The synopsis of Phyllanthus section Phyllanthus in Brazil treats 4 subsections (2 of them newly described) including 30 species. A new subsection Almadenses is described based on Phyllanthus almadensis Müll. Arg., a species from Bahia, Brazil with a unique growth pattern. Another new subsection Clausseniani is proposed; it includes 17 species, all endemic to Brazil. Seven new species in this subsection are described: Phyllanthus allemii, P. atalaiensis, P. caparaoensis, P. carvalhoi, P. mocotensis, P. piranii, and P. sincorensis. A key to the Brazilian sections of subgenus Phyllanthus is provided, as well as synoptic keys and descriptions for the two new subsections and subsections Niruri and Swartziani.
Phyllanthus L., the third largest genus of Euphorbiaceae, with more than 800 species (Govaerts et al., 2000), is represented in the Neotropics by 8 subgenera, 33 sections, and approximately 200 species (Webster, ined.). A considerable number of new sections and species have been described since the synopsis of the West Indian taxa (Webster, 1956–58). In that study, it was shown that pollen morphology provides critical characters for determining phylogenetic affinity; those light microscopic observations have recently been confirmed and extended using scanning electron microscopy (Webster and Carpenter, 2002).
In the current circumscription of Phyllanthus subgenus Phyllanthus (Webster, 1970), most of the species belong to section Phyllanthus, in which the habit is herbaceous or shrubby, branching is phyllanthoid (Webster, 1957) and the androecium consists of 2 or 3 (rarely 4) stamens. Four of the five American subsections of section Phyllanthus occur in Brazil, which has a total of approximately 30 species. Since the treatment of Muller in the Flora Brasiliensis (1873), there has not been an overall review of the Brazilian species of section Phyllanthus, although a few species have been treated by Webster (1959), Jablonski (1967), Smith et al. (1988), and Cordeiro (1989, 1992). The overall similarity in habit and the very small flowers of many species cause difficulties in identification; a high percentage of the specimens in herbaria are misdetermined. There appear to be polyploid complexes in subsections Clausseniani, Niruri, and Swartziani (Mangenot et al., 1977), and a considerable number of fragmentary specimens cannot be determined with certainty. The relationships of the two autochthonous subsections, Almadenses and Claussenianiy await clarification..
It is now clear that subsection Claussenianiy with 17 species—7 of which are here described as new—is the most speciose group of herbaceous and suffruticose species of Phyllanthus in Brazil. It is notable that the distribution of the subsection is entirely extra-Amazonian; in the Amazon basin, it is replaced by species of subsection Swartziani. Discussions of the characteristic non-Amazonian habitats in eastern Brazil may be found in Eiten and Goodland (1979) and in Rizzini (1979).
Species of section Phyllanthus have long been of considerable ethnobotanical interest, particularly for their use in folk medicine to treat liver or bladder ailments (Uphof, 1968). For Brazil, Correa (1969) listed medicinal species that belong to three subsections: Niruri—Phyllanthus niruri L. (as “Phyllanthus lathyroides”); Swartziani— P. amarus Schumach. & Thonn. (as “Phyllanthus niruri”), P. lindbergii Müll. Arg., and P. stipulatus (Raf.) G. L. Webster (as “Phyllanthus diffusus”); and Clausseniani— P. acutifolius Poir. ex Spreng. Corréa used the common names “Herva Pombinha” and “Quebra-Pedra.” The prevalent taxonomic confusion of names of species in sect. Phyllanthus in Corréa's synonymy is shared by most works on medicinal plants earlier in the 20th century. In contrast to Corréa, whose illustration of Phyllanthus niruri is correct while his description (based on P. amarus) is misapplied, Matos (1989) correctly illustrates and applies the name Phyllanthus amarus. Matos considers that the vernacular name “Quebra-Pedra” is mainly applied to Phyllanthus urinaria L. (section Urinaria) in the Amazon, to P. amarus in the Nordeste, and to P. niruri (s. str.) in central and southern Brazil. The widespread confusion in applying Latin names to species in section Phyllanthus has become distressing due to the recent discoveries of anti-viral properties in several species, with possible significance for treating Hepatitis B and cancer (Blumberg et al., 1989). A taxonomically oriented survey of potential medicinal applications has shown that within Phyllanthus species of sect. Phyllanthus are of particular interest as potential sources of therapeutic compounds (Unander et al., 1991). It is hoped that this synopsis of the Brazilian species may prove to be useful for investigators of possible medical uses.
In the course of preparing this synopsis, I have become uncomfortably aware of its very provisional nature. At least two species complexes—Phyllanthus niruri/rosellus in subsect. Niruri and P. lindbergii/microphyllus in subsection Swartziani—present so much variability and intergradation that the present treatment does not adequately present the reality; intensive field studies of these problematic populations are needed.
In Brazil there are species of 3 other sections of subgenus Phyllanthus that may be confused with those in section Phyllanthus. In order to avoid uncertainty as to whether or not an unidentified specimen is a species of section Phyllanthus, a key to the sections in Brazil is presented here.
Key to the Sections of Subgenus Phyllanthus in Brazil
1. Branchlets deciduous, with well-developed leaves; axes not dilated into phylloclades.
2. Stamens 5, filaments free; proximal cymules of branchlet bisexual; pollen grains 4-colporate; seeds puncticulate. [P. tenellus Roxb.] Pentandra
2. Stamens 2 or 3 (rarely 4), filaments free or more often partially to completely united; cymules of branchlet unisexual (except P. amarus); pollen grains 3- or 4-colporate; seeds striate, ribbed, or puncticulate.
3. Pistillate flowers proximal on branchlet, staminate distal; seeds transversely ribbed; pollen grains 4-colporate; ovary verrucose. [P. urinaria L] Urinaria
3. Pistillate flowers distal on branchlet, staminate proximal; seeds longitudinally ribbed, striate, or puncticulate; pollen grains 3- or 4-colporate; ovary usually smooth Phyllanthus
1. Branchlets persistent, leaves usually reduced to scales; axes dilated into phylloclades Choretropsis
Conspectus of the Brazilian Taxa of Phyllanthus Section Phyllanthus
Phyllanthus section Phyllanthus. TYPE: Phyllanthus niruri L.
Subshrubs or herbs; monoecious or dioecious; Leaves alternate (opposite in P. almadensis), on penultimate axes usually (at least distally) reduced to scales; Flowers mostly in axillary glomerules; Sepals (4) 5 or 6; Stamens 2 or 3 (4), filaments free to completely connate; Anthers dehiscing vertically to horizontally, connective sometimes enlarged; Pollen grains mostly subprolate to prolate, 3- or 4-colporate, sexine reticulate to finely tectate-perforate; Pistillate disk annular to cupular, sometimes lobed; Ovary 3-locular, glabrous (rarely papillose or pubescent); Styles bifid (sometimes only emarginate); Fruits capsular; seeds striate, finely ribbed, or puncticulate.
One of the largest sections of the genus, section Phyllanthus is pantropical, and includes nearly 100 species in 7 subsections (Webster, ined.). Five subsections occur in the neotropics, of which 4 are represented in Brazil (subsection Pentaphylli G. L. Webster is confined to the Caribbean).
Key to the Brazilian Subsections of Phyllanthus Section Phyllanthus
1. Branchlets with only 2 subopposite leaves, terminating in a racemose inflorescence; anther connective enlarged; pollen grains tectate-perforate. Subsect. 2. Almadenses
1. Branchlets with >10 alternate leaves, flowers in axillary cymules; anther connective enlarged or not; pollen grains reticulate to tectate-perforate.
2. Seeds puncticulate; leaf blades mostly inaequilateral at base; cymules unisexual; pollen grains prolate, 4-colporate, exine reticulate; style - branches capitate Subsect. 1. Niruri
2. Seeds striate, ribbed, or puncticulate; leaf blades not (or scarcely) inaequilateral at base; cymules unisexual or bisexual; pollen grains subspheroidal to prolate, 3- or 4-colporate, exine reticulate to tectate-perforate; style branches usually not capitate.
3. Filaments connate; anther connective not appreciably enlarged, not deeply emarginate; pollen grains subspheroidal to prolate, 3- (4-) colporate; sexine finely reticulate to tectate-perforate. seeds longitudinally striate or ribbed (rarely puncticulate); branching strictly phyllanthoid, the distal leaves on penultimate axes reduced to cataphylls; branchlet axes usually not zig-zag Subsect. 3. Swartziani
3. Filaments usually free; anther connective distinctly enlarged, deeply emarginate, the thecae discrete and sometimes stipitate; pollen grains mostly subspheroidal, 4-colporate, coarsely reticulate; seeds mostly puncticulate (sometimes striate); branching not phyllanthoid, leaves on penultimate axes not reduced to cataphylls in some species; branchlet axes often zig-zag Subsect. 4. Clausseniani
Subsect. 1. Niruri G. L. Webster, Contr. Gray Herb. 176: 52. 1955; J. Arnold Arbor. 38: 399. 1957. Niruri Adanson, Fam. Pl. 2: 356. 1763. Type: Phyllanthus niruri L.
Herbs or shrubs; cataphyllary stipules linear-lanceolate; Leaf blades obliquely inaequilateral at base; Sepals 5; stamens 3, filaments free or connate proximally; anthers dehiscing obliquely or horizontally; Pollen grains prolate, 4-colporate; pistillate disk patelliform; Ovary smooth and glabrous; Styles free, bifid, tips subcapitate; Seeds puncticulate.
This subsection of 5 species is restricted to America, although it is possible that the west African species Phyllanthus benguelensis may prove to belong here.
Key to the Brazilian Species of Subsection Nlruri
1. Fruiting pedicels > 10 mm long; staminate sepals 1.9–2.4 mm long; stamens free. 1. P. augustinii
1. Fruiting pedicels < 10 mm long; staminate sepals not over 1.8 mm long; stamens free or connate.
2. Staminate sepals (1-) 1.2–1.8 mm long; seeds > 1 mm long.
3. Staminate flowers with pedicels mostly 1–3 mm long, calyx yellowish or greenish; leaf blades mostly 7–17 mm long 2. P. niruri
3. Staminate flowers with pedicels mostly 4–6 mm long, calyx ± reddish; leaf blades mostly 3–5 (-8) mm long 3. P. rosellus
2. Staminate sepals 0.5–1 mm long; seeds < 1 mm long 4. P. perpusillus
1.1. Phyllanthus augustinii Baill., Adansonia 1.5: 354. 1865; Müll. Arg., Fl. Brasil. 11(2): 65. 1873. Type: BRAZIL [Rio de Janeiro:] Rio de Janeiro, “montagne de S. Diogo,” A. St. Hilaire A1743 (holotype: P!).
This species remains poorly known; Muller (1873) stated that he had not seen Baillon's specimen. A collection from Rio de Janeiro at Kew (Glaziou 17757) has unusually large flowers and may represent Phyllanthus augustinii.
1.2. Phyllanthus niruri L., Sp. Pl. 981. 1753; Webster, J. Arnold Arbor. 37: 300, pi. XIX figs. A, B. 1957; Ann. Missouri Bot. Gard. 54: 225. 1968; Brittonia 22: 66. figs. 9, 10. 1970; Allem, Rev. Brasil. Biol. 37: 103.1977; Smith, Downs, & Klein, Fl. Ilustr. Catar. EUFO 43, pi. 6 figs. j-o. 1988; Burger & Huft, Fieldiana, Bot. n.s. 36: 145. 1995; Webster, Fl. Nicaragua 1: 898. 2001. Type: Herb. Hort. Cliffort. (holotype: BM!).
Phyllanthus lathyroides H.B.K., Nov. Gen. Sp. Pl. 2: 110. 1817. Phyllanthus niruri ssp. lathyroides (H.B.K.) Webster, Contr. Gray Herb. 176: 52. 1955. Type: VENEZUELA. [Monagas:] “Novae Andalusiae prope coenobium Caripense,” A. Humboldt & A. Bonpland s.n. (holotype: P!). Additional synonyms of P. niruri are cited in Webster (1957, 1970). Distribution and habitat: Widespread in tropical America from Mexico and the West Indies south to Argentina and Brazil, mostly in rain forest, but also in deciduous forest or cerrado, 0–1000 m.
Representative Specimens: Acre: Reserva Extrativista Chico Mendes, Fereira & Ming 107 (NY). Amazonas: Ilha do Yarary, Kuhlmann 388 (RB). Bahia: Mun. Santo Amaro: 10 km W of Santo Amaro, Webster, Dehgan, & Queiroz 25820 (DAV). Ceará: Fortaleza, Serra de Maranguape, Egler 756 (MG). Distrito Federal: 20 km N of Brasília, Irwin et al. 11307 (UB). Fernando Noronha: Ilha Rata, Moseley (K), Miranda et al 840 (ESA). Maranhão: Mun. Loreto, Eiten 10440 (US). Mato Grosso do Sul: Mun. Corumbá, near Corumbá airport, Webster & Pott 25372 (DAV, UEC, NY). Minas Gerais: Serra Mantiqueira, cerca Ouro Preto, Calago 272 (DAV). Paraná: Curitiba, Hatschbach 3655 (DAV). Pernambuco: Arcoverde, Estação Experimental, Andrade Lima 71-6374 (IPA). Rio de Janeiro: Parque Nacional Serra dos Orgaos, Webster, Dehgan, & Oliveira 25865 (DAV, R). Rio Grande do Sul: Mun. Sao Vicente do Sul: Cerro Loreto, Webster, Dehgan, & Sobral 25965 (DAV, ICN). Santa Catarina: Mun. Florianópolis, Riberão da Ilha, Klein & Bresolin 5417 (US). Sño Paulo: Mun. Cananéia, Ilha Cardoso, Webster, Cordeiro, & Kirizawa 25560 (DAV, SP).
This highly variable and widespread neotropical species appears to represent a polyploid complex, with reported chromosome numbers of 2n = 14, 26, 36 (Mangenot et al., 1977). The circumscription of the species as adopted here must be regarded as tentative, and no attempt is made to formally recognize subspecific taxa. The specimens cited from Bahia and Rio de Janeiro represent Phyllanthus lathyroides var. microcarpus Müll. Arg., a delicate form with smaller parts that may prove to be distinguishable at a subspecific level.
1.3. Phyllanthus rosellus (Müll. Arg.) Müll. Arg., Fl. Brasil. 11(2): 53, pi. 6 fig. I. 1873.
Phyllanthus lathyroides *** [forma] rosellus Müll. Arg., Linnaea 32: 42. 1863. Type: BRAZIL. Minas Gerais: Villarica [Ouro Preto], Vauthier 83 (Lectotype [designated here]: W!).
Distribution and Habitat: Cerrado and cerradão, often on sandy soils, 8002000 m, extending into adjacent Argentina and Paraguay.
Representative Specimens: Minas Gerais: Baipendi, Brade & Apparicio 20456 (RB); Diamantina, Brade 13584 (RB); Serra do Caraça, Pohl (M), Pereira 2395, Pabst 2543 (RB); Serra da Moeda, Duarte 9697 (RB); 35–40 km E of Belo Horizonte, Irwin, Harley, & Onishi 30219, 30453 (DAV, NY). Paraná: 7 km E of Contendas, Krapovickas & Cristóbal 39631 (CTES); 10 km W of Guarapuava, Reitz & Klein 17639 (US); Jaguariaiva, Smith, Klein, & Hatschbach 14748 (US). Rio de Janeiro: Itirapina, Toledo 594 (RB); Petrópolis, Góes & Dionisio 488, 599 (RB). São Paulo: 15 km NW of Angatuba, J.R. & N.F. Mattos 8172 (US).
It is with considerable diffidence that Phyllanthus rosellus is listed here as a distinct species. Muller (1863, 1866) originally regarded it as a form of P. lathyroides [P. niruri], but later elevated it to specific rank because of its solitary, “slightly larger” flowers, smaller leaves, and a dubious distinction in seed sculpturing. In Minas Gerais, P. rosellus appears clearly distinguishable from P. niruri, but small-leaved forms of the latter from Bahia and Rio de Janeiro are problematic. Allem (1977) has pointed out several problems in the taxonomic characterization of P. rosellus, and suggested that it be treated as a synonym of P. niruri.
1.4. Phyllanthus perpusillus Baill, Adansonia I. 5: 358. 1865; Müll. Arg., Fl. Brasil. 11(2): 55. 1873; Webster, Sellowia 11: 169. 1959; Smith, Downs, & Klein, Fl. II-ustr. Catar. EUFO 51. 1988. Type: BRAZIL. Minas Gerais: Ponte Alta, A. St. Hilaire Cat. D, 301d (holotype: P!; isotype fragment: A!). Webster (1959) pointed out that Muller's description (1873) was based mainly on a collection of Pohl that appears not to be conspecific with St. Hilaire's type collection.
Distribution and Ecology: Recorded only from Minas Gerais and Santa Catarina, in campos, 750–1000 m. The apparently disjunct distribution of Phyllanthus perpusillus may be an artifact of collecting, since this diminutive species may have escaped detection (perhaps through confusion with species of subsection Swartziani).
Representative Specimens: Santa Catarina: Irani, Reitz & Klein 4707 (US); Mafra, Reitz & Klein 11502 (HBR, US); Rio Negrinho, Reitz 5200 (US).
Subsection 2. Almadenses G. L. Webster, subsect. nov. TYPE: Phyllanthus almadensis Müll. Arg.
Ab aliis subsectionibus sectionis Phyllantho differt ramulis bifoliatis, foliis suboppositis, racemis terminalibus; anthers malliforis.
Herbs, monoecious, glabrous, with single main stem; Branchlets with a terminal pair of subopposite leaves, terminating in a raceme; bracts stipulate; proximal cymules staminate, distal bisexual; Flowers pedicellate; sepals 5; staminate disk segments 5; stamens 3, filaments free, anthers dehiscing laterally, with enlarged flattened connective, Pollen grains prolate, 3-colporate, exine tectate-perforate; Pistillate disk cupular, entire; Ovary smooth; Styles bifid, free; Seeds unknown.
This subsection takes its name from the type (and only known) species, which was collected by Von Martius near Almada, in Bahia, Brazil. Because of the distinctiveness of the type species, a full description is provided. When Muller described Phyllanthus almadensis in the “Flora Brasiliensis” (1873), he noted its characteristic inflorescence, but grouped it with a number of unrelated species such as P. attenuatus Miq. and P. sellowianus Müll. Arg. Because he did not have a clear grasp of the significance of phyllanthoid branching, he failed to emphasize the extraordinary vegetative morphology of P. almadensis, in which the branchlets bear a single pair of opposite leaves and terminate in a racemose axis. This morphological pattern is unique in the entire genus Phyllanthus, and thus sets P. almadensis apart from all other species. A few New Caledonian species in subg. Gomphidium have branchlets with only 1 or 2 leaves (Schmid, 1991), but these do not subtend a terminal raceme as in P. almadensis. However, the flowers of this Brazilian species are similar in size and number of parts to species of section Phyllanthus as defined by Webster (1967, 1970). Furthermore, SEM observations on pollen morphology of P. almadensis (Webster and Carpenter, 2002) indicate a general resemblance to pollen grains of species in sect. Phyllanthus such as those described (using light microscopy) by Punt (1962) and illustrated using SEM by Punt (1987) and by Punt and Rentrop (1974) and (for Mauritian species) by Bor (1979). It therefore appears reasonable to assign Phyllanthus almadensis to a separate subsection in subgenus Phyllanthus section Phyllanthus.
The enlarged anther connective of Phyllanthus almadensis suggests a possible affinity with subsection Clausseniani; however, the anther thecae are not distinctly separated, and the finely tectate-perforate pollen exine of P. almadensis, as illustrated by Webster and Carpenter (2002), is quite different from the pollen exine of species of subsection Clausseniani, which is coarsely reticulate with luminar baculae. Species of subsection Swartziani have pollen grains more similar in shape (i.e., more prolate) and with finely tectate-punctate exine. However, in that subsection there is no approach to the vegetative, inflorescence, and anther morphology of Phyllanthus almadensis. Thus, at present the relationships of subsection Almadensis have to be regarded as uncertain.
2.1 Phyllanthus almadensis Müll. Arg., Fl. Brasil. 11(2): 28. 1873. TYPE: BRAZIL. Bahia: sylvis ad lacum da Almada, I 1819. C. F. P. von Martius (Holotype: M!; Isotype: G!). Taking into account the description of the Rio Almada by Paynter & Traylor (1991), the lake described by Martius appears on the “milionésimo” map (“Carta do Brasil ao Milionésimo”) at approximately 14°35′ S, 39°06′ W, c. 25 km NNW of Ilhéus. (Fig. 1)
Monoecious herb, annual or shortlived perennial, glabrous, 20–40 cm high, with single usually unbranched main stem. Branchlets 1.5–2.5 cm long and 0.4–0.8 mm thick, terminating in a pair of opposite leaves that subtend a terminal raceme. Leaf blades chartaceous, oblong-ovate, tapering to an obtuse or subacute tip, slightly paler abaxially, adaxially smooth (minutely alveolate); veins 5–10 per side, straightish, ± brochidodromous, scarcely raised, lacking a fine reticulum; margins plane; petiole 1–1.5 mm long; stipules dark, lanceolate, entire, 1.5–2.5 mm long. Racemes terminal, slender, 4–8 cm long, with 7–15 nodes; proximal cymules staminate, mostly with 3–8 flowers, at distal nodes with solitary pistillate flower; bracts 1.2–1.5 mm long, narrowly lanceolate, appearing trifid due to stipules. Staminate flowers with pedicel 1.7–2.7 mm long; sepals 5, obovate, 1-veined, 1.7–2.2 mm long, 1–1.5 mm broad; disk segments 5, thickened, 0.3–0.5 mm across; stamens 3, filaments free, 0.7–1.7 mm long; anthers 0.3–0.5 mm long and broad, with flattened expanded connective, dehiscing laterally (± horizontally), Pollen grains prolate, 3-colporate, exine finely tectate-reticulate. pistillate flowers with pedicels 3–4 mm long; sepals 5, elliptic, 2.3–3.5 mm long; disk patelliform, slightly cupular, 0.2 mm high, 1.0 mm broad; styles free, erect, ca. 1 mm long, shortly bifid, tips subcapitate. Fruits and seeds not seen.
Additional Specimens Examined: Bahia: Mun. Ilhéus, Pirataquisse, Pimenta Vielloso 903 (R); Mun. Itabuna, bairro Pedro Jerônimo, dos Santos 3614 (CEPEC, DAV).
Phyllanthus almadensis appears to be restricted to a small area (ca. 30 km across) of the coastal rain forest (“mata atlantica”) in the vicinity of Ilhéus, Bahia (vegetation mapped by Brazão & Araújo, 1981). Although Sr. dos Santos (personal communication) reported that it occurred within the city limits of Itabuna, my attempts to locate it there were unsuccessful.
Subsect. 3. Swartziani G. L. Webster, Contr. Gray Herb. 176: 53. 1955; J. Arnold Arbor. 38: 306. 1957; Brittonia 22: 68. 1970. Type: Phyllanthus swartzii Kostel. (= P. amarus Schumach. & Thonn.).
Monoecious or dioecious annual or perennial Herbs; branching strictly phyllanthoid (distal leaves on penultimate axes reduced to cataphylls); Leaves aequilateral at base (or nearly so); Stamens 2 or 3, filaments partially or wholly united; anthers mostly dehiscing horizontally, connective not enlarged, not deeply emarginate between the thecae; Pollen grains 3-colporate, exine finely reticulate or tectate-perforate; Pistillate disk entire to lobed or dissected; ovary smooth; styles bifid, branches not capitate; Seeds longitudinally striate or ribbed (puncticulate in Phyllanthus leptophyllus).
This subsection of ca. 25 species is pantropical in distribution, with the main concentrations of species in America, Africa, and India. Most of the Brazilian species have been recently characterized in the treatment of species in Venezuelan Guayana by Webster (1999). In contrast to subsection Clausseniani, there are no species of subsection Swartziani endemic to Brazil, except for P. leptophyllus, whose position in this subsection is doubtful. Only 5 of the 8 Brazilian species are commonly encountered; Phyllanthus caribaeus, P. debilis, and P. leptophyllus are only recorded from single specimens. The single introduced species, Phyllanthus debilis, might be confused with the exotic species P. tenellus Roxb. (section Pentandra), but the latter is easily distinguished by its longer fruiting pedicels and androecium of 5 free stamens. It is possible that an additional introduced species, Phyllanthus fraternus G. L. Webster, has been found in Rio Grande do Sul (C. A. Machado, in litt.). It is vegetatively similar to P. amarus, but differs in its unisexual cymules and sharply angled scabridulous branchlet axes.
Key to the Brazilian Species of Subsection Swartziani
1. Cymules unisexual, the staminate flowers at proximal nodes of the branchlet; sepals obtuse to rounded at tip.
2. Stamens 3 (rarely 2); monoecious or dioecious; plants relatively robust, stems mostly at least 1–1.5 mm in diameter; staminate pedicels usually at least 0.5 mm long, sepals > 0.5 mm long; seeds at least 1.1 mm long.
3. Monoecious; filaments partially or completely united; anthers dehiscing horizontally or obliquely; seeds ribbed or finely striate; leaves mostly obtuse or rounded at tip (except in P. debilis).
4. Deciduous branchlets terete; leaf blades obtuse to truncate at tip, often minutely scabridulous abaxially; seeds with 10–12 very fine longitudinal striae.
5. Pistillate disk 5-angled or lobed; filaments completely connate; stems often with basal aerenchyma; pistillate sepals and leaf margins sometimes reddish-tinged 1. P. stipulatus
5. Pistillate disk asymmetrically 3-lobed; filaments connate only in lower half; stems never with basal aerenchyma; pistillate sepals and leaf margins not reddish 2. P. caribaeus
4. Deciduous branchlets sharply angled; leaf blades ± pointed at tip, not minutely scabridulous abaxially; seeds with 6 or 7 longitudinal ribs 3. P. debilis
3. Dioecious or subdioecious; filaments usually completely united; anthers dehiscing vertically or obliquely; seeds finely striate or puncticulate; leaves obtuse to pointed at tip.
6. Anthers dehiscing vertically; staminate sepals 1.5–2.3 mm long; staminal column > 0.5 mm high; seeds 1.8–2 mm long. 4. P. lindbergii
6. Anthers mostly dehiscing obliquely; staminate sepals not over 1 mm long; staminal column mostly < 0.5 mm high; seeds < 1.5 mm long.
7. Seeds striate, 1.1–1.3 mm long; style branches not capitate 5. P. microphyllus
7. Seeds puncticulate, 1.3–1.4 mm long; style branches capitate 6. P. leptophyllus
2. Stamens 2 (rarely 3); monoecious; plants delicate, the stems mostly < 1 mm in diameter, staminate pedicels 0.2–0.5 mm long, sepals 0. 4–0.5 (-0.7) mm long; seeds 0.9–1 mm long. 7. P. minutulus
1. Cymules all bisexual, each of 1 pistillate and 1 staminate flower; sepals acute 8. P. amarus
3.1. Phyllanthus stipulatus (Raf.) G. L. Webster, Contr. Gray Herb. 176: 53. 1955; J. Arnold Arbor. 38: 315. 1957, pi. XIX, figs. L-M; Ann. Missouri Bot. Gard. 54: 226. 1968; Burger & Huft, Fieldiana, Bot. n.s. 36: 146, fig. 8. 1995; Webster, Fl. Nicaragua 1: 898. 2001. Moeroris stipulata Raf., Sylva Tellur. 91. 1838. Type: JAMAICA. O. Swartz (Lectotype [designated by Webster, 1955]: S!).
Phyllanthus diffusus Klotzsch, Bot. Voy. Herald 105. 1853. Type: Panama, B. C. Seemann s.n. (holotype: BM, not seen).
Phyllanthus hoffmannseggii Müll. Arg., Linnaea 32: 45. 1863. Type: BRAZIL. Pará, Hoffmansegg s.n. (holotype: B 17983!).
Distribution and Habitat: Mexico and the West Indies south to Argentina and Brazil, moist forest and swamps, often growing in inundated areas, 0–1100 m.
Representative Specimens: Acre: Mun. Manoel Urbano, Colocação Nova Olinda, Daly et al 9164 (DAV). Amazonas: Rio Purus, 2 km N of Labrea, Prance et al. 13719 (US). Bahia: Mun. Santa Cruz Cabrália: Estação Pau Brasil, Webster 25018 (CEPEC, DAV, NY, UEC, UEFS). Distrito Federal: Reserva Ecológica do Guará, Pereira da Silva et al. 1694 (CEN, DAV). Goiás: Mun. Uruaçu, 13 km NE of Vila Agua Branca, Walter et al. 1987 (CEN, DAV). Maranhão: Loreto, Eiten 3844 (SP). Mato Grosso: Mun. Cuiabá, Córrego Pindaival, Hatschbach et al. 36068 (DAV, MBM). Mato Grosso do Sul: Mun. Corumbá, Fazenda Nhumirim, pantanal, Webster & Pott 25356 (DAV). Minas Gerais: Corinto, Mexia 5662 (TEX, US). Pará: Belém, São Joaquim, Barbosa da Silva 60 (US). Rondônia: Mun. Sta. Bárbara, Teixeira etal. 723 (DAV). Roraima: Serra Tepequem, Maguire 40006, Prance et al. 4256 (US). São Paulo: Mun. Paulínia, Calago 256 (DAV). Santa Catarina: Pilhões, Palhoça, Reitz & Klein 2781 (HBR).
Other than the ubiquitous Phyllanthus amarus, this is the most widespread Brazilian species of section Phyllanthus. Although not as diversified as P. niruri, it does show considerable variation in leaf shape. Sporadically occurring forms have elongated, more pointed leaves.
3.2. Phyllanthus caribaeus Urb., Symb. Ant. 5: 382. 1908; Webster, J. Arnold Arbor. 38: 318, pi. XIX figs. N, O. 1957; Ann. Missouri Bot. Gard. 54: 227. 1968; Fl. Nicaragua 1: 896. 2001. Type: TRINIDAD & TOBAGO. Tobago, H. F. A. von Eggers 5733 (Lectotype [designated by Webster, 1957]: F!; Isolectotype: NY!).
Distribution and Habitat: Widespread in lowland rain forests, Nicaragua and Lesser Antilles south to northern South America, 0–700 m. Presently known only from one Brazilian record, but to be expected in other localities in western Amazonian Brazil.
Specimen Examined: Acre: Mun. Sena Madureira, Prance et al. 7540 (US).
3.3. Phyllanthus debilis Klein ex Willd., Sp. Pl. 4: 582. 1804; Webster, J. Arnold Arbor. 38: 307, pi. XIX figs. E, F; 1957; Fl. Ceylon 11: 224. 1997. Type: INDIA. Madras: Tranquebar, D. Klein 3081 (Lectotype [designated by Webster in Webster & Wheeler, 1997]: Herb. Willdenow 17978, B!).
Distribution and Habitat: Native to east Asia, introduced sporadically into the neotropics, often in rice fields, 0–100 m.
Specimen Examined: Rio de Janeiro: Morro da Babilônia, Hoehne 25049 (SP).
Only a single collection has been recorded from Brazil in this study, but Phyllanthus debilis is also recorded from the Lesser Antilles (Guadeloupe) and Hawaii (Webster, 1957).
3.4. Phyllanthus lindbergii Miill. Arg., Fl. Brasil. 11(2): 35, pl.4 fig. II. 1873; Webster, Sellowia 11: 168. 1959; Fl. Venez. Guayana 5: 197. 1999; Smith, Downs, & Klein, Fl. Ilustr. Catar. EUFO 43, pi. 6 figs. a-h. 1988. Type: BRAZIL. Minas Gerais: Caldas, 1855, G. A. Lindberg 438 ( Lectotype [designated here]: S!).
Distribution and Ecology: Colombia and Venezuela to Brazil and Paraguay, in swamps and wet savannas, 50–1100 m. In the northern part of its range, Phyllanthus lindbergii is sympatric in swampy areas with three other species of the subsection: P. microphyllus, P. minutulus, and P. stipulatus. A number of plants with ambiguous characters occur here, suggesting the possibility of a hybrid complex.
Representative Specimens: Amazonas: Mun. Humaitá, Cemtachójnicov & Janssen 389 (US). Goiás: Luziana, Heringer 17675 (US). Mato Grosso: Rio Brilhante, Hatschbach 30997 (DAV, US). Minas Gerais: Caldas, Regnell I 404, 404a (US). Paraná: Foz de Iguaçú, Schinini 8146 (CTES); Rio 10 km W of Guarapuava, Reitz & Klein 17614 (US); Jamaracarú, Black et al. 5719539 (DAV). Santa Catarina: Mun. Abelardo Luz, Smith & Klein 13847 (US); Mun. Campo Erê, Smith & Klein 13727 (HBR); Mun. Chapecó, Smith & Klein 9405, 11577 (HBR). São Paulo: Mun. Moji Guaçú, 3 km NNW of Pádua Sales, J.& N. Mattos 8226 (SP); Rio do Peixe, Edwall (SP 13722).
The “typical” form of Phyllanthus lindbergii, with large staminate flowers, vertically dehiscing anthers, and large seeds, is very distinct from the other species of subsection Swartziani. However, particularly in the northern part of its range (Amazonian Brazil, Venezuela), the flowers and seeds are smaller. It is not clear whether this is due to hybridization with P. microphyllus and/ or P. stipulatus.
3.5. Phyllanthus microphyllus H.B.K., Nov. Gen. Sp. 2: 109. 1817; Müll. Arg., Fl. Brasil. 11(2): 56. 1873; Webster, Fl. Venez. Guayana 5: 198. 1999. Type: VENEZUELA. “In ripa Orinoci?”, A. Humboldt & A. Bonpland s. n. (holotype: P!).
Distribution and Ecology: Pantanal and other swampy or inundated areas, Venezuela south to Brazil, 100–250 m.
Representative Specimens: Goiás: Goiânia, Brade 15406 (RB). Mato Grosso: Mun. Corumbá, Fazenda Marilândia, Pereira et al. 224 (RB). Mato Grosso do Sul: Campo Grande, Nienstedt 142 (RB). Minas Gerais: Parque Nacional Sertão Vereda, Rodrigues da Silva, Filguieras, & Oliveira 291 (DAV). Paraá: Alto Tapojós, Missño Velha, Egler 996 (MG). Rondânia: 2–4 km E of Abuna, Prance et al. 8584 (US). Roraima: Serra Tepequem, Boca da Mata, Prance et al. 4256 (MG).
This species has long been misunderstood and confused with other species, such as Phyllanthus lindbergii and P. stipulatus. Muller (1873) cited no specimens from Brazil except for Riedel 2810, on which he based his variety radicans (which may be a form of P. leptophyllus), and he did not describe the staminate flowers.
3.6. Phyllanthus leptophyllus Müll. Arg., DC. Prodr. 15(2): 411. 1866; Fl. Brasil. 11(2): 57. 1873. TYPE: BRAZIL. Minas Gerais: Serra da Lapa, I 1825, L. Riedel 1008 (Holotype: B, destroyed; photograph, F!).
? Phyllanthus microphyllus var. radicans Müll. Arg., Fl. Brasil. 11(2): 54. 1873. Type: BRAZIL. Minas Gerais, L. Riedel 2810 (not seen).
Distribution: known with certainty only from the type, a pistillate specimen from Minas Gerais. Müller (1873) distinguished it from Phyllanthus microphyllus by unconvincing vegetative characters and by the capitate style branch tips. However, since the seeds of P. leptophyllus, according to Müller's description, are puncticulate, it is possible that it belongs to subsect. Clausseniani.
3.7. Phyllanthus minutulus Müll. Arg., Fl. Brasil. 11(2): 54. 1873; Cordeiro, Bol. Bot. Univ. São Paulo 13: 180, figs. 42–47. 1992; Webster, Fl. Venez. Guayana 5: 198, fig. 185. 1999. Type: BRAZIL. Goiás: Porto Imperial, 1828/29, W. J. Burchell 8486, 8533; Minas Gerais, G. H. von Langsdorff (Syntypes, G? , not seen). Despite the fact that I have been unable to locate the types, Müller's description seems so unambiguous that there can be little doubt about the typification of the name.
Phyllanthus hoffmannseggi var. oblongifolius Müll. Arg., Linnaea 32: 45. 1863. P. diffusus var. oblongifolius (Müll. Arg.) Müll. Arg., DC. Prodr. 15(2): 410. 1866; Fl. Brasil. 11(2): 55. 1873. Type: BRAZIL. Para: Santarem, 1850, R. Spruce 797 (Lectotype [designated here]: W!).
Distribution and Habitat: Colombia, southern Venezuela, and Brazil, in wet swamps and savannas (pantanais), 50–1100 m.
Representative Specimens: Distrito Federal: Bacio do Rio São Bartolomeu, Heringer et al 6075 (US). Goías: Mun. Alto Paraíso, Chapada dos Veadeiros, Irwin et al 24726 (DAV); Mun. Caldas Novas, Serra da Arnica, Hatschbach & Ramamoorthy 38143 (DAV, MBM); Mun. Cristalina, estrada para Cachoeira do Arrojado, Hatschbach & Cordeiro 51819 (DAV); Mun. Gurupi, Eiten & Eiten 10008 (DAV). Maranhão: Mun. Loreto, Serra do Penitente, G. & L. T. Eiten 4210 (DAV). Minas Gerais: Mun. Diamantina, 9–13 km NW of Diamantina, Arbo et al 5067 (CTES); Mun. Santana do Riacho, Henrique et al CFSC 6893 (UEC). Pará: Mun. Conceiçã do Araguaia, 20 km W of Redenção, Plowman et al 8715 (DAV); Marabá, Serra dos Carajás, da Silva et al 1807 (DAV). Pernambuco: Goiana, Mello 4144 (US). Rio de Janeiro: Morro do Cruzeiro, Damazio 1333 (RB; locality somewhat uncertain). Rondonia: Mineração Campo Novo, 100 km SW of Ariquemes, Zarucchi et al 2724 (DAV); 2–4 km E of Abuna, Prance et al 8584 (DAV). Roraima: between Surumu and União, Ule 8199 (US). Tocantins: Ilha do Bananal, Aparecida da Silva et al 4031 (DAV).
Phyllanthus minutulus appears to be one of the commonest and most widespread Brazilian species of section Phyllanthus, but it has often been overlooked or misdetermined because of its small size and superficial resemblance to small plants of P. stipulatus or P. microphyllus. Jablonski (1967) misidentified the Spruce collection from Santarem as P. microphyllus.
3.8. Phyllanthus amarus Schumach. & Thonn., Beskr. Guin. Pl. 421. 1827; Danske Vidensk. Selsk. Skr. 4: 195. 1829; Webster, J. Arnold Arbor. 38: 313 pi. IX, figs. I-K. 1957, Ann. Missouri Bot. Gard. 54: 226. 1968; Burger & Huft, Fieldiana, Bot. n. s. 36: 143. 1995. Type: GUINEA. H. C. F. Schumacher & P.Thonning s.n. (Holotype: C!). Synonyms are cited in Webster (1957).
Distribution and Habitat: Apparently native to the Americas, but now pantropical, it is the weediest and most widespread species in the genus; in Brazil common in ruderal habitats, 0–1000 m. In publications before 1950, this species was usually called Phyllanthus niruri L., a misidentification made by Muller (1873).
Representative Specimens: Amapá: Macapá, Rabelo, Silveira, & Penafort 3250 (DAV). Amazonas: São Pelipe [Eirunepé], Schultes & López 9199 (US). Bahia: CEPEC station, between Ilhéus and Itabuna, Paixao 137 (NY). Ceará: Serra de Baturité, Eugenio 760 (RB). Distrito Federal: 12 km S of Planaltina, Irwin et al. 26458 (US). Goiás: Mun. Porangatu, G. & L. T. Eiten 9991 (DAV, SP, US). Maranhão: Maranhão, Ozimo da Carvalho (RB 105963, SP). Mato Grosso: Mun. Corumbá, Urucum, Pereira et al. 429 (RB). Minas Gerais: Corinto, Mexia 5662 (US, WIS). Pará: Belém, Cavalcante 2705 (MG). Paraíba: Alagoinha, Deslandes 6061 (SP). Pernambuco: Olinda, Bento 30 (RB). Rio de Janeiro: Quinta da Boa Vista, Kuhlmann (R 25998, US). Roraima: Serra Tepequém, Prance et al. 4256 (US). Santa Catarina: Brusque, Limeira, Reitz & Klein 11232 (US). Sergipe: Aracaju, Vieira & Skorupa 1202 (DAV).
Subsection 4. Clausseniani G. L. Webster, subsect. nov. Type: Phyllanthus claussenii Baill.
Differt ab aliis subsectionibus sectionis Phyllantho filamentis liberis, connectivo antherae dilatato, axes ramulorum saepe fractiflexis.
Monoecious or dioecious perennial Herbs or subshrubs; Branches and leaves smooth or papillose-scabridulous; leaves on main axis reduced to cataphylls (in the majority of species); Branchlets usually pinnatiform (sometimes branched in P. subemarginatus); Flowers pedicellate, in axillary clusters; sepals 6; Staminate Flowers with 3 stamens, filaments free (connate in P. allemii and P. fastigiatus); anthers deeply emarginate or the thecae distinct and stipitate; Pollen grains 4-colporate, subspheroidal, exine reticulate; Pistillate flowers with patelliform disk; ovary smooth or papillose; styles free, bifid, branches not capitate; Seeds puncticulate or striate.
As here defined, subsection Clausseniani clearly belongs in section Phyllanthus as indicated by its suffruticose habit, androecium of 3 stamens, and non-prolate pollen grains with relatively coarse reticulum. The flowers and pollen are suggestively similar to those of species in the “leafless” broomlike species of sect. Choretropsis. Although it is included here because of its characteristic androecium and pollen, Phyllanthus subemarginatus and related species are aberrant vegetatively, displaying little or no reduction in leaf lamina on the primary axes and generally ill-defined branchlets. Phyllanthus heteradenius and P. atalaiensis are also aberrant in appearance, resembling species of sect. Loxopodium (in subgenus Isocladus).
This subsection of 17 species appears to be entirely confined to Brazil, where it occurs in a variety of habitats: restingas, coastal rain forest, seasonal forest, montane forests, and subalpine campos rupestres. The subsection is similar in many respects to the West Indian subsection Pentaphylli, but differs in its cataphyllary stipules (which are not cordate and are blackened only in Phyllanthus fastigiatus); furthermore, species of subsection Clausseniani differ from the West Indian taxa in having puncticulate seeds, usually free filaments, and (in some species) non-phyllanthoid branching.
Key to the Species of Subsection Clausseniani
1. Distal leaf blades on penultimate axes reduced to cataphylls; branchlets smooth or papillose.
2. Branchlet axes smooth; anthers deeply emarginate (except in P. mocotensis) or thecae separate and stipitate.
3. Leaf blades mostly 3–6 cm long; stipules 4–5 mm long; fruiting pedicels 15–20 mm long, seeds 1.5–1.8 mm long 1. Phyllanthus glaziovii
3. Leaf blades, stipules, and fruiting pedicels, and seeds shorter.
4. Leaf blades obtuse or rounded at tip, abaxially lacking a whitish or yellowish coat of minute papillae.
5. Styles free (except sometimes at base), < 1 mm long; leaf blades symmetrical or asymmetrical at base.; fruiting pedicels 2–25 mm long; fruiting sepals 0.8–2.5 mm long.
6. Leaf blades symmetrical at base, not falcate, obtuse to rounded at apex; fruiting pedicels 2–15 mm long.
7. Leaf blades mostly 1 cm long or more, oblong to ovate or orbicular; branchlets not winged; anthers deeply emarginate or thecae separate and stipitate.
8. Staminate calyx purplish-tinged; anthers deeply emarginate but thecae not stipitate; fruiting sepals ca. 2 mm long 2. P. blanchetianus.
8. Staminate calyx not purplish; anther thecae distinctly stipitate; fruiting sepals 0.8–1.5 mm long.
9. Monoecious; sepals mostly 6; fruiting pedicels 6–18 mm long, sepals 0.8–1 mm long; seeds 1.3–1.5 mm long 3. P. claussenii
9. Dioecious; sepals mostly 5; fruiting pedicels 2–5 mm long, sepals 1–1.5mm long; seeds 1–1.2 mm long 15. P. subemarginatus
7. Leaf blades < 1 cm long, oblong; branchlet axis winged; anthers slightly emar ginate 4. P. mocotensis
6. Leaf blades falcate, asymmetrical at base, rounded to truncate at apex; fruiting pedicels 13–27 mm long 5. P. carvalhoi
5. Styles connate into a column ca. 1 mm high; leaf blades mostly asymmetrical at base; fruiting pedicels 3–5 mm long, fruiting sepals 2.5–3 mm long 6. P. sincorensis
4. Leaf blades acute at tip, abaxially with whitish or yellowish coat of minute papillae; fruiting pedicels 9–12 mm long, sepals ca. 2 mm long 7. P. hypoleucus
2. Branchlet axes papillose or scabridulous; anthers lightly to deeply emarginate.
10. Branchlet axes ± terete, at least not sharply winged.
11. Leaf blades ovate, rigid, acute, papillose or hirtellous abaxially; ovary papillose; stamens usually 2 8. P. arenicola
11. Leaf blades oblong, thin, obtuse, falcate, smooth or indistinctly papillose abaxially; ovary smooth; stamens 3 9. P. itatiaiensis
10. Branchlet axes sharply angled or winged.
12. Leaf blades mostly ovate, acuminate, 10–20 mm long; monoecious; fruiting pedicels 6–11 mm long 10. P. acutifolius
12. Leaf blades broadly elliptic to suborbicular, obtuse or rounded at tip, 3–5 mm long; dioecious or subdioecious; fruiting pedicels 1.5–2 mm long 11. P. caparaoensis
1. Distal leaf blades on penultimate axes not reduced to cataphylls; branchlets smooth.
13. Stems, leaves, and flowers copiously reddish-hirsutulous; stamens 2; ovary hispidulous 12. P. piranii
13. Stems, leaves, and flowers glabrous; stamens 2 or 3; ovary glabrous.
14. Stamens 3, filaments free; anthers emarginate.
15. Monoecious; branchlets not ramified; fruiting pedicel 1–2.5 mm long.
16. Staminate disk-segments falcate-acuminate; bracts of staminate cymules neither glandular nor lobed/dissected; staminate pedicels 0.1–0.5 mm long; seeds 1.1–1.4 mm long, puncticulate-striate 13. P. heteradenius
16. Staminate disk-segments obtruncate; bracts of staminate cymules glandular-lobed or-dissected; staminate pedicels 2–3 mm long; seeds 1.7–1.8 mm long, finely striate 14. P. atalaiensis
15. Dioecious; branchlets often ramified; fruiting pedicel 2–5 mm long; staminate disk segments rounded 15. P. subemarginatus
14. Stamens 2 or 3, filaments united in a column; anthers not clearly emarginate.
17. Stamens 3; branchlets conspicuously papillate-scabridulous; leaves pointed, not rigid; fruiting pedicel not clavate; seeds puncticulate, without coarse cross-striae 16. P. fastigiatus
17. Stamens 2; branchlets smooth; leaves apically truncate to emarginate, rigid; fruiting pedicel clavate; seeds puncticulate-striate with coarse cross-striae 17. P. allemii
4.1. Phyllanthus glaziovii Müll. Arg., Fl. Brasil. 11(2): 41, pi. 8. 1873. Type: BRAZIL. Rio de Janeiro: A.-F.-M. Glaziou 2892 (holotype: G!).
Distribution and Habitat: Parana to Rio de Janeiro and southern Minas Gerais, in evergreen montane forest (mata atlantica or matinha nebular), 50–1100 m.
Representative Specimens: Minas Gerais: Serra de Ibitipoca, Kreiger & Urbano 9315 (SPF). Rio de Janeiro: Mun. Nova Friburgo, Serra dos Orgãos, Castelo Diagua, Pereira 219 (RB); Santa Magdalena, Rio Muribecca, 1100 m, Santos Lima & Brade 13256 (RB); Teresópolis, Barria 98, Brade 9278 (R); Itatiaia, Maromba, Gottsberger 110–15471 (DAV, MBM), Pereira 6997 (US). Paraná: Serra do Mar, San Juan, Dusén 3570 (MO); Mun. Antonina, Rio Cotia, Hatschbach, Lindeman & Haas 13597 (US), Hatschbach 1451 (DAV, MBM); Mun. Morretes, Véu Noiva, Silva & Cordeiro 187 (DAV); Serra Marambi, Hatschbach 25356, 44589 (MBM). São Paulo: Paranapiacaba, Kuhlmann 3195 (SP).
Phyllanthus glaziovii is notable for its large acuminate leaves and large seeds, having the aspect of species of subgenus Xylophylla. However, the colporate pollen grains and deeply emarginate anthers suggest that its closest affinity is in subsect. Clausseniani.
4.2. Phyllanthus blanchetianus Müll. Arg., Linnaea 32: 38. 1863; Fl. Brasil. 11(2): 45. 1873.
Type: BRAZIL. “parte meridionale provinciae Bahia,” J. S. Blanchet 3158A (holotype: G!; isotype: P!).
Distribution and Habitat: Endemic to Bahia; not recollected as far as can be determined. A poorly known species, possibly not belonging to this subsection.
4.3. Phyllanthus claussenii Müll. Arg., Linnaea 32: 40. 1863; DC. Prodr. 15(2): 401. 1866; Fl. Brasil. 11(2): 61, pi. 9 fig, I. 1873. Type: BRAZIL. Minas Gerais: P. Claussen 1530 (Lectotype [designated here]: W!). Müller cited no collection numbers in 1863 and 1866, but cited three Claussen collections in 1873; the specimen at Vienna was the only one that could be located.
Phyllanthus claussenii var. oblongifolius Müll. Arg., Fl. Brasil. 11(2): 61. 1873. Type: BRAZIL, Minas Gerais, Riacho Fundo, 1834, L. Riedel 1364 (K!; perhaps the holotype).
Distribution and Habitat: Common in thickets and woodland (sertão, cerradão), Ceará to Pernambuco, Minas Gerais, and south to Paraná, 500–1100 m.
Representative Specimens: Ceará: Mun. Novo Oriente: Ibiapaba, Araújo 322 (PEUFR); Sertão da Caridade, Ducke 2110 (MG). Espiríto Santo: Goitacases, Rio Doce, Kuhlmann 6606 (RB). Minas Gerais: Mun. Santana do Riacho, Cerra do Cipó, Cordeiro et al. CFSC 8228 (UEC), 43546 (SPF); Serra da Espinaço, Pico do També, Anderson, Stieber, & Kirkbride 35931 (DAV, NY); Viçosa, Mexia 5359 (MO, US). Paraná: Mun. Senges, Rio Itararé, Hatschbach 26741 (DAV). Pernambuco: Arcoverde, Chiopetto 938 (SP); Serra das Varas, Lima 56–2560 (PEUFR). São Paulo: Itaporanga, Pickel 360 (US).
Specimens of Phyllanthus claussenii have often been confused with P. subemarginatus, but P. claussenii is usually monoecious, with 6-merous rather than 5-merous perianth, and larger seeds (1.3–1.4 mm long). Furthermore, in P. claussenii the distal leaves on penultimate axes are reduced to cataphylls, and the branchlets are normally unramified. However, fragmentary specimens without fruiting pedicels can be difficult to distinguish.
4.4. Phyllanthus mocotensis G. L. Webster, sp. nov.
Type: BRAZIL. Rio do Janeiro: Mun. Santa Maria Madalena, Alto Mocotó, X 1933, J. Santos Lima 194 (Holotype: RB 26003!). (Fig. 3A)
A P. itatiaiensi differt ramulis laevibus ancipitibus, foliis non falcatis, pedicellis longioribus; ab P. fastigiato differt staminibus liberis, ramulis laevibus.
Glabrous monoecious Herb with single main stem; Branchlets terete, smooth, decurrent-winged on one side, 5–9 cm long, 0.5–0.6 mm thick, with 35–45 leaves. Leaf BLADES oblong-obovate, obtuse-apiculate at the tip, cuneate at base, 4–6 mm long, 2–4 mm broad; petioles < 1 mm long; stipules subulate-acuminate, truncate at base, c. 1.5–2 mm long. Staminate flowers 1–3 in proximal glomerules: sepals 5, ± orbicular, 1–1.2 mm long; stamens 3, filaments free, 0.6–0.8 mm long; anthers slightly emarginate, dehiscing horizontally, c. 0.3 mm broad. Pistillate Flowers solitary at distal nodes of branchlet, pedicel 5–6 mm long in fruit; fruiting sepals 5, obovate, 1.21.8 mm long; ovary 3-locular, styles bifid, c. 0.5 mm long, branch tips subcapitate. Fruits and seeds unknown.
Known only from the type collection made by Santos Lima. The habit of Phyllanthus mocotensis is very similar to that of P. fastigiatus and P. itatiaiensis, but it appears morphologically distinct from the former in its typical phyllanthoid branching and the from latter in its non-falcate leaves.
4.5. Phyllanthus carvalhoi G. L. Webster, sp. nov.
Type: BRAZIL. Bahia: Mun. Itamarajú, Fazenda Pau-Brasil, mata higrófila sul Baiana, 160 m, 3 XI 1983, André M. de Carvalho, R. Callellas & L. A. Mottos Silva 2022 (Holotype: CEPEC!; isotypes: DAV!, NY!). (Fig. 2, 3B)
Ab aliis speciebus subsect. Clausseniani differt foliis grandis falcatis ad basin inaequaliteribus; a Phyllanthus sincorensis differt stylis liberis, pedicellis fructiferis longioribus.
Suffruticose monoecious perennial, with a single main stem; main axis densely scabridulous; Branchlets 7–12 cm long, 0.8–1 mm broad, with c. 20–30 leaves, axis terete, broadly winged, obscurely scabridulous in lines. Leaves sessile or subsessile; leaf blades obovate, falcate-asymmetrical 1.0–1.5 cm long, 0.5–1.0 cm broad, truncate to rounded at apex, strongly inaequilateral at base, adaxially minutely alveolate, abaxially purplish; lateral veins 3 or 4 pairs, arching, obscure adaxially but slightly raised abaxially; stipules lanceolate, 0.8–1 mm long, truncate at base. Staminate flowers on bracteate brachyblasts up to 3 mm long at proximal axils, pistillate flowers solitary at apical 2 or 3 nodes of branchlet. Staminate pedicel capillary, 5–8 mm long; sepals 5, free, obotate to suborbicular, entire, 1-veined, 1.5–2 mm long; disk segments 5, ± obcuneate, crenulate; stamens 3, filaments free, 0.8–1.2 mm long; anthers with enlarged deltoid connective but not emarginate, 0.5–0.6 mm broad, dehiscing horizontally. Pistillate Flowers solitary and axillary at 2–6 distal nodes of branchlet; pedicel 13–27 mm long; sepals 5, obovate, entire, midrib branching, carinate abaxially,2–2.5 mm long in fruit; disk annular, ca. 1.3 mm broad; ovary smooth; styles spreading, deeply bifid, ca. 0.7–0.8 mm long, branches recurving. Capsules not seen entire; valves ca. 2.5–3 mm long; columella ca. 1.5 mm long; seeds ca. 2 mm long, puncticulate-striate.
This highly distinctive species, named for Andre de Carvalho (student of the Bahian flora at CEPEC), is immediately separable from all other species of subsection Clausseniani by its winged branchlets with truncate, falcate, sessile leaves. Phyllanthus itatiaiensis, from Rio de Janeiro, has much smaller pointed, falcate leaves, but does not appear to be closely related.
4.6. Phyllanthus sincorensis G. L. Webster, sp. nov.
Type: BRAZIL. Bahia: Campos gerais, região Serra Sincorá, entre Brejão e Iracema, 18 II 1943, Ricardo Lemos Fróes 20172 (holotype: US!). (Fig. 3C)
Ab aliis speciebus subsect. Clausseniani differt stylis connatis columna sylorum 0.8–1 mm alto; ab P. carvalhoi differt foliis petiolatis non falcatis; ab P. claussenii differt foliis ad basin inaequalis.
Monoecious Shrub, glabrous; Branchlets terete, smooth, 10–12 cm long, 0.30.4 mm thick, with c. 25 leaves. Leaf BLADES ovate to oblong, 10–18 mm long, 5–11 mm broad, obtuse-apiculate at tip, obtuse to rounded or truncate and ± inaequilateral at base, veins prominent abaxially but not raised; petiole 1–1.5 mm long; stipules 1–2 mm long, acuminate. Flowers axillary, staminate in glomerules at proximal axils, pistillate solitary at distal axils. Staminate pedicels 3–7 mm long; sepals 5 or 6, oblong, c. 2 mm long and 1.2 mm broad; disk segments 5 or 6, angular, ca. 0.3 mm long, stamens 3, filaments free, 1–1.5 mm long; anthers c. 0.4 mm broad, with enlarged connective, thecae discrete, dehiscing ± horizontally. Pistillate pedicel 3–4 mm long in fruit; sepals 5, obovate, 1-veined, 2.3–3 mm long, 0.8–1.3 mm broad and reflexed in fruit; disk patelliform, ca. 0.8–1 mm broad; ovary 3-locular; ovary smooth; styles erect, 1–1.2 mm high, connate into a column 0.8–1 mm high, tips bifid. Seeds puncticulate, 1.4 mm long.
Known only from the type collection. In its basally oblique leaf blades, Phyllanthus sincorensis appears similar to species of subsection Niruri. However, the anthers with enlarged connective suggest placement in subsection Clausseniani. The prominent stylar column separates it from all the other Brazilian species of section Phyllanthus.
4.7. Phyllanthus hypoleucus Müll. Arg., Linnaea 32: 40. 1863. Phyllanthus lacteus Müll. Arg., DC. Prodr. 15(2): 402. 1866; Fl. Brasil. 11(2): 63. 1873 (nom. superfl.). Type: BRAZIL. Bahia: prope Nazare, F. Sellow (Holotype: G!; isotype: K!). Muller (1866) needlessly renamed his species because of Anisonema hypoleucus Miq. (1861), which he transferred to Phyllanthus as P. hypoleucus (Miq.) Müll. Arg. [= Glochidion lutescens Blume].
Distribution and Habitat: eastern Bahia and Espírito Santo, restinga and seasonal woodland (mata de cipó), near sea level to 1000 m.
Representative Specimens: Bahia: Mun. Maracás, 8–18 km SE Maracás, dos Santos, Mattos Silva, & Brito 3463 (CEPEC, DAV); Mun. Maraú, ca. 20 km S of Ponta do Mutá, Carvalho & Plowman 1407 (CEPEC, DAV). Espírito Santo: Mun. Serra, Estação Biológica do Mestre Álvaro, Mattos Silva et al. 167 (SP).
As noted below, Phyllanthus hypoleucus is very similar to P. acutifolius; however, it occurs at lower elevations in drier vegetation types.
4.8. Phyllanthus arenicola Casar., Nov. Stirp. Bras. Dec. 88. 1824; Müll. Arg., Fl. Brasil. 11(2): 64. 1873; Cordeiro, Bol. Bot. Univ. São Paulo 13: 182. 1992. Type: BRAZIL. Rio de Janeiro: Taipú, 1839/40, G. Casaretto 1793 (Holotype: G!).
Distribution and Habitat: Minas Gerais to Espirito Santo and Rio de Janeiro, restingas (lowland) and montane scrub, sea level to 1500 m. Cordeiro (1992) has noted that the plants of matas ciliares in Minas Gerais differ from the coastal populations in their pilose leaves; it is possible that they can be distinguished as an inland subspecies.
Representative Specimens: Espírito Santo: Vargem Alta, Fruteira, Pereira 2288 (HBR, US). Minas Gerais: Serra do Cipo, Palacio, 150 km N of Belo Horizonte, Irwin, Maxwell, & Wasshausen 20576 (NY); Mun. Santana do Riacho, Vale do Córrego Véu da Noiva, Cordeiro et al. 40735 (SPF), Rossi et al. 7005 (UEC), 36404 (SPF). Rio de Janeiro: Pedra Itauna, Maas & Carauta 3137 (UB); Recreio dos Bandeirantes, Hoehne 5705 (SP, UEC); Restinga de Itaipu, Schwacke 7107 (RB), Warming 1697 (C); Restinga de Maricá, Webster et al. 25446 (DAV, R); Restinga da Barra da Tijuca, Kuhlmann s.n. (RB 111186), Pereira & Lima 5752 (DAV, MBM, R).
4.9. Phyllanthus itatiaiensis Brade, Arch. Jard. Bot. Rio de Janeiro 15: 9, t. 7 figs. 12–16. 1957. Type: BRAZIL. Rio de Janeiro: Serra do Itatiaia, Pedra do Altar, 2400 m, III 1937, A. C. Brade 15575 (Holotype: RB 32920!).
Distribution and Habitat: Coastal range (Serra do Mantiqueira), Rio de Janeiro, subalpine scrub associated with Chusquea pinifolia, 2400–2600 m.
Additional Specimens Examined: Rio de Janeiro: Maçiço de Itatiaia, Brade 20362 (RB), Glaziou 8927 (US), Gottsberger 11015471 (UB), 134–17471 (DAV).
This diminutive species appears to be a narrow endemic to the Itatiaia massif. The falcate leaves distinguish it from those of most of the other species of subsect. Clausseniani, except for the much larger leaves of P. carvalhoi..
4.10. Phyllanthus acutifolius Poir. ex Spreng., Syst. Veg. 3: 21. 1821; Müll. Arg., Fl. Brasil. 11(2): 64. 1873. Type: BRAZIL. Herb. Poiret (Holotype: P!).
Phyllanthus oxyphyllus Müll. Arg., Linnaea 32: 40. 1863 (now. illeg.; non P. oxyphyllus Miq., 1861). TYPE: “Brasilia meridional” F. Sellow (Holotype: B; destroyed).
? Phyllanthus lagoensis Müll. Arg., Fl. Brasil. 11(2): 65. 1873. Type: BRAZIL. Minas Gerais: Lagoa Santa, 1863/66, J. E. B. Warming 1586 (Holotype: G!; ISOTYPE: C!).
Distribution and Habitat: Serra de Mantequeira, Rio de Janeiro and adjacent Minas Gerais and São Paulo, montane vegetation, 1100–1200 m. In Itatiaia, the species occurs at lower elevations than Phyllanthus itatiaiensis.
Representative Specimens. Minas Gerais: Baependi, São Tomé, Brade & Apparicio 20467 (RB). Rio de Janeiro: Parque Nacional Itatiaia, Altamiro & Walter 37 (MG), Brade 10322 (R), Sampaio 4687 (R), Ule 213 (R); Lago Azul, Pereira, Egler, & Maciel Barroso 69 (RB); Maromba, Pereira 6997 (US). São Paulo: São José do Barreiro, Hoehne & Gert 17645 (SP).
Phyllanthus acutifolius is distinctive within subsect. Clausseniani in its relatively large ovate leaves and roughened branchlets. It is extremely similar in many respects to P. hypoleucus of Bahia, which is clearly its allopatric sister species. Although Muller compared Phyllanthus lagoensis with P. claussenii and P. subemarginatus, it appears to be synonymous with P. acutifolius; the characters in Muller's key are not diagnostic.
4.11. Phyllanthus caparaoensis G. L. Webster, sp. nov.
Type. BRAZIL. Minas Gerais/Espírito Santo: Serra do Caparaó, 1879 (?), H. Wawra 1039 (Holotype: US 1234018!) (Fig. 3D)
Ab aliis speciebus subsect. Clausseniani differt plantis dioeciis, foliis parvulis rotundatis coriaceis; a P. arenicola differt foliis glabris, ramulis ancipitibus.
Suffruticose, dioecious or subdioecious, glabrous, 20–30 cm high; stems reddish-brown, finely channelled; Branchlet axes ± compressed, winged, scabridulous, 3–12 cm long, 0.3–0.5 mm thick, with 1025 leaves. Leaf blades broadly elliptic to suborbicular, rigid, 3–5 mm long, 0.25–5 mm broad; rounded or obtuse-apiculate at tip, rounded to truncate or subcordate at base, veins obscure on both faces; petiole 0.3–0.8 mm long; stipules acuminate, 0.7–1.5 mm long. Flowers axillary, the staminate in glomerules, the pistillate solitary. Staminate pedicel (1-) 1.5–2.7 mm long; sepals 5, rounded, 0.7–1.1 mm long; disk segments 5, elliptic, smooth, 0.2–0.3 mm in diameter; stamens 3; filaments free, 0.4–0.6 mm long; anthers deeply emarginate, 0.30.4 mm in diameter. Pistillate pedicel 1.5–2 mm long; sepals 5, 1–1.2 mm long, margins scarious; disk patelliform, 1–1.2 mm in diameter; ovary 6-sulcate; styles free, ascending, 0.4–0.5 mm long, bifid, branches not capitate. SEEDS puncticulate, 1.25 mm long.
Distribution and Habitat: Endemic to the Serra da Caparaó on the border of Minas Gerais and Espírito Santo, in subalpine vegetation, to 2400 m.
Additional Specimens Examined: Minas Gerais: Serra do Caparaó, Brade 16989 ex p. [mixed with P. subemarginatus] (DAV), Wawra s.n. (US); Campos do Caparaó, VII 1888, Schwacke (R), other than P. itatiaensis.
Phyllanthus caparaoensis is distinctive in its pale channelled stems, small rigid rounded leaves, and its occurrence on the slopes of Pico da Bandeira at the highest elevations of any Phyllanthus in Brazil.
4.12. Phyllanthus piranii G. L. Webster, sp. nov.
Type: BRAZIL. Espírito Santo: Mun. Conceitção de Castelo, km 116 da BR 262, 24 XI 1982, J. R. Pirani O. Yano,, D. F. Santos 230 (Holotype: SP 183289!). (Fig. 4, 5C)
Ab aliis speciebus subsectionis Clausseniani differt ramulis foliisque hirsutulis, stipulis aciculis longioribus, pedicellis fructiferis brevibus.
Herb, monoecious, 20–30 cm high; Stems persistent (lateral branches not deciduous), sparsely branching, terete, slightly zigzag, up to 0.5 m long, 0.8–1 mm in diameter; axes and leaves hirsutulous with reddish-brown multicellular trichomes 0.20.7 m long. Leaf blades broadly elliptic to orbicular, obtuse to rounded at apex, rounded at base, hirsutulous adaxially and abaxially, lateral veins 4–6 per side, arching, obscure adaxially, prominulous abaxially; petiole 1–1.5 mm long; stipules subulateacuminate, entire, hirsutulous, 2.5–3.5 mm long. Flowers axillary, solitary, the staminate and pistillate at separate axils. Staminate pedicel ca. 3 mm long; sepals 5, 1.2–1.5 mm long; disk segments 5, elliptic, ca. 0.25 mm in diameter; stamens 2; filaments free, ca. 0.5 mm long; anther thecae spheroidal, 0.2 mm across, dehiscing horizontally. Pistillate pedicel 1.5–1.7 mm long in fruit, hirtellous; sepals in fruit 1.8 2.2 mm long; disk annular, thin; ovary hispidulous with flattened trichomes ca. 0.1 mm long; styles 0.6 mm long, bifid, tips subcapitate. Seeds puncticulate in longitudinal rows, 1.2 mm long.
Known only from the type collection in southern Espírito Santo. Phyllanthus piranii is named in honor of the collector, J. R. Pirani (Herbário, Depto. de Botanica, Universidade de São Paulo), a student of Brazilian campos rupestres. The species is very isolated in subsect. Clausseniani by the dense pubescence and the completely nonphyllanthoid branching; its assignment to the subsection must be regarded as provisional.
4.13. Phyllanthus heteradenius Müll. Arg., Fl. Brasil. 11(2): 63. 1873. Type: BRAZIL: locality unknown, C. F. P. von Martius s. n. (holotype: M!).
Distribution and Habitat: northeastern Brazil, in caatinga and cerrado, 250500 m.
Representative Specimens: Bahia: Espigão Mestre, 5 km. S of Cocos, Anderson et al 37089A (DAV); Mun. Glória, 10 km N of Macuraré, G. & L. T. Eiten 4965 (DAV). Paraíba: regiões xerófitas, Coêlho de Moraes 2123 (US); Areia, near Alagoa de Remígio, Coêlho de Moraes 741 (MG, RB). Pernambuco: 15 km S of Afrânio, Heringer et al 333 (PEUFR).
Phyllanthus heteradenius is an unusual species, both for its habit which is very mimetic of the completely unrelated P. caroliniensis (sect. Loxopodium), and the remarkable falcate staminate disk segments. Its only close relative within subsection Clausseniani appears to be P. atalaiaensis, which has similar staminate disk segments.
4.14. Phyllanthus atalaiensis G. L.Webster, sp. nov.
Type: BRAZIL. Goiás: Mun. Monte Alegre de Goiás, Serra da Atalaia, 25 km by road SW of Monte Alegre, forest on granitic rock, 600–800 m, 12 III 1973, W. R. Anderson 6911 (Holotype: US 2869354!). (Fig. 5A, B)
A P. heteradenius differt bracteis, glanduloso-dissectis, pedicellis longioribus; ab aliis speciebus subsect. Clausseniani differt segmentis disci falcatis.
Annual herb, monoecious, glabrous, 3–4 dm high; leaves on penultimate axes not reduced to cataphylls; Branchlets terete, smooth, 4–7 cm long, 0.25–0.3 mm thick, with 7–10 leaves. Leaf Blades broadly obovate to suborbicular, 5–10 mm long, 5–8 mm broad, margins densely scabridulous, petiole 1–1.5 mm long; stipules 0.5–1 mm long, apically glandular-dissected. Flowers axillary, staminate in proximal glomerules, pistillate solitary and distal; bracts glandular-dissected as stipules. Staminate pedicel 2.5–3 mm long; sepals 5, broadly elliptic or suborbicular, 1-veined, 0.8–1.2 mm long; disk segments 5, massive, obtruncate, ca. 0.3 mm in diameter; filaments free, 0.3–0.4 mm long; anthers deeply emarginate. Pistillate pedicel ca. 2.5 mm long; sepals 5, obovate, pinnately veined, 2.7–3 mm long. Seeds 1.7 mm long, with 5 or 6 dark narrow longitudinal stripes, very finely cross-striate.
Known only from the type collection, from northeastern Goiás near the borders with Tocantins and Bahia. Although in appearance Phyllanthus atalaiensis resembles a form of P. heteradenius, it differs from that species in its unusual laciniate stipules and bracts, a feature unique in sect. Phyllanthus.
4.15. Phyllanthus subemarginatus Müll. Arg., Linnaea 32: 39. 1863; Fl. Brasil. 11(2): 66. 1873; Webster, Sellowia 11: 166, fig. 3 a-f. 1959; Smith, Downs, & Klein., Fl. Ilustr. Catar. EUFO 34, pi. 5 figs. a-f. 1988. Type: BRAZIL Rio de Janeiro: Vauthier 84 (Lectotype [designated here]: W!).
Phyllanthus amoenus Müll. Arg., Fl. Brasil. 11(2): 66. 1873. Type: BRAZIL. Minas Gerais: Serra da Piedade, J. E. B. Warming 1585 (Lectotype [designated here]: Cl).
Phyllanthus retroflexus Brade, Arch. Jard. Bot. Rio de Janeiro 15: 8, t 1 (figs. 9–11), 6. 1957. Type: BRAZIL. Espírito Santo: Mun. Cachoeiro do Tapemerim, Vargem Alto, Morro de Sal, 700 m, IV 1949, Brade 19768 (Holotype: RB 67063!).
? Phyllanthus janeirensis Müll. Arg., Fl. Brasil. 11(2): 45. 1873. TYPE: BRAZIL. Rio de Janeiro: A. F. M. Glaziou 1467 (Holotype: G, not seen). Müller (1873) placed this species far away from Phyllanthus subemarginatus, apparently because the specimen lacked staminate flowers and did not show the characteristic anthers.
Distribution and Habitat: widespread and common in eastern Brazilian rain forests (mata atlántica) from Minas Gerais and Espírito Santo south to Santa Catarina, 400–1000 m.
Representative Specimens: Bahia: Serra da Pioneira, Santa Terezinha, Noblick, Lemos, & Valdomiro 3211 (DAV). Ceará: between São José do Belmonte and Jati, Heringer et al. 738 (PEUFR). Espiríto Santo: Mun. Domingos Martins, Pedra Azul, Hatschbach 49959, Hatschbach & Guimarães 46893 (DAV, MBM). Minas Gerais: Belo Horizonte, Porto & Fagundes 2186 (RB); Serra do Caraça, 10 km W of Barão do Cocais, Irwin, Harley, & Onishi 28858 (DAV). Paraná: Mun. Quatro Barras, Rio Taquari, Hatschbach 19938 (DAV, MBM). Pernambuco: Serra dos Cavalos, Caruaru, Miranda & Araujo 364 (ESA). Rio de Janeiro: Serra da Carioca, Estrada do Sumaré, Pabst 5707 (M), 6779 (US); Mun. Teresópolis, Serra dos Orgãos, Rio Beija-Flor, Martinelli & Simonis 9056 (US). Santa Catarina: Palho9a, Morro do Cambirela, Klein & Bresolin 9406 (US). São Paulo: Ilha dos Alcatrazes, Luederwaldt & Fonseca 13816 (SP).
Phyllanthus subemarginatus is the most problematic species of subsect. Clausseniani It is the only widespread species in the subsection that has leaves generally not reduced to cataphylls on the penultimate axes; it is deviant also in the repeated branching of the branchlets. However, In some specimens (e.g., Martinelli & Simonis 9056 from Serra dos Orgãos) there is a progression of increasingly reduced leaf blades on the main axis, with the uppermost leaves reduced to cataphylls. This suggests that the rather labile branching pattern in P. subemarginatus may reflect a “degeneration” from species of subsection Clausseniani with typical phyllanthoid branching. Specimens with mostly unramified branchlets are superficially similar to P. claussenii and the two species are often confused in herbaria; however, P. subemarginatus can be distinguished by its dioecious flower production and shorter fruiting pedicels (2–5 mm long vs. 7–18 mm in P. claussenii).
The status of Phyllanthus amoenus Miill. Arg. appears questionable. Muller distinguished it from P. subemarginatus by having true phyllanthoid branching and larger differently veined leaves. However, it appeaars to fall within the overall range of variation of P. subemarginatus. Phyllanthus retroflexus Brade appears to represent an unusually diffusely branching form.
4.16. Phyllanthus fastigiatus Mart, ex Müll. Arg., Linnaea 32: 45. 1863; Fl. Brasil. 11(2): 57. 1873. TYPE: BRAZIL. Minas Gerais: Serra Itacolomi, Morro de Vila Rica, IV 1818, C. F. P. von Martius s.n. (lectotype [designated here]: Ml; holotype at B destroyed). The typification is somewhat problematic because Müller in 1863 ascribed the species to “Mart. mss. in herb. Berol.”, but cited only collections by Riedel and Sellow (presumably the reference to a Martius specimen was implicit, because an annotation label by Muller of the specimen at M indicates the original collection was made by Martius).
Distribution and Habitat: Scrub on rocky slopes, Minas Gerais, two disjunct localities in Serra da Moeda and Serra do Espinaço, near Ouro Preto, to 1750 m.
Specimens Examined: Minas Gerais: Mun. Itabirito, Monte Itabira, II 1835, Lund 1439 (G); Mun. Mariana, Serra do Itacolomi, Damazio (RB 18685), Damazio 1606 (RB 7870), Schwacke 14483 (RB), Ule 2686 (R).
Phyllanthus fastigiatus, which appears to be narrowly endemic to the region around Ouro Preto, is readily recognized among species of subsect. Clausseniani by its virgate habit (somewhat resembling Greater Antillean species of subsect. Pentaphylli) with narrow pointed leaves and by its muticous (non-emarginate) anthers.
4.17. Phyllanthus allemii G. L. Webster, sp. nov.
Type: BRAZIL. Goiás: Mun. Taguatinga, 7 km NE of Taguatinga towards Barreiras, cerrado, 850 m, 19 XI 1984, A. C. AUem, G. L. Webster, & W. E. Wemeck 3041 (Holotype: CEN!; ISOTYPE: DAV!). (Fig. 5D).
Ab aliis speciebus Brasiliensibus sectionis Phyllantho differt foliis coriaceis, pedicellis fructiferis clavatis, seminibus majoribus, testa longitudinaliter puncticulato-striata transversaliter hispidulo-strigosa.
Perennial herb, monoecious, glabrous, sparsely ramifying, 0.5–1 m high; leaves on penultimate axes not reduced to cataphylls; BRANCHLET axes terete smooth, 3–5 cm long, with 10–20 leaves. Leaf BLADES suborbicular, rigid and coriaceous, 3–5 mm long and broad, rounded to emarginate at tip, obtuse to truncate at base; petiole c. 0.5 mm long; stipules acuminate, 0.5–0.8 mm long. Flowers axillary, cymules unisexual, pistillate flowers solitary in distal axils, staminate flowers 1 or 2 at proximal axils. Staminate pedicel 1–1.2 mm long; sepals 5, obovate, 1.0–1.1 mm long, 0.5–0.7 mm broad; disk segments 5, elliptic, 0.25–0.3 mm in diameter; Stamens 2, filaments completely connate into a column ca. 0.6 mm high; anthers sessile on the column, dehiscing obliquely, 0.25–0.3 mm broad. Pistillate pedicel in fruit clavate, 3–5 mm long and 0.5–0.7 mm in diameter apically; sepals 5 or 6, elliptic, the outer 1.1–1.2 mm long, the inner 1.4–1.6 mm long; disk annular, pitted, ca. 1.3 mm in diameter; styles spreading, bifid, ca. 0.5 mm long. Capsule oblate, 4 mm in diameter; columella ca. 1.5 mm long; Seeds trigonous, brownish, 1.8–2 mm long, longitudinally finely puncticulate-striate, with coarse strigose cross-striae.
Known only from the type collection. Although its general habit suggests smallleaved specimens of P. subemarginatus, the clavate fruiting pedicels, stamen number, and large seeds with different ornamentation of P. allemii clearly set it apart. In its lack of reduction of the leaf blades on penultimate axes and its androecium of two connate stamens, P. allemii seems closest to P. fastigiatus. However, it is easily distinguished from that species by its smooth (non-scabrid) axes, thicker and blunter leaves, and much larger seeds. It is a pleasure to dedicate this distinctive species to Dr. Antonio Costa Allem, of EMBRAPA/ CENARGEN (Brasília), a leading expert on Euphorbiaceae of Brazil, who was both guide and colleague on the extended botanizing trip from Brasilia to remote stretches of Minas Gerais, Bahia, and Goiás during which this species was discovered.
This study has been made possible through the cooperation of many collaborators during my trips to Brazil, including Atonio Costa Allem (EMBRAPA/CENARGEN, Brasília); André M. de Carvalho (CEPEC, Itabuna); Inês Cordeiro (Instituto de Botânica, São Paulo); Margarete Emmerich and associates (Museu Nacional, Rio de Janeiro); Gert Hatschbach (Museu Botanico Municipal, Curitiba); Afrânio Gomes Fernandes and F. J. Abreu Matos (Universidade Federal do Ceará, Fortaleza); Fernando and Eleana Martins, and Yoshio Tamashiro (Departamento de Botânica, Universidade Estadual de Campinas); Arnildo and Vali Pott (Herbário EMBRAPA, Corumbá); Luciano P. de Queiroz (Universidade Federal de Feira de Santana); Maria J. N. Rodal and colleagues (Universidade Federal Rural de Pernambuco); Ricardo Secco (Museu Paraense Emílio Goeldi, Belém), and Marcus Sobral (Universidade Federal do Rio Grande do Sul, Porto Alegre). Logistic support was provided by grants from the National Science Foundation, the National Geographic Society, the Smithsonian Institution, and from the Academic Senate, University of California, Davis. Ellen Dean, Director of the Herbarium (DAV) at the University of California, Davis, and Jean Shepherd, the Curator, displayed infinite patience with the process of handling a large number of loans of specimens. Thanks are also due to the curators of the herbarium who loaned or made available specimens for study: A, B, C, CEN, CEPEC, CTES, DAV, EAC, ESA, F, G, HBR, IBGE, ICN, K, LL, M, MG, MBM, MO, NY, P, PEUFR, R, RB, S, SP, SPF, TEX, UB, UC, UEC, UEFS, ULM, US, W, and WIS.