SystematicBotany (1992), 17(2): pp. 311-323 (? Copyright1992 by the American Society of Taxonomists

Revision of (Euphorbiaceae)

GRADY L. WEBSTER Department of Botany, University of , Davis, California 95616

ABSTRACT. Astrocasia,a neotropical belonging to the Euphorbiaceae, tribe , occupies an isolated position in the tribe; a new subtribe,subtribe Astrocasiinae, is described to accommodate it. The genus is interpretedas including five species, of which two require new combinations:Astrocasia jacobinensis and A. austinii. A key, descriptions,distribution maps, and citationsof representativespecimens are provided. Astrocasiatremula is shown to have a remarkable disjunct distribution,with populations recorded from the Greater Antilles, Yucatan peninsula, western , , and .

AstrocasiaRobinson and Millspaugh, a small (A. austinii,A. neurocarpa(Muell. Arg.) I. M. genus of fiveneotropical species, belongs to the Johnstonex Standley). subfam. Phyllanthoideae. The genus was de- A particularlyuseful diagnostic foliar char- scribed by Robinson and Millspaugh (in Mills- acter in Astrocasiais the cuticular wax pattern paugh 1905) on the basis of a plant collected in on the abaxial (under) side of the leaf blades. Yucatan by Seler. The authors noted that their As Rogers and Appan (1973) found forManihot new genus resembled species of PhyllanthusL., Miller, each species of Astrocasiahas a distinc- but differedin having a well-developed corolla. tive cuticular"fingerprint." The prevalent pat- The Yucatan species had, in fact,been earlier tern in most species is the pustulate (fig. 1A), described by Grisebach (1859) as with rounded, often closely packed, waxy pro- tremulus,based on a collection from . tuberances. In the widespread A. tremula(Gri- Mueller Argoviensis (1863, 1866, 1873) created seb.) Websterof and West In- a new sect. Ciccopeltandrawithin Phyllanthusto dies, the pattern is distinctlyverruculose (fig. accommodate his Phyllanthusjacobinernsis from 1B); this is particularlyuseful in distinguishing Bahia, ; this Brazilian plant is clearly con- A. tremulafrom the closely related A. peltata.In generic with the Jamaican one. Neither Rob- A. neurocarpa,the most xeromorphic species, inson and Millspaugh (in Millspaugh 1905) nor thereis only a scantywaxy layer,and the abaxial Pax and Hoffmann(1922) appear to have been surface appears smooth (fig. 1C). Although aware of these earlier treatmentsof the genus. methodsof specimen preparation(perhaps tem- Johnston(1923) and Standley (in Ferris 1927) perature during drying) may modifyor some- added two more species from Mexico, giving times obscure these cuticular characters,they the impressionthat Astrocasia is a genus of Mex- nevertheless appear to be valuable for identi- ico and the GreaterAntilles. Study of specimens fyingvegetative material and in separatingspe- now available, however, shows that Astrocasia cies. extends to and includes a total The inflorescencein Astrocasiais always ax- of at least five species. illary, and flowers are produced from highly condensed axes thatmay be described as "glom- erules"; staminateflowers may be produced in MORPHOLOGICALLYIMPORTANT CHARACTERS considerable numbers (often more than 10), In overall vegetative characters species of As- whereas thereare only 1-3 pistillateflowers per trocasiaare mostlysimilar, with alternate,stip- cluster.In general, species of Astrocasiaappear ulate, long-petiolate blunt leaves; only A. aus- to be dioecious; at least, herbarium specimens tinii(Standley) Webster, comb. nov., diverges are nearly always unisexual. A curious devia- notably in its relativelyelongated caudate-acu- tion, however, is found in A. tremula,where minate leaf blades. The leaves of A. peltata both in Jamaicaand in Central America collec- Standley mostly have a peltate leaf-base, al- tors have observed what may be termed"mod- though this characteris variable, and very nar- ular dioecism": staminateand pistillateflowers rowly peltate leaf bases occur in other species can occur on the same plant, but always on dif-

311

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 312 SYSTEMATIC BOTANY [Volume 17

F 1 C ws

E a-I '

lmm

FIG. 2. Pistillate flower of Astrocasiajacobinensis (MattosSilva 2838, DAV).

- 'A k within the tribe Phyllantheae in having five anthersencircling a discoid pistillode atop the staminalcolumn. The Guatemalan A. austiniiis similarexcept that there are only threestamens instead of five. Most divergent is the androe- cium of A. jacobinensis(Muell. Arg.) Webster, comb. nov., in which the four or five anthers are distinctlystalked, and the pistillode is stip- itate.Judging by the prevalentandroecial con- figurationsin the Wielandieae and other Phyl- lantheae, it appears that the androecium of A. jacobinensismay represent the least modified facesin Astrocasia;bar indicates100 ,uM. A. Pustu- condition. late; A. jacobinensis(Webster et al. 25829, DAV). B. The pistillateflower of Astrocasiahas a peri- Verruculose;A. tremula et al. 17538, C. (Webster DAV). anth and disk similarto those of the staminate. Smooth;A. neurocarpa(Webster et al. 13121,DAV). The sepals and petals are early deciduous, as in Savia Willd., but in contrastto FlueggeaWilld. ferentbranches. In other species, such as A. or Phyllanthus,in which the pistillatesepals are neurocarpa,all appear to be strictlyuni- usually persistentunder the fruit.In this re- sexual, although it is impossible to be certain spectAstrocasia appears somewhatintermediate about this until more observationsare made in between the tribes Wielandieae and Phyllan- the field. theae. The gynoeciumin Astrocasiais similarin The flowersare similar in all species of As- most species, with a glabrous 3-locular ovary trocasia(figs. 2, 3). The calyx of both staminate and threemore-or-less free bifid styles; A. trem- and pistillateflowers is characteristic,with small, ula is divergent in having deflexed, more-or- more indurate outer sepals that contrastwith less stigmatiformstyle-branches. the largerand thinnerinner sepals; this recalls The fruitof Astrocasiais an explosively de- the heterogeneityin the sepals of Savia dictyo- hiscent capsule that shattersand leaves a per- carpa Muell. Arg. (Webster 1982) and sistentcolumella. The seeds may be single or schuechiana(Muell. Arg.) Webster (Webster paired in each coccus, and have a smooth to 1984). The androecium of the type species, As- slightlyroughened seed-coat except in A. aus- trocasiatremula, and its related Mexican con- tinii,where the seed coat is deeply and promi- genersA. neurocarpaand A. peltata,is distinctive nentlyrugose.

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 1992] WEBSTER: ASTROCASIA 313

A ; B

mmi1 mm

D ,-. I Y I

E 1~~~~~~~~~~~~~mm E~~~~~~

~~~~~~~~~~~~

1mm1m

FIG.3. Flowers of Astrocasia. A. A. austinii,pistillate (Standley 72981, NY). B. A. austinii,staminate (Standley 72951, F). C. A. tremula,pistillate (Websteret al. 8573, DAV). D. A. tremula,staminate (Websteret al. 8573, DAV). E. A. jacobinensis,staminate (Pedra do Cavalo 810, UEFS). F. A. neurocarpa,staminate (Webster& Arm- bruster20502, DAV).

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 314 SYSTEMATICBOTANY [Volume 17

110 100 90 0 70 00 s0 40

- --

- 5~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-

00 ~ ~ ~ ~ ~ ~ 0 so 80 . os 4

autni N,PoopooodbyHoo~~~~~~~~~~~~~~~~~~~~~~~~~~~~dok0Rypkooo,o A.nuoap;P ~ .pett;T~ ~ ~ ~ 0.teua~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ ~ 2 20~ ~~ ~ ~~~~~~~~~00 00

00 0070 by 000 UoooooootyofUtooot Poblohod by 000 Stoto Uoooooooty0o0o0of Utoooht, 000 Nothoolooodo Opootoooootof 000000000000

FIG. 4. Geographical distributionof Astrocasia. Dots represent A. jacobinensis.Letters represent: A, A.~~~~~~~~~~~~~~~~~~~~~~~~~~~~1

austinji; N, A. neurocarpa;P. A. peltata;T, A. tremula.~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~

ECOLOGY AND GEOGRAPHICAL DISTRIBUTION rock type. Only A. austinii,in the lowland rain- forestsof northernGuatemala, diverges mark- Most of the species of Astrocasiaare found in lowland tropical forestor woodland with pro- edly in its ecological niche. The of nounced seasonality in precipitation.In Mexi- overall distribution Astrocasiais bi- co, A. tremulaand its vicariant A. peltataoccur modal, with a Mexican center extending into in deciduous woodlands ("selva baja caducifo- and the GreaterAntilles, and a dis- lia" of Rzedowski 1978). According to Rze- junct South American area (figs.4, 5). The mor- dowski (1959), A. neurocarpain San Luis Potosi phologically least specialized species, A. jacob- is characteristicof the "matorralsubmontano," inensis,appears to have a disjunct distribution, a scrubbyvegetation of higher elevations (800- with a single station in and a rather 1700 m). In Bahia, Brazil, A. jacobinensisgrows restrictedoccurrence in the lowland seasonal in tropical deciduous woodland reminiscentof forestsof northeasternBrazil. Between Guate- that found in Mexico. All of these species ap- mala and Bolivia, Astrocasiawas not recorded pear to be distinctlycalciphilous, forlimestone until recent collections in Panama and Vene- is invariably noted by collectors as the parent zuela that appear to representdisjunct stations

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 1992] WEBSTER: ASTROCASIA 315

.~~~~~~~~~~~~~~~~~

SCALE~~~~~~~

1000FOOT CONTOUR J\\) / EQA-AEPROJECTIO.

BO_.E_ P; FIG. 5. Geographical distributionof Astrocasiain Mexico and Guatemala. Squares, A. austinii;triangles, A. neurocarpa;open circles,A. peltata;dots, A. tremula. of A. tremula.It is difficultto provide a plausible tween Astrocasiaand genera of the tribe Phyl- scenario for this evidently relict distribution, lantheae. I now concur with thisviewpoint, but but I think it most likely that Astrocasiaorigi- would suggestthat Astrocasia should be referred nated in South America and later spread to Me- to a monotypicsubtribe of Phyllantheae. soamerica, where it differentiatedmainly in Among the genera of tribePhyllantheae, the Mexico. The Panamanian and Venezuelan pop- closest morphologicallyappears to be ulations of A. tremulamight perhaps reflectdis- Baillon, which is similar in aspect and flowers, placements that occurred during Pleistocene and differsmainly in its apetaly and deeply climatic changes. pitted seeds. There is a strikingresemblance in habit between Astrocasiaaustinii and the geo- AFFINITIES graphicallyproximate species Meineckiabartlett- ii (Standley) Webster,and M. capillipes(Blake) Because Astrocasiais endemic to the New Webster; it remains unclear whether this re- World, its affinitiesmight be expected to fall flectsclose phylogenetic affinityor ecological with other neotropical genera of subfam.Phyl- convergence. lanthoideae. In my systemof tribesof that sub- family (Webster 1975), Astrocasiawas referred SYSTEMATICTREATMENT (with some hesitation) to tribeWielandieae be- cause of some floralresemblances with genera Astrocasiinae Webster, subtribus nov.-TYPE: such as HeywoodiaSim, and Savia. However, the AstrocasiaRobinson and Millsp. palynological studies of Punt (1962) and Kohler (1965), and the leaf anatomical investigations Genus ab generibus aliis tribusPhyllantheae of Levin (1986), support a closer affinitybe- differtpetalis praesentia, ovulo anatropo, ru-

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 316 SYSTEMATIC BOTANY [Volume 17

dimento pistillum peltato, stipulis caducis cos- walled; columella slender,persistent; seeds 1 or tulatis. 2 per locule; seed-coat dry, smooth or rough- Dioecious shrubs without phyllanthoid ened, the raphe conspicuous; endosperm co- branching; stipules deciduous, ribbed; flowers pious; embryostraight, the cotyledonsthin, flat, with conspicuous petals; disk annular in both without chlorophyll,much longer and broader sexes; pistillode insertedatop staminal column; than the radicle. anthers extrorse;ovules anatropous. As here interpreted,Astrocasia is a neotropical The subtribeincludes the single genus Astro- genus of five species, extending from Mexico casia. It has much in common with subtribeSe- (Sinaloa, ) and south to Bolivia curineginae Muell. Arg. as circumscribedby and Brazil. Webster(1975), but differsfrom all of the genera of that subtribe in the well-developed petals. KEY TO THE SPECIES OF ASTROCASIA

1. Leaves rounded to obtuse or acute at the tip, ASTROCASIARobinson and Millsp., Beibl. Bot. never long-acuminate; stamens 4 or 5; seeds Jahrb.80:19. 1905.-TYPE: Astrocasiaphyl- smooth to rugose. lanthoidesRob. and Millsp. [=A. tremula 2. Abaxial cuticular pattern of the leaf blade (Griseb.) Webster]. verruculose; pistillate petals 2.5-3.5 mm Phyllanthussect. AmphiandraGriseb., Fl. Br. W. long; styles appressed to the ovary, de- Ind. 34. 1859.-TYPE: Phyllanthustremulus flexed, the branches thickened and stig- Griseb. [=Astrocasiatremula (Griseb.) Web- matiform;seeds smooth,3.4-5 mm long . . ster]...... 3. A. tremula Phyllanthussect. CiccopeltandraMuell. Arg.,Lin- 2. Abaxial cuticularpattern of the leaf blade not naea 32:2. 1863.-TYPE: Phyllanthusjacobi- verruculose; pistillate petals 1.5-2.5 mm nensisMuell. Arg. [=Astrocasiajacobinensis long; styles spreading, the branches slen- der. (Muell. Arg.) Webster]. 3. Abaxial cuticularpattern of the leaf blade Glabrous trees or shrubs; ultimate axes not pustulate; leaves thinly chartaceous, deciduous; foliage deciduous or evergreen; mostlyat least twice as long as broad. leaves alternate, petiolate; blade entire, pin- 4. Leaves not peltate; stamens 4 or 5, the anthers and pistillode stipitate,the nately veined or sometimes 3-5-veined at the anthers deflexed; seeds rugose, 2.8- base, chartaceousto coriaceous, sometimespel- 3.6 mm long...... 1. A. jacobinensis tate or stipellate at the base; stipules ribbed, 4. Leaves distinctly peltate (except for deciduous. Monoecious or dioecious (when smallestleaves on a branch); stamens monoecious the staminateand pistillateflowers always 5, the anthers and pistillode oftenon separate branches); flowersaxillary, in sessile atop the staminal column; glomerules of several to many staminateor 1- seeds smooth, 3.9-4.8 mm long...... 2. A. peltata 3 pistillate flowers. Staminate flower pedicel- late; sepals 5, usually unequal, the outersmaller 3. Abaxial wax deposition on the leaf cuticle and more indurate; petals 5, longer than the scanty,the surfacesmooth; leaves rigid, broadly elliptic to orbicular,mostly not sepals, prominently veined; disk annular or over 1.5 times as long as broad; seeds filaments shallowly patelliform;stamens 3-5; minutelyroughened, 3.7-4.8 mm long. connate into a column; anthers sessile or stip- ...... 4. A. neurocarpa itateatop the column, extrorsein bud, horizon- 1. Leaves long-acuminate at the tip, chartaceous, tal or deflexed;pollen grains subglobose, 3-col- abaxial wax patternpustulate; stamens 3; seeds porate,the exine reticulate,the colpi elongated deeply rugose, 3.5-4.3 mm long .. 5. A. austinii with well-defined germ pores, pistillode disci- form,sessile or stipitateatop the staminal col- 1. Astrocasia jacobinensis (Muell. Arg.) Web- umn. Pistillate flower long-pedicellate; sepals ster, comb. nov.-Phyllanthus jacobinensis and petals similarto the staminate,soon decid- Muell. Arg.,Linnaea 32:6. 1863; DC. Prodr. uous; disk annular to cupular; ovary 3-locular 15(2):334. 1866; Fl. Brasil. 11(2):29. 1873.- (veryrarely 4-locular); stylesfree, bifid, the tips TYPE:Brazil, Bahia, Jacobina,Blanchet 3291 spreading or deflexed;ovules 2 per locule, anat- (holotype: G!). Figures 1A; 2; 3E; 6B. ropous, obturatormassive. Fruitcapsular, thin- Phyllanthusinaequalis Rusby, Mem. Torrey Bot.

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 1992] WEBSTER: ASTROCASIA 317

E

A A b i

FIG. 6. Habit of Astrocasia species. A. A. aulstinii(Standley 39300, F). B. A. jacobinensis (Noblick 3266, UEFS).

Club 6:118. 1896.-TYPE: Bolivia, Dept. La 5-10 mm long; sepals 5 (rarely4), unequal, 0.8- Paz, between Tipuani and Guanai, Bang1704 1.2 mm long, 0.5-1 mm broad; petals obovate- (holotype: NY!; isotypes: GH!, US!). oblong, 2.2-2.5 mm long, 1-1.3 mm broad; disk entire, undulate, nearly flat,1-1.5 mm across; Shrub or small tree 2-5 m high; flowering stamens5 (rarely4), androeciumca. 1 mm across; branches 1-2 mm thick; foliage deciduous. staminalcolumn 0.6-0.8 mm high; anthersmore Leaves with stipules2-4 mm long; petiole (0.5)1- or less stipitateand deflexed,0.3-0.5 mm broad; 2.5 cm long; blade chartaceous, elliptic to ob- pistillode 0.4-0.6 mm across, more-or-lessstip- ovate, obtuse or rounded at the tip, cuneate to itate (0.1-0.3 mm) above the top of the staminal rounded or subcordate at the base, (2)4-11 cm column. Pistillate flower: pedicel becoming long, (1.5)2.5-5.5 cm broad; lateral veins 7-10 (20)30-45 mm long, slender (0.15-0.25 mm thick per side, ascending, tertiaryveinlet networkir- medianly,0.5-0.7 mm thickdistally); sepals lan- regular (not markedly scalariform), ultimate ceolate, 1-1.5 mm long; petals broadly elliptic, veinlets formingangular areoles; abaxial cutic- 2.5-2.8 mm long, 1.9-2.2 mm broad; stylesslen- ular pattern pustulate; margins slightly thick- der, bifid,0.6-0.7 mm long. Fruitsnot seen in- ened, plane; stipels narrowlylanceolate, up to tact;valves distinctlyveiny, ca. 6 mm long; seeds 0.5 mm long, often reduced or absent. Flowers ellipsoidal, rugose, 2.8-3.6 mm long, 2-3 mm solitaryor in clusters.Staminate flower: pedicel broad.

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 318 SYSTEMATIC BOTANY [Volume 17

Distribution.Deciduous forest,disjunct be- sion of the filamentsand pistillode, so thatboth tween Bolivia and Bahia, Brazil. The species is anthers and pistillode appear stipitate;the an- common in the "mata seca" (deciduous or thorn therstend to be deflexed,quite unlike those of forest),where it is found at low elevations (be- otherspecies, which are strictlyhorizontal. The low 100 m) on limestone. seeds, mostly less than 3.5 mm long, are dis- tinctlysmaller than those of the other species. Representativespecimens examined. BRAZIL. Bahia: As the key indicates,the closest species to A. Mun.Almadina, rodovia para Ibitupa, Raimundo SP1140 jacobinensisin overall morphological featuresis (CEPEC);Mun. Boa Vistado Tupim,Itaberaba to Len- the Mexican A. peltata.However, that species qois, Mattos Silva et al. 1580 (DAV); Mun. Cachoeira, Estacaode Mata,Barragem Bananeiras, Pedra do Cavalo differsin a number of features,including its 319,810 (BAH, UEFS), 895 (UEFS); MorroBelo, Pedra usually peltate leaf base, androecium with sub- do Cavalo et al. 958 (BAH, UEFS); Mun. Conceiqao de sessile anthers,and larger seeds. The gap is suf- Feira,NE da BarragemBananeiras, Carvalho et al. 527 ficientlylarge thatA. peltatacannot be unqual- (CEPEC); Mun. Feirade Santana,Serra de Sao Jose, ifiedlyregarded as the sister species. Fazenda Boa Vista, Noblick3244, 3268, 3287 (DAV, Noblick& Hahn 3375 Roca da UEFS), (DAV); Fazenda 2. ASTROCASIAPELTATA Standley, Contr. Dudley Serra,Noblick 3213 (DAV); FazendaMonte Verde, 11 Herb. 1:74, pl. 1, fig. 4. 1927.-TYPE: Mex- km NW of Jaguara,Queiroz et al. 1147(UEFS); lime- stonehill 34 km W of Feira de Santana,Webster & ico, , Islas Tres Marias, Isla Maria Queiroz25829 (DAV, UEFS); Mun. Ipira,Fazenda Re- Madre, Ferris5571 (holotype: DS!). Figure creio,Estrada Feijao km 43, Queirozet al. 994 (UEFS); 4; 5; 7B. Mun. Itambe,rod. BA-265,km 8 do trechoBR-415, Shrub or small tree 2-5 m high; foliage de- Mori et al. 9382 (CEPEC). ciduous. Leaves with stipules (2.5)4-7.5 mm This species has been confused in the botan- long; petiole slender,1.5-4.5 mm long, peltately ical literaturebecause Phyllanthusinaequalis and inserted on the blade (except some on smaller P. jacobinensiswere originallyknown only from leaves); blade firmlychartaceous to subcoria- staminate specimens. Mueller Argoviensis ceous, suborbicular to broadly obcuneate, ob- (1863) made P. jacobinensisthe type of his sect. tuse to rounded at the tip, rounded below the Ciccopeltandra,the choice of name indicatinghis peltate insertionof the petiole, (1)2-10 cm long, intuitionthat the plant was related to his sect. (1)2.5-8 cm broad (often broader than long); Peltandra(=Meineckia). Rusby, in describing P. major veins on each side above, veinlets prom- inaequalis,compared it with P. nutansSw. of Ja- inentlyreticulate beneath; abaxial cuticularpat- maica and the Amazonian P. martiiMuell. Arg., tern pustulate (or sometimes nearly smooth); neither of which are particularlysimilar. Both marginsthickened but scarcelyrevolute; stipels Mueller and Rusby, perhaps because of inade- absent. Flowers mostlysolitary. Staminate flow- quate material,interpreted the flowersas being er: pedicel 6-7 mm long; sepals oblong, thin, apetalous. ca. 1.5-1.6 mm long and 1.8 mm broad; disk ca. Recent collections fromBahia, and in partic- 0.3 mm high, 1.2 mm broad; stamens 5, an- ular the field assistance of Sr. Luciano Queiroz, droecium ca. 1.2 mm across; staminal column now permit for the firsttime an accurate de- ca. 0.4 mm high, pistillode more or less sessile, scriptionof the floralmorphology; this clearly 0.6 mm broad. Pistillate flower:pedicel becom- indicates that the plant belongs to Astrocasia. ing (25)30-60 mm long, 0.4-0.5 mm medianly, Despite the great geographical disjunction,the 1-1.5 mm thick distally; sepals and petals not single collection fromBolivia does not appear seen. Fruit oblate, 10-10.5 mm across; valves to differfrom the Bahia specimens in any sig- 8.3-8.7 mm long; columella 4.5-4.8 mm long; nificantway. Both the Bolivian and Brazilian seeds plano-convex or ellipsoidal, smooth,3.9- specimens show flowerswith fourand five sta- 4.8 mm long, 3.4-3.9 mm broad. mens on the same branch, an unusual feature Distribution.Deciduous or semi-deciduous not observed in any other species (Rusby re- forestor woodland, coastal limestone hills of ported "6 or more" stamens,but this could not ,Islas Tres Marias, and with an isolated be confirmed). population disjunct in Sinaloa near Mazatl'an. The staminate flowers of A. jacobinensisare distinctivenot only in the variation in stamen Representativespecimens examined. MEXICO. Jalisco: number,but also in the less-than-completefu- Mun.La Huerta,deciduous woodlands, Estaci6n Bio-

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 1992] WEBSTER: ASTROCASIA 319

A /

B T

FIG. 7. Habit of Astrocasiaspecies. A. A. tremula(Webster et al. 8573, DAV). B. A. peltata(Magellanes 279, MICH). C. A. neurocarpa(Webster and Armbruster20590, DAV).

l6gica Chamela, Lott 1574 (CAS), Lott& Bullock1442 Jamaica,Purdie s.n. (lectotype: K!). Figure (MO, TEX), Magellanes279 (MICH), 3996 (TEX), Perez 1B; 3C,D; 4; 5; 7A. J. 1786 (CAS), 1804 (MO). Nayarit: Islas Tres Marias, Astrocasiaphyllanthoides Robinson and Mills- Isla Maria Madre, Chavez & Villamar(WISC), Nelson paugh, Bot. Jahrb.36, Beibl. 80:19. 1905.- 4215 (US); Isla Maria Magdalena, 1962,Grant s.n. (DS). TYPE: Mexico, Yucatan, Mrida, Seler 3943 Sinaloa: Isla Creston, near Mazatlan, Howell 10564 (holotype: F!; isotype,GH!). (US). Shrub or tree 2-10 m high; branches terete 3. ASTROCASIA TREMULA (Griseb.) Webster, J.Ar- or obscurely angled, smooth, pale; foliage de- nold Arb. 39:208. 1958.-Phyllanthus tremu- ciduous. Leaves with stipules 4-6 mm long; pet- lus Griseb., FL. Br. W. Ind. 34. 1859. -TYPE: ioles 2-6 cm long; blade chartaceous,ovate or

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 320 SYSTEMATIC BOTANY [Volume 17

elliptic, acute or obtuse at the tip, broadly cu- US), 28018(MO). St. Elizabeth:Great Pedro Bay, neate to rounded and sometimesvery narrowly Britton1213 (NY); Round Hill, Harris9707 (NY); Mal- peltateat the base, 5-13 cm long, 2.5-9 cm broad; vernto Mountainside,Harris 9903 (NY); 1 mi SSW of major veins 5-9 on a side, ascending brochi- GiddyHall, Proctor 11700 (NY). St. James:Montego Bay,Britton & Hollick2354 (NY); Rose Hall, Miller1412 dodromous; veinlets prominent beneath; ab- (US); Ironshore estate, Proctor 23224 (DAV, pattern verruculose; margins axial cuticular MICH). Trelawny:2 mi ESE of Falmouth,coastal plane or narrowlyrevolute; stipels scarious, 0.3- woodland,Proctor 11765 (US). Westmoreland:Little 0.5 mm long, freeor fused,sometimes obsolete. Bay, Adams8538 (MO); Negril, Britton& Hollick2051 Flowers in glomerules, expanding with the (NY); Negril Hill, Websteret al. 8512 (DAV). leaves; staminateflowers many per glomerule, MEXIcO. Campeche:Campo ExperimentalFores- pistillateflowers 1-3. Staminateflower: pedicel tal"El Tormento,"Chavela et al. ES238 (MEXU, MICH), 8-20 mm long; sepals broadly ellipticto oblong- ES-3087(MEXU); Tuxpefia, Lundell 827 (A, GH, MICH, lanceolate, 1.2-1.5 mm long, 1.2-1.8 mm broad; MO, NY, US); entreCeibaplaya y Champot6n,Mi- petals elliptic to oblong-lanceolate,2.5-3.5 mm randa 8005 (MEXU); Xpujil, Shepherd204 (WISC). Jalisco:Mun. El Tuito,35 km E of Cabo Corrientes, long, 0.8-1.1 mm broad; disk cupuliform,fluted, Anderson& Anderson6064 (MICH). Nayarit:Mun. 0.4-0.5 mm high, 0.9-1.8 mm broad; stamens 5, Compostela,20 miW ofCompostela, Graber 22 (TEX); the androecium 0.8-0.9 mm across; anthershor- QuintanaRoo, 6 km N of Pto. Morelos,Cabrera & izontal, 0.4 mm across; pistillode sessile, 0.5-0.6 Cabrera3587 (CAS); SE side ofL. Chicnancanab,Dar- mm across. Pistillate flower:pedicel becoming win 2473 (MO); Coba, E of ruins,Lundell & Lundell (25)30-55 mm long, 0.4-0.6 mm broad medi- 7827(MICH, TEX); Chichankanab, Gaumer 1261 (CAS, anly, 1.5-2.2 mm broad distally,often lenticel- F, MO, NY, US), 1794(MO); caminoa Xcaret,Morena late; sepals orbicularto elliptic,1.7-2.5 mm long, 279 (NY); PuertoMorelos, Tellez & Cabrera1314 (DAV); 1.5-2 mm broad; petals obovate-oblong,3.5-4.2 Cancuin,Tellez & Cabrera1552 (DAV). Yucatan: Chi- mm long, 1.5-1.7 mm broad; disk cupular, the chen Itzi, Bequaert7 (US), Fogg22447 (US), Steere1318 (MICH, MO), Webster& Lynch18583 (DAV, MEXU); marginundulate, 0.5-1 mm high and 2-2.5 mm Izamal, Gaumer475 (CAS, DS, GH, NY), Greenman392 thickened, the broad; styles 0.5-0.6 mm long, (GH); Dzilbilchaltun,Bradburn & Darwin 1139 (DAV); recurved tips clavate. Fruits oblate, 3-angled, Mayapan,Darwin & Sundell2133 (MO); Ticul,Darwin reticulate-venose,ca. 10 mm in diameter; col- 2436 (NY); Silam, Gaumeret al. 23346 (A, F, GH); Apa- umella 3.2-3.5 mm high; seeds plano-convex, zoli, Goldman491 (US); Piste, Lundell& Lundell7345 smooth, yellowish, 3.4-5 mm long, 3-4.5 mm (A, LL, MICH); Mrida, Schott319 (US). broad. PANAMA. Colon: forestalong shoresof Madden lake,near Madden Dam, Knapp1299 (DAV, F, MO), Distribution.Deciduous forest, usually on Witherspoon8805 (DAV, MO); 6 km N of Chilibre, limestone, sea level to 500 m, Mexico, , along MaddenLake, Knapp 2715 (MO). northernGuatemala, Jamaica, eastern Cuba, and Astrocasia tremula has a characteristically to Venezue- ; disjunct Panama, northernCaribbean distributionin the Yucatan la. peninsula and GreaterAntilles, except for the anomalous occurrencesin western Mexico, Pa- Representativespecimens examined. BELIZE. El Cayo: nama, and Venezuela. The species is well sep- El Cayo, Gentle2179 (A, CAS, MICH, TEX), Lundell 6150 (MICH, NY, TEX, US). arated from its congeners by its larger leaves CAYMAN ISLANDS. Grand Cayman: Grape Tree with verruculose abaxial cuticularwax pattern, Point, Proctor11975, 15140 (GH). elongated petals, large cupulate pistillate disk, CUBA. Oriente: El Dudoso cliffs,Pil6n, Alain 1094 and peculiarly thickened style branches. It is (US); Santiago, Playa Siboney, Ekman8715 (NY). nevertheless similar to A. peltata in habit, and GUATEMALA.Peten: Uaxactun, Bartlett12528 (A, these two taxa could perhaps be interpretedas MICH); MilpaGrande, Macanche, Contreras 5488 (MO, allopatric subspecies of a single species. The NY); Santa Elena, Ortiz1786 (MO, NY). Retalhuleu: occurrence of A. tremula in mainland Nayarit 87526, between Nueva Linda and Champerico,Standley and in westernJalisco is surprising,since these 87643 (F). populations are in the same region as A. peltata JAMAICA. Clarendon: Cockpit,Read 1999c(MICH, and fromthe nearest eastern US); Round Hill, Websteret al. 8251 (DAV). Hanover: markedlydisjunct Lance's Point, Websteret al. 8573 (DAV, MICH, Mexican populations in Campeche. The Pana- US). St. Ann: Salem, Britton2547 (NY). St. Cath- manian plants, although known only from erine: 1 mi W of Riverhead,Proctor 19837 (DAV, NY, fruitingspecimens, appear to representA. trem-

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 1992] WEBSTER: ASTROCASIA 321 ula, despite obscurityin the cuticular wax pat- side, ascending, weakly brochidodromous,the tern. Specimens probably of A. tremulafrom veinlets forming a prominent reticulum be- Venezuela (Lara) were also observed in the her- neath; abaxial cuticular pattern smooth; mar- barium at Barquisimeto, but have not been gins narrowly revolute; stipels free or fused, available for this study. Because A. tremulais dark and scarious,0.2-0.5 mm long, or obsolete. evidently an obligate calciphile, some disjunc- Flowers mostlysolitary or the staminatepaired. tion in its distributioncan be expected, since it Staminateflower: pedicel 6-12 mm long; sepals will be found on limestone soils but not en- elliptic, entire,the outer often smaller and ba- countered on volcanic soils. sally indurate,1-1.7 mm long, 0.9-1.5 mmbroad; Fawcett and Rendle (1919, 1920) applied the petals elliptic,2-2.5 mm long, 1.2-1.5 mmbroad; name Phyllanthusglabellus (L.) Fawcettand Ren- stamens 5, the androecium 1-1.2 mm across; dle to the species here called Astrocasiatremula. staminal column 0.8-1 mm high; anthers ses- This, however, was in my opinion based on sile, dehiscing horizontally,0.5-0.6 mm broad; faulty interpretationof typificationand no- pistillode sessile, 0.5-0.8 mm broad. Pistillate menclature.To begin with,they used the basio- flower: pedicel becoming (10)15-25(35) mm nym Crotonglabellus L. twice for two different long, 0.4-0.5 mm thick medianly, 0.8-1.5 mm plants-surely a heterodox proceeding under broad; sepals deltoid (outer) to elliptic (inner), any scheme of nomenclature.They correctlyin- 1.0-1.8 mm long and broad; petals narrowly terpretedCroton glabellus L. (Syst. Nat., 10th ed., obovate-oblong, parallel-veined, 3.3-3.7 mm p. 1275, Oct. 1759), as a synonymof Crotonlu- long, 1.3-1.5 mm broad; disk shallowly cupular, cidusL., based on the presumed type specimen 2-2.2 mm across; styles bipartite,0.6-1.0 mm in Herb. Sloane (BM; H.S. 5, fol. 109, fideDandy long, the branches slender. Capsule oblate, 1958). However, they then again used Croton 3-lobed, reticulate-venose,9-10 mm across; col- glabellusL. (Fl. Jam.Pugill, p. 409, Nov. 1759), umella ca. 4 mm high; seeds planoconvex or based on a Jamaican collection by Patrick ellipsoid, plump, minutely verruculose, 3.7- Browne,as the basionym fortheir combination 4.4(4.8) mm long, 3-3.7 mm broad. Phyllanthusglabellus. This is clearlyillegal under currentrules of nomenclature. Distribution.Sclerophyllous scrub (matorral) on limestone hillsides, 1100-2100 m, north- 4. ASTROCASIANEUROCARPA (Muell. Arg.) I. M. eastern Mexico. Johnstonex Standley,Contr. Dudley Herb. Representativespecimens examined. MEXICO. Puebla: 1:74. 1927.-Phyllanthusneurocarpus Muell. Mun. Acatlan, 3.5 mi NW of Petlalcingo, Webster& Arg., Linnaea 34: 69. 1865.-TYPE: Mexico, Armbruster20746 (DAV, MEXU, NY, TEX). San Luis between Victoria and Tula, Berlandier2216 Potosi: 4-11 mi W of Ciudad del Maiz, Henrickson& (holotype: G!; isotype:MO!). Figure 1C; 3F; Lee17586a (TEX), Johnston & Graham 4475, 4476 (MICH), 4; 5; 7C. Webster& Armbruster20590 (DAV, MEXU),Webster et Jatrophacercidiphylla Standley, Contr. U.S. Natl. al. 13121 (DAV, MEXU). Tamaulipas: Jaumave Val- Herb. 23:639. 1923.-TYPE: Mexico, be- ley, Nelson 4455 (F); 33 km E of Jaumave,Henrickson tween San Luis Potosi and Tampico, Palmer & Hess 19095, 19100(TEX); 27 km WNW of Jaumave, 1140 (holotype: US 42743!) Henrickson& Hess 19255(NY, TEX); 4 mi S of Jaumave, Stanfordet al. 2264 (CAS, GH, MO Astrocasia(?) populifoliaI. M. Johnston,Contr. NY); Bustamante, Harriman11629 (DAV); 6.3 mi N of Hwy. 101 on road Gray Herb. 68:84. 1923.-TYPE: Mexico, Ta- to Bustamante,Johnson et al. 11183(DAV, LL); 6 mi maulipas, JaumaveValley, Nelson4455 (ho- SW of Palmillas, Webster& Armbruster20502 (DAV, lotype: GH!; isotype: F!). MEXU); 11 mi S of Palmillas, Crutchfield& Johnston 5613B N Shrub 1-2 m high; branches terete or ob- (MICH,TEX); ofLa Presita,23 mi N ofTula, Dorr & Atkins2345 (CAS, GH, TEX); Mun. Tula, 1 km scurely angled, tan to grey; foliage deciduous. E of La Presita, GarciaMendoza 2235 (DAV, MEXU). Leaves with stipules blackish, 2-4(5) mm long; Additional localities in San Luis Potosi and Tamau- petioles (0.3)0.8-2 (2.5) cm long; blade firmly lipas are cited by Rzedowski (1959). chartaceousto subcoriaceous,elliptic to broadly ovate or suborbicular, obtuse to rounded or Astrocasianeurocarpa is the most xeromorphic emarginate at the tip, rounded or truncate to species in the genus, and also attainsthe highest subcordateat the base, (1)1.5-5 cm long, (0.8)1.5- elevations. Withlong-petioled, roundish leaves 3.5 cm broad; major veins mostly 4 or 5 on a the plants have the aspect of aspens (PopulusL.),

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 322 SYSTEMATIC BOTANY [Volume 17

as noted by a number of collectors.The plants This species is very distinctivebecause of its in the disjunct populations in Puebla occur in stronglyacuminate leaves and staminate flow- a riparianhabitat and have largerthinner leaves, ers with only three stamens.The seeds also ap- but neverthelessappear to show moresimilarity pear to be more deeply rugose than in other to A. neurocarpathan to otherspecies; until flow- species. Ecologically,A. austiniiis very different eringspecimens are collected,they appear most fromits congeners in inhabitingevergreen rain reasonably assigned here. forestinstead of monsoonal vegetation. There is a particularlystriking similarity in 5. Astrocasia austinii (Standley)Webster, comb. habit between A. austiniiand Meineckiabartlettii, nov.-Phyllanthus austiniiStandley, Publ. which occurs in driervegetation in Chiapas and Field Mus. Nat. Hist. Bot. 22:38. 1940. Belize. However, in M. bartlettiithe flowersare TYPE: Guatemala, Izabal, Escoba, Standley apetalous and the fruitingpedicel is articulate; 72968 (holotype: F; isotype: US!). Figure furthermore,the staminateandroecium has five 3A, B; 4; 5; 6A. stamenswith stipitateanthers. Despite a similar gestalt,therefore, the two species Shrub 1-5 m high; floweringbranches slen- do not appear to be der, 1-2 mm thick,terete, pale; foliage evidently immediate relatives or even congeners. Nevertheless,this evergreen. Leaves with stipules 2-5(12) mm similarityunderscores the fact that Meineckiaseems to be closer to long; petiole slender (0.5)0.9-1.7 cm long; blade Astrocasia in general morphological characters chartaceous, ovate- or elliptic-lanceolate, dis- than any other genus in tribe Phyllantheae. tinctlyacuminate at the tip, obtuse to rounded and sometimesnarrowly peltate at the base, 5- ACKNOWLEDGMENTS. Herbariumspecimens were 12 cm long, 2.3-5.5 cm broad, pinnatelyveined examined through the courtesy of the curators of a with ca. 5-7 straightishlaterals on each side, numberof herbaria:A, BAH, CAS, CEPEC, DS, F, GH, tertiaryveinlets more or less scalariform,ulti- LL, MEXU, MICH, MO, NY, TEX, UEFS, and WISC. mate veinlets formingsquarish areoles; abaxial Vital assistance in field work in Bahia, Brazil, was cuticular patternpustulate; margins plane; sti- provided by Sr. Luciano de Queiroz, of the Univer- pels absent. Flowers apparently solitary. Sta- sidade Estadual de Feira de Santana. SEM photo- minate flower: pedicel 4.5-5 mm long; sepals graphs of leaf surfaceswere supplied by Dr. Jerome elliptic, somewhat unequal, 1.2-1.4 mm long, Ward. The drawings were made by Ms. Leslie Randall (except for flowersof A. tremulaby Mr. Norm Dees- 1-1.2 mm broad; petals obovate, 1.5-1.7 mm ing). Field work in Brazil and Mexico was supported long, 0.8-1 mm broad; disk entire, nearly flat, by grantsfrom the National Science Foundation. 1.3-1.8 mm across; stamens 3, the androecium 0.5-0.7 mm across; staminalcolumn 0.7-0.8 mm high; anthers horizontal,sessile; pistillode ses- LITERATURE CITED sile, 0.3-0.4 mm across. Pistillateflower: pedicel DANDY,J. E., ed. 1958. The Sloane Herbarium,and becoming 25-40 mm long, 0.3 mm medianly, annotated list of the Horti Sicci composing it. 0.8 mm thick distally; sepals, petals, and gy- London: BritishMuseum (Natural History). noecium not seen. Capsule not seen entire; FAWCETT,W. and A. B. RENDLE. 1919. Notes on Ja- valves ca. 6-7.5 mm long; columella ca. 3.5 mm maica plants. J.Bot 57:65-68. long; seeds roughened and deeply rugose, yel- and . 1920. Flora of Jamaica,vol. 4. lowish, 3.5-4.3 mm long, 2.7-3.2 mm broad. London: BritishMuseum (Natural History). FERRIS,R. S. 1927. Preliminaryreport on the flora Distribution.Restricted to lowland rain for- of the Tres Marias Islands. Contr. Dudley Herb. ests (below 150 m), northern Guatemala adja- 1:57-88. cent to Bahia de Amatique. GRISEBACH,A. H. R. 1859. Flora of the BritishWest Indian Islands. Goettingen. Representativespecimens examined. GUATEMALA. Iza- JOHNSTON,I. M. 1923. Diagnoses and notes relating bal:Guatemala-Peten road, Cienaga, bordering R. Zula, to the spermatophyteschiefly of North America. Contreras10781 (LL); Escoba,near sea level,Standley Contr. Gray Herb. 68:80-104. 72951 (F); Rio Escobas,Montafia Escobas, across bay KOHLER,E. 1965. Die Pollenmorphologie der bio- fromPuerto Barrios, 20-50 m, Steyermark39886 (A, vulaten Euphorbiaceae und ihre Bedeutung fur US); Rio Bonita,top of lowestpeak, 30-150 m Stey- die Taxonomie. Grana Palynol. 6:26-120. ermark41715 (F). LEVIN,G. A. 1986. Systematicfoliar morphology of

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions 1992] WEBSTER: ASTROCASIA 323

Phyllanthoideae (Euphorbiaceae). I. Conspectus. ROGERS,D. J.and S. G. APPAN. 1973. Manihot,Ma- Ann. Missouri Bot. Gard. 73:29-85. nihotoides(Euphorbiaceae). Flora Neotropica 13: MENNEGA, A. M. W. 1987. Wood anatomy of the 1-272. Euphorbiaceae, in particular of the subfamily RZEDOWSKI,J. 1959. Notas sobre la flora y la vege- Phyllanthoideae. J.Linn. Soc., Bot. 94:111-126. taci6n del Estado de San Luis Potosi. VIII. Cinco MILLSPAUGH, C. F. 1905. Plantae a clariss.Ed. et Caec. faner6gamasinteresantes de sur y centro del Es- Seler in Yucatan collectae. Bot. Jahrb.36(Beibl. tado. Ciencia [Mexico] 19:77-86. 80):11-30. 1905. . 1978. Vegetaci6n de Mexico. Mexico City: MUELLER ARGOVIENSIS, J. 1863. Euphorbiaceae. Vor- Editorial Limusa. laiifige Mitteilungen aus dem furDe Candolle's WEBSTER, G. L. 1965. A revision of the genus Mei- Prodromus bestimmten Manuscript iiber diese neckia(Euphorbiaceae). Acta Bot. Neerl. 14:323- Familie. Linnaea 32:1-126. 365. . 1866. Euphorbiaceae [except Euphorbieae]. . 1975. Conspectus of a new classificationof In Prodomussystematis naturalis regni vegetablis, 15 the Euphorbiaceae. Taxon 24:593-601. (2):1-126, ed. A. Candolle. . 1982. Systematicstatus of the genus Klei- 1873. Euphorbiaceae [firstpart]. In FloraBra- nodendron(Euphorbiaceae). Taxon 31:535-539. siliensis,11(2):1-292, ed. Martius. . 1984. A revision of Flueggea (Euphorbi- PAX, F. and K. HOFFMANN. 1922. Euphorbiaceae- aceae). Allertonia 3(4):259-312. Phyllantheae. In Das Pflanzenreich,IV, 147, and M. J. HUFT. 1988. Revised synopsis of XV(Heft 81):1-349, ed. A. Engler. Panamanian Euphorbiaceae. Ann. Missouri Bot. PUNT, W. 1962. Pollen morphology of the Euphor- Gard. 75:1087-1144. biaceae with special referenceto .Wen- tia 7:1-116.

This content downloaded from 169.237.8.7 on Tue, 8 Oct 2013 12:12:54 PM All use subject to JSTOR Terms and Conditions