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Synopsis of the Genera and Suprageneric Taxa of Euphorbiaceae

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Synopsis of the Genera and Suprageneric Taxa of Euphorbiaceae 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. This content downloaded from 169.237.8.36 on Mon, 29 Dec 2014 19:11:36 PM All use subject to JSTOR Terms and Conditions 34 Annals of the MissouriBotanical Garden 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 binucle- ate 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- This content downloaded from 169.237.8.36 on Mon, 29 Dec 2014 19:11:36 PM All use subject to JSTOR Terms and Conditions Volume 81, Number1 Webster 35 1994 Synopsis of Taxa of Euphorbiaceae metophyte binucleate; pistillode present or absent;
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