A Synopsis of Phyllanthus Section Nothoclema (Euphorbiaceae)

NUMBER 6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 19

A SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA (EUPHORBIACEAE)

Grady L. Webster Herbarium, Section of Plant Biology, University of California, Davis, California 95616

Abstract: Phyllanthus section Nothoclema, after revision, includes 10 species that are notable in the genus for their use as "barbascos" (fish poisons). The section is remarkable for its palynological diversity, which provides important diagnostic char acters for distinguishing species. Two new species are described, both from Vene zuela: P. liesneri from Falcon and Yaracuy; and P. meridensis from Merida. Al though pollen material is lacking for study, both species appear related to congeners with 3-colporate pollen having a vermiculate-reticulate exine. Phyllanthus graveolens is treated as a polytypic species, including P. micrandrus as a subspecies. Two sub species are recognized within P. brasiliensis, the species most commonly used as a fish-poison; ssp. brasiliensis throughout most of its range includes mostly sterile semi domesticated clones.

Keywords: Euphorbiaceae, Phyllanthus, subgenus Conami, section Nothoclema, Ve nezuela, barbascos.

INTRODUCTION widespread use as fish-poisons in South America, where they are commonly re Phyllanthus, the third largest genus of ferred to as "barbascos," they also have Euphorbiaceae, has not been revised at the many indigenous names (Radlkofer, 1886; species level on a worldwide basis since the Acevedo-Rodriguez, 1990). Uses as a fish monograph of Mi.iller (1866). About a poison have been recorded for three spe quarter of the 800 species of Phyllanthus cies: Phyllanthus acuminatus, P. anisolobus, are neotropical, but only the West Indian and P. brasiliensis but in the Amazon re species have been revised monographically gion all reports of use as barbascos are (Webster, 1956-58). In preparation for a based on P. brasiliensis. References to pis synopsis of all of the neotropical species, a cicidal species of Phyllanthus in the litera number of revisions have appeared (Web ture are confused because species delimi ster, 1970, 1991, 1999, 200la, 200lb, tations and nomenclature have remained 2002). unclear. It is not surprising that a large Phyllanthus section Nothoclema G. L. percentage of the specimens of section Webster was described (Webster, 1957) to Nothoclema in all herbaria are misidenti include woody plants having compound fied, because the morphological variations (bipinnatiform) deciduous branchlets with in foliar and floral morphology are rather axillary cymules of flowers mostly on the subtle, and it is often difficult to identify lateral (ultimate) axes (Fig. 1). The hex species without flowers or fruit. Prepara amerous flowers have a well-developed tory to a monographic revision of the sec disk, and the androecia each consist of 3 tion (Webster, ined.), this synopsis is pre connate stamens dehiscing more or less sented here with validation of new taxa horizontally. With 10 species and 5 sub and new combinations. As treated in this species, it is the largest of the three sections new revision, 10 species are recognized, in of the neotropical subgenus Conami addition to several subspecific taxa. (Aubl.) Webster. A notable aspect of taxonomic diver The species of section Nothoclema are sity within sect. Nothoclema is the striking of ethnobotanical interest because of their variation in pollen morphology. As dem-

LUNDELLIA 6:19-36. 2003 20 LUNDELLIA DECEMBER, 2003

MISSOURI BOTANICAL GARDEN HERBARIUM 11 ~111111111111111111 !ill N? 4361313

lln-bmriM, Unh~,,.lly of CaUfom!a. Dnb {DAV)

The New Yorl-. Botanical Ga rden Plants of BOLIVIA

Phyllanthus brasilicnsis (Aubl.) Poir, dct. I. Vorgns C., 1992

Depto, Santa Cruz , Prov , Andres !bane!'.!, along road from &ints CTuz to Samaipsn1, 1 km. SI.' o f Angostura. ca. JB 0 09'S, 63°3I'W. alt . 650 tn,

Subtropical semi-deciduous forest and road sides in gorge of Rio Pirni as it emerges from the castt>rmrost foothills of the Andes.

Young s hrub I m. tail. Flowers hangfng , pfnkish.

Coll.: H. Nee 33502 13 Jan. 1987

FIG. 1. Habit of Phyllanthus brasiliensis ssp. glaber (Nee 33502, MO). NUMBER 6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 21

onstrated by Webster and Carpenter KEY TO THE SPECIES OF SECTION (2002), pollen grains of several species NOTHOCLEMA (Phyllanthus anisolobus, P. graveolens, and 1. Branchlets and leaves glabrous (petioles rarely P. mocinianus) have tricolporate pollen sparsely scab rid or hirtellous). grains with elongated colpi and exine 2. Branchlets simple or occasionally with 1 or sculpturing that varies from reticulate to 2 lateral axes; seeds smooth or nearly so. vermiculate; two species (P. acuminatus 3. Branchlets strongly compressed-ribbed; leaf blades narrowly elliptic to obovate, and P. mcvaughii) have crassimarginate cuneate at base, mostly < 1 cm broad; colpi and pilate exines; and in two species fruiting pedicel winged-angled, 5-7 mm (P. brasiliensis and P. pavonianus) the pol long; pollen grains pantoporate, exine len grains are pantoporate with pilate ex pilate ...... 10. P. pavonianus ines. These three palynological groups ap 3. Branchlets angled, not ribbed; leaves ovate, rounded at base, mostly > 1 cm pear to be diagnostic in recognizing three broad; fruiting pedicel terete, 8-12 mm clades within sect. Nothoclema. However, long; pollen grains colporate, exine re- because pollen morphology has not been ticulate ...... 6. P. caymanensis determined for all taxa, and the floral and 2. Branchlets with 2 to 20 or more lateral axes palynological characters appear imperfectly (rarely unbranched); seeds puncticulate. 4. Styles emarginate, erect, coherent into a correlated, formal subsections are not pro column; pistillate sepals 2.5-4 mm long posed here...... 5. P. meridensis 4. Styles bifid, spreading or ascending, free or basally connate; pistillate sepals 1-2.5 SYSTEMATIC TREATMENT mm long. 5. Bracts of cymules inconspicuous, Phyllanthus section Nothoclema G. L. not scarious-indurate; leaf blades Webster, Contr. Gray Herb. 176: 56. obtuse or rounded at apex, mostly 1-4 X 1-3 cm. 1955; J. Arnold Arb. 38: 363. 1957. 6. Stamina! column 0.7-1.2 mm TYPE: Phyllanthus acuminatus Vahl. long; anthers distinctly deflexed; styles 0.7-1.1 mm long; branch lets with 5 to 10 lateral axes; vein MONOECIOUS SHRUBS OR TREES with lets of leaf blade abaxially prom phyllanthoid branching: leaves of the per inulous . . . . . 1. P. anisolobus manent axes reduced to cataphylls; decid 6. Stamina! column < 0.7 mm uous branchlets mostly compound (bipin long; anthers not distinctly de flexed; styles not over 0.7 mm natiform). LEAVES of the branchlets disti long; branchlets with 2 to 15 or chous; blades chartaceous, elliptic to orbic more lateral axes; veinlets of ular, entire, pinnately veined. FLOWERS in leaf blade not abaxially promi axillary, mostly bisexual cymules, on lateral nulous (except in P. mocini branchlet axes and sometimes on main anus ). 7. Branchlets with mostly 2 to axes. STAMINATE FLOWERS distinctly pedi 4(-10) lateral axes; fruiting cellate; sepals 6, ± biseriate; disk entire to sepals 1.5-2.5 mm long. dissected; stamens 3, filaments connate, an 8. Staminate pedicels 5- thers dehiscing horizontally. POLLEN 15 mm long; sepals 1.7-2.2 mm long; an GRAINS spheroidal, 3-colporate or panto thers emarginate, not porate, exine reticulate or vermiculate to pi compressed; seeds 1.7- late. PISTILLATE FLOWERS pedicellate; se 2.1 mm long; veinlets pals similar to the staminate; disk usually of leaf blades promi cupular; ovary 3-locular; styles mostly free nulous abaxially ..... or basally connate, bifid or bipartite. . . . . 2. P. mocinianus 8. Staminate pedicels 1.5- FRUITS capsular; columella persistent; seeds 4 mm long; sepals 0.8- trigonous, smooth or puncticulate. 1. 7 mm long;· anthers 22 LUNDELLIA DECEMBER, 2003

acute, compressed; 1. PHYLLANTHUS ANISOLOBUS Miill. Arg., seeds 2.1-2.5 mm long; DC. Prodr. 15(2): 382. 1866; Webster, Ann. veinlets of leaf blades not (or scarcely) prom Missouri Bot. Gard. 54: 229. 1968. TYPE: inulous abaxially ... "Peruvia," J. A. Pavon (HOLOTYPE: G!)...... 7. P. mcvaughii The species has not been recorded from (glabrous form) Peru, and Pavon' s specimen was doubtless 7. Branchlets with mostly collected in Ecuador. (10-) 15 to 20 lateral axes; fruiting sepals 0.8-1.3 mm long. Phyllanthus pittieri Pax, Anal. Inst. Fis. 9. Leaf blades mostly Geogr. Nae. Costa Rica 9: 1895; in Pit broadly elliptic to sub orbicular, apically ob tier, Prim. FL Costaric. 2(5): 327. 1900. tuse or rounded; fruit TYPE: COSTA RICA. CARTAGO: Canton ing pedicels 3-20 mm Cartago, Aguacaliente, H. F. Pittier 2505 long .. 3. P. graveolens (LECTOTYPE, chosen here: B!; ISOTYPES: 9. Leaf blades mostly BR!, US!). broadly ovate, obtuse to acute at the apex; Phyllanthus mexiae Croizat, J. Washington fruiting pedicels 3-7 Acad. Sci. 33: 14. 1943. TYPE: ECUA mm long ...... DOR. "Leon" [Cotopaxi]: Canton Pujili, .... 9. P. brasiliensis near Santa Rosa, 1000 m, Nov. 1934, ssp. glaber 5. Bracts of cymules pale, scarious-in Ynes Mexia 6718 (HOLOTYPE: US!). durate, persistent; leaf blades elliptic to ovate, acute to breviacuminate, DISTRIBUTION AND HABITAT: Costa mostly 3-6 X 2.5-3.5 cm ...... Rica to western Ecuador, in lowland to up ...... 4. P. liesneri 1. Branchlets and leaves (at least the petioles) usu land rain forest or cloud forest, 100-1700 m. ally scabridulous or hirtellous. PHENOLOGY: Collected in flower April 10. Seeds puncticulate (rarely almost smooth); to October; in fruit July to August, Novem staminate disk segments 3, distinct; leaves ber to January. glabrous or if hirtellous (in P. mcvaughii) then anthers compressed, acute. 11. Leaf blades glabrous or hirtellous; an REPRESENTATIVE SPECIMENS EXAM thers compressed, acute; pollen grains COSTA RICA. colporate, exine pilate; leaves often red INED: NORTH AMERICA. dish ...... 7. P. mcvaughii Alajuela: Canton Naranjo, Austin Smith 11. Leaf blades glabrous (or hirtellus, not P2423 (A, UC). Cartago: Canton Turrialba, reddish); anthers not compressed, ob 6 km SE of Tuis, Webster & Raveret 22239 tuse or emarginate; pollen exine vermic (CR, DAV, MO). Puntarenas: Canton Coto u!ate-reticulate ...... 3. P. graveolens 10. Seeds smooth; staminate disk segments Brus, Finca Las Cruces, Webster 21946 sometimes coalescent; leaf blades hirtellous (DAV); Canton Golfito, Osa Peninsula, or scabrid at least adaxially. Liesner 3118 (MO). San Jose: Canton Tur 12. Leaf blades distinctly and abruptly acu Grayum et al. minate, scabridulous at least on midrib rubares, Cerro Turrubares, and petiole, and usually abaxially; sta 10533 (DAV, MO). NICARAGUA. Mata minate disk of 3 distinct ± bilobed seg galpa: Quebrada Santa Cruz, Stevens & Mo ments; styles spreading, ± flattened and reno 22325 (DAV, MO). PANAMA. Bocas dilated; main axis of branchlet without del Toro: Dto. Chaguinola, Rio Teribe, staminate cymules . . 8. P. acuminatus 12. Leaf blades acute or cuspidate, not dis Kirkbride & Duke 553 (MO). Chiriqui: Dto. tinctly acuminate, hirtellous rather than Gualaca, Fortuna Dam, McPherson 11686 scabridulous; staminate disk annular or (DAV, TEX). Code: Dto. Anton, El Valle lobed; styles ascending, subulate; main de Anton, Allen 3626 (F, MO, NY). Darien: axis of branchlet often with staminate cymules ...... Dto. Chepigania, between Cerro Mali and . . . . . 9. P. brasiliensis ssp. brasiliensis Cerro Tacarcuna, Gentry & Mori 13861 NUMBER6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 23

(DAV, MO). Veraguas: Dto. Santa Fe, Her REPRESENTATIVE SPECIMENS EXAM nandez et al. 744 (F). INED: BELIZE. Corozal: Lawry's Bight, SOUTH AMERICA. COLOMBIA. Ris Crane 202, 370, 486 (LL), 543 (TEX). EL aralda: Mistrato, Franco et al. 3455 (MO). SALVADOR. Ahuachapan: El Imposible, Sucre: Coloso, Gentry & Cuadros 68123 San Benito, Sandoval ]BL 1337 (MO). (MO). Valle del Cauca: Mpio. Buenaven GUATEMALA. Escuintla: 2 km N of San tura, Timbiqui, Lehmann 9016 (K). EC Jose, Morrison 8834 (UC). Huehuetenango: UADOR. Carchi: Canton Chical, Madison Quen Santo, Seler 3088 (GH, NY, US). et al. 4679 ( QCA). Esmeraldas: Canton Quetzaltenango: Rio Ocosito, Donnell Quininde, Bilsa Biological Station, Clark Smith 2622 (F, GH, US). Retalhuleu: 4 km 374 (US), Pitman 1148 (DAV, QCNE). Im W of Retalhuleu, Standley 87448 (A, F). babura: Canton Ibarra, Lita to Cachaco, Suchitepequez: 2-3 km N of La Maquina, Boyle et al. 3693 (DAV, MO). Los Rios: Proctor 27452 (DAV). MEXICO. Campe Canton Quevedo, Rio Palenque Station, che: Mpio. Champoton, Tuxpefia, Lundell Gentry 9918 (MO, US), 9977 (DAV, MO, 821 (GH, MO, UC, US, WIS). Chiapas: Q). Pichincha: Canton Quito, Bosque Pro Mpio. Ocozocuautla, 8 km S of Berriozabal, tector Maquipucuna, Webster et al. 32940 Breedlove 23071 (MICH). Chihuahua: (DAV, GH, QCNE, TEX). Mpio. Batopilas, Batopilillas, H. S. Gentry 2619 (A, MO, UC, US). Colima: Manzanil la, Ferris 6066 (A, DS, US). Guanajuato: This species was combined with Phyl Mpio. Atarjea, Mangas Cuatas, Ventura & lanthus mocinianus by Burger and Huft Lopez 8739 (DAV). Guerrero: Mpio. La (1995), but is retained here as a separate Union, El Cedral, Langlasse 304 (GH, US). species because of its morphological dis Jalisco: Mpio. Autlan de Navarro, 9-10 mi tinctiveness, with larger flowers, the stami SW of Autlan, McVaugh 14201 (MICH, na! column to 1 mm high, and anthers de US); Mpio. La Huerta, Estacion Chamela, flexed and distinctly emarginate. Lott 534 (DAV), 1661 (US), 3671 (DAV). Mexico: Mpio. Apatzingo, Sierra Nanchiti 2. PHYLLANTHUS MOCINIANUS Baillon, tla, Hinton et al. 5330 (GH, MO, TEX, US, Adansonia I. 1: 35. 1860; Mull. Arg., DC. WIS). Michoacan: Mpio. Coahuayano, 8 Prodr. 15(2): 382. 1866; McVaugh, Bot. Re km NW of Aquila, Rzedowski 17923 (ENCB, sults Sesse and Moc;:ifio Exped. 243. 2000; MICH). Morelos: Mpio. Yautepec, Canon Webster, Contr. Univ. Michigan Herb. 23: de Lobos, Rzedowski 12571 (ENCB), Web 383. 2001. TYPE: MEXICO. M. Sesse & J. ster & Armbruster 20673 (DAV, MEXU). M. Mo~iiio (HOLOTYPE: G, Delessert; n.v.). Nayarit: Islas Tres Marias, Maria Madre, Ferris 5569 (A, DS, US); Mpio. Compostela, La Cucaracha, McVaugh 19218 (MICH). Phyllanthus pringlei S. Wats., Proc. Amer. Oaxaca: Mpio. San Miguel Puerto, 3 km Acad. 26: 147. 1891. TYPE: MEXICO. SW of Copalita, Torres et al. 5226 (DAV, SAN LUIS POTOSI: June 1890, C. G. Prin MEXU). Puebla: Mpio. Tepexi de Rodri gle 3532 (HOLOTYPE: GH!). guez, 15.5 km S of Molcaxac, Chiang & Val iente 2007 (DAV, MEXU). Queretaro: DISTRIBUTION AND HABITAT: Western Mpio. Pinal de Amoles, Las Adjuntas, Za Mexico to Belize, Guatemala, and western mudio et al. 9045 (DAV). Quintana Roo: El Salvador, in tropical deciduous or semi Isla Mujeres, Webster & Lynch 17621 deciduous forests or woodlands, often on (DAV); Isla Cozumel, Gaumer 68 (GH). limestone, 0-1200 m. San Luis Potosi: Mpio. Valles, 9 mi E of PHENOLOGY: Collected in flower May Valles, Webster & Breckon 15475 (DAV, to August; in fruit August to January. MEXU). Sinaloa: Mpio. Rosario, Colomos, 24 LUNDELLIA DECEMBER, 2003

Rose 1774 (GH, US). Tamaulipas: Mpio. 3. Seeds 1.8-2.4(-2.7) mm long; fruiting ped Mante, Cafi6n

from which it differs mainly in lack of pu Subspecies novogalicianus was earlier bescence. referred to Phyllanthus micrandrus (Web ster, 200la), and it does seem to be most 3b. PHYLLANTHUS GRAVEOLENS ssp. nov similar to ssp. micrandrus, from which it ogalicianus G. L. Webster, subsp. nov. differs in its shorter staminate pedicels. TYPE: MEXICO. JALISCO: Mpio. Gua dalajara, Barranca de Huentitan el Alto, 3c. PHYLLANTHUS GRA VEOLENS ssp. ben 1350 m, 4 July 1990, B. Chazaro 6260 (Ho thamianus (Miill. Arg.) G. L. Webster, stat. LOTYPE: MEXU; ISOTYPES: DAV!, TEX!, & comb. nov. WIS!). Phyllanthus benthamianus Miill. Arg., Lin Frutex glaber, a subspeciei benthamiano differt pedicelis

3d. PHYLLANTHUS GRAVEOLENS ssp. subspecies. In Cajamarca, Peru, both gla GRA VEOLENS. brous (Sanchez 197) and hirtellous (Brand byge 145) forms occur, and the pointed leaf Phyllanthus millei Standl., Field Mus. Nat. blades are similar to those of Peruvian pop Hist., Bot. 22: 87. 1940. TYPE: ECUA ulations of P. brasiliensis. DOR. MANABi: Canton Sucre, Bahia de Caraquez, Jan. 1938, L. Mille 1061 (HO 4. Phyllanthus liesneri G. L. Webster, sp. LOTYPE: F!; photograph, AAU). nov. (Fig. 2a, b). TYPE: VENEZUELA. FALCON: Dto. DISTRIBUTION AND HABITAT: South Acosta, Parque N acional de la Cueva El ern Ecuador to northern Peru, mostly in Toro, trail to La Piedra, 10°N, 69°07'W, semideciduous woodlands or scrub, 100- 600-900 m, 22 June 1979, R. Liesner, A. 2400 m. Gonzalez, & R. Wingfield 7802 (HOLOTYPE: PHENOLOGY: Collected in flower Jan MO!; ISOTYPE: DAV!). uary, February, May, August to October; in Frutex vel arbor 4-5 m alto, caulibus foliisve fruit March. glabris laevis, ramulis lateralibus plerumque 7 ad 10; ab aliis speciebus sectionis differt foliis majoribus REPRESENTATIVE SPECIMENS EXAM ovatis acutis subacuminatisve, bracteis floralibus sca INED: ECUADOR. Guayas: Canton Gua riosis insigniter persistentibus. yaquil, 24 km W of Guayaquil, Dodson & GLABROUS TREES OR SHRUBS to 5 m Gentry 9639 (AAU, MO); Isla Puna, Madsen high; branchlets bipinnatiform, mostly 20- 63552, 63993 (AAU), 63847 (AAU, MO). 30 cm long; main axis 1.5-3 mm in diam Loja: Canton Gonzanama, Gerinoma, Pas eter, with 5 to 10 nodes; lateral axes aca 1656 (LOJA); Macara, Tambo Negro, smooth, subterete-angled, 8-20 cm long, 1- Harling & Andersson 18382 (AAU), Kessler 2 mm in diameter, with mostly 8 to 10 2724 (DAV); Canton Paltas, Catacocha to leaves. LEAF BLADES mostly ovate (occa Macara, Klitgaard et al. 399 (QCNE). Man sionally suborbicular), acute or breviacu abi: Canton Jipijapa, Cerro Cabeza de Vaca, minate, 2.5-5.5 X 2-3.5 cm; veins and Cornejo & Bonifaz 833 (DAV); Canton veinlets slightly prominulous adaxially and Montecristi, Montecristi Hill, Fagerlind & abaxially; petioles 2-2.5 mm long, smooth Wibom 609 (S); Canton Sucre, Bahia de Ca or obscurely papillate; stipules on branch raquez, Cornejo & Bonifaz 1737 (DAV). lets 1-1.5 mm long, persistent. FLOWERS in PERU. Cajamarca: Prov. Contumaza, Chi axillary bisexual cymules on branchlet lat letes, Brandbyge & Astholm 145 (AAU), eral axes, each cymule with 1 central pistil Sanchez Vega 197 (F), Prov. Jaen, Pucara, late and 10 to 15 lateral staminate flowers; Sagastegui 880, 5851 (US). Lambayeque: bracts scarious-indurate, pale, persistent, Prov. Chiclayo, Cahuageno, Llatas 777 forming nodules 1-4 mm in diameter. STA (MO); Prov. Lambayeque, Purculla ad Ol MINATE pedicel ca. 4 mm long; sepals 6, mos, Woytkowski 6780 (CTES, MO). Piura: biseriate, 1-1.5 mm long; staminal column Prov. Huancabamba, Quebrada Beatita de ca. 0.5 mm high; anthers ca. 0.3 mm long. Humay, Lopez et al. 8998 (AAU, MO). PISTILLATE pedicel 13-17 mm long in fruit; Tumbes: Prov. Tumbes, E of Hacienda Chi sepals ca. 1-1.5 mm long; styles free, bifid, cama, Weberbauer 7644 (NY, US). 0.3 mm long. FRUIT capsular, ca. 4 mm in diameter; columella 2 mm long; seeds tri Subspecies graveolens diverges from the gonous, 2.2-2.4 mm long, puncticulate. other three subspecies in its hirtellous axes, but otherwise appears morphologically sim DISTRIBUTION AND HABITAT: North ilar; however, in Ecuador at least it appears ern Venezuela, semideciduous forests on to occur in drier habitats than the other limestone hills, 10-600 m. NUM BER 6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 27

FIG. 2. Habit of Phyllanthus liesneri (Liesner et al. 7780, DAV). 2a, Branch with branchlets. 2b. Part of branchlet with node of indurated floral bracts.

PHENOLOGY: Collected in flower April, 5. Phyllanthus meridensis G. L. Webster, June; in fruit July. sp. nov. (Fig. 3a, b ). TYPE: VENEZUELA. MERIDA: Dto. Ar zobispo Chacon, between La Cumbre, San ADDITIONAL SPECIMENS EXAMINED: Jose, and Mucutuy, 1820-2590 m, 3 May VENEZUELA. Falcon: Dto. Acosta, Parque 1944, f. A. Steyermark 56256 (HOLOTYPE: F Nacional Quebrada de la Cueva El Toro, 1245696!). Liesner et al. 7780 (DAV, MO); Dto. Silva, La Soledad, Steyermark & Manara 110825 (VEN); Lizarde, R. Smith V8338 (VEN); Li Frutex glaber monoicus, ab aliis speciebus sec tionis differt stylis subintegris erectis coherentibus, zarde to Mallorquines, 10-25 m, Steyermark sepalis grandioribus. 110744 (VEN); Sanare, 10 m, Webster 23576 (DAV). Yaracuy: Prov. Bolivar, zona El MONOECIOUS SHRUBS to 3 m high; Chara!, R. Smith V9899 (MO). branchlets bipinnatiform; lateral axes 7 to 10, subterete, hirsutulous, 8-10 cm long, This species is named for the collector 0.7-1 mm thick, bearing ca. 10 to 15 leaves. of the type, Ronald Liesner, who has made LEAF BLADES glabrous, slightly ovate to many interesting collections of Phyllanthus mostly obovate or orbicular, obtuse to in Venezuela. It appears to be closely related rounded at apex and base, 1.5-2.5 X 1.2- to P. graveolens, but the habit of Phyllanthus 2.2 cm; midrib and major lateral veins 5 to liesneri appears strikingly different due to its 7 on a side, straightish but irregular, prom larger pointed leaves and to the persistent in ulous ad axially and ab axially; tertiary indurate floral bracts. veinlets slender, not raised; petiole smooth, 28 LUNDELLIA DECEMBER, 2003

\I.

~ .... l'l..A.Nn °"VO

1.5- 2 mm long; stipules lanceolate-acumi the highest elevation of any species in sec nate, scarious, brownish, 2.5-3 mm long. tion Nothoclema, and is perhaps the only FLOWERS in mostly bisexual cymules on species to extend into the Matorral Andino lateral branchlet axes; staminate flowers 5 (Ceja). to 10 per cymule, subtended by inconspic uous bracts. STAMINATE pedicel 4-5 mm In its general facies, Phyllanthus meri long; sepals oblong, the larger ca. 1. 7 mm densis appears quite similar to P. graveolens long, the shorter 1.2-1.3 mm long; stamina! ssp. micrandrus. However, it is clearly dis column ca. 0.5 mm high; anthers deflexed, tinct in its larger flowers, and especially in obtuse, ca. 0.8 mm long; disk segments 3, its remarkable gynoecium, quite different obcuneate, c. 0.7 mm long. PISTILLATE from that of any other species in the sec pedicel 10-12 mm long at anthesis; sepals tion. broadly elliptic, the larger 3.5-4 X 2-2.5 6. PHYLLANTHUS CAYMANENSTS G. L. mm, the shorter ca. 2.5 X 1.5 mm; disk Webster & G. R. Proctor, Rhodora 86: 121, cupular, pitted, rim irregularly dentate, c. figs. l, 2. 1984; Proctor, Kew Bull. Add. Ser. 0.7 mm high, 1.7-2 mm broad; ovary 11: 524. 1984. TYPE: CAYMAN ISLANDS. smooth; styles erect, subentire ( emarginate Cayman Brae, Foster Land District, 0. 7 mi. to slightly bifid distally), cohering in a col NW of Portland Bay, ca. 30 m, 7 Aug. 1975, umn 1.3-1.5 mm high. FRUITS and seeds G. R. Proctor 35151 (HOLOTYPE: JAM; ISO not seen. TYPES: DAV!, US!).

DISTRIBUTION AND HABITAT: Known DISTRIBUTION AND HABITAT: Endem only from the type collection in the Vene ic to the Cayman Islands, in woodlands on zuelan Sierra Nevada. It appears to occur at rocky substrate, near sea level. NUMBER 6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 29

PHENOLOGY: Collected in flower and Teopisca, Sousa & Magallanes 6697 (UC); fruit August. Mpio. Zinacantan, Laughlin 1216 (US).

ADDITIONAL SPECIMENS EXAMINED: 8. PHYLLANTHUS ACUMINATUS Vahl, CAYMAN ISLANDS. Little Cayman: N of Symb. Bot. 95. 1791; Mull. Arg., DC. Prodr. Wend of airstrip, Proctor 35145 (JAM, US). 15(2): 381. 1866; FL Brasil. 11(2): 42, t. 7 fig. 2. 1873; Webster, J. Arnold Arb. 38: 364, This is the only endemic West Indian t. 23 figs. C-F. 1957; Ann. Missouri Bot. species in section Nothoclema, and its rela Gard. 54: 228. 1968; FL Nicaragua 1: 896. tionships are unclear. Although similar to 2001; Burger & Huft, FL Costaricensis 113 Phyllanthus mocinianus in its habit, promi (Fieldiana Bot. II 36): 142. 1995. TYPE: nulous leaf venation, and pollen type, it is TRINIDAD. J. Ryan (HOLOTYPE: C!). closer to P. mcvaughii in its pointed anthers and it approaches P. acuminatus in anther Phyllanthus lycioides Kunth in H.B.K., Nov. form and smooth seeds. Gen. Sp. 2: 112. 1817. TYPE: COLOM BIA. [Bolivar] Mompox [Momp6s], F. 7. PHYLLANTHUS MCVAUGHII G. L. Web W H. A. Humboldt & A. J. A. Bonpland ster, Brittonia 18: 339, figs. 1, 7. 1966. TYPE: (HOLOTYPE: P!). MEXICO. CHIAPAS: Mpio. Ixtapa, 30 mi by Phyllanthus mucronatus Kunth in H.B.K., road E ofTuxtla Gutierrez, 1400 m, 24 June Nov. Gen. Sp. 2: 112. 1817. TYPE: VE 1962, G. L. Webster, WP. Adams, K. Miller, NEZUELA. "Provincia Caracasana," F. & L. Miller 11698 (HOLOTYPE: DAV!; ISO W H. A. Humboldt & A. J. A. Bonpland TYPES: GH!, MEXU!, MICH!, TEX!). (HOLOTYPE: Pl). Phyllanthus ruscoides Kunth in H.B.K., Nov. DISTRIBUTION AND HABITAT: Eastern Gen. Sp. 2: 113. 1817. TYPE: VENEZUE Mexico (Chiapas) to Guatemala and El Sal LA. [Monagas] "in sylvis prope Caripe et vador, mainly in oak/pine forest on lime in devexis montis Cocollar," F. W H. A. stone, 850-1800 m. Humboldt & A. J. A. Bonpland (HOLO PHENOLOGY: Collected in flower July TYPE: Pl). to September, in fruit September to Feb ruary. DISTRIBUTION AND HABITAT: North ern Mexico and Cuba south to northern REPRESENTATIVE SPECIMENS EXAM Argentina, Paraguay, and Brazil; mostly in INED: EL SALVADOR. Morazan: Mt. Ca secondary moist woodlands, gallery forests, caguatique, Tucker 679 (DAV, F, UC, US). or montane rain forests, 0-2250 m. GUATEMALA. Chimaltenango: between PHENOLOGY: Collected (Northern Chimaltenango and San Martin Jilotepeque, Hemisphere) in flower mainly May to No Standley 64499 (A, F, US). Huehuetenango: vember; in fruit July to December; (South between Las Palmas and Chacula, Steyer ern Hemisphere) in flower and fruit mainly mark 51756 (F, GH). Jalapa: 2 mi E of San November to March. Pedro Pinula, Steyermark 32993 (F). Solola: Lago Atitlan, Panajachel, Molina et al. REPRESENTATIVE SPECIMENS EXAM 16264 (NY), Webster et al. 11817 (DAV, INED: NORTH AMERICA. BELIZE. Belize: MEXU, TEX), Williams et al. 25333 (GH, Gracie Rock, Liesner & Dwyer 1529 (GH, US). MEXICO. Chiapas: Mpio. Chiapa de MO). Cayo: Vaca, Gentle 2520 (MO). Corzo, 5.6 mi E of Chiapa, Breedlove 10680 Orange Walk: Orange Walk, Lundell 4901 (MICH, US); Mpio. Cintalapa de Figueroa, (UC, US). COSTA RICA. Alajuela: Oroti 5 km W Rizo de Oro, Breedlove 24645 na, Holway 327 (US). Cartago: Canton (MICH, MO); Mpio. Teopisca, 5 km SW of Turrialba, Juan Vifias, Pittier 3655 (US). 30 LUNDELLIA DECEMBER, 2003

Guanacaste: Canton Liberia, 11.5 mi SE of Atlantico Sur: NW of Cano Angostura, Liberia, Webster & Millers 12443 (DAV). Sandino 4628B (DAV, MO). Carazo: Di Heredia: Universidad Nacional campus, riamba, Reyes 77 (MO). Chinandega: Los Unander & Poveda 1520 (DAV). Limon: Balcones, Moreno 11515 (DAV, MO). Canton Guacimo, 7 km NW of Siquirres, Chontales: Hae. San Martin, Stevens 21823 Wilbur 20751 (GH). Puntarenas: Canton (DAV, MO). Jinotega: Cerro Penas Blancas, Osa, 5 km E of Puerto Cortez, Wilbur 20980 Grijalva & Moreno 1046 (DAV, MO). Ma (GH, MO, US). San Jose: Canton Turru saya: Volcan Masaya, Neill 2810 (MO). Ma bares, Reserva Biologica Carara, Zuniga & tagalpa: 5-10 km W ofMatagalpa, Williams Jimenez 152 (DAV). EL SALVADOR. Cus et al. 23791 (MO, US). Rivas: Isla Ometepe, catlan: Las Pavas, Hernandez 211 (VS). San Robleto 1014 (DAV). PANAMA. Bocas del Salvador: Tonacatepeque, Standley 19520 Toro: Dto. Bocas del Toro, Cayo Agua, von (US). GUATEMALA. Escuintla: Finca El Wedel 870 (GH, MO). Chiriqui: Dto. Baru, Balli, Tonduz & Rojas 74 (MO). Izabal: Los Progreso, Cooper & Slater 222 (US). Code: Amates, Blake 7733 (US). Peten: Tikal, Or 12 mi NE of Penonome, Lewis et al. 1522 tiz 478 (MO). Retalhuleu: San Felipe, Da (DAV, MO, UC, US). Colon: Portobelo, nell Smith 2620 (US). Zacapa: Gualan, Correa & Dressler 1751 (DUKE). Panama: Deam 6301 (GH). HONDURAS. Atlantida: Dto. Capira, Cerro Campana, Duke 5978 Cerro Puerto Arturo, Saunders 1056 (MO). (US), Webster & Breckon 16522 (DAV). Comayagua: Talauba, Molina 13017 (US, Veraguas: Dto. Santiago, between Atalaya WIS). Copan: Hacienda Grande to Ostu and Santiago, Dwyer & Kirkbride 7421 man, Molina 24760 (US, WIS). Lempira: (UC). Lepaera, Hazlett 774 (GH, MO). Santa Bar WEST INDIES. CUBA. Cienfuegos: bara: San Pedro Sula, Kamb 3165 (US). Pico Sombrero, Webster 4769 (GH). Ha Valle: Amapala, Rodriguez 3371 (GH). bana: Santiago de las Vegas (cult. ?), Baker Yoro: Quebrada Olotillo, Molina 6828 (GH, 104 (UC). Pinar del Rio: Vinales, Palmar US). MEXICO. Baja California Sur: La ito, Ekman 16624 (S). JAMAICA. St. Mary: Burrera, 27 km E of Todos Santos, Tenorio Wag Water Valley, Harris 11147 (F, JAM, & Romero 10506 (DAV, MEXU). Campe NY). TRINIDAD & TOBAGO. Tobago: St. che: Mpio. Champoton, Tuxpena, Lundell George, Easterfield, Broadway 4555 (US). 805 (MO, US, WIS). Chiapas: Mpio. Villa Trinidad: Chaguaramas naval base, Webster Corzo, Breedlove 52041 (CAS). Colima: & Miller 9957 (DAV). Mpio. Manzanillo, Playa Santiago, Mc SOUTH AMERICA. ARGENTINA. Ju Vaugh 15593 (MICH, US). Guerrero: Atoy juy: Dto. Ledesma, Parque Nacional Cali ac de Alvarez, Hinton 14606 (US). Jalisco: legua, Bartlett 20354 (MICH, US), Mulgara Mpio. La Huerta, Rancho Cuixmala, Lott et et al. 1376 (MO). Salta: Dto. San Ramon, al. 3565 (DAV). Michoacan: Aquila, Hinton Rio Blanco, Venturi 5561 (MO, UC, US). 25965, 16109 (MICH, US), Nayarit: Aca BOLIVIA. Beni: Prov. Vaca Diez, Cachuela poneta, Rose 1906 (NY, US). Oaxaca: Mpio. Esperanza, White 533 (A, US). Chuquisaca: San Juan Guichicovi, Mogone, Orcutt 5214 Prov. Tomina, between Monteagudo and (DAV). Quintana Roo: Coba, Lundell 7614 Sucre, Beck 6346, Moretti 1502 (DAV). La (MICH, US). San Luis Potosi: Tamasopo Paz: Prov. Nor Yungas, Coripata, Bang Canyon, Pringle 3708 (UC, US). Tabasco: 2086 (MO). Santa Cruz: Prov. Florida, 6 Tenosique, Cowan & Magana 3226 (DAV). km SW of Angostura, Nee 48145 (MO). Tamaulipas: Mpio. Gomez Farias, 3 km Tarija: Prov. Arce, N of Sidras, Solomon NW of Gomez Farias, Diggs & Nee 2430 10141 (MO). BRAZIL. Alagoas: Serra Pao (DAV, MICH). Yucatan: Mayapan, Cabrera de As;ucar, de Lyra et al. 287 (MBM). Bahia: & Cabrera 9121 (MO). NICARAGUA. At Mun. Feira de Santana, Ipiras;u, Pedra de lantico Norte: Siuna, Narvaez 3180 (MO). Cavalho 741 (BAH). Ceara: Mun. Pacoti, NUMBER6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 31

Serrinha, Fernandes & Matos EAC 12061 Guayaquil, Reserva Faunfstica Churute, (DAV). Distrito Federal: Corrego Landim, Cornejo 1132 (MO, QCNE). Loja: Bosque Irwin & Wasshausen 19444 (M, MO). Petrificado Puyango, Cornejo & Bonifaz Goias: 35 km NE of Formosa, Irwin et al. 5034 (LOJA). Manabi: Canton Pajjon, Gu 15697 (UB, US). Maranhiio: Ilha Sao Luis, ale, Haught 3063 (US). Pastaza: Mera, As Fr6es 11816 (MICH). Minas Gerais: Mun. plund 18602 (B, US). Tungurahua: Banos, Curvelo, Gruta do Maquine, Beringer 7397 Fagerlind & Wibom s.n. (MICH, MO). Za (U). Para: Mun. Alenquer, Ducke 4946 mora-Chinchipe: Canton Chinchipe, Zum (RB). Paraiba: Areia, Coelho de Moraes 733, ba, Vivar & Merino 3018 (LOJA). GUYA 923 (US). Pernambuco: Mun. Vitoria de NA. Rupununi: between Kato and Para Santo Antao, Tapera, Pickel 1274 (US, makatoi, Gillespie & Persaud 879 (US). WIS). Rio de Janeiro: Rio de Janeiro, Sao PARAGUAY. Amambay: Sierra Amambay, Cristovao, Glaziou 7820 (US). Rondonia: Hassler 11435 (MICH, MO, UC). Gran 100 km SW of Ariquemes, Zarucchi et al. Chaco: Loma Clavel, Hassler 6699 (MICH, 2711 (DAV). Roraima: Porto Alegre, Rio MO). PERU. Ayacucho: Prov. La Mar, Amajary, Fr6es 23085 (RB). Sao Paulo: Ayna, Killip & Smith 22822 (US). Cajamar Mun. Campinas, Reserva Santa Genebra, ca: Prov. San Ignacio, Mandriga, Campos & Leitao Filho & Morellato 22894 (UEC). CO Diaz 2423 (DAV, MO). Cuzco: Prov. Calca, LOMBIA. Antioquia: Mun. Anori, Tona to La Quebrada, van der Hoogte & Roersch Liberia, Callejas et al. 4561 (MO, US). Bo 1819 (MO). Junin: Prov. Chanchamayo, La livar: Cartagena, Heriberto 375 (US). Boy Merced, Killip & Smith 23417 (US). San aca: Mt. Chapon, Lawrence 189 (MICH, Martin: Tarapoto to Yurimaguas, Rimachi US). Caldas: Mun. Manizales, Zuluaga 6 4091 (F), 5177 (MO). SURINAME. Suri (HUA). Cauca: La Manuelita, near Palmira, name River, Kappler 1688 (MO). VENE Pittier 812 (MO, US). Choc6: Mun. Istmi ZUELA. Aragua: Dto. San Sebastian, Banos na, below Andagoya, Forero et al. 5077 de San Juan de Los Morros, Pittier 12296 (MO). Cundinamarca: Mun. San Francis (MO, US). Barinas: Dto. Bolivar, Barinitas, co, Guaduas, Garca-Barriga 10961, 11703 Breteler 4238 (MO, US). Carabobo: Dto. (US). Guajira: Mun. Barrancas, Rio Ran Bejuma, Canoabo, Meier & Flauger 8790, cherfa, Roldan et al. 908 (MO). Huila: Mun. 8805 (DAV). Cojedes: Dto. Girardot, El Algeciras, Little 7511 (US). Magdalena: Baul, Trujillo et al. 16589 (MO). Distrito Santa Marta, 8 km E of Banda, H. H. Smith . Federal: Dto. Vargas, La Guaira, Curran & 375 (UC). Norte de Santander: Sarare, La Hamman 978 (US). Guarico: Dto. Roscio, Cabuya, Cuatrecasas et al. 12071 (US). Pu San Juan de los Morros, Meier & Elsner tomayo: Puerto Limon, Plowman 2152 6145 (DAV). Lara: Dto. Palavecino, Sanare, (US). Risaraldo: Mun. Mistrato, between Div. Ejuc. Progr. 5 (MY). Miranda: Dto. Ac Geguadas and San Antonio de Chami, Fer evedo, E of Caucagua, Berry 920 (MO). nandez et al. 10018 (MO, US). Santander: Monagas: Dto. Maturin, W of Jusepin, Mun. Betulia, between Lebrija and San Vi Seigler 1110 (DAV). Portugesa: Dto. Arau cente, Langenheim 3170 (UC, US). Tolima: re, Camburito, Ortega & Aymard 1722 Mun. Espinal, Chicoral, Haught 6343 (UC, (MO). Tachira: San Cristobal, Archer 3210 US). Valle del Cauca: Mun. Buenaventura, (US). Trujillo: Dto. Bocono, SW Agua Frfa, El Papayo, Cuatrecasas 15999 (F). ECUA 550 m, Dorr et al. 8503 (DAV, US). Zulia: DOR. Carchi: Canton Mira, between Ibarra Dto. Mara, Destacamento Guasare, Bunting and San Lorenzo, Dodson & Thien 1566 & Leon 12574 (MO). (MO). El Oro: Palo Solo, Vivar & Merino 3018 (LOJA). Esmeraldas: Canton Eloy Al Phyllanthus acuminatus is the common faro, Comuna de Calle Mansa, Yanez et al. est and most widespread species of sect. 1487 (DAV, MO, QCNE). Guayas: Canton Nothoclema, but it shows remarkably little 32 LUNDELLIA DECEMBER, 2003

geographic variation in morphological without staminate cymules; seeds rarely produced; characters. Although it has been confused widespread in Amazon lowlands . . ssp. brasiliensis 1. Branchlets and flowers (including ovary) glabrous; by collectors with P. brasiliensis, pollen main branchlet axis usually lacking staminate cy characters indicate a closer relationship mules; seeds commonly produced; uplands of Bo- with the Central American P. mcvaughii livia ...... ssp. glaber (Webster & Carpenter, 2002).

9. PHYLLANTHUS BRASILIENSIS (Aubl.) 9a. PHYLLANTHUS BRASILIENSIS ssp. BRAS Poir. in Lam., Encycl. 5: 296. 1804; Mull. ILIENSIS. Arg., DC. Prodr. 15(3): 383; Fl. Brasil. 11(2): 44. 1873. Conami brasiliensis Aubl., Phyllanthus subglomeratus Poir., Encycl. Hist. Pl. Guiane 2: 926, t. 354. 1775. Phyl Method. 5: 304. 1805. TYPE: MARTI lanthus conami Sw., Prodr. 28. 1788 (as to NIQUE. Herb. Lamarck (HOLOTYPE: Pl). type only). Conami conami (Sw.) Britton, Phyllanthus piscatorum Kunth in H.B.K., Bot. Porto Rico 5: 475. 1924. TYPE: GUI Nov. Gen. Sp. 2: 90, 113. 1817. TYPE: ANE FRANyAISE. Aublet (HOLOTYPE: VENEZUELA. [Amazonas] "cataractum BM!). Aturensium," F. W H. A. Humboldt & A. ]. A. Bonpland (HOLOTYPE: Herb. The delimitation of Phyllanthus brasi Humboldt, Pl). liensis in botanical literature has been dif Phyllanthus pseudo-conami Mull. Arg., FL ficult for both taxonomic and nomencla tural reasons. Swartz (1788, 1800) intro Brasil. 11(2): 43. 1873; var. glaber Mull. duced confusion that lasted for two centu Arg., ibid. TYPE: VENEZUELA. AMA ries by describing P. conami, derived from ZONAS: San Carlos, R. Spruce 3016 (HO Aublet's name but based on a description LOTYPE: Herb. Martius, M). of P. acuminatus. As a consequence, the Phyllanthus pseudo-conami var. pubescens name P. brasiliensis has been misapplied to Mull. Arg., Fl. Brasil. 11(2): 43. 1873. specimens of P. acuminatus by authors into TYPE: PERU. [LORETO]: Yurimaguas, the 20th century (e.g., Standley & Steyer Poeppig 2758 (LECTOTYPE, selected here: mark, 1949). Muller (1866, 1873) avoided W!). that confusion but recognized three species, P. brasiliensis, P. piscatorum, and P. pseu DISTRIBUTION AND HABITAT: Ama doconami, for populations in the Brazilian zonian lowland forests from Colombia and Amazon, where only a single species is ac Venezuela south to Peru, Brazil, and north cepted in the present treatment. ern Bolivia, 10-500(-1000) m; introduced The delimitation of Phyllanthus brasi to the West Indies from Trinidad to Guad liensis is further complicated as far as pop aloupe (Webster, 1957, 1991). ulations from Bolivia are concerned. The PHENOLOGY: Collected in flower Bolivian highland populations have been throughout the year; in fruit (rare) April, assigned to P. graveolens by Pax and Hoff June, July, October. mann, but except for lack of indumentum appear to have many of the characteristics REPRESENTATIVE SPECIMENS EXAM (especially in pollen morphology) of P. INED: WEST INDIES. LESSER ANTILLES. brasiliensis. Consequently two subspecies of Dominica: Grand Bay, Eggers 662 (GH, P. brasiliensis are recognized here. GOET, L, US, W), Imray s.n. (GOET). Gua deloupe: Matouba, Duss 212 (P), 2446 (GH, KEY TO THE SUBSPECIES OF MO, US); Grande Terre, L'Herminier s.n. PHYLLANTHUS BRASILIENSIS (A, P); St. Claude, Quentin 527 (F), Stehle 1. Branchlets and flowers mostly hirtellous; ovary 63 (A); Trois-Rivieres, Quentin 671 (P). glabrous to hirtellous; main branchlet axis with or Martinique: Ajoupa-Bouillon, Duss 2047 NUMBER6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 33

(NH); St. Pierre, Belanger 294 (G, P). To (U). GUYANA. Essequibo: Kabakaburi, bago: Valencia Road, Baker (TRIN 14864). Hoffman & Roberts 2466 (DAV, TEX). Trinidad: without locality, Ryan (C). Northwest: Koriabo River, Archer 2355 SOUTH AMERICA. BOLIVIA. Beni: (US); Moruka River, Mora Landing, de la Prov. Gral. Jose Ballivian, NE San Borja, Cruz 962 (US). PERU: Huanuco: Prov. Oviedo 41 (DAV). BRAZIL. Acre: Mun. Huanuco, Cerros del Sira, Wolfe 17193 (F); Cruzeiro do Sul, Penereiro 11063 (DEC); Tingo Maria, Belshaw 3068 (F, LL, MICH, Mun. Manoel Urbano, Silveira 1520 (DAV, MO, UC, US); Prov. Leoncio Prado, Dto. NY); Mun. Santa Rosa, Daly et al. 9913 Rupa Rupa, King & Ramirez 350 (F, MO); (MO); Mun. Tarauaca, Ehringhaus et al. 378 Dto. Puerto Inco, Seiderschwarz 3774 (F). (DAV). Amapa: Amapa, Ducke (MG 4840). Loreto: Prov. Alto Amazonas, Balsapuerto, Amazonas: Mun. Barcelos, Prance & Ramos Klug 2913 (MO, US); Yurimaguas, Poeppig 23616 (MO, U); Mun. Boca do Acre, Born 2758 (W); Pongo de Manseriche, Mexia Lugar, Goeldi (MG 3978); Mun. Manaus, 6253 (TEX, UC, US); Prov. Maynas, Intuto, Froes 20521 (US); Mun. Sta. Isabel do Rio McDaniel 10841 (DAV, MO); Prov. Coro Negro, Prance et al. 15556 (F, MG, U, US); nel Portillo, Bosque Nacional von Hum Mun. Umarituba, Rio Xi6, Froes 20521 (F). boldt, Diaz & Aronson 736 (DAV, F, MO); Para: Mun. Belem, Jardim Botanico, Ducke Pongo de Canarachi, Ule 6408 (MG, RB). 1261 (MG, RB, US); Mun. Breves, Black 48- Madre de Dios: Prov. Marni, Cocha Cashu 3019 (NY). Roraima: Mun. Boa Vista, Station, Foster 9847 (MO); Prov. Tambo Auaris, Prance et al. 21351 (MG, U). CO pata, Barosola, Vargas 18704 (MO, US). LOMBIA. Amazonas: La Chorrera, Schultes San Martin: Prov. Lamas, Distr. Alonso de 3898 (AAU, GH). Boyaca: Mun. Chiscas, Alvarado, Schunke 9725 (AAU, DAV, F, Cuatrecasas 13030 (US). Caqueta: La Maria, MO); Prov. Mariscal Caceres, Juan Jui, Klug Hermann 11233 (COL, US). Putumayo: 3806 (F, MO, US); Prov. Moyobamba, Rio Mocoa, Archer 3416 (COL); between Puerto ja, Woytkowski 6063 (MO); Rioja to Pedro Ruiz, van der Werff et al. 15480 (DAV, Asis and Puerto Leguizamo, Garcia-Barriga MO); Prov. San Martin, Pucacaca, Zegarra et al. 18702 (US). Vaupes: Jinogoje, Schultes 9 (NY); Tarapoto, Spruce 4939 (NY). SU & Cabrera 16832 (GH); confluence of Rio RINAME. Nickerie: Sipaliwini, Oldenburger Vaupes and Rio Kubiyli, Soejarto & Lock et al. 1275 (BBS, MG, NY, U). Saramacca: wood 2432 (F, HUA). ECUADOR. Napo: between Voltzberg and Ralegh Vallen, Web Cant6n Archidona, Mexia 7246 (U, US); ster 24145 (DAV, U). VENEZUELA. Ama Cant6n La Hoya de los Sachas, Baker et al. zonas: Dto. Atabapo, Culebra, Perez & Sosa 5981 (MO); Cant6n Tena, Ahuano, Rios & JPB-602 (MO), Steyermark & Delascio Vivanco 375 (QCA); Jatun Sacha, Ceron 129155 (MO); base of Duida near Culebra, 1140 (QCA); E of Puerto Napo, Webster & Liesner 17878 (DAV); S of San Juan de Richerson 28494 (DAV, MO, QCA, QCNE). Manapiare, Berry 1605 (MO); Dto. Casa Orellana: Cant6n Aguarico, Nuevo Roaca quiare, Rio Guainia, Williams 14800 (F); fuerte, Alarcon 101 (QCA); Cant6n Orel Dto. Rio Negro, San Carlos de Rio Negro, lana, San Pablo de los Secoyas, Brandbyge Spruce 3016 (W); Tamatama, Williams & Asanza 32853, 32941 (AAU). Pastaza: 15865 (F, RB, US). Anzoategui: Dto. Ced Cant6n Pastaza, near Puyo, Skutch 4441 eno, Sta. Maria de Erebato, Steyermark et (MO, US), Webster 23170 (DAV, MO, Q, al. 109862 (F); Dto. Freites, Bajos del Rio TEX, WIS). Sucumbios: Cant6n Lago Tigre, Cristovero, Pittier 14822 (UC, US). Agrio, Shushufindi, Vickers 45 (F); Cant6n Bolivar: Dto. Roscio, Mata Negra, S de El Putumayo, Puerto El Carmen, Andrade Tigre, Williams 13392 (F, US). 33115 (AAU). GUIANE FRANyAISE: De grad Kaw, riviere Cainama, Oldeman B-775 Subspecies brasiliensis is the most wide (NY, U); Riviere Orapu, Moretti CM1046 spread taxon in sect. Nothoclema after P. 34 LUNDELLIA DECEMBER, 2003

acuminatus, and is the main plant used in Surutu, Steinbach 7315 bis (MO); 4 km NW South America as a fish poison. Examina Buena Vista, Nee 39061 (MO). Tarija: Prov. tion of specimens across the Amazon basin Arce, 2 km NW Bermejo, Nee 46166 (TEX). has shown that except for southern Ecuador and northern Peru, nearly 100% of the The resemblance of Bolivian specimens plants in the rest of the distribution pro of ssp. glaber to the glabrous populations of duce no fruits and seeds. Apparently most Phyllanthus graveolens i~ so apparent that populations of ssp. brasiliensis represent Pax and Hoffmann's attribution was not sterile clones that have been cultivated by unreasonaable. However, the discovery that indigenous peoples in a semi-domesticated the pantoporate pollen grains with pilate state. exine are nearly indistinguishable from those of ssp. brasiliensis (Webster & Car 9b. PHYLLANTHUS BRASILIENSIS ssp. glaber penter, ined.) justifies the taxonomic dis (Pax & K. Hoffm.) G. L. Webster, stat. & position made here. comb. nov. (Fig. 1). The choice of glaber as subspecific ep ithet is undesirable because of the prior var. Phyllanthus graveolens var. glaber Pax & K. glaber used by Miiller (1873) for his Phyl Hoffm., Meded. Rijksherb. Leiden 40: 18. lanthus pseudoconami var. glaber; even 1921. TYPE: BOLIVIA. SANTA CRUZ: be though Miiller's name is illegitimate, it tween Santa Cruz and Rio Pirai, T.K.]. could lead to confusion. However, use of Herzog 1338 (LECTOTYPE: W!; chosen Rusby's P. ichthyomethius as the basis of a here to replace destroyed holotype at B). name is also undesirable. Specimens from Phyllanthus ichthyomethius Rushy, Mem. N. El Beni and La Paz have a distinctly differ Y. Bot. Gard. 7: 282. 1927. TYPE: BO ent leaf shape from those in Santa Cruz and LIVIA. EL BEN!: Prov. Gral. Jose Balli Tarija departments, and furthermore, it ap vian, Rurrenabaque, White 886 ex p. pears that P. ichthyomethius is apparently (HOLOTYPE: NY!; ISOTYPES: Kl, MICH!, founded on a mixed collection. The de US!). scription and the holotype at NY cited Rur renabaque, in El Beni, as the location; but DISTRIBUTION AND HABITAT: Bolivia, the label on the sheet at US indicates Hu upland subtropical forests from southern achi in La Paz as the locality. It appears pos Dpto. El Beni to Dptos. Santa Cruz and sible that White 886 may include plants col Tarija, 300-700 m. lected at two locations over 100 kilometers PHENOLOGY: Collected in flower and apart! Although it would be acceptable to fruit October, November. create an entirely new epithet at the sub specific rank, it appears that the least un REPRESENTATIVE SPECIMENS EXAM satisfactory alternative is to use Pax and INED: BOLIVIA. La Paz: Prov. Sud Yungas, Hoffmann's epithet as the basionym of the Alto Beni, San Miguel de Huachi, White 886 subspecies. ex p. (US); Sapecho, Seidel 2788 (DAV); The differences between the northern Santa Ana de los Mosetenes, Hinojosa & and southern populations of ssp. glaber sug Bozo 1056, Hinojosa & Huasang 1146 gest that the taxon might eventually be sub (DAV). Santa Cruz: Prov. Andres Ibanez, 1 divided even further. However, until further km SW Angostura, Nee 33502 (MO); Rio sampling can be done, it appears best to Salado, Nee 38047 (U); Rio Pirai, Jardin Bo refer all of the non-Amazonian Bolivian tanico, Nee 30480, Zuloaga et al. 1425 specimens to ssp. glaber. (MO); 6 km NW Santa Cruz, Solomon 13841 (MO); Prov. Florida, 4.5 km ENE La 10. PHYLLANTHUS PAVONIANUS Baill., Ad Guardia, Nee 45943 (TEX); Pro. Ichilo, Rio ansonia I. 1: 30. 1860; Miill. Arg., DC. NUMBER6 WEBSTER: SYNOPSIS OF PHYLLANTHUS SECTION NOTHOCLEMA 35

Prodr. 15(2): 380. 1866; Macbride, Field bes: Prov. Contralmirante Villar, moun Mus. Nat. Hist., Bot. 13 (3A, 1): 44. 1951. tains E of Hacienda Chicama, 700-800 m, TYPE: PERU. ]. A. Pavon 314 (LECTOTYPE, Weberbauer 7664 (NY, US). designated here: Herb. Lessert, G!). Because of its elongated leaf blades and Phyllanthus oxycladus Mtill. Arg., Linnaea ribbed-compressed branchlet axes, Phyllan 32: 26. 1863. TYPE: Herb. Pavon (LEC thus pavonianus is one of the most mor TOTYPE, designated here: G-Boiss.). The phologically distinctive species in sect. data provided by the Flora Huayaquilen Nothoclema. However, the pantoporate, sis (Estrella, 1989) suggest that the type pollen grains with pilate exine are very sim specimens for both species names were ilar to those of P. brasiliensis. collected at Colonche, Prov. Guayas, Ec uador, by Juan Tafalla, in 1801 (not far ACKNOWLEDGEMENTS from the type locality of P. haughtii). However, the illustration in Flora Hu I wish to thank the curators of the in ayaquilensis (lamina CLXXI) appears stitutions that provided loans and access to more similar to P. anisolobus. Apparently collections: A, AAU, B, BBS, BAH, BR, the type locality of P. pavonianus, as sug CAS, COL, DAV, DS, ENCB, F, GH, gested by Estrella (and verified by L. GUAY, HUA, IBUG, IEB, JAM, K, LL, Holm-Nielsen) is correct, but was mis LOJA, MBM, MEXU, MG, MICH, MO, takenly associated with a different illus MY, NY, P, Q, QCNE, RB, S, TEX, U, UB, tration. UEC, US, VEN, W., WIS, ZEA. Ellen Dean, Phyllanthus haughtii Croizat, Caldasia 3: 22. Director of the University of California, Da 1944. TYPE: ECUADOR. GUAYAS: Can vis, Herbarium, provided space and logistic ton Pedro Carbo, 15 km W of Pedro assistance. Kevin Carpenter made signifi Carbo, 17 Jan. 1940, 0. Haught (HOLO cant SEM photographs of pollen. Comple TYPE: GH!; ISOTYPES: K!, US!; photo tion of the work was expedited by a grant graph, AAU). from the UC Davis Washington Center, in cooperation with the National Herbarium, DISTRIBUTION AND HABITAT: South Smithsonian Institution, Washington. ern Ecuador and northwestern Peru, in de ciduous shrubbery along banks of intermit LITERATURE CITED tent streams, 200-1500 m. PHENOLOGY: Collected in flower Jan Acevedo-Rodriguez, P. 1990. The occurrence ofpis uary, February, April; in fruit January, Au cicides and stupefactants in the plant kingdom. gust. Adv. Econ. Bot. 8: 1-23. Aublet, F. 1775. Histoire des plantes de la Guiane franfoise. Didot, London. ADDITIONAL SPECIMENS EXAMINED: Burger, W., and M. Huft. 1995. Euphorbiaceae, in ECUADOR. El Oro: Canton Arenillas, Flora Costaricensis. Fieldiana IL 36: 1-169. Quebrada Los Zabalos, 330 m, Cornejo & Estrella, E. 1989. Flora Huayaquyilensis: sive descrip Bonifaz 4077 (DAV); Canton Zaruma, be tiones et icones plantarum huayaquilensium secun dum systema Linnaeanum digestae. 2 vols. Insti tween Guishahuina and Curtincapa, Rio tuto ad Conservandam Naturam y Horto Regio Luis, 850 m, Steyermark 5381 (AAU, NY). Matritense, Madrid. Loja: Canton Catamayo, Quebrada Grande, Meewis, B., and W. Punt. 1983. Pollen morphology Gerinoma, 1500 m, Pasaca 1656 (LOJA); and taxonomy of the subgenus Kirganelia (Jus Canton Puyango, Mercadillo, between Cel sieu) Webster (genus Phyllanthus, Euphorbiaceae) from Africa. Rev. Palaeobot. Palynol. 39: 131-160. ica and Alamor, f. F. Smith 1971 (MO). Mfiller, J. 1866. Phyllanthus, in De Candolle, Prod PERU. Piura: Prov. Huancabamba, 900- romus Systematis Universalis Regni Vegetabilis. 1000 m, Weberbauer 6019 (GH, US). Tum- 15(2): 274--436. 36 LUNDELLIA DECEMBER, 2003

---. 1873. Phyllanthus, in Martius, C. P. F. (ed.), phorbiaceae) from Central America. Brittonia 18: Flora Brasiliensis 11(2): 23-76. 336-342. Pax, F. and K. Hoffmann. 1921. Phyllanthus in Die ---. 1970. A revision of Phyllanthus (Euphorbi von Dr. Th. Herzog auf seiner zweiten Reise aceae) in the continental United States. Brittonia