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- Annals v,is(i- of the Missouri Botanical Garden 1988 #

Volume 75 Number 1 Volume 75, Number ' Spring 1988

The Annals, published quarterly, contains papers, primarily in systematic , con- tributed from the Missouri Botanical Garden, St. Louis. Papers originating outside the Garden will also be accepted. Authors should write the Editor for information concerning arrangements for publishing in the ANNALS. Instructions to Authors are printed on the inside back cover of the last issue of each volume.

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Editorial is. \I,,S ouri Botanu •al Garde,,

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Henl : van der W< ?rff V//.S.S ouri Botanic

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i Botanical Garden 1988 REVISED SYNOPSIS Grady L. Webster2 and Michael J. Huft" OF PANAMANIAN EUPHORBIACEAE1

ABSTRACT induded in \ • >,H The new taxa ai I. i i " I ! I _- i II • hster, correi //,-," |1 U !. I :,.(, , uw .<• r K:nil. ,.J>s;. I i. .lu< \ v|, h, f I/,/..'/ l/f . It ,„„',, ,W/.,/' hi".-lh.il r .,.,„,-[, //u/f, Tetrorchidium microphyllum //u/j, pachypodus Webster, Croton speciosus Muell. Arg. subsp. taca runcnsis II ebster, Croton draco Cham. & Schldl. subsp. panamensis (Klotzsch) Webster, Croton billbergian Muell. Arg. subsp p\rdiiud.ilis (J l> Smith ii i ///, panamensis Webstt Gymnanthesdrrs.sl.ri il /•>». ,/'.-./ Cwin nil i • " ' (f .••.,',, ' . neu or updated ke arc provided where appropriate, as well as d, • . imen citations.

When the treatment of the for In order to in the of Panama was originally published changes, the g< (Webster & Burch, 1968), it was anticipated that new keys to sp it would prove to be incomplete, but collections genera. Citatior made during the past 20 years show that it was species already included in the original treatment even more provisional than we had thought. Not unless they represent new records for provinces or only has Pausandra been found as predicted, but considerable range extensions. Descriptions are no fewer than nine other genera new to Panama: provided for most species new to Panama, but in hlcnoplnictlra. Astrocasia, Croizatia, Drypetes, a few cases, if the Panamanian material is not Gymnanthes, Maprounea, , and Sene- adequate for description or if the species has been feldera, and an unpublished from Cerro Ta- recently described elsewhere, a literature reference genera to 45. In addition, species new to Panama The preparation of this paper has involved us have been found in a number of genera, including in the study of much extra-Panamanian material , , , Croton, Dale- and has led to the resolution of a number of an- champta. , Hyeronima, Mabea, Man- ciliary taxonomic and distributional problems. We ihot, Sapium, Sebastiania, Tetrorchidium, and have not hesitated to discuss these additional items Tragia. The arrangement of genera within the where appropriate. The central position of Panama family has become obsolete since the publication in the Neotropics certainly renders this account of of a new classification (Webster, 1975); there are Panamanian Euphorbiaceae of value to an under- now five subfamilies recognized, of which four oc- standing of the family throughout cur in Panama. This new treatment, with the order and northern . It seems appropriate, of the genera now following the revised classifi- therefore, to include peripheral items that, while cation, includes the taxa new to Panama as well not concerning Panamanian species directly, are as references to recent publications on these taxa. definitely of relevance.

1 We thank the i m the loan of specimens. Part of the work of the senior author uus suppotttd b\ /.-',,< ,, "-> , ,, > ' .stoii, I niiejstt\ of Calif,i nia, Hans, consul, ruble assistance was provided by the staff of the Missouri Botanical Garden, particularly lh Cordon McPherson, who supplied material of some critical taxa. Drawings were provided by Wan-Ling Peng, Clara Richardson, and Steve Wilson. Ilppailim ni ot ii ' • s I I". ''<««'( key to genera and new treatments of genera 1, 3, 4, 5, 6, 8, 9, 23, 27, 32, 34, 36, and 37. The treatment of genus 28, Cnidoscolus, is furnished by Dr. Gary Breckon, University of Puerto Rico, Mayagiiez. Missouri IU,i,i'i>< ,!•:,•, I >, 1,1 \L> . > ;n, .»/ \,i!,il„'' II: In',. Chicago, Illinois (»t)/,or,, I S / . primarily responsible for new treatments of genera 2, 12, 13, 14, 17, 19, 20, 26, 30, 33, 39, and 42; the treatments of genera 16, 22, and 35 are the joint responsibility of both authors.

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j8j ENUMERATION OF TAXA 1 Jill Subfamily I. PHYLLANTHOIDEAE Asch. 1 ?«2|2 1. Astasia i 1^33 Astrocasia Robinson *S. Millsp., Rot. Jahrb. Syst. 36, Reibl. 80: 19. 1905. TYPE: Astrocasia lil phyllanthoides Robinson & Millsp. = Astro- j casia tremula (Griseb.) Webster. 11 I t| Dioecious, glabrous or . al- J"S ternate, petiolale: iij ili i MM deciduous; blades $ J "i entire, pinnately veined. axillary; nil 4 in cluster > • . ,, >cd II i V * 5, sometimes unequal; 5, longer than ItI Z& I the sepals; disk annular; 3 or 5, the fil- ;lj| •ffs aments connate into a , anthers extrorse in 1 2 ^ , dehiscing horizontally; grains tricol- porate, reticu lo ilated at tip into a *il peltate disk capping the staminal column. Pistillate III! flowers long-pedicellate; sepals 5, articulated, de- • s *^ 11 « . i petals 5, longer than sepals; disk cupuli- 111! U pels; 2 per locule, anatropous; styles free, Ill's hil'ul. funis eapsiilai : columella slender, persistent: I^SI !l 1 or 2 per locule, ecarunculate; coat ^I'i- 13 dry, thin, smooth, raphe conspicuous; e lospene copious; enibi vo straight: cot vledoiis thin. ilal. n icl longer and broader than the radicle.

This neotropical genus of four species was not reported from Panama in the original treatment. Astrocasia is one of the more primitive genera of Knphorltiaeeae ami Ii i ^ its do r-l :. latives in Yfnea :u,,,| int., sHiHl and Madaga-- >im, 11 ielandia Rail- Ion).

1.1. Astrocasia tremula (Griseb.) Webster, J. Arnold Arbor. 39: 208. 1958. Phyllanthus tremulus Griseb., Fl. Rrit. W.I. 34. 1859. TYPE: : Purdie, Wullschlaegel (syn- types, K).

Astrocasia fj/nlIni . Millsp., Bot. Jahrb. Syst. 36, Beibl. 80: 19. 1095. TYPE: Mexico. Yu- catan: Merida, Seler 3943 (holotype, F).

Shrub or 2-10 m high; branches terete or obscurely angled, pale; foliage deciduous. Leaves not chlorenchyma-sheathed; main axis no villi slender petiol 2 <> cm long; stipules lanceo- nen, in 5 lateral cymes; styles usually bifid; . Leaves opposite, stipulate, inequilateral at ices pseudanthial, i.e., the 5 involucral brae thia more or less radially synnnc • r: late, chartaceous, 4-6 mm long; blades charta- ceous, ovate, acute or obtuse at tip, broadly cu- |fc3 u neate at base, 5-12 cm long, 3-7.5 cm broad; Missouri Botanical Garden

ers with pedicels 8 15 mm Ion-; sepals hmadb embed*led glands. Flowers in axillary clusters. 5/a- elliptic to obovate, entire, I .1! I ..r) nun limp, 1.2 niinate Jlowcis pedicellate; sepals 5, imbricate; 1.8 mm broad; petals elliptic lanceolate, 2.1 2.7 petals 5. inucli shorter than sepals, pubescent; disk mm long, 0.8-1.1 mm broad; disk « upuliform. annular; stamens 5, free or connate, the anthers fluted, 0.4-0.5 mm high, 0.9 1 mm broad; an- ± introrse; pollen grains 3-colporate, the sexine droecium 0.7-0.9 mm across; stamens 5; anthers echinate; pistillode .'Mid. Pistillate flowers pedi- 0.4 mm across; pistillode head -ncular, 0.5 0.6 cellate; sepals 5, imbricate; petals 5, much shorter mm across. Pistillate flowers with slender pedicels than sepals, pubescent; disk annular; pu- becoming 2.5-5.5 cm long; sepals suborbicular to bescent; styles free, twice bifid; ovules paired in elliptic, 2 2.2 mm long, 1.8 2 mm broad; disk each locule. heinitropous. capsular; colu- cupuliform, its margin undulate, ca. 1 mm high mella distally expanded into 3 broad papery wings; and 2 mm broad; styles thickened, 0.6 mm long, seeds paired or solitary in each locule, smooth, not bifid, the tips clavale. Fruits oblate. /I angled, re- fleshy, ecaruru ulate; absent; cotyledons ticulate-venose, cocci ribbed on ba.'k; columella greenish, contortuplicate, much broader than and cylindric, 3.2 3.5 mm long; seeds plano-convex, about as long as the radicle, smooth, yellowish, 4.4 5 mm long, 3.8-4 mm

The recent discovery of istrocasia in Panama WKBSTKK, C. I.., L. Gil. is one of the most surprising additions to the llora, Systematic* of Cro\ particularly since it was found near Madden Dam

in what is surely one of the most heavily botanized The affini,ies 0f this small neotropical genus of

locations in the country, .istrocasia tremula has tnree S[)ecjes [,ave remained questionable because

a broad but greatly disjunct distribution from Mex- of fragmentary material. The recent discovery of

ico and Jamaica to , Venezuela, and Bra- staminate flowers of Croizatia naiguatensis Stey-

zil. The Madden Dam locality, however, is the only erm. (Webster et al., 1 987) has not made it possible known station in Central America south of Belize to determine the alfin.ties of the genus more closely,

and Guatemala. jn th(, proto]()gll(. ,„ ,|„, orig,nal description of

Specimens examined. PANAMA, COLON: forests along Ooizatin. Steyermark proposed a relationship to dH.ie.ut Madden lake, •„•.„ \l.i,l,ln, D.n„. "»(i ,„. knap), the genus Blume on the basis /2W(DAV, F, MO), Witherspoon H«c>.» (DAY. MO): 6 of a suggestion by Dr. Leon Croizat. That sugges- Chilibre, along Madden lake K,i'i/>/> .

and . On the basis of gross morphology,

Aublet, Hist. PI. Cuiane 256. 1775. from Actephila, es[)ecially by virtue of its pubes- : guianensis Aublet. cent petals and ovary and its twice-bifid styles. In tin.- latter character it is similar to the African Aublet, Hist. PI. Peutubrachium Muell. Arg.; however, in the Af- Guiane 256. 1775. TYPE: French Cruana: hi- riean genu-, the seeds ha\e abundant endosperm hid (possibly at BM, not seen). and the is not contorted as in Croizatia. The echinate pollen grams ol (,i ,,i ..alia are \crv Idditionnl specimens examined. PANAMA, (,OI.6N: ra. „ , c , , ... n t . ,g Ri„ Chagres and associated tidal channels off side ,,lff<*•1 [ro• ,hose ol -^tephlla or Pentabn 1 ') km from Fort Lorenzo turnoff, 9°40'N, }

3.1. Croizatia panamensis Webster, Syst. Bot. 12: 7. 1987. TYPK: Panama. Panama: primary forest along road from El Llano to Carti-Tu- , Bot. 28: 308, fig. pile, 300 500 m, 30 Mar. 1973, Liesner 57. 1952. TYPE: Croizatia neotropica Stey- 1279 (holotype, MO; isotype, DAV).

Shrub or small tree 16 m high, usually with Dioecious trees or shrubs; indumentum simple. a single main stem. I.cares with petioles 0.5-1 cm Leaves alternate, petiolate; stipules persistent or long, 3 1 mm thick; stipules ± persistent, oblong- deciduous; blades entire, pinnately veined, without lanceolate, acuminate, ribbed, sericeous, 10 20 Volume 75, Number 3

mm long, 6-7 mm broad; blades chartaceous, gla- ers sessile; calyx 3-5-lobed, the 1 brous or spar-eK • . . . !. i oh- disk segments 3-5; stamens 3-6, ovate, abruptly short-acuminate, basally attenuate, exserted from calyx; anthers intror 22-47 cm long, 5-15 cm broad, with ca. 15 dehiscing longitudinally; connectiv arcuate-ascending lateral nerves connected by in- pollen grains prolate, 3-colporate, : train,irginal loops, the veins and (to some extent) ti< <11.1i• . | n< li v», •• i< ell,iie; calyx 3-5-lobed, the lobes imbricate; disk with sparsely pubescent pedicels 3-4 mm long; cupulate; ovary 3-locular, glabrous or pubescent; sepals (4-)5, elliptic, entire, 1.7-2.5 mm long, 1- styles short, bifid; ovules 2 per locule, anatropous. 1.5 mm broad; stamens 5, the filaments 2-4 mm Fruits capsular (somewhat fleshy and tardily de- long, connate at the base for 0.5-1.5 mm, the hiscent); columella slender, upwardly dilated, with column long-pubescent; anthers 0.6-0.8 mm long; papery wings; seeds solitary in each locule, pistillode 1.5 2.5 mm lorn with ecarunculate, the outer testa fleshy pubescent pedicels ca. 1.5 cm long, becoming 2.5 present; cotyledons broad, plane, basally 3.5 cm long in fruit; sepals 5, elliptic-lanceolate, A neotropical genus of five closely related species, ± acute, 8-12 mm long, 3-4 mm broad, i i , u pre\ ionsK mm > I Iron m i'md North Amer- ulous without, persistent and becoming reflexed in ica. Richeria appears to be most closely related fruit; ovary 3.5-5 mm diam., densely hirsutulous; to the African genus Marsohotrya li'-nlh. and lo styles 3, 3-4 mm long, connate basally into a >.".;., •,., I Jlil'iM ol - nil . a: < • II V la J'l ! Ma a-. 1.1 column ca. 1 mm high, three times bifid. Fruit capsular, 10- 1 5 mm broad; columella ca. 8-9 mm differs from that of Mueller (1866) and Jablonski ' 'I Vo-in*"""? ^^q ""*TTUB' °"1U""'' (1967), since section (Klotzscb) Muell. msn, 1.1 mm long, b.d rj.b mm broad. ^ ^^ ^ Richena loranthoides (Klotzsch) aiiitoicsls. Panama and Colombia. Muell. Arg.)-I n.M I u ed as the monotypic genus Podocalyx Klotzsch, which, in fact (as in- This more complete species de- icription of Croiz- dicated by the spinose pollen), belongs in the atia panamensis has been mad e possible by re- I i lil I » I deae rather than the Phyllan- cently collected flowering specim ens and data pro- vided by Dr. Gordon McPherson. It is now apparent There are two species of Richeria in Panama. that C. panamensis is clearly different from C. neither previously reported. naiguatensis in floral characters: staminate flowers with stamens connate in C. par mmensis (free in ('.. naiguatensis), staminate pet; lis more long-cil- iate and styles more divided in C. panamensis. Since flowering material of C. r wotropica is still or nearly so; stipules less than I unknown, it remains difficult lo i tssess its relation- folate 1. H. obov ships wiili ( «... distinctly puberiilent: si vies on long

I from El Llano to Carti, beyond Nusagandi, " '15'N, 79°00'W, ca. 300 m MO), ill. |H-t,l!atr;, IK) If, 11041 (DAV, MO). Richeria obovata (Muell. Arg.) Pax & K. Hoffm., Pflanzenreich IV. 147. XV(Heft 81): 29.ZV. 1922;tvzz; Jablonski,jablonski, Mem. Newl>ew YorkI one Bot.not. 4. Richeria Card. 17(1): 126. 1967. Richeria Vahl, Eclog. Amer. 1: 30, tab. 4. 1797. f obovata Muell. Arg. in DC, Prodr. 15(2): TYPE: Richeria grandis Vahl. 468. 1866; Fl. Bras. 11(2): 16. 1873. TYPE: "Brazil,1' Rio Casiquiari. Spruce ,'i.'>2f) (not Trees or shrubs; dioecious; indumentum simple seen). or absent. Leans allereale. j>ctiolal< : stipules de ii o i Mi i i .1 i i 1 < renulate, pin- A species description is not offered here, since nately veined, sometimes with basal laminar glands. the Panamanian specimens are incomplete, and it Inflorescences axillary, racemose or spicate; sta- is not possible to expand the description of Pax & minate flowers several per in sessile or pe- Hoffmann. In the absence of flowers, it is not dunculate glomei ll.it. 1 acts subtending entirely certain that the Panamanian specimen- / flowers; flowers ape I . • • . itht Missouri Botanical Garden

ally ovoid-ellipsoid, tapering with reddish, fleshy ribbed-st mm long, 4.8-5.3 PANAMA. PANAMA: ca. 5-6 mi. \ ,,i 1:1 I In,, , Gentry 5796 (GH, MO, SCZ). Bharpl, cloud forest, Cerro Tute, NW of Sa. Hunk, yju-i (DAV, MO). noecium with distinct styles; t are similar to th <>| /»'.•< itenu ^uuuhs toil ddlei 4.2. Richeria dressleri Webster, sp. nov. TYI in the more slender cylindrical pistillode. The acu- Panama. Panama: Santa Rita Ridge, road minate leaves often copiously hirtellous beneath Estaci6n Calibrar el Agua Clara, 9°22' and the large foliaceous stipules also appear dis- 79°42-45'W, 1,000-1,500 ft., 26 June tinctive. The • oil.-, lions hum ( !os|j Rica are mor- 1971, Webster & Dressier 16744 (holotype, pho o i III divergent but mav tenlalivelv be DAV; isotype, MO). grouped with the Panamanian . It seems ap- i lifl'ert stylis elongati capsulifl 2-locularis; foliiss aacuminatis eglan ' ' > sericeo-hispido.

Tree to 15 m high, 3.5 dm thick; twigs s made a significant contribution mostly densely appressed-hirtellous when j to our knowledge .I Mesoamencan Kuphorbia

A number of collections made in cloud forests prismalie. apiealK hirtellous. 0/2 0.3 mm high; in Panama indicate that there is at least one ad stamens T)( ()); filaments free, 2 3 mm long; an- I • - • • ii< the lowland thers ellipsoid, ca. 0.4 mm I i [ i I, lui //. laxiflora. However, as often happens, these ducal. denselv hirtellous, 1-1.2 mm high, 0.6 0.9 mm broad. Pistillate flowers subsessile; sepals taxonomically; the following revised treatment is mostly 4(-5). elliptic, lometitulose outside, sen li lil i i ending revisionary studies of this ceous within, 1.5-2 mm long, 0.8-1.3 mm broad; across; ovary of 2 carpels, sericeous; styles 2- or 3-fid, 1.5-1.7 mm long. Capsules ellip Old, (not venose), ca. 10 13 mn ; pistillode slender, bid,I. 0.7 0.8 n I; columella flattened, papery s sparsely lepidotc; blades inosilv I' long; seeds somewhat asv in g with petioles of 3-9 cm; stipu long; ovary densely lepidote; endo- cupulate, shallowly 5-lobed, 1 -1.4 mm high, densely over mm ong lepidote; disk massive, 0.6-0.8 mm high, densely ; pis.illodr S.O..I. nm l.ilid. to 0.5 mm "" lepidote on top. Pistillate/towers subsessile (ped- es sparsely to densely lepidote be- icels equaling or shorter than the ); calyx cupulate, shallowly 5-lobed, densely lepidote, 1- of 1-2 cm; stipules apparently obsolete; ovary 1.3 mm high; disk cupulate, subentire, glabrous, glabrous to lepidote; endocarp of fruit at least 4 mm long 2. H. oblonga ca. 0.4-0.5 mm high; ovary ovoid, ca. 1.5 mm high, glabrous or nearly so; stigmas punctiform. 5.1 Hyeronima laxiflora (Tul.) Muell. Arg., FruUs elllPsoid' acute at both ends' coarsely bul" Linnaea 34: 67. 1865. Stilaginella laxiflora late-rugose, 5-6 mm long (endocarp 4-5.5 mm Tub, Ann. Sci. Nat. Bot. Ill, 15: 244. 1851. lon6>- TYPE: Guyana: "British Guiana," Schom- Montane rainforests, Guatemala to Panama and burgk 879, Hostmann 391 (syntypes, P). South America.

The specimens originally cited under this name With some reluctance we are referring all of were correctly referred to H. laxiflora, which is the high-elevation (cloud forest) populations of Hy- apparently widespread in lowland rain forests in eronima in Panama to a single species. There is a northern South America. All of the lowland pop- striking amount of variation in pubescence, and ulations of Hyeronima in Panama belong to this the literature might lead one to recognize two, species, three, or even more species. Plants with densely lepidote leaves, pale axes, and the 5.2 Hyeronima oblonga (Tul.) Muell. Arg., ovary glabrous or nearly so could be referred to Linnaea 34: 66. 1865; in DC., Prodr. 15(2): H. scabrida (Tul.) Muell. Arg., and plants with 271. 1866. Stilaginella oblonga Tub, Ann. sparsely lepidote leaves and densely lepidote ovary

Sci. Nat. Bot. Ill, 15: 248. 1851. TYPE: to //. oblonga s. str. However, specimens from Guyana: "British Guyana," Schomburgk 805 Darien in particular have the pale inflorescence (P, not seen). axes of H. scabrida combined with the sparsely lepidote leaves of //. oblonga. Both "species" oc- cur in the vicinity of El Valle. Plants from the vicinity of Cerro Campana, divergent in having SYNTYPES: Mexico. Oaxaca: Hartweg 513 (P), Gal- densely lepidote leaves (with prominulous venation ... eot l , • o • i r> /- ,r* above) and larger flowers, appear to match the Hieronyma guatema TI. D. S • Bot. Gaz. (Craw- ; 8 . ^ fordsville) 54: 241. 1912. TYPE: Guatemala. Alta descriptions of H. oblonga var. benthamu (Tul.) Verapaz: Tuerckheim 423, II 2228 (not seen). Muell. Arg. However, it is not clear whether that _ , _ ,. , , , , , variety can h. isfad oil I "limited from other Tree to 10 m high; young twigs angled densely popuktions 0n,y cntlca, field gtudies can ^^ lepidote (scales ca 0.15 0.25 mm across). Leaves whethef the broad deiimitation of H oblonga with petioles mostly 10-15 mm long; stipules ap- ^ ^ .g ^^ parently absent; blades mostly obovate, abruptly cuspidate or short-acuminate, cuneate at base, gen- Representative specimens examined. PANAMA, BOCAS erally 4 8 cm long, 2.5-5 cm broad; major veins DEL TORO: between Criollo and Quebrada Higueron on ca. 5-7 on a side, divergent, straight, brochido- |1| ,,, i( ^{] _ f , ,,

0.25 mm across, with pale center (lower sur- M00 m Gentry & Mori ]3629 (DAV< M0). Cerro face much paler than upper). Panicles densely Tacarcuna, 1,800-1,850 m, Gentry & Mori 13989, lepidote with whitish scales; lateral axes mostly 2- 14025 (DAV, MO). PANAMA: Cerro Campana, Webster & %,4 thetne staminate ones 5-10o IU cm lonelong, theuie pismiaieDistillate BreckonQ 16 1^ I/ • y ;/ ^ Ay xs: 3-4 ^^ km W ^of ^^Santa ones ca. 1.5-2.5 cm long; bracts densely lepidote, Arizona N of> , _ , ff(mmd 4?4] (MQ); acute, ca. 0.7-1 mm long. Staminate flowers with Cerro Tute, just W of Santa Fe, Knapp & Dressier 5390 rigid stout pedicels ca. 0.4-1.2 mm long; calyx (MO). Additional specimens examined. PANAMA. PANAMA: Kino Colorado I., \nnour Trail, hosier X (mat 2308 II) \\); vi.-initN ol Vrnioiir Trail. Croat 14843, 14849, H>.,lt> (|)\\. Mill: S ol /,-tek I I. hosier 1122 (DAV, Dl KK. MO) COLON: Santa Hila lumber roa.l. 0.4 km 1'rccs or shrubs, dioec from Transisthmiai, Highway, Dressier 38I

iminal disk; stamens mostly 4-5(-50), fila- s tree; anthers basifixed, extrorse to i grains tricolporate, reticulate; pi-iillode pros >r absent. Pistillate flowers pedicellate; disk [ or 2 (rarely 3 or 4) carpels; -ivies obsolete or nearlv so. dilated stigmas capping the ovary; ovules 2 in each locule, anatropous. Fruits iiidelnseent, ± drupaceous, the exocarp i L., Sp. PI. 981. 1753. fleshy or leathery, the endocarp crustaceous or bony; seeds usually solitary in each locule. eca- nineulate. the testa smooth; endosperm copious; embryo straight, the cotyledons broad and flat.

A large circumtropical genus of about 150 species, best represented in the Old World; about mique du genre Phyllanthus (Euphorhiaeees). Hois- 20 neotropical species have I n described. 'I he single Panamanian species was discov ei ed on li.it i o ( a.lo.ado Island shorllv alter the publication of our 8.11. Phyllanthus anisolobus Muell. Arg. in original treatment. DC, Prodr. 15(2): 382. 1866. TYPE: Peru: Paeon (holotype, C).

The collection from La Palma, Darien (Pittier (>(>()(), US) mentioned with doubt in 1968 now appears to represent Phyllanthus anisolobus on the ha-i.- ol il- i-i•semblance to tile Darien specimens i lied below. Drypetes standleyi \ , Madrono 24: 65, fig. 1. 1977; Croat, Fl. Barro Colorado Additional specimens examined. PANAMA. I«K:AS DFX I. 529, fig. 321. 1978. TYPE: Panama. Canal I oiio: Ki'o Trnli.-. UT.YN I'IICIO I'.ilen.pie, knUuidc X Hale ,...•< |\|()| IIMIIIN: M.mono. Kirkhnde X II,,Man Zone: Barro Colorado I., Armour Trail, Fos- !><>:> AID): Km Balsa, between Manene and Guayabo, ter <£ Croat 2307 (holotype, DAV; isotypes, Puke .V \ol.ason 11'/.,!! (MO); bio Pnc.ru. between DUKE, F, F neg. 62358, MO). Cerro Mali and ( erro lacarcuna, Centry& Mori 13861 (MO). u:n\u v,: :' ., km \\\ ol Santa Ye on road to bio CalouTo,,,. :,()() .()() ,„. Ilrmandc: et ah 744(F). levi may occur over a broad area in Panama. A barren collection from the Burica Peninsula, Chi- 8.12. Phyllanthus gentryi Webster, sp. nov. rtqui Province {liuse\ fpeiaes niav, also MO). occur in Costa Mica; a specimen from La Selva (Hartshorn 1009, DAV) resembles I), standleyi, although it is divergent in having more slender Webster & Huft Panamanian Euphorbiaceae de< i.h hnmehleis ;>11.11.i i: t• >i in. at least 5 dm long, obtusely angled, brownish, minntelv scahrid- ulous. I.cares with petioles 5 8 mm Ion; : !qnil nmi li'ii;;., 1>I id> i > imm • !l , n i I i 12-28 cm long, 6-7.5 cm broad, acuminate, 9. Caperonia lloueis. St,unuiatc /lowers with pedicels 15-20 Caperonia A. St. Hil., Hist. PI. Remarq. Bresil nun long; scpab •. lu-iMtlh i-lliplii oi oh • M \ .' • 244. 1820. LECTOTYPE: Caperonia castanei- 3 mm long, 2.2 2.7 mm broad; disk entire, angled, folia (L.) A. St. Hil. (Croton castaneifolius L.) (chosen by Britton & Wilson, Bot. Porto Kimh: 180. 1925). and ca. 0.5 mm broad; anthers suborbicular, flat- tened, dehiscing [ii.n. fill; II', , <>>; i I.') i it i ong 11.1 broad. I'istilla I dicels 8-13 mm long; -M-p.il . . .. , ' nni. mostly 3 4 nun long. 2.5 3 mm broad; disk uiassi\e, angled. C. paludosa Klotzsch. It is extremely difficult to separate that species I mm ('. castancifolia (L) A. 3 car(iels; styles in-arb free, dilated, 1-1.2 mm Si. Nil., and we now believe that our Panamanian long, 0.7-1 mm across, ('.apsides reddish, valves specimens of (.'. paludosa probably represent ..nh 12.5-13 mm long; columella massive, 4.5-5 mm forms of that more wide-ranging species. However, long, 4.2 4.5 mm broad; seeds trigonous-umbo- the narrower leaves of plants referred to C. pal- nate, 0.5-7.1 mm long, 5.2-5.3 mm broad, udosa are distinctive, and furlhei studs in the held smooth, with irregu n \\a\ • horizontal dark brown l i« -1 d to eslal i h Itl p. • I. - < oneepl I i i I i i light hi mi !>.!• [ I'lound. the apex some can be upheld.

1.5 mm across); hi triangular, broad, 10. Argythamnia i long and broad. Argythamnia P. Browne, Civ. Nat. Hist. Jamaica •Uiilitional specimens examined. PANAMA. DARIEN: trail u|> Orm I'nre. (,c,,!ry 4589 (MO); razorback ridge mi Criiu I'niv, PII/.C t, ).)/) (MO); Serraiua de 1'irre, trail (to„; O IVnviiiu.) t„ Cerro I'irn-, ca. 10 km airline SSE of HI Heal, in subtropical moist-to-wet forest, 300-750 .... Reccul X Duke 40/<>(MARY. MO); around Rancho Frio, halfway up slope of Cerro Pirre from Piji Vasal, holsom ol'/o (K, MO); S of El Real on trail up Cerro 1805: TYPE: Alchorneopsls floribunda I'nre e.i H°00'N, 77°45'W, 550-1,030 m. Ud'h, >,,•;, (Benth.) Muell. Arg. { var. '^floribunda Benth.). I'liyllanthus gentryi is the first representativ of subgenus Xylopliylla (L.) Pers. discovered i .1 Alchorneopsis floribunda (Benth.) Panama. It clearly belongs in se( Muell. Arg., Linnaea 34: 150. 1805; in DC, (Klotzsch) Muell. Arg. by virtue of its Prodr. 15(2): 705. 1800. Alchornea glan- dulosa Poeppig var. floribunda Benth., Hook- P. juglandifolius Willd. in general a.' er's J. Bot. Kew Card. Misc. 0: 331. 1854. though il could be interpreted as a sub : Spruce 2681 (holo- /'. juglandifolius, it is distinctive in its la I'he • original treatment, from Darien, ' the key on p. 221 a mined. COSTA Rl( TAGO: 24 km NE of Turrialba on hwy. to Limon, at Tres Equis on jeep road 1.5 km, 9°58'N, 83 450 525 m, Liesner et al. 15354 (MO), HEREDIA: Finca La Selva, the OTS Field Station on the Rio Puerto Viejo just E of its junction with the Rio Sarapiqui, ca. 100 m. Hammel 9425, 11083 (F, MO); Istaru Farm, Hrimbina, ' •" ... /.- "I \1 MM* -.,.) ,,| I.omas ,lc Sicrpe, NE of terminus of road from Villa franca. I0°19'N, 83'34'W, Grayum et al. 3;>JO (!'. MO), SAN JOSI•:: 2 kin N of Dominical along CR 223, 40 This poorly known genus, hitherto considered 100 in, I llry & I tley 4938 (F). PANAMA. BOCAS DEL to be South American, includes only three species. TORO: Cerro Pila de Arroz, along road to Chiriqui Grar 1 10 road-mi from Continental Divide and 2 mi. al Croi I ' i ported aedra from Pan- pipeline access road E of highway, ca. 8°55'N, 82°08' ama on the basi- 350-500 m, McPherson 8750 (F). but in the original treatment (Webster & Burch, 1968: 278) it was pointed out that his original generic disposition (J. Arnold Arbor. 24: 167. 1943)

jndron Kars ten, Fl. Columb. 1: 91 ol tins i- iiannm was cor-

45. 1860. TYPE: Caryodendron :cn« i • ough that species is now known to be Kar anonymous with C. membranaceum Pax & K.

12.1. Caryodendron angustifolium fii nil; r t.-.bh-b the piesen •(• ol Idenophai

The collection h li«- i I ' in nulci ' 13.1. Adenophaedra grandifolia (Klotzsch) in the original collection is now known to be Se- Muell. Arg. in Mart., Fl. Bras. 11(2): 386. ne/eblera lesticulata Pittier (q.v.). No additional 1874. Tragia grandifolia Klotzsch, London collcclioris el < ('.< wi./i 'Hi/on ! :/<>!, •, ».< nc J. Bot. 2: 46. 1843. TYPE: Guyana: "British known from Panama or elsewhere. Guiana," Sehomburgk 948 (presumably K, not seen) H Klot/scb) Muell. Arg., Linnaea 34: 173. 1865; in DC, Prodr. 15(2): 918. 1866. Adenophaedra (Muell. Arg.) Muell. Arg. in Mart., Fl. Bras. 11(2): 385. 1874. Bernardia sect. >l

iinbncalc; |>etals absent; disk 3-lobed; ovary of 3 to 25; pistillate spikes thicker, not flexuous. 5 I 2 14.1. Bernardia macrophylla Siandlcv, I • in long, wild 4-7 solitary flowers. Staiim-n, Wash. Acad. Sci. 15: 103. 1925. TYPE: Pa- em to 12 per glomeruli earl\ dehiscent, leaving nama: Rio Tocumen, near sea level, 3 Jan. persistent pedicels ca. 1 mm long; calyx lobes mem- 1924, Standley 29389 (holotype, US). hranoiis. ilcll,ilc. spreading at anthesis, ca. 0.5 mm No Panamanian specimens of this species have long. Pistillate /loners not seen; pedicels at ma- been found in addition to those cited in ihc original turity ca. I mm long, reflexed, sericeous. Capsules treatment. The recent collection cited helow ex- depressed globose. deepK 3 i<>ht » mm high. tend- llie range to Co-la Iv'ica. I hat .specimen has 12-18 mm diam., sericeous, glabrate at maturity, the persistent calyx lobes ca. 2 mm lout:, deltale. ranamainan eolleclioiis and was taken at a con- sericeous; seeds suhglobose to ovoid, ca. 1 mm siderably higher altitude but seem- otherwise idcn long, ca. 0.7 mm diam., yellowish, mottled. tical. As indicated in the original treatment, the closest relative of Bernardia macrophvlla seems to be H. jaequiniana Muell. Arg. of Venezuela. hnl the killer differs in having retrorse rather than as. ciidine. pubescence on the stems, MIMIC promi <;,;„„•: el „l. :'.' nenl \enatioii on the uiidersurlace ol the leaves. and 9 12 vs. 14 stamens. fll.in (I ): Cerro Pila de Arroz along road i.> Chin,,-,, TAKKNAS: foothills of the Cordillera de III Crande, 10 read-miles from Continental Divide and 2 mi. Tres Colinas, 9°07'N, 83°04'W, 1,800 ,,,/s, el a! _'.>6// (F).

82°()8'W< 350 500 m, MePherson 8752 (F). 8777, (K,

\\\ olSan'.a le , •>7t>'(GllM0). 1 .. Svst.

Miller, Card. Diet. abr. ed. 4, 28 54. LECTOTYPE: Bernardia earpinijidic \ Ichor.ua Svv., Prodr. 98. 1788; Fl. Ind. Occ. iseb. (see Buchheim, 1962). 2: 1 153. 1800. TYPE: Alchomea latifolia Sw.

Several recent collections of Alehornea indicate that there are some additional taxa in Panama, but the material is still inadequate lor a satistactorv treatment. The taxa that appear lo be present mas be treated as follows.

tische Hemerkungen iiher die Cattung Bernardia PAX, F. & (Fuphorhiaceae). Willdenowia 2: 291 318. lyphe; . I%2. ('her die 'f\|iusarl der Cattung Her- /enrcicl, l\. I 1.. \ 11(11, it ( miidn, (Kii|.horhiafeae). Willdenowia 3: 217-220.

ciilum only moderately

3a. heaves M p| • mostly axillary

4a. Styles f> 20 mm long, relatively slender. 5a. Leaves coriaceous, usually with 2-4 basal glands; spikes mostly cauliflorous. 6a. Leaves mostly 8 20 cm long, acuminate, with mostly 5-8 main lateral veins, ent or crenate-dentate; foliar glands mostly 2(-4); pistillate sepals 2-2.8 mm long .. Webster & Huft

glands 2 4; sp;l II , 21). Leaves. >\ ei 20 « i I [,i„ n I- I, II above and dis >ei il Pistillate spikes mostly 25 cm long or lorn

with this leaf form c

235. 1914. TYPE: (

arianls within /. latijblia, hut i

ordinary amplitude »i I i 1 studies in the field ade, the situation will record from < olon hi (< ,1 i: Ordonez et at. 58, nain unsatisfactory. MO) The plant described from Colombia by Croizat (Caldasia 2: 357. 1944) as A. umboensls may hl:!(i;„ial specimens examined. PANAMA, CHIRIQUI: prove to be a form of A. costaricensis. San Felix, Croat 33416 (MO). PANAMA: Cerro Campana, Croat 14673, Duke 10742, Sullivan 434 (MO), Mendez Idditional specimens examined. PANAMA, CHIHIOI i: 19, 49(F); Cerro Jefe, Buyer et al. 501H. 50 19, (,entry \iemil\ of San Bartolo l.iinite, 11 mi. W of Puerto Ar- (MO Gentrj & Dwye, i.i it (Oil. MO). II ehste, muelles. Croat 21973a (MO). COCLE: along Rio San Juan & Dressier 16454 (DAV); between Cerro Jefe and Cerro below its junction with Rio Tife, Hammrl .tin, (MO). Azul, Tyson et al. 4325, Mori et al. 65 13 (MO): Cerro COLON: Rio ltui|iit i. v i. n • i • •>••. b Rio Escandaloso, Azul, Dwyer 5042 (MO), Lao & Holdridg, .i.i (1)\\. MO), Stimson et al. 5158 (GH, MO), Tyson & Blum along liui 1.ii,m li i ,i il ' c« lii ,1, i //,-;// A 4081 (MO); N of El Llano, Gentry 5105 (MO); El Llano- I /."' I\K)I: .I mi. from Portobelo, Correa di Dressier Carti road, 7.8-8.6 mi. from Pan-American Hwy., Fol- I7V)(C,\\. MO), DAKIKN: Kio Ucurganti. liristait Il'i7 som 3572, Mori & Kallunki 6405 (MO). (MO): luo Inqueza, below Quebrada Venado, Bristan it><> . (I) \\ , MO). LOS SANTOS: Loma Prieta, Cerro Grande, Hide I 187,9. I, ,r,s <•! ,//. 2208(MO). PANAMA: Chiltepe, 16.3. (Spn ngel) MueB Holdridge 6471 (MO), SAN BLAS: Canagangi, forest up- Arg. in DC, Prodr. 15(2): 909. 1866; Pax >lrenm of village, 9°24'N, 79°24'W, 100 m, de Nerers & K. Hoffm., Pflanzenreich IV. 147. VII (Heft et al. 5720 (F). 63): 227. 1914. Antidesma trip/inert turn Sprengel, Neue Entdeck. 2: 116. 1821. TYPE: ».2. Alchornea latifolu 5w., Prodr. 98. Brazil. Rio de Janeiro: Serra do Mar, Gardner 617 (neotype, G; chosen here). I Peru

Further botanical exploration has shown thai Alchornea latijblia is widely distributed in Pana ma. iiiiludiiiL- Hnrroi (,|«, :..., I-and, whence it was correctly recorded by Croat (1978). On severa peaks and ridges in central Panama C hose of Alchornea trip- forms that appear very different from typical A. latijblia of lowland Central America and the West tlchomea previously known from Panama. Another Indies. For example, plants with entire leaves and eolleclion that may represent •/. Iriplincrria is niiiisiialU short petioles are found on Cerro Jefe Hamrnel 7252 (MO) from Cerro Sapo, Darien; this and Santa Rita Ridge (e.g., Gentry & Dwyer 5536, has much larger leaves and somewhat resembles Croat 15309). These specimens somewhat suggest some of the aberrant forms here treated as A. the South American A. pearcei Britton, but their for the present, its assignment must be relalncK lon;;; f >«• 111 d«• - ami -liort s|> kes ::in-, tfiei « regarded as dubious. closer to A. latijolin. >[>• nam- -,\ ith very unusual The collections from Code do not fit any of the narrowly obovate leaves have been collected on varieties recognized by Pax & Hoffmann (1914: Santa Rita Ridge (e.g., Croat 13844, Duke 15264); 228-230), but the variation within A. triplinervia has not yet been critically studied, and it would Although it has been confused with A. glan- certainly be premature to assign the Panamanian dulosa, the relatively short thick styles and stiff material to a new variety. The typification of Al- glandular leaves distinguish A. grandiflora from chornea tripliner pun .u « nm n i / i>Jli i / folia. Mueller (loc. Sprengel apparently left no type specimen. Mueller cit.) reported A. grandiflora from Costa Rica and (1866: 909) designated what may be regarded as Venezuela, so its occurrence in Panama is not the typical clement of the species as Alchornea surprising.

, r .•,.,•„,,*, */>ec,mens ruunilied. PANAMA. CHIR1QU1: Cerro there is a good microfiche image (G, Prodromus Colorado> li690 mj Croat 37195 (MO), numv Cm, Herb.) of Gardner 617, probably from the general Tacarcuna, elfin forest, 1,800 1,850 in, Gentry & Mori area of the collection that was available to Sprengel, 13995 (DAV, F, MO). ; appropriate to designate that as neotype.

Specimens rxamim-d. PANAMA. <:O<:I,K: h 16.6. Alchornea grandis Benth., Bot. Voy. alle, E slope of Cerro Gaital, 900-1,000 m, Knapp Sulphur 164. 1844. TYPE: Colon 351 (MO), McPherson 11242, 11260 (MO). Tumaco, Barclay & Hinds (K,

he specimens cited below, and several collec- 16.4. Alchornea glandulosa Poeppig 3 from Choco Province, Colombia {Fernandez pittieri (Pax) Pax, Pflanzenreich VII(Heft <3): W"l ;iT Alchornea pitted 'W;' KilUl> & '<'<«'"•-• 3W<\ both UC) fur- Syst. 33: 291. 1903. TYPE: msl' l"VN'uusK ".iknown characters for the sta- _:—^ pj taminatc spikes nmstlv compound, Costa Rica: Canas Cordas, Pittier 11101 e ant: s cm long, with 1-6 lateral axes; staminate otype, US). glabrous, sepals ca. 1.2 mm long: stamens he Darien collection of Terry & Terry 8, anthers 0.7-0.8 mm long, blunt. was reported in our original PANAMA, VKKACUAS: Isla ot additional specimens oad from Canipameiito Jimc.il io Coloma IVi confirm the widespread irrenee ot I , V(il || glandulosa in montane forests of Panama. Ex- amination ..I till | IIMl pittieri, originally described from Costa Rica. Tl variety is very similar to var. glandulosa of t -Hk. upper Amazon but differs in the smaller glandul spots at the base of the leaf (mostly 0.5 mm long Tree to ca. 10 m high; trunk 1.5 dm diam.; or less in var. pittieri, reaching 1-1.5 mm long twigs subterete, smooth, glabrous. Leaves with siout in var. glandulosa). At present, var. pittieri is petioles 0.5-1.5 cm long, glabrous, plicate; stipules known only from Costa Rica, Panama, and adja- inconspicuous, ca. 1.5 mm long or shorter, dark, cent Colombia (Choco). triangular, pubescent; blades becoming subcoria- tic to obovate, abruptly t apex (the acumen ca. 12 cm (MO); along pipeline long), ca. 25-40 cm long, 7-15 cm broad, glabrous road in area of rortuna Dam, near end of road, .a. or dabrescent (minute scattered stellate hairs on , ' ' * '" ;•«, '' ' niih d „ I to midrib); basal (|-> 1»MtM V (...n:, C;i;>si I r.-nl, lrn\ A /-•/••. -' .>, .>(MO); ..,.,,, , • , i • Cerro Pure, linsta, M I le Nique, Cerro foliar S,ands obscure or absent' maJor lateral ve,ns Pirre massif, 1,300 1,520m, Gentry et al. 2864 7 (DAV, ca. 10-15 on a side, straight, ascending, raised MO), (eiro I... in II II i . i I I'll beneath, connected by ladderlike prominulous vein-

•Acu•'ir^ aPex abruptly short-acuminate (acumen ca. lunki 6219 (MO). ' ' cm lon^,) >;•,:,e-. « .mliiloroiis, |>endulous, stel-

16.5. Alchornea grandiflora Muell. Arg., spikes ca. 50-75 cm long, with ca. 20 30 flowers. Linnaea 34: 170. 1865; in DC, Prodr. 15(2): Staminate flowers not seen. Pistillate flowers sub- 907. 1866. SYNTYPES: Venezuela: Fendler sessile; calyx ca. 3.5 cm broad, 4-lobed, pubescent; 1272 (C), Moritz 1497a (G). Costa Rica: ovary copiously pubescent with minute stellate hairs; 20-25 mm long, Webster & Huft

connate for 2-4 mm, basally stellate, api- nooth and long-attenuate. Capsules reddish stellate-pubescent, not seen entire; seeds , plump, pale brown, coarsely tuberculate,

Rainforests, Panama and Colombia. 17.1. Cleidion membranaceum Pax & K. Hoffm. in Engler, Pflanzenreich IV. 147. This sinking species stands out from all other XIV(Heft 68): 23. 1919. TYPE: Venezuela. Panamanian taxa by virtue of its long, pendulous, cauliflorous inflorescences and its large, coriaceous, Lara: around Palmosola, in forest along Rio Aroa, near sea level, 26-28 June 1913, Pit- more or less oblanceolate leaves. It resembles A. tier 6375 (US, photo F neg. 44609). grandis in a number of respects but differs in leaf hap< MI sparseness of the laminar pubescence. Cleidion woodsonianum Croizat, J. Arnold Arbor. 24: 167. 1943. TYPE: Panama. Panama: vicinitj id Sal Specimens examined. PANAMA. DARIEN: La Laguna amanca Hydrographic Station, Rio Pequeni, ca. 80 ridg.- brtwe.-n Pucuro and Tapalisa rivers, 820-840 m, m, Woodson et al. 1587 (holotype, A; isotypes, F, Centra cv Mori 13560 (DAV, MO); top of Cerro Mali, F neg. 62417, MO, F neg. 62356, NY). 1,400 m. Gentry & Mori 13693 (DAV, MO); Cerro Tacarcuna, Gentry & Mori 13938 (MO); Alturas de There appear to be no differences between the Nique, S of El Real, 900-1,250 m, McPherson 11614 Panamanian plants and the Venezuelan collections (MO). tnaceum. The three known Venezuelan collections are all from a restricted area near the junction of the provinces of Falcon, Ya- racuy, and Lara. No substrata i il di< < I i n ih« labels of these collections, but it is known that much of this area is underlain by limestone (van der Werff Recent collections in Panama and further study & Smith, 1980), which is also true of the Pana- he South American species have greatly altered manian collections. The recent disjunct collection picture of Cleidion in Panama. Largely as a from Peru strengthens the probability that this ult of problems encountered in the preparation this account, the junior author has undertaken accounts for the peculiar disjunctions in its range. evisionary study of the neotropical species of Further study may show that Cleid (Ca- lclusions expressed here must sar.) Baillon, of Brazil from Bahia I San Paulo, which has leaves recent collection irien by Dr. Gordon to those of the Panamai plant! ^ Pherson may bel

ilchornea oblongifolia Standley, Carnegie Inst. Wash. Publ. 461(Botany of the Maya Area 4): 66. 1935. Cleidion ohlo,-. .(-land I. MM it I >.,„!, >istillate ra

abruptly caudate-acuminate at tip, acute or cu- 'a„ of"these

neate at the somewhat inequilateral base, 15-26 a(,ldwn castanet folium was described from cm long, 6.5-9.5 cm broad, minutely pustulate, -peru - lm| ., |„,^(N(. isu|M„. sh(.H a| p {Huiz &

glabrous, the veins 8-10 on a side; margins shal- Pav{)n sn) [s |ahe]ed as haying been co„ecled at

lowly dentate, the teeth callose, 15-28 on a side. (;u.lvaqiliK E(.uador (another Ruiz & Pavon spec.

Inflorescences unisexual, axillary; pistillate ra- imen a{ f has [1(( ,(((.all|v da,a) J]w spedes ^ n()t

cemes to 16 cm long, widely divergent from the d(,fmitt.|v blowll )rom Peru? but there are two

stem, the rachis glabrous, or puberulent toward the modem',()l,(M,l()Ils lrum lM.lwri.„ Santo Dornjngo

apex, with ca. 3 doners occurring smgU (3 fruits an() Quillinde in Ksmeraldas Province, Ecuador

pistillate seen); staminate (.irosla Solfs UO-W, Little 6/96, both at F, both tin rses 5 9 cm long, wi distributed as Alchornea). at each of the 20-50 nodes, the rachis densely There appear to be no salient differences sep- puberulent. S/„,ninote /loners on pedicels to 1 arating Cleidion castaneifolium from I'anama and mm long; calyx lobes cucullate. reflexcd, 1.5 2 South \nicnca from the Mexican and Central \IIICI nan populations that have long been referred to ('. ohlon^ijoliiim. In addition to the enmiuon -ecu: ln.ni- narrouK Ian late, rigid, divergent populations all ca. 2 mm long, puberulent; fruiting pedicels 10 frequeruly

12 mm long, slightly clavale. puherulous. jointed; ||u> |(1.lv(/s calyx lobes (in fruit) 3 5. somewhat reflexed, del- PANAMA. DARlfcN: Serrar the margins; styles (persistent on mature fruits) by Cerro Pirre, along Q Peredngo (I'.irasrinm) i 12 mm long, deeply bifid, densely strigose. Ca,,- ['^ ',^'liu''plt r^'",'", /"Vn" air km S O|"KI'H'( .... les 3-locular (of which often only 2 fully devel- 77° 1 V\\ . He, ml X Duke 4875 (MARY, MO); ed), deeply lobed. dor-alls rami.He. ca. I cm SE of Pijibasal .... Rio Perasenico, *h, 14-18 mm diam., densely puberulent, drying i:'i<;;;{\-, MOi. ick; columella 6 8 mm long, trigonous, narrowly nged, the seed scars elongate, conspicuous; seeds

d light and dark brown.

Rainforests, souther)

Acalypha L., Acalypha

19.10. Acalypha cuneata Poeppig in Poeppig lections), and since the one on the type & End| N()V Gen Sf) p, 3. 22 1841. TYPE: photo appears crushed and misshapen (and thus ,>eru Maynas: Yurimaguas, Poeppig (not made to appear six-lobed), it seems that Mueller seen) was merely careless in his description. Since these collections are in perfect agreement with all other •i'-alyp/iu ohovuta Ifcnth. in Seemann, Bot. Voy. Sulphur : d d,,,„gu1Sh,„g charace^c, of ,hU species

Shrub or small tree 2-5(-8) m high; monoe- KEY TO THE SPI cious; stems nearl\ glahrous. Leaves with petioles la. Leaves pa 17 cm long, glabrous; stipules lanceolate, 4-7 drical, slei mm long, strongly keeled, caducous; blades obovate ^.5 mm bl or obovate-oblong, cuspidate-acuminate at the tip ' ^vespi (the acumen 1.5-3 cm long), acute at the base, 15-30 cm long, 5-13 cm broad, 2-3.2 times as long as broad, glabrous, pinnately veined, the sec- 20.1. Pluk ondary veins 11-15 per side, arcuate, prominent 1753. r above and below, connected by a prominulous re- at BM). Iiculuni; margins shallowly crenate-denticulate. In- florescences axillary, spicate, unisexual; staminate This *Pet'ies is more widespread in eastern Pan- spikes to 15 cm long, densely flowered, densely ama ,han was indicated by the single collection puberulenl, solitary and pedunculate or 2 4 and (lU>(1 in the original treatment. These new collec- sessile on a slender rachis; pistillate spikes 7-15 t,orls also

4-5 mm diam., hispid, verrucose; seeds obovo.d- (;uan(.h(, () km s of Portobei0, 0-10 m, Nee & Gentry ellipsoid, 3-3.5 mm long, ca. 2.5 mm diam., 8686 (MO). DARIEN: Cerro Pirre, valley between Pirre smooth, brown, the caruncle nearly obsolete. and next most southerly peak, Folsom 4426 (F, MO). PANAMA: 4-5 hours walk upriver from Torti Arriba, 200 This is a widespread species of lowland rainfor- 300 m. h,lsom et „l. 6845 (F, MO), ests in northern South America, and its discovery in eastern Panama is not surprising. It is easily 20.2. Plukenetia penninervia Muell. Arg., recognized by the long-petiolate obovate leaves with Linnaea 34: 158. 1864; in DC, Prodr. 15(2): pinnate venation and axillary pistillate inflores- 770 1866 TYPE; Venezuela, near Biscaina, Fendler 2412 (holotype, C, not seen; photo of Panama treati Fneg. no. 7110).

Plukenetia angustifolia Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 314. 1929. TYPE: Honduras. tire or shallowly dentate. Atlantida: Lancetilla Valley, 8 Mar. 1928, Standley '. Spikes mostly bisexual, 56708 (holotype, F, F neg. 52742). 1 or 2 pistillate bracts , it base, these sub- tending 2 or 3 flowers A. diversifolii ; twigs | li it. glalues. cut. '. Spikes unisexual, to 15 cm long, the pis- Leaves with petioles ca. I cm long, puberulent; stipules brownish, glabrous, rigid, deltate-lanceo- late, 1-1.7 mm long; blades chartaceous-oblong, !li Qiptic, or oblong-lanceolate, acute to acu- minate at tip, abruptly acuminate-truncate at base, 5-10 cm long, 2-4.5 cm broad, glabrous and I ding ibove, glabrate or with a few hairs along the nerves and pa i i ion ieni -land above at the base, often with 1 3 pairs of -mallei ones in a row above them, pinnately veined, the midrib and secondary veins (6 1 1 on a side) prom- inent below; margins shallowly crenate-denticulate. The discovery of a second species of Plukenetia Inflorescences axillary, bisexual or staminate, 0.5- in Panama makes it necessary to provide the fol- 3 cm long; pistillate flowers solitary at lower nodes lowing key. of bisexual inflorescences, the staminate flowers few at the distal nodes. 5Uaminate ftowersmth .'TV) \\ , St/w„„ ///. lb. MO) I'..-. ,M\. .|,„i.. the I I Llano Cart; rd., ca. 10 mi. N of Pan Am Hwy., 500 m, | .< ni i, '-•!:- -| I. , rt-pilose, 4-7 n im long; calyx segments Centn ri al K«/«(M<». -\\ iti \x F.l l.lanoCarti road, r.ri:. I.I IS V . , obovate, acu te, 1.2-1.6 mm long; receptacle ( •ylindrical, 1.6 -1.8 mm high; disk ob- solete; stam ensca. 18-25, inserted spirally on the receptacle, • In- hlan rni , ca. 0.1 mm long, the i with 22. Tragia ..,'.|i. . |: |.. i i.until: . 2 I'd inn, lout:, ill. M- narn.wh Tragia L., Sp. PI. 980. 1753. LECTOTYI-K: T,,lfiia clavate, strigose when young, glabrate to sparsely volubilis L. (chosen by Small in Britton & lis.11 i nilii- . ml a I maturity; calyx lobes lanceolate, Brown, III. Fl. N. U.S. ed. 2, 2: 458. 1913). 1 1.2 mm long, ca. 0.5 mm broad, strigose in a km.: along the center; ovary of 4 carinate carpels, The discovery of three additional species in Pan- strigose on the keels, otherwise glabrous, the stylar ama makes it necessary to provide a key to the column to 2 turn bmonl.ll hid unlobed. Capsules deeply 4-lobed, oblate, to ca. 1 t-m high. 1 .5 fin broad, the cocci thick and rigid;

seeds subglobose, only slightly compressed later PAX.F.&K. HOHMVNV ]<>|<). KupliorliKiccac \ca- ally, reticulate-venose, brownish mottled, ca. 5 mm lypheae —Plukenetiinae. In: A. Engler, Das Pflan- long, 3-4 mm thick. zenreich IV. 147. IX (Heft 68): 1-108 (Tragia, pp.

Lowland evergreen rainforests, Mexico (Oaxaca, Yucatan Peninsula) to Colombia and Northern Bra- .- ( I' !•• .i i. II • !. i epoi !i

wen des. nhed b\ Standlev as /' aiii; 3a. Pistillate flew. tr long-pedicellate; styles no salient differences between these plants and those of South \meru a are apparent. 3b. Pistillate ale: styles free

Specimens examined. PANAMA, COLON: Santa Rita Huli-'i- in i i • IM. I.nn f.-.a \-na I l.u; ; ami. il .laLna Tragia bailloniana Muell. Arg., Linnaea 10(1 ..00 ,„, Foster & Morion 2222 (F); Santa Rita i: 178. 1865; in DC, Prodr. 15(2): 927. Ridge, ca. 9°20'N, 79°45'W, ca. 500 ,„, \L-l>hc,so„ 366. TYPE: Mexico. Tabasco: Teapa, Linden 84b 1 (F). PANAMA: 1-2 mi. S of Pan American Highway, 3.0.,.,. KofCana/a cla-c.;, , ..I! ,t N-r, ; i2'N, 78°I.V\\. 0 50 ,n, Km,, I

co-.ei-.-ii .vilh stinging hairs. Leans will |..-liole- 8-14 cm long; stipules ovate-lanceolate, acumi-

nous, broadly ovate, unlobcd, i mu I.- I 21.1 nicaraguensis (Hemslev) lobe, or shallowly 3-lobed, Webster, Ann. Missouri Bot. Card. 54: 191. at tip, deeply cordate at base, 12-25 cm long, 1967. Cleidion ?nicaraguensis Hemsley, Biol. 11-18 cm wide, sparsely beset above and In-low Cent.-Amer., Bot. 3: 130. 1883. TYPE: Nic- v.illi Minting hairs. usu.ilK 7-\eined at the base. aragua. Chontales: Tate 352, 455 (syntypes, the margins renmleK denticulate. Inflorescences presumably K, not seen).

Additional specimens examined I'WWIA. COLON: Sanla Hita, Cone,

petioles 4-11 cm long; stipules lanceolate, 5 mm long, reflexed; pedic long or more; blades thinly chartaceous, oblong- glabrate; sepals 5,1 or elliptic-obovate, rather abruptly short-acuminate 6 mm long; stamens ca. 40; pyrifoi at tip, distinctly cordate at base with open to closed I'istillolr Jlt,t<;-rs sulilais in the axil ol c sinus, mostly 8-16 cm long, 4-7 cm broad, sparse- ly hispidulous on both faces with stinging and non- stinging hairs, mostly 5-nerved at base; the margins . i- ;> i long; mar bluntly and coarsely crenate (teeth 15-25 on a gins ciliate with long, stiff 1 •y densely side). Inflorescences opposite the leaves, becoming ii juti I i Miff hairs ca. 1 n i long; styl. black, ca. 10-15 cm long, distinctly bifurcate and pro- 6 8 mm long, fused togynous, the lower pistillate branch with 5-10 branches slightly spreading. Capsuh < ! flowers; pistillate bracts entire, 3-4 mm long, the 3-lobed (one lobe sometimes abortive), ca. 15 mm .5 2 i • ong. diam., ca. 8 mm high, densely hirsute with stiff th .... • -In r ii.Hi i in, |,M!,MJ ,, pedicels ca. 1.5- hairs; columella 6-7 mm long, with 3 prominent, 2 mm long, articulate near the base (stumps re- narrow wings at tip; seeds nearly globose, ca. 6 maining after dehiscence of much shorter •otli. with i • 9 light longitudinal than subtending bract); sepals 3 or 4, obovate, acute, strigose without, 1.5-3 mm long, 1.8-2.2 mm broad; disk glands 5, erect, cylindric im k r Forests, southern Mexicc ) western Pan

I' \luell. Arg., dis- flowers with hispidulous pedicels up to 2.5 mm tinguished from all other American species long in fruit; sepals 6, lanceolate, asymmetric, green, is (ca. 40 vs. 2 or 3 reflexed in fruit, becoming 3.5-6 mm long, 1-1.5 , 20) ially lobed mm broad; ovary densely hispidulous with stinging hairs; styles basally connate or nearly free, 2-2.5 considerable range xtension, for mm long, distinctly papillate. Capsules copiously n only from s hispid with stinging hairs, cocci ca. 8 mm long; (Veracruz, Chiapas, columella 2.4-2.7 mm long; seeds globose, mottled duras. Earlier reports of Tragia bailioniana from brownish and gray, 3.6-3.7 mm across. Costa Rica (Standley, 1937: 622) are erroneously The single specimen of this species, previously based on collections of Dalei hampu haul unrecorded from Panama, is in poor condition and without flowers, so there is some doubt regarding collections establish the presence of the former its assignment. The Bristan collection matches a photograph of the type specimen from Venezuela, Specimens examined. COSTA RICA, ALAJUELA: along although the basal leaf sinus is not as open in the Panamanian plant. There is also some resemblance C to Colonia Virgen del Socorro, 10 18'N, 84°10'W, ca. to T. japurensis Muell. Arg., described from Am- 800 m, Burger et al. 11850 (F); lower NE slope of Arenal Volcano, 10°29'N, 84°42'W, 500 m, Lent 2947 azonian Brazil. However, it seems probable that (!• | [H hi Is 2 ail be km SSE of Islas Buena Vista in the Brazilian species is synonymous with the one the Rio Colorado, 14 airline km SW of Barro del Colorado; from Venezuela; at least, no convincing differences 83°40'W, 10°40'N, 10-120 m, Davidse & Herrera are given by Pax & Hoffmann (1919: 36). UOLKiO MO) i .ternmost ridge of Cerro Coronel, NW-facing slope, just S of the Rio Colorado, 10°40'30"N, lined. PANAP 83°39'30"W, 10-80 m, Davidse & Herrera 31388 (F, MO); Cerro Coronel, E of Laguna Danto; 10°41'N, 83°38'W, 20-170 m, Stevens 24383 (F, MO). PANAMA. CHIWQUl: Fortuna Dam site, 1,400-1,600 m, Folsom et 22.4. Tragia correae Huft, sp. nov. TYPE: Pan- al. 5612 (MO). ama. Panama: Picada da Estrada Panama- San Bias entre 320-420 m, 9.1.1973, Sucre, Braga, Dressier & Correct 9832 (holotype, 22.2. Tragia fendleri Muell. Arg., Linnaea 34: RB-165572, F neg. 62359). 178. 1865; in DC, Prodr. 15(2): 928. 1866. TYPE: Venezuela: Biscaina, Fendler 1208 (G). Caulis volubilis 1 u. - mi il ruhelli dense pilosi. Folia alterna elliptica-oblonga 6-12 cm longa septemner- via, infra dense pilosa supra sparsia, basi corda i margin I mostly nonstinging h 1919). This species bear dense hispiduloso, sivli- paj.ill.i11>. volubilis but differ- b\ densely pilose leaves, and Twining woody ; twigs reddish, densely pi- lose, tardilv fzlaln.il.-. ihe older twig- with loose. .III- dis exfoliating bark, leans with petioles 0.5 4 em species for Profesora Mireya Correa of the I m- long, densel) pilose; stiptiliv- delta versity of Panama. Tragia correae is known only acute, () 10 nun Ion-, pilo-e below. -I from the type collection. Dra. Correa has kindly blades membranous, elliptic-oblong, searched for a duplicate at the of the tip. cordate at base, 6 12 cm long, 3 4.5 em I niversitv of Panama (PMA), but so far, unfor- broad, 2.2-2.7 times as long as broad, sparsely ateK. pilo-e above, more densely so below, usually 7-nerved at has.-; margin- remotely denticulate (teeth 18-22 on a side). Inflorescences opposite ia I.., Sp. PI. 1054. the leaves, racemose, ea. 3 cm long (immature), ip hamnia scandens L with a -ingle basal pi-lillali- Mower, the remaining nodes (ca. 20-25) with staminate Mowers; bracts trilid; bract- entire. Staminate flowers on short. hispiduloUS pedicels; • a|\ \ lube- 3. ohovale. aeule. c/mmpta (Euphorbiaceae) I ca. 1.3 mm long, ea. 1.2 mm broad, lu-piduhm- Bot. 9: 272-278. without, cucullate; stamens 3, the filaments thick & A. HERZIC. 1984. and tle-li\ . tree. ea. 0.8 nun long; anther- elliptic. 0.2-0.3 mm long, extrorse. Pistillate flowers with pilose - ca. 2 mm long; calw lobes 5. lau- eeolat.-. acute, ca. 3 mm long; ovarv densely ln-- »uri Bot. Gard. 71:341. |)idulou- with stinging hairs, the styles free to the PAX, F. & K. HOFFMANN. Euphorbiaceae — Dale- base, spreading, papillate, ca. 2 nun long. Mature cliauipicae. ///. A. En litor), Das Pflanzenreich

\\'rust i u. (J. I.. & B. WKHSTKR. 1 972. The morphology In aspect Tragia correae resembles the species phorbiaceae). Amer. J. Bot. 59: 573-586. that Pax & Hoffmann (I<)1<)) placed in section Hut, particularly such species as '/'. fcna'lcri. T. As a result of additional collecting in Panama, japnrensts Muell. \rg.. and '/'. flillux Muell. Arg. the species of Ihilcchampi a are m,w , on-iderabl\ The new -pe, le- i- e\«ln,|e,| Iroin that affinity, better unilerstood. and the number of species has however, by its racemose rather than bifurcate laminate Mowers with three (vs. key. The order of species has been modified to ens. The entire sepals and extrorse reflect better then -\-lemalic relationship- ( \\ eb ate its placement in sect Tragia unpubl. synopsis).

Stem tips and inflorescence- den.-el\ golden-lin - < oil MI tmvolueral bracts narrowly spathulate and apically mate involucel free) 1. D. sha,

iaJ] 1-5.

apically lacerate; leaves unlobed (occasionally with 1 or 2 small lateral teeth); involucral brae creamy or pinkish, marginally lacerate; seeds more or less rugulose. 4a. asymmetric, slightly dilated, not over 1.2 mm across; involucral bracts creamy . white with greenish veins; hairs of stem spreading at least in part; leaves persistenth pube-cc beneath, attenuate-acuminate, basal sinus narrow or lobes overlapping 2. D. canescens subsp. friedricl 4b. Stigma peltate, 1-3.5 mm across; involucral bracts pink or purple to white with pink veil (rarely white with greenish veins); hairs of stem appressed; leaves glabrate beneath, cuspidal with broad open sinus 3. I). diosco, Volume 75, Number 3

EE.

23.1. Dalechampia shankii (A. Molma) lh.lt. queron, 6-8 km upstream from Peluca Hydro Station, Ann. Missouri Bot. Card. 71: 341. 1984. Siri Km,, (DAY), SAN ISI AS: Puerto Obaldia, sea level, Knapp & Mallet 4627 (DAV, MO). Tragia shankii A. Molina, Ceiba 11: 68. 1965. TYPE: Costa Rica. Limon: Rio Reventazon, 15 It now appears that Dalechampia friedrichs- m, 23 Oct. 1951, Shank & Molina 4427 thalii is excessively close morphologically to I). (holotype, F). th (Nov. Gen. Sp. 2: 98. 1817). rom San Bias in particular seems This species, mil de : mbed from Costa Rica, has now been recorded from Nicaragua to biologically to treat the tw Colombia. It may easily be distinguished from our replacing subspecies of a : ngle species. The two other Panamanian species b\ it.- large and diminu- subspecies may be keyed ( tive golden hairs, as well as yellowish, narrowly ii n i\ oh HI I lie free bracts of the staminate involucel and the lacerate sta- minate bractlets indicate that the species belongs

Leaf blades mo-tl- ; i i\W;:--|.-i (N. \ri,il,i(!-i i-. iii.-i;.;. long with the two pubescent beneath (many haii lollo in I'anai HI I \;i I In i ollt i tioiis ol Bai - long, overlapping in sinuses b ry Hammel show that D. shankii is polymorphic in leaf shape, since leaf blades from I he single The Colombian taxon, subsp. canescens, was locality vary from unlobed to having one lateral described from Tolima Province (Mariquita) and lobe to three-lobed. Additional collections from Cos- cited from Narino by Pax & Hoffmann (1919: 52). ta Rica, Panama, and Colombia are cited by Huft The additional collections cited below indicate that subsp. canescens is widely distributed in the low- lands and foothills of the western Andean region 16.7 km N of turnoff to Coclesito from Llano Grande, 700 ft., Hammel 1811, 1812, 1813 (MO); 12 mi. from Llano Grande, 200 m, Churchill rt al. 4148 (F, MO). Sfx-cimrris cXtituilied C Mill \. • \l.l Yeguas, 20 km N of Honda, MH) m, Cent, i (DAV, MO), CAUCA: Rio Patia, 590 m, 23.2. Dalechampia canescens Kunth subsp. Vaughan 5354 (DAV). SANT\M>KK: 29 k friedrichsthalii (Muell. Arg.) Webster & Huft, stat. mi' Muell. Arg., Flora 55: 45. 1872. TYPE: Nic- aragua. Rio San Juan: Friedrichslhal 683 23.3. Dalechampia dioscoreifolia IWppig (not seen; locality erroneously cited by Mueller in Poeppig & Endl., Nov. Gen. Sp. PL 3: 20. as Guatemala). 1841. TYPE: Peru. Maynas: Poeppig 2163

Several additional collections of this plant have One additional locality merits noting: l'\\\\l\. DARIEN: near Rio Canglon, Duke & Bristan 378

d. ml • ' in . * nnli - A Smith 11,0 (MO). (MO); Rio Fato, Pittier 3866 (GH, NY, US); Rio Bo- An unusual specimen from Playon Chico, San Mas(r,Y///M (>3(>5. MO) appears lo be mt 5: 299, tab. 8. 1784. between D. dioscoreifoiia and D. canesc if i ,!n, 11.i-, iiii; the broad stigi former and the pubesc< Subfamily III. CROIONOIDKAK Pax leaves of the latter. 26. Tt'trorchidium

23.4. Dalechampia websi I < iroi i Indium I ' . I Syst. Bot. 9: 272. 1984. TYPE: Costa Rica. Nov. Sp. PI. 3: 23, tab. 227. 1841. TYPE: Heredia: La Selva, 3 km SE of Puerto Viejo, Armbruster & Herzig 79-207 (DAV).

This species, recently described from Costa Rica, has been identified from Panama on the basis of ew province records \ the single record that was attributed (with doubt) to I) rissifolia in our treatment of 1968.

.5. Dalechampia cissifolia Poeppig subsp. with persistent swollen stipules; panamensis (Pax & K. Hoffm.) Webster, less than 7 cm long, the paired glands I the apex of the petiole Ann. Missouri Bot. Card. 54: 193. 1967. I), 1. T. microphyllui panamensis Pax & K. Hoffm., Pflanzenreich IV. 147. XII (Heft 68): 19. 1919. SYNTYPES: Costa Rica: Tonduz 8089, Guatemala: Cub- ilgiiitz, Tuerckheim II. 211, 7978, Mexico. 2. T. costar Chiapas: Escuintla, Donnell Smith 2079. Stems smooth, the apex appressed-pubescent o Panama: Oersted; Pittier 2311, 3775. glabrescent; leaves mosth more ilian , i m lone

This species still requires ,u study- A long. variant with simple, unlobed lea 3a. Basal foliar glands ea. ()..r> ().(. mm tin. k. c n:ii|M»uiid i>nrs, lo wliirli I In- name / .hi irriuan pm heteromorpha Pax & K. Hoffm. has ben n: IS. moslb 1T> cm long or more, with o or 7 occurs in Panama am: ill*.- i-'i ml < Antral \mcrica sessile ' 3. T. euryphyllum but does not appear in be s(„-rib. a b di unci from lib. liasal foliar glands ca. 0.15 0.3 mm thick, attached at junction of lamina and petiole There is one new provincial record for D. cis-

li l.li \. II pi III! It! I .... I . lit, i I Additional specimen examined. PANAMA. BOCAS DEL pedicels mostly 1.5-2 mm long TOKO: 10 mi. NW of Almirante, D'Arey llL'dl I (MO).

24. .1. Tetrorchidium microphyllum Huft, sp. nov. TYPE: Panama. Chiriqui: 3.5 mi. NE of Boquete, end of road along Rio Alto, 6,200 ft., 18 Nov. 1978, Hammed 5721 (holotype, MO; isotype, E, F neg. 62357). Figure 2.

24.1. Omphalea , praeter apicem dense liirsntinn, ghilnis; loliis 1377. 1763. r. sumably BM, no

anches brittle, gla- 25. Vera ensely hirsute, ap- Pera Mutis, Kongl. Vetei f persistent, swollen Based on Hammel

Stipules, leaves short-petiolate, crowded near ends of branches; petioles 0.5-1.2(-1.5) cm long, densely appressed-pubescent with malpighiaceous hairs to glabrate, with massive, paired, thick-stalked, opposite or subopposite glands near the tip, these This distinctive new species of Tetrorchidium 0.6-0.8 mm long, 0.8-1 mm thick, stipules glan- appears to be most closely related to T. brevifolium duliform, tumid, broadly triangular, 1.5-2 mm long, Standley & Steyerm., described from the province densely pubescent, persistent, glabrate soon after of Alta Verapaz, Guatemala (Publ. Field Mus. Nat. leaf-fall; blades chartaceous, oblanceolate, acumi- Hist., Bot. Ser. 23: 126. 1944), from which it nate at tip, cuneate at base, (2.5)4 5.5(-7) cm differs by the smaller leaves (7-12 cm long in T. long, (0.9-1)1.2-2 cm broad, sparsely pubescent brevifolium), densely pubescent apices and with malpighiaceous hairs to glabrate on both sides, inflorescences (both glabrous in T. Inc., 'hum) the midrib and primary veins (3-4 on a side) prom- and shorter inflorescences (4-7 cm long in T. brevi- inently raised below, the veinlets forming a prom- folium). The type specimen of the Guatemalan inent reticulum; margins entire. Inflorescences ax- species (Rubelpec, Finca Seamay, Wilson 188, F) lacks the persistent tumid stipules that are so char- long, the rachis densely strigose; pistillate inflores- acteristic of T. n > ill these are pres- cences unknown s / s ^ubsessile; ca- ent on several collections of the former species lyx lobes 3, triangular, glabrous, ca. 1-1.2 mm made in 1974 and 1975 from Baja Verapaz, Gua- long; petals lacking; anthers subsessile, 0.8-0.9 temala (Lundell & Contreras 19173, 19436 (both F, LL); milifims et al. 13277, F). The type spec- I isiillati i accrues to 5 cm long. Stanunatc lloiccis leaves and stamiiiiite inflorescences that appear to in glomerules ol 2 5. sessile; sepals 3, obovate. have been taken from rapidly growing long . cucullate, glabrous without, pilose within: anthers , described from subsessile, 1.3-1.5 mm broad. Pistillate flowers (loud forests in the inoimtains above San Juancilo. subsessile. the pedicels heeomiug 0.5-3 mm long Honduras (Fieldiana, Bot. 29: 348. 1961, based in fruit; sepals broadly ovate, obtuse, 3 3.5 mm on Williams & Molina 170()8, F), is similar in all long, glabrous or sparingly short-strigose without, respects to T. brevifolium and should be relegated densely hispid toward base within: disk segment- to the syiionvmv of that species. A pa rat \ pe spet - free, narrowly ligulate, ca. 2 mm long; ovary imen of T. molinae from the same area. U illiams smooth, 2-locular, glabrous above. den-civ long & Molina 13980 (F), has long shoots without slngo e below: st\le eap at maluntv 0.5 0.8 mm stipules attached to a normal -hoot with persi-tent high, 1.5-1.8 mm diam. Capsule 3-5 mm high, stipule-. I hi- inalche- the pattern of the type col- lection of T. brcri/olii/rii. thus . onfirming the sus- ovoid-lenticular. 5-6 mm long, proniineniK and pi. ion voiced above concerning the nalme of that coarselv reticulate, the caruncle an irregular vellow papery keel running halfwav from the Inluni to ihe

Additional specimen examined. PANAMA. CHIRiyni: end of road past Palo Alto NE of Boquete in forest along Known only from Costa Rica and extreme west- ridge, 0.21)0 (,.«()() It.. H„mmci(,t>:iO(Y, MO). ern Panama, this distinctive species is easily dis- tinguished by the large, dark green, lanceolate 26.2. Tetrorchidium costaricense Huft, sp. leaves that are conspicu.»usl\ venose, the paired nov. TYPE: Costa Rica. Puntarenas: Cordillera glands near the middle of the rather long petioles. de Tilaran, Monteverde Reserve, near Con- and the densely and minutely strigose braiichlel tinental Divide on Pacific side, 1,520 1,580 tips, petioles, and leave-. m, Dryer 1403 (holotype, CR, F neg. 62351; isotypes, F, F neg. 62350, MO, F neg. 62349). of species with free ligulate disk segments in the pistillate flower that include- 7 lotundulitm Stand ley and T. brevifolium Standley & Steyerni. in northern Central \iii<-rica. lake I casta/icc/ise. T. rolundifolium has paired glands near the middle ii brevipedicellatis, segmenti.- disci .1 of long petioles, but differs in its completely gla- brous stems and leaves, mostly unbranched sta- iinnate tin rses thai have larger glomerules. densely Dioecious tree to 16 m; branches densely stri- pubescent pistillate calve.--, and di-tuicil\ pedicel gose tow.iid tip. appealing kuolili\ from the raised late fruits. Tetrorchidium brevifolium differs in its leal s, .ii- and oo, asionallv from persi-tent. indili glabrous stems, leaves, and calyces, short petioles ate stipules. I.raves long-petiolate, not crowded with paired glands near the tip. and unbranched toward ends of branches; petioles 2.5 (> cm long, staminate thyrses. The widespread South \inerican glabrous or minutely strigose, with paired, -ubop species T. rubrivenium Poeppig also belongs to this posite, sessile, patelliform glands near the middle. group but has idahroiis -|, rns ,\w>\ leaves, crenate these ca. 1 mm (rarely to 2.2 mm) diam.; stipules or denticulate leaf margins, sometimes lone petiole- oblong. 1.5 2.5 mm long, 1 1.5 mm broad, dense wilh the paired gland- near the lip. densely pu- ly strigose, persistent, sometimes indurate aftei bescent pistillate calyces, uniformly puberulent leaf fall; blades membranous, narrowly oblong, 8 ovary and capsule, and pedicellate fruits. 16 cm long, 3-7 cm broad, 2.2-3.3 times as long as broad, abruptK cuspidate al lip with an acumen M vll 1,1 v. Cordillera de Tilaran, Monteverde IvY-erve. 5-10 mm long, acute to attenuate a I base. inmiitelv Ulaiiiie -ide. 1,500-1,580 m, Dryer 1071 (CR, Y). puberulent below will) -cattep-,| short malpighia- cvmvco: Reserva de Tapanti', 1,300 I ,800 ,„. <„,„„•. 1877*2 (F). ITMVHKWS: Cordillera de Tilaran. Monte- v erde Re-erv e. Pacific side, en orilla de Pantano Chomogo, ate. prominent below, obscure above; 1,600-1,620 m, Dryer 659 (CR), 887 (CR. 10. M.»,, teverde Reserve, at field station, 1,500 in, l/aher 49 I tire, .-glandular, Inflorescences axillary, (10: Monteverde lie-ene. 1,570 til. //,//.,•/ S. Hell., InlU •nsel\ slrigo-e with -hort malpighiaceous if), 1.500 ,„, Uaber & Bella 2457(F). ALAJI KI \ ITVI v- minate thyrses 1 9 cm long, freely RKNAS: on and near the Continental Divide, ca. 2-5 km Webster & Huft

K .in.i si: ,,I \I i. i i • 'i ,., . i /1. 1.7(1(1 in. liu/fier cv Cent, v !U,()!!(¥): Monteverde, Dryer 1/3 1 |K). S\N JOSE: bajo de La Hondura, Poveda 862 iCR. I Ml. P\W\I\. iso, \s hi i rORO: along Continental I >i\ ulc. t rail to headwaters of Rio Mali, to W of Oleoducto Road, 8°47'N, 82°13'W, 1,200 m, Churchill 5276 (K); nt • Rita Ri \% Roa< I I an I ransisthmian Hwy., Kortuna Dam region, along Continental Divide W of high- Dwver X Gentry 9338 (MO, 2 sheets); Santa Rita Ridge way pass, ca. 8°45'N, 82°15'W, ca. 1,200 m, McPherson Road 4 mi. from Transisthmian Hwy. to Agua Clara 9695 (K). weather station, ca. 500 m. Gentry el al. 8841 (MO, 2

300-700 ft., Hammel 4894 (MO). PANAMA: El Llano- 26.3. Tetrorchidium euryphyllum Stand- av. 17-20 km N of El Llano, Dressier 4629 ley, Publ. Field Columbian Mus., Bot. Ser. 4: (K, MO); 10 km N of Margarita on road to Madrono, 219. 1929. TYPE: Panama. Bocas del Toro: vicinity of Almirante, 1928, Cooper 621 (ho- (MO); Cerro Jefe region 2.5 mi! N of'tur tower along road, 2,400 ft., Hammel (,300 (MO); Kl lotype, F). Llano-Carti road, 9.6-11 km from Inter-American Hwy., When the original treatmenl was written, this 350 i„. Man * Kallunh 3531 (MO): 5 10 km NKof Altos de Pacora on trail at end of road, 700 800 m. >|icnrs was known only from Costa Rica and ex- Mori & Kallunki 6058 (MO); El Llano-Carti road, 8 km treme western Panama, hut recent coll i n N of Pan Am. Hwy. at El Llano, ca. 450 m. Ace & Warmbrodt 10391 (MO).

27. Manihot

[ono: between Quebrada and Buena Vista, Klrkbride & Manihot Miller, Card. Diet. abr. ed. 4. 1754. Duke i)i,2 (MOl. < HiHloi i: Koriiina Dam region, along TYPE: Maniliol esciilcntu Crantz (Jatropha Quebrada Arena, ca. 8CI5'\. 82°I5'\V, ca. 1,100 m, n 8 194 (K). cot 11.: slopes of Cerro Pil6n near manihot L.). El Valle, 700-900 m, Duke 12196 (MO, 2 sheets); Cerro Pilon. Dwver 8330 (MO); La Mesa, 8.5 mi. from Club Campestre (El Valle), Dwyer 10515 (MO); Margarita near chicken farm, Dwyer & Duke 8280 (MO); La Mesa, above El Valle, Dwver & AW 11938 (MO); La Mesa, 4 km N of El Valle," 850-875 m, Xee * Dwver 9212 (MO). I) Mill-\: Cerro Sapo, ca. 2,5(111 ft., Hammel 1240 metric methods used. Econ. Bot. 27: 1-112 (MO). s-\\ m AS: Cerro Brewster, 9°18'N, 79°16'W, 850 m, de Xevers el al. 5408(F). \KK\.a AS: Caribbean slope The discovery of an additional I'amiir above Rio Primero Brazo, 5 mi. NW of Santa Ke, 700- 1,200 m, Croat 23233 (MO); NW of Santa Pe, 4.2 km hum I.-emit Vu.i i< oi; \1|. , ,- I'M ;|,;1 »/,,,, ,v A ,-,•//„.,•/,,. 4831 (MO); ca. 2.7 km hum I•:,< uela Agricola Alto de , 6208 (MO); ', ia. weaves giouruus, muouj wim .-* IUUCS, nta Ke on road past agricultural school, 2. ' «•• glabrous within (12-14 mm long; disk e 11216 (MO).

26.4. Tetrorchidium gorgonae Croizat subsp. Leaves pubescent or v robledoanum (Cnatrec.) Webster, Ann. 2a. Leaves pubescent, Missouri Bot. Card. 54: 199. 1967. T. roble- doanum Cuatrec., Brittonia 9: 81. 1957. TYPE: Colombia. Antioquia: 23 Jan. 1947, Gutierrez 35556 (holotype, CAL, not seen).

This species is still unknown in Central America outside of Central Panama. Several recent collec- 3. M. brachyloba tions allow a description of the fruit to be made for the first time. They are on densely strigose 27.1. Manihot aesculifolia (Kunth) Pohl, PL pedicels 7-10 mm long and jointed below the mid- Bras. Icon. Descr. 1: 55. 1827. Janipha aes- dle. The capsule is green, drying to brown, globose, culifolia Kunth, Nov. Gen. Sp. 2: 85, tab. shallowly 3-lobed, rugulose, 4-6 mm high, 4.5-7 109. 1817. TYPE: Mexico. Campeche: Hum- boldt & Bonpland (P, not seen). short (0.2-0.6 mm) malpighiaceous hairs. The 3 styles are deeply bifid, 0.6-0.7 mm long, and tu- This plant was called XL gualanensis Blake in mid. The seeds are ovoid. 1 5 nun long, 3-4 mm 28.1b. Cnidoscolus urens .subsp. adenophi- t monograph of the genus hy Rogers & Appan lus (Pax & K. Hoffm.) Breckon, stat. nov. Jatropha adenophila Pax & K. Hoffm., Pflanzenreich IV. 147. VIRHeft 63): 409. 27.3. Manihot brachyloba Muell. Arg., Fl. 1914. Cnidoscolus adenophilus (Pax & K. Brasil 11(2): 451. 1874; Rogers & Appan, Hoffm.) Pax & K. Hoffm., Nat. Pflanzenfam. Fl. Neotrop. 13: 190-192. 1973. TYPE: Bra- ed. 2, 19c: 166. 1931. TYPE: Panama. Pa- zil. Para: Marlins (syntype, G; microfiche nama: Chepo, Pit tier 1740 (isotype, US).

i specimens have flores Locks, Stem et al. 81 (MO): Pipeline Road. ( mat 12732 (MO); Curundu, McDaniel 5180 (MOl. /,>..« vegetatively matt !<>!., (MO); l|. \„ui,io, Wands. Issnn 5113 (MO). i given by Rogers i.viiifv Kl Real. I.a.o, A (one, 3.U, I (MO), v,-,-, ,t al. 454 (MO), LOSSANTOK: 5 mi. NW of Guararc, // dhui et al. 120,1 (MO); Monagre Reach. Lewis et al. 1673 (MO), PWWIA: Jenine, Rio Canita, Duke 3821 (M7 I i ,,1, i . I i i (MO); Puente de Pacora, De floyos 18 (MO), \hK\..i\s: f Rio Tuqueza, between Quebrado Venado and Peje 2-4 mi. E of Santiago. I hike 123., 1 (MO); 12 km E of wamp, Bristan 1001 (DAV, MO); between Manene and Santiago, Dwyer & Kirkbride 7450 (MO). 'joCoasi, Hartman 12127 (MO). 28.2. Cnidoscolus aconitifolius (Miller) I. M. Johnston, Contr. Cray Herb. 68: 86. 1923. subsp. aconitifolius. jatropha aconitifolius Cnidoscolus Pol,I. I'l. Ihusil. bun. Descr. 1: 56. Miller, Card. Diet. ed. 8. 1768. TYPE: Herb. 1827. EECTOTYPE: Cnidoscolus hamosus Pohl ' :, »' ' i " rsimiahK at I>\1. 11< • I seen). (chosen by Small in Britton & Brown, Illust. Fl. N. U.S. ed. 2, 2: 462. 1913). In the present interpretation, the Panamanian .ii.i •> lis all belong 28.1. Cnidoscolus urens (L.) Arthur, Tor lo the nominate subspecies, winch is not native to reya 21: 11. 1921. Jatrupha urens L., Sp. PI. 1007. 1753. TYPE: "America calidiori, in Brasilia & c." (not seen, possibly in Hortus 29. Jatropha I lim.rli i ii- Herbarium, BM). L., Sp. PI. 1006. 1753. I.ECTOTYPE:

Since the treatment of I"',., h, lr Jatropha gossypiifolia L. (see McVaugh, e Panamanian specimens of ('.. uren leads e conclusion that the two variants discussed there ?rit taxonomic recognition. Pending a more de- iled revision of the C. urens complex (Breckon,

. Bot. 09: 128.'{ I 2«>:>. . 1984. Phylogenetic significance ol inlerspe- ciiic Inhndi/ation in J,ilro/>li,i (Kuphorhiai cae). SvM. 28.1a. Cnidoscolus , Bot. 9: 467 478. & G. L. WKHSTKR. 1979. Morphology and mlrayencnc relalioiiships ol the genus Jain,/,ha (Eu- Specimens examined. PANAMA. PANAMA: 1 phorbiaceae). Univ. Calif. Publ. Bot. 74: 1-73. Beach area, Correa et al. 1586, Dwyer 3065, cVAl-CH, R. 1944. The genus Cnidoscolus: gencne 1803 (MO). coa.F.: between Aguadulce and San t 1 Woodson et al. 1226a (MO); between Anton and 71: I .: 174. D'Arcy & Croat 4117 (MO); banks of Rio Grande, , et al. 1157 (MO), HERRKRA: Chitre to Divisa, Bu, al. 1357 (MO); Sal Salinas de Chitre, Croat 9692 PANAM\: San Carlos, de McPherson 11 (MO). Pausandra Radlk.. Flora 53: 92, tab. 2 TYPE: I'alisiintha nioiisiunn (Casar.) (Thouinia morisiana Casar.). Webster & Huft

nate, simple; petiole swollen distally; blades pin- r.s horizontally aligned; petiole 1.5- nately veined, biglandular at base, the margins 3.5 cm long, 2-4 mm thick, terete, strigose with serrate. Inflorescences axillarv. spiciform; stami- short (to 0.5 mm long) malpighiaceous hairs; glands nate flowers in glomi I ules ; pistillate flowers solitary at apex of petiole (3 )4, cylindrical, 1-1.5 mm at each node; bracts inconspicuous, eglandular. Staminate flou.t- -ul>~.—ilr: . al\x lobes 5, im- • I i iilate, rounded, obtuse, or abruptly short- bricate; petals 5 or rarely 6, connate (at least cuspidate at tip, long-attenuate at base, 20-50 cm below), adaxially villous; disk extrastaminal, ur- long, 7-18 cm broad, 2.6 4 times as long as broad, celoate, lobate, glabrous; stamens (3-)5-7; fila- ments free; anlli.

This primarily South American genus is here disk cupulate. Pistillate flowers not seen. Capsule reported from Panama for the first time. smooth, strigose with short malpighiaceous hairs, apparently subglobose, ca. 1 mm diam. (fragments

BAIIION, H. 1873. Nouvelles observations sur les Eu- phorbiacees. Adansonia 11: 72-138. LANJOUW, J. 1936. The genus Pausandra Radlk. Re- 9 mm long, c cueil Trav. Bot. Neerl. 33: 758-769.

30.1. Pausandra trianae Baillon, Adansonia I I OIK:! i ra-. i 11: 92. 1873, proposed without reference to f'o^oiK'fihon! tf:a>:ir \ii:•."!•.. Arg. TYPE: Co- The description is based on the Panamanian lombia: Bogota, pla ol San Martin, RioMeta, specimens cited below and supplemented by col- Triana 2597 (holotype, P, not seen; isotypes, lections at F from Costa Rica and Nicaragua. G, not seen, photo F neg. 24574, K, not seen, The genus Pausandra remains poorly under- holotype of Pogonophora trianae Muell. Arg.). stood. The most recent revision (Lanjouw, 1936) was based upon only 33 collections and resulted Pogonophora trianae Muell. Arg.. Flora 47: 434. 1864. in the recognition el rune >pccie>. most known from TYPE: Colombia. Bogota: plains of San Martin, Rio only a single sex. Lanjouw admitted that several Meta, Triana 2597 (holotype, K, not seen, isotypes, G, not seen, photo F neg. 24574, P, not seen, of the species might need to be united as more holotype of Pausandra trianae Baillon). material became available. Pausandra trianae dor- lulosa Pax & K. Hoffm., Pflan- appear, however, to be one of the better-delimited zenreich IV. 147. XIV(Heft 68): 43. 1919. TYPE: species in the genus and is certainly the most Brazil: Rio Acre, Seringal S. Francisco, I /<• 9.5.W (holotype, B, not seen, photo F neg. 5406). widespread. The Central American plants readily ausandra e indley, Trop. I " key to that species in Lanjouw's revision, where 1929; Publ. h I. f ,.!.„„ ,i.„, \lu , But S.-r. P. extorris and P. quadriglandulosa are reduced Oct. 1929. rvi'i;: Nicaragua: Bragman's Bluff, En- to synonymy. The identity of C.lavija septentrio- glesing 216 (holotype, F, F neg. 52719; sam- nalis as P. trianae was first pointed out by R. L. ple, Y no. 13301). Imtrionalis L. (). Williams, Fieldiana. B;>1. Liesner of the Missouri Botanical Garden (pers. 32: 205. 1970. TYPE: Nicaragua: Cabo Gracias a Dios, Laimos Creek, ca. 15 km SW of Waspam, 7 The for Pausandra trianae lias Mar. 1961, Bunting cv Lieht 390 (holotype, F; been almost universally given for the last century isotypes, NY, US). as Pausandra trianae (Muell. Arg.) Baillon, car- rying the implication that Baillon had transferred 11 - an trianae Muell. Arg. to its correct berulent with malpighia place in Pausandra. It is (dear from Baillon"s text. :U) 100 in. (If \rrns A II, una ,(., I | I ). along uruK cut road from El Llano to Carti-Tupile, near Continental Divide. ."»00 .',00 in. I.irsnrr 12H<) (DAY, F, MO), \U,n cv Kallunki 5535 (MO); seasonal low- Mueller's name, land rainforest on the Aila Tilar (Hi.. Via), iTIH'Y and none would he c\pe. led. One would not nor- 77°40'30"W, 25 KM) ,„. Sullen 121. 5fiti [Mi)). mally look in I'oiiuiiojihoKi lor species pertaining to Pausandra: the j;i'iicra air too different lor

The fact that the names of Mueller and of Bailie ire both based on the same collection, although c hfferent specimens, is merely a coincidence. Th 31.1. Garcia nutans Yahl in Hobr. turhist.-Selsk. 2: 217, lab. 9. 17< hut not \cr\ surprising, given the common practice Colombia. Magdalena: near Santa M ol iuiniiiiii species after llie colic, lor ol the l\pe. Hofn (C. not seen). Thus, although Baillon ideally should have made a transfer of Mueller's name, he was understand- ably unaware that Mueller had pre\ iousl\ described ihe spec ics at hand, and dnl nol in.ike the transfer.

the rules would mandate a transfer of Mueller's epithet to create a new combination in Pausandra Croton L., Sp. PI. 1004. 1753. LECTOTYPE: supplanting Baillon's name. But thai solution is ton aromatieus L. (chosen by Webste, closed because it would create a later homonym. Arnold Arbor. 48: 354. 1967). Article 63 of the Code (Voss, 1983) might mislead one to reject Baillon's name as superfluous, hrcnuse Seven species new to Panama (incl tiding lv it seems to be based on the type of a name whose new species and one new subspecies) an • report. epithet ought to have been adopted under the rules. here; a few new province records are in d.cated i A correct reading of the (lode, however, makes it well. These additions make it necessarv to provi. clear that a type is a specimen, and not a gathering, win. h usiialK consists of several duplicale speci- mens. An isotvpe has no olli. 1.11 standing as long as the holotype exists, although its value is un- questioned. Thus, for a name to be rejected as Webster & Huft

5a. Leaves more or 1<^ >M<>i lipi |>i I in I; mil i s mostly 2 cm long or longer; pedicels of |.i hi I « i- MM I ! i ,, lung or longer 1. C. schledea 5b. Leaves ovate, more or less cordate, (3-)5-veined at base; inflorescences 1 cm long or shorter; pedicels of pistillate flowers 1-2 mm long 2. C. pseudoniv Stamens 14-17, or if fewer, then filaments hirsutulous; anthers 1-1.2 mm long; petals of staminate flowers lepidote; seeds 15 mm long or longer.

• :"l» ::.: .nni long 3. C. tenu.

'Mi. Leaves with basalU

7a. Seeds 3-5 mm I - > • , ' ' 5. C. lanjouwensis 7b. Seeds 16-17 mm long; fruiting pedicels not over 7 mm long 6. ('.. /><:.

gland- at junction with bl lei pi m< in I - . inl • rescence (at least in part) terminal. 8a. Leaves deeply 3-5-lobed; stan I b distinct] imbl < ate in bud; staminate receptacle

8b. Leaves unlobed oi ball< I lobed (1< ban! n ); staminate calyx lobes mostly valvate in bud; staminate receptacle sparsely to densely villoa 5 eds not tetragonal. 9a. Stamens over 50; styles more than 5 mm long, twice bifid; stipules tomentose, dentate, 3-6 mm long; seeds ventrally ribbed 7. C. speciosus

10a. Lower cymules of inflorescence with both staminate and pistillate flowers at the same

11a. Styles bifid; pistillate calyx lobes not reduplicate-valvate; stamens 13 or more; filaments glabrous or nearly so; leaves unlobed, without scattered Uutninai : and 12a. Pistillate flowers distinctly pedicellate, the pedicels mostly 3-6 mm long;

12b. Pistillate flowers subsessile, the pedicels in fruit not over 2 mm long; inflorescences mostly 1-2 dm long; stamens 25-45; styles dis'in i) i ill ie pubescent; seeds at least 5 mm long 9. C. pun, Styles niultifid. .stellate-pubescent: pistillate calyx lobes reduplicate-valvate; sta- mens 11 or 12; filaments hirsutulous; leaves more or less 3-lobed, with scattered peltate laminar glands above 10. C. smlthu 'X nodes of bisexual inflorescences with solitary pistillate flowers (not with staminate rs at the same nodes), or inflorescences unisexual. Styles twice bifid to multifid. 14a. Shrubs or trees; pistillate calyx lobes not strongly unequal, the abaxial ones not deeply lacerate.

mostly 7-9-veined at base, entire: petioles moMlv 5 20 cm long 11. C. billbergianus 17b. Pistillate calyx lobe- ovate, reduplicate-valvate, accres- cent, stellate-tomentose within, becoming 7-8 mm long and broad; staminate petals 4.5-5 mm long; leaves most- ly 5-veined at base, entire to denticulate; petioles 1-4 cm long 12. C. fragrant Stamens 10-12; seeds smooth; staminate receptacle sparsely villose. 18a. Leaves mostly alternate (occasionally opposite at 1 or 2 distal nodes), 7-9-veined at base, the margins distinctly dentate (teeth 15-40 on a side), the basal laminar glands

i 3-12 pis-

glands long-stipitate; stipules a 15b. Pistillate bracts

Annual herbs; pistillat reduced or obsolete, 6-8 mm long

Styles once bifid; annual herbs. 19a. Leaves coarsely and sharply serrate (major teeth usually not over 10 per side); Miliums H 10; styles less than 1.5 mm long, spreading IT ( trimtaU* 1%. Leaves more finch and or bluntly toothed; stamens mostly 11 or 12; styles 1.5 mm long or more, ascending or erect; seeds minutely beaked. 20a. Stems coarsely hispid; leaves mostly ovate, pointed at the lip, petiolar glands stipitate; bracts with Ion I a ip] ! pi -esses .. 18. C. hirtus 20b. Stems not coarse! In \> >l ' I. , i. n oblong, blunt at tip, the petiolar glands sessile; bracts eglandular [C. glandulosus)

i Lundell, Phyto receptacle villose; petals obovate or narrowly ellip- logia 1: 451. 1940. TYPE: Costa Rica. Sai tic, ca. 2 mm Ion du n tali . detiscK Jose: vicinity of El General, Skutch 2575 (ho villose on margins, glabrous on back: ,, 12 lotype, MICH). or 13; filaments glabrous or sparsely hirMitulou.-; anthers elliptic, 0.6-0.7 mm long. Pistillate flow- ): along road to Chiriqui Grande, 10 road mi. fron ers with stout lepidote pedicels becoming 8 1 5 mm long; calyx lobes 5, equal, triangular, ± redupli- cate-valvate, lepidote without, ± stellal- • ..i. within on the recurved margins, 1.5 5 mm long: disk 5-lobed, adnate to calyx; petals rudimentary 32.5. Croton lanjouwensis Jabl., Mem. New (shorter than 1 mm long); ovary densely lepidote York Bot. Gard. 12: 158. 1965. C. matou- rensis Aublet var. benthamianus Muell. Arg., ca. 4 mm long, stellate proximallv (brandies gla Linnaea 34: 95. 1865. C. benthamianus lii. Capsules not seen entire; cocci lepidote (Muell. Arg.) Lanjouw, Euphorb. Surinam 17. ca. 5 mm long; columella slender, ca. 4 mm long; 1931, non C. benthamianus Muell. Arg., Fl. Bras. 11(2): 106. 1874. TYPE: Brazil: Rio 3.5 , Negro, Spruee Croton 2 (isotype, NY). The Panamanian plants appear to agree in most

Tree to ca. 12 m high; monoecious: twigs deiisdv respects with C. lanjouwensis as defined |,s | ;,n lepidote. Leans wild [.choirs lepidote, 1.5-3.5 cm jouw (1931: 12-17; as C. benthamianus); the long listal • ill II iiit • i.wish, subsessile, ii I i lillate calyx with adaxiallv stellate lobes patelliform glands ca. 1.5-2 mm across; stipules is apparently diagnostic in separating the species linear-lanceolate, entire, densely lepidote, 8-10 from C. matourensis Aublet. However, the Pan- mm long, early deciduous; Ida* i M.I i plants occurring in cloud forest at 800 liptic to elliptic-oblong, acute 1,000 m would appear to differ ecologically from at tip, cuneate at base, mostly 8-15 cm long, 3.5- the Amazonian plants, which have been colic, led 6.5 cm broad, s < il ml II. <>us above, densely in lowland rainforests. Jablonski (1965: 157-158) l< I .1 ;. beneath (scales denticulate-margined, ca. 0.25-0.4 mm across and nearly or quite contig- \ ei • , nd; . ,. Pan- uous); venation distinctly pinnate, the major lateral veins (ca. 11-15 on a side) straight; margin* entire needed. The specimens from Panama Pros in. e dd .ri.: If| dole margin, h- /;;//."- r.s, , nccs terminal. fer rather strikingly in their duplex petiolar glands, racemose, 13-15 cm long, In MI I; ( n ij. ! ». sparsely lepidote upper leaf surfaces, and shorter ers solitary at 3 6 proximal axils; staminate flowers in cymules of 2 or 3 at distal axils; bracts entire, lepidote, up to 3 mm long. Staminate flowers with lepidote pedicels 2 5 mm long; calyx lobes 5, val- vate, ovate-triangular, lepidote, 2-2.5 mm long; Valle, ca. 900 m, Duke & Correa 14718 (DAV, MO); glabrous; petals obsolete, represented by whitish cloud forest, hills N of El Valle de Ant6n, Dressier 4083 tufts of hairs; ovary densely lepidote (scales 0.5- (DAV, MO); cloud forest, El Valle, 800-1,000 m, Duke 13166 (DAV, MO), Dwyer & Correa 7951 (DAV, MO), 0.8 mm across, denticulate, 50 70-radiate, with Correa 311 (MO). PANAMA: primary forest, road from El 20-30 darkened radii); styles blackish, twice-bifid, Llano to Carti-Tupile, 200-500 m, Croat 22905 (DAV, 3.5-4 mm long, nearly glabn . Ca\ ."•'• D I een MO). entire; valves of cocci ca. 23-25 mm long; colu- mella slender, ca. 20 mm long; seeds elliptic, some- 32.6. Croton pachypodus Webster, sp. nov. what compressed, flattened and obscurely carinate TYPE: Panama. San Bias: km 18 of El Llano- on the back, distinctly keeled on inner face, grayish Carti road, 9°19'N, 78°55'W, 350 m, 1 Oct. brown, smooth and shining, 16.2-17.2 mm long, 1984, de Severs & Herrera 3980 (holotype, 10.3-11.3 mm si . MO; isotypes, DAV, F). obscure, tenuous, ca. 2.5 mm long, 1.5 mm broad. Collections of this species have been determined as C. lanjouwensis, to which indeed it is related and superficially very similar. However, it is dis- tinguished by leaves very spa i eath Tree 7-25 m high; twigs obtusely angular, gla- and with margins free of scales, shorter petioles brate. Leaves with petioles lepidote, 0.8-1.5 cm with smaller darker glands, and especially by the long, distally (near base of blade) with 2 subsessile much larger fruits borne on greatly thickened ped- short-stipitate (to 0.7 mm) blackish patelliform icels. Among species earlier reported Iron I'ai a ma. rids 0.5 the new species resembles C. tcnuicaudatiis; how- ' lepidote, 4-10 mm long, 0.9-1.2 ever, that species has eglandular leaves lepidote ; blades chart; on both faces, strongly lepidote staminate petals, tic-oblong to somewhat obovate, subacute or acute and more slender pistillate pedicels. Croton pyri- to abruptly short-acuminate at tip, cuneate to rounded at base, 8-22 cm long, 3.5-7 cm broad, ovate palmately veined eglandular leaves, longer smooth and glabrate above (with sparse scales on and more slender (1.5 mm or thinner) pistillate l young), evenly and sparsely lep- pedicels, and verruculose capsules; however, the i (scales 0.25- seeds, although larger, are similar in shape to those ca. 50 radii), the scales widely sep- of C. pachypodus. on disiineiK pinnule, the major laleral >n a side) straight or slightly curving, 12 mi. above Pan-American Hwy., 200-500 r 22905 (MO); 20.7 km from Pan-American Hv m, Mori & Kallunki 5116 (MO).

32.7. Croton speciosus Muell. Arg. [Linnaea 34: 83. 1865] subsp. tacarcunensis Web- inflorescences, staminate flowers solitary or paired ster, subsp. nov. TYPK: Panama. Darien: Cerro at distal axils; bracts triangular, blackish, sparsely Tacarcuna, S slope, premontane wet forest on lepidote, 0.5-1 mm long. Si, | ud ridge below summit, 1,250-1,450 m, Gentry only observed) with lepidote pedicels 1.5-2.5 mm & Mori 13925 (holotype, MO; isotype, DAV). long; calyx lobes 5, valvate, triangular, lepidote, ca. 3 mm long; receptacle villose; petals narrowly Haec a subsp. s pecioso differt stipulis r.,m..M|,i-, _4m dulis petioli brevioj ribus, carunculo seminis ca. 2 mm lato. elliptic, ca. 2.5 mm long, densely villose on mar- gins, sparsely lepidote (often a single scale) on the Monoecious tree 5 m high; t\ vigs subterete, back, densely hirsutulous adaxially; stamens 14- densely tawny-v illose with dendritic : hairs. Leaves 16, the filaments glabrous, the anthers 0.8-0.9 with petioles (2 -)3-ll cm long, t omentose, api- mm long. Pistillate flowers with stout lepidote ped- cally with 4-6 i stalked glands on v< •nlral side, the icels ca. 2.5-3.5 mm long, becoming in fruit 4.2- 6.5 mm long, 3.2 4 mm broad; calyx lobes 5, stipules lanceolate, densely tomento se, toothed, 3- equal, triangular-ovate, reduplicate-valvate, dense- 6 mm long; bl; ades membranous, mostly ovate, ly lepidote without, densely loin a IOM \illosc with- long-acuminate at tip, rounded to subcordate at in, 3-3.5 mm long, 2.5-3 mm broad; disk shal- base, the larger ones shallowly 3-lobed, 12-21 cm lowly 5-lobed, nearly 4 mm across, smooth and long, 7-12 cm 1broad; lamina abov< i copiously pu- ellate-tufted hairs, beneath copiously Mueller ga\e the -tnmcii number

subulate stipules ca. 3 5 mm long. Starninute Jlou - tervening collections mas close the gap. However, ers with stellate-tomeutose pedicels 5 8 mm long; the Panamanian plants differ strikingly in their receptacle densely tomeiitose; calyx lobes 5, fill- much smaller and less lacerate stipules, as well as

vous-tomentose. obtuse, entire. 3.5 5 nun long: m haying distmclK shorter -talked glands at the petals obovate, 4.5-5 mm long, densely appressed- apex of the petiole, furthermore, the caruncle in pubescent without, glabrous within, woolly-villose seeds from Panama is roundish and ca. 2 mm along margins: stamen- ca. 60 70; filaments slen- broad, whereas it is distinctly laterally expanded der, glabrous, ca. 4-5 mm long; anthers oblong, and ca. 3 mm broad in seeds from Venezuela, apiculate, 1.2-1.6 mm long, 0.5 0.7 mm broad. Proy isionally, therefore, it seems best to designate

I'istillatcJ/oiccis with -lout toiiienlose pedicels ca. the Cerro Tacarcuna plants as a distinct subspecies.

ns examined. PAr> t long in fruit); CJ ilyx segments 5, valvate, oblong, peak of Cerro Tacarcuna, 1,100- densely whitish - to fulvous-tomentose without, l.:!lHI,n. dent,, rial Hu'i7.' i M()): r..l«.-lo,. l.rkm Ml,, sparsely toment ose in distal third within, ca. 10 de Nique base camp, Gentry el al. 28727 (MO). mm long, 3-5 nrim broad; disk inconspicuous, ad- natetobaseofc; alyx, crenulate, stellate-pubescent. 32.8. Croton draco Cham. & Schldl. [Linnaea ca. 5 mm across ;; petals rudimentary, densely hir- 6: 360. 1831]subsp. panamensis (Klotzsch) sute, ca. 1-1.5 mm long; ovary densely fulvous- Webster, stat. nov. Cyclostigma panamense tomentose; style: s twice-bifid near the base, densely Klotzsch in Seem., Bot. Voy. Herald 105. stellate-pubescei it below (and with scattered stellate 1853. Croton panamensis (Klotzsch) Muell. hairs distallv ne; irly to tips), ca. 7 9 mm long, the Arg. in DC, Prodr. 15(2): 546. 1866. TYPE: Panama. Chiriqui: Volcan Cbiriqui, Seentann tips. Capsules si ibglobose, fulvous-hispidiiloiis. ca. (K, not seen), (see Webster & Burch, 1968: 1 cm long and broad; seeds plump, plumbeous 254 for additional synonymy.) brown, distinctly • costate ventrally, obscurely cos- tateonback,ca. 7 mm long, 5 mm broad; caruncle Further examination of Mexican and Central round, h low i. 2 mm across. American specimens of C.ioton draco indicate- lfi.it the Panamanian plant- cannot reasonably be main This striking Croton from the cloud forests at tained as a separate -pecie- Kxcept for the larger, the , rest of Cerro Tac.m appears o )e con- broader stipules (mostly 2 mm or more across), the specific with C. specosus. which was described Mexican populations here assigned to subsp. - with a single species. The stamen number in the Tree to 18 m high; twigs angled or sulcat Venezuelan plants varies from 40 to 80 and hence yellowish-scurfy with pedicellate stellate hair includes the number for the Cerro Tacarcuna plants. Leaves with petioles mostly 5 15 cm long; pate Volume 75, Number 3 Webster & Huft

; stipules lint •ar-lancn. laic to s ;:a:liul, h ng,l 2 mm ides mo stly ovate 3-lobed, blu nt to acu subcordate ait base, the larger ones 15 iri indumentum, and a specimen from < ihoeo 5, 10-30 cm broad; i palmate {Archer 2062, US) has the characteristic leaf form major vein s at base . 5-8 1, aterals 01 and margin, but the are not di.-tmelK pedicellate as in the Panamanian plants. The Pan- chotomizing towards t ts prominulous on bo Croton killlplanus Croizat, described from Bo- riform; trichomes on yaca; however, the type collection of that species .,.,11, 1:11 (Lawrance 588; isotype, US) he ibentir leaf IHLII t.cncatli denser and + floeeose: -mall palel- margins and an appressed, rather sparse indumen- hi: ii i idand- (0.1 I iniii arrows) occasional on tum more characteristic of C benthamianus Muell. upper surface; margins denticulate, with occasional Arg. Croton niintians Croizat from Guyana is small stalked glands. Inflorescences terminal, most- somewhat similar but differs in its smaller fruiting ly 20-50 cm long; 5-15 proximal eyniules hise\- calyx and shorter fruiting pedicel. Until this -pe, ie- late, entire, deciduous, ca. 1-2.5 mm long, to C. smlthianus. Several collections from Nica- subtending sewral (lowers. Slaiiiinalc flowers with ragua and Costa Rica are also referred to that -lellate lomentose pedicels 1.5 5 mm long; calyx distinctly gamopln lions. 3.5 1 mm long; calyx lobes Specimens examined. COSTA RICA. HEREDIA: Finca ovate, acute, valvate, 2.2 3.7 mm long, 2-2.8 La Selva, Hammel & Trainer 12849, 13044 (DAV, mm broad; petals narrowly spathulate, 3.2-4.5 III Kb), I-I M Mil \\>: forest remnants along highway 4 mm long. 0.5 1 mm broad, densely villose ven- mi. SE of turnoff to Buenos Aires, 9°07'N, 83°17'W, trally, strigose-hirsutulous dorsally; receptacle 1(10 in. II chster 21883 (DAV, MO); rainforest 17 mi. SE of San Isidro General, 700 m, Webster c\ Miller densely villose; stamens 11 or 12; filaments flat- 12394 (DAV). NiCARA',. 3 1.5 mm quistain 3223 (DAV, MO). ZEI.AYA: Bluefields, \, •,// 27,'tH long; anthers elliptic, the connective glandular-pus- (DAV, MO). PANAMA, COLON: Rio Salud, Howell 128. tulate, 1.1-1.5 mm long. Pistillate flowers with Lao & Holdndge 224 (MO). DARIEN: between Manene stellate-lepidote pedicels becoming 9-14 mm long; and Tusijuanda, Duke 13576 (DAV). PANAMA: SE slopes of Cerro Trinidad, Kirkbridc & Ihtkc 1665 (MO). calyx lobes 5. valvate (not distinctly reduplicate),

32.11. Croton billbergianus Muell. Arg., Lin- tip and along margins, 5-7.5 mm long, 3.5-6.5 naea 34: 98. 1865; subsp. billbe17.ia.ms. TYPE: Panama. Colon: Portobelo, Billberg 316 ovary yellowish-si I! r or stellate hispid, trichomes 0.5-1.5 mm across in fruit; styles free, multifid, sparsely to copiously stellate-hispid, ca. 5 mm long. Capsules subglobose. yellowish with appressed scales; columella ca. 1 mm long; seeds plump, Additional specimens examined. F \NA\I \. w >< \S DI-.I TORO: Siirsuba, Rio Changuinola, Divyer s.n (MO). -\N lenticular, brownish, finely costate-rugulose, blunt- itl,\s: Puerto Obaldia to ba lionga, knupp & Mallet 4667 ly pointed at both ends, 3.9-4.1 mm long, 3.3 (MO), VERACUAS: Coquyito mine to Rio Barrera, Hammel 3.5 mm broad; caruncle flat, bilobed, 1.7-2.1 mm 5221 (MO); Santa be. Folsnm X Eduards 3392 (MO).

The description in the original treatment (Web- Croton smit/ih ~ b, !< I i ,\ mil ster & Burch, 1968: 257-258) applies only to montane forests, up to ca. 1,500 m elevation, ib . her^ianiis \- suggested at that time, Nicaragua to Cob i- ii July to Septem- C. pyramidalis J. D. Smith, extending lrom Ve- racruz, Mexico, to Honduras, does not appear to The Panamanian representative of the wide- be a distinct species. It may be retained at the spread and variable South American species com- subspecific level because of its apparently larger plex centering on ( rot mi K b seeds (5.7-6.2 mm in the Veracruz population vs. i- heie re b Tree, n. < :>>••: I,,-;;,,, , « m/ai because 4.3-5.5 mm in If.. I ".man an •.,!• plan: • : a, id I- aiL'ei

Volume 75, Number 3 Webster&Huft 1123 1988 Panamanian Euphorbiaceae

stipules (7-15 mm long vs. 5-7 mm in the Pan- nerved at base, the secondary veins 5-8 per side; margin entire to remotely denticulate, Inflores- ican plant is therefore necessary.^ terminal, the pistillate ones axillary, occurring only 32.13. Croton hircinus Vent., Jard. Malmai-

cences 17-22 cm long, densely brown stellate- villous; nodes 15-30; flowers 1-3 at each node on stellately pubescent pedicels 6-9 mm long, the l additional provincial record is cii bracts subulate, 2-3 mm long, stellate below, gla- brous above; pistillate inflorescences 5-11 cm long, r obre hri.lpe, 2, . mi. W of Tole, Divyei densely brown-v ill I >• Ha ellate, deeply la- ciniate, 10-12(-14) mm long, at least the lower ones loosely end n llie II > I 32.14. Croton santaritensis Huft, sp. nov. flowers: sepals 5, debate, joined at base, valvate, TYPE: Panama. Colon: Santa Rita Ridge Road, stellately pubescent, the lobes 2.5 3 mm long, ca. 21-26 km from Transisthmian Highway, 2 mm broad; petals 5, only slightly exceeding the tropical wet forest, 500 550 m, 9°25'N, calyx lobes, ca. 6 mm long, ca. 2 mm broad below 79°37'W, 4 July 1982, Knapp 5882 (holo- the tip, the tip abruptly expanded, ca. 2.5 mm type, MO; isotypes, DAV, F, F neg. 62353, broad, coarsely erose; stamens 12-15; disk con- PMA). Figure 3. sisting of 5 nearly separate glands; receptacle gla- ne): sepals 5, del- , fleshy; ovary densely stellate-1 twice-divided. Mature frui and »

all other Cen

conspicuous fimbri it< I nl< ml the oblong, la- ciniate bracts that loosely enclose the young pis- Leaves -villi petioles ilenseK villous ; 5-20 mm long; petiolar glands sever; petiole, stipitate. tnimpet-sliaped. 0.5 Because of its combination of pentamerous calyces 0.3-0.5 mm across; stipules ovate-d< in both pistillate and staminate flowers, five petals branous, eglandular, appressed. 8-1 and glabrous receptacles in the staminate flowers, 4-6 mm broad, the margins fimbi iate; caudate at apex, rounded to subcorc ognized by Mueller (1866: 511-700) in 10-15 cm long, 4-8 cm broad, 1.7 recent worldwide account of Croton. N< long as broad, sparsely to moderatel) closely resembling Croton santaritensis bescent above, sparsely tomentose found among the large holdings of South /

Croton billbergianus s 32.15. Croton brevipes Pax, Bot. Jahrb. Syst. Smith) Webster, stat. nov. Croton pyn 33: 290. 1903. TYPE: Costa Rica. Rio del Smith, Bot Gaz. 35: 7. 1903. nn- <; Verapaz: Rio Dolores near Cubilguitz, Tw Convento, Pittier 12117 (isotype, US, photo, (holotype, US; not seen).

FIGURE 3. Croton santaritensis -i . , . • >ieseet„-es. — B. Detail of leaf

; petals, abaxial view on left, adaxiul on n^ht. //. Pistil' view.—I. Pistillate flow. i Knapp 5882. Illustration by Clare larger laminar glands; tin- stipules, bracts, and ca- pressed-slellate. Leaves alternate below, mostly op- lyx lobes lack the glandular serrations of C. hir- posite or teniate abo\e: petiole-. densef appreused stellate, 3-20(30) mm lone (less than '/ length dontiis Muell. Arg. from Mexico shows that it is of blade): petiolar glands (at base of blade) con- extremely close to ('.. hrevipes. Although -pecimen- spicuous. e\ lindrii al. apiealK Innicate and dilated. from Costa Rica and Panama may be easily rec- 1 2.5 mm long, 0.3 0.5 mm across; stipules subu- ognized by their distally opposite, less coarsely late to narrowly lanceolate, dark, entire, eglan- toothed lea'- e- with more rounded h.i-e- and -hortei petioles, they are \er\ similar to the \le\i. an pi.mi- ihinK chartaceous. elliptic to ovate-elliptic or ob- ni most details, including pubescence, floral details, and bulls. The seed- of the Mexican plants are at base, 4- I 3 em long, (1 )2 5 cm broad. sparsely somewhat larger, but this difference ma\ di appear stellate or appressed hispid abo\e with leu-rased upon further sampling. Provisionally, the two -p,-. i,-- trichomes. sparsely appressed-slellato and ineon- may be kept distinct on the basis of the foliar -ph nou-ly glandular-punctate beneath. pinnalely characters, and because no intermediate popula \ en ied (or inconspicuous! \ I riplmery ed) with mostly 5 7 veins on each side: margins subentne to rather i Costa lii.a . Mex- coarsely am! irregularly deiil.ite (teeth ca. 8-15 Specimens examined. I'wun. ininv Nu.la Kit.. on a >ide). with stalked viands between some of the Ridge, ca. 300 in, •tnloiim /.Y.i'MD W, f, MO). Co,tea teeth. In/loieseem e.s iu.r-.lK terminal and bisexual A Dressle, <) 12 (F, MO). Coat I Mi'lH (MO). Put,, /..."'/ (\K». I hi w7 ,->'.,/.•; | \IO), lh>,e, At.enlis ">'"-. :;< i.:,ie •eU :>,,|, If. Mm. haste, 177*1 (D\\, 1)1 Kf, f), Gentry 1X74 (1)\\. f. MO), ken ne,lYl>7.-,<> (WO), knnpp .',«•/.» A\ . pistillate flowers, the siaminale (lowers I or 2 per I. MO). SMsmu 20.17 (MO). 2()7>l (f, MO). II .•/,•,/,•/ A bract at distal axils: bracts narrow, entire, eglan- Ihes.le, 10727(DAV, MO, I S);Eas1 Ridge, Duke l.,2"l (DAV). r\\\\l\: Cerro .lef'c, 700 750 in, Dnsslei it! I I sliorler. Stnminate J/oicers with sparsely stellate (DAV, MO. I'S), Webster A Dressier 10477 (DAV, or nearly glabrous pedicels I 2 nun long: calyx DfKf. MO. IS): T..rt[ \rr ,1M, /• olsom et ui r,o •/••/ I l)A\ . MO). lobes 5. elliptic-lanceolate, acute, stellate-pubes- cent, glandular-punctate, 1.2 1.7 mm long, 0.9- 32.16. Croton argenteus I ., Sp. PI. 1004. I . I mm broad: recept.n le moderately \ illo-e; pelal- 1753. Juloerntim argenteus (L.) Didr., Vi- olxnalc-spathulale. 1.1 1.8 iiim long, glandular- densk. Meddel. Dansk Naturhist. Foren. Kjflbenhavn 1857(8 10): 134. 1857; Muell. mens 10-12; filaments glabrous, 1.8-2.5 mm long; Arg. in DC, Prodr. 1 5(2): 703. 1866; Croizat, anthers ovate, 0.5-0.7 mm long. Pistillate Jlouers Revista Argent. Agron. 10: 125. 1943; Cor- with -lout appressed -lellale pedicel- heeoining I .'.', rell & Johnston, Man. Vase. Plants Texas 3.5 mm long; calw lobe- 5. subequal, narrowly 939. 1970. TYI'K: America (not seen; presum- ably in Hortus Cliffortianus Herbarium, BM; 1140.8 in LINN). . 3.5 i long. . glab, Annual herb 2 10 dm high; stems pseudodi-

van sparsely stellate-pube-ccul apically. glabrous chotomizing. with long internodes ,md psendover- clow; stvles free. ca. 2 mm long, twice bifid. ticels of leaves, appressed stellate-pubei iilenl labrous to hispidulous. Capsules subglobo.-e. Leaves with petioles 1 5 cm long, these without parseK -lellate-pubescent or glabrescent, ca. 5 paired apical gland-: -dpule- subulate, (2.5 )5 10 mi diam.; columella slender, 3.2 4 mm long; seeds mm long; blades chartaceous, ovate or the upper roadly ellipsoid, compressed, apically beaked, ones oblong-o • or rounded to subacute rownish. nearly smooth (minutely striolate), 3.8- 7( I5)c ong, 2.5-5 cmi broad,b 5-veined at h mall, ca. 0.5-0.8 mm across. e green and finely appressed-stel- Rainforest below 1,000 m, Costa Rica and Pan- late. beneath grayish and more densely stellate; margins finely serrulate. Inflorescences terminal. bisexual, ca. 1-4 cm long; bracts subtending sol- itary flowers, the pistillate (lowers 4 6 at base of Webster & Huft

! U'L s nl i rls ca. 1.5-2.5 mm long; calyx lobes lanceolate, acute, valvate, ca. 1.5-2 mm long; petals linear, 2.1-2.3 mm long, 0.3-0.4 mm broad, glabrous except for the li MI < j nsl) \illose; sta- mens usually 11; filaments sparsely to rather co- |:i nun broad. !!•> i; laciniate, provided on each side with 5 10 33.1. Mabea teeth ca. 0.5-3 mm long, the 2 abaxial lobe- tnin-li Prodr. 15(2): 1151. 1866. TYl'K: Colombia: smaller, nearly or quite obsolete: pelals absent: disk Schlim 1132. Venezuela: Fendler 24. Pan- strongly asymmetrical, with larger adaxial lobes ama: Sutton Hayes 715 (syntypes, not seen). 0.9-1.2 mm long, 0.5 0.7 nun broad, the 2 adax- ial lobes very small; ovary stellate-tomentellous; Additional collections examined. PANAMA, VKH- styles erect, distally quadrifid, hispid-stellate, ca. e between Santa Fe and San JOM-. I) lrc\ 2-4 mm long. Capsules ca. 5 mm long; columella 10314 (MO); : 3 km S of Santa Fe, below 500 m, Folsom X Collins 10 1 4 (MO); a 1 km del puente sobre el desvio 3-4.5 mm long; seeds ellipsoid, smooth, mottled del Rio San Ju an, Luna 47 (MO). gray and brown, apically beaked, 3.1-4 mm long, 2.4-2.9 mm broad; caruncle ca. 1.5 mm broad. 33.3. Mabea jefensteH I rtologia62: 341. Scattered in weedy habitats from extreme south- 1987. TYF'K: Panama. Panama: newly bull- ern Texas to Panama, reappearing in \ enezuela. dozed trail off Cerro Jefe Road, 0.4 km beyond Paraguay, and Argentina, but not reported from turnoff to Alto de Pacora, 29 Sep. 1975, J. T. most of tropical South America. The two Pana- & F. Witherspoon 8570 (bolotype, MO, F neg. 62352). Figure 4.

Specimens examined. P \v\\i\. 1'VWMA: marsh area 2 mi. S of Tocumen Airport, Tvson & Clewed 5899 (MO); RioTapia. linrllett X l.asse, H>629 (MO). Mabea jefensis is known from abundant colh tions from both the Cerro Jefe area and from t Continental Divide north of El Llano in easte Subfamily IV. Ki'i'iioitmoiDi-: \t- Panama Province, as well as from a single eola- tion from the Cafiazas mountain range in the we em part of the province. It seems likely that tl

in, M. J. 1987. Notes on Mabea (Euphorbiaceae) Additional specimens examined. PANAMA. PANAMA: Cerro Jefe area. Intomo el a I. 33'X, (F|. Correa el al. Apodae in Brazil. Phytologia 62: .'.'>') 313. 1601, 1610(MO). Com COM. I 1333 (MO), /i'lm ihiM.K,K.E. i":; -li: in. n - i.' & D'Arcy 6253 (MO, 2 sheets), D'Arcy 121X3 (MO). (Euphorbiaceae) in Panama. Syst. But. 8: 105-117. 12201 (F), D'Arcy X Svtsma 14733 (F, MO), Duke 9474 (MO), Dwrer et al. 7296 (MO, 2 sheets), 7364 The discovery of a distinctive new species of \l(» ' >m el al f>/<>9. , 163 (MO). Hummel i. >'' labia in Panama makes it necessary to provide IM , 36, (I , tAe,nc, , '< (MO). Porter et al. 3072 (MO), Svtsma 1473, 4112 (F), Sytsma et al. 2849 (r). Sytsma X l>C,y 3660 (F), Tyson et al. 3204 (MO). 1333 (MO. 2 d„-ets), l\s„n 33<><» (MO. SC/.), Wilbur & Weaver 11360 (MO); La Eneida, Correa & Dressier 824 (MO), Mans et al. 17,67 (MO); El Llano Carti road, 6-22 km N of Inter-American Highway, 300- 500 m, Croar 25127, 33805 (MO), Hanunel 867 (MO), Huft & Knapp 1391, 1613 (MO). Huft et al. 1868

I MO), Kiuipp I.VHtiY). Km,f,f>rl ,,l t728[V). l.nsnr, cm long; lateral axes 5 8, the peduncles 5-11 mm 1393 (MO), Mans el al. /..,•; |MO), Man A Kaltank, long. Staminate flowers 2 or 3 per node; bracts 1864 (MO), Nee et al. 8752 (MO), Sytsma 960 (F, MO); Canazas mountain chain, near Rancho Chorro, above ca. 1-1.5 mm long, acute, the glands infolded on Torti Arriba, 400-700 in, Folsom et al. 6709 (MO). adaxial side; pedicels 0.5 mm long or shorter, sub- tended by 1 or more bractlets within the bract; 34. Senefeldera calyx segments 3, unequal, acute, as broad as or broader than long, ca. 0.5-0.7 mm long; stamens Senefeldera C. Martius, Flora 24 (Beibl.): 29. 5; anthers apiculate, ca. 0.4-0.5 mm long (much 1841. TYPE: Senefeldera multiflora C. Mar- longer than the very short filaments). Pistillate olitary at the 2 or 3 lowermost nodes of Monoecious shrubs or trees without evident m each lateral axis, sessile; bracts apiculate, 1-1.3 latex; glabrous throughout. Leaves alternate mm long, with glands 0.8-1 mm across; calyx pseudoverticillate at ends c segments 3, slightly imbricate, broadly ovate, apic- iolate; stipules deciduous; blad* nately veined, usually glandular margins entire. Inflorescences terminal, paniculi (of compound sp ruinate flowers solitary or in glome) axils of infloreso dular bracts; calyx 3-5- metrical, not covering anthers in bud; petals and disk absent; stamens 5-12; anthers subsessile on an elevated receptacle, extrorse, dehiscing longi- tudinally; pollen grains subglobose, tectate, 3-col- raphe conspicuous. porate; pistillode absent. Pistillate flowers solitary Lowland evergreen rainforest, Panama to Ven- at proximal nodes of inflorescence, sessile; calyx ezuela; here reported from Panama for the first 3-parted, segments distinctly imbricate; petals and disk absent; carpels 3, unappendaged, each with a single ; styles unbranched, free, or basally The Panamanian specimens are a rather good connate. Fruits capsular, thin-walled; columella match tor collcrii,,ns »,t ,S /.--,/,, >//,-.<<; lioiti Xulia, slender, usually not persistent; seeds solitary in Venezuela (Stcymnark 997,70. 90917, VEN). Al- though they differ from the South American plants in having more acute leaf bases and shorter inflo- As treated by Jablonski (1965: 171-174), Se- /,-, feldera is a genus of nine rather poorly under- stood South American species. It is here recorded Specimens examined. PANAMA, DARIKN: Rio Can- from North I i for the first time. glon, Duke & Bristan 363 (DAV, MO); Rio Perrecenico off Rio Pirre, Duke & Bristan 8236 (DAV, MO); ridge 2 hours by piragua upstream above Tucuti, Duke 5261 34.1. Senefeldera testiculata Pittier, Contr. (DAV, MO; incorrectly cited as Caryodendron angusti- Fl. Venez. 2: 31. 1923. TYPE: Venezuela. Zu- lia: Perija, Pittier 10910 (US). Shrub or small tree to ca. 4 m high; twigs subterete, channeled, smooth. Leaves with petioles variable in length, 0.5-4 cm long, adaxially chan- 35. Sebastiania neled; stipules triangular, ca. 2 mm long, deciduous Sebastiania Sprengel, Entd. Pflanzenk. 2: (leaving conspicuous scars); blades chartaceous, base, 12-24 cm long, 4-10 cm broad, usually with a ventral median swollen gland ca. 0.5-1 mm long; major lateral veins ca. 10-15 on a side, slightly Montane rainforests , or cloud forests, western Panama. flowers borne distichously on the rachis; seeds not over 2.5 mm long 1. S. corniculata This shrubby species, now represented by sev- Shrub 2 -3 in high; leaves elliptic-lanceolate; eral collections from montane rainforests in Chi- staminate flowers borne spirally on the rachis; riqui and Veragua-. doe- ; not appear to have been seeds ca. 4 mm long 2. 5. panamensis overall appearance and floral characteristics it re scinblc;- the weedy N. cor- 35.2. Sebastiania panamensis Webster, sp. nicithittiiA section Micro stachvs:(Adr. Juss.) Muell. nov. TYPE: Panama. Chiriqui: N of San Felix w I\ habit. Il- -piialK at Chiriqui Moras del Toro border, on Cerro Colorado copper mine road, 5,000-5,500 ft., 3 May 1975, Mori & Kallunki 5786 (holo- Pax (Pflanzenrcch 85: 8<>fl'. I O | 2) it would key type, DAV; isotype, MO). to section Klachocroton (V. Muell.) Pax, but it does not resemble any of the species in that -e« lion except possibly N. slipulocea (Muell. \rg.) Muell. \rg.: that -|.ecie-. however, i- entirely glabrous, with broader leaves and multicornute ovaries. The Shrub ca. 2 3 in high, I he trunk ± unbranched; Panamanian plant- do not lit into section lilrno twig- -lender. -uhlerele. antmr-ely ± appressed gyne (Klotzscb) Benth. because the staminate ca- pubescent. I,ciiics with petioles 3 7 mm long, a[»- Ivx is not asymmetric, the fruit is echinate. and pressed puhe-cent: stipules triangular-lanceolate, the blanche- arc not p111x . Nor do the Panamanian dark, 0.8-1.2 mm long; blades thinly chartaceous plants agree with species in section Sebastiania, or membranous, elliptic-lanceolate, ± caudate- because of their leaf-opposed spikes, broader sta-

long, (1 )2 4 cm broad, concolorous, without lam- The sectional divisions used by Pax do not seem mar glands, glabrou- to di-lmetl\ hirsutulou.- on to be very well founded, -o thai a -a ti-la.[oi v both laces; midrib plane above, distinctly raised beneath; major lateral \eins ca. 8 20 on a side. seem to be feasible unt -traiglili-h. -lightly proiuiiiuloii- beneath, distalh At present, it appear anaslamosiiig into intramarginal loops; vemlei- represent a possible < lonniiig a delicate iiicon-piciiou- reticulum; mar gins finely crenulate with ca. 8 25 appie—ed teeth on a side. Inflorescences opposite leaves (some- times pseuiloteriuiual). -pieiloriu. usually bisexual. 2-3 cm long, rachis ± hirtcllous; pistillate flowers Additional specimens examined. PAN Wl \. i Ulliioi I: -olilar\ at base, -laminate Mower- I or 2 per distal Cerro I olorad.., 1,20(1 1,50(1 in, Man X Dressier 78J7 bract; bract- -pirall\ arranged, lanceolate, dark, (MO. dnpl. .U SC/ -ecu bv \1. Hull). Snlhran .<"/lM<». ca. 1 mm long, on each side with a short-stipitate Antonio 1409 (MO, dupl. at PMA seen by M. Unit); above San Felix along mining road, above Chame, 1,200 (ca. 0.5 mm)cyathiform gland 0.5 -0.8 mm across. 1,500 m, Croat 33044 (DAV, \IO). ,c.;: silM Fel.x.Mon Stamina!,- /lowers: pedicel less than 0.5 mm long: 1,200 m, Croat 33437 ( M<)): Cl.iri.pii I rail, pre,„„,) calyx 3-lobed, the lobes obovate, ca. 0.6 -0.8 mm rainforest between I'niol.i ,uid Ouehrada Honda, Kirk- long, the anthers ca. 0.3 mm long. Pistillate flou- bride X Puke 898 (DAV, MO), Churchill cv Chinch,II eis: -ub-e-sile or the pedicel up to 1.5 mm long ,,(>:!3(i: MO): I .a fortune hydroelectric pn.j.-.-t. I.Kill 1,200 m, Hammei\2165 (DAV, MO), Knapp 4980(MO), at anthesis, becoming up to I mm lour in fruit: Mendozaetal. 7 70(MO); E of Fortuna campsite, balsam cal\\ lohe- .'.. imbricate (coxering o\ar\ in bud). X /Vcs.s/,7 >:> mm high, ca. 7 mm broad, with 0 subapical lower triangular processes; columella ca. 1.5 mm high; seeds ovoid- 35.3. Sebastis ellipsoid, reddish brown and mottled, smooth, ca. Several ttuitin Webster & Huft

1-1 I I.Mhi. icellate; calyx of 1 lobe or rudimentary or absent; nhn.n e, broadly ovi petals and disk absent; stamens mostly 2-5; fila- ...ii', i« . ments free or basally connate; anthers extrorse, . Sebasti- dehiscing longitudinally; pollen grains subglobose, ania from southern Brazil and Argentina as S. tectate. 3-eolporate; pistillode absent. I'tstillatc brasiliensis Sprengel. ,S. anisandra (Griseb.) Lillo, floiccrs sessile or pedicellate; calyx moslK 3-lohed. and S. warmingii (Muell. Arg.) Pax. Even closer the lobes sometimes reduced or obsolete, eglandular is .S. macrocnrpa Muell. Arg. of Ceara in north- within; petals and disk absent; carpels 3. each with i i llrazil. with which it shares a large capsule a single ovule; styles free or basally connate, un- (to 15 mm in diameter). branched. Fruits capsular; columella ± persistent; Until flowering material becomes available, it seeds solitary in each loeule, carunculate; testa -ii th: endosperm copious. ndescribed species or belongs to o As here circumscribed, Gymnanthes includes : Braz Actinostcmon Klotzsch and Dartylostemon The Costa Rican specimen cited below is the Klotzsch. Although most authors have upheld at basis for the record of Ophellantha spinosa Stand- least l< tinostemon as a distinct genus, this seems ley cited by Standley from Costa Rica (Standley, 1938: 1557). That species is definitely known only Pax from northwestern Mexico to Honduras. One of & Hoffmann (1912: 13) did not provide any con- the Colombian collections {Gentry & Guadros vincing distinctions in their key, nor did Jablonski 47466A, MO) has an immature inflorescence with (1967: 164, 178), who candidly stated that "the a -ingle pistillate flower and the multiparled brac- distinction between \ctinoslemon and Gymnan- Ira glands ill .il air \ ha car lerisl ir of several species thes is very vague." Even when Actinostcmon is merged with Gymnanthes, the enlarged genus is difficult to distinguish from Scbastiama. Mueller San Ramon, Los Loros, Brenes 22679 (CK, \ i I. I'w (1866: 1164-1165), in fact, combined the two, \M\. I.os SANTOS: 1 2 mi. W of Candelaria, Duke 12443 but called the composite genus Schasiiama be- (MO, US); Las Tablas, Dwyer 1100 (MO); Los Santos, cause the name Gymnanthes (which has pnorilx) :',() MI, l.uo .HMMOI. CoioMHix. VIIANTKX): Puerto Co- lumbia, 5U~K)0 in, Dufruml '>:'>, (| . distributed as S. seemed inappropriate to him. The distinctly re- granatvnsis Muell. Arg.). UOI.iVA " ' duced staminate calyx ol Gymnanthes fumi-hes only a tenuous difference from Scbastiama. in A' t „„,.'/<. r/56,

well developed. As Pax & Hoffmann (1912: 89- 90), , Sebasliania mnanthes Sw., Prodr. 95. 1788. LECTOTYPE: nus, part of which -houM he combined with Gym- Gymnanthes lucida Sw. (chosen by Grise- tithes. Clarification of these difficulties will have bach, Fl. Br. W. Ind. 50. 1859). trepid monographer, Rothmalr. (1944), follow ig a suggestion by

recently (Webster, 1983); Aterc posed of by lectotypifying it so 1

stly persistent; blades ropical genus of about 40 spec ulate, glandular or s are reported hen eglandular. Inflorescences terminal or axillary, bi- sexual (or less commonly unisexual), spiciform, of 1 -several basal solitary pistillate flowers and many BROWNE, P. 1756. The (a\i 1 and Natural History of

ilitary pistillate flowers and 1-several HALLIER, H. 1918. Uber Patrick Browne's Gattungei, . Meded. Rijks-Herh. Leiden 3h:

I'it.'i areae. I. Monograph of the genus Aetmoslemon. I'hslulofiia 18: 213-240. PAX, F. & K. HOFFMANN. 1912. Euphorbiaceae — Hip- ulent, ca. 2-4 r l long ; pomaneae. In: A. Engler, Das Pttanzenn-i, h l\ . I I ,". 13-21 long in fruit; calyx segments 3. V(Heft 52): 1 319 (Gymnanthes, pp. 81-88). pointed, not imbricate, 0.6 0.7 mm long; o\ar\ Hoiiimn u. \V. 1044. Nomina generica neglecta 1753- 1703. K.,,,,. 53: 1 37. ii j laged, smooth, puberulent; styles .3. ha sally connate for ca. '/,-'/, their length, 2.5, 5 mm long. Capsules not seen entire; cocci 11-13 mm long, smooth: seeds plump, ovoid, ca. 0.3 6.6 mm long, 5.8-6 mm broad, brownish, -mooih. oh-cure- lv beaked; caruncle 0.8 1.3 mm broad.

The -ingle Panamanian collection of this species lb. Stanimate bracts (at least in pari) with paired bears only fruits, so the identification of Bristan's glands on each side, each bract subtending 1 plant with a Mexican species must be provisional. flower; leaves eglandular on margins; staminate However, the characteristic leaf venation and es- calyx absent; fruiting pedicels 40-50 mm long; seeds ca. 5.5 mm long 2. G. dressleri pecially the distinctive marginal foliar glands sug- gest that our plant is conspecihe with specimens from Veracruz. Mexico (such as I'urpiis 37<)5. 36.1. Gymnanthes actinoslemoides Mu. II. 8060 from Zacuapan, I 110 from bortin. all at Arg., Linnaea 32: 103. 1863; Pax & Hoff- UC, from which the floral characters have been mann, Pflanzenreich IV. 147. V(Heft 52): 85. 1912 (as G. aetinostemonoides). Sebastiania actinoslemoides Muell. Arg. in DC, Prodr. 15(2): 1184. 1866. TVI'K: Mexico. Veracruz: Zacuapan, Linden 1357 (holotype, G, mi- crofiche seen). 36.2. Gymnanthes dressleri Webster, sp. nov. TYPE: Panama. Panama: La Kneida, region of Cerro Jefe, 3 Jan. 1968, Dressier 3323 (ho- lotype, MO; isotype, DAV). Leaves with petioles 3- i long, ± puberuler im long; blades cha taceous, elliptic-oblong, mostly ide- and pu- mm long; blades chartaceous, broadly elliptic, berulent [iro\uuall\: major lateral veins ca. 10 15 on a side, arcuate: veinlet- distinciU pronii- ;umen obtuse, 0.5-1 cm long), cuneate at base, nulous beneath (diiditlv -o above), forming irreg- -8 cm long, 1.5-4 cm broad, appearing eglan- ular areoles; margins plane, remotek crenulale (teeth mostly 12 17 on a side), the distal teeth lar glands, dis- will i a mini lie deciduous glandular lip. I be pro \ una I tinctly paler and glaucous beneath; major lateral teeth with depressed cyathiform glands. In/lores- veins ca. 6-8 on a side, straightish or dislally (c/ii-e.\axillarv. racemilorm, unisexual oi bisexual. arcuate; veinlets distinctly prominulous beneath, 2-7 cm long, the rachis puberulent. Staminatc often as prominent a- the lalerals. forming areole- (loners in cvmules of .'». subtended b\ iimhonate partly parallel to the lalerals; margins entire, with bracts ca. 1 mm long. each hr.nl with a pair of a plane or slightly reflexed suheartilagiiiou run. cyathiform glands 0.6 0.8 mm across attached Intloieseenees axillary, mostly at the base or lower above the base (-iibupical); pedicels 0.5 1.5 nun axils of annual increment-, racemilorm. unisexual long (longer in central flower); calyx usually of I or bisexual, where bisexual with I or 2 basal pis- Volume 75, Number 3 Webster & Huft

gland on each side, and in its well-developed i ; >ate calyx. Among the species with bracts sub- ious, ca. 0.5-0.7 mm long, mostly with paired tending solitary flowers, the closest to G. dressleri subsessile cyathiform glands on each side at base, appears to be the Cuban species G. albit « <• a the larger gland of each pair ca. 0.3 mm across. seb.) Urban; however, in that species the leaves Staminate flouers with pedicels ca. 0.4-1.2 mm are more elongated and lack cuspidate tips, the long, articulated at the top; calyx, petals, and disk absent; stamens 2-4; filaments free or basally have 5-12 stamens. united, 0.4 0.7 mm long; anthers 0.4-0.5 mm long. I'istillate flowers with pedicels ca. 3-5 mm 37. Maprounea long at anthesis, increasing to 40 50 mm long in fruit; calyx lobes 3, ovate, not overlapping, ca. Maprounea Aublet, Hist. PI. Guiane 2: 895. 1775. 0.5-0.7 mm Ion • _ .. i

Shrubs >rous throughout; latex nei- rtyles 3 a. 1.5-2 mm long, thick monoecious. I.cures alter- nnate at base. Capsules r stipules small. persistent: 5.5 () nun loni!. -111.|.«-1 blade I , entire, glandular or eglan- eeds i 4.5 Infl, Li I!-, smooth, dark brown i I soliturv. pedicellate, pistillate Mowers at basal apically beaked; caruncle ca. 1 mm across. nodes, the staminate flowers densely aggregated in This new species from Cerro Jefe appears to be a strobiliform mass at the end of the fleshy enlarged i If i I ) > I