Hoffrn. study. [= Acidoton nicaraguensis(Hemsl.) Webster].

A neotropical genus of six or seven species, in 200. Tragia Plumier ex L., Sp. P1. 980. 1753; the Greater Antilles (Jamaica and Hispaniola), Gen. P1. ed. 5, 421. 1754; Muell. Arg., DC. southern Central America, and northern South Prodr. 15(2): 927. 1866; Bentham, Gen. P1. America. 3: 329. 1880; Prain, Fl. Trop. Afr. 6(1): 964. 1913; Pax & Hoffm., Pflanzenr. 147. IX (Heft 68): 32. 1919; Prain, Fl. Capensis 5(2): 202. Pachystylidium Pax & Hoffmann,Pflan 502. 1920; Lourteig & O'Donell, Lilloa 6: zenr. 147. IX (Heft 68): 108. 1919; Backer 347. 1941; Johnston,Rhodora 64: 137. 1962; & Bakh., Fl. Java 1: 491. 1963; Airy Shaw, Miller & Webster, Rhodora 69: 241. 1967; Kew Bull. 23: 115. 1969. TYPE: Pachystylid- Webster, J. Arnold Arbor. 48: 376. 1967; ium hirsutum (Bl.) Pax & Hoffm. [Tragia Leandri, Adansonia II. 11: 437. 1971; Rad- hirsuta Bl.]. cliffe-Smith,Fl. E. Trop. Afr. Euphorb. 1: A monotypic genus of tropical Asia (India to 291. 1987; Webster & Huft, Ann. Missouri Indochina, Java, and the Philippines; not reported Bot. Gard. 75: 1106. 1989. TYPE: Tragia from Sumatra or Borneo). volubilis L. [lectotype, designated by Small, in Britton & Brown, Ill. Fl. N. U.S., ed. 2, Subtribe 31c. DALECHAMPIINAE (Muell. 2: 458. 1913]. Arg.) Webster, stat. nov. Tribe Dalecham- pieae Muell. Arg., Bot. Zeit. 22: 324. 1864; Bia Klotzsch,Arch. Naturgesch. 7(1): 189. 1841. TYPE: DC. Prodr. 15(2): 1232. 1866. TYPE: Dale- Bia alienata Didr.[= Tragia alienata (Didr.)Mulg. & M. Guttierrez,Candollea 46: 523. 1991; this champia Plum. ex L. nameunfortunately must replace Tragia sellowiana Muell. Arg., 1866; lectotype]. Monoecious undershrubs or more often clam- Leptorhachis Klotzsch,Arch. Naturgesch.7(1): 189. bering or twining;indumentum of simple and sting- 1841. TYPE: Leptorhachishastata KI. [= Tragia ing hairs (sometimes almost absent); leaves alter- hastata (Kl.) Muell. Arg.]. nate, blades simple, lobed or partite, usually Leucandra Klotzsch,Arch. Naturgesch. 7(1): 188. 1841. stipellate; stipules persistent; inflorescences pseu- TYPE: Leucandra betonicaefoliaKI. [= Tragia leucandra Pax & Hoffm.]. danthial, subtended by a pair of usually showy CtenomeriaHarvey, Hook. LondonJ. Bot. 1: 29. 1842. involucral bracts; pistillate flowers 3 per pseudan- TYPE: Ctenomeriacordata Harv. [= Tragia capen- thium, in a basal cymule; staminate pleiochasium sis Thunb.]. terminal, with mostly 8-12 flowers; staminate Lassia Baillon,Etude Gen. Euphorb.464. 1858. TYPE: bractlets mostly stamens 8-100 or Lassia scandens Baill. [= Tragia scandens (Baill.) resiniferous; Muell. Arg.]. more, filaments connate; pollen grains prolate, ZuckertiaBaillon, Etude Gen. Euphorb.495. 1858. TYPE: 3-colporate, sexine with prominent raised reticu- Zuckertia cordata Baill. [= Tragia bailloniana lum; pistillodeabsent; pistillatesepals entire to pin.- Muell. Arg.]. natifid; ovary 3-locular; stylar column elongated, stigmaticsurface extending down upper 1/3 to 34; A large and diverse genus of ca. 125 species, fruitcapsular. mainly in Africa and America. The genus is pal- ynologically heterogeneous, and some taxa such as This subtribe contains only the single genus Dal Zuckertia may prove to merit segregation on fur- echampia. Although it has been treated at tribal ther study. rank by almost all workers since Mueller (1866), Dalechampia has links to both the Tragiinae (stinging hairs) and Plukenetiinae col 201. Acidoton Swartz, Prodr.6,83.1788 (nom. (elongate stylar umn). cons.); Fl. Ind. Occ. 2: 952, t. 18. 1800; Muell. Arg., DC. Prodr. 15(2): 914. 1866; Bentham, Gen. P1. 3: 328. 1880; Urban, 203. Dalechampia Plumier ex L., Sp. P1. 1054. Symb. Ant. 7: 513. 1913; Pax & Hoffm., 1753; Gen. P1. ed. 5,473. 1754; Muell, Arg.,

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DC. Prodr. 15(2): 1232. 1866; Fl. Bras. 11(2): whose affinitieshave been controversial for many 633. 1874; Bentham, Gen. P1. 3: 330. 1880; years. Early workers, such as A. Jussieu (1824) Pax & Hoffm.,Pflanzenr. 147. XII (Heft 68): and Baillon (1858), associated it with the Hippo- 3. 1919; Leonard, Fl. Congo Rwanda-Burundi maneae. Mueller (1866) kept it in the tribe Hip- 8(1): 194. 1962; Jablonski, Mem. New York pomaneae, in subtribeGelonieae. Pax & Hoffmann Bot. Gard. 17: 145. 1967; Webster, Ann. (1912) created a special subtribe of Hippomaneae Missouri Bot. Gard. 54: 308. 1968; Webster for Omphalea, but later (1931) abandoned their & Armbruster,Brittonia 31: 352. 1979; Syst. subtribe and inserted the genus in the Gelonieae. Bot. 7: 484. 1982; Armbruster,Syst. Bot. 9: Croizat (1942), in contrast, pointed out affinities 272. 1984; Radcliffe-Smith,Fl. E. Trop. Afr. withthe tribe Plukenetieae, and Punt (1962) noted Euphorb. 1: 285. 1987; Armbruster, Syst. that pollen morphology supports Croizat's opinion. Bot. 13: 303. 1988; Webster, Ann. Missouri Although Omphalea shows striking resem- Bot. Gard. 75: 1108. 1988; Armbruster,Brit- blances to the Plukenetieae in a number of char- tonia 41: 44. 1989; Webster, Brittonia 41: acters, it also approaches the tribe Stomatocaly- 1. 1989; Webster & Armbruster,Bot. J. Linn. ceae, and thus provides a link between subfamilies Soc. 105: 137. 1991. TYPE: Dalechampia Acalyphoideae and Euphorbioideae. Clarificationof scandens L. its affinitiesappears to be critical in establishing the phylogeny of the uniovulate Euphorbiaceae. CremophyllumScheidweiler, Bull. Acad. Roy. Sci. Brux. 9(1): 23. 1842. TYPE: Cremophyllumspathulatum spathulata (Scheidw.) Scheidw. [= Dalechampia Nat. ed. 10, 1264. Baillon]. 204. Omphalea L., Syst. Megalostylis Spencer Moore, J. Bot. 54: 208. 1916. 1759 (nom. cons.); Muell. Arg., DC. Prodr. TYPE: Megalostylispoeppigii Sp. Moore [= Dal- 15(2): 1134. 1866; Bentham, Gen. PI. 3: echampia micranthaPoepp.]. 332. 1880; Pax & Hoffm., Pflanzenr. 147. V (Heft 52): 14. 1912; Fawc. & Rend., Fl. A genus of slightlyover 100 species, about 90 4: 319. 1920; Croizat, Bull. Jard. Bot. of which are American, with a few in Africa, Mad- Jam. Buit. III. 17: 204. 1941; Alain, Fl. Cuba 3: agascar, and India, and a single species reaching Webster, Ann. Missouri Bot. Gard. southern China and Java. The genus is diverse, 109. 1953; Airy Shaw, Kew Bull. 23: and Webster & Armbruster (1991) have recog- 54: 295. 1968; Mem. New York Bot. Gard. nized 6 sections for the 90-odd neotropical species. 130. 1969; Alain, 21(2): 124. 1971; Airy Shaw, Kew Bull. Add. Ser. 8: 177. 1980; Liogier, Fl. Espafiola 4: Tribe 32. OMPHALEAE (Pax & Hoffmann) 178. 1986; Radcliffe-Smith,Fl. E. Trop. Afr. Webster, Taxon 24: 598. 1975. Hippoma- Euphorb. 1: 371. 1987; Gillespie, Rev. Phyl. neae subtribe Omphaleinae Pax & Hoffm., Anal. Omphalea 144 (dissert.). 1988. Om- Pflanzenr. 147. IV (Heft 52): 14. 1912. TYPE: phalandria P. Browne, Civ. Nat. Hist. Ja- Omphalea L. maica 334. 1756 (nom. rej.). Duchola Adan- Monoecious trees, shrubs, or lianas; stems with son, Fam. PI. 2: 357. 1763 (nom. superfi.). reddish latex; indumentumsimple; leaves alternate, TYPE: Omphalea triandra L. (typ. cons.). simple or lobed, pinnately or palmately veined, usually with 2 sessile basal glands; stipules decid- Ronnowia Buchoz, P1. Nouv. Decouv. 6, t. 4. 1779. uous. Inflorescences terminal or axillary, panicu- TYPE: Ronnowia domingensisBuchoz (nom. illeg.) [= Omphalea triandraL.]. late, bracts ? foliose and biglandular; flowersapet- Hecatea Thouars,Hist. P1. Iles AfriqueAustr. 27, t. 5. or alous. Staminate sepals 4 or 5, decussate 1804. TYPE: Hecatea oppositifoliaWilld. [= Om- imbricate; disk annular or lobed, often massive; phalea oppositifolia(Willd.) Gillespie;lectotype, stamens 2 or 3, filaments connate, anther con- designatedby Gillespie]. Kongl.Vetensk. Akad. Handl. 1854: nectives free or fused; pollen grains oblate, Hebecocca Beurling, 146. 1856. TYPE: Hebecocca panamensis Beurl. pis- 3-colporate, sexine finely punctate-spinulose; [= Omphalea diandra L.]. tillode absent. Pistillate sepals 4 or 5, imbricate; Neomphalea Pax & Hoffm.,Pflanzenr. 147. XIV (Heft disk tenuous or obsolete; ovary 3-locular; styles 68): 54. 1919. TYPE: Neomphalea papuana (Pax connate into a stout column, stigmas undivided. & Hoffm.)Pax & Hoffm.[= Omphalea papuana Pax & Hoffm.]. Fruit thick-walled, capsular or indehiscent; seeds large, globose, ecarunculate, testa dry; endosperm A genus of ca. 15 species in both the Old World copious. and New World, but with the greatest species con- This tribe contains the single genus Omphalea, centrationsin the Greater Antillesand Madagascar.

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Subfamily IV. CROTONOIDEAE Pax, Bot. much narrower circumscription than that of Pax Jahrb.5: 413. 1884; Natiirl. Pflanzenfam. (1890), which took in nearly all of the uniovulate ed. 1, 3(5): 14. 1890. TYPE: CrotonL. taxa of Euphorbiaceae. It is considerably broader than the subfamily Crotonoideae of Hurusawa or herbs;latex coloredor absent; Trees, shrubs, (1954), which included only the tribe Crotoneae. or leaves indumentumsimple, stellate, lepidote; In the classic system of Mueller (1866), genera or rarelywhorled, stipulate or alternate,opposite, of the presently accepted Crotonoideae were dis- simpleor palmatelylobed or com- exstipulate;blade persed between tribes Crotoneae, Acalypheae, and oftenwith foliar glands. Inflorescences ax- pound, Hippomaneae. Bentham (1880) placed many of or terminal,dichasial to racemoseor spicate. illary the Crotonoideae in the firsttwo subtribes of his sepals imbricateor valvate; pet- Staminateflower: tribe Crotoneae, but in other subtribes had a mix- or absent; stamens(3-)5- als and disk present ture of Acalyphoideae and Crotonoideae. Hutch- freeor connate;pollen grains bi- many,filaments inson (1969) placed many Crotonoideae in his tribes tricolporateto more com- nucleateor trinucleate, 13-20, 38, and 39; but 5 of those tribes also or sexine mostlywith monlyporate inaperturate, included genera of either Acalyphoideae or Eu- of knobsin hexagonalpattern; pis- ornamentation phorbioideae. Pistillateflower: sepals tillodepresent or absent. As predicted by Erdtman (1952), pollen mor- or valvate,often connate; petals (2)3-6, imbricate phology has provided decisive evidence in estab- or absent; ovary mostly2-4- and disk present lishinglines of phylogenetic affinitywithin the Cro- bifidto multifid(rarely entire); ovules locular;styles tonoideae. Punt (1962), followingErdtman's lead, Fruitcapsular solitaryin each locule, anatropous. grouped most of the genera of Crotonoideae under seeds carunculateor ecarun- (rarelyindehiscent); his "Croton configuration." However, "crotonoid" fleshy;endosperm usually culate, testa sometimes pollen is useful mainly for establishingmembership copious,often oily. in the subfamily,and except in the more primitive As definedhere, following Webster (1975), subtribes does not help to clarify problems of tribal familyCrotonoideae includes 12 tribes.This is a delimitation.

KEY TO THE TRIBES OF SUBFAMILY CROTONOIDEAE 1a. Pollen grainscolporate or porate,sexine reticulate to clavate; petals absent;indumentum mostly simple. 2a. Laticifersarticulated; pollen grains 3-nucleate; plants mostly monoecious; basic chromosomenumber x = 9. 3a. Pollen grainscolporate; styles bifid; seeds carunculateor ecarunculate;endosperm oily. 33. MICRANDREAE 3b. Pollen grainsperiporate; styles mostly multifid; seeds carunculate;endosperm starchy...... M.A.N...I...... 4.. M AN IH OTEAE 2b. Laticifersnonarticulated; pollen grains 2-nucleate; plants mostly dioecious; basic chromosomenumber x = 11. 4a. Pollen grainscolporate; leaves non-pellucid-punctate,stipules free; inflorescences axillary, mostly spicateor paniculate 35. ADENOCLINEAE 4b. Pollengrains periporate; leaves pellucid-punctate,stipules connate; flowers in clustersopposite the leaves GELON.EAE 36. lb. Pollen grainsinaperturate, sexine with"crotonoid" pattern; petals usuallypresent; indumentum simple, malpighiaceous,or stellate;laticifers nonarticulate (except in some Jatropheae). 5a. Seeds withoutendosperm, cotyledons massive; inflorescences dichasial; petals absent; stylesundivided,dilated; latex milky 37. ELATERIOSPERMEAE 5b. Seeds withcopious endosperm, cotyledons thin; petals present at least in staminateflowers (except in Neoboutonieae);inflorescences various; styles bifid to multifid,less commonlyunlobed. 6a. Staminatesepals imbricate,free or ifconnate then not completelycovering petals in the bud; seeds carunculateor ecarunculate. 7a. Indumentumsimple. 8a. Stamensmostly more than 5, freeor connate;pollen sexine coarsely clavate. 9a. Inflorescencesterminal, mostly dichasial; leaves mostlypalmately lobed or compound;fruits capsular or drupaceous,seeds carunculateor ecarunculate -...... 38. JATR O PHEA E 9b. Inflorescencesterminal or axillary,racemose or spicate to paniculate;leaves unlobed,usually without basal glands;fruit capsular 39. CODIAEAE 8b. Stamens3-5, filamentsconnate; pollen sexine finely clavate; monoecious;inflorescences terminalor axillary,racemose or thyrsoid;seeds ecarunculate. 40. TRIGONOSTEMONEAE 7b. Indumentum(at least in part) of stellateor lepidotehairs.

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1Oa. Fruitcapsular, seeds ecarunculate;petals free or absent. 1la. Pollensexine with spinose or reducedprocesses; leaves exstipulate;filaments oftenconnate, at least in part; cotyledonsbroad or narrow. 41. RICINOCARPEAE 1lb. Pollen sexinous,processes not spinoseor reduced;leaves usuallystipulate; filamentsfree; cotyledons broad 42. CROTONEAE lOb. Fruitdrupaceous, seeds ecarunculate;petals ? coherent 43. RICINODENDREAE 6b. Staminatesepals fused in thebud, splitting valvately or irregularlyinto segments; fruit capsular or drupaceous;seeds ecarunculate 44. ALEURITIDEAE

Tribe 33. MICRANDREAE (Muell.Arg.) Web- KEY TO THE GENERAOF SUBTRIBE MICRANDRINAE ster,Taxon 24: 598. 1975. Crotoneaesub- 1a. Sepals free;staminate disk present; stamens 5- tribe MicrandreaeMuell. Arg., DC. Prodr. 8; floralbracts small. 15(2): 709. 1866. Gelonieae subtribeChae- 2a. Antherselliptical; indumentum simple tocarpinae series MicrandriformesPax & 205. Micrandra 2b. Antherslinear; indumentum stellate Hoffm.,Pflanzenr. 147. XIV (Heft 68): 49...... _.__....._.___.__.__..._..._..._....._.._._...... 206.Micrand rropsis 1919. TYPE: Micrandra Benth. lb. Sepals connate;staminate disk absent; stamens 8-10; floralbracts large 207. Cunuria Monoecious or dioecioustrees; stems withar- ticulatelaticifers and usually whitishlatex; indu- 205. Micrandra Bentham, Hook. Kew J. Bot. or stellate;leaves alternate,un- mentumsimple 6: 371. 1854 (nom. cons.); Muell. Arg., DC. or triplinerved,with lobed,entire, pinnately veined Prodr. 15(2): 709. 1866; Fl. Bras. 11(2): stipulesoften laminar glands at apex of petiole; 289. 1873; Bentham, Gen. P1. 3: 289. 1880; or caducous. Inflorescencesterminal axillary,di- Pax, Pflanzenr. 147. I (Heft 42): 18. 1910; bracts small or large; flowers chasial-paniculate, Schultes, Bot. Mus. Leafl. 15: 201. 1952. apetalous. Staminatesepals 5, imbricate,free or TYPE: Micrandra siphonioides Benth. (typ. or absent;stamens connate;disk dissected,lobed, cons.). 5-10, free or connate; pollen grains 3-nucleate, 3-colporate,colpi operculate, sexine reticulate; pis- PogonophyllumDidr., Vidensk.Medd. Dansk Naturh. tillodepresent or absent. Pistillatesepals 5, im- Foren. Kj0benhaven1857: 144. 1857. TYPE: Po- gonophyllumelatum Didr. [= Micrandra elata bricate,free, deciduous in fruit;disk 5-lobed; ovary (Didr.) Muell. Arg.]. 3-locular; stylesfree, bifid. Fruit capsular; columella persistent;seeds large, carunculateor eca- A poorly understood genus of about seven Am- runculate,endosperm oily. azonian species.

As treatedby Webster (1975), thetribe includes 206. Micrandropsis Rodrigues, Acta Amazon- threeor fourgenera distributed into two subtribes. ica 3(2): 5. 1973. TYPE: Micrandropsis scle- The Micrandreaeclearly form a linkbetween sub- roxylon (W. Rodr.) W. Rodr. familiesAcalyphoideae and Crotonoideae. A monotypic genus of Amazonian Brazil.

KEY TO THE SUBTRIBES OF TRIBE MICRANDREAE 207. Cunuria Baillon, Adansonia I. 4: 287. Aug. 1a. Leaves simple;stamens free, sometimes inflexed in bud; sepals discrete,imbricate or valvate. 1864; Muell. Arg., DC. Prodr. 15(2): 1123. .------33a. Micrandrinae 1866; Bentham, Gen. P1. 3: 289. 1880; Pax, lb. Leaves palmatelycompound; stamens connate; Pflanzenr. 147. I (Heft 42): 16. 1910; Bald- sepals connate,valvate -33b. Heveinae win & Schultes, Bot. Mus. Leafl. Harvard Univ. 12: 325. 1947. TYPE: Cunuria spru- Subtribe 33a. MICRANDRINAE Muell. Arg., ceana Baillon. DC. Prodr. 15(2): 709. 1866. TYPE: Micran- dra Benth. ClusiophyllumMuell. Arg., Flora 47: 518. 25 Oct.1864. TYPE: Clusiophyllumsprucei Muell. Arg. [= Cun- Leaves simple,unlobed. Panicles with bracts uria crassipesMuell. Arg.]. conspicuousor inconspicuous.Staminate sepals im- Baldwin & Schultes (1947) recognized five spe- bricate,free or nearlyso; disk dissected;stamens cies of Cunuria, all from Amazonian South Amer- 5-7, filamentsfree, ? inflexedin bud; pistillode ica. Schultes (1952) on reconsideration reduced present;pistillate disk annular. Cunuria to a synonymof Micrandra, and because A subtribeof threeSouth Americangenera. of his intimateknowledge of these South American

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trees his opinion must be weighted heavily. The tum simple(stinging hairs sometimespresent); leaves listingof Cunuria as a separate genus is done here alternate, mostly palmately lobed or dissected; pet- mainly to call attentionto the problem,not to affirm ioles glandular or stipellate at apex; stipules usually its generic distinctness. present. Inflorescences terminal or pseudoaxillary, most racemose-paniculate or dichasial-paniculate, Subtribe 33b. HEVEINAE Muell. Arg., Lin- usually bisexual; flowersapetalous. Staminate flow- naea 34: 202. 1865; DC. Prodr. 15(2): 716. er: calyx synsepalous, sepals 5, imbricate, ? pet- 1866. TYPE: Hevea Aubl. aloid; disk intrastaminal or extrastaminal; stamens 8-10(-25), filamentsfree or connate; pollen grains Leaves palmately compound. Panicles with in- globose, 3-nucleate, periporate, sexine withCroton- conspicuous bracts. Staminate sepals connate into pattern; pistillodepresent or absent. Pistillate flow- a cup, lobes valvate; disk lobed or dissected; sta- er: sepals 5, free or connate, sometimes deciduous; mens 5-10, filamentsconnate into a column, an- disk annular, sometimes with staminodia; ovary thers sessile; pistillode present; pistillate disk dis- 3-locular (rarely 5-locular); styles free, mostlymul- sected or obsolete; seeds ecarunculate. tifidor lacerate (rarely bifid). Fruit capsular; col- As originallydelimited by Mueller (1865, 1866), umella often persistent; seeds carunculate, testa subtribe Heveinae is here interpreted to include dry. the single genus Hevea. Pax (1910) placed Hevea As delimitedhere, tribe Manihoteae includes two in subtribe Jatrophinae, a disposition that is con- genera, both confinedto the New World. The close tradicted by pollen morphology,chromosome num- affinitybetween Manihot and Cnidoscolus was in- ber, and other characters. dependently confirmed on palynological grounds by Punt (1962) and Miller & Webster (1962). 208. Hevea Aublet, Hist. PI. Guiane Fr. 2: 871, t. 335. 1775; Muell. Arg., DC. Prodr. 15(2): KEY TO THE GENERA OF TRIBE MANIHOTEAE 716. 1866; Fl. Brasil. 11(2): 297. 1874; Pax, Pflanzenr.147. I. (Heft 42): 117. 1910; 1a. Stinginghairs absent; stamensfree, staminate disk intrastaminal;staminate perianth Ducke, Arch. Inst. Biol. Veg. Rio Janeiro2: usually yellowishor greenish;leaf blades stipellate(not 217. 1935; Schultes,Bot. Mus. Leafl. Har- glandular)at base; inflorescenceracemose or vard Univ. 12: 1. 1945; 13: 1. 1947; 14: racemose-paniculate...... 209. Manihot 79. 1950; 15: 111,247, 255. 1952; 16:21. lb. Stinginghairs present;stamens connate (very 1953; 25: 243. 1977; Bot. Review36: 197. rarelyfree); staminate disk extrastaminal,annular; staminateperianth whitish; leaf blades 1970; Econ. Bot. 41: 125. 1987; Malaysian glandular(not stipellate)at base; inflorescence RubberRes. Dev. Board Mon. 14: 5. 1990. dichasial-paniculate...... 210. Cnidoscolus TYPE: Hevea guianensis Aubl. SiphoniaL. C. Richard,in Schreber,Gen. PI. 2: 656. 209. Manihot Miller, Gard. Dict. ed. 4. 1754; 1791. TYPE: Siphonia elastica Pers. [= Hevea gui- anensisAubl.; lectotype]. Pohl, P1. Bras. Ic. 1: 17. 1827; Muell. Arg., CaoutchouaJ. F. Gmelin,Syst. Nat. 2: 677. 1791. TYPE: DC. Prodr. 15(2): 1057. 1866; Bentham, Gen. Caoutchoua elastica Gmel. [= Hevea guianensis P1. 3: 306. 1880; Pax, Pflanzenr. 147. II Aubl.]. (Heft 44): 21. 1910; Croizat, J. Arnold Arbor. According to Schultes (1990), Hevea includes 23: 216. 1942; Rogers & Appan, Fl. Neotrop. ten species with several varieties, all found within 13: 1. 1973; Rogers & Fleming, Econ. Bot. Amazonian South America. Because of the eco- 27: 1. 1973; Allem, Rev. Brasil. Biol. 49: 1- nomic importance of the genus, there is extensive 26. 1989. Mandioca Link, Handb. 2: 436. literature on its systematics; references may be 1831. TYPE: Manihot esculenta Crantz [Ja- found in Schultes (1970, 1990). tropha manihotL.]. JaniphaHBK, Nov. Gen. Sp. 2: 106, t. 109. 1817. Tribe 34. MANIHOTEAE (Muell. Arg.) Pax, TYPE: Janiphaaesculifolia HBK [= Manihotaes- culifolia(HBK) Pohl]. Natiirl. Pflanzenfam. ed. 1, 3(5): 77. 1890. ManihotoidesRogers & Appan,Fl. Neotrop.13: 247. Hippomaneae subtribeManihoteae Muell. Arg., 1973. TYPE:Manihotoides pauciflora (Bdge.) Rog- DC. Prodr. 15(2): 1056. 1866. TYPE: Ma- ers& Appan[= Manihotpauciflora Bdge.]. nihot Miller. A neotropicalgenus of ca. 60 species,mostly Monoecious (rarely dioecious) trees, shrubs, or nativeto Brazil.Rogers & Appan(1973) created herbs; laticifers articulated, latex white; indumen- a newgenus, Manihotoides, for an aberrantspe-

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cies in Mexico characterized by reduced inflores- Most of the genera of this tribe were referred cences, and trifoliolateleaves clustered on short to the Gelonieae by Pax & Hoffmann(1931), but shoots. However, the recent discovery of another were scattered in differentsubtribes and associated Mexican species (ined.) with unlobed leaves on withgenera belongingto three differentsubfamilies. short shoots largely erases the supposed distinction. Following the arrangement of Webster (1975), the six genera are grouped into two subtribes. 210. Cnidoscolus Pohl, P1. Bras. 1: 56. 1827; Pax & Hoffm., Natiirl. Pflanzenfam. ed. 2, KEY TO THE SUBTRIBES OF TRIBE ADENOCLINEAE 19c: 1964. 1931; Lourteig & O'Donell, Lilloa la. Indumentumsimple; stamens free; fruit dehis- 9: 105. 1943; McVaugh, Bull. Torrey Bot. cent or indehiscent,2-3-locular. Club 71: 457. 1944; Lundell, Bull. Torrey 35a. Adenoclininae stellate; stamens connate; fruit Bot. Club 72: 319. 1945; Webster, J. Arnold lb. Indumentum baccate, 1-7-locular 35. Endosperminae Arbor. 48: 349. 1967; Breckon, Brittonia31: 125. 1979. TYPE: Cnidoscolus hamosus Pohl Subtribe 35a. ADENOCLININAE Muell.Arg., [lectotype, designated by Small, in Britton & Linnaea 34: 203. 1865; DC. Prodr. 15(2): Brown, Ill. Fl. N. U.S. ed. 2, 2: 462. 1913]. 1139. 1866. TYPE: Adenocline Turcz. JussieuiaHoustoun, Reliq. Houst.6. 1781 (nonJussiaea Gelonieaesubtribe Tetrorchidiinae Pax, Pflanzenr.147. L., 1753). TYPE: Jatrophaherbacea L. [= Cnidos- IV 52): 29. 1912. TYPE: TetrorchidiumPoepp. colus herbaceus(L.) I. M. Johnston]. (Heft Bivonea Raf., Specchio 1: 156. 1814 (nom. rej.). TYPE: Trees, shrubs, or herbs; indumentum simple or Bivonea stimulosa (Michx.) Raf. [= Cnidoscolus stimulosus(Michx.) Engelm.& Gray]. malpighiaceous; inflorescence diverse; stamens 3- VictoriniaLe6n, Mem. Soc. Cub. Hist. Nat. Felipe Poey 30, free; ovary 2- or 3-locular; stigmas distinct; 15: 242. 1941. TYPE: Victoriniaregina(Le6n)Le6n fruitcapsular or drupaceous. [Jatropharegina Le6n; = Cnidoscolusregina (Le6n) McVaugh]. This subtribe includes 5 genera with ca. 30 species of the Neotropics and Africa. A neotropical genus of ca. 50 species, with the greatest concentration in Mexico. The genus was KEY TO THE GENERA OF SUBTRIBE ADENOCLININAE reviewed by McVaugh (1944), and his reduction disk present,at least as staminodia. of Victorinia to synonymy appears justified. la. Pistillate 2a. Stamens25-30; fruitsdrupaceous; ovary 2-locular;leaves entire, with laminar glands Tribe 35. ADENOCLINEAE (Muell. Arg.) -...... 21 1. Glycydendron Webster, Taxon 24: 598. 1975. Hippoma- 2b. Stamensless than20; fruitscapsular; ova- neae subtribe Adenoclineae Muell. Arg., Lin- ry 2-3-locular; leaves ? dentate,with or withoutlaminar glands. 203. DC. Prodr. 15(2): 1139. naea 34: 1865; 3a. Trees or shrubs;seed-coat fleshy; sta- 1866. TYPE: AdenoclineTurcz. minatedisk not of interstaminalprocesses. Monoecious or dioecious trees, shrubs, or herbs; 4a. Stamens8-10; anthers2-locular, latex clear (? colored); indumentum simple, mal- not peltate;staminate disk pres- pighiaceous, or stellate; leaves alternate (rarely op- ent; leaves eglandular,stipules caducous; indumentumsimple; posite), simple, penninerved or triplinerved,with endotestasmooth. or withoutlaminar or petiolar glands; stipules per- 212. Klaineanthus sistent or deciduous. Inflorescences terminal, ax- 4b. Stamens 3; anthers 4-locular, illary, or opposite the leaves, spicate to paniculate peltate; staminatedisk absent; leavesusually with laminar glands, or glomerular; flowersapetalous. Staminate sepals stipulespersistent; indumentum 3-5, imbricate, free or connate; disk dissected or malpighiaceous(at least in part); absent; stamens 3-30, filamentsfree or connate; endotestafoveolate or echinulate pollen grains 2-nucleate, 3-colporate, colpi oper- -...... 213. Tetrorchidium culate or inoperculate, sexmiewith Croton-pattern; 3b. Herbs; seed-coatdry; staminatedisk of interstaminalprocesses. Pistillate sepals 3-5, pistillode present or absent. -...... 214. Adenocline imbricate, free or connate; disk dissected or an- lb. Pistillatedisk absent;leaves glandular-dentate, nular, sometimes stammodial; ovary 2-6-locular; withoutlaminar glands 215. Ditta styles bifidor stigmatiform.Fruit capsular or drupaceous; seeds ecarunculate, seed-coat dryor fleshy; 21 1. Glycydendron Ducke, Arq. Jard. Bot. Rio endosperm present. Janeiro 3: 199. 1922; 4: 107, pl. 10 figs. a-

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i. 1925. TYPE: Glycydendron amazonicum sirable at this time due to confusion in Turcza- Ducke. ninov's application of names to specimens].

A monotypicgenus of AmazonianBrazil. Pax Diplostylis Sonder,Linnaea 23: 113. 1850. TYPE: not & Hoffmann(1931) mistakenlyreduced it to a designated. synonymof Nealchornea because they misread Paradenocline Muell. Arg., DC. Prodr. 15(2): 1141. TYPE: Paradenocline procumbensMuell. Arg. [ the plate (Ducke, 1925), whichcontains both gen- Adenoclineviolifolia (Kunze) Prain]. era. As treatedby Prain (1920), Adenocline includes eightspecies of the Cape regionof South Africa. 212. Klaineanthus Pierreex Prain,Kew Bull. 1912: 105; Fl. Trop. Afr. 6(1): 963. 1913; 215. Ditta Grisebach, Mem. Amer. Acad. Arts. Hook. Ic. P1. 30: t. 2985. 1913; Pax & Sci. II. 8: 160. 1861; Muell. Arg., DC. Prodr. Hoffm.,Pflanzenr. 147. VII (Heft 63): 408. 15(2): 1138. 1866; Pax & Hoffm., 147. V 1914; Leonard, Fl. Congo 8(1): 130. 1962. (Heft 52): 270. 1912; Urban,Symb. Ant. 7: TYPE: Klaineanthus gaboniae Pierreex Prain. 261. 1912; Alain, Fl. Cuba 3: 112. 1953; A monotypicgenus of West Africa(Nigeria to Borhidiet al., Acta Agron.Acad. Sci. Hungar. Gabon). 27: 432. 1978; Liogier, Fl. Espaniola 4: 135, 348. 1986; Descr. Fl. Puerto Rico 2: 383. 1988. TYPE: Ditta myricoides Griseb. 213. Tetrorchidium Poeppigin Poeppig& En- dlicher,Nov. Gen. Sp. 3: 23, t. 227. 1842; A genus of one or two species of the Greater Muell. Arg., DC. Prodr. 15(2): 1132. 1866; Antilles (Cuba, Hispaniola, Puerto Rico). Bentham,Gen. P1. 3: 288. 1880; Pax & Hoffm.,Pflanzenr. 147. IV (Heft 52): 1912; Subtribe 35b. ENDOSPERMINAE Pax & Natiirl.Pflanzenfam. ed. 2, 19c: 184. 1931; Hoffmann,Pflanzenr. 147. XIV (Heft 68): 53. Cuatrecasas,Brittonia 9: 76. 1957; Leonard, 1919 (nomen); Natiirl. Pflanzenfam. ed. 2, Fl. Congo 8(1): 133. 1962; Radcliffe-Smith, 19c: 42. 1931 (in clavi). TYPE: Endospermum Fl. E. Trop. Afr.,Euphorb. 1: 373. 1987; Bentham. Huft, Ann. MissouriBot. Gard. 75: 1110. Dioecious trees or shrubs, indumentum stellate; 1989. TYPE: Tetrorchidium rubrivenium inflorescences axillary; staminate calyx gamophyl- Poepp. lous; stamens 6-10, filaments connate; pistillode HasskarliaBaillon, Adansonia I. 1: 51. 1860 (nonMeis- small or absent; ovary 2-6-locular; styles stig- ner,1843). TetrorchidiopsisRauschert, Taxon 31: matiform,confluent into a cap; fruitcapsular. 559. 1982. TYPE: Hasskarlia didymostemonBaill. [= Tetrorchidiumdidymostemon (Baillon) Pax & The subtribe includes only the type genus En- Hoffm.]. dospermum. A genus of ca. 20 species, mostlyin tropical 216. Endospermum Bentham,Fl. Hongkong. Americabut ca. 5 species in Africa.The African 304. 1861 (nom. cons.); Muell. Arg., DC. specieswere referred to a separategenus Hasskar- Prodr. 15(2): 1131. 1866; Bentham, Gen. P1. lia by Pax & Hoffmann(1912), but they later 3: 322. 1880; Beccari, Malesia 2: 45, t. 2. (Pax & Hoffmann,1931) reducedit to a section 1884; Pax & Hoffmann, Pflanzenr. 147. IV of Tetrorchidium. (Heft 52): 33. 1912; Docters van Leeuwen, Treubia 10: 431, tt. 12, 13. 1919; Pax & 214. Adenocline Turczaniov, Bull. Soc. Imp. Hoffm.,Natiirl. Pflanzenfam. ed. 2, 19c: 184. Nat. Moscou 16: 59. 1843; Muell. Arg.,DC. 1931; Schodde, Blumea 15: 397. 1967; Prodr.15(2): 1139. 1866; Bentham,Gen. P1. Schaeffer,Blumea 19: 171. 1971; AiryShaw, 3: 310. 1880; Prami,Ann. Bot. 27: 404. Kew Bull. Add. Ser. 8: 78. 1980. TYPE: En- 1913; Pax & Hoffm.,Pflanzenr. 147. VII dospermum chinense Benth. (Heft 63): 409. 1914; Prain, Fl. Cap. 5(2): 488. 1920; Milne-Redhead,Kew Bull. 5: 349. Capellenia Teijsmann& Binnendijk,Natuurk. Tijdscr. 1951; Dyer, Gen. S. Afr. Fl. P1. ed. 3, 1: Ned.-Ind. 29: 238. 1867. TYPE: Capellenia mol- 315. 1975. Mercurialis sect. Adenocline uccana Teijsm. & Binn. [= Endospermummoluc- canum (Teijsm. & Binn.) Kurz]. (Turcz.) Baillon,Adansonia I. 3: 159. 1864. TYPE: not designated[lectotypification not de- A paleotropical genus of ten species, occurring

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from China to Malaya, New Guinea, and tropical Tribe 37. ELATERIOSPERMEAE Webster, Australia (Queensland). Taxon 24: 599. 1975. TYPE: Elateriosper- mum Blume. Tribe 36. GELONIEAE (Muell. Arg.) Pax, Na- Monoecious trees; stems with whitishlatex; in- tiirl. Pflanzenfam. ed. 1, 3(5): 88. 1890. Hip- dumentum simple; leaves alternate, pinnately pomaneae subtribeGelonieae Muell. Arg., Lin- veined, entire, petiole biglandular at apex; stipules naea 34: 202. 1865; DC. Prodr. 15(2): 1124. caducous. Inflorescences terminal or axillary, cy- 1866. TYPE: Gelonium Roxb. ex Willd. [= mose, bisexual; flowersapetalous. Staminate sepals Suregada Roxb. ex Rottl.]. 5, free, imbricate; disk lobed, pubescent; stamens Dioecious (very rarely monoecious?) trees or 10-18, filaments free; anthers apiculate; pollen shrubs; latex not evident; indument simple, usually grains globose, inaperturate, with Croton-pattern; very scant or absent; leaves alternate, simple, pen- pistillode rudimentary or absent. Pistillate sepals ninerved, lamina usually pellucid-punctate; stipules 5, imbricate, caducous; disk cupular, pubescent; caducous. Inflorescences opposite the leaves; flow- ovary 2-4-locular; styles unlobed, stigmas dilated. ers apetalous, in glomerules. Staminate sepals usu- Fruit capsular but exocarp fleshy; seeds ecarun- ally 5 or 6, imbricate, free, sometimes glandular culate endosperm scanty, cotyledons massive. abaxially; disk extrastaminal or intrastaminal, an- A monotypic tribe including only Elateriosper- nular or dissected; stamens (6-)10-25(-60), fil- mum. The genus was placed between Suregada aments free; anthers dehiscing longitudinally;pol- and Endospermum by Mueller (1866), while Ben- len grains 2-nucleate, subglobose, 3-6-porate, exine tham (1880) and Pax (1910) positioned it with with Croton-pattern;pistillode absent. Pistillate se- Cunuria and Micrandra. The inaperturate pollen pals mostly 5(4-8), imbricate,sometimes glandular grains contradict both these positions, however. abaxially; disk annular, sometimes withstaminodes; Airy Shaw (1975) included it in his subtribe Ja- ovary 3-(rarely 2- or 4-)locular; styles bifid(rarely trophinae along with Aleurites, Annesijoa, Ta- multifid).Fruit capsular or sometimes indehiscent poides, and Loerzingia. The genus appears to be and drupaceous; seeds ecarunculate, seed-coat + a relict, without any close affinities,but perhaps it fleshy; endosperm present. is a distant relative of Micrandra and Manihot. A monogeneric paleotropical tribe, as treated 218. Elateriospermum Blume, Bijdr. Fl. Jav. here. Pax & Hoffmann(1931), in their finaltreat- 620. 1826; Muell. Arg., DC. Prodr. 15(2): ment, construed Gelonieae as a large tribe with 18 1130. 1866; Bentham, Hook. Ic. P1. 13: t. genera in 7 subtribes; a considerable number of 1294. 1879; Gen. P1. 3: 288. 1880; Pax, those genera are here included in the Adenoclineae. Pflanzenr. 147. I (Heft 42): 17. 1910; J. J. Smith, Med. Dept. Landb. 10: 571. 1910; Backer & Bakh. f., Fl. Java 1: 497. 1963; 217. Suregada Roxburgh ex Rottler, Ges. Na- Whitmore, Tree Fl. Malaya 2: 91. 1973; Airy turf. Freunde Berlin, Neue Schriften 4: 206. Shaw, Kew Bull. Add. Ser. 4: 108. 1975; 1803; Baillon, Etude Gen, Euphorb. 395. Kew Bull. 36: 292. 1981. TYPE: Elateriosper- 1858; Croizat, Bull. Bot. Gard. Buitenzorg III. mum tapos Bl. [lectotype; the other species 17: 212. 1942; Merrill, J. Arnold Arbor. 37: of Blume, E. tokbrai, was transferredto Blu- 79. 1951; Leonard, Bull. Jard. Bot. Brux. 28: meodendron]. 443. 1958; Fl. Congo 8(1): 124. 1962; Rad- cliffe-Smith,Fl. E. Trop. Afr., Euphorb. 1: A monotypic genus of tropical southeast Asia: 376. 1987. TYPE: Suregada roxburghii Rot- Thailand to Java and Borneo. tler. Tribe 38. JATROPHEAE (Meisner) Pax, Na- GeloniumRoxb. ex Willd., Sp. P1. 4: 831. 1806 (non tiurl. Pflanzen fam. ed. 1, 3(5): 72. 1890; TYPE: Willd. Gaertn.,1791). Geloniumlanceolatum Pflanzenr. 147. I (Heft 42): 1. 1910. Cro- [= Suregada lanceolata (Willd.) Croiz.]. Meisn., P1. Vasc. ErythrocarpusBlume, Bijdr. Fl. Jav. 604. 1825. TYPE: toneae subtribe Jatropheae Erythrocarpus glomerulatus BI. [= Suregada Gen. 1: 341. 1841. Jatropheae subtribe Ja- glomerulata(BI.) Baillon;lectotype]. trophinae Pax, Pflanzenr. 147. I (Heft 42): CeratophorusSonder, Linnaea 23: 120. 1850. TYPE: 21. 1910. TYPE: Jatropha L. Ceratophorusafricanus Sond. [= Suregada africana (Sond.) Croiz.]. Joannesieae(Muell. Arg.) Pax, Bot.Jahrb. 59: 142. 1924. Acalypheae subtribe Joannesieae ('Johanneseae') A paleotropical genus of ca. 40 species, the Muell. Arg., Linnaea 34: 201. 1865. TYPE: Joan- majority in tropical Asia. nesia Vell.

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Monoecious (less commonlydioecious) trees, lb. Leaves palmatelycompound; monoecious; seeds shrubs,or herbs; stems with articulatedand/or ecarunculate. nonarticulatedlaticifers, latex clear, white,or red- 5a. Stylesdilated, subentire to lacinate;ovary 2-locular;anthers apiculate; foliagewith dish; indumentumsimple, sometimesglandular; malpighiaceoushairs. leavesunlobed or palmately lobed or parted,usually 6a. Staminatecalyx cupular, not covering withglands at apex of petiole; stipulespersistent thepetals in bud;stamens 7- 10, inner or deciduous. Inflorescencesterminal and some- filamentsconnate; staminate disk dissected; fruitdrupaceous. 223. Joannesia times axillary as well, dichasial-paniculatebut 6b. Staminatecalyx fused in bud, com- sometimesreduced. Staminatesepals usually 5, pletelycovering the petals; stamens distinct,imbricate; petals 5, freeor sometimesco- 18-33, innerfilaments connate; sta- herentor connate;disk entire or dissected;stamens minatedisk annular and intrastaminal; (6)8-30, filamentsat least partlyconnate; pollen fruitunknown . 224. Leeuwenbergia 5b. Stylesslender, bifid; ovary 3-locular;an- grainsglobose, binucleate, inaperturate, sexine with thersmuticous; foliage glabrous; staminate Croton-pattern;pistillode present or absent.Pistil- calyx 5-lobed;stamens 15-25, innerfila- late sepals 5(6), free,imbricate, ? persistentin mentsconnate; staminatedisk dissected; fruit;disk annular,lobed, or dissected,sometimes fruitcapsular .225. Annesiloa withstaminodia; ovary mostly 2-3-locular; styles unlobedor bifid,sometimes dilated. Fruit capsular 219. Jatropha L., Sp. P1. 1006. 1753; Gen. or drupaceous;seeds carunculateor ecarunculate, P1. ed. 5, 437. 1754; Muell. Arg., DC. Prodr. endospermpresent. 15(2): 1076. 1866; Bentham, Gen. P1. 3: 290. 1880; Pax, Pflanzenr. 147. I (Heft 42): As circumscribedhere, tribe Jatropheae includes 21. 1910; Hutchinson, Fl. Trop. Afr. 6(1): onlytwo of the genera(Jatropha and Joannesia) 775. 1912; Fawc. & Rend., Fl. Jam. 4: 310. assignedby Pax (1910). None of the genera enu- 1920; Pax & Hoffm., Naturl. Pflanzenfam. meratedfor the Jatropheaeby Hutchinson(1969) ed. 2, 19c: 160. 1931; McVaugh, Bull. Tor- appear to belongin the same tribewith Jatropha. rey Bot. Club 72: 271. 1944; Webster, J. My previouscircumscription (Webster, 1975) of Arnold Arbor. 48: 340. 1967; Dehgan & the Joannesieae(a name mistakenlyadopted for Webster, Univ. California Publ. Bot. 74: 1. the Jatropheae)was broaderthan the presentJa- 1979; Radcliffe-Smith,Fl. Trop. E. Afr., Eu- tropheae,since it includedtwo genera, Givotia and phorb. 1: 343. 1987. Adenoropium Pohl, P1. Ricinodendron,that are now referredelsewhere. Bras. Ic. Descr. 1: 12. 1827. TYPE: Jatropha It mightbe possibleto maintaintwo subtribes within gossypiifolia L. [lectotype]. the Jatropheae,based on Jatrophaand Joannesia respectively;but Deutzianthus and Oligocerasare Curcas Adanson,Fam. P1. 2: 356. 1763. TYPE: Jatropha curcas L. intermediatein many respects,and it now seems CastiglioniaRuiz & Pav6n, Fl. Peruv. 139. 1794. TYPE: betternot to recognizeany subtribeswithin the Castiglionia lobata Ruiz & Pav6n, nom. illeg. [= Jatropheae. Jatrophacurcas L.]. Loureira Cavanilles,Icon. Descr. P1. 5: 17. 1799. TYPE: Loureira glanduliferaCav. [= Jatropha cordata KEY TO THE GENERA OF TRIBE JATROPHEAE (Ort.) Muell. Arg.]. Mozinna Ortega,Nov. Rar. P1. Hort.Bot. Matr. 8: 104. la. Leavessimple (lobed or unlobed);monoecious 1798. TYPE: Mozinna spathulata Ort.[= Jatropha or dioecious;staminate sepals free or connate; dioica Sesse]. latexusually reddish. Zimapania Engler & Pax, Natiurl.Pflanzenfam. ed. 1, 2a. Stylesbifid or unlobed; sepals distinct; mon- 3(5): 119. 1891. TYPE: Zimapania schiedeana En- oecious(rarely dioecious). gler & Pax [= Jatrophadioica Sesse]. 3a. Fruitcapsular; sepals not horned on Collenucia Chiovenda,Fl. Somala 1: 177. 1929. TYPE: back. Collenucia paradoxa Chiov. [= Jatropha para- 4a. Seeds carunculate;leaves often doxa (Chiov.) Chiov.]. lobedor dentate, without distinct pairedglands at apexof petiole; A diverse genus of ca. 175 species of tropics stipulesusually persistent, not in- and subtropics in the Old World and New World; fra-axillary 219. Jatropha 4b. Seeds ecarunculate;leaves un- native species absent fromAustralia and the Pacific lobed,entire, with paired glands islands. The assignment of species into subgenera at apexof petiole; stipules infra- and sections is given by Dehgan & Webster (1979). axillary,caducous - 220. Vaupesia 3b. Fruitdrupaceous; sepals horned on back-221. Oligoceras 220. Vaupesia Schultes, Bot. Mus. Leafl. Har- 2b. Stylestwice bifid; staminate sepals connate vard Univ. 17: 27. 1955. TYPE: Vaupesia below;dioecious 222. Deutzianthus cataractarum Schultes.

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A monotypic genus of the Rio Negro region, A neotropical genus of two South American spe- Amazonian Colombia. Vaupesia is a remarkable cies. genus with latex and foliage of tribe Micrandreae but inaperturate pollen very similar to that of Ja- 224. Leeuwenbergia Letouzey & HallM,Adan- tropha. It may therefore represent a connecting sonia II. 14: 379. 1974. TYPE: Leeuwenber- link between tribes Micrandreae and Jatropheae; gia letestui Letouzey & HallM. comm.) regards it as better placed Schultes (pers. A tropical west African genus of two species in Micrandreae. in the Cameroon, Congo, and Gabon. The genus is in many ways intermediate between Joannesia and 221. Oligoceras Gagnepain, Bull. Soc. Bot. Annesijoa, but the type species (L. letestui) differs 872. 1925; Fl. Indochine5: 467. France 71: strikinglyfrom both in having a completely gam- AiryShaw, Kew Bull. 14: 392. 1960. 1926; ophyllous calyx that splits open on anthesis as in TYPE: Oligoceras eberhardtii Gagnep. genera of Aleuritideae. Unfortunately,the stami- A monotypic genus of Indochina; closely related nate flowersof the second species, Leeuwenbergia to Deutzianthus but distinguished by its monoe- africana Let. & Halle, are still unknown. cious flowerproduction and curious horned sepals. 225. Annesijoa Pax & Hoffmann, Pflanzenr. 222. Deutzianthus Gagnepain, Bull. Soc. Bot. 147. XIV (Heft 68): 9. 1919; Airy Shaw, France 71: 139. 1924; Fl. Indochine 5: 297. Kew Bull. 16: 345. 1963; Hook. Ic. P1. 38: 1925; Wu, Acta Phytotax. Sin. 6: 245. 1957; t. 3713. 1974; Kew Bull. Add. Ser. 8: 27. Airy Shaw, Kew Bull. 14: 362. 1960; 16: 1980. TYPE: Annesijoa novoguineensis Pax 346. 1963. TYPE: Deutzianthus tonkinensis & Hoffm. Gagnep. A monotypic genus endemic to New Guinea. Loerzingia AiryShaw, Kew Bull. 16: 365. 1963; 25: 543. 1972; 36: 312. 1980, syn. nov. TYPE: Loer- Tribe 39. CODIAEAE (Pax) Hutchinson, Amer. zingia thyrsifloraAiry Shaw [= Deutzianthus J. Bot. 56: 747. 1969. Clutieae subtribeCodi- thyrsiflorus(Airy Shaw) Webster,comb. nov.]. aeinae Pax, Pflanzenr. 147. III (Heft 47): 10. A genus of two species, in Indochina and Su- 1911. TYPE: Codiaeum Rumph. ex A. Juss. matra. Airy Shaw compared his Loerzingia with Monoecious or dioecious trees or shrubs; stems Tapoides Airy Shaw from Borneo; however, the with nonarticulated laticifers, latex clear (some- staminate calyx, with distinctimbricate lobes, sug- times reddish) and nontoxic; indumentum simple gests a closer affinityto tribe Jatropheae. Com- or stellate; leaves penninerved or triplinerved,un- parison of the illustrations of Gagnepain (1926) lobed, usually eglandular; stipules often caducous, and Airy Shaw (1981) indicates that in fact the sometimes obsolete. Inflorescences terminal or ax- Sumatran Loerzingia is extremely similar to the illary,racemose or paniculate. Staminate sepals 4- Indochinese Deutzianthus, and their differences 6, free or connate, imbricate or valvate; petals seem of specific rather than generic value. mostly 5, free, usually imbricate; disk dissected or lobed; stamens (5-) 10- 100 or more, free or basally 223. Joannesia Vellozo, Alogr. Alkalis 199. connate; pollen grains binucleate, globose, inaper- 1798; Muell. Arg., DC. Prodr. 15(2): 715. turate, sexine with Croton-pattern; pistillode ab- 1866; Bentham,Gen. P1. 3: 290. 1880; Pax, sent. Pistillate sepals 4, usually imbricate, some- Pflanzenr.147. 1 (Heft42): 116. 1910; Ducke, timesaccrescent; petals usually 5, sometimesabsent; Arch.Jard. Bot. Rio de Janeiro3: 198. 1922; disk annular (rarely dissected or absent); ovary Schultes, Bot. Mus. Leafl. Harvard Univ. 17: mostly 3-locular; styles unlobed to bipartite. Fruit 25. 1955; Smith et al., Fl. Ilustr. Catar., capsular; columella usually persistent; seeds ca- EUFO 165. 1988. TYPE: Joannesia princeps runculate or ecarunculate, testa sometimes fleshy; Vell. endosperm present.

Anda A. Juss., Tent. Euphorb. 3. 1824. TYPE: Anda This tribe of 15 genera with over 100 species gomesiiA. Juss.,nom. illeg. [= Joannesiaprinceps has the greatest generic diversity in the Crotono- Vell.]. ideae, and relationshipsbetween the genera are not Andicus Vellozo,Fl. Flum. 80, t. 86. 1825. TYPE: An- to attempt to recognize dicus pentaphyllus Vell. [= Joannesia princeps clear; it seems premature Vell.]. formal subtribes at this time.

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KEY TO THE GENERA OF TRIBE CODIAEAE la. Petalspresent, at leastin staminateflowers. 2a. Petalspresent in pistillateflowers. 3a. Staminatecalyx lobed; fruit capsular. 4a. Leaveswithout beadlike glands beneath; anthers and petalsnot gland-tipped; inflorescences terminalor axillary,not opposite the leaves. 5a. Inflorescencesterminal; seeds carunculate; staminate disk and receptacle glabrous. 6a. Ovary3-locular; petals glabrous outside ...... 226. Baloghia 6b. Ovary2-locular; petals golden-sericeous outside ...... 227. Hylandia 5b. Inflorescencesaxillary; staminate disk or receptaclepilose. 7a. Indumentumsimple; styles bifid 228. Ostodes 7b. Indumentum(in inflorescence) malpighiaceous; staminate disk cupular. 8a. Stylesbifid; staminate disk glabrous; stamens mostly 5-7; petalspubescent adaxially;seeds carunculate .229. Pausandra 8b. Stylesmultifid; staminate disk pubescent; stamens 14-16; petalsnot pubescent adaxially;seeds ecarunculate .230. Dodecastigma 4b. Leaveswith beadlike glands beneath; anthers and petalswith apical glands; inflorescences oppositethe leaves .231. Pantadenia 3b. Staminatecalyx almost truncate. 9a. Pistillatecalyx accrescent; petals glabrous; fruit capsular .232. Dimorphocalyx 9b. Pistillatecalyx not accrescent; petals tomentose; fruit drupaceous .233. Fontainea 2b. Petalsabsent in pistillateflowers. 10a. Inflorescencesaxillary. 1la. Flowersin racemes;stipules caducous or absent;anthers muticous. 12a. Stylesentire, slender .234. Codiaeum 12b. Stylestwice bifid .235. Sphyranthera 11b. Flowersin axillaryclusters; stipules persistent, ? spinose;anthers apiculate; styles bifid or dilated..236. Acidocroton 10b. Inflorescencesterminal. 13a. Pistillatesepals accrescent, not recurved; floral disk present. 14a. Pistillatesepals eglandular; seeds mostly ecarunculate . .237. Blachia 14b. Pistillatesepals glandular; seeds carunculate .238. Strophioblachia 13b. Pistillatesepals recurved, not accrescent; floral disk absent . .239. Sagotia lb. Petalsabsent; styles bifid; seeds carunculate; leaves glandular . .240. Baliospermum

226. Baloghia Endlicher,Prodr. Fl. Norf.84. Loerzingia (Deutzianthus), especiallythe latter. 1833; Bentham,Gen. P1. 3: 300. 1880; Pax, These suggestionsare generallycompatible with Pflanzenr. 147. III (Heft 47): 12. 1911; thepresent treatment, but it appears that Baloghia Maiden, For. Fl. N. S. W. 1: 165, t. 28. may prove to be the closestgenus. 1904; White, Proc. Roy. Soc. Queensl. 53: 226. 1942; AiryShaw, Kew Bull. 35: 598. 228. Ostodes Blume,Bijdr. Fl. Ned. Ind. 619. 1980; McPherson& Tirel, Fl. Nouv.-Caled. 1825; Muell. Arg. 15(2): 1114. 1866; Ben- 14(1): 43. 1987; James& Harden,Fl. N. S. tham,Gen. P1. 3: 299. 1880; Hook. f., Fl. W. 1: 410, 1990. TYPE: Baloghia lucida Brit.Ind. 5: 400. 1887; Pax & Hoffm.,Pflan- Endl. zenr. 147. III (Heft 47): 17. 1911; Gagne- pain, Fl. Indochine5: 322. 1925; AiryShaw, SteigeriaMuell. Arg., Linnaea 34: 215. 1865;DC. Prodr. Kew Bull. 20: 409. 1966; 26: 311. 1971; 15(2): 1121. 1866.TYPE: Steigeriamontana Muell. 35: 334. 1981. TYPE: Ostodespaniculata Bl. Arg.[= Baloghiamontana (Muell. Arg.) Pax]. As circumscribedby AiryShaw (1966), Ostodes An Australasiangenus of 15 species, 12 in New is a genus of four species of tropicalAsia, from Caledonia and the othersin NorfolkIsland, Lord theeastern Himalayas to northernBorneo. A num- Howe Island, and Australia. ber ofspecies included by Pax & Hoffmann(1911) were removedby Airy Shaw (1966) to Fahren- 227. Hylandia AiryShaw, Kew Bull. 29: 329. heitia, a dispositionthat is accepted here pending 1974; 35: 643, fig.4. 1980. TYPE: Hylandia furtherstudy. dockrilliiAiry Shaw. A monotypic genus of tropical Australia 229. Pausandra Radlkofer,Flora 53: 92, t. 2. (Queensland).Airy Shaw (1974) suggesteda re- 1870; Baillon, Adansonia I. 11: 92. 1873; lationship with Dimorphocalyx, Ostodes, and Muell. Arg., Fl. Brasil. 11(2): 503, t. 99.

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1874; Pax & Hoffm., Pflanzenr. 147. III 1975; Kew Bull. 35: 624. 1980. TYPE: Di- (Heft 47): 41. 1911; Lanjouw, Rec. Trav. morphocalyx glabellus Thw. Bot. Neerl. 33: 758. 1936; Jablonski, Mem. A genus of ca. 15 species, mainlyin tropical New York Bot. Gard. 17: 153. 1967; Secco, Asia (India and Sri Lanka to Hainan and Borneo) Bol. Mus. Par. Emilio Goeldi, Bot. 3: 59. but extendingto Australia. 1987; Huft, Ann. Missouri Bot. Gard. 75: 1115. 1989; Secco, Rev. Gen. Anomalocalyx 233. Fontainea Heckel, Etudes sur Fontainea ... Amer. Sul 58. 1990. TYPE: Pausandra pancheri. 1870; Baillon, Hist. P1. 5: 194. morisoniana (Casar.) Radlk. 1874; Pax, Pflanzenr. 147. III (Heft 47): 30. As revised by Secco (1990), Pausandra is a 1911; AiryShaw, Kew Bull. 35: 632. 1980; neotropical genus of ca. eight species, extending Jessup & Guymer, Austrobaileya2: 112. from Honduras to Brazil. The genus is very dis- 1985; McPherson& Tirel, Fl. Nouv.-Caled. tinctive and isolated fromother genera of the tribe 14(1): 74. 1987. TYPE: Fontainea pancheri except for Dodecastigma, which is similar in in- (Baillon)Heckel. dumentum and staminate disk, but very different An Australasiangenus of six species,extending in a number of other characters. fromAustralia to New Guinea,New Caledonia,and Vanuatu. 230. Dodecastigma Ducke, Notizbl. Bot. Gart. Berlin 11: 343. 1932; Arq. Jard. Bot. Rio 234. Codiaeum Rumphiusex A. Jussieu,Eu- Janeiro 6: 58, t. 5. 1933; Sandwith, Kew phorb.Tent. 33. 1824 (nom. cons.); Muell. Bull. 1950: 134. 1951; Jablonski,Mem. New Arg., DC. Prodr. 15(2): 1116. 1866; Ben- York Bot. Gard. 17: 154. 1967; Secco, Rev. tham,Gen. P1. 3: 299. 1880; Pax, Pflanzenr. Gen. Anomalocalyx ... Amer. Sul 42. 1990. 147. III (Heft 47): 23. 1911; Airy Shaw, TYPE: Dodecastigma amazonicum Ducke. Kew Bull.Add. Ser. 4: 88. 1975; 8: 62. 1980; A genus of two species of the Brazilian Amazon A. C. Smith,Fl. VitiensisNova 2: 549. 1981; and the Guianas. McPherson& Tirel, Fl. Nouv.-Calkd.14(1): 95. 1987. TYPE: Codiaeum variegatum (L.) 231. Pantadenia Gagnepain, Bull. Soc. Bot. A. Juss. France 71: 873. 1925; Fl. Indochine 5: 470. Phyllaurea Loureiro,Fl. Cochinch. 575. 1790. TYPE: 1926; Airy Shaw, Kew Bull. 28: 122. 1969; Phyllaurea codiaeum Lour. [nom. illeg., = Codi- 26: 312. 1972. TYPE: Pantadenia adenanth- aeum variegatum(L.) A. Juss.]. era Gagn. Synapisma Endlicher,Gen. P1. 1110. 1840. TYPE: Cro- zophora peltata Labill.[= Codiaeumpeltatum (La- Parapantadenia Capuron,Adansonia II. 12: 206. 1972. bill.) P. Green]. TYPE: Parapantadenia chauvettaeLeandri [= Pan- JunghuhniaMiquel, Fl. Ind. Batav. 1(2): 412. 1859 tadenia chauvetiae (Leandri) Webster, comb. (non Corda, 1842). TYPE: Junghuhniaglabra Miq. nov.]. [ Codiaeum variegatum(L.) A. Juss.]. A paleotropical genus of two widely disjunct An Australasiangenus of ca. 15 species, from species, one endemic to Madagascar, the other Borneoand thePhilippines to New Guinea,tropical found in Thailand and Indochina. The genus Para- Australia,New Caledonia,and the Pacificislands. pantadenia was distinguishedby the 2-locular ova- Mueller(1866) adoptedan extremelybroad con- ry that develops into a 1-seeded, indehiscent fruit. cept of Codiaeum,including Baloghia and Blach- However, the species are otherwise so similar in ia as sections;however, recent workers concur in the peculiar indumentum and inflorescence that it the narrowerdefinition. seems best to regard them as congeneric. 235. Sphyranthera Hooker f., Hook. Ic. P1. 18: t. 1702. 1887; Chakrabarty& Vasudeva 232. Dimorphocalyx Thwaites, Enum. PI. Zeyl. Rao, J. Econ. Tax. Bot. 5: 959. 1984; 6: 278. 1861; Bentham,Gen. PI. 3: 301. 1880; 429. 1985. TYPE: Sphyranthera capitellata Hook. f., Fl. Brit.Ind. 5: 403. 1887; Trimen, Hook. f., nom. illeg. [= Sphyranthera lutes- Handb. Fl. Ceylon 4: 54, pl. 84. 1898; Pax, cens (Kurz) Pax & Hoffm.]. Pflanzenr. 147. III (Heft 47): 31. 1911; Airy Shaw, Kew Bull. 23: 123. 1969; 26: 251. An endemic genus of the Andaman/Nicobar 1972; Whitmore,Tree Fl. Malaya 2: 86. archipelago;there are two species, the type and 1973; AiryShaw, Kew Bull. Add. Ser. 4: 95. the recentlydescribed S. airyshawii Chakrab. &

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Vasud. The relationshipsof the genusare stillun- ouslyinterpreted. Mueller (1866) createdfor it a certain;Hooker suggested that Sphyranthera might subtribeplaced betweenPogonophora and Clutia. belongto the Acalypheae. Bentham(1880) suggestedan affinitywith Jatro- pha; and Pax (1910), apparentlyfollowing Ben- 236. Acidocroton Grisebach,Fl. Br. W. Ind. tham's lead, referredAcidocroton to tribeJatro- 42. 1859 (nom. cons. prop.; proposedhere); pheae, but located it in subtribeMicrandrinae Muell. Arg., DC. Prodr. 15(2): 1042. 1866; adjacentto Garcia. Althoughthe matter still needs Bentham,Gen. P1. 3: 291. 1880; Pax, Pflan- morestudy, the totality of floral characters support zenr. 147. I (Heft 42): 13. 1910; Urban, a placementin tribeCodiaeae. Symb. Ant. 7: 513. 1913; Fawc. & Rend., Fl. Jam. 4: 316. 1920; Urban, Symb. Ant. 237. Blachia Baillon,Etude GCn. Euphorb. 385. 9: 208. 1924; Ark.Bot. 20A(15): 62. 1926; 1858; Bentham,Gen. P1. 3: 301. 1880; Hook. Alain,Fl. Cuba 3: 73. 1953; Fl. Espafiola4: f.,Fl. Brit.Ind. 5: 402. 1887; Pax, Pflanzenr. 69. 1986. TYPE: Acidocroton adelioides Gri- 147. III (Heft47): 36. 1911; Gagnepain,Fl. seb. Indochine 5: 410. 1926; Airy Shaw, Kew Bull. 23: 121. 1969; 26: 223. 1972; Kew OphellanthaStandley, J. Wash. Acad. Sci. 14: 97. 1924; Bull. Add. Ser. 4: 57. 1975; Thin, Tap Chi syn. nov. TYPE: Ophellantha spinosa Standl.,op. cit. 98 [= Acidocroton spinosus (Standl.)Web- Sinh Hoc 11(3): 16. 1989. TYPE: Blachia ster,comb. nov.]. umbellata (Willd.) Baill. [Croton umbellatus Willd.]. 2 from A mainly Caribbean genus of 12 species, A genusof ca. ten species,in tropicalAsia from the re- Central America (Chiapas/Guatemala) and southernIndia to Hainan,the Philippines,and per- Antilles,with the ma- mainderfrom the Greater haps Borneo. The genus appears relatedto Aci- in undescribedspecies is reported jority Cuba (an docrotonof the New World,especially the section comm.). The genus from Colombia by Huft; pers. Ophellantha. has been subject to both nomenclatural and taxonomic problems since its description. Grisebach 238. Strophioblachia Boerlage,Handl. Fl. Ned. cited Acidoton P. Brown and Adelia acidoton L. Ind. 3(1): 235. 1900; Pax, Pflanzenr.147. as the basis for his genus and type species; unfor- III (Heft 47): 35. 1911; Merrill,Univ. Cali- tunately, the Jamaican plant of Browne and Lin- forniaPubl. Bot. 10: 425. 1925; Gagnepain, naeus representsFlueggea acidoton (L.) Webster Fl. Indochine5: 408. 1926; AiryShaw, Kew (Allertonia3: 299. 1984). AlthoughGrisebach's Bull. 25: 544. 1971; Thin,Tap Chi Sinh Hoc description is indeed based on material of the genus 11(3): 15. 1989. TYPE: Strophioblachiafim- accepted by later authors as Acidocroton, by typ- bricalyxBoerl. ificationhis genus is a synonym of Flueggea. Con- In- servation of this genus that is common and wide- A genus of two species in southeastAsia: spread in the Greater Antilles would seem to be dochinato Hainan, the Philippines,and Celebes. the best solution. 239. Sagotia Baillon,Adansonia I. 1: 53. 1860 Although Ophellantha, described from Central (nom. cons.); Muell. Arg., DC. Prodr. 15(2): America, has been accepted as a distinct genus 1113. 1866; Fl. Bras. 11(2): 504. 1874; (e.g., by Standley & Steyermark, Fieldiana Bot. Bentham,Gen. P1. 3: 302. 1880; Pax, Pflan- 24(6): 139. 1949), it has never been compared zenr. 147. III (Heft47): 39. 1911; Jablonski, with Acidocroton. While the two taxa appear dif- Mem. New York Bot. Gard. 17: 151. 1967; ferentin aspect because of the reduced leaves and Secco, Acta Amazonica 15(1-2, suppl.): 81. greater spinosity of the West Indian plants, they 1985; Rev. Gen. Anomalocalyx ... Amer. are very similar in floral characters except for the Sul 99. 1990. TYPE: Sagotia racemosa Bail- more accrescent calyx in the Central American lon. plants. It therefore appears that Ophellantha is best treated as Acidocroton sect. Ophellantha Accordingto Secco (1985), Sagotia includes (Standl.) Webster, comb. nov. The only other spe- two species of AmazonianSouth America; Levin cies in the section besides the type is Acidocroton (pers. comm.) also reportsthe genus fromCosta steyermarkii (Standl.) Webster, comb. nov. Rica and Panama. The systematicposition of the [Ophellantha steyermarkii Standley, Field Mus. genus is ratherdoubtful; it shows some resem- Bot. 23: 123. 1944]. blances to Ostodes but also to Crotonand related The affinitiesof Acidocroton have been vari- genera.

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240. Baliospermum Blume, Bijdr. Fl. Ned. Ind. Sci. 41: 329. 1930; Jablonski, Brittonia 15: 603. 1826; Decaisne in Jacquemont, Voy. 151. 1963; Airy Shaw, Kew Bull. 23: 126. Inde Atlas 2: t. 155. 1844; Muell. Arg., DC. 1969; 25: 545. 1971; Whitmore, Tree Fl. Prodr. 15(2): 1125. 1866; Hook. f., Fl. Brit. Malaya 2: 134. 1973; Airy Shaw, Kew Bull. Ind. 5: 461. 1887; J. J. Smith, Meded. Dept. Add. Ser. 4: 201. 1975; Kew Bull. 35: 352. Landb. 10: 599. 1910; Pax & Hoffm.,Pflan- 1981. TYPE: Trigonostemon serratus Bi. zenr. 147. IV (Heft 52): 24. 1912; Gagne- EnchidiumJack, Malayan Misc. 2(7): 89. 1822 (nom. pain, Fl. Indochine 5: 429. 1926; Airy Shaw, rej.). TYPE: Enchidium verticillatumJack [= Tri- Kew Bull. 26: 222. 1972; 36: 267. 1981; gonostemonverticillatus (Jack) Pax]. Thin, Tap Chi Sinh Hoc 11(3): 16. 1989. Silvaea Hook. & Arn.,Bot. Beechey'sVoy. 211. 1837. TYPE: Baliospermumaxillare Bl. TYPE: Silvaea semperflorens(Roxb.) Hook. & Arn. [= Trigonostemonsemperfiorens (Roxb.) Muell. An Asian genus of ca. 12 species, distributed Arg.]. from India to China (Yunnan) and Java. Mueller AthroismaGriffith, Not. P1. As. 4: 477. 1854 (non A. P. DeCandolle,1833. TYPE: not designated(Griffith (1866) associatedBaliospermum with Gelonium, citedtwo species, neither positively identified by Pax which it somewhat resembles in habit, and has been & Hoffmann[1911]). followed in this by Pax & Hoffmann (1912) and TelogyneBaillon, Etude Gen. Euphorb.327. 1858. TYPE: other later workers. However, the nonaperturate Telogyneindica Baill. [= Trigonostemonverticil- latus (Jack)Pax]. pollen is quite differentfrom that of Suregada, TritaxisBaillon, Etude Gen. Euphorb.342. 1858. TYPE: and Airy Shaw (1975) seems closer to the mark Tritaxis gaudichaudii Baillon [= Trigonostemon in referringit to his "subtribe Codiaeinae" along gaudichaudii (Baill.) Muell. Arg.]. with Codiaeum, Blachia, and Strophioblachia. TylosepalumKurz ex Teysm.& Binn.,Natuurk. Tijdschr. Thin (1989) has described a new section, Dioicea, Ned.-Indie27: 50. 1864. TYPE: Tylosepalumau- rantiacumKurz ex Teysm. & Binn. [= Trigonos- on based Baliospermum balansae Gagnep. temonaurantiacus (Kurz ex Teysm.& Binn.)Boerl.]. NepenthandraSp. Moore,J. Bot. 43: 149. 1905. TYPE: Tribe 40. TRIGONOSTEMONEAE Webster, Nepenthandralanceolata Sp. Moore [= Trigonos- Taxon 24: 599. 1975. TYPE: Trigonostemon temonlanceolatus (Sp. Moore) Pax]. Bi. ProsartemaGagnepain, Bull. Soc. Bot. France 71: 875. 1925. TYPE: Prosartemastellaris Gagnep. [= Tri- Monoecious trees or shrubs; indumentum sim- gonostemonstellaris (Gagnep.) AiryShaw]. ple; leaves alternate, unlobed, pinnately veined, Poilaniella Gagnepain,Bull. Soc. Bot. France 72: 467. sometimesstipellate at base but lacking paired glands 1925. TYPE: Poilaniella ftagilis Gagnep. [= Tri- at apex of petiole; stipules persistentor deciduous, gonostemonftagilis (Gagnep.) AiryShaw]. NeotrigonostemonPax & Hoffm.,Notizbl. Bot. Gart. often small. Inflorescences terminal or axillary, ra- Berlin10: 385. 1928. TYPE: Neotrigonostemondi- cemose or thyrsoid. Staminate sepals 5, free, im- versijbliusPax & Hoffm.[= Trigonostemonvirn- bricate; petals 5, exceeding the calyx; disk cupular dissimus(Kurz) AiryShaw]. or dissected; stamens 3(-5), filamentsconnate, an- KurziodendronBalakrishnan, Bull. Bot. Surv. India 8: 68. 1966. TYPE: Kurziodendronviridissimum (Kurz) ther connective enlarged, often elongated; pollen Balakr.[= Trigonostemonviridissimus (Kurz) Airy grains globose, inaperturate, with Croton-pattern, Shaw]. sexinous processes small; pistillodeabsent. Pistillate sepals 5, free, imbricate; disk cupular; ovary A rather diverse genus of tropical Asia, extend- 3-locular; styles free or nearly so, unlobed or bifid. ing from India and Sri Lanka to New Guinea. The Fruit capsular; seeds ecarunculate, testa dry; en- diversity of the genus is clearly indicated by the dosperm present. remarkably extensive generic synonymy. The re- The tribe contains the single paleotropical genus lationships of the genus have been differentlyun- Trigonostemon. derstood. Mueller (1866), who accepted a broad circumscription (including Dimorphocalyx), re- 241. Trigonostemon Blume, Bijdr. Fl. Ned. ferred Trigonostemon to the Jatropheae. Pax & Ind. 600. 1825 (as Trigostemon);Fl. Javae Hoffmann(1911) included it in tribe Clutieae sub- Praef. viii. 1828 (nom. cons.); Muell. Arg., tribe Clutiinae, where it is clearly out of place. Airy DC. Prodr.15(2): 1105. 1866; Bentham,Gen. Shaw (1975) placed it in his informal "subtribe" PI. 3: 298. 1880; Hook. f., Fl. Brit. Ind. 5: Ostodinae along with Fahrenheitia and Dimor- 395. 1887; Pax & Hoffm.,Pflanzenr. 147. phocalyx. This placement has much to recommend III (Heft 47): 85. 1911; Gagnepain, Fl. In- it, and it is possible that the tribeTrigonostemoneae dochine 5: 309. 1925; Quisumbing, Phil. J. cannot be maintained as distinctfrom the Codiaeae.

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Tribe 41. RICINOCARPEAE Muell. Arg., Bot. An Australasiangroup of threegenera, two en- Zeit. 22: 324. 1864; DC. Prodr. 15(2): 199. demic to Australia. 1866. Subfam. Ricinocarpoideae Pax, Natiirl. Pflanzenfam. ed. 1, 3(5): 113. 1890. Family KEY TO THE GENERA OF SUBTRIBE RICINOCARPINAE Ricinocarpaceae (Muell. Arg.) Hurusawa, J. 1a. Stylesbifid to multifid;stamens at leastpartially Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: 224. connate. 1954. TYPE: Ricinocarpos Desf. 2a. Staminatecalyx lobed halfwayor more; filamentsall connate;petioles short. Monoecious (rarely dioecious) shrubs or herbs; ...... 242. Ricinocarpos latex scanty or apparently absent; indumentum 2b. Staminatecalyx lobed less than halfway; outerfilaments free; petioles long. stellate, glandular hairs often present; leaves al- ...... 243. A lphandia ternate, entire, often ericoid, eglandular, exstipu- lb. Stylesunlobed, dilated, forming a cap over the late. Inflorescences terminal or axillary, racemose ovary;stamens all free.. 244. Beyeria or reduced to clusters or solitaryflowers. Staminate sepals 4-6, free, imbricate, often petaloid; petals 242. Ricinocarpos Desfontaines,MWm. Mus. usually 5, or absent; disk dissected or absent; sta- Hist. Nat. Paris 3: 459, t. 22. 1817; Muell. mens mostly 15 or more, filamentsconnate into a Arg., DC. Prodr. 15(2): 203. 1866 (as Ri- column or short and apparently free; anthers ex- cinocarpus); Bentham & Mueller, Fl. Austral. trorse; pollen grains globose, inaperturate, sexine 6: 68. 1873; Bentham, Gen. P1. 3: 263. 1880; with modifiedCroton-pattern (processes spinose, or Maiden, Ill. N. S. W. P1., t. 16. 1908; Grin- reduced); pistillode absent. Pistillate sepals (4)5, ing, Pflanzenr. 147 (Heft 58): 37. 1913; Pax imbricate; petals 5, sometimes reduced or absent; & Hoffm., Natiirl. Pflanzenfam. ed. 2, 19c: disk dissected or absent; ovary 3-locular; styles 226. 1931; Stanley, Fl. S.-E. Queensland 1: unlobed, bifid, or multifid. Fruit capsular, 1-3- 432. 1983. TYPE: Ricinocarpos pinifolius locular; seeds carunculate; endosperm present; em- Desf. bryo cylindrical, with narrow cotyledons. Roeperia Sprengel,Syst. Veg. 3: 147. 1826. TYPE: Roe- As indicated by pollen characters, tribe Ricin- peria pinifolia (Desf.) Spreng. [= Ricinocarpos ocarpeae definitelybelongs in subfamily Crotono- pinijblius Spreng.]. ideae and not in a separate group of "Stenolobeae" An Australian genus of ca. 15 species grouped as treated by Mueller (1866), Bentham (1880), into 4 sections by Grining. Althoughreported from and Pax & Hoffmann (1931). The tribe appears New Caledonia, it does not occur there; the sup- rather closely related to the Crotoneae, and further posed New Caledonian species has been transferred work may show that the two groups should be to Baloghia by McPherson & Tirel (1987). combined.

KEY TO THE SUBTRIBESOF TRIBE RICINOCARPEAE 243. Alphandia Baillon, Adansonia I. 11: 85. 1873; Pax & Hoffmann, Pflanzenr. 147. III 1a. Pollen grains with large clavate sexinous processes; petals and disk usually present; ovary (Heft 47): 22. 1911; Airy Shaw, Kew Bull. 3-locular; leaves alternate - 41 a. Ricinocarpinae Add. Ser. 8: 27. 1980; McPherson & Tirel, lb. Pollen grains with reduced sexinous processes; Fl. Nouv.-Caled. 14(1): 86. 1987. TYPE: Al- petals and disk absent; ovary 1-3-locular; leaves phandia furfuracea Baillon. alternate, opposite, or whorled- 41 b. Bertyinae A Melanesian genus of three species in New Subtribe 41a. RICINOCARPINAE Webster, Guinea, New Caledonia, and Vanuatu. Taxon 24: 599. 1975. TYPE: Ricinocarpos Desf. 244. Beyeria Miquel,Ann. Sci. Nat. III. 1: 350, Trees, shrubs, or undershrubs; leaves alternate, t. 15. 1844; Muell. Arg., DC. Prodr. 15(2): sometimes resinous; flowersin terminal or axillary 201. 1866; Bentham& Mueller,Fl. Austral. racemes or clusters (or solitary); sepals distinct or 6: 63. 1873; Bentham,Gen. P1. 3: 263. 1880; connate; petals usually present, longer or shorter Grining,Pflanzenr. 147. (Heft58): 63. 1913; than sepals; stamens 20 or more, filaments free Pax & Hoffm.,Natiirl. Pflanzenfam. ed. 2, (or inner connate at base); pollen grains with large 19c: 227. 1931; James& Harden,Fl. N. S. clavate processes; ovary 3-locular; styles bifid or W. 1: 412. 1990. TYPE: Beyeria viscosa (La- unlobed; fruit capsular, with 3 seeds. bill.) Miq.

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CalyptrostigmaKlotzsch in Lehm., P1. Preiss. 1: 175. A monotypic genus restricted to northwestern 1845. TYPE: Calyptrostigmaviscosum (Labill.) K1. New Caledonia. [= Beyeria viscosa (Labill.) Miq.]. Beyeriopsis Muell. Arg., Linnaea 34: 56. 1865. TYPE: Beyeriopsisbrevifolia Muell. Arg. [= Beyeria brev- 247. Cocconerion Baillon, Adansonia 11: 87. ifolia(Muell. Arg.)Benth.; lectotype, designated by 1873; Guillaumin,Fl. Syn. Nouv.-Cal6d.189. Wheeler, 1975]. 1948; AiryShaw, Kew Bull. 25: 503. 1971; 382. 1978; McPherson& Tirel,Fl. Nouv.- A genus of ca. 15 species endemic to Australia. 32: Cal6d. 14(1): 38. 1987. TYPE: Cocconerion balansae Baillon. Subtribe 41b. BERTYINAE Muell. Arg., Lin- naea 34: 56. 1865; DC. Prodr. 15(2): 208. A genus of two species endemicto New Cale- 1866. TYPE: Bertya Planchon. donia. Trees or shrubs; leaves alternate, opposite, or 248. Borneodendron Airy Shaw, Kew Bull. verticillate; flowersin axillary racemes or clusters 16: 359. 1963; Hook. Ic. PI. 7(2): t. 3633. (or solitary); sepals distinctor absent; petals present Kew Bull. Add. Ser. 4: 60. 1975. TYPE: or absent; stamens numerous, filaments connate 1967; Borneodendron aenigmaticum Airy Shaw. into a column, anthersoften pubescent; pollen grains with sexinous processes reduced; ovary 2-3 loc- A monotypic genus endemic to Borneo. In the ular; styles bifidto multifid;fruit 1-3-seeded. original description, Airy Shaw (1963) compared Borneodendron to Baloghia, a genus that it some- This Australasian subtribe includes one genus what resembles in foliar characters. Later (Airy from Australia, two from New Caledonia, and one Shaw, 1975), he regarded it as most closely allied from Borneo. with Cocconerion, a disposition that is followed here. KEY TO THE GENERA OF SUBTRIBE BERTYINAE la. Staminatesepals present;petals present or absent; ovary 3-locular. Tribe 42. CROTONEAE Dumort., Anal. Fam. 2a. Leaves alternate;anthers glabrous; fruit PI. 45. 1829 ("Crotonieae"). TYPE: Croton L. usually 1-seeded 245. Bertya 2b. Leaves oppositeor whorled;anthers pu- Monoecious or dioecious trees, shrubs, or herbs; bescent;fruit 3-seeded. stems with nonarticulated laticifers; latex clear or 3a. Leaves opposite;inflorescences race- reddish, often scanty or apparently absent; indu- mose, terminal;styles multifid 246. Myricanthe mentum stellate or lepidote; leaves alternate, simple 3b. Leaves whorled; inflorescences to palmately lobed, sometimes pellucid-punctate; 1-flowered,axillary; styles 2-4-fid petiole withor withoutapical paired glands; stipules 247. Cocconerion present or absent. Inflorescences terminal or ax- lb. Staminatesepals absent; petals absent; ovary illary,racemose or spicate. Staminate sepals mostly 2-locular;staminate flowers in terminalbrac- teatecatkins, pistillate solitary, axillary; anthers 5, imbricate to valvate, free; petals 5, imbricate, pubescent;styles bifid 248. Borneodendron sometimes absent; disk annular or dissected, receptacle often villose; stamens 3-400, filaments 245. Bertya Planchon, Hook. Lond. J. Bot. 4: free, ofteninflexed in bud; anthers muticous; pollen 472. 1845; Muell. Arg., DC. Prodr. 15(2): grains globose, inaperturate, with sexinous Croton- 208. 1866; Bentham & Mueller, Fl. Austral. pattern; pistillodeabsent. Pistillatesepals 4-7(-10), 6: 74. 1873; Bentham, Gen. P1. 3: 264. 1880; + free, sometimes accrescent, imbricate to redu- Grining, Pflanzenr. 147. (Heft 58): 49. 1913; plicate-valvate; petals 5, mostly reduced or obso- Pax & Hoffm., Natiirl. Pflanzenfam. ed. 2, lete; ovary mostly 3-locular; styles free or nearly 19c: 226. 1931; Guymer, Austrobaileya 2: so, once to several times bifid.Fruit capsular; seeds 427. 1988; James & Harden, Fl. N. S. W. carunculate, testa dry; endosperm copious. 1: 414. 1990. TYPE: Bertya rosmarinifolia The circumscription of tribe Crotoneae is ex- Planchon [lectotype, chosen here]. panded from my previous treatment (Webster, A genus of ca. 20 species endemic to Australia. 1975), in which only Croton and the two satellite genera Crotonopsis and Eremocarpus were in- 246. Myricanthe AiryShaw, Kew Bull. 35: 390. cluded. The suggested inclusion of Mildbraedia, 1980; McPherson & Tirel, Fl. Nouv.-Cal6d. Fahrenheitia, and Moacroton in the tribe is still 14(1): 72. 1987. TYPE: Myricanthe discolor rather shakily supported and requires furthereval- Airy Shaw. uation, but at least has the merit of not leaving

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Crotonin a taxonomic"black box" withoutany As definedby Croizatand Alain, Moacrotonis apparentrelatives. a genusof six species endemicto Cuba. It appears closely relatedto Croton,and its status requires KEY TO THE GENERA OF TRIBE CROTONEAE evaluation(Webster, 1992). 1a. Filamentsnot distinctly inflexed inbud; pistillate flowerspetaliferous. 252. Croton L., Sp. P1. 2: 1004. 1753; Gen. 2a. Stamens10-30; staminatepetals thinner P1., ed. 5, 436. 1754; Klotzsch,Hook. Lon- intexture than sepals; pistillate flowers pet- don J. Bot. 2: 48. 1843; Muell. Arg., DC. aliferous;styles bifid. 3a. Stamensfree; staminate petals free; Prodr. 15(2): 512. 1866; Fl. Bras. 11(1): 81. inflorescencesaxillary; seed coat dry 1873; Bentham,Gen. P1. 3: 293. 1880; Hook. _...._..___..._._.._....._.._._.___._....._._...... _._._._. 249. Mildbraedia f., Fl. Brit.Ind. 5: 385. 1887; Ferguson,Rep. 3b. Stamenswith filaments connate; sta- MissouriBot. Gard. 12: 33. 1901; Hutchin- minatepetals coherent; inflorescences terminal;seed coat fleshy . son, Fl. Trop. Afr.6(1): 746. 1912; Gagne- ...... 250. Fahrenheitia pain, Fl. Indochine 5: 256. 1925; Pax & 2b. Stamens3-6; staminatepetals similar in Hoffm.,Natiirl. Pflanzenfam. ed. 2, 19c: 83. textureto sepals;pistillate flowers apetal- 1931; Leandri,Ann. Inst. Bot.-Geol.Colon. ous;styles unlobed or emarginate. Marseille,V. 7(1): 1. 1939; Carabia, Carib. .._...... ___. .._.._....__._...... - ....._. ... _.._..._. .__.._. 2 5 1. M oa cro to n lb. Filaments? distinctlyinflexed in bud; pistillate For. 3: 114. 1942; Croizat,Darwiniana 5: petalsmostly reduced to absent 252. Croton 417. 1941; 6: 442. 1944; Johnston& War- nock, Southw.Nat. 7: 1. 1962; Webster,J. ArnoldArbor. 48: 353. 1967; Leandri,Adan- 249. Mildbraedia Pax, Bot. Jahrb.43: 319. sonia II. 10: 191. 1970; Liogier,Fl. Espaiiola 1909; Pflanzenr.147. III (Heft47): 11. 1911; 4: 108. 1986; Thin,Tap Chi Sinh Hoc 8(2): Hutchinson,Fl. Trop. Afr.6(1): 798. 1912; 28. 1986; Webster,Ann. Missouri Bot. Gard. Leonard, Fl. Congo Belge 8(1): 85. 1962; 75: 1116. 1989; Novon 2: 270. 1992; Taxon Radcliffe-Smith,Fl. E. Trop. Afr.,Euphorb. 42: 793. 1993. TYPE: Croton aromaticus L. 1: 340. 1987. TYPE: Mildbraedia paniculata [lectotype, designated by Webster, J. Arnold Pax. Arbor. 48: 354. 1967; this replaced the ear- NeojatrophaPax, Pflanzenr.147. I (Heft42): 114. lier but incorrectselection of Crotontiglium 1910. TYPE: Neojatrophacarpinifolia (Pax) Pax L. by Small, in Britton& Brown,Ill. Fl. N. [= Mildbraediacarpinifolia (Pax) Hutch.]. U.S. ed. 2, 2: 254. 1913]. A tropicalAfrican genus of fourspecies. CiecaAdanson, Fam. P1. 2: 355. 1763 (nom.rej.). TYPE: Crotonargenteus L. CrotonopsisMichaux, Fl. Bor.-Amer.2: 185. 1803. Lep- 250. Fahrenheitia Reichb. f. & Zoll., Linnaea temonRaf., Med. Repos. II. 5: 353. 1808 (nom. 28: 599. 1856; Muell.Arg., DC. Prodr.15(2): superfl.).TYPE: Crotonopsislinearis Michaux [ Crotonmichauxii 1256. Kew Bull. 20: Webster]. 1866; AiryShaw, 409. Friesia Sprengel,Anleit. Kenntn. Gewachse, ed. 2, 2(2): 1966; 26: 270. 1972; Whitmore,Tree Fl. 885. 1818. TYPE: Friesia argentea Sprengel[= Malaya 2: 97. 1973; AiryShaw, Kew Bull. Crotonmichauxii Webster]. Add. Ser. 4: 114. 1975; Kew Bull. 36: 298. Decarinium Raf., Neogenyton1. 1825. TYPE: Decari- niumglandulosum (L.) Raf. [= Crotonglandulosus 1981. TYPE: Fahrenheitia collina Reichb. f. L.]. & Zoll. [= Fahrenheitia pendula (Hassk.) DrepadeniumRaf., Neogenyton 2. 1825. TYPE: Dre- AiryShaw]. padenium maritimum(Walt.) Raf. [= Crotonmar- itimusWalt.]. A tropicalAsian genus of four species, found Heptallon Raf., Neogenyton1. 1825. TYPE: Heptallon fromsouthern India and Sri Lanka to the Philip- graveolens Raf. [= Crotoncapitatus Michx.; lec- pines and Borneo. Pax & Hoffmann(1911) in- totype,chosen here]. cludedthe then-knownspecies within Ostodes, but JulocrotonMartius, Flora 20(2) Beibl.:119. 1837.TYPE: Julocrotonphagedaenicus Mart. [= Croton tri- AiryShaw (1966) showedthat Fahrenheitiawas queterLam.]. quitedistinct, although he later(Airy Shaw, 1975) AstrogyneBentham, PI. Hartweg.14. 1839. TYPE: As- referredit to the same informal"subtribe" as Os- trogynecrotonoides Benth. [= Crotondiocus Cav.]. todes. Astraea Klotzsch,Arch. Naturgesch. 7: 194. 1841. TYPE: Astraea lobata (L.) Ki. [= Crotonlobatus L.]. Cleodora Klotzsch,Arch. Naturgesch.7: 196. 1841. 251. Moacroton ArnoldArbor. 26: Croizat,J. TYPE: Cleodora sellowiana Ki. [= Crotonsphaero- 189. 1945; Alain,Fl. Cuba 3: 85. 1953. TYPE: gynus Baill.]. Moacroton leonis Croiz. Eutropia Klotzsch,Arch. Naturgesch.7: 196. 1841.

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TYPE: Eutropia brasiliensisKi., nom. illeg.[= Cro- The relationships of Croton within the family ton polyandrusSpreng.]. have been obscured by the fact that it has generally Medea Klotzsch,Arch. Naturgesch. 7: 193. 1841. TYPE: been referred to a separate tribe associated only Medea hirtaKlotzsch [= Crotontimandroides (Didr.) satellite genera Crotonopsis, Eremocar- Muell. Arg.]. with the Ocalia Klotzsch,Arch. Naturgesch. 7: 195. 1841. TYPE: pus, and Julocroton. However, these commonly Crotonperdicipes St. Hil. [= Crotonantisiphili- accepted segregate genera cannot be maintained ticusMart., lectotype, designated by Wheeler, 1975]. under a phylogenetic circumscription of Croton PilinophytumKlotzsch, Arch. Naturgesch. 7: 255. 1841. (Webster, 1992). The African genus Mildbraedia TYPE: Pilinophytumcapitatum (Michx.) Ki. [= Cro- ton capitatus Michx.]. seems to be a likely candidate for the sister group, PodostachysKlotzsch, Arch. Naturgesch. 7: 193. 1841. since it has in common characters of indumentum, TYPE: Podostachyssubfioccosa Didr. [= Crotonlun- flowers, and seeds, but it diverges in its axillary, (Didr.) dianus (Didr.) Muell. Arg. var. subfioccosa unisexual, more or less cymose inflorescences. The Muell. Arg.; lectotype,designated by Wheeler 3-nucleate pollen grains of Croton suggest a pos- (1975)]. Timandra Klotzsch,Arch. Naturgesch.7: 197. 1841. sible relationship with the tribe Ricinocarpeae. TYPE: Crotonserratus Muell. Arg. [lectotype,designatedby Wheeler(1975)]. Tribe 43. RICINODENDREAE (Pax) Hutch- HeterochlamysTurcz., Bull. Soc. Imp. Nat. Moscou 16: 61. 1843. TYPE: Heterochlamys quinquinervia inson, Amer. J. Bot. 56: 749. 1969. Cluytieae Turcz. [= Crotonargenteus L.]. subtribe Ricinodendrinae Pax, Natiirl. Pflan- Lasiogyne Klotzsch,Nov. Act. Acad. Caes. Leop.-Carol. zenfam. ed. 1, 3(5): 87. 1890. TYPE: Rici- 418. 1843. TYPE: Lasiogyne Nat. Cur. 19(suppl.)1: nodendron Muell. Arg. brasiliensisKl. [= Crotoncompressus Lam.]. Tiglium Klotzsch,Nov. Act. Acad. Caes. Leop.-Carol. Dioecious trees or shrubs; indumentum stellate; Nat. Cur. 19(suppl.)1: 418. 1843. TYPE: Tiglium leaves alternate, unlobed to lobed or palmatisect; officinaleKl. (nom. illeg.)[= Crotontiglium L.]. Eremocarpus Bentham,Bot. Voy. Sulphur53. 1844. stipules entire to lobed, or absent. Inflorescences Piscaria Piper, Contr.U.S. Nat. Herb. 11: 382. axillary or the pistillate terminal, paniculate or 1906 (nom.superfl.). TYPE: Eremocarpussetigerus reduced to clusters or solitary flowers. Staminate setigerusHook.]. (Hook.) Benth.[= Croton sepals 4 or 5, imbricate; petals 5, ? coherent into Angelandra Endlicher,Gen. PI. Suppl. 5: 91. 1850. a tube; disk dissected or lobed; stamens (3)5-20, TYPE: Crotonellipticus Nutt., non C. ellipticusGei- seler [= Crotonlindheimerianus Scheele]. filamentsunited below; pistillode absent. Pistillate Barhamia Klotzsch,in Seem., Bot. Voy. Herald 104. sepals 4 or 5, imbricate; petals 5, coherent; disk 1853. TYPE: Barhamia panamensis Kl. [= Croton lobed; ovary 1-3-locular; styles bifid. Fruit dru- hircinusVent.; lectotype,designated by Wheeler paceous, 1-3-seeded; seeds ecarunculate; endo- (1975)]. CyclostigmaKlotzsch, in Seem., Bot. Voy. Herald 104. sperm copious, oily. 1853. TYPE: Cyclostigmapanamensis Kl. [= Cro- This paleotropical tribe of two genera is entirely tondraco Schlecht.ssp. panamensis(Ki.) Webster; in designatedby Wheeler(1975)]. African or Madagascan except for one species GynamblosisTorrey, Rep. Marcy Exped. 295. 1853. India. Relationships of the tribe are uncertain. Gi- TYPE: Gynamblosismonanthogyna (Michx.) Torr. votia suggests Jatropha in its habit, but diverges monanthogynusMichx.]. [= Croton in its stellate indumentum and differentinflores- CentrandraKarsten, Linnaea 28: 440. 1857. TYPE: Cen- may be relat- trandra hondensis Karst. [= Croton hondensis cence; nevertheless, Ricinodendreae (Karst.) Webster]. ed to Jatropheae. MyriogomphosDidrichsen, Vidensk. Medd. Dansk Na- 142. 1857. TYPE: turh.Foren. Kjoebenhavn1857: KEY TO THE GENERA OF TRIBE RICINODENDREAE MyriogomphosffuscusDidr. [= Crotonffuscus(Didr.) Muell. Arg.]. la. Leaves unlobedto 5-lobed;stipules small, un- HeterocrotonSpencer Moore, Trans. Linn. Soc. II. 4: lobed 253. Givotia 461.1895. TYPE: Heterocrotonmentiens Sp. Moore lb. Leaves palmatelyparted. [= Crotonmentiens (Sp. Moore) Pax]. 2a. Stipulesflabellate-lobed, persistent; ovary mostly2-3-locular; endocarp smooth, thin- walled 254. Ricinodendron A large and highly diverse but definitelymono- 2b. Stipulessmall, unlobed, deciduous; ovary phyletic genus of at least 800 species, the majority mostly 1-locular;endocarp pitted,thick- American but also with more than 200 paleotrop- walled 255. Schinziophyton ical species. Only a selection of the more important references has been given here. The complexity of 253. Givotia Griffith,Calcutta J. Nat. Hist. 4: the genus is indicated by the 40 sections recently 88. 1843; Muell. Arg., DC. Prodr. 15(2): discriminated (Webster, 1993). 11 12. 1866; Bentham,Gen. P1. 3: 297. 1880;

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Pax & Hoffm., Pflanzenr. 147. III (Heft 47): TribeAleuritideae is circumscribedas previously 44. 191 1; Radcliffe-Smith,Kew Bull. 22: 493. (Webster,1975) to include5 subtribeswith a total 1968; Fl. Trop. E. Afr. Euphorb. 1: 329. of 15 generaand 45-50 species. It is overwhelm- 1987. TYPE: Givotia rottleriformisGriffith ex inglypaleotropical, with only two American genera. Wight. The tribeas presentlyconstituted is stillnot sat- isfactorilydefined. Bentham (1880) A genus of four species, one from Africa, two includedAleu- rites and othergenera of from Madagascar, and one from India. Aleuritideaewithin the Jatropheae.There are indeedmany points of sim- ilaritybetween Aleurites and 254. Ricinodendron Muell. Arg., Flora 47: Jatropha,and Ben- tham'sdisposition is much 533. 1864; DC. Prodr. 15(2): 1111. 1866; betterthan that of Pax & Hoffmann(1931), who Bentham,Hook. Ic. PI. 13: t. 1300. 1879; includedAleurites in a veryartificial subtribe along with Gen. PI. 3: 297. 1880; Pax, Pflanzenr.147. Agrostistachys. Joannesia has III (Heft 47): 45. 1911; Mildbraed,Notizbl. charactersthat suggestboth the Jatropheaeand Bot. Gart. Berlin 12: 516. 1935; Leonard, Aleuritideae,and it may provede- sirableto resurrect Fl. Congo 8(1): 116. 1962; Radcliffe-Smith, tribeJoannesieae (Pax, 1924) in orderto arriveat a Fl. E. Trop. Afr. Euphorb. 1: 325. 1987. morenatural classification. TYPE: Ricinodendron africanus Muell. Arg.

A genus of two African species, closely related KEY TO THE SUBTRIBES OF TRIBE ALEURITIDEAE to Givotia and apparently differingonly in vege- la. Monoecious. tative characters. 2a. Petals 5, glabrous;stamens 8-20, filamentsconnate; inflorescence paniculate; leavespalmately veined or lobed . 255. Schinziophyton Hutchinson ex Radcliffe- 44a. Aleuritinae Smith, Kew Bull. 45: 157. 1990. TYPE: Schin- 2b. Petals6-13, sericeous;stamens 30-100, ziophyton rautanenii (Schinz) Radcl.-Sm. filamentsfree; inflorescence a cymose clus- ter;leaves pinnately veined 44b. Garciinae A monotypic African genus, discriminated by l b. Dioecious. Hutchinson in an unpublished manuscript and 3a. Petalspresent, at leastin staminate flower. 4a. Indumentumsimple or characterized by Radcliffe-Smith absent;petals as distinct from free;inflorescence mostly terminal. Ricinodendron in a number of characters, includ- -...... 44c. Grosserinae ing its truly compound leaves, small stipules, and 4b. Indumentumstellate or lepidote;pet- large fruitswith deeply pitted endocarp. als oftencoherent or connate;inflo- rescenceterminal or axillary. 44d. Crotonogyninae 3b. Petalsabsent; indumentum stellate; leaves Tribe 44. ALEURITIDEAE Hurusawa, J. Fac. withbeadlike glands - 44e. Neoboutoninae Sci. Univ. Tokyo, Sect. 3, Bot. 6: 309. 1954. TYPE: Aleurites G. Forster. Subtribe 44a. ALEURITINAE Monoecious trees or shrubs; stems with nonar- (Hurusawa) Webster,Taxon 24: 599. 1975. TYPE: Aleu- ticulated laticifers, latex scanty or absent; indu- ritesJ. R. & G. Forster. mentum simple or stellate; leaves alternate, pinnately to palmately veined or lobed, eglandular or Monoecioustrees or shrubs;indumentum with laminar glands; stipules present or absent. simple or stellate;leaf blades palmatelyveined or lobed, Inflorescences terminal or axillary, dichasial-pa- with glands at apex of petiole; inflorescence? niculate or reduced to cymose clusters. Staminate paniculate;staminate calyx segments 2 or 3; petals calyx closed in bud, splittinginto valvate segments; 5, free,glabrous; disk-segments 5; stamens8-20, petals mostly 5(6-13), free, imbricate; disk dis- innerfilaments connate; ovary 2- or 3-locular;fruit sected or of intrastaminal segments; stamens 8- drupaceousor capsular; seeds ecarunculate. 20(- 100), filaments free; pollen grains globose, inaperturate,with Croton-pattern; pistillode absent. The subtribeAleuritinae is here equivalentto Pistillate sepals and petals as in staminate; disk the tribeAleuritideae of Hurusawa(1954), except lobed, dissected, or obsolete; ovary 2-5-locular; that the genus Aleuritesis treatedas subdivided styles bifid. Fruit drupaceous or capsular and tar- into three genera, followingAiry Shaw ('1967). dily dehiscent; seeds ecarunculate, testa dry; en- There is considerabledoubt whetherthese segre- dosperm copious, oily. gates need to be recognized,as I indicatedearlier

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(Webster, 1967); however, they are provisionally 1866; Pax & Hoffm., Pflanzenr. 147. XIV accepted here, pending furthercritical study. (Heft 68): 9. 1919. TYPE: Garcia Vahl in Rohr. KEY TO THE GENERA OF SUBTRIBE ALEURITINAE Monoecious trees or shrubs; indumentum sim- 1a. Ovary2-locular; fruit drupaceous; stamens 15- ple; leaves entire, pinnately veined, withoutglands 20, anthers introrse;indumentum distinctly at apex of petiole, exstipulate; inflorescences ter- stellate.256. Aleurites lb. Ovary3-5-locular; fruit dehiscent; stamens 7- minal, reduced to cymose clusters; petals 6-13, 12. sericeous; staminate disk dissected, intrastaminal; 2a. Indumentumstellate; anthers extrorse; in- stamens 30-100, filamentsfree; ovary 3-locular; florescencedensely pubescent, bracts con- styles bifid; fruitcapsular; seeds ecarunculate. spicuous 257. Reutealis 2b. Indumentumof bifidor simplehairs; an- Although Mueller (1866) included three other thers introrse;inflorescence not densely genera (Aleurites, Crotonogyne, and Mannioph- pubescent,bracts inconspicuous . yton) in his subtribe Garciinae, Pax & Hoffmann ...... 258. Vernicia (1919) removed them. In the treatmentof Webster (1975), this monogeneric delimitationwas followed, 256. Aleurites J. R. & G. Forster,Charact. and it is still accepted here. Gen. PI. 111, t. 56. 1776; Muell. Arg., DC. Prodr. 15(2): 722. 1866; Bentham, Gen. PI. 259. Garcia Vahl in Rohr, Skriv. Naturh.-Selsk. 3: 292. 1880; Hemsley, Hook. Ic. PI. 29: t. Kjobenh. 2: 217, t. 9. 1792; Muell. Arg., 2801, 2802. 1909; Pax, Pflanzenr.147. I DC. Prodr. 15(2): 721; Lundell, Wrightia 1: (Heft 42): 128. 1910; Gagnepain,Fl. Indo- 1. 1945; Webster, Ann. Missouri Bot. Gard. chine 5: 290. 1925; AiryShaw, Kew Bull. 54: 238. 1968. TYPE: Garcia nutans Vahl ex 20: 393. 1967; Webster,J. ArnoldArbor. Rohr. 48: 342. 1967; Walker, Fl. Okinawa 644. 1976; Smith,Fl. VitiensisNova 2: 547. 1981; A neotropical genus of two species distributed Radcliffe-Smith,Fl. E. Trop. Afr.,Euphorb. from Mexico to Colombia. 1: 176. 1987. TYPE: Aleurites triloba J. R. & G. Forster [= Aleurites moluccana (L.) Subtribe 44c. GROSSERINAE Webster,Tax- Willd.]. on 24: 600. 1975. TYPE: Grossera Pax.

In the restricted sense adopted by Airy Shaw Dioecious trees or shrubs; indumentum simple (1967), Aleurites is a genus of two species ex- or absent; leaves pinnately veined or triplinerved, tending from India to the Pacific islands. glandular or eglandular; stipules deciduous or absent; inflorescences terminal, racemose or panic- 257. Reutealis Airy Shaw, Kew Bull. 20: 394. ulate; staminate petals 4 or 5, free; stamens 6- 1967. TYPE: Reutealis trisperma (Blanco) Airy 40, filamentsfree or connate; pistillate petals 4 or Shaw. 5, free; ovary 3-5-locular; styles bifid; fruitcapsular; seeds ecarunculate. A monotypic genus endemic to the Philippines; more or less intermediatebetween Aleurites (sensu The seven genera of this subtribeare American/ stricto) and Vernicia. African in distribution,except for Tapoides from Indonesia. 258. Vernicia Loureiro, Fl. Cochinch. 586. KEY TO THE GENERA OF 1790; Hemsley, Hook. Ic. PI. 29: t. 2801, SUBTRIBE GROSSERINAE 2802. 1906 (under Aleurites); Airy Shaw, la. Sexinousprocesses of pollen rounded to obtuse; leaves pinnatelyveined. Kew Bull. 20: 394. 1967; Radcliffe-Smith, 2a. Staminatepetals glabrous; leaves pellucid- Fl. E. Trop. Afr., Euphorb. 1: 178. 1987. punctate. TYPE: Vernicia montana Lour. 3a. Bracts large, imbricate, caducous, forminga cone beforeanthesis; leaves An Asian genus of three species distributedfrom entire;stipular scars subannular,con- Burma to Indochina, China, and Japan; found in spicuous;pistillate sepals unthickened more temperate latitudes or elevations than Aleu- ...... 260. Cavacoa rites and Reutealis. 3b. Bracts small,persistent, not forming a cone beforeanthesis; leaves denticulate; stipulescars verysmall; pistil- Subtribe 44b. GARCIINAE Muell. Arg., Lin- late sepals mediallythickened. naea 34: 143. 1865; DC. Prodr. 15(2): 719...... 261. Grossera

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2b. Staminatepetals pubescent; leaves notpel- and it possibly belongs close to Dodecastigma and lucid-punctate. Sagotia. 4a. Inflorescencesaxillary; seeds ecarunculate; staminatedisk annular. 5a. Stamens6-8; capsuletomentose 264. Sandwithia Lanjouw, Kew Bull. 1932: ...... 262. Tapoides 184. 1933; Secco, Bull. Mus. Par. Emilio 5b. Stamens over 20; capsule gla- Goeldi Bot. 3: 157. 1987; 4: 177. 1988. brous. 263. Anomalocalyx TYPE: Sandwithia guyanensis Lanj. 4b. Inflorescencesterminal; seeds carunculate; staminatedisk dissected;sta- A genus of two species of the Guyana region capsule tomentose.. mensover 20; and Amazonian Brazil and Venezuela. Secco (1988) ...... 264. Sandwithia genus, lb. Sexinous processes of pollen echinate; leaves regarded Sagotia as the most closely related triplinerved;staminate disk dissected. an opinion that needs careful consideration. Sand- 6a. Pistillatesepals 4 or 5, imbricate;stami- withia is here provisionally retained in subtribe nate receptacleglabrous ...... 265. Tannodia Grosserinae because of its resemblance to Anom- 6b. Pistillatesepals 2 or 3; staminaterecep- alocalyx, but it is quite possible that all three tacle pilose ...... 266. Domohinea genera may prove to belong to the same tribe or subtribe. 260. Cavacoa Leonard, Bull. Jard. Bot. Brux. 25: 320. 1955; Fl. Congo Belge 8(1): 191. 1962; Elffers & Taylor, Hook. Ic. P1. 36: t. 265. Tannodia Baillon, Adansonia I. 1: 251. 3561. 1956; Radcliffe-Smith, Fl. E. Trop. 1861; Muell. Arg., DC. Prodr. 15(2): 728. Afr., Euphorb. 1: 174. 1987. Grossera sect. 1866; Bentham, Gen. P1. 3: 304. 1880; Pax Racemiformes Pax & Hoffm.,Pflanzenr. 147. & Hoffm.,Pflanzenr. 147. VI (Heft 57): 110. 50: 125. 1912; Leonard, VI (Heft 57): 105. 1912. TYPE: Cavacoa 1912; Prain, J. Bot. Fl. quintasii (Pax & Hoffm.) Leonard [Grossera Bull. Jard. Bot. Brux. 25: 300. 1955; Fl. quintasii Pax & Hoffm.]. Congo 8(1): 186. 1962; Radcliffe-Smith, E. Trop. Afr., Euphorb. 1: 172. 1987. Tan- A genus of three species of central and east donia Baillon, Adansonia I. 1: 184. 1861 (non Africa. Tandonia Moq.-Tand., 1849). TYPE: Tanno- dia cordifolia (Baill.) Baill. [Tandonia cor- 261. Grossera Pax, Bot. Jahrb. 33: 281. 1903; difolia Baill.]. Pflanzenr. 147. VI (Heft 57): Pax & Hoffm., Holstia Pax, Bot. Jahrb.43: 220. 1909. Tannodia sect. 105. 1912; Cavaco, Bull. Mus. Hist. Nat. Holstia (Pax) Prain, J. Bot. 50: 127. 1912. Neo- Paris 21: 272. 1949; Leonard, Bull. Jard. holstia Rauschert,Taxon 31: 559. 1982 (nom. Bot. Brux. 25: 316. 1955; Bull. Jard. Bot. superfl.).TYPE: Holstia tenuifoliaPax [= Tannodia (Pax) Prain; lectotype]. Brux. 28: 118. 1958; Fl. Congo 8(1): 188. tenuijolia 1962. TYPE: Grossera paniculata Pax [lec- A genus of three or four species in east Africa totype, designated by Leonard, 1955]. and the Comoro Islands. Although Radcliffe-Smith maintained Holstia (as Neoholstia) as a An African genus of seven species. (1987a) distinct genus, it differsonly in a relatively trivial perianth character, and Prain's reduction of it to 262. Tapoides Airy Shaw, Kew Bull. 14: 473. a section of Tannodia appears justified. 1960; 20: 412. 1966; Hook. Ic. P1. 37: t. 3632. 1967; Kew Bull. Add. Ser. 4: 200. 266. Domohinea Leandri, Bull. Soc. Bot. 1975. TYPE: Tapoides villamilii (Merr.) Airy France 87: 285. 1940. TYPE: Domohinea per- Shaw [Ostodes villamilii Merr.]. rieri Leandri.

A monotypic genus of Borneo. A monotypic genus from Madagascar. In de- scribing it, Leandri noted its affinityto Grossera 263. Anomalocalyx Ducke, Notizbl. Bot. Gart. and Tannodia, and this dispositionappears correct. Berlin 11: 344. 1932; Arq. Jard. Bot. Rio Janeiro 6: 60. 1933; Secco, Rev. Gen. Anom- Subtribe 44d. CROTONOGYNINAE Web- alocalyx . . . Amer. Sul 39. 1990. TYPE: An- ster, Taxon 24: 600. 1975. TYPE: Crotono- omalocalyx uleanus (Pax) Ducke. gyne Muell. Arg.

A monotypic genus of Amazonian Brazil. The Trees or shrubs, sometimes scandent; indumen- phylogenetic position of Anomalocalyx is dubious, tum stellate or lepidote; leaves pinnately or pal-

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mately veined, biglandular at juncture withpetiole, 269. Manniophyton Muell. Arg., Flora 47: stipulate; inflorescences axillary, spicate or race- 530. 1864; DC. Prodr. 15(2): 719. 1866; mose to paniculate; staminate petals coherent or Bentham, Hook. Ic. P1. 13: t. 1267. 1878; connate; staminate disk dissected; stamens 7-40, Gen. P1. 3: 297. 1880; Pax & Hoffm., Pflan- filamentsfree or connate; anthers often apiculate; zenr. 147. VI (Heft 57): 120. 1912; Keay, pistillate sepals 4 or 5, imbricate or valvate; disk Fl. W. Trop. Afr., ed. 2, 1: 400. 1954; Leon- cupular; ovary 3-locular; styles bifid or multifid; ard, Bull. Jard. Bot. Brux. 25: 290. 1955; fruitcapsular; seeds ecarunculate. Fl. Congo 8(1): 171. 1962; Adam, MWm.Mus. An African tribe of three genera. Crotonogyne Hist. Nat. Paris 20: 495, t. 183. 1971. TYPE: and Manniophyton were originallyreferred to sub- Manniophyton africanum Muell. Arg. tribe Garciinae by Mueller (1866); however, they A monotypic genus of west Africa (Sierra Leone differfrom Garcia in having stellate indumentum, to Angola and Principe). dioecious inflorescences, and fewer petals that are more or less coherent. The Crotonogyninae seem Subtribe 44e. NEOBOUTONINAE (Hutch- most closely related to the Grosserinae. inson) Webster, Taxon 24: 600. 1975. Aca- lypheae subtribe Mercurialinae ser. Neobou- KEY TO THE GENERA OF SUBTRIBE CROTONOGYNINAE toniiformes Pax & Hoffm., Pflanzenr. 147. 1a. Trees or shrubs;leaves pinnatelyveined; peti- VII (Heft 63): 71. 1914. Tribe Neoboutonieae oles lackinginflated trichomes. Hutch., Amer. J. Bot. 56: 752. 1969. TYPE: 2a. Stylesbifid; inflorescences terminal, panic- Neoboutonia Muell. Arg. ulate;staminate petals free; staminate disk receptacularand extrastaminal,of more Tribe BenoistieaeRadcliffe-Smith, Kew Bull. 43: 632. than 10 ...... segments 267. Cyrtogonone 1988. TYPE: Benoistia Perrier& Leandri. 2b. Stylesmultifid; inflorescences axillary, racemoseor spicate;staminate petals mostly Dioecious trees or shrubs; indumentum stellate coherentor staminate connate; disk ex- or lepidote; leaves unlobed, pinnately or palmately trastaminal,of 5-8 segments ...... 268. Crotonogyne veined, glandular-dotted, stipulate; inflorescences lb. Lianas; leaves palmatelyveined; petioleswith terminalor axillary, racemose to paniculate; calyx- inflatedtrichomes; styles bifid; staminate petals segments 2 or 3; disk dissected or absent; stamens connate.269. Manniophyton 15-40, filaments free, shorter than anthers; anthers with glandular connective; ovary 3-locular; 267. Cyrtogonone Prain,Kew Bull. 1911: 231. styles bifid; fruit capsular; seeds carunculate or 1911; Pax & Hoffm.,Pflanzenr. 147. VI (Heft ecarunculate. 57): 111. 1912; Prain, Hook. Ic. P1. 31: t. 3008. 1915; Keay, Fl. W. Trop. Afr., ed. 2, A subtribe of two genera, one African and one 1: 399. 1958. TYPE: Cyrtogonone argentea Madagascan. The two genera are quite distinct, (Pax) Prain [Crotonogyne argentea Pax]. and Radcliffe-Smith(1988) has proposed to place Benoistia in a separate tribe. However, Neobou- A monotypicgenus of west Africa(Nigeria to tonia and Benoistia stillseem more closely related Cameroon). to one another than to other taxa, so they are provisionally kept in the same subtribe. 268. Crotonogyne Muell. Arg., Flora 47: 535.

1864; DC. Prodr. 15(2): 720. 1866; Ben- KEY TO THE GENERA OF SUBTRIBE NEOBOUTONINAE tham,Gen. P1. 3: 305. 1880; Pax & Hoffm., 1a. Leaves palmatelyveined; stipulespersistent; Pflanzenr.147. VI (Heft 57): 111. 1912; seeds carunculate.270. Neoboutonia Prain,Hook. Ic. P1. 31: t. 3019. 1915; Leon- l b. Leaves pinnatelyveined; stipules deciduous; ard, Fl. Congo 8(1): 174. 1962; Adam, Fl. seeds ecarunculate 271. Benoistia Mt. Nimba 468, t. 165. 1971. TYPE: Croto- nogyne manniana Muell. Arg. 270. Neoboutonia Muell. Arg., J. Bot. 2: 336. NeomanniophytonPax & Hoffm.,Pflanzenr. 147. VI 1864; DC. Prodr. 15(2): 892. 1866; Ben- (Heft57): 115. 1912. TYPE: Neomanniophytonim- tham, Hook. Ic. P1. 13: t. 1298, 1299. 1879; peditum (Prain) Pax [= Crotonogyneimpedita Gen. P1. 3: 317. Pax & Pflan- Prain; lectotype,chosen here]. 1880; Hoffm., zenr. 147. VII (Heft 63): 71. 1914; Robyns, A genus of 15 species in west Africa(Sierra Fl. Sperm. Parc Nat. Albert 1: 451, t. 44. Leone to Congo and Angola). 1948; Radcliffe-Smith,Fl. E. Trop. Afr., Eu-

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phorb. 1: 231. 1987. TYPE: Neoboutonia af lb. Pollen sexine perforate-tectate;inflorescences ricana Muell. Arg. mostlyracemose, spicate, or pseudanthial;bracts mostlyglandular or adnate to rachis; stamens A genus of three African species. 1-many, free or connate; monoecious(rarely dioecious)trees, shrubs or herbs,not scandent. 2a. Inflorescencesmostly 271. Benoistia Perrier & Leandri, Bull. Soc. racemoseor spicate (if capitate,then not pseudanthial);styles Bot. France 85: 528. 1938; Radcliffe-Smith, mostlyundivided; staminate calyx usually Kew Bull. 43: 632. 1988. TYPE: Benoistia developed. perrieri Leandri [lectotype, chosen here]. 3a. Floral bracts usually biglandularat base, not peltate nor adnate to the A Madagascan genus of three species. rachis;stamens free or connate. 4a. Staminate calyx open in bud; leaves mostlynot spinulose-den- Subfamily V. EUPHORBIOIDEAE. TYPE: tate; stamens free or filaments Euphorbia L. connate; seeds carunculate or ecarunculate 46. HIPPOMANEAE SubfamilySapioideae Hurusawa, J. Fac. Sci. Univ.Tokyo, 4b. Staminatecalyx closed in bud, Sect. 3, Bot.6: 310. 1954. TYPE: Sapium P. Browne. valvatelydehiscent into 2 segments;leaves spinose-dentate;fil- Monoecious or less commonly dioecious trees, aments and anthers connate; shrubs, or herbs (rarely scandent); laticifers non- seeds ecarunculate articulated, latex usually whitish,sometimes scanty 47. PACHYSTROMATEAE or apparently absent; indumentumsimple or absent 3b. Floralbracts eglandular, peltate or adnate to the rachisand coveringflow- (dendritic in Mabea); leaves alternate or opposite, ers; seeds ecarunculate 48. HUREAE simple and unlobed, entire or dentate, usually pin- 2b. Inflorescencespseudanthial (cyathial), usu- nately veined, often with glands at base of blade; ally witha singleterminal pistillate flower stipules usually small, often reduced or absent. and 4 or 5 lateral staminatemonochasia or dichasia; perianth Inflorescences terminalor axillary, spicate to race- reduced or absent; stylesmostly bifid; seeds carunculateor mose, paniculate, or pseudanthial (cyathium); bracts ecarunculate 49. EUPHORBIEAE often biglandular at base; flowers apetalous. Sta- minate sepals 3-6, imbricate to valvate, mostly open in bud, commonly reduced or obsolete; disk Tribe 45. STOMATOCALYCEAE (Muell. absent; stamens 1-20(-70), filamentsfree or con- Arg.) Webster,Taxon 24: 600. 1975. Hip- nate; pollen grains oblate to prolate, 3-colporate, pomaneae subtribeStomatocalyceae Muell. colpi usually marginate, sexine usually tectate-per- Arg., Linnaea 34: 202. 1865. TYPE: Stoma- forate; pistillode absent. Pistillate sepals 3-6, free tocalyx Muell. Arg. [= Pimelodendron or connate, imbricate or open in bud, sometimes Hassk.]. reduced or obsolete; disk absent; ovary 2-3(-20)- Dioecious trees,shrubs, or lianas; latex yellow- locular; styles nearly always unlobed, free or com- ish,often scanty; indumentum usually absent; leaves monly connate into a column. Fruit capsular (rarely alternate,unlobed, pinnately veined, without glands drupaceous); seeds carunculate or ecarunculate, at base of blade; stipulessmall and caducous or testa dry or fleshy; endosperm copious. absent. Inflorescencesaxillary, racemose, bracts The circumscription of subfamily Euphorbioi- eglandular.Staminate sepals 4-8, freeand imbri- deae and its five tribes is essentially the same as cate or connateand calyx 2-lipped;stamens 10- that adopted earlier (Webster, 1975). It corre- 30, free, filamentsshort; pollen grains coarsely sponds roughly to the tribes Hippomaneae and reticulateor reticulate-perforate.Pistillate sepals Euphorbieae in the sense of Pax & Hoffmann 5 or 6, freeor connateand 2-lipped;ovary 2-10- (1931), except that the tribe Stomatocalyceae (in- locular; stylesunlobed, sometimes dilated or stig- cluded in the Gelonieae by Pax & Hoffmann) is matiform.Fruit capsular or indehiscent,1-3-seed- added. ed; seeds ecarunculate;endosperm copious, oily. This tribeincludes one neotropicaland three KEY TO THE TRIBES OF SUBFAMILY EUPHORBIOIDEAE paleotropicalgenera, distributed into two subtribes. 1a. Pollen sexine reticulate;inflorescences race- Some characteristicswithin the Stomatocalyceae, mose or paniculate,bracts eglandular; stamens 10-30, free;staminate calyx 3-8-lobed;styles such as the liana habit, colored latex, and oily undivided;seeds ecarunculate;dioecious trees endosperm,suggest possible affinity with Ompha- or lianas 45. STOMATOCALYCEAE lea, in the Acalyphoideae.

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KEY TO THE SUBTRIBESOF TRIBE STOMATOCALYCEAE 45b. Subtribe HAMILCOINAE Pax & Hoff- 1a. Pollen grainsfinely reticulate-perforate; fruits mann, Pflanzenr.147. VII (Heft 63): 419. indehiscent;styles abbreviated; ovary 2-1 0-loc- 1914. TYPE: Hamilcoa Prain. ular.45a. Stomatocalycinae lb. Pollen grainscoarsely reticulate; fruits capsu- Trees or lianas; leaves long-petiolate, entire or lar; styles? elongated;ovary 2-3-locular. crenate; racemes simple or branched; staminate ...... 45b. Hamilcoinae sepals 4 or 5, distinct,imbricate; stamens 10-20, free, filaments very short; pollen grains coarsely 45a. Subtribe STOMATOCALYCINAE Muell. reticulate; pistillate sepals 4-6, + distinct; ovary Arg., Linnaea 34: 202. 1865; DC. Prodr. 2- or 3-locular; styles connate below, erect or 15(2): 1142. 1866. TYPE: Stomatocalyx spreading; fruitcapsular. Muell. Arg. [= Pimelodendron Hassk.]. A group of two genera, one American and one Trees or shrubs; leaves entire or dentate; ra- African. Although included in two differentsub- cemes simple or slightlybranched; sepals distinct tribesof the Gelonieae by Pax & Hoffmann(1931), or connate; stamens 10-20; pollen grains finely Hamilcoa and Nealchornea share similar distinc- reticulate; ovary 1-1 0-locular; styles stigmatiform; tively coarse-reticulate pollen grains that are cu- fruitindehiscent, 1-seeded. riously similar in ornamentation to those of Dal- This subtribeincludes two genera, one of In- echampia. donesia,the otherof Africa. KEY TO THE GENERAOF SUBTRIBE HAMILCOINAE KEY TO THE GENERAOF SUBTRIBE STOMATOCALYCINAE la. Ovary3-locular; styles erect; stamens 18-20; la. Sepals 6-8, free;ovary 1-locular. pistillatesepals not glandular; stems ? scandent ...... Plagi.P ostyles 274. Hamilcoa lb. Sepals connate,staminate calyx 2-lipped; ovary lb. Ovary2-locular; styles spreading; stamens 10; 2-10-locular 273. Pimelodendron pistillatesepals with large basal glands; stems notscandent .275. Nealchornea 272. Plagiostyles Pierre,Bull. Mens. Soc. Lmin. Paris 2: 1326. 1897; Prain, Kew Bull. 1912: 274. Hamilcoa Prain, Kew Bull. 1912: 107. 107. 1912; Fl. Trop. Afr.6(1): 1001. 1913; 1912; Fl. Trop. Afr. 6(1): 1000. 1913; Pax Pax & Hoffm.,Pflanzenr. 147. VII (Heft63): & Hoffm.,Pflanzenr. 147. VII (Heft 63): 419. 420. 1914; Stapf,Hook. Ic. PI. 31: t. 3010. 1914; Stapf,Hook. Ic. P1. 31: t. 3009. 1915; 1915; Leonard,Fl. Congo 8(1): 131. 1962. Keay, Fl. W. Trop. Afr.,ed. 2, 1: 413. 1958. TYPE: Plagiostyles klaineana Pierre [= Pla- TYPE: Hamilcoa zenkeri (Pax) Prain [Plu- giostyles africana (Muell. Arg.) Prain]. kenetia zenkeri Pax]. A monotypicgenus of west Africa (southern A monotypic genus of west African rainforests Nigeriato Gabon and Congo). (Cameroon).

273. Pimelodendron Hasskarl, Versl. Med. 275. Nealchornea Huber, Bol. Mus. Goeldi 7: Afd.Natuurk. Kon. Akad. Wetensch.4: 140. 297. 1913; Pax & Hoffm.,Pflanzenr. 147. 1856; Benth.,Gen. PI. 3: 331. 1880; Pax & XIV (Heft68): 51. 1919; Ducke, Arch.Jard. Hoffm.,Pflanzenr. 147. V (Heft52): 54. 1912; Bot. Rio de Janeiro4: 107. 1925; Macbride, J. J. Smith,Bull. Jard.Bot. Buit. III. 6: 100. Field Mus. Nat. Hist., Bot. 13(IIIA, 1): 180. 1924; Whitmore,Tree Fl. Malaya 2: 124. 1951. TYPE: Nealchornea yapurensis Huber. 1973; AiryShaw, Kew Bull.Add. Ser. 4: 186. A monotypicgenus of the Amazonian rainforests 8: 196. 1980; Kew Bull. 36: 339. 1975; (eastern Colombia and Peru to Brazil). 1981. TYPE: Pimelodendron amboinicum Hassk. Tribe 46. HIPPOMANEAE A. Jussieu ex StomatocalyxMuell. Arg., DC. Prodr.15(2): 1142. 1866. Spach, Hist. Nat. Veg. 2: 522. 1834. TYPE: TYPE: Stomatocalyxgriflithianus Muell. Arg. [= Pimelodendrongriflithianum (Muell. Arg.) Benth. Hippomane L. ex Hook. f.]. Monoecious (less commonly dioecious) trees, An Australasian genus of six to eight species shrubs, or herbs; latex usually milky, commonly distributedfrom Malaya to New Guinea, tropical acrid or toxic; indumentum simple or absent (den- Australia(Queensland), and the SolomonIslands. driticin Mabea); leaves alternate (rarely opposite),

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simple and unlobed, pinnately veined (rarely pal- dular; staminate sepals 3-6, imbricate; stamens 2- mately veined), commonly withlaminar or petiolar 70, free from an expanded receptacle or connate glands; stipules conspicuous to reduced or absent. into a column; pistillate sepals 3-6, sometimes Inflorescences terminal or axillary, racemose or glandular; ovary 3-locular, exappendiculate; styles spicate (sometimespaniculate), usually bisexual with connate into a column, tips slender and unlobed; one or a few pistillateflowers at base; bracts mostly fruitcapsular; seeds carunculate. glandular. Staminate calyx open in bud (exposing This neotropical subtribe includes three genera anthers), sepals distinct or connate, imbricate to as conventionally defined. valvate, sometimes vestigial or absent; stamens 1- 70, free or filaments connate below; anthers dis- KEY TO THE GENERA OF SUBTRIBE MABEINAE crete, exrorse; pollen grains 3-colporate, colpi usu- columnelongate; style tips slender,un- ally marginate, sexine perforate-tectate. Pistillate la. Stylar lobed; inflorescencerachis ? flexuous;stami- sepals 3-6, imbricate, sometimes reduced or ab- nate flowerslong-pedicellate; stamens (3-)10- sent; ovary 2-3-locular (rarely 4-10); styles free 70; leaves and stemswith dendritic hairs...... or connate, unlobed. Fruit capsular or drupaceous; ...... 276. M abea seeds carunculate or ecarunculate, testa dry or lb. Stylar column short,style tips rather thick; inflorescencerachis rigid; stamens 3-12. fleshy; endosperm copious, not markedly oily. 2a. Latex veryscanty or absent;indumentum This diverse complex of 400-500 species was absent;stamens 5-12; leaves chartaceous ...... 277. Senefeldera subdivided by Pax & Hoffmann(1912) into four 2b. Latex copious,milky; indumentum of den- tribes, but these are so poorly defined that it seems dritichairs present on inflorescenceaxes; impossible to maintain them. Generic delimitations stamens2 or 3; leaves rigid...... are uncertain and controversial, and may change ...... 278. Senefelderopsis drastically when a monographic revision is completed. 276. Mabea Aublet, Hist. P1. Guyane Fr. 2: 867, t. 334. 1775; Muell. Arg., DC. Prodr.

KEY TO THE SUBTRIBES OF TRIBE HIPPOMANEAE 15(2): 1148. 1866; Fl. Bras. 11(2): 515. 1874; Bentham, Gen. P1. 3: 363. 1880; Pax 1a. Staminatereceptacle convex or conical; staminatesepals 3-6, imbricate;stamens mostly & Hoffm., Pflanzenr. 147. V (Heft 52): 26. 6-70, free; seeds carunculate;inflorescences 1912; Jablonski, Mem. New York Bot. 17: terminal,paniculate; leaves stipulate,without 164. 1967; den Hollander & Berg, Proc. Kon. petiolarglands ...... 46a. Mabeinae Ned. Akad. Wet. 89C: 147. 1986; Huft, Phy- lb. Staminatereceptacle plane; staminatesepals 62: 339. 1987; Ann. Missouri Bot. mostly2 or 3, sometimesonly 1 or absent; tologia stamens1-6(-20), filamentsfree or united;in- Gard. 75: 1125. 1989. TYPE: Mabea piriri florescencesterminal or axillary,racemose or Aubl. [lectotype, designated by Pfeiffer,No- spicate(axes not compound);leaves oftenwith mencl. Bot. 2(1): 191. 1874]. petiolarglands. 2a. Staminatecalyx laterally compressed; sta- A diverse and difficultgenus of ca. 50 neotrop- mens 5-20, free; leaves with petiolar ical species. glands,stipules large and deciduous;latex watery;seeds carunculate- 46b. Carumbiinae 2b. Staminatecalyx not laterallycompressed; 277. Senefeldera Martius, Flora 24 (Beibl. 2): stamens1-20, filamentsfree or connate; 29. 1841; Muell. Arg., DC. Prodr. 15(2): leaves withor withoutpetiolar glands, stip- 1153. 1866; Fl. Bras. 11(2): 529. 1874; ules small or absent; latex usuallymilky; Bentham, Gen. P1. 3: 332. 1880; Pax & seeds carunculateor ecarunculate. 23...... 46c. Hippomaninae Hoffm.,Pflanzenr. 147. V (Heft 52): 1914; Jablonski, Mem. New York Bot. Gard. 12: 171. 1965; Webster, Ann. Missouri Bot. Gard. Subtribe 46a. MABEINAE Pax & Hoffmann, 75: 1127. 1989. TYPE: Senefeldera multi- Pflanzenr.147. V (Heft52): 22. 1912. TYPE: flora Mart. Mabea Aubl. A neotropical genus of 8-10 species, mainly in Monoecious trees or shrubs; latex milky, innoc- Amazonian South America but reaching north to uous; indumentum of dendritic hairs, or absent; Panama. leaves alternate, pinnately veined, withoutlaminar or petiolar glands, stipulate; inflorescences termi- 278. Senefelderopsis Steyermark, Bot. Mus. nal, bisexual, racemose or paniculate, bracts biglan- Leafl. Harvard Univ. 15: 45. 1951; Jablonski,

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Mem. New York Bot. Gard. 12: 174. 1965; W[artmanniaMuell. Arg., Linnaea 34: 218. 1865. TYPE: Gillespie, Brittonia 45: 92. 1993. TYPE: Se- WartmanniastillingiiJblia (F. Muell.) Muell. Arg. [= Omalanthusstillingiifolius F. Muell.]. nefelderopsis croizatii Steyerm. DibrachionRegel, Index Sem. Hort.Bot. Petropol.1865: A genus of four species endemic to South Amer- 51. 1866. TYPE: Dibrachion peltatum Regel [ Omalanthusfastuosus (Linden) Villar]. ica (tepui areas of Colombia, Venezuela, northern Brazil). An Old World genus of ca. 30 species, extending from Malaya through Indonesia to the Pacific is- Subtribe 46b. CARUMBIINAE Muell. Arg., lands (as far as Tahiti and the Austral Islands). Linnaea 34: 203. 1865; DC. Prodr. 15(2): 1142. 1866. TYPE: Carumbium Reinw. [ Subtribe 46c. HIPPOMANINAE Grisebach, Omalanthus A. Juss.]. Fl. Brit. W. Ind. 49. 1859; Pax, Natiirl. Pflan- zenfam. ed. 1, 3(5): 91. 1890. TYPE: Hip- Hippomaneae subtribeHomalanthinae Pax & Hoffm., Pflanzenr.147. V (Heft52): 42. 1912. TYPE: Oma- pomane L. lanthusA. Juss. Hippomaneae subtribeAdenopeltinae Pax & Hoffm., Monoecious or dioecious trees or shrubs; latex Pflanzenr.147. V. (Heft52): 57. 1912. TYPE: Ad- enopeltisBert. watery, innocuous; leaves entire, pinnately to pal- Hippomaneae subtribeExcoecariinae Pax & Hoffm., mately veined, petiolate, glandular at apex of pet- Pflanzenr.147. V (Heft 52): 153. 1912. TYPE: iole, stipulate; inflorescence terminal, racemose, Excoecaria L. usually bisexual, bracts biglandular; staminate ca- Hippomaneae subtribeGymnanthinae Pax & Hoffm., lyx compressed and 2-lipped, or of only 1 lobe; Pflanzenr.147. V (Heft52): 57. 1912. TYPE: Gym- nanthesSw. stamens 5-50, filamentsfree; pistillate calyx 2- or Hippomaneaesubtribe Stillingiinae Pax & Hoffm.,Pflan- 3-lobed; ovary 2-locular (rarely 3-locular), exap- zenr. 147. V (Heft52): 174. 1912. TYPE: Stillingia pendiculate; styles free or nearly so, unlobed to Gardenex L. bifid,usually glandular near tip; fruitdehiscent or Monoecious or dioecious trees, shrubs, or herbs; indehiscent; seeds with large caruncle. latex milky, often caustic; indumentum simple or As interpretedearlier (Webster, 1975), subtribe absent; leaves alternate (rarely opposite), usually Carumbiinae is here treated as including only the unlobed, pinnately veined, often with laminar or type genus Omalanthus. Pax & Hoffmann(1912) petiolar glands; stipules mostly small, sometimes also included Pimelodendron in the subtribe,prob- obsolete or absent; inflorescences terminal or ax- ably because of the similar staminate calyx. How- illary, mostly spicate (sometimes racemose), bracts ever, Pimelodendron differs in its pollen sexine usually biglandular; staminate calyx open in bud, ornamentation, eglandular bracts, and ecaruncu- sepals 0-3; stamens 1-20, free or connate; pis- late seeds. tillate sepals mostly 2 or 3, sometimes obsolete or absent; ovary 2-4-locular (6-10-locular in Hip- 279. Omalanthus A. Jussieu, Tent. Euphorb. pomane), often carinate or horned; styles ? con- 50. 1824; Bentham, Gen. P1. 3: 331. 1880 nate, unlobed; fruitcapsular or drupaceous; seeds (as Homalanthus); Pax & Hoffm., Pflanzenr. carunculate or ecarunculate. 147. V (Heft 52): 42. 1912; J. J. Smith, Nova This subtribe is the major taxon withinthe sub- Guinea 8: t. 140, 141. 1912; 12: t. 228, familyother than the tribe Euphorbieae. The 500- 229. 1917; AiryShaw, Kew Bull. 21: 409. odd species are distributedamong about 15 genera, 1968; Kew Bull. Add. Ser. 8: 115. 1980; many of which are notoriously poorly delimited. Sykes, New Zeal. J. Bot. 7: 302. 1969; A. Pax & Hoffmann (1912) proposed an unworkable C. Smith,Fl. Vit. Nova 2: 558. 1981; St. system in which the genera were assigned to five John, Nordic J. Bot. 4: 53. 1984. TYPE: Oma- subtribes; these cannot be satisfactorily distin- lanthus leschenaultianus A. Juss. [= Oma- guished. Future monographic work may provide lanthus populneus (Geisel.) Pax]. some clues that will make it possible to subdivide CarumbiumReinwardt, Syll. P1. Nov. 2(1): 6. 1826. subtribe Hippomaninae, but at present, a subdi- TYPE: Carumbiumpopulifolium Reinw. [= Oma- vision seems premature. lanthuspopulneus (Geisel.) Pax].

KEY TO THE GENERA OF SUBTRIBE HIPPOMANINAE la. Ovary 2-3-locular; fruitscapsular or drupaceous. 2a. Staminatespikes not capitular(or if so, seeds not foveolate).

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3a. Staminatespikes conspicuously bracteate; staminate sepals mostly3-5; stamens3; seeds ecarunculate...... 280. Spirostachys 3b. Staminatespikes usually not conspicuouslybracteate; staminate sepals 2 or 3 (or obsolete). 4a. Seeds generallycarunculate, testa dry. 5a. Spikes usuallyterminal or oppositeleaves; staminatecalyx 2-3-lobed; pistillateflowers sessileor short-pedicellate(fruiting pedicels less than 1 cm long). 6a. Columellanot 3-hornedat base; pistillateflowers often pedicellate . 281. Sebastiania 6b. Columella3-horned at base; pistillateflowers sessile .282. Stillingia 5b. Spikes axillary;staminate sepals 1 or 2, oftenreduced or obsolete;pistillate flowers distinctlypedicellate (1 cm long or morein fruit)...... 283. Gymnanthes 4b. Seeds generallyecarunculate, testa dryor fleshy. 7a. Fruitcapsular (or if drupaceousthen bracts glandular). 8a. Dioecious (rarelymonoecious); petioles usually eglandular; inflorescences axillary; staminatesepals distinct;ovary not appendiculate;caruncle deciduousfrom seed and persistenton columella...... 284. Excoecaria 8b. Monoecious;petioles often glandular; inflorescences terminal or axillary. 9a. Bractsglandular; leaves mostlywith petiolar glands. 10a. Seed coat dry; calyx suppressed. 1la. Columellanot 3-horned. 12a. Stamens2-4; leaves pinnatelyveined, unlobed .- ... .-.-.-.----.-----..----- ...-.---...- 285. Colliguaja 12b. Stamen1; leaves palmatelyveined, ? lobed 286. Dalembertia 1lb. Columella3-horned; leaves pinnatelyveined .- . 287. Adenopeltis 10b. Seed coat fleshy;staminate calyx present ...... 288. Sapium 9b. Bractseglandular. 13a. Spikes capitular,pseudoaxillary; stamens 3, filamentsexserted beyond gamophyllouscalyx; stylesconnate ...... 289. Glyphostylus 13b. Spikes elongated. 14a. Spikes terminal;filaments connate; leaves withglands at apex of petiole .---- .--...-. . 290. Grimmeodendron 14b. Spikes axillary;filaments free; leaves eglandular . 291. Bonania 7b. Fruitdrupaceous; inflorescences axillary; bracts eglandular .--...------292. Duvigneaudia 2b. Staminatespikes capitular, terminal; bracts glandular; seeds foveolate,with large caruncle ...... 3...... Mun...... a...... 2 9 a pr o lb. Ovary 6-10-locular; fruitsdrupaceous; spikes terminal; bracts glandular ...... 294. Hippomane

280. Spirostachys Sonder, Linnaea 23: 106. MicrostachysA. Jussieu,Euphorb. Tent. 48. 1824. TYPE: 1850; Pax & Hoffm.,Pflanzenr. 147. V (Heft Microstachysbicornis (Vahl) A. Juss.[= Sebasti- Arg.; lectotype]. Afr. 6(1): ania corniculata(Vahl) Muell. 52): 153. 1912; Prain, Fl. Trop. CnemidostachysMartius & Zuccarini,Nova Gen. Sp. 1: 1005. 1913; AlfaroCardoso, Serv. Agric. Mo- 66. 1824. TYPE: not designated(Martius described vamb., Publ. A18: 1. 1964; Dyer, Gen. S. a largenumber of taxa of Cnemidostachys,and the Afr.Fl. P1. ed. 3, 322. 1975; Radcliffe-Smith, choice of lectotypeis best leftto a futuremonog- Fl. E. Trop. Afr.,Euphorb. 1: 385. 1987. rapherof thisgroup). DitrysiniaRaf., Neogenyton2. 1825. TYPE: Ditrysinia TYPE: Spirostachysafricana Sond. ligustrina(Mx.) Raf.[= Sebastiania ligustrinaMx.]. AdenogyneKlotzsch, Arch. Naturgesch.7: 183. 1841. An Africangenus of one species (or two, if TYPE: Adenogynepachystachya Kl. [= Sebasti- Excoecaria veneniferaPax is included). ania pachystachys(Kl.) Muell. Arg.]. SarothrostachysKlotzsch, Arch. Naturgesch.7: 185. 281. Sebastiania Sprengel, Neue Entdeck. 1841. TYPE: Gymnanthesmultiramea Baillon [ Pflanzenk.2: 118, t. 3. 1820; Muell. Arg., Sebastiania multiramea(Baillon) Muell. Arg.]. ElachocrotonF. Mueller,Hook. J. Bot. Kew Gard. Misc. 1866; Fl. Bras. 11(2): DC. Prodr.15(2): 1164. 9: 17. 1857. TYPE: Elachocrotonasperococcum F. 544. 1874; Bentham,Gen. P1. 3: 336. 1880; Muell.[= Sebastiania chamaelea (L.) Muell.Arg.]. Pax & Hoffm.,Pflanzenr. 147. V (Heft 52): TragiopsisKarsten, Wochenschr. Gaertnerei Pflanzenk. 88. 1914; Natiirl.Pflanzenfam. ed. 2, 19c: 2: 5. 1859. TYPE: TragiopsisftuticulosaKarst. [ Sebastiania corniculata(Vahl) Muell. Arg.]. 192. 1931; Smith,Downs & Klein,Fl. Illustr. Dendrocousinsia Millspaugh,Field Mus. Publ. Bot. 2: Catar. EUFO 242. 1988; Webster & Huft, 374. 1913. TYPE: Dendrocousinsiaspicata Millsp. Ann. MissouriBot. Gard. 75: 1127. 1989. [= Sebastiania spicata (Millsp.)Pax & Hoffm.]. TYPE: Sebastiania brasiliensisSpreng. A genus of ca. 100 species, mainly neotropical, GussoniaSprengel, Neue Entdeck.Pflanzenk. 2: 119. Asia, 1820. TYPE: Gussonia discolor Spreng.[= Sebas- but withoutliers in temperate North America, tiania discolor(Spreng.) Muell. Arg.]. and Australasia. The herbaceous species of section

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Microstachysappear very different from the woody Actinostemonas a distinctgenus, there appear to ones and may possiblyrepresent a distinctgenus. be no satisfactorydistinguishing characters. In the However,there are connectinglinks such as Se- previouscentury, Baillon (1858) upheldActinos- bastiania panamensis Webster(Webster & Huft, temonas distinct,while Mueller (1866) maintained 1988), so that it seems preferableat presentto Actinostemon and Dactylostemon as separate gen- maintainthe broad (and probablyunnatural) ge- era but combined Gymnanthes with Sebastiania. nericcircumscription of Pax & Hoffmann(1931). Genericdelimitation clearly remains a majorprob- lem in thiscomplex of neotropicalHippomaninae 282. Stillingia Gardenex L., Syst.Nat. ed. 12, withreduced staminate flowers. 2: 637. 1767; Mant. P1. 19, 126. 1767; FollowingRothmaler (1944), a numberof Amer- Muell. Arg., DC. Prodr. 15(2): 1155. 1866; ican botanistshave adoptedthe name Ateramnus Fl. Bras. 11(2): 537. 1874; Bentham,Gen. P. Brownein place of Gymnanthes.However, this P1. 3: 334. 1880; Pax & Hoffm.,Pflanzenr. is based on a very questionableidentification of 146. V (Heft 52): 180. 1912; Rogers,Ann. Browne's plant fromJamaica with Gymnanthes Missouri Bot. Gard. 38: 207. 1951; van lucida Sw.; it appearsmore likely that Browne had Steenis,Blumea Suppl. 5: 302. 1966; Web- a species of Sapium (Webster,1983). ster,J. ArnoldArbor. 48: 388. 1967; A. C. Smith, Fl. Vit. Nov. 2: 565. 1981. TYPE: 284. Excoecaria L., Syst. Nat. ed. 10, 1288. Stillingia sylvatica L. 1759; Bentham, Gen. P1. 3: 337. 1880; Hook. GymnostillingiaMuell. Arg., Linnaea 32: 89. 1863. f.,Fl. Brit.Ind. 5: 472. 1888; Pax & Hoffm., TYPE: Gymnostillingiaacutifolia (Benth.) Muell. Pflanzenr.147. V (Heft52): 157. 1912; Gag- Arg.[= Stillingiaacutifolia (Benth.) Hemsl.; lec- nepain,Fl. Indochine5: 402. 1926; Airy totype,designated by Wheeler, 1975]. Shaw, Kew Bull. 26: 268. 1971; Whitmore,Tree A mainlyAmerican genus of ca. 25 species, Fl. Malaya 2: 96. 1973; AiryShaw, Kew Bull. witha fewdisjunct taxa in the Mascarene Islands, Add. Ser. 4: 112. 1975; 8: 88. 1980; Kew easternMalaysia, and Fiji. Bull. 35: 630. 1980; 36: 296. 1981; Rad- cliffe-Smith,Fl. E. Trop. Afr. Euphorb. 1: 283. Gymnanthes Swartz,Prodr. Veg. Ind. Occ. 382. 1987. TYPE: Excoecaria agallocha L. 6, 95. 1788; Grisebach,Fl. Brit.W. Ind. 50. Commia Loureiro,FH. Cochinchin. 605. 1790. TYPE: 1859; Bentham,Gen. P1. 3: 337. 1880; Sar- Commia cochinchinensis Lour. [= Excoecaria gent,Silva N. Amer.7: t. 309. 1895; Pax & agallocha L. ?]. Hoffm.,Pflanzenr. 147. V. (Heft 52): 81. Anomostachys(Baillon) Hurusawa, J. Fac. Sci. Univ. 1912; Fawc. & Rend.,Fl. Jam.4: 329. 1920; TokyoBot. 6: 311. 1954. TYPE: Stillingia lastelhii Webster,J. ArnoldArbor. 48: 387. 1967; Baillon[= Excoecaria lastelhii(Baillon) Muell. Arg.]. Taxon 32: 304. 1983; Ann. MissouriBot. A paleotropical genus of ca. 40 species, mostly Gard.75: 1129. 1989. TYPE: Gymnanthes in tropicalAsia but a few taxa in Africa,Mada- lucida Sw. [lectotype,chosen by Grisebach, gascar,and tropicalAustralia. Mueller (1866, 1974) 1859]. gave Excoecaria a verybroad circumscriptionin- ActinostemonMartius ex Klotzsch,Arch. Naturgesch. 7: cludingSapium, but thishas not been followedby 184. 1841; Pax & Hoffm.,Pflanzenr. 147. V (Heft recentauthors. 52): 57. 1912;Jablonski, Phytologia 18: 213. 1969. TYPE: Actinostemonconcolor (Spreng.) Muell. Arg. 285. Colliguaja Molina, Saggio Chile 158. [Gussoniaconcolor Spreng. = Gymnanthescon- 1781; Hooker, Bot. Misc. 1: 138. 1830; Muell. color(Spreng.) Muell. Arg.; lectotype, designated byJablonski, Mem. New York Bot. Gard. 17: 177. Arg., DC. Prodr. 15(2): 1226.1866; Fl. Bras. 1967; Farret al. (1979) incorrectlycited Actinos- 11(2): 630, t. 85. 1874; Bentham, Gen. P1. temongrandifolius KI., an invalidname]. 3: 338. 1880; Pax & Hoffm.,Plfanzenr. 147. DactylostemonKlotzsch, Arch. Naturgesch. 7: 181. 1841. V (Heft 52): 265. 1912; L. Smithet al., Fl. TYPE: Dactylostemonschomburgkii Kl., Hook. Lon- don J. Bot. 2: 44. 1843 [= Gymnanthes schom- Illustr.Catar. EUFO 326. 1988. TYPE: Col- burgkii (K1.) Webster,comb. nov.; lectotype,cho- liguaja odorifera Mol. sen here]. A South Americangenus of fivespecies. As circumscribedhere following Webster (1989), Gymnanthesis an Americangenus of ca. 40 spe- 286. Dalembertia Baillon, Etude G6n. Eu- cies. Althoughmost 20th-centuryworkers have phorb. 545. 1858; Muell. Arg., DC. Prodr. followedPax & Hoffmann(1912) in recognizing 15(2): 1225. 1866; Bentham, Gen. P1. 3:

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339. 1880; Pax & Hoffm., Pflanzenr. 147. 317. 1954. TYPE: Shirakiajaponica (Sieb. & Zucc.) & Zucc.) Pax]. V (Heft 52): 268. 1912; Standley & Stey- Hurus.[= Sapium japonicum (Sieb. erm., Fieldiana Bot. 24(6): 86. 1949. TYPE: As here delimited,Sapium is a large and diverse Dalembertia populifolia Baillon. genus of ca. 90-100 species, the majority neo- Alcoceria Fernald,Proc. Amer.Acad. ArtsSci. 36: 493. tropical, a minorityAsian. Triadica, accepted by 1901. TYPE: Alcoceria pringlei Fern. [= Dalem- some Asian workers, is distinctivelydifferent from bertiapopulifblia Baillon]. the neotropicalspecies in such featuresas its non- arillate seeds; furtherinvestigation may show that A well-marked genus of five species in Mexico it should be generically separate from Sapium. and Guatemala, distinguishedby its compact inflorescence and highly reduced staminate flowers. 289. Glyphostylus Gagnepain, Bull. Soc. Bot. France 71: 871. 1925; Fl. Indochine5: 402. 287. Adenopeltis Bertero ex A. Jussieu, Ann. 1926; AiryShaw, Kew Bull. 16: 372. 1963; Sci. Nat. I. 25: 24. 1832; Muell. Arg., DC. 26: 281. 1972. TYPE: Glyphostylus laoticus Prodr. 15(2): 1164. 1866; Bentham, Gen. P1. Gagnep. 3: 338. 1880; Pax & Hoffm.,Pflanzenr. 147. V (Heft 52): 264. 1912. TYPE: Adenopeltis A monotypic genus of southeast Asia (Thailand colliguaya Bert. ex A. Juss. (nom. illeg.) [= to Vietnam). Excoecaria serrata Ait., Hort, Kew. ed. 2, 5: 418. 1813; = Adenopeltis serrata (Ait.) 290. Grimmeodendron Urban,Symb. Ant. 5: Webster, comb. nov.]. 397. 1908; Pax & Hoffm.,Pflanzenr. 147. V (Heft 52): 258. 1912; Fawc. & Rend., Fl. A monotypic genus of Chile. Jam.4: 326. 1920; Alain, Fl. Cuba 3: 119. 1953; Correll& Correll,Fl. Bahama Arch. 288. Sapium P. Browne, Civ. Nat. Hist. Jam. 818. 1982; Liogier, Fl. Espafiola 4: 153. 338. 1756; Jacquin,Select. Stirp. Amer. Hist. 1986: TYPE: Grimmeodendron jamaicense 249. 1763; Bentham, Gen. P1. 3: 334. 1880; Urban [lectotype,designated by Wheeler, Hemsley, Hook. Ic. P1. 27: t. 2647-2650, 1975]. 2677-2684. 190 1-1902; 29: t. 2878-2900. A West Indian genus of two species. 1909; Pittier, Contr. U.S. Natl. Herb. 12: 159. 1906; Pax & Hoffm., Pflanzenr. 147. V (Heft 52): 199. 1912; Fawc. & Rend., Fl. 291. Bonania A. Richard,Hist. Fis. Cuba, 2, Jam.4: 324. 1920; Gagnepain,Fl. Indochine 11: 201. 1853; Bentham,Gen. P1. 3: 335. 5: 394. 1926; Leonard, Bull. Jard. Bot. Brux. 1880; Pax & Hoffm.,Pflanzenr. 147. V (Heft 29: 133. 1959; Fl. Congo 8(1): 151. 1962; 52): 259. 1912; Alain,Fl. Cuba 3: 115. 1953; Webster, J. Arnold Arbor. 48: 391. 1967; Borhidi,Acta Bot. Acad. Sci. Hungar. 22: Jablonski,Phytologia 14: 441. 1967; 16: 393. 305. 1976; Correll & Correll,Fl. Bahama 1968; Radcliffe-Smith,Fl. E. Trop. Afr., Eu- Arch. 784. 1982. TYPE: Bonania cubana A. phorb. 1: 389. 1987; Huft, Ann. Missouri Rich. Bot. Gard. 75: 1132. 1989; Kruijt & Zijlstra, Hypocoton Urban, Symb. Ant. 7: 263. 1912. TYPE: Taxon 38: 320. 1989. TYPE: Sapium jamai- Hypocoton domingensisUrb. [= Bonania dom- cense Sw. ingensis(Urb.) Urb.].

Triadica Loureiro,Fl. Cochinchin.610. 1790. TYPE: A genusof ca. eightspecies, mostly in Cuba but Triadica sinensis Lour. [= Sapium sebiferum(L.) withone each in the Bahamas and Hispaniola. Roxb.; lectotype,chosen here]. Ill. Bot. Himal. 354. 1839. TYPE: Falconeria Royle, Duvigneaudia L6onard,Bull. Jard. Bot. Falconeriainsignis Royle [= Sapium insigne(Royle) 292. Benth.; lectotype,designated by Wheeler, Taxon Brux. 29: 15. 1959; Fl. Congo 8(1): 139. 24: 535]. 1962. TYPE: Duvigneaudia inopinata (Prain) Conosapium Muell. Arg.,Linnaea 32: 87. 1863. TYPE: L6onard. Conosapiummadagascariense Muell. Arg. [= Sa- pium madagascariense (Muell. Arg.) Pax]. A monotypicgenus of west Africa(Cameroon TaeniosapiumMuell. Arg., DC. Prodr.15(2): 1200. 1866. to Congo and Gabon). TYPE: Taeniosapium goudotianum (Muell. Arg.) Muell. Arg. [= Sapium goudotianum(Muell. Arg.) Pax]. 293. Maprounea Aublet,Hist. P1. Guiane Fr. Shirakia Hurusawa,J. Fac. Sci. Univ. Tokyo, Bot. 6: 895. 1775; Muell. Arg., DC. Prodr. 15(2):

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1190. 1866; Fl. Bras. 11(2): 542. 1874; This tribe includes only the type genus Pachy- Bentham, Gen. P1. 3: 333. 1880; Pax & stroma. The position of this tribe has been poorly Hoffm., Pflanzenr. 147. V (Heft 52): 175. understood owing to its initial misplacement by 1912; Leonard, Fl. Congo 8(1): 142. 1962; Mueller (1866) in the Acalypheae because of the Radcliffe-Smith,Fl. E. Trop. Afr. Euphorb. valvate staminate calyx. Bentham, apparently mis- 1: 395. 1987; Webster, Ann. Missouri Bot. led by Mueller, referred Pachystroma to his sub- Gard. 75: 1131. 1988; L. Smith et al., Fl. tribe Adrianeae. Pax & Hoffmann(1919) followed Illustr. Catar. EUFO 316. 1988. TYPE: Ma- Mueller's example by creating a subtribe Pachy- prounea guianensis Aubl. strominaein the Acalphyeae, and finally(Pax, 1924; Pax & Hoffmann, 1931) placed the genus in a AegopricumL., PI. Surinam.15. 1775. TYPE: Aegopri- cum betulinumL. [= Maprounea guianensisAubl.]. separate tribe followingthe Acalphyeae. Curiously, no one seems to have paid attention to the opinion A genus with a disjunct distribution,one or two of Baillon (1865, 1866), who placed his synony- species in South America and Panama, and two in mous genus Acantholoma in the Hippomaneae in west Africa. the vicinity of Stillingia. In support of Baillon's disposition,characters such as the laticiferousstems, 294. Hippomane L., Sp. Pl. 1191. 1753; Gen. spicate inflorescence with glandular bracts, and P1. 499. 1754; Muell.Arg., DC. Prodr.15(2): perforate-tectate pollen grains of Pachystroma 1199. 1866; Bentham,Gen. P1. 3: 333. 1880; clearly rule out a placement in the Acalyphoideae Pax & Hoffm.,Pflanzenr. 147. V (Heft 52): and indicate a position in the Euphorbioideae. In 261. 1912; Fawc. & Rend., Fl. Jam.4: 327. fact, it is rather doubtfulthat Pachystroma should 1920; Webster,J. ArnoldArbor. 48: 393. be kept in a distinct tribe; furtherstudy will prob- 1967; Correll& Correll,Fl. Bahama Arch. ably show that it should be referred to a subtribe 818. 1982; Liogier, Fl. Espafiola 4: 158. of the Hippomaneae. 1986; Webster,Ann. Missouri Bot. Gard. 75: 1131. 1988; Howard,Fl. Lesser Ant. 5: 54. 295. Pachystroma Muell. Arg., Linnaea 34: 1989. Mancanilla Miller,Gard. Dict. Abr. 177. 1865; DC. Prodr. 15(2): 893. 1866; Fl. Ed. 4. 1754. TYPE: Hippomane mancinella Bras. 11(2): 387, t. 54. 1874; Bentham, Gen. L. [lectotype,designated by M. L. Green,Prop. P1. 3: 307. 1880; Pax, Pflanzenr. 147. II Brit.Bot. 195. 1929]. (Heft 44): 99. 1910; Pax & Hoffm., Natiirl. A neotropical genus of three species, one wide- Pflanzenfam. ed. 2, 19c: 151. 1931; Smith spread in the Caribbean and Galapagos, the other et al., Fl. Ilustr. Catar. EUFO 225. 1988. two endemic to Hispaniola. TYPE: Pachystroma ilicifolium Muell. Arg. [= Pachystroma longifolium (Nees) I. M. Jhnt.].

Tribe 47. PACHYSTROMATEAE (Pax & Acantholoma Gaudichaud ex Baillon, Adansonia I. 6: Hoffmann)Pax, Bot. Jahrb.59: 145. 1924. 231. 1866. TYPE: Acantholoma spinosumGaud. [= Acalypheae subtribePachystromatinae Pax & ex Baillon Pachystroma longijblium(Nees) I. M. Jhnt.]. Hoffm.,Pflanzenr. 147. XIV (Heft 68): 3. 1919. TYPE: Pachystroma Muell. Arg. This monotypic Brazilian genus includes a single variable species found from Minas Gerais to Rio Monoecious trees or shrubs; stems with milky Grande do Sul. latex; indumentum absent; leaves alternate, unlobed, pinnately veined, margins entire to mostly Tribe 48. HUREAE Dumortier, Anal. Fam. P1. spinose-dentate, stipulate. Inflorescence spicate, 45. 1829 (as Huraceae); Webster, Taxon 24: terminal, pistillate flowersproximal on spikes or in 600. 1975. Hippomaneae subtribe Hureae upper axils; staminate bracts biglandular. Stami- Muell. Arg., Linnaea 34: 203. 1865. TYPE: nate flowerssubsessile; sepals 5 or 6, valvate; disk Hura L. absent; stamens 3, filamentsconnate into a column, anthers extrorse; pollen grains 3-colporate, per- Monoecious trees or shrubs; latex + toxic; in- forate-tectate; pistillode absent. Pistillate flowers dumentum absent; leaves alternate, unlobed, pin- subsessile; sepals 3, persistent; disk absent; ovary nately veined, biglandular at base of blade, stipu- 3-locular; styles basally connate, unlobed. Fruit late. Inflorescences terminal or axillary, spicate; capsular; columella 3-horned at base; seeds eca- bracts ? adnate to rachis of inflorescence, eglan- runculate, endosperm present. dular. Staminal calyx 2-3-lobed; disk absent; sta-

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mens 1-20, free or filamentsconnate; pollen grains A neotropicalgenus of fivespecies, endemicto 3-colporate, colpi somewhat marginate, sexine very easternBrazil. The generaAlgernonia and Tetra- finely perforate-tectate. Pistillate sepals 5 or 6, plandra appear veryclosely related, and it might free or connate; ovary 3-20-locular, exappendic- be preferableto treatthem as subgeneraof a single ulate; stylesunlobed, connate. Fruit capsular; seeds genus. ecarunculate, testa dry. 298. Ophthalmoblapton Fr. Allemao,PI. No- This strictlyAmerican tribe of four genera, with vas Brasil,1849; Ann. Sci. Nat. Bot. III. 13: its center of diversity in eastern Brazil, closely 119. 1849; Muell. Arg. 15(2): 1155. 1866; approaches the tribe Hippomaneae via Dalember- Fl. Bras. 11(2): 531. 1874; Bentham,Gen. tia. PI. 3: 333. 1880; Pax & Hoffm.,Pflanzenr. 147. V. (Heft 52): 278. 1912; Emmerich, KEY TO THE GENERA OF TRIBE HUREAE Bol. Mus. Nac. Rio de Janeiro,Bot. 62: 1. la. Sepals 3-5, imbricate;styles united halfway or 1981; L. Smithet al., Fl. Illustr.Catar. EUFO less. 2a. Filamentsof stamensconnate; styles con- 325. 1988; Howard, Fl. Lesser Ant. 5: 57. nate less than halfway;petioles mostly 1 1989. TYPE: OphthalmoblaptonFr. Allem. cm or longer 296. Tetraplandra 2b. Filamentsfree; stylesconnate halfway or A neotropicalgenus of fourspecies endemicto more; petiolesmostly 1 cm or shorter. easternBrazil. . . . 297. Algernonia lb. Sepals notimbricate; styles completely connate intoa long column. 299. Hura L., Sp. PI. 1008. 1753; Gen. PI. ed. 3a. Ovary 3-locular; stylar column apically 5, 439. 1754; Muell. Arg.,DC. Prodr. 15(2): 3-lobed; stamen 1; staminatecalyx rup- 1228. 1866; Bentham,Gen. P1. 3: 339. 1880; or turinginto 2 3 segments;inflorescences Pax & Hoffm.,Pflanzenr. 147. V (Heft 52): axillary;leaves oblanceolate,cuneate at base 298. Ophthalmoblapton 271. 1912; Fawc. & Rend., Fl. Jam.4: 333. 3b. Ovary 5-20-locular; stylar column api- 1920; Standley& Steyerm.,Fieldiana Bot. callydilated into a lobeddisk; stamens 10- 24(6): 124. 1949; Burch,Ann. MissouriBot. 80, filamentsconnate; staminate calyx cu- Gard. 54: 330. 1968; Correll& Correll,Fl. pulate,truncate; inflorescences terminal (or pistillateflowers axillary); leaves ovate, Bahama Arch. 821. 1982. TYPE: Hura cre- roundedto cordateat base 299. Hura pitans L. A neotropicalgenus of two species, the type 296. Tetraplandra Baillon, Ann. Sci. Nat. IV. widespreadfrom Cuba and Mexico southto Brazil, 9: 200. 1858; Etude G6n. Euphorb. 549. theother (H. polyandra Baill.)recorded from Mex- 1858; Muell. Arg., DC. Prodr. 15(2): 1230. ico to Costa Rica; widelyintroduced throughout 1866; Fl. Bras. 11(2): 533. 1874; Pax & the tropics. Hoffm., Pflanzenr. 147. V (Heft 52): 275. 1912; Emmerich, Arq. Mus. Nac. Rio de Ja- Tribe 49. EUPHORBIEAE [Blume,Bijdr. 631. neiro 56: 94. 1981. TYPE: Tetraplandra lean- 1825]; Pax & Hoffmann,Natiirl. Pflanzen- dri Baill. fam.ed. 2, 19c: 207. 1931; Wheeler,Amer. Midl. Nat. 30: 477. 1943. TYPE: Euphor- A neotropical genus of seven species, mostly in bia L. eastern Brazil, but also recorded from the Ama- zonian region as far west as Peru (Huft, pers. Monoecious(or less commonlydioecious) trees, comm.). shrubs,or herbs;latex milky,innocuous or toxic; indumentumsimple or absent; leaves alternate, opposite,or whorled,mostly unlobed and pinnately 297. Algernonia Baillon, Ann. Sci. Nat. IV. 9: veined,without laminar or petiolarglands; stipules 198. 1858; Etude Gen. Euphorb. 546. 1858; presentor absent.Inflorescences pseudanthial, the Muell. Arg. 15(2): 1230. 1866; Bentham, connate bracts forminga cyathiumenclosing a Gen. P1. 3: 339. 1880; Pax & Hoffm.,Pflan- singleterminal pistillate flower and 4 or 5 lateral zenr. 147. V (Heft 52): 276. 1912; Emmer- staminatemonochasia; bracts eglandular.Stami- ich, Arq. Mus. Nac. Rio de Janeiro 56: 92. nate flowerwith calyx reducedor absent; stamen 1981; Bradea 3: 148. 1981. TYPE: Alger- 1; pollengrains 3-colporate, colpi marginate,sex- nonia brasiliensis Baill. ine perforate-tectate.Pistillate flower with calyx

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3-6-lobed or absent; ovary (2-)3-(4-)locular; styles KEY TO THE GENERA OF SUBTRIBE ANTHOSTEMINAE free or basally connate, bifid(rarely entire). Fruit 1a. Cyathiabisexual; involucral bracts laterally con- capsular (rarely drupaceous); seeds carunculate or nate into an open 4-lobed involucre;cyathial ecarunculate; testa dry, smooth or ornamented. glands at marginsof internalbracts; inflorescences axillary;ovary 3-locular. 300. Anthostema A large tribe of more than 1500 species, here l b. Cyathiamostly unisexual; outer involucral bracts divided (followingWebster, 1975) into 3 subtribes connateinto a closedtube; cyathial glands fused by pairsinto 4 lobesalternating with involucral with 11 genera. A catalogue of the more than 2000 lobes; inflorescencesterminal; ovary 4-locular species names in tribe Euphorbieae has recently ...... 301. Dichostemma been published by Oudejans (1990). 300. Anthostema A. Jussieu, Euphorb. Tent. KEY TO THE SUBTRIBESOF TRIBE EUPHORBIEAE 56. 1824; Baillon, Ann. Sc. Nat. 9: 193. 1a. Staminatecalyx present; involucre of 4 bracts, 1858; Boissier, DC. Prodr. 15(2): 188. 1862; partiallyfused; cyathial glands commissural, not Bentham, Gen. P1. 3: 261. 1880; Hutchinson, on rimof cyathialtube 49a. Anthosteminae Fl. Trop. Afr.6(1): 607. 1912; Pax & Hoffm., lb. Staminatecalyx absent;involucral bracts usually 5, connateinto a tube. Natiirl. Pflanzenfam. ed. 2, 19c: 207. 1931; 2a. Petaloid appendages bracteal (dilatedin- Keay, Fl. W. Trop. Afr. ed. 2, 1: 416. 1958; volucral lobes), opposite the staminate Berhaut, Fl. Ill. Senegal 3: 379. 1975. TYPE: monochasia;cyathial glands commissural; Anthostema senegalense A. Juss. bractsof staminatemonochasia large, imbricate,enclosing staminate flowers; pistil- A paleotropical genus of three species, two in late sepals 5 or 6, large,imbricate. west Africa and one in Madagascar...... 49b. Neoguillaum iniinae 2b. Petaloidappendages interbracteal (i.e., op- positecyathial glands), alternate with sta- 301. Dichostemma Pierre, Bull. Mens. Soc. minatemonochasia (sometimes absent); cy- Linn. Paris 1 (159): 1259. 1896; Hutchinson, athialglands on rimof cyathium;bracts of Fl. Trop. Afr. 6(1): 605. 1912; Pax & Hoffm., staminatemonochasia small or obsolete Natiirl. Pflanzenfam. ed. 2, 19c: 207. 1931; ...... E uphorbiinae Keay, Fl. W. Trop. Afr. ed. 2, 1: 416. 1958. TYPE: Pierre. Subtribe 49a. ANTHOSTEMINAE (Baillon) Dichostemma glaucescens Webster, Taxon 24: 600. 1975. Anthostem- A genus of three species in tropical west Africa. ideae Baillon, Ann. Sci. Nat. IV. 9: 192. 1858. Anthostemeae K1. & Gcke., Monatsber. Kon- igl. Preuss. Akad. Wiss. Berlin 1859: 247. Subtribe 49b. NEOGUILLAUMINIINAE 1859. TYPE: Anthostema A. Juss. Croizat, Phil. J. Sci. 64: 408. 1938; Webster, Taxon 24: 601. 1975. TYPE: Neoguillaumi- Monoecious or dioecious trees; leaves alternate, nia Croiz. pinnately veined, stipules inconspicuous; cyathia in terminal or axillary dichasia; cyathial involucre of Trees or shrubs; indumentum absent; leaves al- 4 outer bracts subtending or enclosing 4 subflo- ternate or opposite, entire; stipules obsolete; cy- rescences; pistillateflower solitary, central and ter- athia in axillary pedunculate cymes; involucral minal (or apparently lateral); staminate flowersin bracts 4(5), dilated or petaloid, connate near base; dichasia ? enclosed by 4 secondary (inner) bracts; glands 4, 8, or 10, in 4 or 5 pairs, partitioning cyathial glands projecting inward from infolded the staminate inflorescences; pistillate flower cen- margins of outer bracts; staminate flowersand pis- tral, staminate flowers in bracteate monochasia; tillate flowers with gamophyllous calyx; ovary 3- staminate calyx absent; pistillate sepals 5 or 6, 4-locular; styles emarginate to bifid;fruit capsular; imbricate; ovary 3-locular; styles connate at base, seeds carunculate or ecarunculate. unlobed, dilated; fruitcapsular; seeds smooth, carunculate. This paleotropical subtribe includes two genera of Africa and Madagascar. The cyathia in the An- As defined by Webster (1975), this subtribe thosteminae are apparently tetramerous and some- includes two Australasian genera. In their pseu- what bilaterallysymmetrical; the tetramerycarries dopetals developed from the involucral bracts, the over to some extent in the Neoguillauminiinae, cyathia of the Neoguillauminiinae are distinctively suggesting that the pentamerous condition in the differentfrom those in Euphorbia, with append- Euphorbiinae may be a derived condition. ages developed from the interbracteal glands.

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KEY TO THE GENERA OF SUBTRIBE phorbieaesubtribe Pedilanthinae (KI. & Gcke.) Hu- NEOGUILLAUMINIIAE rusawa,J. Fac. Sci. Univ. Tokyo, Sect. 3, Bot. 6: 226. 1954. TYPE: PedilanthusPoit. la. Cyathialglands 8 or 10, massive;petaloid in- Euphorbieaesubtribe Anisophyllae Ki. & Gcke., Mon- bracts large (ca. 1 cm long); leaves volucral atsber.Preuss. Akad. Wiss. Berlin1859: 247. 1859. 302. Neoguillauminia spiral TYPE: AnisophyllumHaw. [= Chamaesyce S. F. if lb. Cyathialglands 4, small; involucralbracts, Gray]. petaloid,less than 1 cm long; leaves opposite Euphorbieaesubtribe Tithymalinae Ki. & Gcke., Mon- 303. Calycopeplus atsber.Preuss. Akad. Wiss. Berlin1859: 247. 1859. TYPE: TithymalusScopoli [= Euphorbia L.]. 302. Neoguillauminia Croizat, Phil. J. Sci. 64: 398. 1938; Bull. Jard. Bot. Buit. III. 17: 206. Monoecious (rarely dioecious) trees, shrubs, or 1941; Guillaumin, Fl. Anal. Synopt. Nouv.- herbs; latex milky,innocuous or toxic; indumentum Caled. 182. 1948; McPherson & Tirel, Fl. simple or absent; leaves alternate, opposite, or lobed), usually pinnately Nouv.-Caled. 14(1): 22, t. 3. 1987. TYPE: whorled, unlobed (rarely Neoguillauminia cleopatra (Baill.) Croiz. veined, stipulate or exstipulate; inflorescences ter- or oftensubtended by paired [Euphorbia cleopatra Baill.]. minal axillary, cyathia bracts; cyathium distinctly cupulate with 4 or 5 A monotypic New Caledonian genus, very dif- interbracteal glands on the rim (rarely glands only ferent in habit from its Australian sister genus. 1 or 2); glands usually discrete, with or without petaloid appendages; pistillate flower central, sta- 303. Calycopeplus Planchon, Bull. Soc. Bot. minate flowersin 4 or 5 monochasia, monochasial France 8: 30. 1861; Boissier, Ic. Euphorb. t. bracts reduced; staminate calyx absent; pistillate 120. 1866; Bentham, Fl. Austral. 6: 52. 1873; calyx 3-6-lobed, or oftenvestigial or absent; ovary Gen. P1. 3: 261. 1880; Pax & Hoffm.,Natiirl. 3-locular; styles distinct or connate, bifid (rarely Pflanzenfam. ed. 2, 19c: 221. 1931; Airy entire); fruit capsular (rarely drupaceous); seeds Shaw, Kew Bull. 35: 603. 1980. TYPE: Ca- carunculate or ecarunculate. lycopeplus ephedroides Planch. The subtribe is here considered to include seven A genus of three species endemic to Australia; genera, although some authors have subdivided although xeromorphic in vegetative morphology, Euphorbia into many segregates. There appears they have cyathia that approach those of the An- to be no necessity for furthersubdivision into sub- thosteminae in structure. tribes such as the Pedilanthinae. Typification of the taxa is reviewed by Wheeler (1943). Subtribe 49c. EUPHORBIINAE. TYPE: Eu- phorbia L.

FamilyPedilanthaceae K1. & Gcke., Monatsber.Preuss. Akad. Wiss. Berlin 1859: 247, 253. 1859. Eu-

KEY TO THE GENERA OF SUBTRIBE EUPHORBIINAE la. Cyathia(?) radiallysymmetrical, the glandsnot hiddenwithin a nectarspur; stylesmostly free or nearly so. 2a. Involucralglands (1-)4 or 5, free,alternating with the lobes of the cyathiumalong the rim. 3a. Leaves alternate,opposite, or whorled,but if opposite then not inaequilateral at base; veinsof leaf not chlorenchyma-sheathed;stipules present or absent;main (embryonic) axis not aborting;seeds carunculateor ecarunculate - 304. Euphorbia 3b. Leaves opposite,stipulate, inaequilateral at base, veins withchlorenchyma sheaths; main axis ceasinggrowth above the cotyledons;seeds usuallyecarunculate -305. Chamaesyce 2b. Involucralglands not 4 or 5 alternatingwith lobes; bractssubtending cyathium connate into a cup. 4a. Pistillateflower calyculate; involucral lobes fused and the 2 glands fused into a shield; seeds ecarunculate;stems not succulent-306. Cubanthus 4b. Pistillateflower naked; involucre and glandsconnate into an annularring; seeds mostly carunculate; stemssucculent. 5a. Cyathiumsomewhat bilaterally symmetrical, the gland witha gap on the lowerside 07-Mium_--__--_----_. __-__--_...-_.--__- _.._- ___-_-.--.--__ __-. -__ __-_---__--3 _.--_-.-__._- ___--. __--__-._ -___--_.--__- _. ___-__-_.--__--_.---__--__--__--__--__.--_.--__--_.--__---_ _0_M07_ o na de n 5b. Cyathiumradially symmetrical, without a gap on one side. 7aMnaenu 6a. Involucralglands conspicuous, ovary without angular crests -308. Synadenium 6b. Involucralglands distinct, located below the crenulaterim; ovary with prominent double crestsat the angles-309. Endadenium

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lb. Cyathiumdistinctly bilaterally symmetrical, the glandshidden within the nectarspur; stylesconnate into a long column Pedilan..th....--.--s 310.

304. Euphorbia L., Sp. P1. 450. 1753; Gen. Galarhoeus helioscopius(L.) Haw. [= Euphorbia P1. ed. 5, 208. 1754; Roeper, Enum. Eu- helioscopia L.]. Medusea Haworth,Syn. P1. Succul. 133. 1812. TYPE: phorb.Germ. 9. 1824; Boissier,DC. Prodr. Medusea major (Ait.) Haw. [= Euphorbia caput- 15(2): 7. 1862; Icones Euphorbiarum,1. medusae L. var. major Ait.; lectotype]. 1866; Muell.Arg., Fl. Bras. 11(2): 666. 1874; Treisia Haworth,Syn. P1.Succul. 131. 1812. TYPE: Trei- Bentham,Gen. P1. 3: 258. 1880; Norton, sia clava (Jacq.) Haw. [= Euphorbia clava Jacq.]. J. 20: 412. 1836. Ann.Rep. MissouriBot. Gard. 11: 85. 1899; Poinsettia Grahm,Edinb. New Phil. TYPE: Poinsettia pulcherrima(Willd.) Graham [ Pax, Bot. Jahrb.34: 61. 1905; Berger,Suk- Euphorbia pulcherrimaWilld.]. kulenteEuphorbien 1. 1907; N. E. Brown, Lacanthis Rafinesque,Fl. Tell. 2: 94. 1837. TYPE: La- Fl. Trop. Afr.6(1): 470. 1911; Fl. Capensis canthis splendens(Boj.) Raf. [= Euphorbia splen- 5(2): 222. 1925; Pax & Hoffm.,Natiirl. dens Boj. = E. milii Desm.]. Rafinesque,Fl. Tell. 4: 116. 1838. TYPE: Eu- 208. 1931; Prok- Agaloma Pflanzenfam.ed. 2, 19c: phorbia corollata L. [lectotype]. hanov,Consp. Syst. Tithymal.Asiae Mediae. AlectoroctonumSchlecht., Linnaea 19: 252. 1847. TYPE: 1933; Croizat,Bull. Jard.Bot. BuitenzorgIII. Alectoroctonumscotanum Schlecht. [= Euphorbia 16: 351. 1940; Hurusawa,J. Jap. Bot. 16: scotana Schlecht.]. 330. 1940; White, A., R. A. Dyer & B. L. AnthacanthaLemaire, Ill. Hort.4: Misc. 73. 1857. TYPE: Euphorbia heptagona L. [lectotype]. Sloane, Succul. Euphorbieae.1941; Wheeler, AdenopetalumKlotzsch & Garcke, Monatsber.K6nigl. Amer. Midl. Nat. 30: 478. 1943; Leandri, Preuss. Akad. Wiss. Berlin1859: 250. 1859 (non Not. Syst. Paris 12: 64. 1946; Vindt,Trav. Turcz., 1858). TYPE: Euphorbia graminea Jacq. Inst. Sci. Ch6rifien6: 23. 1953; Ursch & [lectotype]. Monatsber.K6nigl. Preuss. Leandri, MWm.Inst. Sci. Madag. 5B: 109. ArthrothamnusKl. & Gcke., Akad.Wiss. Berlin1859: 251. 1859 (nonRuprecht, 1954; Dyer, Bull. Jard. Bot. Etat 27: 487. 1848). TYPE: Arthrothamnustirucalli (L.) Kl. & 1957; Dressler,Ann. MissouriBot. Gard. 48: Gcke. [= Euphorbia tirucalliL.]. 329. 1961; Croizat,Webbia 20: 573. 1965; DichrophyllumKlotzsch & Garcke,Monatsber. K6nigl. Webster,J. ArnoldArbor. 49: 395. 1967; Preuss.Akad. Wiss. Berlin1859: 249. 1859. TYPE: Dichrophyllummarginatum (HBK) Kl. & Gcke. Radcliffe-Smith& Tutin, Fl. Europaea 2: 213. [= Euphorbia marginata HBK]. 1968; Allem& Irgang,Fl. Ilustr.Rio Grande EuphorbiastrumKlotzsch & Garcke,Monatsber. K6nigl. do Sul 11: 15. 1975; M. C. Johnston,Wright- Preuss.Akad. Wiss. Berlin1859: 252. 1859. TYPE: ia 5: 120. 1975; Leach, Dinteria12: 1. 1976; Euphorbiastrumhoffimannianum Kl. & Gcke. [= (Kl. & Gcke.) Boiss.; lec- Hassall, Austral.J. Bot. 25: 430. 1977; Su- Euphorbia hoffmanniana totype]. bils, Kurtziana10: 83. 1977; Carter,Hook. SterigmantheKlotzsch & Garcke, Monatsber.K6nigl. Ic. P1. 39: 5. 1982; Huft,Ann. MissouriBot. Preuss.Akad. Wiss. Berlin1859: 252. 1859. TYPE: Gard. 71: 1021. 1984; McPherson& Tirel, Euphorbia splendens Boj. [= Euphorbia milii Fl. Nouv.-Caled. 14(1): 10. 1987; Gilbert, Desm.; lectotype]. TithymalopsisKlotzsch & Garcke, Monatsber.K6nigl. Kew Bull. Kew Bull. 42: 231. 1987; Carter, Preuss.Akad. Wiss. Berlin1859: 249. 1859. TYPE: 42: 673. 1987; Fl. E. Trop. Afr.,Euphorb. Euphorbia corollata L. [lectotype]. 2: 409. 1988; Deil & Miuller-Hohenstein,Eu- TrichosterigmaKlotzsch & Garcke,Monatsber. K6nigl. phorbiaJ. 5: 109. 1988; Lebrun & Stork, Preuss.Akad. Wiss. Berlin1859: 248. 1859. TYPE: Enum. P1. Afr.Trop. 1: 213. 1991; Mayfield, Euphorbiafulgens Karw. [lectotype]. Lyciopsis (Boissier)Schweinfurth, Beitr. Fl. Aethiop.1: Sida 14: 573. 1991; Lewton,Euphorbia J. 8: 37. 1867 (non Spach, 1835). TYPE: Euphorbia cu- 113. 1992. TYPE: Euphorbia antiquorum L. neata Vahl. (lectotype,designated by Millspaugh,Publ. Elaeophorbia Stapf,in Johnston,Liberia 2: 646. 1906. Field ColumbianMus., Bot. 2: 306. 1909]. TYPE: Elaeophorbia drupacea (Schum.) Stapf [ Euphorbia drupacea Schum.]. TithymalusGaertner, Fruct. 2: 115. 1790, nom. cons. DiplocyathiumH. Schmidt,Beih. Bot. Centralbl.22(1): TYPE: Tithymaluspeplus (L.) Gaertner(= Euphor- 40. 1907. TYPE: Diplocyathiumcapitulatum (Rchb.) bia peplus L.]. Schmidt[= Euphorbia capitulata Rchb.]. Dactylanthesanacantha (Ait.) Haw. [= Euphorbia an- EuphorbiodendronMillspaugh, Publ. Field Columbian acantha Ait; lectotype]. Mus. Bot. 2: 305. 1909. TYPE: Euphorbiodendron Esula (Persoon) Hayworth,Syn. P1. Succul. 153. 1812 latazii (HBK) Millsp.[= Euphorbia latazii HBK; (nonMorandi, 1761). TYPE: Esula dalechampiiHaw. lectotype,chosen here; perhaps= Euphorbia laur- [= Euphorbia esula L.]. ifolia Lam.]. Galarhoeus Haworth,Syn. P1. Succul. 143. 1812. TYPE: Ctenadena Prokhanov,Consp. Syst. Tithymal. Asiae Me-

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diae 28. 1933. TYPE: Ctenadena lanata (Sieb.) acters that support generic status for Chamaesyce. Prokh.[= Euphorbia lanata Sieb.]. However, it must be admitted that Euphorbia sub- Consp.Syst. Tithymal. Asiae CystidospermumProkhanov, sect. Acutae is in many ways intermediatebetween Mediae 25. 1933. TYPE: Cystidospermumcheiro- By lepis (Fisch. & Mey.) Prokh. [= Euphorbia chei- Chamaesyce and Euphorbia subg. Agaloma. rolepis Fisch. & Mey.]. strictlyphylogenetic criteria, conservative workers SclerocyathiumProkhanov, Consp. Syst. Tithymal. Asiae are correct in retainingChamaesyce as a subgenus Mediae 30. 1933. TYPE: Sclerocyathiumpopovii of Euphorbia, because its recognition makes the Prokh. [= Euphorbia sclerocyathiumKorov. & latter a paraphyletic group. An obvious solution to Pop.]. this is to further dismantle Euphorbia into seg- A vast genus of over 1000 species, subdivided regate genera, in the tradition of Klotzsch and into many subgenera and sections, a number of Garcke. However, this would not get workers using which have been treated as distinct genera. The cladistic methodology off the hook, because a large number of synonyms cited here (lectotypifi- monophyletic Chamaesyce would probably have to cations by Wheeler, 1943) represents only a frac- include both Euphorbia subg. Agaloma as well as tion of the published names that are referable to Pedilanthus and possibly Cubanthus. Until a de- Euphorbia; they have been chosen because in the tailed phylogenetic analysis of the Euphorbieae is literature they have been treated as distinct at the completed, we have no practical alternative to congeneric or subgeneric level. The literature on Eu- tinuing the use of the present artificial (but rea- phorbia, especially the succulent taxa, is vast, and sonably usable) system. it is not possible to cite here even a fraction of the relevant papers, including many important papers 306. Cubanthus (Boissier) Millspaugh, Field on succulent taxa by Leach and Rauh. Many ref- Mus. Nat. Hist. Bot. 2: 371. 1913; Alain, Fl. erences are provided by Wheeler (1943), Vindt Cuba 3: 127. 1953. TYPE: Cubanthus linear- (1960), Webster (1967), Carter & Radcliffe-Smith ifolius (Griseb.) Millsp. [Pedilanthus linear- (1988), and Lebrun & Stork (1991); and numbers ifolius Griseb.]. of taxa are discussed and illustrated in the eight A genus of threespecies endemicto the West volumes of the Euphorbia Journal. Nevertheless, Indies (Cuba and Hispaniola); very similar to spe- there is great need for a comprehensive biblio- cies of Euphorbia sect. Adenorima, and perhaps graphic guide to the taxonomic work on Euphorbia to be included in that group. that would complement the list of species of Oude- jans (1990). Some segregates, especially Poinset- tia, have been adopted by a number of workers, 307. Monadenium Pax, Bot. Jahrb. 19: 126. and it remains to be seen if futurework will confirm 1894; N. E. Brown, Fl. Trop. Afr. 6(1): 450. the conservative generic circumscription adopted 191 1; Pax & Hoffm., Natiirl. Pflanzenfam. by Boissier and his followers. ed. 2, 19c: 222. 1931; Bally, Genus Mona- denium, 14. 1961; Carter, Kew Bull. 42: 903. 1987; Fl. E. Trop. Afr., Euphorb. 2: 305. Chamaesyce S. F. Gray, Nat. Arrang. Brit. 540. 1988. TYPE: Monadenium coccineum P1. 2: 260. 1821; Croizat & Degener, Fl. Pax. Haw. 1936; Wheeler, Rhodora 43: 97. 1941; Burch, Ann. Missouri Bot. Gard. 53: 90. 1966; StenadeniumPax, Bot. Jahrb.30: 343. 1901. TYPE: Webster, J. Arnold Arbor. 48: 420. 1967; Stenadeniumspinescens Pax [= Monadeniumspi- Hassall, Austral. J. Bot. 24: 633. 1976; Mul- nescens(Pax) Bally]. ligan & Lindsay, Naturaliste Canad. 105: 37- An African genus of ca. 50 species, distributed 40. 1978; Koutnik, S. Afr. J. Bot. 3: 262. from Somalia to the Transvaal. 1984; Allertonia 4(6): 331. 1987. TYPE: Chamaesyce maritima S. F. Gray [= Cha- 308. Synadenium Boissier, DC. Prodr. 15(2): maesyce peplis (L.) Prokh.]. 187. 187. 1862; N. E. Brown, Fl. Trop. Afr. AnisophyllumHaworth, Syn. P1. Succ. 159. 1812 (non 6(1): 462. 1911; Pax & Hoffm., Natiirl. Jacquin,1763). TYPE: Anisophyllumpeplis (L.) Haw. Pflanzenfam. ed. 2, 19c: 221. 1931; White, [= Chamaesycepeplis (L.) Prokh.]. Dyer, & Sloane, Succ. Euphorbieae 951. A widely distributed genus of ca. 250 species, 1941; Carter,Kew Bull. 42: 667. 1987; Fl. most of them in tropical America and Africa; often E. Trop. Afr.,Euphorb. 2: 534. 1988. TYPE: treated as a subgenus of Euphorbia. Koutnik (1987) Synadenium arborescens Boissier [= Syna- presented the most detailed analysis of the char- denium cupulare (Boiss.) L. C. Wheeler].

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A genus of 10-15 species of eastern and south- ed. 2, 19c: 52. 1931; Leonard, Bull. Jard. ern Africa, closely related to Monadenium. Bot. Nat. Belg. 59: 319. 1989. TYPE: Mar- tretia quadricornis Beille. 309. Endadenium Leach, Garcia de Orta 1: This west Africangenus of two species was placed 31. 1973. TYPE: Endadenium gossweileri (N. next to Aporusa by Pax & Hoffmann(1931), but E. Brown) Leach. J. Leonard (1989) proposed that it be placed in a A monotypic genus of Angola; perhaps not dis- new tribe Martretieae K6hler ex Leonard, within tinct from the genus Monadenium. subfamilyPhyllanthoideae. Meeuse (1990) regarded Martretia as more appropriately included in the 310. Pedilanthus Necker ex Poiteau, Ann. Mus. Acalyphoideae (Chrozophoreae). Nat. Hist. Nat. 19: 388. 1812; Boissier, DC. Prodr. 15(2): 4. 1862; Millspaugh, Field Mus. 313. Tacarcuna Huft, Ann. Missouri Bot. Gard. Publ. Bot. 2: 353. 1913; Pax & Hoffm., 76: 1080. 1989. TYPE: Tacarcuna gentryi Natiirl. Pflanzenfam. ed. 2, 19c: 223. 1931; Huft. Wheeler, Contr. Gray Herb. 124: 43. 1939; Dressler, Contr. Gray Herb. 182: 1. 1957; This recently described genus of three species Webster, J. Arnold Arbor. 48: 427. 1967; fromPanama and South America was proposed by Koutnik, Euphorbia J. 3: 38. 1985. TYPE: the author without any indication of affinity,al- Pedilanthus tithymaloides (L.) Poit. though the description of uniovulate locules of the ovary, togetherwith the general aspect, would sug- TithymalusMiller, Gard. Dict. Abr. Ed. 4. 1754. Tithy- gest subfamily Acalyphoideae. However, exami- maloides GomezOrtega, Tabl. Bot. 9. 1773 (nom. nation of material of an isotype of the type species rej.). CrepidariaHaworth, Syn. P1. Succ. 136. 1812. TYPE:Tithymalus myrtifolius Miller [= Pedilanthus (Gentry 16869, DAV) shows that the ovary has tithymaloides(L.) Poit.]. biovulate locules. Althoughfloral structure suggests VentenatiaTrattinick, Gen. P1. 86. 1802 (nonCavanilles, that Tacarcuna belongs to subfamily Phyllanthoi- Trat. [ 1798). TYPE:Ventenatia bracteata (Jacq.) deae, and perhaps to tribe Wielandieae (where it Pedilanthus bracteatus(Jacq.) Boiss.]. Diadenaria Klotzsch & Garcke, Monatsber. K6nigl. may be close to Savia), scanning micrographs of Preuss.Akad. Wiss. Berlin1859: 254. 1859. TYPE: pollen (courtesy of Joan Nowicke; Webster, ined.) Diadenaria pavonis K1. & Gcke. [= Pedilanthus show a striate exine suggestive of pollen in other bracteatus(Jacq.) Boiss.]. tribes such as Antidesmeae or Briedelieae. Hexadenia Klotzsch& Garcke,Monatsber. K6nigl. Preuss. Akad. Wiss. Berlin1859: 253. 1859. TYPE:Hex- adenia macrocarpa (Benth.) K1. & Gcke. [= Pe- 314. Avellanita R. A. Philippi, Linnaea 33: 27. dilanthus macrocarpusBenth.]. 1864. TYPE: Avellanita bustillosii R. A. Phi- A neotropical genus of ca. 15 species, all except lippi. one confinedto Mexico and adjacent Central Amer- The systematic position of this genus has re- ica. mained questionable up to the present. In his original description,Philippi assigned Avellanita to the INCERTAE SEDIS "Crotoneae," and Bentham (1880), on the basis of that desciption alone, placed it in tribe Crotoneae 311. Chlamydojatropha Pax & Hoffmann, between Micrandra and Hevea. Pax (1910), fol- Pflanzenr. 147. VI (Heft 57): 125. 1912; lowingthese earlier dispositions,located Avellanita Natiirl. Pflanzenfam., ed. 2, 19c: 167. 1931. withintribe Jatropheae followingGarcia; however, TYPE: Chlamydojatropha kamerunica Pax & he noted that its habit was aberrant and its leaves Hoffm. suggestive of Argythamnia. Pax & Hoffmann This mysteriousplant, known only frompistillate (1931), in an abrupt shift of opinion, assigned specimens from Cameroon, was placed next to Avellanita to tribe Acalypheae, in juxtaposition Mildbraedia by Pax & Hoffmann (1931). Until with the Old World genera Trewia and Melano- the staminate flowers are described, it cannot be lepis. Croizat, in manuscript notes (1960) on a definitelyplaced. specimen in the Chilean Museo Nacional de His- toria Natural (SGO), remarked on the vegetative 312. Martretia Beille, Compt. Rend. Hebd. Se- similarityof Avellanita to genera of Chrozopho- ances Acad. Sci. (Paris) 145: 1294. 1907; reae; however, the apetalous flowers and polyan- Bull. Soc. Bot. France 55 (Mem. 2) 8: 64. drous androecium of the Chilean plant would be 1908; Pax & Hoffmann,Natiirl. Pflanzenfam. quite discordant with that tribe.

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Examination of scanning micrographs from a of Bischofia). Examination of authentic material specimen in the Museo Nacional (SGO 051523) (kindly furnished by Rizzini) shows that the plant clearly contradicts an assignment of Avellanita to belongs to the Meliaceae and, followingthe treat- subfamilyCrotonoideae, as the exine does not show ment of Pennington (1981), clearly represents Tri- a Croton-pattern.The 3-colporate grains withrath- chilia lepidota Mart. subsp. lepidota. er coarse muri topped by nanospinules are somewhat similar to those in tribe Adelieae subfam. 2. Casabitoa Alain, Phytologia 47: 175. 1980; Acalyphoideae, but they are also not dissimilar to Fl. Espafiola 4: 92. 1986. TYPE: Casabitoa the grains found in tribe Acalypheae, e.g., Trewia. perfae Alain. Although the precise affinityof Avellanita remains This inadequately characterized genus, based on ambiguous, the evidence from pollen and floral pistillate specimens of a single species from His- characters clearly points to its placement within paniola, is not euphorbiaceous. According to Tom subfamily Acalyphoideae. Zanoni (in press), who visited the type locality, the plant is a species of Simarubaceae (Picramnia dic- 315. Cubacroton Alain, Candollea 17: 116. tyoneura (Urb.) Urb.). 1960. TYPE: Cubacroton maestrense Alain.

I have not been able to examine material of this 3. Polygonanthus Ducke, Notizbl. Bot. Gart. enigmatic genus, which is known only from the Berlin 11: 345. 1932. TYPE: Polygonanthus type collection in the Sierra de Maestra, Cuba. amazonicus Ducke. Judging from the description, the plant probably The genus is not euphorbiaceous, although its belongs in tribe Crotoneae; it appears similar to systematic position has been controversial. Al- the Cuban endemic genus Moacroton, and possibly though Baehni & Dansereau (1939) referredPoly- may prove to be an aberrant species of that genus. gonanthus to the Saxifragaceae (sensu lato), placement in the Anisophylleaceae by Pires and ADDENDUM Rodrigues (1971) probably has the most support 316. Dendrothrix Esser, Novon 3: 245. 1993. (e.g., in Cronquist, 1981). TYPE: Dendrothrix yutajensis (Jabl.) Esser. 4. Pseudocroton Muell. Arg., Flora 55: 24. This genus, referred by its author to subtribe 1872; Huft, Ann. Missouri Bot. Gard. 69: Mabeinae, was published too late to be accom- 429. 1982. TYPE: Pseudocroton tinctorius [ modated in the main sequence of this synopsis. It Capparis indica (L.) Fawc. & Rend.]. is related to Senefelderopsis, and thereforeshould be inserted in subtribe Mabeinae following that 5. Skutchia Pax & Hoffmann, J. Wash. Acad. genus. Esser provides several keys to distinguish Sci. 27: 306. 1937; Burger, Fieldiana Bot. Dendrothrix from the other genera of Mabeinae. 40: 212. 1977. TYPE: Skutchia caudata Pax & Hoffm. [= Trophis mexicana (Liebm.) 317. Cladogelonium Leandri, Bull. Soc. Bot. Bur.]. France 85: 530. 1939. TYPE: Cladogelonium madagascariense Leandri. 6. Trisyngyne Baillon, Adansonia 11: 136. This monotypicgenus fromMadagascar remains 1874; Bauman-Bodenheim, Bull. Mus. Hist. poorly known. Judging from the description and Nat. Paris II. 25: 420. 1953; Steenis, Adan- illustration, it does appear to be similar to the sonia II. 1 1: 615. 1971. TYPE: not designated widespread paleotropical genus Suregada, so it (Baillon published 2 species: Trisyngyne bal- may prove to belong to tribe Gelonieae. ansae and T codonandra; but neither Bau- man-Bodenheim nor van Steenis designated a type). EXCLUDEDTAXA The New Caledonian genus Trisyngyne was re- 1. Burseranthe Rizzini, Leandra 4/5: 5. 1974. duced by van Steenis (1971) to Nothofagus sub- TYPE: Burseranthe pinnata Rizz. sect. Bipartitae. This monotypic genus, based on a single species from eastern Brazil (Alagoas), was referredto subfamily Phyllanthoideae by Rizzini. The plant, how- LITERATURE CITED ever, has pinnate leaves, which are otherwise un- AIRY SHAW, H. K. 1961. Notes on Malaysian Eu- known in the family (except for the dubious case phorbiaceae.Kew Bull. 14: 353-397.

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. 1963. A synopsisof the genus Ptychopyxis la regionamazonienne. III. Arq. Jard.Bot. Rio Ja- Miq. Kew Bull. 14: 363-374. neiro4: 1-208. . 1965. Diagnosesof newfamilies, new names, ERDTMAN, G. 1952. PollenMorphology and PlantTax- etc. forthe seventhedition of Willis's"Dictionary." onomy.Almqvist & Wiksells,Uppsala. Kew Bull. 18: 249-272. FARR, E. R., J. A. LEUSSINK & F. A. STAFLEU (editors). . 1966. Willis' Dictionaryof the Flowering 1979. Index NominumGenericorum (Plantarum). Plantsand Ferns,Addenda [pp. xxi,xxii]. Cambridge 3 vols. Bohn,Scheltema & Holkema,Utrecht. Univ. Press, Cambridge. FORMAN, L. L. 1966. The reinstatementof Galearia * 1967. Notes on Malaysianand otherAsiatic Zoll. et Mor. and MicrodesmisHook. f. in the Pan- Euphorbiaceae.Kew Bull. 20: 379-414. daceae. Kew Bull. 20: 309-321. * "1971" [1972]. The Euphorbiaceaeof Siam. 1971. A synopsisof Galearia Zoll. and Mor. Kew Bull. 26: 191-363. (Pandaceae). Kew Bull. 26: 153-165. 1974. NoteworthyEuphorbiaceae from trop- GAGNEPAIN, F. & L. BEILLE. 1926. Euphorbiacees. Fl. ical Asia (Burmato NewGuinea). Hook. Ic. P1. 38(1): GeneraleIndo-Chine 5: 373-516. 3701-3725. GEHRMANN. 1908. Vorarbeitenzu einer Monographie . 1975. The Euphorbiaceaeof Borneo. Kew der GattungBridelia mit besondererBeriicksichti- Bull. Add. Ser. 4: 1-245. gungder afrikanischenArten. Bot. Jahrb.41(Beibl. . 1980. A partial synopsisof the Euphorbi- 95): 1-42. aceae-Platylobeae ofAustralia (excluding Phyllan- GILG, E. 1908. FlacourtiaceaeAfricanae. In: A. Engler thus,Euphorbia, and Calycopeplus).Kew Bull. 35: (editor),Beitrage zur Flora von Afriken32. Bot. 577-700. Jahrb.40: 444-518. * 1981. The Euphorbiaceaeof Sumatra.Kew GILLESPIE, L. 1993. A synopsisof neotropicalPluke- Bull. 36: 239-374. netia(Euphorbiaceae) including two new species. Syst. BAEHNI, C. & P. DANSEREAU. 1939. Polygonanthus, Bot. 18: 575-592. genre de Saxifragacees.Bull. Soc. Bot. France 86: GRUNING, G. 1913. 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LEANDRI, J. 1958. Euphorbiacees.Fl. Madagascar 11 1: . 1874. Euphorbiaceae[part 2]. In: Martius, 1-209. Fl. Brasiliensis11(2): 293-750. LEBRUN, J.-P. & A. L. STORK. 1991. Enumerationdes OUDEJANS, R. C. H. M. 1990. World Catalogue of Plantesa Fleursd'Afrique Tropicale, Vol. I. General- Species Names Publishedin the Tribe Euphorbieae ites et Annonaceae a Pandaceae. Conservatoireet (Euphorbiaceae)with Their GeographicalDistribu- JardinBotaniques de la Ville de Geneve,Geneva. tion.Oudejans, Utrecht[self-published]. LEONARD, J. 1951. Especes congolaisesnouvelles ou PAX, F. 1890. Euphorbiaceae. In: A. Engler & K. interessantes.Bull. Soc. Roy. Bot. BeIg. 84: 47-60. Pranti,Die NatiirlichenPflanzenfamilien, ed. 1, 3(5): 1956. NotulaeSystematicae 21. Observations 1-119. surles genresOldfieldia, Paivaeusa et Cecchia (Eu- * 1910. Euphorbiaceae-Jatropheae.In: A. En- phorbiaceaeafricanae). Bull. Jard. Bot. Brux. 26: gler (editor),Das PflanzenreichIV. 147. [I] (Heft 335-343. 42): 1-148. 1959. 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Engler(editor), Das PflanzenreichIV. Flored'Afrique Centrale: Spermatophytes. Jardin Bo- 147. IX. (Heft68): 1-108. taniqueNational de Belgique,Bruxelles. & . 1922. Euphorbiaceae-Phyllan- LEVIN, G. A. 1986a. Systematicfoliar morphology of theae. In: A. Engler(editor), Das PflanzenreichIV. Phyllanthoideae(Euphorbiaceae), I. Conspectus.Ann. 147. XV. (Heft81): 1-349. MissouriBot. Gard. 73: 29-85. & . 1931. Euphorbiaceae.In: A. En- *1 986b. Systematicfoliar morphology of Phyl- gler & K. Prantl(editors), Die NatiirlichenPflan- lanthoideae(Euphorbiaceae). II. Pheneticanalysis. zenfamiliened. 2, 19c: 11-233. Ann. MissouriBot. Gard. 73: 86-98. PENNINGTON, T. D. 1981. A monographof neotropical *1 986c. Systematicfoliar morphology of Phyl- Meliaceae. Fl. Neotrop.28: 1-470. lanthoideaeIII. Cladisticanalysis. Syst. Bot. 11: 515- PFEIFFER, L. 1874. NomenclatorBotanicus. T. Fischer, 530. Cassell. & M. G. SIMPSON. 1994. Phylogeneticimpli- PIRES, J. M. & W. A. RODRIGUES. 1971. Notas s6bre cationsof pollenultrastructure in the Oldfieldioideae os generos Polygonanthuse Anisophyllea. Acta (Euphorbiaceae).Ann. Missouri Bot. Gard.81: 203- Amazonica1(2): 7-15. 238. PRAIN, D. 1920. Euphorbiaceae[in part]. In: W. T. LINDLEY, J. 1836. A NaturalSystem of Botany,2nd Thiselton-Dyer(editor), Flora Capensis,Vol. 5, sect. ed. Longman,Rees, Orme,Brown, Green, and Long- 2: 410-516. man, London. PUNT, W. 1962. Pollen morphologyof the Euphorbi- LIOGIER, A. H. 1971. Novitatesantillanae. Mem. New aceae withspecial referenceto taxonomy.Wentia York Bot. Gard. 21: 107-157. 7: 1-116. RADCLIFFE-SMITH, A. 1973. An account of the genus MCPHERSON, G. & C. TIREL. 1987. Euphorbiacees. Fl. Nouv.-Caled.14(1): 1-226. CephalocrotonHochst. (Euphorbiaceae).Kew Bull. 28: 123-132. MCVAUGH, R. 1944. The genus Cnidoscolus:Generic . 1987a. Euphorbiaceae,Part 1. Fl. Trop. E. limitsand intragenericgroups. Bull. Torrey Bot. Club Afr.1-407. 71: 457-474. - 1987b. Segregatefamilies from the Euphor- MEEUSE, A. D. auct. J. 1990. The Euphorbiaceae plur.: biaceae. Bot. J. Linn. Soc. 94: 47-66. An Taxon. Delft. Unnatural Eburon, 1988. Notes on Madagascan Euphorbiaceae MENNEGA, A. M. W. 1987. Wood anatomyof the I. On the identityof Parageloniumand the affinities Euphorbiaceae,in particularof the subfamilyPhyl- ofBenoistia and Claoxylopsis(Euphorbiaceae). Kew lanthoideae.Bot. J. Linn. Soc. 94: 111-126. Bull. 43: 625-647. MERRILL, E. D. 1935. A commentaryon Loureiro's ROGERS, D. J. & S. G. APPAN. 1973. Manihot, Ma- "Flora Cochinchinensis."Trans. Amer.Philos. Soc. nihotoides(Euphorbiaceae). Fl. Neotrop.13: 1-272. 24: 1-49. ROTHMALER, W. 1944. Nominagenerica neglecta 1753- MILLER, K. & G. L. WEBSTER. 1962. Systematicpo- 1763. Repert.Sp. Nov. 53: 1-37. sitionof Cnidoscolus and Jatropha. Brittonia14: SCHULTES, R. 1952. Studiesin the genus Micrandra 174-180. I: The relationshipof the genus Cunuriato Micran- MUELLER, J. 1865. Euphorbiaceae. Linnaea 34: 1- dra. Bot. Mus. Leafl.Harvard Univ. 15: 201-222. 224. . 1970. The historyof taxonomicstudies in . 1866. Euphorbiaceae.In: DC., Prodromus Hevea. Bot. Rev. 36: 197-276. 15(2): 189-1261. 1990. A brieftaxonomic view of the genus

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Hevea. MalaysianRubber Research & Development WHEELER, L. C. 1943. The genera of livingEuphor- Board Monograph14: 1-57. bieae. Amer.Midl. Naturalist30: 456-503. SECCO,R. 1985. Notas sobre o novo conceitode Sa- . 1975. Euphorbiaceousgenera lectotypified. gotia racemosaem relacao as suas variedades.Acta Taxon 24: 534-538. Amazonica15(1-2, suppl.): 81-85. * 1988. Dialissepalia do genero Sandwithia APPENDIX 1. Outlineof taxa as treatedin the text. Lanj. (Euphorbiaceae):Uma novidadebotanica do alto Rio Negroe da Venezuela.Bol. Mus. Par. Emilio I. SubfamilyPHYLLANTHOIDEAE GoeldiBot. 4: 177-185. 1. Tribe WIELANDIEAE * 1990. Revisao dos generos Anomalocalyx 1. Heywoodia Ducke, Dodecastigma Ducke, Pausandra Radlk., 2. Savia Pogonophora Miersex Benth.e Sagotia Baill. (Eu- 3. Gonatogyne phorbiaceae-Crotonoideae)para a Americado Sul. 4. Petalodiscus Museu Paraense EmilioGoeldi, Belem. 5. Blotia & G. L. WEBSTER. 1990. Materiaispara a 6. Actephila floraamazonica IX. Ensaio sobre a sistematicade 7. Discocarpus generoRicheria. Bol. Mus. Para. EmilioGoeldi, Bot. 8. Lachnostylis 6: 141-158. 9. Chonocentrum STEENIS, C. G. G. J.VAN. 1971. Revisionof Nothojagus 10. Wielandia in New Caledonia.Adansonia II. 11: 615-624. 2. Tribe AMANOEAE TAKHTAJAN, A. L. 1980. Outlineof the classificationof 1 1. Pentabrachion floweringplants (Magnoliophyta). Bot. Rev. 46: 225- 12. Amanoa 359. 3. Tribe BRIEDELIEAE THIN, N. N. 1984. TribusAlchornieae (Euphorbiaceae) 13. Cleistanthus of VietnameseFlora. Tap Chi Sinh Hoc 6(3): 26- 14. Briedelia 29. 4. Tribe PHYLLANTHEAE 1988. Tribe Epiprineae(Muell. Arg.) Huru- 4a. SubtribeASTROCASIINAE sawa (Euphorbiaceae)in Vietnam.Tap Chi SinhHoc 15. Astrocasia 10(2): 30-33. 4b. SubtribeLEPTOPINAE . 1989. Tribus Codiaeae (Pax) Hutch. in Vi- 16. Leptopus etnam.Tap Chi Sinh Hoc 11(3): 14-17. 4c. Subtribe PSEUDOLACHNOSTYLIDI- THOMAS, D. W. 1990. Conceveiba Aublet(Euphorbi- NAE aceae) new to Africa.Ann. MissouriBot. Gard. 77: 17. Chascotheca 856-858. 18. Zimmermannia VINDT, J. 1960. Monographiedes Euphorbiacees du 19. Zimmermanniopsis Maroc. Index Bibliographique.Trav. Inst. Sci. Cher- 20. Meineckia if.,Bot. 19: 482-521. 21. Pseudolachnostylis WEBSTER, G. L. 1956. A monographicstudy of the 22. Keayodendron WestIndian species of Phyllanthus. J. Arnold Arbor. 4d. SubtribeSECURINEGINAE 37: 91-122, 217-268, 340-359. 23. Securinega * 1957. A monographicstudy of the West 4e. SubtribeANDRACHNINAE Indianspecies of Phyllanthus.J. Arnold Arbor. 38: 24. Andrachne 51-80, 170-198, 295-373. 4f. SubtribeFLUEGGEINAE 1958. A monographicstudy of the West 25. Flueggea Indianspecies of Phyllanthus.J. Arnold Arbor. 39: 26. Richeriella 49-100, 111-212. 27. Aerisilvaea . 1967. The genera of Euphorbiaceaein the 28. Margaritaria southeasternUnited States. J. ArnoldArbor. 48: 29. Phyllanthus 303-430. 30. Reverchonia 1975. Conspectusof a new classificationof 31. Sauropus the Euphorbiaceae.Taxon 24: 593-601. 32. Breynia 1983. A botanicalgordian knot: The case of 33. Glochidion Ateramnusand Gymnanthes(Euphorbiaceae). Tax- 5. Tribe DRYPETEAE on 32: 304-305. 34. Lingelsheimia * 1987. The saga of the spurges:A reviewof 35. Drypetes classificationand relationshipsin the Euphorbiales. 36. Sibangea Bot. J.Linn. Soc. 94: 3-46. 37. Putranjiva * 1992. Realignmentsin AmericanCroton (Eu- 6. Tribe ANTIDESMEAE phorbiaceae).Novon 2: 269-273. 6a. SubtribeSPONDIANTHINAE . 1993. A provisionalsynopsis of the sections 38. Spondianthus of the genus Croton (Euphorbiaceae).Taxon 42: 6b. SubtribeUAPACINAE 793-823. 39. Uapaca & W. S. ARMBRUSTER. 1991. A synopsisof 6c. SubtribeSCEPINAE the neotropicalspecies of Dalechampia (Euphorbi- 40. Protomegabaria aceae). Bot. J. Linn. Soc. 105: 137-177. 4 1. Maesobotrya & M. J. HUFT. 1988. Revised synopsisof 42. Richeria PanamanianEuphorbiaceae. Ann. Missouri Bot. Gard. 43. Jablonskia 75: 1087-1144. 44. Baccaurea

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45. Apodiscus 14. Tribe POGONOPHOREAE 46. Ashtonia 90. Pogonophora 47. Aporusa 15. Tribe CHAETOCARPEAE 6d. SubtribeANTIDESMINAE 91. Trigonopleura 48. Thecacoris 92. Chaetocarpus 49. Phyllanoa 16. Tribe PEREAE 50. Celianella 93. Pera 51. Leptonema 17. Tribe CHEILOSEAE 52. Antidesma 94. Cheilosa 53. Hyeronima 95. Neoscortechinia 6e. SubtribePORANTHERINAE 18. Tribe ERISMANTHEAE 54. Poranthera 96. Erismanthus 55. Oreoporanthera 97. Moultonianthus 7. Tribe HYMENOCARDIEAE 98. Syndyophyllum 56. Didymocistus 19. Tribe DICOELIEAE 57. Hymenocardia 99. Dicoelia 8. Tribe BISCHOFIEAE 20. Tribe GALEARIEAE 58. Bischofia 100. Microdesmis IncertaeSedis (withinPhyllanthoideae) 101. Galearia 59. Centroplacus 102. Panda 60. Meborea 21. Tribe AMPEREAE 103. Monotaxis II. SubfamilyOLDFIELDIOIDEAE 104. Amperea 9. Tribe CROIZATIEAE 22. Tribe AGROSTISTACHYDEAE 61. Croizatia 105. Agrostistachys 10. Tribe PODOCALYCEAE 106. Pseudagrostistachys 10a. SubtribePODOCALYCINAE 107. Cyttaranthus 62. Podocalyx 108. Chondrostylis TETRACOCCINAE 10b. Subtribe 23. Tribe CHROZOPHOREAE Tetracoccus 63. 23a. SubtribeSPERANSKIINAE 10c. SubtribePARADRYPETINAE 109. Speranskia Paradrypetes 64. 23b. SubtribeDITAXINAE 11. Tribe CALETIEAE 110. Caperonia 1la. SubtribeHYAENANCHINAE 111. Philyra 65. Hyaenanche 112. Ditaxis 1lb. SubtribeDISSILARIINAE 113. Argythamnia 66. Austrobuxus 114. Chiropetalum Dissiliaria 67. 23c. SubtribeDORYXYLINAE 68. Canaca 115. Doryxylon 69. Whyanbeelia 116. Sumbaviopsis 70. Choriceras 117. Thyrsanthera 71. Longetia 118. Melanolepis 11c. SubtribePETALOSTIGMATINAE 23d. SubtribeCHROZOPHORINAE 72. Petalostigma 119. Chrozophora PSEUDANTHINAE IId. Subtribe 24. Tribe CARYODENDREAE 73. Kairothamnus 120. Caryodendron 74. Scagea 121. Discoglypremna 75. Neoroepera 122. Alchorneopsis 76. Micrantheum 25. Tribe BERNARDIEAE 77. Pseudanthus 123. Bernardia 78. Stachystemon 124. Necepsia 12. Tribe PICRODENDREAE 125. Paranecepsia 12a. SubtribePICRODENDRINAE 126. Discocleidion 79. Celaenodendron 127. Adenophaedra Piranhea 80. 26. Tribe PYCNOCOMEAE Picrodendron 81. 26a. SubtribePYCNOCOMINAE 82. Parodiodendron 128. Pycnocoma PAIVAEUSINAE 12b. Subtribe 129. Droceloncia 83. Oldfieldia 130. Argomuellera MISCHODONTINAE 12c. Subtribe 26b. SubtribeBLUMEODENDRINAE 84. Aristogeitonia 131. Blumeodendron 85. Mischodon 132. Podadenia 86. Voatomalo 133. Ptychopyxis 87. Androstachys 134. Botryophora 88. Stachyandra 27. Tribe EPIPRINEAE III. SubfamilyACALYPHOIDEAE 27a. SubtribeEPIPRININAE 13. Tribe CLUTIEAE 135. Epiprinus 89. Clutia 136. Symphyllia

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137. Adenochlaena 31. Tribe PLUKENETIEAE 138. Cleidiocarpon 31a. SubtribePLUKENETIINAE 139. Koilodepas 188. Haematostemon 140. Cladogynos 189. Astrococcus 141. Cephalocrotonopsis 190. Angostyles 142. Cephalocroton 191. Romanoa 27b. SubtribeCEPHALOMAPPINAE 192. Eleutherostigma 143. Cephalomappa 193. Plukenetia 28. Tribe ADELIEAE 194. Vigia 144. Adelia 31b. SubtribeTRAGIINAE 145. Crotonogynopsis 195. Cnesmone 146. Enriquebeltrania 196. Megistostigma 147. Lasiocroton 197. Sphaerostylis 148. Leucocroton 198. Tragiella 29. Tribe ALCHORNEAE 199. Platygyna 29a. SubtribeALCHORNEINAE 200. Tragia 149. Orfilea 201. Acidoton 150. Bossera 202. Pachystylidium 151. Alchornea 31c. SubtribeDALECHAMPIINAE 152. Coelebogyne 203. Dalechampia 153. Aparisthmium 32. Tribe OMPHALEAE 154. Bocquillonia 204. Omphalea 29b. SubtribeCONCEVEIBINAE IV. SubfamilyCROTONOIDEAE 155. Conceveiba 33. Tribe MICRANDREAE 156. Gavarretia 33a. SubtribeMICRANDRINAE 157. Polyandra 205. Micrandra 30. Tribe ACALYPHEAE 206. Micrandropsis 30a. SubtribeRICININAE 207. Cunuria 158. Ricinus 33b. SubtribeHEVEINAE 30b. SubtribeADRIANINAE 208. Hevea 159. Adriana 34. Tribe MANIHOTEAE 30c. SubtribeMERCURIALINAE 209. Manihot 160. Mercurialis 210. Cnidoscolus 161. Seidelia 35. Tribe ADENOCLINEAE 162. Leidesia 35a. SubtribeADENOCLININAE 30d. SubtribeDYSOPSIDINAE 211. Glycydendron 163. Dysopsis 212. Klaineanthus 30e. SubtribeCLEIDIINAE 213. Tetrorchidium 164. Wetria 214. Adenocline 165. Cleidion 215. Ditta 166. Sampantaea 35b. SubtribeENDOSPERMINAE 30f. SubtribeMACARANGINAE 216. Endospermum 167. Macaranga 36. Tribe GELONIEAE 30g. SubtribeCLAOXYLINAE 217. Suregada 168. Erythrococca 37. Tribe ELATERIOSPERMEAE 169. Claoxylon 218. Elateriospermum 170. Claoxylopsis 38. Tribe JATROPHEAE 171. Mareya 219. Jatropha 172. Discoclaoxylon 220. Vaupesia 173. Micrococca 221. Oligoceras 174. Amyrea 222. Deutzianthus 30h. SubtribeLOBANILIINAE 223. Joannesia 175. Lobanilia 224. Leeuwenbergia 30i. SubtribeROTTLERINAE 225. Annesijoa 176. Mallotus 39. Tribe CODIAEAE 177. Deuteromallotus 226. Baloghia 178. Cordemoya 227. Hylandia 179. Coccoceras 228. Ostodes 180. Trewia 229. Pausandra 181. Neotrewia 230. Dodecastigma 182. Rockinghamia 231. Pantadenia 183. Octospermum 232. Dimorphocalyx 30j. SubtribeACALYPHINAE 233. Fontainea 184. Acalypha 234. Codiaeum 30k. SubtribeLASIOCOCCINAE 235. Sphyranthera 185. Lasiococca 236. Acidocroton 186. Spathiostemon 237. Blachia 187. Homonoia 238. Strophioblachia

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239. Sagotia 287. Adenopeltis 240. Baliospermum 288. Sapium 40. Tribe TRIGONOSTEMONEAE 289. Glyphostylus 241. Trigonostemon 290. Grimmeodendron 41. Tribe RICINOCARPEAE 291. Bonania 41 a. SubtribeRICINOCARPINAE 292. Duvigneaudia 242. Ricinocarpos 293. Maprounea 243. Alphandia 294. Hippomane 244. Beyeria 47. Tribe PACHYSTROMATEAE 41b. SubtribeBERTYINAE 295. Pachystroma 245. Bertya 48. Tribe HUREAE 246. Myricanthe 296. Tetraplandra 247. Cocconerion 297. Algernonia 248. Borneodendron 298. Ophthalmoblapton 42. Tribe CROTONEAE 299. Hura 249. Mildbraedia 49. Tribe EUPHORBIEAE 250. Fahrenheitia 49a. SubtribeANTHOSTEMINAE 251. Moacroton 300. Anthostema 252. Croton 301. Dichostemma 43. Tribe RICINODENDREAE 49b. SubtribeNEOGUILLAUMINIINAE 253. Givotia 302. Neoguillauminia 254. Ricinodendron 303. Calycopeplus 255. Schinziophyton 49c. SubtribeEUPHORBIINAE 44. Tribe ALEURITIDEAE 304. Euphorbia 44a. SubtribeALEURITINAE 305. Chamaesyce 256. Aleurites 306. Cubanthus 257. Reutealis 307. Monadenium 258. Vernicia 308. Synadenium 44b. SubtribeGARCIINAE 309. Endadenium 259. Garcia 310. Pedilanthus 44c. SubtribeGROSSERINAE 260. Cavacoa IncertaeSedis (withinEuphorbiaceae) 261. Grossera 262. Tapoides 311. Chlamydojatropha 263. Anomalocalyx 312. Martretia 264. Sandwithia 313. Tacarcuna Avellanita 265. Tannodia 314. Cubacroton 266. Domohinea 315. Dendrothrix 44d. SubtribeCROTONOGYNINAE 316. 267. Cyrtogonone 317. Cladogelonium 268. Crotonogyne APPENDIX 2. List of new taxa. 269. Manniophyton 44e. SubtribeNEOBOUTONINAE CROIZATIEAE Webster,trib. nov. 270. Neoboutonia LEPTOPINAE Webster,subtr. nov. 271. Benoistia PODOCALYCEAE Webster,trib. nov. V. SubfamilyEUPHORBIOIDEAE PODOCALYCINAE Webster,subtr. nov. 45. Tribe STOMATOCALYCEAE PYCNOCOMINAE Webster,subtr. nov. 45a. SubtribeSTOMATOCALYCINAE TETRACOCCINAE Levin,subtr. nov. 272. Plagiostyles DALECHAMPIINAE (Muell. Arg.) Webster,stat. nov. 273. Pimelodendron PICRODENDRINAE (Small) Webster,stat. nov. 45b. SubtribeHAMILCOINAE PORANTHERINAE (Muell. Arg.) Webster,stat. nov. 274. Hamilcoa SCEPINAE (Lindl.)Webster, stat. nov. 275. Nealchornea SPONDIANTHINAE (Webster)Webster, stat. nov. 46. Tribe HIPPOMANEAE Acidocrotonsect. Ophellantha(Standl.) Webster,comb. 46a. SubtribeMABEINAE nov. 276. Mabea Acidocrotonspinosus (Standl.) Webster,comb. nov. 277. Senefeldera Acidocrotonsteyermarkii (Standl.) Webster,comb. nov. 278. Senefelderopsis Adenopeltisserrata (Ait.) Webster,comb. nov. 46b. SubtribeCARUMBIINAE Deutzianthusthyrsiflorus (Airy Shaw) Webster, comb. 279. Omalanthus nov. 46c. SubtribeHIPPOMANINAE Gymnanthesschomburgkii (KI.) Webster,comb. nov. 280. Spirostachys Leptopusdiplospermus (Airy Shaw) Webster,comb. nov. 281. Sebastiania Leptopusphyllanthoides (Nutt.) Webster, comb. nov. 282. Stillingia Lingelsheimiamanongarivensis (Leandri) Webster, comb. 283. Gymnanthes nov. 284. Excoecaria Orfileamultispicata (Baillon) Webster, comb. nov. 285. Colliguaja Orfileaneraudiana (Baillon) Webster, comb. nov. 286. Dalembertia Pantadeniachauvetiae (Leandri) Webster, comb. nov.

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Phyllanthusanamalayanus (Gamble) Webster, comb. nov. Amanoa 12 Phyllanthuschevalieri (Gagnep.) Webster,comb. nov. AMANOEAE T2 Sauropuspierrei (Beille) Webster,comb. nov. Amanoinae T2 Amperea 104 APPENDIX 3. Index to genera and highertaxa. This AMPEREAE T21 indexincludes all the generaand highertaxa mentioned Amyrea 174 in this paper. Accepted taxa are shown in bold, with Anabaena 191 supragenerictaxa all capitalized.Synonymized taxa are Anabaenella 191 italicized,and thosemerely mentioned are shownin reg- Anaua 35 ular text.Taxa are numberedas theyappear in the text, Anda 223 withthose in Roman numeralsreferring to subfamilies, Andicus 223 those precededby "T" referringto tribesor subtribes, Andrachne 24 those precededby "E" referringto excludedtaxa, and ANDRACHNINAE T4e othersreferring to genera. Androstachys 87 Acalypha 184 Angelandra 252 Acalyphaceae III Angostyles 190 ACALYPHEAE T30 Angostylidium 193 ACALYPHINAE T30j Anisonema 29 ACALYPHOIDEAE III Anisophyllae T49c Acalyphopsis 184 Anisophyllum 305 Acanthocaulon 199 Annesijoa 225 Acantholoma 295 Anomalocalyx 263 Accia 194 Anomospermum 6 Acidocroton 236 Anomostachys 284 AcidotonP. Browne 25 Anthacantha 304 Acidoton Sw. 201 Anthostema 300 Aconceveibum 176 Anthostemeae T49a Actephila 6 Anthostemideae T49a Actinostemon 283 ANTHOSTEMINAE T49a Adelia 144 Antidesma 52 ADELIEAE T28 ANTIDESMEAE T6 Adenochlaena 137 ANTIDESMINAE T6d Adenocline 214 Aonikena 114 ADENOCLINEAE T35 Aparisthmium 153 ADENOCLININAE T35a Aphora 112 Adenocrepis 44 Apodandra 193 Adenogyne 281 Apodiscus 45 Adenogynum 140 Aporosella 29 Adenopeltinae T46c Aporusa 47 Adenopeltis 287 Aporuseae T6c Adenopetalum 304 Arachnodes 29 Adenophaedra 127 Argomuellera 130 Adenoropium 219 Argythamnia 113 Adisca 176 Aristogeitonia 84 Adriana 159 Arthrothamnus 304 Adrianeae T30b Ashtonia 46 ADRIANINAE T30b Asterandra 29 Aegopricum 293 Astraea 252 Aerisilvaea 27 Astrocasia 15 Agaloma 304 ASTROCASIINAE T4a AGROSTISTACHYDEAE T22 Astrococcus 189 Agrostistachys 105 Astrogyne 252 Agyneia 33 Astylis 35 Alchornea 151 Athroandra 168 Alchorneiformes T29a Athroisma 241 ALCHORNEAE T29 Austrobuxus 66 ALCHORNEINAE T29a Avellanita 314 Alchorneopsis 122 Axenfeldia 176 Alcinaeanthus 95 Baccaurea 44 Alcoceria 286 Baccaureopsis 48 Alectoroctonum 304 Baliospermum 240 Aleurites 256 Baloghia 226 ALEURITIDEAE T44 Barhamia 252 ALEURITINAE T44a Beltrania 146 Alevia 123 Bennettia 101 Algernonia 297 Bennettieaceae T20 Allenia 76 Bennettieae T20 Alphandia 243 Benoistia 271

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Benoistieae T44e Centrostylis 137 Bernardia 123 Cephalocroton 142 BERNARDIEAE T25 Cephalocrotoneae T27a Bernardiiformes T25 Cephalocrotonopsis 141 Bertya 245 Cephalomappa 143 Bertyeae T41 CEPHALOMAPPINAE T27b BERTYINAE T41b Ceramanthus 29 Bessera 25 Ceratococcus 193 Bestram 52 Ceratogynum 31 Beyeria 244 Ceratophorus 217 Beyeriopsis 244 Chaenotheca 17 Bia 200 CHAETOCARPEAE T15 Bischofia 58 Chaetocarpus 92 Bischofiaceae T8 Chamaesyce 305 BISCHOFIEAE T8 Chascotheca 17 Bivonea 210 Cheilosa 94 Bivonia 123 CHEILOSEAE T17 Blachia 237 Cheilosiformes T17 Bleekeria 151 Chiropetalum 114 Blotia 5 Chlamydojatropha 311 BLUMEODENDRINAE T26b Chlorocaulon 114 Blumeodendron 131 Chloropatane 168 Bocquillonia 154 Chondrostylis 108 Bonania 291 Chonocentrum 9 Borneodendron 248 Choriceras 70 Bossera 150 Choriophyllum 66 Botryanthe 194 Chorisandra 29 Botryophora 134 Chorisandrachne 16 Bradleja 33 Chorizotheca 77 Brexiopsis 35 Chrozophora 119 Breynia 32 CHROZOPHOREAE T23 Breyniopsis 31 CHROZOPHORINAE T23d Briedelia 14 Chrysostemon 77 BRIEDELIEAE T3 Cicca 29 Briedelioideae T3 Cladodes 151 Bureavia 66 Cladogyniformes T27a Burseranthe El Cladogelonium 317 Caelebogyne 152 Cladogynos 140 Caletia 76 Claoxyleae T30g CALETIEAE Tll Claoxyliformes T30g Calococcus 28 CLAOXYLINAE T30g Calpigyne 139 Claoxylon 169 Calycopeplus 303 Claoxylopsis 170 Calyptosepalum 35 Clarorivinia 133 Calyptroon 44 Clavistylus 196 Calyptrostigma 244 CLEIDIINAE T30e Canaca 68 Cleidiocarpinae T27a Candelabria 14 Cleidiocarpon 138 Caoutchoua 208 Cleidion 165 Capellenia 216 Cleistanthus 13 Caperonia 110 Cleodora 252 Caperonieae T23b Clonostylis 186 CARUMBIINAE T46b Clusiophyllum 207 Carumbium 279 Clutia 89 CARYODENDREAE T24 CLUTIEAE T13 Caryodendron 120 Cluytiandra 20 Casabitoa E2 Cluytieae T13 Castiglionia 219 Cluytiinae T 13 Cathetus 29 Cnemidostachys 281 Caturus 184 Cnesmone 195 Cavacoa 260 Cnidoscolus 210 Cecchia 83 Coccoceras 179. Celaenodendron 79 Coccoglochidion 33 Celianella 50 Cocconerion 247 Cenesmon 195 CODIAEAE T39 Centrandra 252 Codiaeinae T39 Centrodiscus 120 Codiaeum 234 Centroplacus 59 Coelebogyne 152

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Coelodisceae T30i Diplomorpha 31 Coelodiscus 176 Diplostylis 214 Collenucia 219 Discocarpus 7 Colliguaja 285 Discoclaoxylon 172 Colmeiroa 25 Discocleidion 126 Cometia 35, 48 Discoglypremna 121 Commia 284 Dispermae I Conami 29 Dissiliaria 67 Conceveiba 155 DISSILIARIINAE T1 lb Conceveibastrum 155 DITAXINAE 23b CONCEVEIBINAE T29b Ditaxis 112 Conceveibum 153 Ditrysinia 281 Conosapium 288 Ditta 215 Cordemoya 178 Dodecastemon 35 Corythea 184 Dodecastigma 230 Cremophyllum 203 Domohinea 266 Cremostachys 101 DORYXYLINAE T23c Crepidaria 310 Doryxylon 115 Croizatia 61 Drepadenium 252 CROIZATIEAE T9 Droceloncia 129 Croton 252 DRYPETEAE T5 CROTONEAE T42 Drypetes 35 Crotonogyne 268 Duvigneaudia 292 CROTONOGYNINAE T44d DYSOPSIDINAE T30d Crotonogynopsis 145 Dysopsis 163 CROTONOIDEAE IV Echinocroton 176 Crotonopsis 252 Echinus 176 Ctenadena 304 Elachocroton 281 Ctenomeria 200 Elaeophora 193 Cubacroton 315 Elaeophorbia 304 Cubanthus 306 ELATERIOSPERMEAE T37 Cunuria 207 Elateriospermum 218 Curcas 219 Eleutherostigma 192 Cyathogyne 48 Emblica 29 Cyclostemon 35 Enchidium 241 Cyclostemoneae T5 Endadenium 309 Cyclostemone's T5 ENDOSPERMINAE T35b Cyclostigma 252 Endospermum 216 Cyrtogonone 267 Enriquebeltrania 146 Cystidospermum 304 EPIPRINEAE T27 Cyttaranthus 107 EPIPRININAE T27a Dactylanthes 304 Epiprinus 135 Dactylostemon 283 Epistylium 29 Dalechampia 203 Eraclissa 24 Dalechampieae T31c Eremocarpus 252 DALECHAMPIINAE T31 c Eriococcus 29 Dalembertia 286 ERISMANTHEAE T18 Danguyodrypetes 34 Erismanthus 96 Decarinium 252 Erythrocarpus 217 Dendrocousinsia 281 Erythrococca 168 Dendrophyllanthus 29 Erytrochilus 169 Dendrothrix 316 Esula 304 Desfontaena 114 Euphorbia 304 Deuteromallotus 177 Euphorbiastrum 304 Deutzianthus 222 EUPHORBIEAE T49 Diadenaria 310 EUPHORBIINAE T49c Dibrachion 279 Euphorbiodendron 304 Dichelactina 29 EUPHORBIOIDEAE V Dichostemma 301 Euphyllantheae T4f Dichrophyllum 304 Eutropia 252 Dicoelia 99 Everettiodendron 44 DICOELIEAE T19 Excoecaria 284 Diderotia 149 Excoecariinae T46c Didymocistus 56 Fahrenheitia 250 Dimorphocalyx 232 Falconeria 288 Dimorphocladium 29 Flueggea 25 Diplochlamys 176 FLUEGGEINAE (or PHYLLANTHINAE) 4f Diplocyathium 304 Flueggeopsis 29

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Foersteria 32 Holstia 265 Fontainea 233 Homalanthinae T46b Fragariopsis 194 Homalanthus 279 Freireodendron 35 Homonoia 187 Friesia 252 Humblotia 35 Galarhoeus 304 Hura 299 Galearia 101 HUREAE T48 GALEARIEAE T20 Hyaenanche 65 Garcia 259 Hyaenancheae T1la GARCIINAE T44b HYAENANCHINAE Ti la Gatnaia 44 Hyeronima 53 Gavarretia 156 Hylandia 227 Geblera 25 Hymenocardia 57 GELONIEAE T36 Hymenocardiaceae T7 Gelonium 217 HYMENOCARDIEAE T7 Geminaria 29 Hypocoton 291 Genesiphylla 29 Irregulares T23 Gentilia 14 Jablonskia 43 Gitara 201 Janipha 209 Givotia 253 Jatropha 219 Glochidion 33 JATROPHEAE T38 Glochidionopsis 33 Jatrophinae T38 Glochisandra 33 Joannesia 223 Glycydendron 211 Joannesieae T38 Glyphostylus 289 Julocroton 252 Godefroya 13 Junghuhnia 234 Gonatogyne 3 Jussieula 210 Grimmeodendron 290 Kairothamnus 73 Grossera 261 Keayodendron 22 GROSSERINAE T44c Kirganelia 29 Guarania 42 Klaineanthus 212 Gussonia 281 Kleinodendron 2 Guya 35 Koelera 35 Gymnalypha 184 Koilodepas 139 Gymnanthes 283 Kunstlerodendron 108 Gymnanthinae T46c Kurziodendron 241 Gymnostillingia 282 Lacanthis 304 Gynamblosis 252 Lachnostylis 8 Gynoon 33 Laneasagum 35 Haematostemon 188 Lasiococca 185 Halliophytum 63 LASIOCOCCINAE T30k Hamilcoa 274 Lasiocroton 147 HAMILCOINAE T45b Lasiogyne 252 Hancea 176 Lasiostyles 165 Hasskarlia 123 Lassia 200 Hebecocca 204 Laurembergia 149 Hecatea 204 Lautembergia 149 Hedraiostylus 193 Lebidieropsis 13 Hemicicca 29 Leeuwenbergia 224 Hemicyclia 35 Leidesia 162 Hemiglochidion 33 Leiocarpus 47 Heptallon 252 Leiopyxis 13 Hermesia 15i Lepidanthus 16 Heterocalymnantha 31 Lepidostachys 47 Heterocalyx 105 Lepidoturus 151 Heterochlamys 252 Leptemon 252 Heterocroton 252 Leptonema 51 Hevea 208 LEPTOPINAE T4b HEVEINAE T33b Leptopus 16 Hexadena 29 Leptorhachis 200 Hexadenia 310 Leucandra 200 Hexaspermum 29 Leucocroton 148 Heywoodia 1 Limacia 35 Hieronymeae T6d Lingelsheimia 34 Hippocrepandra 103 Linostachys 184 Hippomane 294 Liodendron 37 HIPPOMANEAE T46 Liparena 35 HIPPOMANINAE T46c Lobanilia 175

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LOBANILIINAE T30h Myricanthe 246 Loerzingia 222 Myriogomphos 252 Longetia 71 Nageia 37 Loureira 219 Nanopetalum 13 Lumanaja 187 Nealchornea 275 Lyciopsis 304 Necepsia 124 Mabea 276 Nellica 29 MABEINAE T46a Neoboutonia 270 Macaranga 167 Neoboutonieae T44e Macarangeae T30f Neoboutoniiformes T44e MACARANGINAE T30f NEOBOUTONINAE T44e Macraea 29 Neochevaliera 92 Maesobotrya 41 Neogoetzea 14 Malloteae T30i Neoguillauminia 302 Mallotus 176 NEOGUILLAUMINIINAE T49b Mancanilla 294 Neoholstia 265 Mandioca 209 Neojatropha 249 Manihot 209 Neomanniophyton 268 MANIHOTEAE T34 Neomphalea 204 Manihotoides 209 Neopalissya 124 Manniophyton 269 Neoroepera 75 Mappa 167 Neoscortechinia 95 Maprounea 293 Neotrewia 181 Mareya 171 Neotrigonostemon 241 Mareyeae T30g Neowawraea 25 Mareyopsis 171 Nepenthandra 241 Margaritaria 28 Nephrostylus 139 Martretia 312 Niedenzua 137 Martretieae 312 Niruri 29 Meborea 60 Nymania 29 Mecostylis 167 Nymphanthus 29 Medea 252 Ocalia 252 Medusaea 304 Octospermum 183 Megabaria 38 Odonteilema 184 Megalostylis 203 Oldfieldia 83 Megistostigma 196 OLDFIELDIOIDEAE II Meineckia 20 Oligoceras 221 Melanolepis 118 Omalanthus 279 Melanthesa 32 Omphalandria 204 Melanthesopsis 32 Omphalea 204 Menarda 29 OMPHALEAE T32 Mercurialiiformes T30c Omphaleinae T32 MERCURIALINAE T30c Ophellantha 236 Mercurialis 160 Ophthalmoblapton 298 Mettenia 92 Orbicularia 29 Micrandra 205 Oreoporanthera 55 MICRANDREAE T33 Orfilea 149 Micrandriformes T33 Ostodes 228 MICRANDRINAE T33a Pachystemon 167 Micrandropsis 206 Pachystroma 295 Micrantheum 76 PACHYSTROMATEAE T47 Micrococca 173 Pachystromatinae T47 Microdesmis 100 Pachystylidium 202 Microelus 58 Paivaeusa 83 Microsepala 44 PAIVAEUSINAE T12b Microstachys 281 Palenga 37 Mildbraedia 249 Palissya 124 Mischodon 85 Panda 102 MISCHODONTINAE T12c Pandaceae T20 Moacroton 251 Pantadenia 231 Moeroris 29 Paracleistus 13 Molina 163 Paradenocline 214 Monadenium 307 Paradrypetes 64 Monotaxideae T21 PARADRYPETINAE T10c Monotaxis 103 Paragelonium 84 Moultonianthus 97 Paranecepsia 125 Mozinna 219 Parapantadenia 231 Muricococcum 143 Parodiodendron 82

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Passaea 123 Pseudagrostistachys 106 Pausandra 229 PSEUDANTHINAE TI Id Paxiuscula 112 Pseudanthus 77 Pedilanthaceae T49c Pseudocroton E4 Pedilanthinae T49c Pseudoglochidion 29 Pedilanthus 310 PSEUDOLACHNOSTYLIDINAE T4c Peltandra 20 Pseudolachnostylis 21 Pentabrachion 11 Pseudotragia 193 Pera 93 Psilostachys 165 Peraceae T16 Pterococcus 193 PEREAE T16 Ptychopyxis 133 Perula 93 Putranj iva 37 Petalodiscus 4 Putranjiveae T5 Petalostigma 72 Pycnocoma 128 Petalostigmateae TI Id PYCNOCOMEAE T26 PETALOSTIGMATINAE Tl lc PYCNOCOMINAE T26a Phaedra 123 Pycnosandra 35 Philyra 111 Quadrasia 169 Phocea 167 Ramelia 154 Phyllanoa 49 Ramsdenia 29 Phyllanthaceae I Redia 165 Phyllanthaceae T4 Regnaldia 92 PHYLLANTHEAE T4 Regulares T23 Phyllanthidea 24 Reidia 29 Phyllanthinae T4f Reutealis 257 Phyllanthodendron 29 Reverchonia 30 PHYLLANTHOIDEAE I Rhytis 52 Phyllanthus 29 Richeria 42 Phyllaurea 234 Richeriella 26 Picrodendraceae T 12 Ricinella 144 PICRODENDREAE T12 RICININAE T30a PICRODENDRINAE T12a Ricinocarpaceae T41 Picrodendron 81 RICINOCARPEAE T41 Pierardia 44 RICINOCARPINAE T41a Pierardieae T5 Ricinocarpoideae T41 Pilinophytum 252 Ricinocarpos 242 Pimelodendron 273 RICINODENDREAE T43 Piranhea 80 Ricinodendrinae T43 Plagianthera 176 Ricinodendron 254 Plagiostyles 272 Ricinus 158 Platygyna 199 Riseleya 35 Pleiostemon 25 Rockinghamia 182 Plukenetia 193 Roeperia 242 PLUKENETIEAE T31 Roigia 29 PLUKENETIINAE T31 a Romanoa 191 Podadenia 132 Ronnowia 204 PODOCALYCEAE T1O Rottlera 176 PODOCALYCINAE T1Oa ROTTLERINAE F30i Podocalyx 62 Sagotia 239 Podostachys 252 Sajorium 193 Poggeophyton 168 Samaropyxis 57 Pogonophora 90 Sampantaea 166 POGONOPHOREAE T14 Sandwithia 264 Pogonophyllum 205 Sapioideae V Poilaniella 241 Sapium 288 Poinsettia 304 Sarothrostachys 281 Polyandra 157 Sauropodeae T4f Polyboea 123 Sauropus 31 Polydragma 186 Savia 2 Polygonanthus E3 Scagea 74 Poranthera 54 Scepa 47 Poranthereae T6e Scepaceae T6c PORANTHERINAE T6e Scepasma 29 Porphyranthus 102 SCEPINAE T6c Prosartema 241 Schinziophyton 255 Prosopidoclineae T16 Schismatopera 93 Prosorus 28 Schistostigma 13 Protomegabaria 40 Schousboea - 151

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Sclerocyathium 304 Tetraglossa 165 Scortechinia 95 Tetraplandra 296 Sebastiania 281 Tetrorchidiinae T35a Securinega 23 Tetrorchidiopsis 213 SECURINEGINAE T4d Tetrorchidium 213 Seidelia 161 Thecacoris 48 Senefeldera 277 Thelypetalum 16 Senefelderopsis 278 Thyrsanthera 117 Serophyton 112 Tiglium 252 Shirakia 288 Timandra 252 Sibangea 36 Tithymalinae T49c Silvaea 241 Tithymaloides 310 Sinopimelodendron 138 Tithymalopsis 304 Siphonia 208 TithymalusGaertner 304 Skutchia E5 TithymalusMiller 310 Spathiostemon 186 Toxicodendrinae TI la Speranskia 109 Toxicodendrum 65 SPERANSKIINAE T23a Traganthus 123 Sphaerostylis 197 Tragia 200 Sphragidia 35 Tragiella 198 Sphyranthera 235 TRAGIINAE T31b Spirostachys 280 Tragiopsis 281 Spixia 93 Treisia 304 Spondiantheae T6a Trewia 180 SPONDIANTHINAE T6a Trewiaceae T30i Spondianthus 38 Trewiiformes T30i Stachyandra 88 Triadica 288 Stachystemon 78 Tricarium 29 Staphysora 41 Trichosterigma 304 Staurothyrax 29 Trigonopleura 91 Steigeria 226 Trigonostemon 241 Stelechanteria 35 TRIGONOSTEMONEAE T40 Stenadenium 307 Trisyngyne E6 Stenonia 112 Tritaxis 241 Sterigmanthe 304 Tylosepalum 241 Stilagineae T6, T6d Tyria 123 Stilaginella 53 Tzellemtinia 14 Stilago 52 Uapaca 39 Stillingia 282 Uapaceae T6b Stillingiinae T46c UAPACINAE T6b Stipellaria 151 Uranthera 29 STOMATOCALYCEAE T45 Vaupesia 220 STOMATOCALYCINAE T45a Veconcibea 155 Stomatocalyx 273 Ventenatia 310 Strophioblachia 238 Vernicia 258 Stylodiscus 58 Victorinia 210 Sumbavia 115 Vigia 194 Sumbaviopsis 116 Voatamalo 86 Suregada 217 Wartmannia 279 Symphyllia 136 Wetria 164 Synadenium 308 Wetriaria 130 Synapisma 234 Whyanbeelia 69 Syndyophyllum 98 Wielandia 10 Synexemia 29 WIELANDIEAE TI Synostemon 31 Wielandiidees TI Tacarcuna 313 Wielandiinae TI Taeniosapium 288 Williamia 29 Tandonia 265 Wurtzia 28 Tannodia 265 Xylophylla 29 Tapoides 262 Zarcoa 33 Telephioides 24 Zimpania 219 Telogyne 241 Zimmermannia 18 Tetracarpidium 193 Zimmermanniopsis 19 TETRACOCCINAE TlOb Zuckertia 200 Tetracoccus 63 Zygospermum 28 Tetraglochidion 33

Volume81, Number1, pp. 1-144 of the ANNALS OF THE MISSOURI BOTANICAL GARDEN was publishedon March 21, 1994.

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