v,is(i- of the Missouri Botanical Garden 1988...Missouri Botanical Garden, 2345 Tower Grove Av-...

Annals of the Missouri Botanical Garden 1988 #

-

v,is(i-

Volume 75

Number 1

Volume 75, Number ' Spring 1988

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REVISED SYNOPSIS Grady L. Webster2 and Michael J. Huft" OF PANAMANIAN EUPHORBIACEAE1

ABSTRACT

species induded in \ • >,H The new taxa ai I. i i " I ! I _- i II • hster, Tragia 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, Croton 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 ///, Sebastiania 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 Euphorbiaceae for In order to in

the Flora 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, Richeria, and Sene- adequate for description or if the species has been

feldera, and an unpublished genus from Cerro Ta- recently described elsewhere, a literature reference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

Acalypha, Alchornea, Cleidion, Croton, Dale- and has led to the resolution of a number of an-

champta. Euphorbia, 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 Central America

cur in Panama. This new treatment, with the order and northern South America. 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>< ,<! (••,,., M.i'',^ ,i,l,',, ,s D, pfr.'m, at ,,( II,>!•:,•, 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.

ANN. MISSOURI BOT. GARD. 75: 1087-1144. 1988.

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Volume 75, Number 3 Webster & Huft 1091 1988 Panamanian Euphorbiaceae

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.

lil 36, Reibl. 80: 19. 1905. TYPE: Astrocasia

j phyllanthoides Robinson & Millsp. = Astro-

casia tremula (Griseb.) Webster.

11 I t| Dioecious, glabrous trees or shrubs. Leaves al-

J"S ternate, petiolale: iij ili i MM deciduous; blades

nil It

$ "i entire, pinnately veined. Inflorescences axillary;

4J flowers in cluster > • . ,, >cd II i

V * sepals 5, sometimes unequal; petals 5, longer than

I Z& I the sepals; disk annular; stamens 3 or 5, the fil-

;lj| •ffs aments connate into a column, anthers extrorse in

1 2 ^ bud, dehiscing horizontally; pollen grains tricol-

porate, reticu lo ilated at tip into a

III! *il peltate disk capping the staminal column. Pistillate

11 flowers long-pedicellate; sepals 5, articulated, de-

• s *^ « . i petals 5, longer than sepals; disk cupuli-

111! Ill's

U pels; ovules 2 per locule, anatropous; styles free,

!l hil'ul. funis eapsiilai : columella slender, persistent:

I^SI seeds 1 or 2 per locule, ecarunculate; seed 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

:u,,,|

int.,

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

sHiHl and Madaga-- >im, 11 ielandia Rail-

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1.1. Astrocasia tremula (Griseb.) Webster, J.

Arnold Arbor. 39: 208. 1958. Phyllanthus

tremulus Griseb., Fl. Rrit. W.I. 34. 1859.

TYPE: Jamaica: 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 tree 2-10 m high; branches terete or

obscurely angled, pale; foliage deciduous. Leaves

villi slender petiol 2 <> cm long; stipules lanceo-

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;


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; ovary 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. Fruits 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; endosperm 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 Colombia, 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 Old World genus Actephila 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 fruit. On the basis of gross morphology,

Aublet, Hist. PI. Cuiane 256. 1775. from Actephila, es[)ecially by virtue of its pubes-

: Amanoa 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 embryo 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• . |<i ! !!,,tr /Imcers pel

veinlets promiiiuloii b> 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,

± acute, 8-12 mm long, 3-4 mm broad, i i , u

ulous without, persistent and becoming reflexed in

fruit; ovary 3.5-5 mm diam., densely hirsutulous;

styles 3, 3-4 mm long, connate basally into a

column ca. 1 mm high, three times bifid. Fruit

capsular, 10- 1 5 mm broad; columella ca. 8-9 mm

A neotropical genus of five closely related species,

pre\ ionsK mm > I Iron m i'md North Amer-

ica. Richeria appears to be most closely related

to the African genus Marsohotrya li'-nlh. and lo

>.".;., •,., I Jlil'iM ol - nil . a: < • II V la J'l ! Ma a-. 1.1

differs from that of Mueller (1866) and Jablonski

' 'I Vo-in*"""? ^^q ""*TTUB' °"1U""'' (1967), since section Podocalyx (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-

dicated by the spinose pollen), belongs in the

I i lil I » I deae rather than the Phyllan-

There are two species of Richeria in Panama.

neither previously reported.

or nearly so; stipules less than I folate 1. H. obov

distinctly puberiilent: si vies

This more complete species de- icription of Croiz-

atia panamensis has been mad e possible by re-

cently collected flowering specim ens and data pro-

vided by Dr. Gordon McPherson. It is now apparent

that C. panamensis is clearly different from C.

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 unknown, it remains difficult lo i tssess its relation- ships wiili ( «...

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. 1922; Jablonski, Mem. New York Bot. 4. Richeria

ZV. tvzz; jablonski, Mem. l>ew I one not.

Card. 17(1): 126. 1967. Richeria grandis

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:

Trees or shrubs; dioecious; indumentum simple "Brazil,1' Rio Casiquiari. Spruce ,'i.'>2f) (not

or absent. Leans allereale. j>ctiolal< : stipules de seen).

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 bract 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


\ ,,i 1:1 I In,, PANAMA. PANAMA: ca. 5-6 mi.

, Gentry 5796 (GH, MO, SCZ). cloud forest, Cerro Tute, NW of Sa. Hunk, yju-i (DAV, MO).

4.2. Richeria dressleri Webster, sp. nov. TYI

Panama. Panama: Santa Rita Ridge, road

Estaci6n Calibrar el Agua Clara, 9°22'

79°42-45'W, 1,000-1,500 ft., 26 June

1971, Webster & Dressier 16744 (holotype, DAV; isotype, MO).

capsulifl 2-locularis; foliis a i lifl'ert stylis elongati

s acuminatis eglan ' ' > sericeo-hispido.

Tree to 15 m high, 3.5 dm thick; twigs

mostly densely appressed-hirtellous when j

c\ I-I iu.i \ jl.ihi ate: loliage e\ ergi ecu. I.cares vviih

hirtellous petioles 1.5 5 cm long; stipules lanceo-

late, 1-1.5 cm long, densely sericeous, caducous;

blade < el ai laceous. «d«»\ ate. mo ll\ al ip||\ ,i. a

on the petiole, 10-30 cm long, 4 14 cm broad

major veins mostly 10-12 on a side, straight hnx-hidodrouioii.s, I In- mi. J ill saliently raised be

neath; secondaries archingly and irregularly sea

lariform; ultimate veinlets fine, scarcely prominu

Ions; surface of blade above glabrous and flecked

or pitted with minute colored spots, beneath b

colored and densely to sparsely hirtellous (b

ing glabrate in age exce|>t al-mii i h il • . mi ! ny>-i

veins); margins entire, plane or recurved. Inflo-

rcsccnccs spied. <n i nun le . ikes 1.5-5.5 cm

long, pistillate spikes 1.5-7 cm long; a\es den-el\

liiiMiliil.uis without. Shim,naic flowers sessile; ca-

lyx deeply 4 5-lobed, densely hirsutulou without;

ealvv lolies ohlong to obovale or suborliicular. un-

prismalie. apiealK hirtellous. 0/2 0.3 mm high;

stamens T)( ()); filaments free, 2 3 mm long; an-

thers ellipsoid, ca. 0.4 mm I i [ i I, lui

ducal. denselv hirtellous, 1-1.2 mm high, 0.6

0.9 mm broad. Pistillate flowers subsessile; sepals mostly 4(-5). elliptic, lometitulose outside, sen

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

I; columella flattened, papery

long; seeds somewhat asv in

ally ovoid-ellipsoid, tapering

with reddish, fleshy ribbed-st

mm long, 4.8-5.3

Bharpl,

noecium with distinct styles; t

are similar to th <>| /»'.•< itenu ^uuuhs toil ddlei

in the more slender cylindrical pistillode. The acu-

minate leaves often copiously hirtellous beneath

and the large foliaceous stipules also appear distinctive. The • oil.-, lions hum ( !os|j Rica are mor-

pho o i III divergent but mav tenlalivelv be

grouped with the Panamanian plants. It seems ap-

to our knowledge

s made a significant contribution

.I Mesoamencan Kuphorbia<eac

'broi,; h II - in inv . I'lleclioiis and 1 i; monogi iph

of Pedi/anthus (Dressier, 1957) and Euphorbia

subgenus Poinsettia (Dressier, 1961).

Additional specimens examined. COSTA RlCA. HERE- DIA: Finca La Selva, the OTS Field Station on the Rio Puerto Viejo just E of its junction • n- h, \\ • - , • | •• < | • J > 100 ir,(!ni.(,/in;/miv/V,M / I I 7 ( V I. /la,,.:,:;'. < ,'.'•. ««.'*:* (F), Jacobs 2 I 79(F). PIINTARENAS: on roa.l to radio and telecommunications tower 6 km N of Oolfito, .too 100 III. / tte\ X I tier 1902 (K). PANAMA. COCI.E: near El Yalle .le Anion, ca. 8°37'N, 80°07'W, ca. 550 m, Mcl'hcrson .<>!<> (F). COION: Santa Itita lumber ...a.I. 15 km E of Colon, Dressier & Williams .Vtoft (MO). PANAMA: rainforest along El Llano-Cartf road. 1.0 l\.2 mi. N of Panamerican Highway, 350 450 m, lVArcy ///"otMO.ilupl.atSC/s.Tiih\ M. Hull). CV/,/M .OVo

, i" 'I ' D\\ I MO), McPherson 9959(F, MO), Mori & Kallunki 5607 (DAV, MO), SAN BLAS: Cerro Brewster, 9°18'N, 79°16'W. 850 m, de Nevers et al 5414 (F).

5. Hyeronima

Hyeronima Allemao, PI. Novas Brasil 1. 1848.

TYl'K: Hyeronima alchoincoidcs. \ll< inao

A number of collections made in cloud forests

in Panama indicate that there is at least one ad

I • - • • ii< the lowland //. laxiflora. However, as often happens, these

taxonomically; the following revised treatment is

li lil i i ending revisionary studies of this

; pistillode slender, bid,I. 0.7 0.8 n s sparsely lepidotc; blades inosilv I' g with petioles of 3-9 cm; stipu

of 1-2 cm; stipules apparently obsolete; ovary glabrous to lepidote; endocarp of fruit at least 4 mm long 2. H. oblonga

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 bracts); calyx

cupulate, shallowly 5-lobed, densely lepidote, 1-

1.3 mm high; disk cupulate, subentire, glabrous,

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 inflorescence 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 , T • o • i r> /- ,r* above) and larger flowers, appear to match the Hieronyma guatema I. 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- <!•••• Hike 783 (MO), CHIRIQUJ:

drnmrm*- midrih and vpins rai^d heneath and + Ce"° HornitOS, ca. 40 km NW of Gualaca, 2,238 m, dromous, midnb and veins raised beneath and ± ^ & ^^ ^^ ^ (QAV ^ pate

hirsutulous, the veins and veinlets distinctly prom- Macho 4 km N, , . , . ^ e( fl/ 48(j8 (MQ)

inulous above (upper surface scabrous to the touch); COCLE: La Mesa, 2.5 km N of El Valle, 850 m, Mori et lepidote scales on upper surface scattered to absent, al 6610 (DAV, MO); hill 3 km E of El Valle, 2,500 ft., ca. 0.1-0.2 mm across, with reddish center, be- Hammel 4776 (MO); swampy area 5 mi. from El Valle,

. .j J « l • „ „ n o Gentry & Dwyer 3622 (DAV, MO); foothills of Cerro neath sparse to dense and overlapping, ca. 0.2- (>|1| ,,, i( ^{] _ f , ,,

0.25 mm across, with pale center (lower leaf 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 the staminate ones 5-10 cm lone the Distillate & Breckon 16 1 I • xs: 3-4 km W of Santa %, tne staminate ones o IU cm long, uie pismiaie Q ^ / y ;/ ^ Ay ^^ ^ ^^

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).

1'rccs or shrubs, dioec

-of simple hairs. Leaves alternate, short-petiolate,

ipulate; the blades often coriaceous, entire to

; axillary; flowers in axillarv

cauliflorous. Flowers apetal-

sepals usually 4 or 5, imbricate, deciduous.

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

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 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

original treatment.

Drypetes standleyi \

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 from Transisthmiai, Highway, Dressier 38I<H\]0). M K\<a AS: Alto P.edra Santa h-, /.„„ ,V Maasola /.'.'"(MO): CerroTule, Mori et ah 734 I (MO).

, Madrono 24:

65, fig. 1. 1977; Croat, Fl. Barro Colorado I. 529, fig. 321. 1978. TYPE: Panama. Canal

Zone: Barro Colorado I., Armour Trail, Fos-

ter <£ Croat 2307 (holotype, DAV; isotypes,

DUKE, F, F neg. 62358, MO).

levi may occur over a broad area in Panama. A

barren collection from the Burica Peninsula, Chi- rtqui Province {liuse\ f<l)J, \IO) may possibly rep-

resent I), standleyi, although it differs from the

prominent \einlet reticulum. I'he >peiaes niav, also

occur in Costa Mica; a specimen from La Selva

(Hartshorn 1009, DAV) resembles I), standleyi,

although it is divergent in having more slender

i L., Sp. PI. 981. 1753.

mique du genre Phyllanthus (Euphorhiaeees). Hois-

8.11. Phyllanthus anisolobus Muell. Arg. in

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.

Additional specimens examined. PANAMA. I«K:AS DFX I oiio: Ki'o Trnli.-. UT.YN I'IICIO I'.ilen.pie, knUuidc X Hale ,...•< |\|()| IIMIIIN: M.mono. Kirkhnde X II,,Man !><>:> AID): Km Balsa, between Manene and Guayabo, Puke .V \ol.ason 11'/.,!! (MO); bio Pnc.ru. between 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).

8.12. Phyllanthus gentryi Webster, sp. nov.

TYPE: Panama. Darien: lower slopes of Cerro

Pirre, 200-500 m, Gentry & Clewell 7017

(holotype, K, F neg. 62354; isotypes, DAV,

MO).

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,

lloueis. St,unuiatc /lowers with pedicels 15-20

nun long; scpab •. lu-iMtlh i-lliplii oi oh • M \ .' •

3 mm long, 2.2 2.7 mm broad; disk entire, angled,

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.

3 car(iels; styles in-arb free, dilated, 1-1.2 mm

long, 0.7-1 mm across, ('.apsides reddish, valves

12.5-13 mm long; columella massive, 4.5-5 mm

long, 4.2 4.5 mm broad; seeds trigonous-umbo-

nate, 0.5-7.1 mm long, 5.2-5.3 mm broad,

smooth, with irregu n \\a\ • horizontal dark brown

I i i I i i light hi mi !>.!• [ I'lound. the apex some

1.5 mm across); hi

i long and broad.

triangular, broad,

•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 I'nre e.i H°00'N, 77°45'W, 550-1,030 m. Ud'h, >,,•;,

I'liyllanthus gentryi is the first representativ

of subgenus Xylopliylla (L.) Pers. discovered i

Panama. It clearly belongs in se(

(Klotzsch) Muell. Arg. by virtue of its

P. juglandifolius Willd. in general a.'

though il could be interpreted as a sub

/'. juglandifolius, it is distinctive in its la

9. Caperonia

Caperonia A. St. Hil., Hist. PI. Remarq. Bresil

244. 1820. LECTOTYPE: Caperonia castanei-

folia (L.) A. St. Hil. (Croton castaneifolius

L.) (chosen by Britton & Wilson, Bot. Porto

Kimh: 180. 1925).

C. paludosa Klotzsch. It is extremely difficult to

separate that species I mm ('. castancifolia (L) A.

Si. Nil., and we now believe that our Panamanian

specimens of (.'. paludosa probably represent ..nh

forms of that more wide-ranging species. However,

the narrower leaves of plants referred to C. pal-

udosa are distinctive, and furlhei studs in the held

l i« -1 d to eslal i h Itl p. • I. - < oneepl

can be upheld.

10. Argythamnia

Argythamnia P. Browne, Civ. Nat. Hist. Jamaica

1805: TYPE: Alchorneopsls floribunda (Benth.) Muell. Arg. {Alchornea glandulosa

var. '^floribunda Benth.).

.1 Alchorneopsis floribunda (Benth.)

Muell. Arg., Linnaea 34: 150. 1805; in DC,

Prodr. 15(2): 705. 1800. Alchornea glan-

dulosa Poeppig var. floribunda Benth., Hook-

er's J. Bot. Kew Card. Misc. 0: 331. 1854.

: Spruce 2681 (holo-

I'he •

the key on p. 221 original treatment, from Darien, '

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, 40 100 in, I llry & I tley 4938 (F). PANAMA. BOCAS DEL TORO: Cerro Pila de Arroz, along road to Chiriqui Grar 10 road-mi from Continental Divide and 2 mi. al pipeline access road E of highway, ca. 8°55'N, 82°08' 350-500 m, McPherson 8750 (F).

45.

Kar

jndron Kars

1860. TYPE:

ten, Fl. Columb.

Caryodendron

1: 91 :cn«

12.1. Caryodendron angustifolium dley, Publ. Field Columbian Mus., Bot

4: 217. 1929. TYPE: Panama. Chiriqui

greso, Cooper & Slater 192 (holotype neg. 59913).

Scr.

Pro-

F, F

The collection h li«- i I ' in nulci '

in the original collection is now known to be Se-

ne/eblera lesticulata Pittier (q.v.). No additional

collcclioris el < ('.< wi./i 'Hi/on <nt<:<!>! :/<>!, •, ».< nc

known from Panama or elsewhere.

Adenophaedra (Muell. Arg.) Muell. Arg. in Mart.,

Fl. Bras. 11(2): 385. 1874. Bernardia sect.

idenophaedra Muell. Arg., Linnaea 34: 172. 1865; in DC, Prodr. 15(2): 918. 1866. TYPE: llcrnniilt*; . . , Muell. Arg. =

Adenophaedra megalophylla (Muell. Arg.) Muell. Arg.

Dioecious trees and shrubs; indumentum of sim-

ple Incliomes. I,rates alternate, petiolate, stipu

late; blades pinnately veined, without embedded

laminar glands, dentate Inflorescences axillary or

brae , ii ,(,, , i u II filiate or several

calvx splitting into 3 valvate lobes at anthesis;

petals and disk absent; stamens 2(-3); filaments

short; anthers with enlarged connectives, dehiscing

IIIIIOIMK <IM 1 iii:uitii(lin II j llode absent. Pis-

iinbncalc; |>etals absent; disk 3-lobed; ovary of 3

This poorly known genus, hitherto considered

to be South American, includes only three species.

Croi I ' 1i ported aedra from Pan-

ama on the basi-

but in the original treatment (Webster & Burch,

1968: 278) it was pointed out that his original

generic disposition (J. Arnold Arbor. 24: 167. 1943)

ol tins plant i- iiannm was cor-

i • ough that species is now known to be

anonymous with C. membranaceum Pax & K.

fii nil; r t.-.bh-b the piesen •(• ol Idenophai <ii<: in

pear- that lh< ;.l m II in the original treatment is actually / ^i

13.1. Adenophaedra grandifolia (Klotzsch)

Muell. Arg. in Mart., Fl. Bras. 11(2): 386.

1874. Tragia grandifolia Klotzsch, London

J. Bot. 2: 46. 1843. TYPE: Guyana: "British

Guiana," Sehomburgk 948 (presumably K, not seen) H Klot/scb)

Muell. Arg., Linnaea 34: 173. 1865; in DC,

Prodr. 15(2): 918. 1866.

Mus., Bot. Ser. 4: //ru/n/n (Standlev)V 54: 200. 1967. T

>l<j| • f'lK.L 1'ick 1 .luilll.i'111

Shrub or small tree to 8 m; twigs smooth, red-

dish, thinly puberulenl. tardily glabrate. Leaves

with petiole 3-6 mm long; stipules lanceolate, ca.

1 mm long, ca. 1 mm broad, caducous; blades

12-35 cm long, 3-12 cm

long as broad, glabrous, or

below on the principal veins,

to 25; pistillate spikes thicker, not flexuous. 5 I 2

• in long, wild 4-7 solitary flowers. Staiim-n,

em to 12 per glomeruli earl\ dehiscent, leaving

persistent pedicels ca. 1 mm long; calyx lobes mem- hranoiis. ilcll,ilc. spreading at anthesis, ca. 0.5 mm

long. Pistillate /loners not seen; pedicels at ma-

turity ca. I mm long, reflexed, sericeous. Capsules depressed globose. deepK 3 i<>ht<l. (> » mm high.

12-18 mm diam., sericeous, glabrate at maturity,

the persistent calyx lobes ca. 2 mm lout:, deltale.

sericeous; seeds suhglobose to ovoid, ca. 1 mm

long, ca. 0.7 mm diam., yellowish, mottled.

<;,;„„•: el „l. :'.'

14.1. Bernardia macrophylla Siandlcv, I Wash. Acad. Sci. 15: 103. 1925. TYPE: Pa- nama: Rio Tocumen, near sea level, 3 Jan. 1924, Standley 29389 (holotype, US).

No Panamanian specimens of this species have been found in addition to those cited in ihc original

treatment. The recent collection cited helow ex-

tend- llie range to Co-la Iv'ica. I hat .specimen has

ranamainan eolleclioiis and was taken at a con-

siderably higher altitude but seem- otherwise idcn

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

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,,-,, Crande, 10 read-miles from Continental Divide and 2 mi.

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

\\\ olSan'.a le , •>7t>'(GllM0).

TAKKNAS: foothills of the Cordillera de III Tres Colinas, 9°07'N, 83°04'W, 1,800 ,,,/s, el a! _'.>6// (F).

Adelia 1 .. Svst.

Miller, Card. Diet. abr. ed. 4, 28

54. LECTOTYPE: Bernardia earpinijidic

iseb. (see Buchheim, 1962). \ Ichor.ua Svv., Prodr. 98. 1788; Fl. Ind. Occ.

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-

miidn, (Kii|.horhiafeae). Willdenowia 3: 217-220. /enrcicl, l\. I 1.. \ 11(11, it (

ciilum only moderately

M p| •

mostly axillary 3a. heaves

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

record from < olon hi (< ,1 i: Ordonez et at. 58,

MO) The plant described from Colombia by Croizat

(Caldasia 2: 357. 1944) as A. umboensls may

prove to be a form of A. costaricensis.

Idditional specimens examined. PANAMA, CHIHIOI i: \iemil\ of San Bartolo l.iinite, 11 mi. W of Puerto Ar- muelles. Croat 21973a (MO). COCLE: along Rio San Juan below its junction with Rio Tife, Hammrl .tin, (MO). COLON: Rio ltui|iit i. v i. n • i • •>••. b Rio Escandaloso,

along liui 1.ii,m li i ,i il ' c« lii ,1, i //,-;// A I /."' I\K)I: .I mi. from Portobelo, Correa di Dressier I7V)(C,\\. MO), DAKIKN: Kio Ucurganti. liristait Il'i7

(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, Holdridge 6471 (MO), SAN BLAS: Canagangi, forest up- >lrenm of village, 9°24'N, 79°24'W, 100 m, de Nerers et al. 5720 (F).

».2. Alchornea latifolu 5w., Prodr. 98.

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

forms that appear very different from typical A.

latijblia of lowland Central America and the West

Indies. For example, plants with entire leaves and

niiiisiialU short petioles are found on Cerro Jefe

and Santa Rita Ridge (e.g., Gentry & Dwyer 5536,

Croat 15309). These specimens somewhat suggest

the South American A. pearcei Britton, but their relalncK lon;;; f >«• 111 d«• - ami -liort s|> kes ::in-, tfiei «

closer to A. latijolin. >[>• nam- -,\ ith very unusual

narrowly obovate leaves have been collected on

Santa Rita Ridge (e.g., Croat 13844, Duke 15264);

ordinary amplitude »i I i

1 studies in the field

nain unsatisfactory.

ade, the situation will

hl:!(i;„ial specimens examined. PANAMA, CHIRIQUI:

San Felix, Croat 33416 (MO). PANAMA: Cerro Campana, Croat 14673, Duke 10742, Sullivan 434 (MO), Mendez 19, 49(F); Cerro Jefe, Buyer et al. 501H. 50 19, (,entry

(MO Gentrj & Dwye, i.i it (Oil. MO). II ehste, & Dressier 16454 (DAV); between Cerro Jefe and Cerro Azul, Tyson et al. 4325, Mori et al. 65 13 (MO): Cerro Azul, Dwyer 5042 (MO), Lao & Holdridg, .i.i (1)\\. MO), Stimson et al. 5158 (GH, MO), Tyson & Blum 4081 (MO); N of El Llano, Gentry 5105 (MO); El Llano- Carti road, 7.8-8.6 mi. from Pan-American Hwy., Fol- som 3572, Mori & Kallunki 6405 (MO).

16.3. Alchornea triplinervia (Spn ngel) MueB Arg. in DC, Prodr. 15(2): 909. 1866; Pax

& K. Hoffm., Pflanzenreich IV. 147. VII (Heft 63): 227. 1914. Antidesma trip/inert turn

Sprengel, Neue Entdeck. 2: 116. 1821. TYPE: Brazil. Rio de Janeiro: Serra do Mar, Gardner

617 (neotype, G; chosen here).

hose of Alchornea trip-

tlchomea previously known from Panama. Another

eolleclion that may represent •/. Iriplincrria is

Hamrnel 7252 (MO) from Cerro Sapo, Darien; this

has much larger leaves and somewhat resembles

some of the aberrant forms here treated as A.

for the present, its assignment must be

regarded as dubious.

The collections from Code do not fit any of the

varieties recognized by Pax & Hoffmann (1914:

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.

16.6. Alchornea grandis Benth., Bot. Voy. Specimens rxamim-d. PANAMA. <:O<:I,K: h 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-

3 from Choco Province, Colombia {Fernandez

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-

_:—^e pjant: staminatc spikes nmstlv compound,

cm long, with 1-6 lateral axes; staminate

glabrous, sepals ca. 1.2 mm long: stamens he Darien collection of Terry & Terry 8, anthers 0.7-0.8 mm long, blunt.

16.4. Alchornea glandulosa Poeppig

pittieri (Pax) Pax, Pflanzenreich

Costa Rica: Canas Cordas, Pittier 11101

otype, US).

was reported in our original

ot additional specimens

confirm the widespread irrenee ot I , V(il ||

glandulosa in montane forests of Panama. Ex-

amination ..I till | IIMl <j UNI' ill [ U>

PANAMA, VKKACUAS: Isla oad from Canipameiito Jimc.il io Coloma IVi

-Hk.

pittieri, originally described from Costa Rica. Tl

variety is very similar to var. glandulosa of t

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<S w'".' '" • <J' ; ,'" lt>^<oi,lra<-,^,oanobtusebase;marginsdistant|y caAC900: •%e7um (SJuS NW of sla F* 2.8 "enulate-dentate (ca. 10-15 glandular teeth on a km Iron, Ks.uela Agricola, Alto de Piedra, Mori & Kal- *',Je)> 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.

This sinking species stands out from all other

Panamanian taxa by virtue of its long, pendulous,

cauliflorous inflorescences and its large, coriaceous,

more or less oblanceolate leaves. It resembles A.

grandis in a number of respects but differs in leaf

hap< MI sparseness of the laminar pubescence.

Specimens examined. PANAMA. DARIEN: La Laguna ridg.- brtwe.-n Pucuro and Tapalisa rivers, 820-840 m, Centra cv Mori 13560 (DAV, MO); top of Cerro Mali, 1,400 m. Gentry & Mori 13693 (DAV, MO); Cerro Tacarcuna, Gentry & Mori 13938 (MO); Alturas de Nique, S of El Real, 900-1,250 m, McPherson 11614 (MO).

Recent collections in Panama and further study

he South American species have greatly altered

picture of Cleidion in Panama. Largely as a

ult of problems encountered in the preparation

this account, the junior author has undertaken

evisionary study of the neotropical species of

lclusions expressed here must

recent collection

Pherson may bel<

(J. Arnold Arbor.

irien by Dr. Gordon

ong to Cleidion prcullum Cn

. 24: 167. 1943), a species

erwise known only from material collected in the

basin of the upper Rio Madeira in Amazonian Brt

zil. The poorly known Polyandra bracteosa Leal

(Arch. Jard. Bot. Rio de Janeiro 11: 64. 1951),

Rio Madeira, now appears to be synonymous with

* DER WERFF, H. & A. R. SMITH. 1980. Pterido- |i'm i< - of the State of Falcon, Venezuela. Opera Bot. 56: 1-34.

17.1. Cleidion membranaceum Pax & K.

Hoffm. in Engler, Pflanzenreich IV. 147.

XIV(Heft 68): 23. 1919. TYPE: Venezuela.

Lara: around Palmosola, in forest along Rio

Aroa, near sea level, 26-28 June 1913, Pit-

tier 6375 (US, photo F neg. 44609).

Cleidion woodsonianum Croizat, J. Arnold Arbor. 24: 167. 1943. TYPE: Panama. Panama: vicinitj id Sal amanca Hydrographic Station, Rio Pequeni, ca. 80 m, Woodson et al. 1587 (holotype, A; isotypes, F, F neg. 62417, MO, F neg. 62356, NY).

There appear to be no differences between the

Panamanian plants and the Venezuelan collections

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

& Smith, 1980), which is also true of the Pana-

manian collections. The recent disjunct collection

from Peru strengthens the probability that this

accounts for the peculiar disjunctions in its range.

Further study may show that Cleid

(Ca-

San sar.) Baillon, of

Paulo, which has leaves

to those of the Panamai

Brazil from Bahia I

plant! ^

nearly filiform pistillate inflorescences, and similar

"• i inn 'is examined. PANAMA. PANAMA: Maje, second growth on limestom I il !• I< ng » boco

' I. ca. 5 mi. up from village of Man ' ••'< ' <<•' Kennedy 2021 (MICH, MO, PMA). PERU, HUANUCO.

II I I Rupa Rupa, Tingo Maria, ca. 9°18'S, 75°59'\\ „ II posite airport, Plow- man et al. 11246 (DAV, F, K). VENEZUELA, FALCON:

Parque Nacional Quebrada de la Cueva El Toro, steep wet valley along river, 10°50'N, 69°07'W, Liesner et al. ::22 (I . MO). YARACUY-FALCON: Reserva Forestal "Rio Tocuyo," a 4 km del Campto. "Canelon," via Tucacas, Blanco 895 (MO).

17.2. Cleidion castaneifolium Muell. Arg.,

Linnaea 34: 184. 1865. TYPE: Peru: Pavon

(holotype, G, F neg. 7159).

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<ernes, and large capsules with

cariuate lobes), their identification seems

The only other Panamanian species of the 10 m high; dioec.ous; Iw.gs glabrous. ^mis (; „„,„, W„„,.„„,, aiuJ lfu, Poruvjan (].

Pax both differ from C. castaneifo- eaves with petioles

sparsely pubescent with short, white, appressed ,,,„„ m theif sma„er ,eaves much shorter petioies

ii long). ard apex; stipule-- not e\ ident:

blades membranous or thinly chartaceous, elliptic,

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 raceme seen); staminate (.irosla Solfs UO-W, Little 6/96, both at F, both

distributed as Alchornea).

There appear to be no salient differences sep-

arating Cleidion castaneifolium from I'anama and

South \nicnca from the Mexican and Central

\IIICI nan populations that have long been referred

to ('. ohlon^ijoliiim. In addition to the enmiuon

populations all

tin rses 5 9 cm long, wi

at each of the 20-50 nodes, the rachis densely

puberulent. S/„,ninote /loners on pedicels to 1

mm long; calyx lobes cucullate. reflexcd, 1.5 2

-ecu: ln.ni- narrouK Ian late, rigid, divergent

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

by Cerro Pirre, along Q Peredngo (I'.irasrinm) i the margins; styles (persistent on mature fruits)

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

ick; columella 6 8 mm long, trigonous, narrowly

nged, the seed scars elongate, conspicuous; seeds

d light and dark brown.

i:'i<;;;{\-, MOi.

Rainforests, souther)

Acalypha L., Acalypha

19.10. Acalypha cuneata Poeppig in Poeppig lections), and since the one capsule 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,,,„gu1Sh,„g charace^c, of ,hU species <la?e ;^Z.•:Sd": «*^ elhptic-ovate leaves, petioles 3 ., cm long and gla- ohovata (Benth) MudL Arg in ^ prodr 15(2).

brou- except for the suhpiiberulous tips, long un- 816. 1866.

Webster & Huft

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 <onfirm the identity of the Panamanian

cm long, 4-7 mm thick, loosely flowered with 15- sPe(:if with Plukenetia volubiUs of the Antilles

the rachis glabrous to densely puberi South America. Recenl collection.-, have .

filiate /hirers solitary; bracts ± reniform, extended the range to Costa Rica (GSme

wide, inconspicuously 8 "'«'• ^ Nicaragua {White 5323, F),

)-lohed, each lobe with a short tuft of bristles, , Mexico {Calzada 103 1. I I.

the bracts otherwise glabrous or lightly short-stri- Additional specimens examined. PANAMA, COLON:

gose; calyx lobes obscure; ovary densely hispid, upstream from bridge over Rio Guanche, 0-100 r

styles free, light I \ strigose.

of Panama treati

/ (F, MO); near Portobelo, Croat 12V6<H\H));

ent,reiengthinto8-12narrowsegments.^.,,/r ^^1^-^

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

Fneg. no. 7110).

Plukenetia angustifolia Standley, Publ. Field Columbian Mus., Bot. Ser. 4: 314. 1929. TYPE: Honduras. Atlantida: Lancetilla Valley, 8 Mar. 1928, Standley 56708 (holotype, F, F neg. 52742).

Liana; twigs | li it. glalues. cut.

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

tire or shallowly dentate. '. Spikes mostly bisexual,

1 or 2 pistillate bracts , it base, these subtending 2 or 3 flowers A. diversifolii

'. Spikes unisexual, to 15 cm long, the pis-

few at the distal nodes. 5 Uaminate ftowersmth | .< ni i, '-•!:- -| I. , rt-pilose, 4-7 n im long; calyx segments

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

.'TV) \\ , St/w„„ ///. lb. MO) I'..-. ,M\. .|,„i.. the I I Llano Cart; rd., ca. 10 mi. N of Pan Am Hwy., 500 m, Centn ri al K«/«(M<». -\\ iti \x F.l l.lanoCarti road,

i with 22. Tragia ..,'.|i. . |: |.. i i.until: . 2 I'd inn, lout:, ill. M- narn.wh

clavate, strigose when young, glabrate to sparsely

lis.11 i nilii- . ml a I maturity; calyx lobes lanceolate,

1 1.2 mm long, ca. 0.5 mm broad, strigose in a

km.: along the center; ovary of 4 carinate carpels,

strigose on the keels, otherwise glabrous, the stylar

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

ally, reticulate-venose, brownish mottled, ca. 5 mm

long, 3-4 mm thick.

Lowland evergreen rainforests, Mexico (Oaxaca,

Yucatan Peninsula) to Colombia and Northern Bra- .- ( I' !•• .i i. II • !. i epoi !i <i li-iptn Panama loi the first

wen des. nhed b\ Standlev as /' aiii;

no salient differences between these plants and

those of South \meru a are apparent.

Specimens examined. PANAMA, COLON: Santa Rita Huli-'i- in i i • IM. I.nn f.-.a \-na I l.u; ; ami. il .laLna 10(1 ..00 ,„, Foster & Morion 2222 (F); Santa Rita Ridge, ca. 9°20'N, 79°45'W, ca. 500 ,„, \L-l>hc,so„ 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

Tragia L., Sp. PI. 980. 1753. LECTOTYI-K: T,,lfiia

volubilis L. (chosen by Small in Britton &

Brown, III. Fl. N. U.S. ed. 2, 2: 458. 1913).

The discovery of three additional species in Pan-

ama makes it necessary to provide a key to the

PAX.F.&K. HOHMVNV ]<>|<). KupliorliKiccac \ca- lypheae —Plukenetiinae. In: A. Engler, Das Pflan- zenreich IV. 147. IX (Heft 68): 1-108 (Tragia, pp.

n 3-lobed; sta:

i ,1 i

ers5-10on § long, 1:

is 2 12. •rate, the

1. T. baillonia, 8 than 10 cm

pistillate flow-

llllioll eerie,

3a. Pistillate

. solit

flew.

mose, not bifurcate; pis-

tr long-pedicellate; styles

3b. Pistillate ale: styles free

Tragia bailloniana Muell. Arg., Linnaea

i: 178. 1865; in DC, Prodr. 15(2): 927.

366. TYPE: Mexico. Tabasco: Teapa, Linden

21.1 Acidoton nicaraguensis (Hemslev)

Webster, Ann. Missouri Bot. Card. 54: 191.

1967. Cleidion ?nicaraguensis Hemsley, Biol. Cent.-Amer., Bot. 3: 130. 1883. TYPE: Nic-

aragua. Chontales: Tate 352, 455 (syntypes,

presumably K, not seen).

Additional specimens examined I'WWIA. COLON:

Sanla Hita, Cone, <v Ihcsslcr 00.' (MO); Santa Kita Ridge Rd., 20 km from Transisthmian Hwy., 9°24'N,

i mu I.- I

co-.ei-.-ii .vilh stinging hairs. Leans will |..-liole-

8-14 cm long; stipules ovate-lanceolate, acumi-

nous, broadly ovate, unlobcd,

lobe, or shallowly 3-lobed,

at tip, deeply cordate at base, 12-25 cm long,

11-18 cm wide, sparsely beset above and In-low v.illi Minting hairs. usu.ilK 7-\eined at the base.

the margins renmleK denticulate. Inflorescences

Webster & Huft

long, reflexed; pedic

glabrate; sepals 5,1

6 mm long; stamens ca. 40; buds pyrifoi

I'istillolr Jlt,t<;-rs sulilais in the axil ol c

. i- ;> i

gins ciliate with long, stiff 1

ii juti I i Miff hairs ca. 1 n i long; styl.

long; mar

•y densely

black,

6 8 mm long, fused

branches slightly spreading. Capsuh < !

3-lobed (one lobe sometimes abortive), ca. 15 mm

diam., ca. 8 mm high, densely hirsute with stiff

hairs; columella 6-7 mm long, with 3 prominent,

narrow wings at tip; seeds nearly globose, ca. 6

• 9 light longitudinal

) western Pan

petioles 4-11 cm long; stipules lanceolate, 5 mm

long or more; blades thinly chartaceous, oblong-

or elliptic-obovate, rather abruptly short-acuminate

at tip, distinctly cordate at base with open to closed

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

bluntly and coarsely crenate (teeth 15-25 on a

side). Inflorescences opposite the leaves, becoming

ca. 10-15 cm long, distinctly bifurcate and pro-

togynous, the lower pistillate branch with 5-10

flowers; pistillate bracts entire, 3-4 mm long, the

.5 2 i • ong.

•otli. with i

Forests, southern Mexicc

I' \luell. Arg., dis-

tinguished from all other American species

, 20) is (ca. 40 vs. 2 or 3

xtension, for

n only from s

ially lobed

considerable range

(Veracruz, Chiapas,

duras. Earlier reports of Tragia bailioniana from

Costa Rica (Standley, 1937: 622) are erroneously

based on collections of Dalei hampu haul

collections establish the presence of the former

Specimens examined. COSTA RICA, ALAJUELA: along

to Colonia Virgen del Socorro, 10C18'N, 84°10'W, ca. 800 m, Burger et al. 11850 (F); lower NE slope of Arenal Volcano, 10°29'N, 84°42'W, 500 m, Lent 2947 (!• | [H hi Is 2 ail be km SSE of Islas Buena Vista in the Rio Colorado, 14 airline km SW of Barro del Colorado; 83°40'W, 10°40'N, 10-120 m, Davidse & Herrera UOLKiO MO) i .ternmost ridge of Cerro Coronel,

NW-facing slope, just S of the Rio Colorado, 10°40'30"N, 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 al. 5612 (MO).

22.2. Tragia fendleri Muell. Arg., Linnaea 34:

178. 1865; in DC, Prodr. 15(2): 928. 1866.

TYPE: Venezuela: Biscaina, Fendler 1208 (G).

th .... • -In r ii.Hi i in, |,M!,MJ ,, pedicels ca. 1.5-

2 mm long, articulate near the base (stumps re-

maining after dehiscence of flower much shorter

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

flowers with hispidulous pedicels up to 2.5 mm

long in fruit; sepals 6, lanceolate, asymmetric, green,

reflexed in fruit, becoming 3.5-6 mm long, 1-1.5 mm broad; ovary densely hispidulous with stinging

hairs; styles basally connate or nearly free, 2-2.5

mm long, distinctly papillate. Capsules copiously

hispid with stinging hairs, cocci ca. 8 mm long;

columella 2.4-2.7 mm long; seeds globose, mottled

brownish and gray, 3.6-3.7 mm across.

The single specimen of this species, previously

unrecorded from Panama, is in poor condition and

without flowers, so there is some doubt regarding

its assignment. The Bristan collection matches a

photograph of the type specimen from Venezuela,

although the basal leaf sinus is not as open in the

Panamanian plant. There is also some resemblance

to T. japurensis Muell. Arg., described from Am-

azonian Brazil. However, it seems probable that

the Brazilian species is synonymous with the one

from Venezuela; at least, no convincing differences

are given by Pax & Hoffmann (1919: 36).

lined. PANAP

I mostly nonstinging h

22.4. Tragia correae Huft, sp. nov. TYPE: Pan-

ama. Panama: Picada da Estrada Panama-

San Bias entre 320-420 m, 9.1.1973, Sucre,

Braga, Dressier & Correct 9832 (holotype,

RB-165572, F neg. 62359).

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

dense hispiduloso, sivli- paj.ill.i11>.

Twining woody vine; twigs reddish, densely pilose, tardilv fzlaln.il.-. ihe older twig- with loose.

exfoliating bark, leans with petioles 0.5 4 em

long, densel) pilose; stiptiliv- delta acute, () 10 nun Ion-, pilo-e below. -I

blades membranous, elliptic-oblong, tip. cordate at base, 6 12 cm long, 3 4.5 em broad, 2.2-2.7 times as long as broad, sparsely pilo-e above, more densely so below, usually 7-nerved at has.-; margin- remotely denticulate (teeth 18-22 on a side). Inflorescences opposite the leaves, racemose, ea. 3 cm long (immature), 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.

ca. 1.3 mm long, ea. 1.2 mm broad, lu-piduhm- without, cucullate; stamens 3, the filaments thick and tle-li\ . tree. ea. 0.8 nun long; anther- elliptic.

0.2-0.3 mm long, extrorse. Pistillate flowers with pilose pedicel- ca. 2 mm long; calw lobes 5. lau- eeolat.-. acute, ca. 3 mm long; ovarv densely ln-- |)idulou- with stinging hairs, the styles free to the base, spreading, papillate, ca. 2 nun long. Mature

In aspect Tragia correae resembles the species that Pax & Hoffmann (I<)1<)) placed in section Hut, particularly such species as '/'. fcna'lcri. T.

japnrensts Muell. \rg.. and '/'. flillux Muell. Arg. The new -pe, le- i- e\«ln,|e,| Iroin that affinity,

however, by its racemose rather than bifurcate laminate Mowers with three (vs.

ens. The entire sepals and extrorse ate its placement in sect Tragia

1919). This species bear

volubilis but differ- b\

densely pilose leaves, and

.III- dis

species for Profesora Mireya Correa of the I m- versity of Panama. Tragia correae is known only from the type collection. Dra. Correa has kindly searched for a duplicate at the herbarium of the I niversitv of Panama (PMA), but so far, unfor-

ateK.

ipia I.., Sp. PI. 1054. hamnia scandens L

c/mmpta (Euphorbiaceae) I Bot. 9: 272-278. & A. HERZIC. 1984.

PAX, F. & K. HOFFMANN.

cliauipicae. ///. A. En

»uri Bot. Gard. 71:341. Euphorbiaceae — Dale-

litor), Das Pflanzenreich

\\'rust i u. (J. I.. & B. WKHSTKR. 1 972. The morphology

phorbiaceae). Amer. J. Bot. 59: 573-586.

As a result of additional collecting in Panama, the species of Ihilcchampi a are m,w , on-iderabl\

better unilerstood. and the number of species has

key. The order of species has been modified to reflect better then -\-lemalic relationship- ( \\ eb

unpubl. synopsis).

Stem tips and inflorescence- den.-el\ golden-lin -

iaJ]

< oil MI tmvolueral bracts narrowly spathulate and apically mate involucel free) 1. D. sha,

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. Stigma 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.

Ann. Missouri Bot. Card. 71: 341. 1984.

Tragia shankii A. Molina, Ceiba 11: 68. 1965.

TYPE: Costa Rica. Limon: Rio Reventazon, 15

m, 23 Oct. 1951, Shank & Molina 4427

(holotype, F).

This species, mil de : mbed from Costa

Rica, has now been recorded from Nicaragua to

Colombia. It may easily be distinguished from our

other Panamanian species b\ it.- large and diminu-

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

i\W;:--|.-i (N. \ri,il,i(!-i i-. iii.-i;.;. long with the two

lollo in I'anai HI I \;i I In i ollt i tioiis ol Bai -

ry Hammel show that D. shankii is polymorphic

in leaf shape, since leaf blades from I he single

locality vary from unlobed to having one lateral

lobe to three-lobed. Additional collections from Cos-

ta Rica, Panama, and Colombia are cited by Huft

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).

23.2. Dalechampia canescens Kunth subsp.

friedrichsthalii (Muell. Arg.) Webster &

Huft, stat. mi'

Muell. Arg., Flora 55: 45. 1872. TYPE: Nic-

aragua. Rio San Juan: Friedrichslhal 683

(not seen; locality erroneously cited by Mueller

as Guatemala).

Several additional collections of this plant have

d. ml • ' in . * nnli - A Smith 11,0

(MO); Rio Fato, Pittier 3866 (GH, NY, US); Rio Bo-

queron, 6-8 km upstream from Peluca Hydro Station, Siri Km,, (DAY), SAN ISI AS: Puerto Obaldia, sea level, Knapp & Mallet 4627 (DAV, MO).

It now appears that Dalechampia friedrichs-

thalii is excessively close morphologically to I).

th (Nov. Gen. Sp. 2: 98. 1817).

rom San Bias in particular seems

biologically to treat the tw

replacing subspecies of a :

subspecies may be keyed (

ngle species. The two

Leaf blades mo-tl- ; i pubescent beneath (many haii long, overlapping in sinuses b

The Colombian taxon, subsp. canescens, was

described from Tolima Province (Mariquita) and

cited from Narino by Pax & Hoffmann (1919: 52).

The additional collections cited below indicate that

subsp. canescens is widely distributed in the low- lands and foothills of the western Andean region

Sfx-cimrris cXtituilied C Mill \. • \l.l

Yeguas, 20 km N of Honda, MH) m, Cent, i (DAV, MO), CAUCA: Rio Patia, 590 m, Vaughan 5354 (DAV). SANT\M>KK: 29 k

23.3. Dalechampia dioscoreifolia IWppig

in Poeppig & Endl., Nov. Gen. Sp. PL 3: 20.

1841. TYPE: Peru. Maynas: Poeppig 2163

One additional locality merits noting: l'\\\\l\.

DARIEN: near Rio Canglon, Duke & Bristan 378

(MO).

An unusual specimen from Playon Chico, San

Mas(r,Y///M (>3(>5. MO) appears lo be mt

between D. dioscoreifoiia and D. canesc

if i ,!n, 11.i-, iiii; the broad stigi

former and the pubesc<

leaves of the latter.

23.4. Dalechampia websi

Syst. Bot. 9: 272. 1984. TYPE: Costa Rica.

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

the single record that was attributed (with doubt)

to I) rissifolia in our treatment of 1968.

5: 299, tab. 8. 1784.

Subfamily III. CROIONOIDKAK Pax

26. Tt'trorchidium

I < iroi i Indium I ' . I Nov. Sp. PI. 3: 23, tab. 227. 1841. TYPE:

ew province records \

.5. Dalechampia cissifolia Poeppig subsp.

panamensis (Pax & K. Hoffm.) Webster,

Ann. Missouri Bot. Card. 54: 193. 1967. I),

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.

Chiapas: Escuintla, Donnell Smith 2079.

Panama: Oersted; Pittier 2311, 3775.

This species still requires ,u

variant with simple, unlobed lea

c n:ii|M»uiid i>nrs, lo wliirli I In- name / .hi irriuan pm

heteromorpha Pax & K. Hoffm. has ben n: IS.

occurs in Panama am: ill*.- i-'i ml < Antral \mcrica

but does not appear in be s(„-rib. a b di unci from

There is one new provincial record for D. cis-

Additional specimen examined. PANAMA. BOCAS DEL TOKO: 10 mi. NW of Almirante, D'Arey llL'dl I (MO).

24. Omphalea

24.1. Omphalea 1377. 1763. r.

sumably BM, no

with persistent swollen stipules; less than 7 cm long, the paired glands I the apex of the petiole 1. T. microphyllui

2. T. costar Stems smooth, the apex appressed-pubescent o glabrescent; leaves mosth more ilian , i m lone

study- A long. 3a. Basal foliar glands ea. ()..r> ().(. mm tin. k.

moslb 1T> cm long or more, with o or 7

sessile ' 3. T. euryphyllum lib. liasal foliar glands ca. 0.15 0.3 mm thick,

attached at junction of lamina and petiole

li l.li \. II pi III! It! I .... I . lit, i I pedicels mostly 1.5-2 mm long

.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.

, praeter apicem dense liirsntinn, ghilnis; loliis

25. Vera

Pera Mutis, Kongl. Vetei

anches brittle, gla-

ensely hirsute, ap-

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 0.6-0.8 mm long, 0.8-1 mm thick, stipules glan- duliform, tumid, broadly triangular, 1.5-2 mm long, densely pubescent, persistent, glabrate soon after leaf-fall; blades chartaceous, oblanceolate, acuminate at tip, cuneate at base, (2.5)4 5.5(-7) cm long, (0.9-1)1.2-2 cm broad, sparsely pubescent with malpighiaceous hairs to glabrate on both sides, the midrib and primary veins (3-4 on a side) prom- inently raised below, the veinlets forming a prominent reticulum; margins entire. Inflorescences ax-

long, the rachis densely strigose; pistillate inflorescences unknown s / s ^ubsessile; calyx lobes 3, triangular, glabrous, ca. 1-1.2 mm long; petals lacking; anthers subsessile, 0.8-0.9

This distinctive new species of Tetrorchidium appears to be most closely related to T. brevifolium Standley & Steyerm., described from the province of Alta Verapaz, Guatemala (Publ. Field Mus. Nat. Hist., Bot. Ser. 23: 126. 1944), from which it differs by the smaller leaves (7-12 cm long in T. brevifolium), densely pubescent shoot apices and inflorescences (both glabrous in T. Inc., 'hum) and shorter inflorescences (4-7 cm long in T. brevifolium). The type specimen of the Guatemalan species (Rubelpec, Finca Seamay, Wilson 188, F) lacks the persistent tumid stipules that are so characteristic of T. n > ill these are present on several collections of the former species made in 1974 and 1975 from Baja Verapaz, Gua- temala (Lundell & Contreras 19173, 19436 (both

F, LL); milifims et al. 13277, F). The type spec-

leaves and stamiiiiite inflorescences that appear to

have been taken from rapidly growing long shoots.

, described from (loud forests in the inoimtains above San Juancilo.

Honduras (Fieldiana, Bot. 29: 348. 1961, based

on Williams & Molina 170()8, F), is similar in all

respects to T. brevifolium and should be relegated

to the syiionvmv of that species. A pa rat \ pe spet -

imen of T. molinae from the same area. U illiams

& Molina 13980 (F), has long shoots without

stipules attached to a normal -hoot with persi-tent

stipule-. I hi- inalche- the pattern of the type col-

lection of T. brcri/olii/rii. thus . onfirming the sus-

pi. ion voiced above concerning the nalme of that

Additional specimen examined. PANAMA. CHIRiyni: end of road past Palo Alto NE of Boquete in forest along ridge, 0.21)0 (,.«()() It.. H„mmci(,t>:iO(Y, MO).

26.2. Tetrorchidium costaricense Huft, sp. nov. TYPE: Costa Rica. Puntarenas: Cordillera

de Tilaran, Monteverde Reserve, near Con-

tinental Divide on Pacific side, 1,520 1,580

m, Dryer 1403 (holotype, CR, F neg. 62351;

isotypes, F, F neg. 62350, MO, F neg. 62349).

ii brevipedicellatis, segmenti.- disci .1

Dioecious tree to 16 m; branches densely stri-

gose tow.iid tip. appealing kuolili\ from the raised

leal s, .ii- and oo, asionallv from persi-tent. indili

ate stipules. I.raves long-petiolate, not crowded toward ends of branches; petioles 2.5 (> cm long,

glabrous or minutely strigose, with paired, -ubop

posite, sessile, patelliform glands near the middle.

these ca. 1 mm (rarely to 2.2 mm) diam.; stipules oblong. 1.5 2.5 mm long, 1 1.5 mm broad, dense

ly strigose, persistent, sometimes indurate aftei

leaf fall; blades membranous, narrowly oblong, 8

16 cm long, 3-7 cm broad, 2.2-3.3 times as long

as broad, abruptK cuspidate al lip with an acumen

5-10 mm long, acute to attenuate a I base. inmiitelv

puberulent below will) -cattep-,| short malpighia-

ate. prominent below, obscure above;

tire, .-glandular, Inflorescences axillary,

•nsel\ slrigo-e with -hort malpighiaceous

minate thyrses 1 9 cm long, freely

I isiillati i accrues to 5 cm long. Stanunatc lloiccis

in glomerules ol 2 5. sessile; sepals 3, obovate.

cucullate, glabrous without, pilose within: anthers

subsessile, 1.3-1.5 mm broad. Pistillate flowers

subsessile. the pedicels heeomiug 0.5-3 mm long

in fruit; sepals broadly ovate, obtuse, 3 3.5 mm

long, glabrous or sparingly short-strigose without,

densely hispid toward base within: disk segment-

free, narrowly ligulate, ca. 2 mm long; ovary

smooth, 2-locular, glabrous above. den-civ long

slngo e below: st\le eap at maluntv 0.5 0.8 mm

high, 1.5-1.8 mm diam. Capsule 3-5 mm high,

ovoid-lenticular. 5-6 mm long, proniineniK and

coarselv reticulate, the caruncle an irregular vellow

papery keel running halfwav from the Inluni to ihe

Known only from Costa Rica and extreme west-

ern Panama, this distinctive species is easily dis-

tinguished by the large, dark green, lanceolate

leaves that are conspicu.»usl\ venose, the paired

glands near the middle of the rather long petioles.

and the densely and minutely strigose braiichlel

tips, petioles, and leave-.

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

of long petioles, but differs in its completely gla-

brous stems and leaves, mostly unbranched sta-

iinnate tin rses thai have larger glomerules. densely

pubescent pistillate calve.--, and di-tuicil\ pedicel

late fruits. Tetrorchidium brevifolium differs in its glabrous stems, leaves, and calyces, short petioles

with paired glands near the tip. and unbranched

staminate thyrses. The widespread South \inerican

species T. rubrivenium Poeppig also belongs to this group but has idahroiis -|, rns ,\w>\ leaves, crenate or denticulate leaf margins, sometimes lone petiole-

wilh the paired gland- near the lip. densely pu-

bescent pistillate calyces, uniformly puberulent

ovary and capsule, and pedicellate fruits.

M vll 1,1 v. Cordillera de Tilaran, Monteverde IvY-erve. Ulaiiiie -ide. 1,500-1,580 m, Dryer 1071 (CR, Y).

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, 1,600-1,620 m, Dryer 659 (CR), 887 (CR. 10. M.»,, teverde Reserve, at field station, 1,500 in, l/aher 49 I (10: Monteverde lie-ene. 1,570 til. //,//.,•/ S. Hell., InlU if), 1.500 ,„, Uaber & Bella 2457(F). ALAJI KI \ ITVI v- 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); Kortuna Dam region, along Continental Divide W of highway pass, ca. 8°45'N, 82°15'W, ca. 1,200 m, McPherson 9695 (K).

26.3. Tetrorchidium euryphyllum Stand-

ley, Publ. Field Columbian Mus., Bot. Ser. 4:

219. 1929. TYPE: Panama. Bocas del Toro:

vicinity of Almirante, 1928, Cooper 621 (ho-

lotype, F).

When the original treatmenl was written, this

>|icnrs was known only from Costa Rica and ex-

treme western Panama, hut recent coll i n

[ono: between Quebrada and Buena Vista, Klrkbride & Duke i)i,2 (MOl. < HiHloi i: Koriiina Dam region, along Quebrada Arena, ca. 8CI5'\. 82°I5'\V, ca. 1,100 m,

n 8 194 (K). cot 11.: slopes of Cerro Pil6n near 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 (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 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); ', nta Ke on road past agricultural school, 2. 11216 (MO).

nt • Rita Ri \% Roa< I I an I ransisthmian Hwy., Dwver X Gentry 9338 (MO, 2 sheets); Santa Rita Ridge Road 4 mi. from Transisthmian Hwy. to Agua Clara weather station, ca. 500 m. Gentry el al. 8841 (MO, 2

300-700 ft., Hammel 4894 (MO). PANAMA: El Llano- av. 17-20 km N of El Llano, Dressier 4629

(K, MO); 10 km N of Margarita on road to Madrono,

(MO); Cerro Jefe region 2.5 mi! N of'tur tower along road, 2,400 ft., Hammel (,300 (MO); Kl Llano-Carti road, 9.6-11 km from Inter-American Hwy., 350 i„. Man * Kallunh 3531 (MO): 5 10 km NKof Altos de Pacora on trail at end of road, 700 800 m. Mori & Kallunki 6058 (MO); El Llano-Carti road, 8 km N of Pan Am. Hwy. at El Llano, ca. 450 m. Ace & Warmbrodt 10391 (MO).

27. Manihot

Manihot Miller, Card. Diet. abr. ed. 4. 1754.

TYPE: Maniliol esciilcntu Crantz (Jatropha

manihot L.).

26.4. Tetrorchidium gorgonae Croizat subsp.

robledoanum (Cnatrec.) Webster, Ann.

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-

tions allow a description of the fruit to be made

for the first time. They are on densely strigose

pedicels 7-10 mm long and jointed below the mid-

dle. The capsule is green, drying to brown, globose,

shallowly 3-lobed, rugulose, 4-6 mm high, 4.5-7

short (0.2-0.6 mm) malpighiaceous hairs. The 3

styles are deeply bifid, 0.6-0.7 mm long, and tu-

mid. The seeds are ovoid. 1 5 nun long, 3-4 mm

metric methods used. Econ. Bot. 27: 1-112

The discovery of an additional I'amiir

ia. weaves giouruus, muouj wim .-* IUUCS, ' «•• glabrous within (12-14 mm long; disk e

Leaves pubescent or v

2a. Leaves pubescent,

3. M. brachyloba

27.1. Manihot aesculifolia (Kunth) Pohl, PL Bras. Icon. Descr. 1: 55. 1827. Janipha aes-

culifolia Kunth, Nov. Gen. Sp. 2: 85, tab.

109. 1817. TYPE: Mexico. Campeche: Hum-

boldt & Bonpland (P, not seen).

This plant was called XL gualanensis Blake in

t monograph of the genus hy Rogers & Appan

27.3. Manihot brachyloba Muell. Arg., Fl.

Brasil 11(2): 451. 1874; Rogers & Appan,

Fl. Neotrop. 13: 190-192. 1973. TYPE: Bra-

zil. Para: Marlins (syntype, G; microfiche

i specimens have

vegetatively matt

i given by Rogers

I i ,,1, i . I i i f Rio Tuqueza, between Quebrado Venado and Peje wamp, Bristan 1001 (DAV, MO); between Manene and 'joCoasi, Hartman 12127 (MO).

Cnidoscolus Pol,I. I'l. Ihusil. bun. Descr. 1: 56.

1827. EECTOTYPE: Cnidoscolus hamosus Pohl

(chosen by Small in Britton & Brown, Illust.

Fl. N. U.S. ed. 2, 2: 462. 1913).

28.1. Cnidoscolus urens (L.) Arthur, Tor

reya 21: 11. 1921. Jatrupha urens L., Sp.

PI. 1007. 1753. TYPE: "America calidiori, in

Brasilia & c." (not seen, possibly in Hortus

I lim.rli i ii- Herbarium, BM).

leads

Since the treatment of I"',., h, lr

e Panamanian specimens of ('.. uren

e conclusion that the two variants discussed there

?rit taxonomic recognition. Pending a more de- iled revision of the C. urens complex (Breckon,

28.1a. Cnidoscolus ,

Specimens examined. PANAMA. PANAMA: 1 Beach area, Correa et al. 1586, Dwyer 3065, 1803 (MO). coa.F.: between Aguadulce and San t Woodson et al. 1226a (MO); between Anton and 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).

28.1b. Cnidoscolus urens .subsp. adenophi-

lus (Pax & K. Hoffm.) Breckon, stat. nov.

Jatropha adenophila Pax & K. Hoffm.,

Pflanzenreich IV. 147. VIRHeft 63): 409.

1914. Cnidoscolus adenophilus (Pax & K.

Hoffm.) Pax & K. Hoffm., Nat. Pflanzenfam.

ed. 2, 19c: 166. 1931. TYPE: Panama. Pa-

nama: Chepo, Pit tier 1740 (isotype, US).

flores Locks, Stem et al. 81 (MO): Pipeline Road. ( mat 12732 (MO); Curundu, McDaniel 5180 (MOl. /,>..« !<>!., (MO); l|. \„ui,io, Wands. Issnn 5113 (MO). 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 (M<M; between Las Margaritas and Rio Mamoni, I hike 3Ht>7 (MO); Puente de Pacora, De floyos 18 (MO), \hK\..i\s: 2-4 mi. E of Santiago. I hike 123., 1 (MO); 12 km E of Santiago, Dwyer & Kirkbride 7450 (MO).

28.2. Cnidoscolus aconitifolius (Miller) I. M.

Johnston, Contr. Cray Herb. 68: 86. 1923.

subsp. aconitifolius. jatropha aconitifolius

Miller, Card. Diet. ed. 8. 1768. TYPE: Herb.

' :, »' ' i " rsimiahK at I>\1. 11< • I seen).

In the present interpretation, the Panamanian .ii.i •> lis all belong

lo the nominate subspecies, winch is not native to

29. Jatropha

L., Sp. PI. 1006. 1753. I.ECTOTYPE:

Jatropha gossypiifolia L. (see McVaugh,

. Bot. 09: 128.'{ I 2«>:>. . 1984. Phylogenetic significance ol inlerspe-

ciiic Inhndi/ation in J,ilro/>li,i (Kuphorhiai cae). SvM. Bot. 9: 467 478. & G. L. WKHSTKR. 1979. Morphology and

mlrayencnc relalioiiships ol the genus Jain,/,ha (Eu-

phorbiaceae). Univ. Calif. Publ. Bot. 74: 1-73. cVAl-CH, R. 1944. The genus Cnidoscolus: gencne

71: I1.: 174.

Pausandra Radlk.. Flora 53: 92, tab. 2

TYPE: I'alisiintha nioiisiunn (Casar.)

(Thouinia morisiana Casar.).

Webster & Huft

nate, simple; petiole swollen distally; blades pin-

nately veined, biglandular at base, the margins

serrate. Inflorescences axillarv. spiciform; stami-

nate flowers in glomi I ules ; pistillate flowers solitary

at each node; bracts inconspicuous, eglandular.

Staminate flou.t- -ul>~.—ilr: . al\x lobes 5, im-

bricate; petals 5 or rarely 6, connate (at least

below), adaxially villous; disk extrastaminal, ur-

celoate, lobate, glabrous; stamens (3-)5-7; filaments free; anlli. <l •! i « nv n .i . i.d .i

gitudinally, the connective not enlarged; pollen

grains globose, inaperturate, clavate; pistillode ab-

sent. Pistillate flowers subsessile; sepals 5, imbri-

cate; petals 5, free, adaxially villous; disk urceolate,

- i i I i'ii I I.ton-; d\ary of 3 carpels;

ovules 1 per locule; styles free, bifid. Fruits cap-

sular; seeds smooth, carunculate; endosperm co-

pious; cmbrvo straight, cotyledons palmatinerved,

much longer than radicle.

This primarily South American genus is here

reported from Panama for the first time.

BAIIION, H. 1873. Nouvelles observations sur les Eu- phorbiacees. Adansonia 11: 72-138.

LANJOUW, J. 1936. The genus Pausandra Radlk. Re- cueil Trav. Bot. Neerl. 33: 758-769.

30.1. Pausandra trianae Baillon, Adansonia

11: 92. 1873, proposed without reference to

f'o^oiK'fihon! tf:a>:ir \ii:•."!•.. Arg. TYPE: Co-

lombia: Bogota, pla ol San Martin, RioMeta,

Triana 2597 (holotype, P, not seen; isotypes,

G, not seen, photo F neg. 24574, K, not seen, holotype of Pogonophora trianae Muell. Arg.).

Pogonophora trianae Muell. Arg.. Flora 47: 434. 1864. TYPE: Colombia. Bogota: plains of San Martin, Rio Meta, Triana 2597 (holotype, K, not seen, isotypes, G, not seen, photo F neg. 24574, P, not seen, holotype of Pausandra trianae Baillon).

lulosa Pax & K. Hoffm., Pflan- zenreich IV. 147. XIV(Heft 68): 43. 1919. TYPE: Brazil: Rio Acre, Seringal S. Francisco, I /<• 9.5.W (holotype, B, not seen, photo F neg. 5406).

indley, Trop. Woods I " ausandra e 1929; Publ. h I. f ,.!.„„ ,i.„, \lu , But S.-r. Oct. 1929. rvi'i;: Nicaragua: Bragman's Bluff, En- glesing 216 (holotype, F, F neg. 52719; wood sam- ple, Y no. 13301).

Imtrionalis L. (). Williams, Fieldiana. B;>1. 32: 205. 1970. TYPE: Nicaragua: Cabo Gracias a Dios, Laimos Creek, ca. 15 km SW of Waspam, 7 Mar. 1961, Bunting cv Lieht 390 (holotype, F; isotypes, NY, US).

berulent with malpighia

r.s horizontally aligned; petiole 1.5-

3.5 cm long, 2-4 mm thick, terete, strigose with

short (to 0.5 mm long) malpighiaceous hairs; glands

at apex of petiole (3 )4, cylindrical, 1-1.5 mm

• I i iilate, rounded, obtuse, or abruptly short-

cuspidate at tip, long-attenuate at base, 20-50 cm

long, 7-18 cm broad, 2.6 4 times as long as broad,

shorl HI dpighiaceous hairs below, the secondary

\ in prominent, 15-23 per side; margin remotely

denticulate. In/Ions,,„, , s -pi. ale an- M IJ

in the upper axils, 10-25 cm long, the rachis

densely puberulent. St ami nate flowers 6-12 per

glomerule, these sessile, widely spaced; sepals 5,

imbricate, densely puberulent, obovate, ca. 1.5 mm

long, ca. 1.5 mm broad, rounded at apex; petals

brous without, densely hirsute toward bast within:

disk cupulate. Pistillate flowers not seen. Capsule

smooth, strigose with short malpighiaceous hairs,

apparently subglobose, ca. 1 mm diam. (fragments

9 mm long, c

I I OIK:! i ra-. i

The description is based on the Panamanian

specimens cited below and supplemented by col-

lections at F from Costa Rica and Nicaragua.

The genus Pausandra remains poorly understood. The most recent revision (Lanjouw, 1936)

was based upon only 33 collections and resulted

in the recognition el rune >pccie>. most known from

only a single sex. Lanjouw admitted that several

of the species might need to be united as more

material became available. Pausandra trianae dor-

appear, however, to be one of the better-delimited

species in the genus and is certainly the most

widespread. The Central American plants readily key to that species in Lanjouw's revision, where

P. extorris and P. quadriglandulosa are reduced to synonymy. The identity of C.lavija septentrio-

nalis as P. trianae was first pointed out by R. L.

Liesner of the Missouri Botanical Garden (pers.

The author citation for Pausandra trianae lias

been almost universally given for the last century

as Pausandra trianae (Muell. Arg.) Baillon, car-

rying the implication that Baillon had transferred 11 - an trianae Muell. Arg. to its correct

place in Pausandra. It is (dear from Baillon"s text.

Mueller's name,

and none would he c\pe. led. One would not nor-

mally look in I'oiiuiiojihoKi lor species pertaining

to Pausandra: the j;i'iicra air too different lor

: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 lowland rainforest on the Aila Tilar (Hi.. Via), iTIH'Y 77°40'30"W, 25 KM) ,„. Sullen 121. 5fiti [Mi)).

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

hut not \cr\ surprising, given the common practice

ol iuiniiiiii species after llie colic, lor ol the l\pe.

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

supplanting Baillon's name. But thai solution is

closed because it would create a later homonym.

Article 63 of the Code (Voss, 1983) might mislead

one to reject Baillon's name as superfluous, hrcnuse

it seems to be based on the type of a name whose epithet ought to have been adopted under the rules.

A correct reading of the (lode, however, makes it

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

31.1. Garcia nutans Yahl in Hobr.

turhist.-Selsk. 2: 217, lab. 9. 17<

Colombia. Magdalena: near Santa M

Hofn (C. not seen).

Croton L., Sp. PI. 1004. 1753. LECTOTYPE: ton aromatieus L. (chosen by Webste,

Arnold Arbor. 48: 354. 1967).

Seven species new to Panama (incl tiding lv new species and one new subspecies) an • report. here; a few new province records are in d.cated i well. These additions make it necessarv to provi.

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. ('.. /><:. </•, ',<

2b. Indumentum not lepidote, the trichomes mostly or entirely stellate or dendritic; petals in pistillate

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; stamens 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 mostly 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

logia 1: 451. 1940. TYPE: Costa Rica. Sai

Jose: vicinity of El General, Skutch 2575 (ho

lotype, MICH).

): along road to Chiriqui Grande, 10 road mi. fron

32.5. Croton lanjouwensis Jabl., Mem. New

York Bot. Gard. 12: 158. 1965. C. matou-

rensis Aublet var. benthamianus Muell. Arg.,

Linnaea 34: 95. 1865. C. benthamianus

(Muell. Arg.) Lanjouw, Euphorb. Surinam 17.

1931, non C. benthamianus Muell. Arg., Fl.

Bras. 11(2): 106. 1874. TYPE: Brazil: Rio

Negro, Spruee Croton 2 (isotype, NY).

Tree to ca. 12 m high; monoecious: twigs deiisdv

lepidote. Leans wild [.choirs lepidote, 1.5-3.5 cm

long listal • ill II iiit • i.wish, subsessile,

patelliform glands ca. 1.5-2 mm across; stipules

linear-lanceolate, entire, densely lepidote, 8-10 mm long, early deciduous; Ida*

liptic to elliptic-oblong, acute

at tip, cuneate at base, mostly 8-15 cm long, 3.5-

6.5 cm broad, s < il ml II. <>us above, densely

l< I .1 ;. beneath (scales denticulate-margined, ca.

0.25-0.4 mm across and nearly or quite contig-

uous); venation distinctly pinnate, the major lateral

veins (ca. 11-15 on a side) straight; margin* entire

.ri.: If| dole margin, h- /;;//."- r.s, , nccs terminal.

racemose, 13-15 cm long, In MI I; ( n ij. ! ».

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;

receptacle villose; petals obovate or narrowly ellip-

tic, ca. 2 mm Ion du n tali . detiscK

villose on margins, glabrous on back: stamen,, 12

or 13; filaments glabrous or sparsely hirMitulou.-;

anthers elliptic, 0.6-0.7 mm long. Pistillate flow-

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

(shorter than 1 mm long); ovary densely lepidote

ca. 4 mm long, stellate proximallv (brandies gla

lii. Capsules not seen entire; cocci lepidote

ca. 5 mm long; columella slender, ca. 4 mm long;

3.5 ,

The Panamanian plants appear to agree in most respects with C. lanjouwensis as defined |,s | ;,n

jouw (1931: 12-17; as C. benthamianus); the ii I i lillate calyx with adaxiallv stellate lobes

is apparently diagnostic in separating the species

from C. matourensis Aublet. However, the Pan- i M.I i plants occurring in cloud forest at 800

1,000 m would appear to differ ecologically from

the Amazonian plants, which have been colic, led

in lowland rainforests. Jablonski (1965: 157-158)

\ ei • , nd; . ,. Pan-

needed. The specimens from Panama Pros in. e dd

fer rather strikingly in their duplex petiolar glands,

sparsely lepidote upper leaf surfaces, and shorter

Valle, ca. 900 m, Duke & Correa 14718 (DAV, MO); cloud forest, hills N of El Valle de Ant6n, Dressier 4083 (DAV, MO); cloud forest, El Valle, 800-1,000 m, Duke 13166 (DAV, MO), Dwyer & Correa 7951 (DAV, MO), Correa 311 (MO). PANAMA: primary forest, road from El Llano to Carti-Tupile, 200-500 m, Croat 22905 (DAV, MO).

32.6. Croton pachypodus Webster, sp. nov.

TYPE: Panama. San Bias: km 18 of El Llano-

Carti road, 9°19'N, 78°55'W, 350 m, 1 Oct.

1984, de Severs & Herrera 3980 (holotype,

MO; isotypes, DAV, F).

Tree 7-25 m high; twigs obtusely angular, gla-

brate. Leaves with petioles lepidote, 0.8-1.5 cm

long, distally (near base of blade) with 2 subsessile

short-stipitate (to 0.7 mm) blackish patelliform

' lepidote, 4-10 mm long, 0.9-1.2

rids 0.5

; blades chart;

tic-oblong to somewhat obovate, subacute or acute

to abruptly short-acuminate at tip, cuneate to

rounded at base, 8-22 cm long, 3.5-7 cm broad,

smooth and glabrate above (with sparse scales on

l young), evenly and sparsely lep-

i (scales 0.25-

ca. 50 radii), the scales widely sep-

on disiineiK pinnule, the major laleral

>n a side) straight or slightly curving,

glabrous; petals obsolete, represented by whitish

tufts of hairs; ovary densely lepidote (scales 0.5-

0.8 mm across, denticulate, 50 70-radiate, with

20-30 darkened radii); styles blackish, twice-bifid,

3.5-4 mm long, nearly glabn . Ca\ ."•'• D I een

entire; valves of cocci ca. 23-25 mm long; colu-

mella slender, ca. 20 mm long; seeds elliptic, some-

what compressed, flattened and obscurely carinate

on the back, distinctly keeled on inner face, grayish

brown, smooth and shining, 16.2-17.2 mm long,

10.3-11.3 mm si .

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

and with margins free of scales, shorter petioles

with smaller darker glands, and especially by the

much larger fruits borne on greatly thickened ped-

icels. Among species earlier reported Iron I'ai a ma.

the new species resembles C. tcnuicaudatiis; how-

ever, that species has eglandular leaves lepidote

on both faces, strongly lepidote staminate petals,

and more slender pistillate pedicels. Croton pyri-

ovate palmately veined eglandular leaves, longer

and more slender (1.5 mm or thinner) pistillate

pedicels, and verruculose capsules; however, the

seeds, although larger, are similar in shape to those

of C. pachypodus.

12 mi. above Pan-American Hwy., 200-500 r 22905 (MO); 20.7 km from Pan-American Hv m, Mori & Kallunki 5116 (MO).

inflorescences, staminate flowers solitary or paired

at distal axils; bracts triangular, blackish, sparsely

lepidote, 0.5-1 mm long. Si, | ud

only observed) with lepidote pedicels 1.5-2.5 mm

long; calyx lobes 5, valvate, triangular, lepidote,

ca. 3 mm long; receptacle villose; petals narrowly

elliptic, ca. 2.5 mm long, densely villose on mar-

gins, sparsely lepidote (often a single scale) on the

back, densely hirsutulous adaxially; stamens 14-

16, the filaments glabrous, the anthers 0.8-0.9

mm long. Pistillate flowers with stout lepidote ped-

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,

equal, triangular-ovate, reduplicate-valvate, dense-

ly lepidote without, densely loin a IOM \illosc with-

in, 3-3.5 mm long, 2.5-3 mm broad; disk shal-

lowly 5-lobed, nearly 4 mm across, smooth and

32.7. Croton speciosus Muell. Arg. [Linnaea

34: 83. 1865] subsp. tacarcunensis Web-

ster, subsp. nov. TYPK: Panama. Darien: Cerro

Tacarcuna, S slope, premontane wet forest on

ridge below summit, 1,250-1,450 m, Gentry

& Mori 13925 (holotype, MO; isotype, DAV).

Haec a subsp. s pecioso differt stipulis r.,m..M|,i-, _4m

dulis petioli brevioj ribus, carunculo seminis ca. 2 mm lato.

Monoecious tree 5 m high; t\ vigs subterete,

densely tawny-v illose with dendritic : hairs. Leaves

with petioles (2 -)3-ll cm long, t omentose, api-

cally with 4-6 i stalked glands on v< •nlral side, the

stipules lanceolate, densely tomento se, toothed, 3-

6 mm long; bl; ades membranous, mostly ovate,

long-acuminate at tip, rounded to subcordate at

base, the larger ones shallowly 3-lobed, 12-21 cm

long, 7-12 cm 1 broad; lamina abov< i copiously pu-

ellate-tufted hairs, beneath copiously Mueller ga\e the -tnmcii number <tl' (',. spaiosas

h whitish dendritic hairs ca. 0.5-1 as ca. 150. which certainly docs not agree with

mm across, 3-5-nerved at base, with 7 10 major Moritz 1.329, in which two buds yielded ca. 70

lateral veins on each side, connected by a scalari- and ca. HO stamens. Possibly the number may

form reticulum of straightish veinlets; margins sub- become higher in some Venezuelan plants, as Muel-

entire (obscurely denticulate). Inflorescences ter- ler (1866: 529) also cited two other collections

minal (or pseudolateral). bisexual, racemose, 2.5 from near Caracas. Fendler 31 and 231 (cited as

6 cm long, with 1 or 3 proximal flowers and 3 7 Linden 34 and 201), which we have not seen,

distal flowers; flowers solitary at each node; bracts The publication of a separate subspecies for the

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 peak of Cerro Tacarcuna, 1,100-

l.:!lHI,n. dent,, rial Hu'i7.' i M()): r..l«.-lo,. l.rkm Ml,, de Nique base camp, Gentry el al. 28727 (MO).

32.8. Croton draco Cham. & Schldl. [Linnaea

6: 360. 1831]subsp. panamensis (Klotzsch)

Webster, stat. nov. Cyclostigma panamense

Klotzsch in Seem., Bot. Voy. Herald 105.

1853. Croton panamensis (Klotzsch) Muell.

Arg. in DC, Prodr. 15(2): 546. 1866. TYPE:

Panama. Chiriqui: Volcan Cbiriqui, Seentann

(K, not seen), (see Webster & Burch, 1968:

254 for additional synonymy.)

Further examination of Mexican and Central

American specimens of C.ioton draco indicate- lfi.it

the Panamanian plant- cannot reasonably be main

tained as a separate -pecie- Kxcept for the larger, 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. <Wo from specimens colleCed nearCara, ,-. \ ene/uela. s|nm M(| t.^.nUai differences from plants with nar-

I have examined Moritz 1329 (A, CH) from Cali- stipules that occur

long in fruit); CJ ilyx segments 5, valvate, oblong,

densely whitish - to fulvous-tomentose without,

sparsely toment ose in distal third within, ca. 10

mm long, 3-5 n rim broad; disk inconspicuous, ad-

natetobaseofc; alyx, crenulate, stellate-pubescent.

ca. 5 mm across ;; petals rudimentary, densely hir-

sute, ca. 1-1.5 mm long; ovary densely fulvous- tomentose; style: s twice-bifid near the base, densely stellate-pubescei it below (and with scattered stellate hairs distallv ne; irly to tips), ca. 7 9 mm long, the

tips. Capsules si ibglobose, fulvous-hispidiiloiis. ca.

1 cm long and broad; seeds plump, plumbeous

brown, distinctly • costate ventrally, obscurely cos-

tateonback,ca. 7 mm long, 5 mm broad; caruncle

round, h low i. 2 mm across.

This striking Croton from the cloud forests at the , rest of Cerro Tac.m

Mueller (incorrectly located by him as in (iolomhi.

rather than Venezuela), as well as three other col-

lections from near Galipan {Allan s.n., Pittier221, 9377. A. CH). These plants in general rather closely

resemble the Cerro Tacarcuna specimens in leaf

shape and pubescence

(and spo- syntypes^c.ted by radica„y in southern Mexico) to Panama and Co-

32.10. Croton smithianus Croizat, J. /

Arbor. 21: 93. 1940. TYPE: Colombia

Mesa de los Santos, Killip &

ub, .ha, « an ,leal,„P ' "« ^"^ A; ^^ "S>-

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 a it base, the larger ones 15

5, 10-30 cm broad; i palmate

major vein s at base . 5-8 1, aterals 01

chotomizing towards t

ts prominulous on bo

riform; trichomes on

.,.,11, 1:11

IHLII t.cncatli denser and + floeeose: -mall palel-

hi: ii i idand- (0.1 I iniii arrows) occasional on

upper surface; margins denticulate, with occasional

small stalked glands. Inflorescences terminal, most-

ly 20-50 cm long; 5-15 proximal eyniules hise\-

late, entire, deciduous, ca. 1-2.5 mm long,

subtending sewral (lowers. Slaiiiinalc flowers with

-lellate lomentose pedicels 1.5 5 mm long; calyx distinctly gamopln lions. 3.5 1 mm long; calyx lobes

ovate, acute, valvate, 2.2 3.7 mm long, 2-2.8

mm broad; petals narrowly spathulate, 3.2-4.5

mm long. 0.5 1 mm broad, densely villose ven-

trally, strigose-hirsutulous dorsally; receptacle

densely villose; stamens 11 or 12; filaments flat-

tened, densely hirsutulous in lower >',. 3 1.5 mm

long; anthers elliptic, the connective glandular-pus-

tulate, 1.1-1.5 mm long. Pistillate flowers with

stellate-lepidote pedicels becoming 9-14 mm long;

calyx lobes 5. valvate (not distinctly reduplicate),

tip and along margins, 5-7.5 mm long, 3.5-6.5

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,

lenticular, brownish, finely costate-rugulose, blunt-

ly pointed at both ends, 3.9-4.1 mm long, 3.3

3.5 mm broad; caruncle flat, bilobed, 1.7-2.1 mm

Croton smit/ih ~ b, !< I i ,\ mil

montane forests, up to ca. 1,500 m elevation,

Nicaragua to Cob i- ii July to Septem-

The Panamanian representative of the wide-

spread and variable South American species com-

plex centering on ( rot mi K b i- heie re b Tree, n. < :>>••: I,,-;;,,, , « m/ai because

iri indumentum, and a specimen from < ihoeo

{Archer 2062, US) has the characteristic leaf form

and margin, but the trichomes are not di.-tmelK

pedicellate as in the Panamanian plants. The Pan-

Croton killlplanus Croizat, described from Bo-

yaca; however, the type collection of that species

(Lawrance 588; isotype, US) he ibentir leaf

margins and an appressed, rather sparse indumen-

tum more characteristic of C benthamianus Muell.

Arg. Croton niintians Croizat from Guyana is

somewhat similar but differs in its smaller fruiting

calyx and shorter fruiting pedicel. Until this -pe, ie-

to C. smlthianus. Several collections from Nica-

ragua and Costa Rica are also referred to that

Specimens examined. COSTA RICA. HEREDIA: Finca La Selva, Hammel & Trainer 12849, 13044 (DAV, III Kb), I-I M Mil \\>: forest remnants along highway 4 mi. SE of turnoff to Buenos Aires, 9°07'N, 83°17'W, 1(10 in. II chster 21883 (DAV, MO); rainforest 17 mi.

SE of San Isidro General, 700 m, Webster c\ Miller 12394 (DAV). NiCARA<a\. Rio SAN Ji AN: Sahalo, Aru- quistain 3223 (DAV, MO). ZEI.AYA: Bluefields, \, •,// 27,'tH (DAV, MO). PANAMA, COLON: Rio Salud, Howell 128. Lao & Holdndge 224 (MO). DARIEN: between Manene and Tusijuanda, Duke 13576 (DAV). PANAMA: SE slopes of Cerro Trinidad, Kirkbridc & Ihtkc 1665 (MO).

32.11. Croton billbergianus Muell. Arg., Lin- naea 34: 98. 1865; subsp. billbe17.ia.ms.

TYPE: Panama. Colon: Portobelo, Billberg 316

Additional specimens examined. F \NA\I \. w >< \S DI-.I

TORO: Siirsuba, Rio Changuinola, Divyer s.n (MO). -\N

itl,\s: Puerto Obaldia to ba lionga, knupp & Mallet 4667 (MO), VERACUAS: Coquyito mine to Rio Barrera, Hammel 5221 (MO); Santa be. Folsnm X Eduards 3392 (MO).

The description in the original treatment (Web-

ster & Burch, 1968: 257-258) applies only to

ib . her^ianiis \- suggested at that time,

C. pyramidalis J. D. Smith, extending lrom Ve-

racruz, Mexico, to Honduras, does not appear to

be a distinct species. It may be retained at the

subspecific level because of its apparently larger

seeds (5.7-6.2 mm in the Veracruz population vs.

4.3-5.5 mm in If.. I ".man an •.,!• plan: • : a, id I- aiL'ei

Volume 75, Number 3 1988

Webster&Huft 1123 Panamanian Euphorbiaceae

stipules (7-15 mm long vs. 5-7 mm in the Pan-

ican plant is therefore necessary.^

32.13. Croton hircinus Vent., Jard. Malmai-

nerved at base, the secondary veins 5-8 per side;

margin entire to remotely denticulate, Inflores-

terminal, the pistillate ones axillary, occurring only

l additional provincial record is cii

obre hri.lpe, 2,r. mi. W of Tole, Divyei

32.14. Croton santaritensis Huft, sp. nov.

TYPE: Panama. Colon: Santa Rita Ridge Road,

21-26 km from Transisthmian Highway,

tropical wet forest, 500 550 m, 9°25'N,

79°37'W, 4 July 1982, Knapp 5882 (holo-

type, MO; isotypes, DAV, F, F neg. 62353,

PMA). Figure 3.

Leaves -villi petioles ilenseK villous ;

5-20 mm long; petiolar glands sever;

petiole, stipitate. tnimpet-sliaped. 0.5

0.3-0.5 mm across; stipules ovate-d< branous, eglandular, appressed. 8-1

4-6 mm broad, the margins fimbi iate;

caudate at apex, rounded to subcorc

10-15 cm long, 4-8 cm broad, 1.7

long as broad, sparsely to moderatel)

bescent above, sparsely tomentose

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

bracts subulate, 2-3 mm long, stellate below, gla-

brous above; pistillate inflorescences 5-11 cm long,

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

flowers: sepals 5, debate, joined at base, valvate,

stellately pubescent, the lobes 2.5 3 mm long, ca.

2 mm broad; petals 5, only slightly exceeding the

calyx lobes, ca. 6 mm long, ca. 2 mm broad below

the tip, the tip abruptly expanded, ca. 2.5 mm

broad, coarsely erose; stamens 12-15; disk con-

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-

Because of its combination of pentamerous calyces

in both pistillate and staminate flowers, five petals

and glabrous receptacles in the staminate flowers,

ognized by Mueller (1866: 511-700) in

recent worldwide account of Croton. N<

closely resembling Croton santaritensis found among the large holdings of South /

Croton billbergianus s Smith) Webster, stat. nov. Croton pyn Smith, Bot Gaz. 35: 7. 1903. nn- <; Verapaz: Rio Dolores near Cubilguitz, Tw (holotype, US; not seen).

32.15. Croton brevipes Pax, Bot. Jahrb. Syst.

33: 290. 1903. TYPE: Costa Rica. Rio del

Convento, Pittier 12117 (isotype, US, photo,

FIGURE 3. Croton santaritensis -i Habit. , . • >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

pressed-slellate. Leaves alternate below, mostly opposite or teniate abo\e: petiole-. densef appreused

stellate, 3-20(30) mm lone (less than '/ length

of blade): petiolar glands (at base of blade) conspicuous. e\ lindrii al. apiealK Innicate and dilated.

1 2.5 mm long, 0.3 0.5 mm across; stipules subu-

late to narrowly lanceolate, dark, entire, eglan-

ihinK chartaceous. elliptic to ovate-elliptic or ob-

at base, 4- I 3 em long, (1 )2 5 cm broad. sparsely

stellate or appressed hispid abo\e with leu-rased

trichomes. sparsely appressed-slellato and ineon-

-ph nou-ly glandular-punctate beneath. pinnalely

\ en ied (or inconspicuous! \ I riplmery ed) with mostly

5 7 veins on each side: margins subentne to rather

coarsely am! irregularly deiil.ite (teeth ca. 8-15

on a >ide). with stalked viands between some of the

teeth. In/loieseem e.s iu.r-.lK terminal and bisexual

:;< i.:,ie •eU :>,,|,

pistillate flowers, the siaminale (lowers I or 2 per

bract at distal axils: bracts narrow, entire, eglan-

sliorler. Stnminate J/oicers with sparsely stellate

or nearly glabrous pedicels I 2 nun long: calyx

lobes 5. elliptic-lanceolate, acute, stellate-pubes-

cent, glandular-punctate, 1.2 1.7 mm long, 0.9-

I . I mm broad: recept.n le moderately \ illo-e; pelal-

olxnalc-spathulale. 1.1 1.8 iiim long, glandular-

mens 10-12; filaments glabrous, 1.8-2.5 mm long;

anthers ovate, 0.5-0.7 mm long. Pistillate Jlouers

with -lout appressed -lellale pedicel- heeoining I .'.',

3.5 mm long; calw lobe- 5. subequal, narrowly

. 3.5 i long. petal . glab, van sparsely stellate-pube-ccul apically. glabrous

clow; stvles free. ca. 2 mm long, twice bifid. labrous to hispidulous. Capsules subglobo.-e.

parseK -lellate-pubescent or glabrescent, ca. 5

mi diam.; columella slender, 3.2 4 mm long; seeds

roadly ellipsoid, compressed, apically beaked,

rownish. nearly smooth (minutely striolate), 3.8-

mall, ca. 0.5-0.8 mm across.

Rainforest below 1,000 m, Costa Rica and Pan-

larger laminar glands; tin- stipules, bracts, and ca-

lyx lobes lack the glandular serrations of C. hir-

dontiis Muell. Arg. from Mexico shows that it is

extremely close to ('.. hrevipes. Although -pecimen-

from Costa Rica and Panama may be easily rec-

ognized by their distally opposite, less coarsely

toothed lea'- e- with more rounded h.i-e- and -hortei

petioles, they are \er\ similar to the \le\i. an pi.mi-

ni most details, including pubescence, floral details,

and bulls. The seed- of the Mexican plants are

somewhat larger, but this difference ma\ di appear

upon further sampling. Provisionally, the two -p,-. i,--

may be kept distinct on the basis of the foliar

characters, and because no intermediate popula

i Costa lii.a . Mex-

Specimens examined. I'wun. ininv Nu.la Kit.. Ridge, ca. 300 in, •tnloiim /.Y.i'MD W, f, MO). Co,tea A Dressle, <) 12 (F, MO). Coat I Mi'lH (MO). Put,, /..."'/ (\K». I hi w7 ,->'.,/.•; | \IO), lh>,e, At.enlis ">'"-. If. Mm. haste, 177*1 (D\\, 1)1 Kf, f), Gentry 1X74 (1)\\. f. MO), ken ne,lYl>7.-,<> (WO), knnpp .',«•/.» <I>A\ . I. MO). SMsmu 20.17 (MO). 2()7>l (f, MO). II .•/,•,/,•/ A 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 (DAV, MO. I'S), Webster A Dressier 10477 (DAV, DfKf. MO. IS): T..rt[ \rr ,1M, /• olsom et ui r,o •/••/ I l)A\ . MO).

32.16. Croton argenteus I ., Sp. PI. 1004.

1753. Juloerntim argenteus (L.) Didr., Vi- densk. Meddel. Dansk Naturhist. Foren.

Kjflbenhavn 1857(8 10): 134. 1857; Muell.

Arg. in DC, Prodr. 1 5(2): 703. 1866; Croizat,

Revista Argent. Agron. 10: 125. 1943; Cor-

rell & Johnston, Man. Vase. Plants Texas

939. 1970. TYI'K: America (not seen; presum-

ably in Hortus Cliffortianus Herbarium, BM;

1140.8 in LINN).

Annual herb 2 10 dm high; stems pseudodi- chotomizing. with long internodes ,md psendover-

ticels of leaves, appressed stellate-pubei iilenl

Leaves with petioles 1 5 cm long, these without

paired apical gland-: -dpule- subulate, (2.5 )5 10

mm long; blades chartaceous, ovate or the upper

ones oblong-o

7( I5)c ong, 2.5-5 cm b

• or rounded to subacute

i broad, 5-veined at h

e green and finely appressed-stel-

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<ni II .. , - ,] iin iiiulous. ca. 2 2.5 mm long;

anthers elliptic-oblong, 0.6-0.8 mm long. Pistil-

late flowers with short pedicels ca. 1-1.5 mm long,

becoming 3-5 mm long in fruit; calyx lobes 5,

imbricate, very unequal, the 3 abaxial lobes much

larger, in fruit 6 8 mm long. 2.5 (> nun broad.

!!•> i; laciniate, provided on each side with 5 10

teeth ca. 0.5-3 mm long, the 2 abaxial lobe- tnin-li

smaller, nearly or quite obsolete: pelals absent: disk

strongly asymmetrical, with larger adaxial lobes

0.9-1.2 mm long, 0.5 0.7 nun broad, the 2 adax-

ial lobes very small; ovary stellate-tomentellous;

styles erect, distally quadrifid, hispid-stellate, ca.

2-4 mm long. Capsules ca. 5 mm long; columella

3-4.5 mm long; seeds ellipsoid, smooth, mottled

gray and brown, apically beaked, 3.1-4 mm long,

2.4-2.9 mm broad; caruncle ca. 1.5 mm broad.

Scattered in weedy habitats from extreme south-

ern Texas to Panama, reappearing in \ enezuela.

Paraguay, and Argentina, but not reported from

most of tropical South America. The two Pana-

icteal glands of the

3. MJejen

33.1. Mabea Prodr. 15(2): 1151. 1866. TYl'K: Colombia:

Schlim 1132. Venezuela: Fendler 24. Pan- ama: Sutton Hayes 715 (syntypes, not seen).

Additional collections examined. PANAMA, VKH-

e between Santa Fe and San JOM-. I) lrc\ 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 del Rio San Ju an, Luna 47 (MO).

33.3. Mabea jefensteH I rtologia62: 341. 1987. TYF'K: Panama. Panama: newly bull-

dozed trail off Cerro Jefe Road, 0.4 km beyond

turnoff to Alto de Pacora, 29 Sep. 1975, J. T.

& 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).

Subfamily IV. Ki'i'iioitmoiDi-: \t-

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

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)

Apodae in Brazil. Phytologia 62: .'.'>') 313. ihiM.K,K.E. i":; -li: in. n - i.'

(Euphorbiaceae) in Panama. Syst. But. 8: 105-117.

The discovery of a distinctive new species of

labia in Panama makes it necessary to provide

Additional specimens examined. PANAMA. PANAMA:

Cerro Jefe area. Intomo el a I. 33'X, (F|. Correa el al. 1601, 1610(MO). Com COM. I 1333 (MO), /i'lm & D'Arcy 6253 (MO, 2 sheets), D'Arcy 121X3 (MO). 12201 (F), D'Arcy X Svtsma 14733 (F, MO), Duke 9474 (MO), Dwrer et al. 7296 (MO, 2 sheets), 7364 \l(» ' >m el al f>/<>9. , 163 (MO). Hummel i. >''

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, 1393 (MO), Mans el al. /..,•; |MO), Man A Kaltank, 1864 (MO), Nee et al. 8752 (MO), Sytsma 960 (F, MO); Canazas mountain chain, near Rancho Chorro, above Torti Arriba, 400-700 in, Folsom et al. 6709 (MO).

34. Senefeldera

Senefeldera C. Martius, Flora 24 (Beibl.): 29. 1841. TYPE: Senefeldera multiflora C. Mar-

Monoecious shrubs or trees without evident m latex; glabrous throughout. Leaves alternate pseudoverticillate at ends c 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 longitudinally; pollen grains subglobose, tectate, 3-colporate; pistillode absent. Pistillate flowers solitary at proximal nodes of inflorescence, sessile; calyx 3-parted, segments distinctly imbricate; petals and disk absent; carpels 3, unappendaged, each with a single ovule; styles unbranched, free, or basally connate. Fruits capsular, thin-walled; columella slender, usually not persistent; seeds solitary in

cm long; lateral axes 5 8, the peduncles 5-11 mm long. Staminate flowers 2 or 3 per node; bracts ca. 1-1.5 mm long, acute, the glands infolded on adaxial side; pedicels 0.5 mm long or shorter, subtended by 1 or more bractlets within the bract; calyx segments 3, unequal, acute, as broad as or broader than long, ca. 0.5-0.7 mm long; stamens 5; anthers apiculate, ca. 0.4-0.5 mm long (much longer than the very short filaments). Pistillate

olitary at the 2 or 3 lowermost nodes of each lateral axis, sessile; bracts apiculate, 1-1.3 mm long, with glands 0.8-1 mm across; calyx segments 3, slightly imbricate, broadly ovate, apic-

As treated by Jablonski (1965: 171-174), Se- /,-, feldera is a genus of nine rather poorly understood South American species. It is here recorded from North I i for the first time.

34.1. Senefeldera testiculata Pittier, Contr. 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 channeled; stipules triangular, ca. 2 mm long, deciduous (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

raphe conspicuous.

Lowland evergreen rainforest, Panama to Ven- ezuela; here reported from Panama for the first

The Panamanian specimens are a rather good match tor collcrii,,ns »,t ,S /.--,/,, >//,-.<<; lioiti Xulia, Venezuela (Stcymnark 997,70. 90917, VEN). Al- though they differ from the South American plants in having more acute leaf bases and shorter inflo-

Specimens examined. PANAMA, DARIKN: Rio Can- 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 (DAV, MO; incorrectly cited as Caryodendron angusti-

35. Sebastiania

Sebastiania Sprengel, Entd. Pflanzenk. 2:

flowers borne distichously on the rachis; seeds not over 2.5 mm long 1. S. corniculata Shrub 2 -3 in high; leaves elliptic-lanceolate; staminate flowers borne spirally on the rachis; seeds ca. 4 mm long 2. 5. panamensis

35.2. Sebastiania panamensis Webster, sp.

nov. TYPE: Panama. Chiriqui: N of San Felix

at Chiriqui Moras del Toro border, on Cerro

Colorado copper mine road, 5,000-5,500 ft.,

3 May 1975, Mori & Kallunki 5786 (holo-

type, DAV; isotype, MO).

Shrub ca. 2 3 in high, I he trunk ± unbranched;

twig- -lender. -uhlerele. antmr-ely ± appressed

pubescent. I,ciiics with petioles 3 7 mm long, a[»-

pressed puhe-cent: stipules triangular-lanceolate,

dark, 0.8-1.2 mm long; blades thinly chartaceous

or membranous, elliptic-lanceolate, ± caudate-

long, (1 )2 4 cm broad, concolorous, without lam-

mar glands, glabrou- to di-lmetl\ hirsutulou.- on

both laces; midrib plane above, distinctly raised

beneath; major lateral \eins ca. 8 20 on a side.

-traiglili-h. -lightly proiuiiiuloii- beneath, distalh

anaslamosiiig into intramarginal loops; vemlei-

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

-olilar\ at base, -laminate Mower- I or 2 per distal

bract; bract- -pirall\ arranged, lanceolate, dark,

ca. 1 mm long, on each side with a short-stipitate

(ca. 0.5 mm)cyathiform gland 0.5 -0.8 mm across.

Stamina!,- /lowers: pedicel less than 0.5 mm long:

calyx 3-lobed, the lobes obovate, ca. 0.6 -0.8 mm long, the anthers ca. 0.3 mm long. Pistillate flou-

eis: -ub-e-sile or the pedicel up to 1.5 mm long

at anthesis, becoming up to I mm lour in fruit: cal\\ lohe- .'.. imbricate (coxering o\ar\ in bud).

0.7-1 mm long, eglandular within; o\ar\ glabrous.

3-carpellate. each carpel with a pair of subapical

horns; styles nearly tree, spreading, tapering, ca.

1.5-2.5 mm long. Capsules ca. (> mm high, ca.

7 mm broad, with 0 subapical lower triangular processes; columella ca. 1.5 mm high; seeds ovoid-

ellipsoid, reddish brown and mottled, smooth, ca.

Montane rainforests , or cloud forests, western

Panama.

This shrubby species, now represented by sev-

eral collections from montane rainforests in Chi-

riqui and Veragua-. doe-

floral characteristics it re

; not appear to have been

overall appearance and

scinblc;- the weedy N. cor-

nicithittiiA section Micro stachvs:(Adr. Juss.) Muell.

w I\ habit. Il- -piialK

Pax (Pflanzenrcch 85: 8<>fl'. I O | 2) it would key

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

Panamanian plant- do not lit into section lilrno

gyne (Klotzscb) Benth. because the staminate ca-

Ivx is not asymmetric, the fruit is echinate. and

the blanche- arc not p111x . Nor do the Panamanian

plants agree with species in section Sebastiania,

because of their leaf-opposed spikes, broader sta-

The sectional divisions used by Pax do not seem to be very well founded, -o thai a -a ti-la.[oi v

seem to be feasible unt At present, it appear represent a possible <

Additional specimens examined. PAN Wl \. i Ulliioi I: Cerro I olorad.., 1,20(1 1,50(1 in, Man X Dressier 78J7 (MO. dnpl. .U SC/ -ecu bv \1. Hull). Snlhran .<"/lM<». Antonio 1409 (MO, dupl. at PMA seen by M. Unit); above San Felix along mining road, above Chame, 1,200 1,500 m, Croat 33044 (DAV, \IO). ,c.;: silM Fel.x.Mon 1,200 m, Croat 33437 ( M<)): Cl.iri.pii I rail, pre,„„,) rainforest between I'niol.i ,uid Ouehrada Honda, Kirk- bride X Puke 898 (DAV, MO), Churchill cv Chinch,II ,,(>:!3(i: MO): I .a fortune hydroelectric pn.j.-.-t. I.Kill 1,200 m, Hammei\2165 (DAV, MO), Knapp 4980(MO), Mendozaetal. 7 70(MO); E of Fortuna campsite, balsam X /Vcs.s/,7 >:><ei IDW. \|()|. uiiV.l \-: Cerro 'lute, 7..0 1.400 m. Iiilama 1817 (MO). Knapp X /v,cv, I3.,'t(\\ MO), knapp X SMsma _'..o2 (F, MO), Mori

.,J3J\] vio

. \1(M. 7h09 (

35.3. Sebastis

Several ttuitin

Webster & Huft

nhn.n

1-1 I I.Mhi.

e, broadly ovi

...ii', i« .

. Sebasti- ania from southern Brazil and Argentina as S.

brasiliensis Sprengel. ,S. anisandra (Griseb.) Lillo,

and S. warmingii (Muell. Arg.) Pax. Even closer

is .S. macrocnrpa Muell. Arg. of Ceara in north-

i i llrazil. with which it shares a large capsule

(to 15 mm in diameter).

Until flowering material becomes available, it

: Braz

ndescribed species or belongs to o

The Costa Rican specimen cited below is the

basis for the record of Ophellantha spinosa Stand-

ley cited by Standley from Costa Rica (Standley,

1938: 1557). That species is definitely known only

from northwestern Mexico to Honduras. One of

the Colombian collections {Gentry & Guadros

47466A, MO) has an immature inflorescence with

a -ingle pistillate flower and the multiparled brac-

Ira glands ill .il air \ ha car lerisl ir of several species

San Ramon, Los Loros, Brenes 22679 (CK, \ i I. I'w \M\. I.os SANTOS: 1 2 mi. W of Candelaria, Duke 12443 (MO, US); Las Tablas, Dwyer 1100 (MO); Los Santos, :',() MI, l.uo .HMMOI. CoioMHix. VIIANTKX): Puerto Co- lumbia, 5U~K)0 in, Dufruml '>:'>, (| . distributed as S. granatvnsis Muell. Arg.). UOI.iVA " '

icellate; calyx of 1 lobe or rudimentary or absent;

petals and disk absent; stamens mostly 2-5; fila-

ments free or basally connate; anthers extrorse,

dehiscing longitudinally; pollen grains subglobose,

tectate. 3-eolporate; pistillode absent. I'tstillatc

floiccrs sessile or pedicellate; calyx moslK 3-lohed.

the lobes sometimes reduced or obsolete, eglandular

within; petals and disk absent; carpels 3. each with

a single ovule; styles free or basally connate, un-

branched. Fruits capsular; columella ± persistent;

seeds solitary in each loeule, carunculate; testa

-ii th: endosperm copious.

As here circumscribed, Gymnanthes includes

Actinostcmon Klotzsch and Dartylostemon

Klotzsch. Although most authors have upheld at

least l< tinostemon as a distinct genus, this seems

A' t „„,.'/<. r/56,

mnanthes Sw., Prodr. 95. 1788. LECTOTYPE:

Gymnanthes lucida Sw. (chosen by Grise-

bach, Fl. Br. W. Ind. 50. 1859).

stly persistent; blades

ulate, glandular or

eglandular. Inflorescences terminal or axillary, bi-

sexual (or less commonly unisexual), spiciform, of

1 -several basal solitary pistillate flowers and many

ilitary pistillate flowers and 1-several

Pax

& Hoffmann (1912: 13) did not provide any con-

vincing distinctions in their key, nor did Jablonski

(1967: 164, 178), who candidly stated that "the

distinction between \ctinoslemon and Gymnan-

thes is very vague." Even when Actinostcmon is

merged with Gymnanthes, the enlarged genus is

difficult to distinguish from Scbastiama. Mueller

(1866: 1164-1165), in fact, combined the two,

but called the composite genus Schasiiama be-

cause the name Gymnanthes (which has pnorilx)

seemed inappropriate to him. The distinctly re-

duced staminate calyx ol Gymnanthes fumi-hes

only a tenuous difference from Scbastiama. in

well developed. As Pax & Hoffmann (1912: 89- 90), , Sebasliania

nus, part of which -houM he

tithes. Clarification of these

Rothmalr. (1944), follow

combined with Gym-

difficulties will have

trepid monographer,

ig a suggestion by

recently (Webster, 1983); Aterc

posed of by lectotypifying it so 1

ropical genus of about 40 spec

s are reported hen

BROWNE, P. 1756. The (a\i 1 and Natural History of

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- pomaneae. In: A. Engler, Das Pttanzenn-i, h l\ . I I ,". V(Heft 52): 1 319 (Gymnanthes, pp. 81-88).

Hoiiimn u. \V. 1044. Nomina generica neglecta 1753- 1703. K. ,,,,. 53: 1 37.

ulent, ca. 2-4 r

13-21

l long ;

long in fruit; calyx segments 3.

pointed, not imbricate, 0.6 0.7 mm long; o\ar\

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.

lb. Stanimate bracts (at least in pari) with paired glands on each side, each bract subtending 1 flower; leaves eglandular on margins; staminate calyx absent; fruiting pedicels 40-50 mm long; seeds ca. 5.5 mm long 2. G. dressleri

36.1. Gymnanthes actinoslemoides Mu. II.

Arg., Linnaea 32: 103. 1863; Pax & Hoff-

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).

Leaves with petioles 3- i long, ± puberuler

im long; blades cha

taceous, elliptic-oblong, mostly <ai

at tip (the acumen acute, ca. 0.5-1.5 cm long),

acutely cuneate at base, ca. 6 16 cm long, 2-6

cm broad, without laminar glands but with 1-3

depressed cvatlulorin glands at proximal marginal

erenulalioiis; midrib raided oti both >ide- and pu-

berulent [iro\uuall\: major lateral veins ca. 10

15 on a side, arcuate: veinlet- distinciU pronii-

nulous beneath (diiditlv -o above), forming irreg-

ular areoles; margins plane, remotek crenulale

(teeth mostly 12 17 on a side), the distal teeth will i a mini lie deciduous glandular lip. I be pro \ una I

teeth with depressed cyathiform glands. In/lores-

(c/ii-e.\axillarv. racemilorm, unisexual oi bisexual.

2-7 cm long, the rachis puberulent. Staminatc

(loners in cvmules of .'». subtended b\ iimhonate

bracts ca. 1 mm long. each hr.nl with a pair of

cyathiform glands 0.6 0.8 mm across attached

above the base (-iibupical); pedicels 0.5 1.5 nun

long (longer in central flower); calyx usually of I

The -ingle Panamanian collection of this species

bears only fruits, so the identification of Bristan's

plant with a Mexican species must be provisional.

However, the characteristic leaf venation and es-

pecially the distinctive marginal foliar glands sug-

gest that our plant is conspecihe with specimens

from Veracruz. Mexico (such as I'urpiis 37<)5.

8060 from Zacuapan, I 110 from bortin. all at

UC, from which the floral characters have been

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).

Tree 2 m or more, glabrous; twigs slender, sub-

rete, smooth, brownish. Leaves with petioles 3-

nim long; stipules ovate, rounded, scarious, ca.

mm long; blades chartaceous, broadly elliptic,

;umen obtuse, 0.5-1 cm long), cuneate at base,

-8 cm long, 1.5-4 cm broad, appearing eglan-

lar glands, distinctly paler and glaucous beneath; major lateral

veins ca. 6-8 on a side, straightish or dislally

arcuate; veinlets distinctly prominulous beneath,

often as prominent a- the lalerals. forming areole-

partly parallel to the lalerals; margins entire, with

a plane or slightly reflexed suheartilagiiiou run.

Intloieseenees axillary, mostly at the base or lower

axils of annual increment-, racemilorm. unisexual

or bisexual, where bisexual with I or 2 basal pis-

Volume 75, Number 3 Webster & Huft

ious, ca. 0.5-0.7 mm long, mostly with paired

subsessile cyathiform glands on each side at base,

the larger gland of each pair ca. 0.3 mm across.

Staminate flouers with pedicels ca. 0.4-1.2 mm

long, articulated at the top; calyx, petals, and disk

absent; stamens 2-4; filaments free or basally

united, 0.4 0.7 mm long; anthers 0.4-0.5 mm

long. I'istillate flowers with pedicels ca. 3-5 mm

long at anthesis, increasing to 40 50 mm long in

fruit; calyx lobes 3, ovate, not overlapping, ca.

0.5-0.7 mm Ion • _ .. i<iu »< lliin, the margins

gland on each side, and in its well-developed i ;

>ate calyx. Among the species with bracts sub-

tending solitary flowers, the closest to G. dressleri

appears to be the Cuban species G. albit « <• a

seb.) Urban; however, in that species the leaves

are more elongated and lack cuspidate tips, the

have 5-12 stamens.

37. Maprounea

Maprounea Aublet, Hist. PI. Guiane 2: 895. 1775.

rtyles 3

eeds i

a. 1.5-2 mm long, thick

nnate at base. Capsules r

5.5 () nun loni!. -111.|.«-1

Li I!-, smooth, dark brown

TYPE: Maprt

Shrubs

4.5 blade I

>rous throughout; latex nei- monoecious. I.cures alter-

stipules small. persistent:

, entire, glandular or eglan-

apically beaked; caruncle ca. 1 mm across.

This new species from Cerro Jefe appears to be

i If i I ) > I <i ' <•<< *" luell. Arg., which

was described (Linnaea 32: 107. 1863) from the

MCIIIIU of Ocana in northern Colombia. Although

the type collection of G. granatensis (Sehlun .,/ih)

has not been examined, study of a photograph of

the type and Mueller's description (in DC, Prodr.

15(2): 1189-1190. 1866) suggests that the Pan-

amanian plant differs in some important particu-

lars: the leaves are broader, more abruptly Cus-

pidate. ;:•-!. \\v\\ •l-\;- •».- hria at I. aaii ei luv ; I

Infl,

. rhes a calyx, whei

duplex glands on each side, while i

species (judging from Mueller's description), the

bracteal glands are single on each side.

In the system of Pax & Hoffmann, G. grana-

triisis would probably fit best into the '"genus"

•mil. although thev lifted it under Se-

bastiania (Pax & Hoffmann, 1912: 150). How-

ever, because of it! glabrou inflorescence and well- developed pistillate calyx, it would not readily fit

m 7M'.S (MMIMI Pax), perliap> tlie species ino-

similar to G. dressleri and G. granatensis i* (,

farinosa (Criscb.) Webster". That West Indiai

species has somewhat similar leaves but differs ii il lli-ee tiov.ci ••.! • M,'!, nal< : ii-'icl.- w it I : - wti

i I soliturv. pedicellate, pistillate Mowers at basal

nodes, the staminate flowers densely aggregated in

a strobiliform mass at the end of the fleshy enlarged

rachis, separated from the pistillate portion by an

elongated internode (pseudopeduncle); bracts bi-

glandular. Staminate flowers mostly 3 per bract;

pedicel very short; calyx ± 3-lobed, distinctly gam-

! i ibrii ate. + < overing the sta-

mens ai hud: p« ;i i i ! 'I . H I uneiis usu

ally 2: filaments completely connate into a slender

tube that is exserted from the calyx at anthesis;

anthers bluntly apiculate, dehiscing extrorsely and

longitudinally; pollen grains subglobose, tectate-

perforate, 3-colporate, colpi marginate; pi^lillode

i in ' iters solitary to each bract,

« ii in pedicellate; calyx 3-parted, segments im-

bricate, eglandular within; petals and disk absent:

carpels 3, each with a single ovule; ovary unap-

pendaged; styles connate into a column, the tips

unbranched, spreading. Fruits capsular; columella

not persistent; seeds solitary in each locule, the

testa distinctly foveolate; caruncle large and partly

occluding top of seed; endosperm copious.

A well-marked genus of three or four species,

one or two in tropical America and two in tropical

Africa, easily distinguished from the genera in the

Gymnanthes-Sehastiania complex by its char-

37.1. Maprounea guianensis Aublet. Hist. I'l. Guiane 2: 895, tab. 342. 1775. TYPF: French (Lilian.i. Inhle! (not seen).

Trees to ca. 12 m high, glabrous; twigs subterete, smooth, brownish. Leaves will) petioles ca. 0.5 1.5 mm long, slender; stipules triangular to lanceolate, scarious. ca. 0.5 1 mm long. persi-teni;

blade- think chart.i< eon-. o\ale to elliptic, rallier

(and nmiutelv auriculate at junction with petiole). ca. (2-)3-7 cm long, 1.5-1 cm broad, usually with

1 or 2 elliptic laminar glands on the underside near midrib (ocoa-ionalK with a tew -mall cm ular glaud-

towards die lip. or souieliitie- entirely eglaudular): midrib plane aho\e. di-liiicily rai-ed beneath: major

let- proniiniiloiis on holh side-, forming areoles ± parallel to lateral veins; margins plane or recurved,

entire. Inflorescences terminal mostly on short lateral branches, 1-2 cm long; pistillate flowers sol-

itary at 1 1 basal nodes; staminate Hower- in head.- ca. 3 9 mm long: staminate bracts rallier Meshy.

the tip acute, less than 0.5 mm long, the cyalhiform

articulate abo\e er> : eels: calyx gamophvllous, basally ( a stipe ca. 0.3 O.M mm long; -laminal column

slender, mostly 0.7 1 mm long; anthers 0.3 0.4 mm long. Pistillate flowers on ascending or recurved pedicels (1 )2 5 mm long at anthesis, these becoming () \\\ mm King in fruit; calyx lobes .'..

l long; ovary s styles 3, 2 3 mm long, united nearly or quite hallway into a stoul column. Capsules t oblate.

not lobed, 6-6.5 mm diam.; seeds ovoid, somewhat compre-sed.giayi-h broun. diiuv. dislinoiU beaked. deeply and coarsely foveolale on both sides, 2.9 3.6 mm long, 2.8 3.5 mm broad (including caruncle): caruncle large. « o\ering nearly hall the lace of the .red venlralK. appearing '2 armed doi

sally.

Ibiinfnre-I- at loss e|ey alion-. ranaina and Trin- idad south to Peru anil Brazil (localities in South America summarized by Jablonski, 1967: 180).

It is curious that this distinctive plant has been ci.Heeled only a single time in a well-known area in the center of the ( .iii.il /one. Presumably it will

esentualK turn up in various lowland areas toward the Colombian border.

Specimen examined. PANAMA. CANAl. /.ONE: 1 mi. N of MHtiniil UN road to 1 A \ radar lower. Tyson rt al. 27<> I

lingia sylvatica L.

39. Sapium

Sapium P. Browne, Civ. Nat. Hist. Jamaica 338. 1750. IVI'I': Sapuini janiaicense Sw.

In the original treatment, the account of Sapium was avowedly tentative, pending the appearance of Jablonski's study of the Caribbean and Central \mericaii -p. -. Jablonski*- work, as well as a

study of more recently collected specimens, ne-

manian specie-. Of the four -pecie- recognized in

the original treatment, only .S. eiilaiidulosum re- main- unchanged. We haye followed Jablonski in referring the -pecie- called .S. aucupar mm by Purch

to S janiaicense and in uniting .V caudatum and S. biglandulosum under the name .S. aticupariittn. Jablonski's action m re-toring the name >'. jamai- i-en^e seems straightforward and is adopted here. The nomeuclatural subtleties coimecled with .S.

blonskfs choice is followed a- a lenlatiye coiicliision

Three additional species of Sapium are now known from Panama, bringing the total to six. Keceiil sterile colic* I .- of a seseiilh. possibly

lUldescribed, species haye been made on Barm Colorado Island.

CKOI/AI. L. 1943. Novelties in American Euphorbi- aceae. J. Arnold Arbor. 24: 165 189.

Hi IT, M. J. 1987. Four new s,„v„s of Sapmm (t.u phorbiaceae) from Central and South America. Phv- tologia 63: 441-448.

tropical Euphorl E. 1968. Notes on

!<,: :\>>:>, i Sapium. Pliytologia

< inflexed or cucullate.

Webster & Huft

less than 2.5 times as long i

II n I ' i, - , ll III Li the secondary vein- usuall) n than 20 per side .... 1. 5. aw

im Jacq., Select. Stirp.

v./ iv, i • , I •' ,, I

//,//,-,

uell. Arg. (Linnaea 32: .' (II li II I //, '•': ;.,,,,!•,,!.• ;

PI. ed. 2, 1431. 1762, correction of H. 'osa L. Sp. PI. ed. 1, 1191. 1753. Kxvoe- •glandulosa (L. ) Muell. Arg. in DC, Prodr. 204. 1866.

in. Sp. PI. 2: 65. t*K lionphuuKW

>ium nwrilziunum KI..I/,, b in Seem., Bot. Voy. Her- ald 100. 1853. Sapium biglandulosum (L.) Muell. Arg. var. moritzianum (Klotzsch) Muell. Arg., Lin- naea 32: 119. 1863. Excoecanu biglandutusa (I..) Muell. Arg. var. montziana (Klotzsrh) \h.rll. \~K. in DC, Prodr. 15(2): 1206. 1866. Sapium aucuparium Jacq. subsp. motil ziariuw (klol/seh) I'IUHT.

Contr. U.S. Natl. Herb. 20: 127. 1918. TYPE: pre- Minuil.K Colombia: \l<>ritz2:tt> I'.m.iiiia: S<-rmann

itary, terminal, to 22 cm long, bisexual or stami-

nate. Staminate flowers in groups of 7-10, the

subtending bract flabellate, ca. 0.5 mm long, bi-

glandular, the glands circular to oblong, 0.5-3 mm

long, 0.5-1 mm broad; calyx ca. 1 mm long,

ri.|Mi|:i . :' lipped: Li I I r ." >?'< -Jilh/lr flowers lo

10, solitary at basal nodes, the bracts and glands

as those of the staminate flowers; calw eupular.

2-lobed; ovary orbicular, the style simple, the style-

branches strongly reflexed, the tips ex[)ande.|. ( u/>-

sules ovoid, to 1 cm long, smooth; seed- moid.

flattened laterally, the surface warty.

There has been considerable controversy con-

cerning the proper name of this species. The name

.S. uiicuparium Jacq. had long been applied to it,

but Croizat (1943: 175), whom Burch followed in

the original treatment, referred that name to the

species usually known as S. jamaieense Sw. and

resurrected -S. bighrndiilosnm for this specie.-. ,|a-

blonski correctly restored Jacquin's name to the

present species, but rejected the name >'. biglari-

dnlosnm a- a rmrrn'ii loii/iisiiin. leasing .S. uucii-

parium as the earliest available name. It would

otologu, v,T,, , I niter, Contr. U.S. Natl. Herb. 20:

,„•,', CamU. .... I' I (chosen by Jablonski, 1968).

s, ..«,, ,, , t&intvum Pittier. Contr. I .S. Natl. Herb. 20: 128. 1918. TYPE: Panama. Col6n: Fato, sea level, 10 Aug. 1911. f'ittirr II 11 (holotype, US).

Stillingia haematnntha Standley, Ann. Missouri Bot. Card. 27: 314. 1940. TYPE:'Panama. Code: N rim of Kl Valle, 9 Julv 1939, Allen I<>15 (holotype, F; isotype, MO, F neg. 62368).

I applic biglcu diilosum can be fixed by a proper lectotypification.

More study is needed before that can reasonably

be done, and, in any case, it is not clear to the present author whether any of the elements orig-

inally cited under S. biglandulosum actually he-

long to this species. The best course, then, seems

to be the tentative acceptance of S. auciifHintim

Jacq. for this species until the matter can be sat-

isfaeloriK resolved. We are following Jablonski (1968) in uniting

Sapium caudatum and .S. bi^lnndiilosiim (-etisii Croizat and Burch) under .S. aueupariitm Ivv

•mnti-ilioti ol a wide i m_.- ol roll, < tiuii- Irom I'an-

iS utl used by Burch to distinguish

ntinuously variable and do not

able species of lowland tropical

long, the 2 apical glands prominent, cylindrical,

ca. 1 mm long; stipules ovate-deltate, 1-1.5 mm

lanceolate to oblong-obovate, 5-40 cm long, 1.5-

8 cm broad, 3-5 times as long as broad, the base

rounded to acute, the margins entire, obscurely

toothed or coarsely serrate, the apex short-acu-

39.2. Sapium eglandulosum Ule, Bot. Jahrb.

Syst. 35: 673. 1905. TYPE: Brazil. Amazonas:

Bom Fin on the Rio Jurua, Nov. 1900, Ule

5356 (holotype, B, F neg. 5522).

The following collection records a considerable

shows that even these meager grounds a

able. Plants with petiolar glands c

in the West Indies (cf. Ekman 5512 from His-

patiiola. I/arris 9156 from Jamaica, and Pringle

104 from Cuba, all F), and thus there is no mor-

•I ulogii ill I -i .inn ii S,. • • jamaicense is

simply a variable species in regard to the presence

or absence of petiolar glands, and there is no jus-

. it/Kirium sensu Croizat, J. Arnold Arbor. 24: tification for recognition of .S. pleiostachys. 174. 1943, sensu Burch, Ann. Missouri Bot. Card. A few new provincial records are recorded be- 54: 325. 1967, non Jacq. 1763. V

Stifiium jilriastacfivs Sclnntiaiin & Pittier, Contr. U.S. OW" Natl. Herb. 12: M,4. 1008. n,,,: Costa Rica. Pun- ^.^ ^.^ ^^ pANAMA DARlfN:

Rio Pirre, flristan 7475 (MO). PANAMA: N of highway ca. 2 mi. E of El Llano, 200 m, Foster & Kennel \ i>>. I if. \Y). umci \s: [sla de Coiba (Penal Colons). Ih, \c, /.566(MO).Coi.o\im\. WIIOOI i\:Miin1n|.i<Ml.-S.1ii l.ms. Canon del Rio Claro, 330-425 m, Cogollo <>0o, 1232

Valverde at Orosi, 1,200 m, Mar. 1902, Pittier (M°)-

s.n., Inst. Pis. Geog. Costa Rica no. 16366 (holotype, US-578045; isotype, F, F neg. 62364). 39.4. Sapium oligoneurum Schumann K I'll

R f w . I i- M„;B„ ,„ P.nom, tier, Contr. U.S. Natl. Herb. 12: 168. 1908. Sapium biglandulosum (L.) Muell. Arg. var. oligoneurum (Schumann & Pittier) Monach.,

The correct name of this species has also been Bull. Torrey Bot. Club 67: 772. 1940. TYPE:

matter of some confusion. Sapium jamaicense Costa Rica: near San Rafael on road from

d been used universally until Croizat (1943: 174) Cartago to Cot, 1,500 m, July 1899, Pittier

>erted that .S. aucufxtiium sensu Jacq., Knum. s.n., Inst. Pis. Geog. Costa Rica no. 13-HPi

. Carib. 31, 1760, properly refers to this plant, (holotype, US-578903).

, (:..i „ ii.K.

Sapium sulciferum Piltier. Conli 169. 1908. "

in contradistinction to Jacquin's

his Selectarum Stirpium Ameria

(1763), which had been followed by most other var. sulciferum (Piuier) Monach., Bull. T..rre> B«lt. authors, who had applied the name S. aucuparium Club 67: 772. 1940. TYPK: Costa Rica. La Palma,

as in this paper. Jablonski (1968), however, cor- i-500 m- 15 Au8- 1898- Tonduz s.n., Inst. Pis.

rectly pouSed out that Jacqum's pubhcat.on of S. %*£&*** ^ 12428**** US"577588;

aucuparium in 1760 is invalid, since a description Sapium schippii Croizat, Amer. Midi. Nat. 29: 477. is lacking, and that the traditional application of 1943. TYPE: Belize. Toledo District: Forest Home, .S. uufupuhum as published b\ Jacquin in 1763 Punta Gorda, Schipp 1049 (holotype, A, F neg. is correct 62361; isotypes, F, F neg. 62365, MO, F neg.

Jablonski (1968) recognized Sapium plriosta-

c/iys as distinct from .S. jamaicense, referring all Monoecious tree to 20 in. Leaves membranous

collections from Panama and Costa Rica, as well or chartaceous; petiole l-3(-6) cm long, the two

as a few from (mate* ! . ' • • . lo the former glands near the apex opposite or suboppo-iie. c_\

species, and restricting the latter to the West Indies lindrical, 1-2 mm long; stipules ovate-deltate,

and northern Central America. The only difference oblique, 2-3 mm long, 1.5-2 mm broad, ap- he adduced was I he presence of peliolar glands on pressed, persistent: l.'ade •; I i : or elliptic oblong.

S. pleiostachys and their absence on S. jamai- 4-10(-18) cm long, 2.5-4.5(-8) cm broad, 1.2-

cense. Even granting this difference, recognition 2.4 times as long as broad; midvein prominent, the

ol S. pleiostachys would be tenuous indeed, given secondary veins 10-15(-20) per side, somewhat

the inadvisability of maintaining a species on the inconspicuous; base rounded lo obtuse; margin ap-

basis of a single i iture and the pearing entire, remotely denticulate with minute

assertion that both variants occur in northern Cen- glandular teeth; apex acute or more often abruptly

tral America (indeed, specimens collected by Ma- short-cuspidate, conspicuously and tightly cucul-

tuda at Escuintla, Chiapas, are cited under each late. Spikes solitary, terminal, to 22 cm long, bi

species by Jablonski). A thorough examination of sexual or staminate. Staminateflowers in groups

collections from throughout the range, however, of 5-7, the subtending bract short, broad, 1-1.2

mm long, 1.8-2.1 mm broad, rounded, hyaline,

slightly erose, biglandular, the glands suborbicular

to oblong, 1.8-3 mm long, 1.8-2.5 mm broad,

flattened; calyx cupular, 1.7-2 mm long, 2-lipped;

stamens 2, the til mrni h •• /'. .,*;• //« ,,,

10-22, borne singly at basal nodes of bisexual

spikes; bracts and calyces as in the staminate flow-

ers; ovary globose; styles simple. Capsules sub- Join.-, I -IIJIIIII. .« nid. -adisessile, 5-9 mm

long, 5—12 mm diam., smooth; seeds subglobose,

slightly compressed, yellowish, the surface warty,

edges, and

stamens 2, the filaments free. Pistillnh

14-22, solitary at basal nodes; bracts as in the

staminate flowers; calyx 1-1.5 mm long, cupular,

2-lipped; ovary globow styles simple, fused for %-

% their length, the free portion strongly coiled.

f globose, subsessile, 7-10 mm long,

smooth; seeds subglobose, flattened laterally, ca. 4

mm diam., the edges short-winged, the surface

Specimens examined. I'WVMV. HOCAS DEL TORO: region of Cerro Colorado, 3.3 mi. above Camp Chami, 8°35'N, 81°45'W, ca. 1,350 m, McPherson 9587 (F). i iiimoi i: K of Boquete on Cerro Azul near Quebrada Jaramillo, 1,500-1,620 m, Croat 26820 (MO, NY); along road between Gualaca and Fortuna Dam site, 10.1 mi. NW of Los Planes de Nornito, 8°45'N, 82°17'W, 1,250 m, Croat 50032 (MO); Boquete, 4,000 ft., Davidson 852 (MO, US; cited in the original treatment as S. au-

• n I' • < i >lorado, ca. 3.5 mi. m iCamp, ca. 8°35'N, 81°45'W, •

.35(1 , Ul'hr, 9\ (F*i I .30(1 ,

goneurum, or S. sulciferum.

39.5. Sapium pachystachys Schumann & Pit- tier, Contr. U.S. Natl. Herb. 12: 168, tab.

16. 1908. TYPE: Costa Rica. San Jose: Dota

Mountains, El Copey, 1,800 m, Feb. 1898,

Tonduz s.n., Inst. Fis. Geog. Costa Rica no.

11875 (holotype, US-333961; isotype, F, F

neg. 62367).

to 25 m; older twigs covered

with < rowded |M rsr-lei:l -lipule- I.circs membra-

nous or chartaceous; petiole 2-5 cm long, the 2

glands near the apex subopposite, cylindrical. 1

3 mm long; stipules deltate, 1 () mm long, 2-3

mm broad. approhsed, p< rsislrnl; blade elliplie. eb

liptic-obovate, or elliptic-lanceolate, 5-20 cm long,

2.5-7.5 cm broad, 1.6-2.4(-4.5) times as long as broad; base rounded or obtuse, rarely acute; mar-

gins < ap. , i aeute. oeea.-ionalk -I iorl aeuinmate. |ha ne. ,S/. ,-/.•<•.•..

solitary at the apex of smooth lateral shoots, to 20 em long, bisexual. Sttiniinntc Ilium s m groups of

7-10(-12), the subtending bract short, broad, to

2 mm long, h\al • >> bid edol I ih ulami-

oblong, 2.5-3 mm long, 1-1.5 mm broad, flat-

tened, calyx 1-1.5 mm long, cupular, 2-lipped;

A related species, S. allenii Huft, has recently

been described from eastern Costa Rica (Huft,

1987) and may eventually be discovered at lower

elevations in western Panama. It differs from S.

- h v.s in having axillary spikes, smaller,

stipitate capsules, and a small membranous calyx

that does not persist on the mature capsules.

border with Chiriqui, Cerro Colorado mine area, from Charm Station to ca. 9 mi. along road, 8°35'N, 81°54'W, 1,100-1,700 m.// Boquete, Finca Collins, Blum & Dwyer 2558 (MO); SO de campamento Fortuna (8°45'N, 82°15'W), sitio de pre- sa, desde la finca Pitti hasta e filo del Cerro Fortuna, 1,000-1,200 m, Correa et al. 2797 (MO, 2 sheets); Cerro Colorado, along road to copper mine 24.1 km beyond bridge ovei •'• • 3anl it km beyond turnoff to Escopeta), 1,390 m, Croat 37305 (MO); Cerro Punta, 2,000 m, Lao 328 (MO); above Los Llanos, 8°47'N, 82°38'W, ca. 2,100 m, McPherson 9258 (F, MO); Cerro Colorado, 50 km N of San Felix on the Continental Divide, 1,200-1,500 m, Mori & Dressier 7819 (MO, NY); be- to., , i, I-!',, | ..,;, I!, n I I , - -i, i , - I ii, p. -milli. i,i -....»•

of Cerro de la Horqueta, 1,200-1,700 m, Pittier 3165 (US); slopes of Volcan Baru near town of Cerro Punta, i., ;M> M S - ", (MO. US). cocLfe: N of El Cope on road past sawmill, 2,400 ft., Antonio 3264 (F, MO). PANAMA: 5-10 km NE of Altos de Pacora, on trail at end of road, 700-800 m, Mori & Kalh.i,!,, f '- (MO, NY, 2 sheets).

39.6. Sapium rigidifolium Huft, Phytologia

63: 444. 1987. TYPE: Costa Rica. Heredia:

pastures above San Rafael, 30 km W of Vaca

Blanca, 1,750 m, 8 Aug. 1971, Lent 2041

(holotype, F; isotypes, MO, NY, US), distrib- ui< «l - , ' •.'/ S, bumaim &

Pittier. Figure 5.

Sapium rigidifolium, which is known only from

high altitudes in Costa Rica and Chiriqui Province

in Panama, belongs to the otherwise wholly South

American subsection Emmenostylum (Mem-lev)

FUJI KK 5. hou-iitfi slam ten- Ifilson

„;.„, umnptrm

Pax (Pflanzenr. IV. 147. V(Heft 52): 211. 1912), ^^ ^J^''"'' Mue"" Arg" ^ characterized by stvles that arc connate for most ol their lengths an.l whose eoluinns persist

mdary veins. The Sou

I include such species as 5. verum

Arriba, above Cerro Punta, 8°52'N, 82°3.VW, 2,100 .. <lr \crcrs & Chutnl.-s »<>:,, (K): Ilc,«,,i«-t.-. Cerro Ho, queta, 5,000-6,000 It.. Ih, w; ,V ll„\,l,;, .»!<.. iMO Cerro Punta, 2,000 ,„, l.fl„ .V>1 (MO. I'M \i. s|,r.-s , Volcan Barii, near town of Cerro Punta, 0,000 h ,S/(7

<\ ('haml)vrs ,'i.> ( \, MO, I'S).

Euphorbia L., Sp. PI. 450. 1753. i.ECTOTYl'K:

Euphorbia antii/itorum L. (chosen by Mills- paugh, Publ. Field Columbian Mus., Bot. Ser.

2: 306. 1909).

Klotzsch) Graham.

Four species new to Panama are reported here.

In addition, the Panamanian endemic, Euphorbia

apocynoides, which was merely mentioned in pac-

ing in the original treatment, is here treated in full.

These changes make it necessary to provide a new

key. Both of the present authors are agreed that

I'ouiscltKi, treated as a separate genus in the orig-

inal treatment, is best regarded as a subgenus of

Euphorbia; thus Poinsettia is suppressed in the

generic key. and the two Panamanian species are

included in the key to Euphorbia.

1836. Euphorh Etud. 284. 1858. Euphorbia subg. Poinsettia ( ham) HmN', New York State Mu>. Hull. J.", I.

<re than 15 cm long, leathery; cyathia 4 <> mm long

of cyathium 1; floral bracts usually with pale or colored nvolucral gland cup-shaped; floral leaves green, white, or ngulate, coarsely tuberculate

2. E. cyathophora

.7. E. ocymoid

... 8. E. xalapen

olucres and capsules pubei-eent. Inflorescence glandular-pilose; leaf blades mostly 1 cm long Inflorescences nonglandular; leaf blades mostly longer than ! 9a. Cyathial glands 4 or 5, plane 9b. Cyathial glands 2, bilabiate

olucres and capsules glabrous. i. Cyathial appendages obsolete; glands with 2 conspicuous lateral horns 10. E. peplus >. Cyathial appendages present; glands elliptic.

1 la. Seeds terete, the surface smooth; cyathia ca. 2 mm in diameter; glands 4, dark; appendages equaling glands, ciliate-pubescent above 11. E. dwyeri

1 1 b. Seeil^ Mrough angled, the surface deeply punctate with pits in regular longitudinal rows; cyathia ca. 1 mm in diameter; glands 4 or 2, green; appendages obsolete (Panamanian specimens) to prominent and white, several times size of gland,

42.3. Euphorbia elata Brandegee, Univ. Calif.

Publ. Bot. 6: 55. 1914. TYPK: Mexico. Chia-

pas: Finca Irlanda. I'urpus 7()2() (holotype,

UC; isotypes. A, BM, F, GH, MO, F neg.

62362, UC, US).

, 600-700 m, Standley & I alerio ype, US; .sotype, US).

ill tree to 3.5 m high, glabrous;

none. Leaves alternate, clustered

petioles 2 4( 6) cm long, ca. 2

es fleshy, light-colored, debate to

im long, 3-4 mm broad; blades

runs. oblanceolate or narrowly el- Bot. Gart. Berlin-Dahlem 11: 137. 193 I ) is known

liptic, the apex rounded to bluiith short-acuminate only from the type collection, and no original ma-

at tip, acute to cunentc at base, 1 5 35 cm long, terial is definitely known to be extant. Until either

4-10(-12) cm broad, 3-5(-7) times as long as new or original material is available, the status of

broad; midrib prominent below, to 2 mm thick; that species cannot be determined. Euphorbia col-

lateral veins 17-30 [.airs on a side, obscure, nearly pansa Ducke (Arq. Inst. Pesq. Agron. 1:21. 1938),

at right angles to midrib; margin entire. In/ions J< . i bed li. •• •-:< n \i c.i/om.iu Hrazil. appears

cences single, terminal or rarely axillary, long- to be synonymous with /*.'. data. Another collector

pedunculate, cymose; peduncle 10-30 cm long, from Bolivia (Hang 619, MO, NY, US) was given

glabrous, minutely brown-puberulent, or covered an unpublished name but will probably also prove

with a waxy brown reticulum; cyme up to 5 times to be E. elata.

divided, the cyathi

groups; bracts opposite, scalelike, attached S peri mens examined. PANAMA. HOCAS OKI. TOKo: road

to Chiriqui Grande, 300 m, McPherson 10085 (MO), stem along a broad base, debate, ca. 4 mm long. I:(,I6N: trail from end of Santa Rita Ridge Rd. to Rio 2.5-3 mm broad; margin entire, sometimes ciliate, Pedros, 600 in, Antonio 3750 (F, MO); Distr. Portohelo, hyaline; apex blunt, somewhat cucullate. Involu- stream off N slope of Rio Gatun, 2,200 ft., .^»ni« :i«04

, . , , . , , (MO): Saul., Hita. I. ol moiiiitaiiioii.s /one, Coma & cres campanulate, glabrous, minutely brown-pu- /W(,r <,,-.,„„ . . , ,.,, K of Colon,/W berulent or covered with a waxy brown reticulum, lrr :u:{8 (PMA)i 3348 (F> PMA). Santa Hita ,umber

green, drying to brown, 4 6 mm high, (3-)4-5.5 road, ca. 15 km K of Colon, Dressier & Lewis 3728 mm diam.; pedicels 1.5 3 mm long, 1 1.5 mm (M<>. PMA); Santa Hita Ridge, 4 hour walk from end of

thick; lobes 5, 1.5-2 mm long, ca. 2 mm broad, •ad' "am^lJ318 2*°* J*'• ^ X, w• i fimbriate, rarely entire, e

• dark red; glands 5, round, 1.7-2 mm d.am., H«v., \hr, A kalian/,, .'/,/ (\IO. Mi -una

:he, 10-20 m, Kennedy & Foster 217 Rita Ridge Rd., ca. 6 km from Boyd-Roosevelt

placed vertically on run of involucre, looking out- Ridge, wards, but positions of lobes making glands appear Mori & Kallunki 3053 (MO, NY), DAKIEN: Cerro Pirre, L 1 • 1 J .• 1111 HI -ill below nm; gland some,,, ere< . and tin, k ^lalk, ,1

so that surface is flat with respect to orifice, the ot a.rro Mt._ 2,400 ft., Antonio et «/. 3426 (F, MO); margin then crisped: ippend: -f none, (.apsulcs jiM helore la f.neida along new trail beside Lopez House,

Tted from the cyathium no more than 1 mm '"'"" ^ /''••-''•' -V / .Ml KK. : ereen elabrous 8-9 mm 816 (M0); E1 Llano-Carti road, l«*-io "' K -'*"" '"_• ' mm »:|„.|1(1, 400 ,„, Correa et al. 18f>7 (I'M U Kl I

high, 10-1 i mm uiam.; siy.es ca. i.j mm .ong, Cart( road ,2 m[ aboye pan.Am Hwy U(,sner [2U

united at base for ca. '/, of their length, strongly (MO); El Llano-Carti road, 11 km from Pan-Am Hwy., recurved, bifurcate: seeds subglnbose, truncate at Mori & Kallunk, 3057 (MO, NY). \KHACI AS: Escuela

regular longiliid

green forests of low to middle

1 diam., ca. 4.5 mm long, ecarun- Agricultura, Alto Piedra near Santa le

iwn. wtlli lighter, low, broken, r to top of Cerro Tute, 3,400-3,800 ft., Antonio 34 (MO); 6.4 km outside of Santa Fe on road past agriculu school, toward the cordillera. Poison, 2<>7<> (V, MO).

42.4. Euphorbia leucocephala Lotsy, Bot.

'" T T^ T" * ' ""<"ru*' Gaz. (Crawfordsville) 20: 350, pi. 24. 1895. Colombia, and may occur as ar south v „ ' TT , F „ ,

LECTOTYI'K: Guatemala. Huebuetenango: Cuil-

rangc from

olivia. It is the most widespread spe

norima (Raf.) Webster, a group man 305 (presumably

by Standley & Steyer- highb divergent species of trees and shrubs lrom ..... J - .,,,.- ,, „, . , ,. . , . ... i - I I ' I I i.fil: 10, . I'» I0|. the West Indies, western and southern Mexico,

and the northern Andes. The species closest to E. Shrub to 3 m high; branches terete, glabrous,

elata are all highly restricted in range and are swollen at the nodes. Leaves verticillale; petioles

poorly understood. Euphorbia sinelairiana Benth. (1 )2-6 cm long, slender, glabrous; stipules glan-

(in Seemann, Bot. Voy. Sulphur 163. 1844), known duliform, 0.3-0.5 mm long; blades glabrous, ellip-

only from the island of Gorgona off the Pacific tic to linear-elliptic, mucronate and rounded or

coast of Colombia, is characterized by a deeply bluntly acute at tip, acute at base, (2)3 7 cm

bifurcate and much-branched inflorescence with long, (0.7 )1.5 2.7 cm broad, 2-3(-4) times as

conspicuous foliaceous bracts, but is otherwise sim- long as broad; margin entire. Inflorescences ter-

ilar to E. elata. The Peruvian E. tcssmannii Mansf. minal, cymose; bracts white, narrowly spatulate;

(Ber. Deutsch Bot. Ges. 46: 674. 1929; Notizbl. petioles 3 7 mm long, thinly pilose; blades 5-10

mm long, 1.2-2 mm broad; base narrowly acute;

margin entire; apex rounds I

on peduncles 1-2.5 mm long; involucre campan-

ulate, 1 -1.8 mm high, 1.1 -1.7 mm diam., densely

tomentosejustbelowtheizl.il i

pilose; glands 5, green, 0.8-1 mm long parallel to

the rim of the cyathium, 0.4-0.5 mm broad, the

center of the inner margin strongly inflexed forming

a deep convex trough; appendages white, linear-

lanceolate, exceeding glands by 2-3(-3.5) mm,

0.5 1 mm broad, the apex narrowly rounded.

Gynophore erect, exserted from cyathium 1-1.5

inn ' - J >u-. smooth. 5-6 mm high,

4.5-5.5 mm diam., shallowly 3-lobed, the cocci

distinctly 3-angled; styles 0.6 0.7 mm long, united

at base, deeply trifid, the style branches strongly

recurved; seeds (immature) ca. 3.5 long, trigonous,

ca. 1.8 mm wide, carunculate.

The report- <>1 V

represent a considerable range extension; the pre-

viously known range is from western —' - '•-

from cultivated tre

Capsules are ur Panamanian collec-

collect Our ilcM-ripli.:

si the i apsules is taken from a specimen collected

in Depto. Hueluietenaiigo. Guatemala I Malum

Specimens examined. P\\\\l\. mci.K: El Valle de Ant6n, 1,000-2,000 ft., Lewis et ai. 2570 (MO); El Valle, Ramos 19 (MO, PMA). PANAMA: Panama Viejo, Giron I (MO); near Cerro Azul, Cambra 49(MO, PM \)

.6. Euphorbia apocynoides Klotzsi

Seemann, Bot. Voy. Herald 99. 1853.

Panama. Darien: Punta Garachine, See,

1096 (holotype, BM; isotype, K; phc

isotype, K, MO).

hrub to 2 m high;

•; petioles slender. [ cm long. Leaves ter-

nm long, very sparsely

ibove, lighter or even somewhat glaucous

Dblong to narrowly obovate, rounded at tip,

t base, 4.5 6 cm long, 1.5-2 cm broad,

times as long as broad, glabrous or with

AT hairs below; margin entire. Inflorescences

n. Cynthia turbinate, ca. 2.5 mm high, ca.

l diam. below the appendages, sparingly to

evenly appressed-pubescent, the hairs short,

straight, nonoverlapping; peduncles 3-4 mm long,

subglabrous; appendages 5, narrowh l 1 «• • 11 •

pul ! i below toward the

base, ciliate on the margins near the juncture with

the involucre, otherwise glabrous, entire in the

lower half, deeply and coarsely crenellate along

the distal margin. Flowers and fruits not seen.

This poorly known species is apparently endemic

to Panama and is still known only from the frag-

mentary type collection and an equally fragmen-

tary recent collection from the type localit] I I <

type is sufficiently complete, however, to enable

the species to be placed with reasonable certainty

in section tlcctom- tonum (Nhldl.) Urn- icr. a , rou|>

characterized by verticillate branching and swollen

nodes, and to say that it is unlike any other species

in the section. Contrary to the statement in the

original treatment (Webster & Burch, 1968: 335),

the cyathium is quite unlike that of sect. I h. hthnin

Boiler. ulu< h is cl .<t acleri/ed l>\ ;i rciluccil nuiii-

ber (usually 2) of bilabiate glands and small, erect,

or often obsolete, appendages. The species of sect. i I .. , i i i i ill.-male leaves

and stems that are pinched just above the nodes.

A plant vegetatively very similar to Euphorbia

apocynoides, collected near Puerto Colombia on

the Caribbean coast of Colombia {Elias 1197, F),

has been identified as E. nudiflora Jacq., a West

lul i |i II athium, however, differs from

the pubescence is crisped with overlappm, t • u

and the appendages are completely glal I

ovate, and with smaller and more numerous cren-

ellations on the distal margin. In all of these

characters it matches E. nudiflora, which differs

vegetatively in its more highly branched habit and

shorter, broader leaves. An elucidation of the re-

lationship of the Colombian plant to the Panama-

nian one, and of both to E. nudiflora, must await

fuller collections from Panama and Colombia.

We are indebted to Mr. A. Radcliffe-Smith of

K. ! i ,i and a sketch of

the cyathium from the isotype of E. apot ynoides

at K, from which our description has been taken.

Additional collection examined. I'wwu. nuniv thorn forest near Punta Garachine, Duke 10485 (MO,

2.7. Euphorbia ocymoidea L., Sp. PI. 453.

1753. TYPE: Mexico. Campeche: Houston s.n.

E. astroites Fisch. & Mey.,

Ill: II. I «-l-r>. ni'i: Mexico: I'ampa. uala, /v»i m|, „ .(.,|. ()f,,.n aborting, ([vat hid on pedicels 1-3 wmsky (LE, not seen). mm [ong. invo|urre (.ampaiiiilate, 0.7-1.3 mm high,

A thorough examination of collections from 0.5-1.3 mm diam., crisp-pubescent; glands 5,

throughout the range of this species (western Mex- green, elliptic to reniform; appendages whitish or

ico to Panama) makes it clear that Euphorbia greenish, broadly ovate, exceeding the gland by

astroites cannot be separated from E. ocyrnouica. 1 2 mm, 1.5 2 mm broad, the margin entire or

The only consistent character separating the two crenate; gynophore glabrous, erect or somewhat

is the glandular-pilose stems of the former, as op- recurved, exserted from the cvathium 1.5-2 mm.

posed to the glabrous or short-pilose, eglandular Capsules sparsely to densely pilose, 1.5 1.9 mm

stems of the latter. The two forms occupy roughly h'Bn< l L5 """ ,liam-: s,vlrs °~ (KB """ long< the same geographical range (the glandular form free to the base, deeply bifid, thinly pilose; seeds

is not yet known from western Mexico, and the ca. 1.5 mm long. 0.9 1 mm diam., ovoid, grayish,

eglandular form has not been collected south of coarsely pitted, tuberculate, ecarunculatc.

Nicaragua) and the same habitats, and they exhibit Euphorbia xalapensis is a common spec.es of

'^arly in the shape f(m,st nor(jers an)) thickets that ranges from west-

exico to Honduras, .iml thus ihe new reports

lere from Panama and Costa Rica represent

siderable range extension. This species was

:ed in the Flora of Guatemala

(Fieldiana, Bot. 24(part 6): 108. 1949) as E. oer-

stedianu (klol/seh \ Carcke) Boissier. a very dif-

lercnl species that has mostly glabrous stems, ar-

ticulated nodes, two cyathial glands that are more

or less bilabiate and that have inconspicuous or

obsolete appendages, and densely while pubescent

capsules. The latter species, winch belongs to sec-

tion Diehilium Boissier, is very rare in Central 42.8. Euphorbia xalapensis kuuth. Nov. Gen. America and is somewhat better known from the

Sp. PI. 2: 61. 1817. Pomsettia xalapensis West Indies and northern South America. It is still

(Kunth) Klotzsch X Carcke, Monatsber. ko- ,mknown from Guatemala, and nearly all of the

nigl. Preuss. Akad. Wiss. Berlin 1859: 253. pil,ative collections from there are referable to E.

1859. TYPK: Mexico. Veracruz: near Xalapa, xalapensis. It has not been re-collected m Panama

Humboldt ,K Honpland s.n. (P, not seen). sinre the collection cited in the original treatment.

Euphorbia enalla Brandegee, Univ. Calif. Publ. Bot. 6: Euphorbia xalapensis is also frequently c~~ 54. 1914. TYPK: Mexico. C quer6n, Purpus 7f>:t.> (h..l..t> neg. 60269, GH, MO, NY:

iinuii ) is le namej. similar in aspect. Euphorbia xalapensis. however, -M.mdlev, Publ. Field (.olum- ,,.,,,,,., . ,

i.ian Mils Bot Sn |- 313 L929 rYPl Honduras r'1" ' ''" " ' ' "iK perennial and Comayagua: near Siguatepec, Feb. 1928, Standley often strongly rlii/omatous habit; the shaggy brown :,(>.i 11 (holotvpe, K: isotypes, F, F neg. 60244, US). pubescence of the stems, leaves, and inflorescence;

Perennial rhizomatous herb to 50 cm high, the |he- °ne-sided appearan

of the leaves, wh ich range Iron i broadly ovate or

deltate to somewhat reniform. As suggested by

McVaugh (1961 : 177), several names based on

collections from western Mexico, particularly E.

subreniformis S. Watson, undou btedly belong here.

The only known Panamaiii an collections are

glandular-pilose, a fact that wa; 3 omitted from the

original treatment , and thus woul d have been placed

under E. astroite x. The species has apparently not

been collected in Panama since the appearance of

the original treat

stems shaggv brown pilose. Leaves opposite 3 due to the frequent

ticillate above, alternate below; petioles 1.5 2.5 ",,(l,': ;""1 ,1"' "v«* Pla»"* ,,,a' have amPle' ^^

cm long, brown curly-pubescent; stipules glandu- Sreenish ^"dages.

liform, brown or black, minute, 0.1-0.2 mm long; Additional specimens examined. COSTA RICA, SAN

base, dark green, 1.5-3.5 cm long, 1 2 cm broad, creek, NK pa. t... • ! • • \. 8 H)3'W, 1.700-

1.5-1.8 times as long as broad. denseK shauuv '•^ ' ': C?'?>•. .* S^% 5'5< ^\ *'"""• i li:,;|. i l \ .4 .a. i ! ill. I.. ., • I

pilose below, more sparsely so above, base rounded t)fill , (lm N],„ ,,M, ,,,„,,,, , - , „,,,„„,„„ ,„„„„„,„ to obtuse; margins entire, ciliate. acute. In/lores- Jacq. (s. lat.)/'

eenei-s lermmal, < \ iiiose. often appearing one-sided

by the abortion of one branch at a node, shaggy- 42.10. Euphorbia peplus L., Sp. PI. 456.

pubescent; bracts similar to the leaves but greatly 1753. TYPK: Europe (presumably in Hortus

; 630.24 43. Cha

lal glabrous herb 18-35 cm high. Leaves

e below, opposite or ternate above, sessile

t-petiolate, numerous, early deciduous be-

pules obsolete; blades bright green, mem-

0 Mi i long, 8-12 r

flexed, congested. Cyathia solitary in forks of up-

per branches, on peduncles 0.6-1 mm long; in-

volucre < ampanulate, light green, ca. 1 mm high,

0.5 0.7 mm diam.; glands 4, green, crescent-

shaped, exappendiculate, ca. 0.5 nun IOIIL' pavall.

to the rim of the involucre, with prolonged narrow

horns 0.5-0.7 mm long; gynophore exserted, ca.

1.3 mm long, recurved. Capsules green, 1.8-2

mm high, 2-2.2 mm diam.. broadest below the

middle, shallowly 3-lobed, the cocci each with 2

nam*-, I,, i- i i, n i 11 ,.•-.-iv |r-cu. 0.1 mm long,

bifurcate, the style branches bifurcate; seeds ovoid-

oblong, weakly 6-angled, gray, ca. 1.5 mm long,

ca. 0.8 mm diam., carunculate, deeply pitted, mi-

This is apparently the first report from southern

Central America of this cosmopolitan weed of tem-

Speetmens examined P\\\\l\. < IIIHIQUI: trail from Paso Respingo to Bajo, Chorro Cerro Punta to Boquete, along stream near Guadalupe, llammch-t «l. 7077 (MO);

Cerro Punta dairy, H,01)0 ft.. />' hey el ai 13193 (F,

Chamaesyce Gray, Nat. Arr. Brit. PI. 2: 260.

1821. TYPE: Chamaesyce maritima Gray =

C. peplis (L.) Prokh. (Euphorbia peplis L.)

(See Wheeler, 1943: 461, for a discussion of

dilanthus Necker ex Poit., Ann. Mus. Natl

Hist. Nat. 19: 388. 1812. TYPE: Pedilanthiu

tithymaloides (L.) Poit. (Euphorbia tithy

maloides L.).

[>AT, T. 1978. Flora of Barro Colorado Island. Stan- ford Univ. Press, Stanford, California.

ESSI.KR, R. L. 1957. The genus Pedilanthus (Eu- phiH l.ii. cu). Contr. Cray Herb. 182: 1-188. . 1961. A synopsis " "

. F. 1971, 48: 32' ;rowth of tropic

I !"• trees exemplified by the Euphorhii 3: 56-62.

MS, A. S. 1973. Chromosomal conspectus of the Euphorhiaceae. Taxon 22: 591-636.

LONSKI, E. 1965-1967. Euphorhiaceae. In: B. Ma- Miiiir (editor). Botam ot tlit- Ouayana 1 lisihlainl \ I. Mem. N.Y. Bot. Gard. 12(3): 150-178. 1965. VII.

VAUGH, R. 1961. Euphorhiaceae novae Novo-Ca- licianae. Brittonia 13: 145-205.

DeCandolle,

ama. Chiriqui: Cerro Horqueta, NW

1979. Acalypha, Croton, and Sapium i. Phytologia 43: 133-195. 1937 1938. Flora of Costa Rica. Publ.

E. G. (Chairman. Editorial Com

— & D. BURCH. 1968. Euphorhiaceae. In: Flora f Panama. Ann. Missouri Bot. Gard. 54: 211-350. — & E. Rl'PERT. 1973. Phylogeneti. significance

ation 27: 524 531.

/ Lopez & Pavon sejm ntrioiidhs L. 0. William

Clei.ii -., illume 1087, 1103 . . uin Pax 1104

obm ata Benth. 1104 membranaceum Pax & K. Hoffm. 1099, 1103, 1104 virginica L. 1104 ?nicaraguensis Hemsley 1106

Acidoton Sw. 1106 ohlongifolium (Standley) Croizat 1103, 1104 iiirariign«Mi.sis (Hemsley) Webster 1106 prealtum Croizat 1103, 1104 urena Sw. 1106 tricoccum (Casar.) Baillon 1103

kctephik Itlurne 1092 woodsonianum Croizat 1099, 1103 Actinostemon Klotzsch 1129 Cnidoscolus Pohl 1114

conrolur (Sprengel) Muell. Arg. 1129 aconitifolius (Miller) I. M. Johnston 1114 Adelia L. 1100 subsp. aconitifolius 1114

ricinella L. 1100 adenophllus (Pax & K. Hoffm.) Pax & K. Hoffm, Adenophaedra (Muell. Arg.) Muell. Arg. 1087, 1099 1114

grandifolia (Klotzsch) Muell. Arg. 1099, 1100 hamosus Pohl 1114 rnegalophylla (Muell. Arg.) Muell. Arg. 1099 wootisoniunti (Croizat) Croizat I !><><>

Alchornea Sw. 1087, 1100 costaricensis Pax & K. Hoffm. 1101 glandulosa Poeppig

var.floribunda Benth. 1098 m glandulosa 1102 var. pittieri (Pax) Pax 1102 panamensis Webster 1095

grandiflora Muell. Arg. 1102 Croton L. 1116 grandis Benth. 1102, 1103 argenteus L. 1124 latifolia Sw. 1100, 1101, 1102 aromaticus L. 1116 rnegalophylla Muell. Arg. 1102 benthamianus Muell. Arg.

- Arthur 111 „.!<,• ././../..- (Pa: K & K 14

jrens 1114 ryerm. 1087, 1092

. 190 i Steyerm. 1092, 1093

pittieri Pax 1102 "'!'!. ^"•-•: Muell. Arg. 1101

-ii. \lu.

brevipes Pax 1123 castaneifolius L. 1098 draco Cham. & Schldl. 1

'-.;•« nr.. illume 1093 Argythamnia P. Browne 1098

candirans Sw. 1098 Astro* a .a Robinson & Millsp. 1087, 1091

I - , n *N M I (. in I tremula (Griseb.) Webster 1091

ll •»,, Browne I \2t) Bernardia P. Miller 1100

• j i n- I Miller 1100 denticulata (Standley) Webster 1099, 1100

r i Muell. Arg. 1099 jacquiniana Muell. Arg. 1100 MUHTophvlla Standley 1 100 ?rnegalophylla Muell. Arg. 1099 Cyclostigma Klotzsch

Capemnu St. llil. 1098 panamensis Klotzsch 1 120 eastaneiiolia (L.) St. Hil. 1098 Dactrlostemon Klotzsch 1129 paludos.1 Klotzsch 1098 l)ale,l,ani|)ia L. 1087, 1108

Car\oden.lfon Karsten 1099 section Dioscoreiloliae Pax & aiigiisiilolinin Standleyy 1099 canescens Kunth

Webster & Huft

i Poeppig 1 frx In • thalii Muell. Arg. 110 f.rtrmnmrnhu Pax & K. Hoffm panamensis Pax & K. Hoffm. 1 scandens L. 1108 shankii(A. Molina) Huft 1107,

(Pax & K. Hoffm.) Webster 1110

guatemalensis J. D. Smith 1095 laxiflora (Tul.) Muell. Arg. 1094, 1095 oblonga (Tul.) Muell. Arg. 1095

var. benthamii (Tul.) Muell. Arg. 1095 scabrida (Tul.) Muell. Arg. 1095

Kunth 1113 atropha L. 1114

icomHfoUui Miller 1114 adenophila Pax & K. Hoffm. 1114 gossypiifol manihot I

puluHa I..

montana Muell. Arg. 1125

section Poinsettia (Graham) Baillon 1137 Maesobotrya Benth. 1093 subgenus Poinsettia (Graham) House 1094, 113' 1 Manihot Miller 1087, 1113 umphimalava Standley 1140 aesculifolia (Kunth) Pohl UK antiquorum L. 1137 brachyloba Muell. Arg. 1114 apocynoides Klotzsch 1137, 1139 esculenta Crantz 1113 astroites Fisch. & Mey. 1139 unnUuiensis Blake 1113 capansa Ducke 1138 Maprounea Aublet 1087, 1131 caracasana Boissier 1141 guianensis Aublet 1131, 1132 dwyeri Burch 1141 Margaritaria L.f. 1096 elata Brandegee 1137 nobilis L.f. 1096 enalla Brandegee 1140 OmphaleaL. 1110 graminea Jacq. 1140 leucocephala Lotsy 1138

diandra L. 1110 triandra L. 1110

nudiflora Jacq. 1139 Ophellantha Standley

f'a.wu-Jra Kadlk. 1087, 11

quadnglandulosa Pax & K. Hoffm ... Baillon 1115

Pedilanthus Necker ex Poit. 1094, 11 tithymaloides (L.) Poit. 1141

Pentabrachium Muell. Arg. 1092

hifiltimliilosti (I..) Muell. Arg. 1133 var. moritziutm (Klotzsch) Muell. Arg.

, , • Griseb. 1131

Gussonia Sprengel concolor Sprengel 1129

Gymnanthes Sw. 1087, 112

dressleri Webster 1131 farinosa (Griseb.) Webste granatensis Muell. Arg. 11

subgenus Xylophylla (L.) Pers. 1098

anisololm Muell. Arg. 1096 gentryi Webster 1096, 1097!

s Willd. 1098

t m I G ;b. 1091 Plukenetia L. 1105

angustifolia Standley 1105 penninervia Muell. Arg. 1105

pulcherrima (Willd. ex Klotzsch) Graham 1 xalapensis (Kunth) Klotzsch & Garcke 11

Polyandra Leal bracteosa Leal 1103

Richeria Vahl 1087, 1093 dressleri Webster 1094 grandis Vahl 1093, 1094

i P. ItmvwL- 1087, 1129, 1132

i Jacq. 1132, 1133,

Senefeldcra C. Martius multiflora C. Martius

Stilaginella Tul. benthamii Tul. 1095

oblonga Tul. 1095

viir. muiitzimmm (KIOI/MI,) Pittier 1 133 • >/,., Sim.lle, 1 1 : aucuparium sensu Croizat 1134 svlvatica L. 1132 higlanilulosum (L.) Muell. Arg. T.-tmr.-l.idi.u.. Poeppig 1087,

hrevifolium Standley & Ste* var. moritzianum (Klotzsch) Muell. Arg. 1133 costaricense Huft 1112

1134 euryphyllum Staiult.-v III.' var. sulnfrrum (Pittier) Monach. 1134 gorgonae Croizat

,„„,/„„„„ Pittier 1132, 1133, 1135 subsp. robledoanui i (1 ual mi.-.-..,,l»ll..m Hut! Ml'

i;'ii;«N/.'i(»( Pit tier 1133 mnlmur I.. 0. Wilham^ 1 1 jamaicense Sw. 1132, 1133, 1134 robledoanum Cuatr. 1113 n r t i / Kl t?s [ 1 1 a rotundatum Standley 1112

oligoneurum Schumann & Pittie: 1 1 3 1 pachwach\-, Schumann i\ Putin 1135 Thouinia Poit. plvmshirhvs Schumann & Pittier 1134 uutumayen.se Croizat 1136 Tragia L. 1087, 1106 rigulifolium Huft 1135, 1136! — II Jia (kl.,l/,chl \lu.-ll salicifolium Kunth 1133 -,-,„,,„ Euttnuio Muell \r> schippii Croizat 1134 section Tragia 1108

i Zuckcrtia (Buillon) Muell. \<r.

conlata Michx. 1 106 correae Huft 1107

v,is(i- of the Missouri Botanical Garden 1988...Missouri Botanical Garden, 2345 Tower Grove Av-...

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Transcript of v,is(i- of the Missouri Botanical Garden 1988...Missouri Botanical Garden, 2345 Tower Grove Av-...