Phyllanthus (Euphorbiaceae)

Transcription

1 Three New Sections and a New Subgenus of Grady L. Webster Davis Herbarium, Section of Plant Biology, University of California, Davis, California 95616, U.S.A. ABSTRACT. Three new sections of Phyllanthus are linis grana grosse reticulata; ab aliis sectionibus subgedescribed: section nero Isoclado differt Antipodanthus (subg. Isocladus; foliispiraliter insertis. type P dictyospermus Muller Argoviensis), 15 spe- Monoecious or dioecious shrubs or subshrubs, cies of South America and Australia; section Pityrbranching unspecialized, branches smooth or paocladus (subg. Emblica; type P symphoricarpoides pillate; phyllotaxy spiral; leaves petiolate, stipules Kunth), 5 species of tropical America; and section persistent; flowers pedicellate, in axillary glomer- Hylaeanthus (subg. Conami; type P attenuatus Miules; sepals 5 or 6, imbricate, entire; staminate disk quel), 6 species of tropical America. Two new spe- of 5 or 6 segments; stamens 3, filaments free or cies are described: P cuatrecasanus (sect. Pityro- connate; anthers vertically or horizontally dehiscladus) and P callejasii (sect. Hylaeanthus). A new cent; pollen grains subglobose, 3- or 4-colporate, subgenus with unique pollen structure, Cyclanthera mostly coarsely reticulate; pistillate disk patelli- (type P lindenianus Baillon), is proposed; it inform; ovary glabrous, 3-locular; styles + free, bifid; cludes sections Callitrichoides and Cyclanthera. fruits capsular, seeds smooth or verruculose. Key words: Euphorbiaceae, Neotropics, Phyllanthus. Phyllanthus, with over 800 species, is the third largest genus of Euphorbiaceae (Govaerts et al., 2000). The remarkable diversity in growth form and floral morphology (Bancilhon, 1971; Nozeran et al., 1984), and especially in pollen (Punt, 1967, 1987), is reflected in its complex classification, with 10 subgenera and over 30 sections (Webster, , 1967). Although about 70% of the species of Phyllanthus are confined to the Old World, the 200- plus neotropical species have not received extensive study except for the treatments of Maller (1873) for Brazil, Jablonski (1967) for the Amazon basin, and Webster ( , 1999) for the West Indies and Venezuelan Guayana. During the course of a revision of the neotropical species, a considerable number of undescribed taxa have been detected. Because the names of three newly recognized sections and a new subgenus are being cited in a study of the pollen morphology of neotropical taxa (Webster& Carpenter, 2002), they are being published here in advance of the overall revision of the neotropical taxa. Bentham, P australis J. D. Hooker, P Phyllanthus subg. Isocladus G. L. Webster calycinus Labillardiere, P Phyllanthus sect. Antipodanthus G. L. carpentariae Maller Argoviensis, Webster, P sect. nov. TYPE: fuernrohrii F. Mueller, P hirtellus Muller Argo- Phyllanthus dictyospermus Muller viensis, P maitlandianus Diels, P oblanceolatus J. Argoviensis. A sectione Paraphyllantho differt disco 9 integro, pol- NovoN 12: About 15 species can be referred to Phyllanthus sect. Antipodanthus, which belongs to subgenus Isocladus by virtue of its unspecialized (non-phyllanthoid) branching. It appears to be the Southern Hemisphere vicariant of section Paraphyllanthus Maller Argoviensis. The two sections seem morphologically similar in many respects, but in section Paraphyllanthus the anthers are always vertically dehiscent and the pistillate disk dissected. Only 5 species have been described from South America, mostly Brazil; the others are native to Australia (species 19, 21, 25-27, 29-32, 35-37, and 39 in Bentham, 1873). A number of the Australian species have recently been treated by Hunter and Bruhl (1996, 2000). The Australian and South American species differ in some characters, the American species having mostly connate filaments and horizontally dehiscing stamens; it is conceivable that cladistic analysis may show that some of the Australian species should be referred to section Lysiandra (F. Mueller) G. L. Webster. The Australian species are therefore listed here but not treated in detail. Australian species included: Phyllanthus aridus T. Hunter & J. J. Bruhl, P scaber Klotzsch, P similis Muller Argoviensis, and P striaticaulis J. T.

2 Volume 12, Number Webster 291 Hunter & J. J. Bruhl. It is possible that P erwinii J. T. Hunter & J. J. Bruhl and P lacunarius F. Mueller are also referable to Phyllanthusect. Antipodanthus. KEY TO THE AMERICAN SPECIES OF PHYLLANTHUS?Phyllanthus cordobensis (Kuntze) K. Schumann, Bot. Jah- SECT. resber. 26(1): Diasperus cordobensis Kun- ANTIPODA4NTHUS tze, Rev. Gen. P1. 3(2): TYPE: Argentina. la. Filaments connate; stems? terete; stipules 1-6 C6rdoba: San Roque, O. Kuntze (holotype, NY). mm long; seeds mm long; monoecious or dioecious. Since the Sellow syntype of Phyllanthus ramil- 2a. Stipules 3-6 mm long; stemsmooth; leaves losus was destroyed, the Herb. Franqueville specon main stems > 5 mm long; dioecious. imen logically should become the lectotype; how- 3a. Leaves on main stems > 5 mm broad; seeds reticulate or irregularly verrucuever, that specimen could not be located in the lose Prodromus Herbarium in Geneva. If the Herb. Frdictyospermus 3b. Leaves on main stems < 5 mm broad; anqueville specimen cannot be located, St. Hilaire seeds foveolate-punctate (P) could be designated as a neotype. It is P. pinifolius also 2b. Stipules mm long; stem smooth or possible that a duplicate of Sellow may yet be papillose; leaves on main stems mostly < discovered. 5 mm long; monoecious or dioecious. Woodlands and scrub, m, Bolivia, 4a. Dioecious; stems papillose; staminate northern Argentina, and Brazil (Santa Catarina and flowers distinctly pedicellate (pedicel > Rio Grande do Sul). The status of Phyllanthus cor- 1 mm long); pistillate sepals 5; leaf blade dobensis remains uncertain; Kuntze's description is marginstrongly revolute, tip blunt with abruptly reflexed acumen. inadequate, and the single specimen is somewhat P ramillosus different from other collections of P. ramillosus; 4b. Monoecious; stems smooth; staminate flowersubsessile (pedicel < possibly it might be validated at subspecific status 1 mm when better known. long); pistillate sepals 5 or 6; leaf blade margins not revolute, apically tapering to a straight acumen P dawsonii 4. Phyllanthus dawsonii Steyermark, Los Angelb. Filaments free; stems decurrent-angled from les Co. Mus. Contr. Sci. 21: TYPE: nodes; stipules 2 mm long or less; seeds Brazil. Goids: Chapada dos Veadeiros, E. Y mm long; dioecious P. rosmarinifolius Dawson (holotype, R). 1. Phyllanthus dictyospermus Miller Argoviensis, DC. Prodr. 15(2): TYPE: Bra- Goiias. zil. Minas Gerais: J. F Widgren 1003 (holotype, G). Phyllanthus regnellianus Milller Argoviensis, Fl. Brasil. 11(2): TYPE: Brazil. Minas Gerais: Caldas, A. E Regnell II 48 (lectotype, designated here, S). Montane scrublands, ca m, Brazil (Minas Gerais). Phyllanthus regnellianus, which was distinguished by Muller only on the basis of a questionable difference in seed sculpturing, shows no clear differences and is here reduced to synonymy. Linnaea 32: TYPE: Brazil, Santa Catarina: Herb. Franqueville (syntype, G); "Brasilia meridionali," F Sellow (syntype, B destroyed). Cerrado and campos rupestres, m, 6. Phyllanthus rosmarinifolius Miller Argoviensis, Fl. Brasil. 11(2): TYPE: Brazil. Rio de Janeiro: Serra dos Orgios, G. Gardner 5852 (holotype, G). Recorded only from rocky slopes, m, Rio de Janeiro, Brazil. Phyllanthus subg. Emblica (Gaertner) Kurz Phyllanthus sect. Pityrocladus G. L. Webster, sect. nov. TYPE: Phyllanthusymphoricarpoides Kunth. A sectione Emblica differt 2. Phyllanthus pinifolius Baillon, Adansonia sepalis plerumque 5, an- I. theris muticis, stylis liberis non dilatatis; a sectione Mi- 5: TYPE: Brazil. Parani: Curitiba, croglochidionis differt foliis non glanduliferis, antheris A. St. Hilaire [15590? ] (holotype, P; muticis, pollinis grana 4- vel 5-colporatis non diporatis. isotype, K). Monoecious (rarely dioecious) shrubs, often Scrublands, m, Minas Gerais, Espir- scandent; branches and branchlets + scabridulous ito Santo, and Parani. or papillate to hirtellous or scurfy; deciduous branchlets pinnatiform (not ramified), with ca Phyllanthus ramillosus Miuller Argoviensis, 50 leaves; leaf blades chartaceous, lacking a sub-

3 292 Novon apical laminar gland; stipules persistent; flowers pedicellate, in axillary unisexual or bisexual cymules; staminate sepals 5 or 6, subequal, free, imbricate; disk segments 5; stamens 2-5(-7), filaments connate (rarely free); anthers muticous, dehiscing horizontally or obliquely; pollen grains subprolate, 4- or 5-colporate, colpi monorate and unbordered, exine reticulate; pistillate sepals 5, imbricate; disk crateriform or dissected; ovary 3-locular; styles free, spreading, bifid to unlobed; fruits dehiscent or indehiscent; seeds smooth. Phyllanthus sect. Pityrocladus, which is a neotropical representative of Phyllanthus subg. Emblica (Gaertner) Kurz, includes 6 described species (and at least 1 or 2 undescribed) mainly of South America, but with one species in Costa Rica. The 4-5-colporate reticulate pollen grains (Webster & Carpenter, 2002) suggest placement in subgenus Emblica, even though the shape of the grains is scarcely prolate. The section, named for the characteristic scabridulous or scurfy indumentum of the axes of some of the species, is similar to section Microglochidion (Muller Argoviensis) Muller Argoviensis in some pollen characters but differs in its tendency to a scandent habit, more or less scurfy indumentum, eglandular leaves, and less elongated pollen grains with fewer colpi. KEY TO THE SPECIES OF PHYLLANTHUS SECT. PITYROCLADUS la. Leaves glabrous on both faces (at most scabridulous or papillate on petiole or midrib); veins distinctly prominulous on abaxial surface; branchlets cm long, mm thick; with leaves; staminate pedicels 3-7 mm long; stamens 3-6; fruiting pedicels 2-4 mm long. 2a. Monoecious; leaf blades elliptic, cm long, apically rounded and apiculate; stamens (3)4-6; fruits indehiscent; styles unlobed or apically bifid; branchlets not distinctly winged P symphoricarpoides 2b. Dioecious; leaf blades ovate-lanceolate, 3-6 cm long, acuminate; stamens 3; fruits dehiscent; styles bifid; branchlets narrowly winged P sponiifolius lb. Leaf blades glabrous or hirsutulous adaxially; veins not prominulous abaxially (or if so, blades distinctly hirsutulous abaxially); branchlets 5-25 cm long, mm thick, with 5-75 leaves; staminate pedicels mm long; stamens 2 or 3; fruiting pedicels 1-4 mm long. 3a. Leaf blades abaxially copiously hirsutulous, with prominent lateral veins; stamens 2; styles unlobed. 4a. Monoecious; branchlets mm thick, with 5-11 leaves; leaf blades cm long; petioles 1-2 mm long; stipules mm long P popayanensis 4b. Dioecious; branchlets mm thick, with leaves; leaf blades cm long; petioles ca. 0.5 mm long; stipules mm long.. 4. P ruscifolius 3b. Leaf blades abaxially glabrous or minutely scabridulous, veins not prominulous; stamens 2 or 3. 5a. Leaf blades elliptic, strongly inaequilateral at base, secondary veins straight, nearly perpendicular to midrib, abaxially separated from midrib; branchlets mm thick; stipules mm long; staminate disk segments discrete, flat P cuatrecasanus 5b. Leaf blades oblong, not inaequilateral at base, secondary veins distant, curving, not abaxially separated from midrib; branchlets mm thick; stipules mm long; staminate disk segments massive, + confluent P. valerii 1. Phyllanthus symphoricarpoides Kunth, in Humboldt, Bonpland & Kunth, Nov. Gen. Sp. 2: Glochidion symphoricarpoides (Kunth) Pax & K. Hoffmann, in Engler & Harms, Nat. Pflanzenfam. ed. 2, 19c: TYPE: Ecuador. Loxa [Loja]: Humboldt & Bonpland (holotype, P; isotype, B). Montane thickets, Peru. m, Colombia to 2. Phyllanthus sponiifolius Miller Argoviensis, Linnaea 32: TYPE: Ecuador. L. Fraser (holotype, G). Montane thickets, Ecuador. m, Colombia to 3. Phyllanthus popayanensis Pax, Bot. Jahrb. 26: TYPE: Colombia. Popay in: Pairamo de Guanacas, E C. Lehmann 4708 (holotype, B destroyed; lectotype, designated here, K). Montane thickets, m, Colombia (Cauca). The specimen at Kew is chosen as lectotype because no other specimens of Lehmann 4708 have been found. 4. Phyllanthus ruscifolius Miuller Argoviensis, DC. Prodr. 15(2): TYPE: Colombia. Prov. Cauca: J. J. Triana 3659 (holotype, P). Montane thickets, m, Colombia (Valle del Cauca). 5. Phyllanthus cuatrecasanus G. L. Webster, sp. nov. TYPE: Colombia. Caqueti: Quebrada del Rio Hacha, m, J. Cuatrecasas 8533 (holotype, US). Figure 1.

4 Volume 12, Number Webster 293 Similis Phyllanthi valerii, differt foliis laevibus, dolabriformis, nervis lateralibus rectis; stipulis minoribus; segmentis disci tenuioribus. Dioecious shrub; branchlets terete, copiously This new section of hirtellous, 8-14 cm long, mm broad, with Phyllanthusubg. Conami ca leaves; leaf blades glabrous, mostly 5-7 (Aublet) G. L. Webster, named for the prevalent distribution of its mm species in lowland Amazonian broad, elliptic, abruptly acute, + inaequilateral forest at base, abaxially alveolate-reticulate, veins (Hylaea), has escaped recognition partly because of its straight, these and reticulum of vegetative resemblance to species of veinlets prominutwo sections of lous; stipules oblong, blunt, glabrous, x 0.3- Phyllanthusubg. Xylophylla (L.) Persoon: section 0.4 mm; staminate flowers 2-5 per axil on brach- Brachycladus G. L. Webster (Webyblasts; pedicel mm long; sepals 5, 1.3- ster, 2001) and section Elutanthos Croizat (Croizat, 1.6 X mm; disk segments 5, elliptic, ); this has led to confusion and failure to rec mm broad; stamens 2, filaments ognize the significance of the diagnosticharacters. completely The resemblance of the connate into a column mm high; anthers < globose pollen grains with 0.5 mm long, dehiscing obliquely; pistillate flowers pilate exine ornamentation to those of section Nothoclema G. L. Webster unknown. clearly supports placement of section Hylaeanthus in subgenus Conami (Web- Known only from the type collection. The distin- ster & Carpenter, 2002). guished collector, Jose Cuatrecasas, described the Miller (1866, 1873) created much of the conperianth segments as "amarillentos con n&rvios ro- fusion with regard to the new section Hylaeanthus jizos." The specimen of Phyllanthus cuatrecasanus when he mistakenly identified the type species, superficially resembles small-leaved forms of Phyllanthus attenuatus Miquel, with P aubletianus ruscifolius, but differs sharply from that species (as P. Radcliffe-Smith (2001; name replacing Meborea well as P valerii) in the glabrous, strongly inae- guianensis Aublet). The affinities of Aublet's Mequilateral apiculate leaf blades with curious glau- borea remain uncertain, but it seems possibly a cous-reticulate bands alternating with green lateral species of section Brachycladus. At present, section veins. Hylaeanthus includes 6 species, although additional species remain unpublished. It is biogeographi- 6. Phyllanthus valerii Standley, Field Mus. Publ. cally parallel to section Pityrocladus in its mainly Bot. 18: TYPE: Costa Rica. He- South American distribution, but with one species redia: Yerba Buena, NE of San Isidro de He- in Costa Rica. redia, 2000 m, P. C. Standley & F L. Valerio KEY TO THE SPECIES OF PHYLLANTHUS SECT. HYLAEANTHUS (holotype, F). Monoecious or dioecious trees or shrubs; branchlets pinnatiform, usually subtended by reduced leaves (but generally not by cataphylls); flowers in axillary unisexual or bisexual cymules borne on brachyblasts; sepals 6 (rarely 5), usually biseriate; disk of discrete segments or cupular; stamens 3, filaments free or connate; anthers muticous, dehiscing horizontally obliquely; pollen grains globose, porate (lacking distinct colpi), exine pilate; pistillate sepals persistent in fruit; ovary 3-5-locular; styles free or connate, bifid to unlobed; fruits baccate; seeds smooth. Montane forests, Costa Rica, m; la. Ovary 3-locular; styles bifid; monoecious or dioecious. very 2a. Leaves completely glabrous; dioecious. similar plants have been collected in Bolivia. 3a. Leaf blades lanceolate, abruptly acuminate; styles connate..... P skutchii Phyllanthus subg. Conami (Aublet) G. L. Webster 3b. Leaf blades usually ovate or elliptic, blunt; styles free P manausensis 2b. Leaves hirtellous at least on petiole; monoecious Phyllanthus sect. Hylaeanthus G. L. Webster, or dioecious. sect. nov. TYPE: Phyllanthus attenuatus Mi- 4a. Fruiting pedicels 2-4 mm long; styles free, quel. bifid or unlobed. 5a. Dioecious; styles bifid; disk entire A sectione Nothoclema differt ramulis pinnatiformibus P attenuatus (non bipinnatiformibus), fructibus baccatis; a sectione 5b. Monoecious; styles unlobed; & disk dis- Apolepis differt filamentis connatis, disco 9 integro. sected P bernardii 4b. Fruiting pedicels 6-7 mm long; styles connate, twice-bifid...5. P. callejasii lb. Ovary 4-5-locular; styles unlobed; filaments connate; monoecious...6. P. madeirensis 1. Phyllanthus skutchii Standley, Publ. Field Mus. Nat. Hist., Bot. 22: TYPE: Costa Rica. San Jos0: El General, A. Skutch 4375 (holotype, F). Evergreen forests, m, southern Costa

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6 Volume 12, Number Webster 295 Rica (Pacific slope); possibly occurring in Colom- petiole 5-7 x mm, copiously scabrid-hirtelbia. lous; stipules blackish, triangular, 2-3 mm long. Flowers in axillary glomerules, the pistillate mostly 2. Phyllanthus manausensis W. A. Rodrigues, 2-5 per axil; bracts and bractlets blackish, persis- Acta Amazonica 1: 17, fig TYPE: tent. Staminate flowerseen only in bud; sepals 6; Brazil. Amazonas: Manaus, W A. Rodrigues disk undivided; stamens 3, filaments connate, an (holotype, INPA 16822). thers dehiscing horizontally. Pistillate flowers with pedicel glabrous or sparsely and minutely hirtel- Lowland Amazonian rain forest, mostly on terra firme, m, Brazil (Amazonas, Pard, Rorai- lous, mm long in fruit; fruiting sepals (5)6, mm long, mm broad; disk patelliform, ma, Amapia). ca. 1.5 mm broad; ovary 3-locular; styles connate into a column ca. 1.5 X 0.7 mm; style branches 3. Phyllanthus attenuatus Miquel, Linnaea 21: mm long, TYPE: Suriname. F W R. Hostman + twice-bifid. Fruits baccate, 7-9 mm diam. (dried); seeds somewhat compressed, & A. Kappler 305 (holotype, L; isotype, W). acute, truncate at base, X mm; testa Terra firme or occasionally inundated rain forest, smooth, brownish m, Colombia, Venezuela, the Guayanas, Cloud forest, m, Cordillera Occiden- Brazil, and Peru. tal, Colombia. Phyllanthus callejasii is clearly related to Phyllanthus attenuatus Miquel by its bifid 4. Phyllanthus bernardii Jablonski, Mem. New styles and dioecious flower production, but is easily York Bot. Gard. 17: TYPE: Venedistinguished by its larger leaf blades and longer zuela. M6rida: Aricagua, L. Bernardi 6258 (ho- petioles, thicker branchlet axes, and longer fruiting lotype, NY). pedicels and sepals. It differs from the lowland R Andean forests, > 2000 m, western Venezuela attenuatus and P manausensis in its upper montane (M6rida). The species deviates from others in cloud forest habitats. Phyl- Phyllanthus bernardii, the lanthus sect. Hylaeanthus in its character combi- only other upland species of Phyllanthusect. Hynation of monoecious flower production, laeanthus, has hirtellous branchlet axes, winged winged pedicels, and small seeds (< 2 mm long). It has pedicels, and smaller seeds. not yet been possible to examine the pollen, and it Additional collections. COLOMBIA. Antioquia: Mun. may prove not to belong to the section. Frontino, Corr. Nutibara, zona de Murri, 5-8 km S de Alto de Cuevas, m, 14 Feb. 1991, R. Callejas, E 5. Phyllanthus callejasii G. L. Webster, sp. nov. TYPE (pistillate): Colombia. Antioquia: Mun. Frontino, Corr. La Blanquita, regi6n de Murri, Alto de Cuevas, 1850 m, 14 July 1988, R. Callejas, A. L. Arbeldez, J. Betancur & L D. Castaiio 6864 (holotype, HUA 52926; isotype, DAV). Figure 2. J. Rolddn & M. V Arbeldez 9940 (staminate paratype: HUA 79857). Narifio: Mun. Piedraancha, Correg. Chucunez, Reserva La Planada, m, 18 May 1991, J. Betancur, S. Churchill & E J. Rolddn 2576 (HUA). 6. Phyllanthus madeirensis Croizat, Trop. Woods 78: TYPE: Brazil. Amazonas: Mun. Humaiti, between Rio Livramento and Rio Ipixuna, B. A. Krukoff 7163 (holotype, A; Ab aliis speciebusectionis differt stylis connatis bis bifidis, sepalis fructiferis majoribus, ramulis isotype, NY). crassioribus. Amazonian rain forest, Dioecious tree 6-8 mostly on terra firme, ca. m high; branchlets terete, m, Brazil smooth (Amazonas, Pard). This species (minutely papillate), cm long, 1.5- is 2.5 mm sharply thick, with 6-9 separated by a number of characters from leaves; leaf blades ovate to the other elliptic, 7-12 cm long, species of the section, but the distinctive cm broad, glabrous except for pollen the incised midrib sculpturing supports placement in Phyllanadaxially and midrib thus sect. Hylaeanthus. and major lateral veins abaxially, veins 5-7 pairs, curving; veinlet reticulum prominulous beneath; Phyllanthus subg. Cyclanthera G. L. Webster, subg. nov. TYPE: Phyllanthus lindenianus Baillon. Figure 1. Phyllanthus cuatrecasanus G. L. Webster. - Left-hand photo. Type specimen (Cuatrecasas 8533, US). -Right-hand photo. Enlarged view of branch apex with deciduous branchlets. Herbae monoicae, ramificatione more sectionis Phyllanthi; flores solitarii, axillares; sepala d 5 vel 6; disci segmentis plerumque discretis; stamina 2 vel 3, connata; antherae discretae vel connatae, horizontaliter dehiscen-

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8 Volume 12, Number Webster 297 tes; pollinis grana exinio clypeato; sepala 9 6; discus dis- Sect. 2. Phyllanthus sect. Cyclanthera G. L. sectus; ovarium 3-loculare; styli bifidi; fructus capsularis; semina scabridula vel verruculosa. Webster, Contr. Gray Herb. 176: TYPE: Phyllanthus lindenianus Baillon. Monoecious annual or perennial herbs; branchlets pinnatiform with one iterative lateral axis; Phyllanthusect. Cyclanthera includes four speflowers axillary, solitary; staminate sepals 5 or cies 6; (and some subspecies) of Hispaniola, two of disk dissected; stamens 2 or which 3, filaments also occur in eastern Cuba connate; (Webster, anthers discrete or completely connate into a 1958). The section is very isolated within Phyllansynthus andrium, dehiscing horizontally; pollen grains by its highly modified androecium with unique spheroidal, clypeate, the exine shields banded or clypeate pollen grains (illustrated by Webster & circular; pistillate sepals 6; disk dissected, often Carpenter, 2002). purplish; ovary 3-locular, styles bifid; fruit capsu- Acknowledgments. I thank Kevin Carpenter for lar; seeds scabridulous or verruculose. his assistance in obtaining critical pollen evidence Phyllanthusubg. Cyclanthera includes two secwith scanning electron microscopy at the Microstions, section Callitrichoides G. L. Webster and sec- copy and Imaging Facility at the University of Caltion Cyclanthera G. L. Webster, both endemic to ifornia, Davis. Photographs of herbarium specimens the Greater Antilles. When these sections were first were taken by Sam Woo, at the Instructional Techtreated (Webster, ), they were included nology and Digital Media Center, University of Calin subgenus Phyllanthus, but it was noted that ifornia, Davis. Thanks are also due to the curators they have extraordinary pollen grains somewhat comof several herbaria (including A, F, GH, NE, NY, parable to the clypeate pollen grains (with exine TEX, UC, US) for the loan or donation of specishields) in subgenus Xylophylla (Halbritter & Hes- mens, and particularly to Ellen Dean, director of se, 1995). Recent studies of the Davis Herbarium, for pollen morphology providing the facilities for (Webster& Carpenter, 2002) graphically indicate this research. The manuscript has been signifithe divergence of the pollen grains in section Cycantly improved by suggestions from Petra Hoffclanthera from all other taxa of Phyllanthus. It mann (Royal Botanic Gardens, Kew) and Peter van therefore seems necessary to remove these odd Welzen (Rijskherbarium, Leiden). West Indian plants from subgenus Phyllanthus into a separate subgenus. The relationships of Literature Cited subgenus Cyclanthera still remain uncertain, although the Bancilhon, L Contribution i l'6tude taxonomique distinctive pollen shows some resemblance to the du genre Phyllanthus (Euphorbiac6es). Boissiera 18: 1- pollen of subgenus Emblica. 81. Bentham, G Phyllanthus. Flora Australiensis 5: L. Reeve, London. Sect. 1. Phyllanthus sect. Callitrichoides G. L. Croizat, L Notes on American Euphorbiaceae with Webster, Contr. Gray Herb. 176: descriptions of eleven new species. J. Wash. Acad. Sci. TYPE: Phyllanthus carnosulus Muller 33: Argoviensis. Govaerts, R., D. G. Frodin & A. Radcliffe-Smith World Checklist and Bibliography of Euphorbiaceae. One species endemic to eastern Cuba; apparent- Royal Botanic Gardens, Kew. ly related to Phyllanthus tenuicaulis Maller Halbritter, H. & M. Hesse The convergent evolu- Argo- tion of exine shields in angiosperm pollen. Grana 34: viensis (in sect. Cyclanthera). This curious plant is vegetatively distinctive in its crisp-succulent leaves Hunter, J. T. & J. J. Bruhl Three new species of and stems rooting at the nodes. The pollen grains, Phyllanthus (Euphorbiaceae: Phyllantheae) in South in contrast to those of section Cyclanthera, have Australia. J. Adelaide Bot. Gard. 17: Phyllanthus. In N. G. Walsh & J. Enelongated exine shields (or perhaps could be intertwisle,. Flora of Victoria, vol. 4: Inkata T. Press, preted as stephanocolporate). Phyllanthus carno- Melbourne. sulus appears not to have been collected since it Jablonski, E Euphorbiaceae. Botany of the Guawas found by Ekman over 70 years ago, so it is not yana Highland-Part VII. Mem. New York Bot. Gard. certain whether it still survives. 17(1): Lanjouw, J The Euphorbiaceae of Surinam. Proefschrift, Rijks-Universiteit, Utrecht. Muller, J Euphorbiaceae [except Euphorbieae]. In A. De Candolle, Prodromus systematis naturalis regni Figure 2. Phyllanthus callejasii G. L. Webster. -Left- vegetabilis 15(2): Victor Masson, Paris. hand photo. Type specimen with pistillate flowers (Callejas Phyllanthus. In F. von Martius (editor), et al. 6864, HUA). -Right-hand photo. Specimen with Flora Brasiliensis 11(2): fruits (Betancur et al. 2576, HUA). Nozeran, R., L. Rossignol-Bancilhon & G. Mangenot.

9 298 Novon Les recherches sur le genre : Acquis et perspectives. Bot. Helv. 94: Punt, W Pollen morphology of the genus Phyllanthus (Euphorbiaceae). Rev. Palaeobot. Palynol. 39: A survey of pollen morphology in Euphorbiaceae with special reference to Phyllanthus. Bot. J. Linn. Soc. 94: Radcliffe-Smith, A Genera Euphorbiacearum. Royal Botanic Gardens, Kew. Webster, G. L A monographic study of the West Indian species of Phyllanthus. J. Arnold Arbor. 37: , , ; 38: 51-80, , ; 39: , The genera of Euphorbiaceae in the southeastern United States. J. Arnold Arbor. 48: Phyllanthus. Pp in P. E. Berry, K. Yatskievych & B. Holst (editors), Flora of the Venezuelan Guayana, Vol. 5. Missouri Botanical Garden Press, St. Louis. ' Synopsis of Croton and in western tropical Mexico. Contr. Univ. Michigan Herb. 23: & K. J. Carpenter Pollen morphology and phylogenetic relationships in neotropical Phyllanthus (Euphorbiaceae). Bot. J. Linn. Soc. 138: