Blumea 63, 2018: 167–198 www.ingentaconnect.com/content/nhn/blumea RESEARCH ARTICLE https://doi.org/10.3767/blumea.2018.63.02.14

Subgeneric delimitation of the plant genus Phyllanthus (Phyllanthaceae)

R.W. Bouman1,2,4, P.J.A. Keβler1,4, I.R.H. Telford 3, J.J. Bruhl 3, P.C. van Welzen2,4

Key words Abstract Over two centuries of taxonomic studies on the species rich genus Phyllanthus have culminated in a broad and complicated classification with many subgenera and (sub)sections. Past taxonomic work has only focused on infrageneric taxonomy local revisions, mostly because of the size of the genus. In this study we aim to summarize most of the taxonomic pantropical work in a list containing the infrageneric delimitations of Phyllanthus. This work will serve as a reference, placing paraphyletic most currently recognized species in subgenera and if possible, in sections for further study. Here we recognize Phyllanthaceae 880 species of Phyllanthus, classified in 18 subgenera, 70 sections and 14 subsections. A few taxonomic changes Phyllanthus are necessary to reconcile published phylogenetic data with the current classification. Subsections Callidisci and Odontadenii are raised to sectional rank, while section Eleutherogynium and section Physoglochidion are reduced to subsections and P. oxycarpus is transferred to the genus Glochidion. A provisional key for the subgeneric classification of Phyllanthus is provided.

Published on 21 November 2018

INTRODUCTION et al. 2009, Jangid & Gupta 2016, Wu et al. 2016) extended the infrageneric groupings to create a working classification for With almost 900 species, the mostly pantropical Phyllanthus L. most Phyllanthus species. is the largest genus in the family Phyllanthaceae (Govaerts et However, recent phylogenetic studies showed that several al. 2000). When considering all vegetative and reproductive subgenera were polyphyletic and even Phyllanthus itself proved organs, Phyllanthus is one of the most diverse groups in the to be paraphyletic (Kathriarachchi et al. 2006). In the follow- Angiosperms (Webster 1956). This diversity is exemplified by ing taxonomic revisions some of the polyphyletic subgenera the multitude of subgenera and (sub)sections defined within the were divided in new monophyletic subgenera (Ralimanana & genus. In the past, most of these subgenera and some sections Hoffmann 2011, 2014, Ralimanana et al. 2013), but discussion were treated at generic rank (De Jussieu 1824, Baillon 1858), remained whether Breynia J.R.Forst. & G.Forst., Glochidion and but were eventually all subsumed in a broad genus concept of Sauropus Blume should be subsumed into Phyllanthus. One Phyllanthus with numerous sections (Müller 1863, 1865, 1866). solution is to subsume these genera in Phyllanthus to create a The last major changes to this concept at genus level have been giant genus (Hoffmann et al. 2006, followed by Chakrabarty & the segregation of the genera Glochidion J.R.Forst. & G.Forst. Balakrishnan 2009, Wagner & Lorence 2011, Kurosawa 2016) (Kurz 1873) and Margaritaria L.f. (Webster 1957, 1979). The and the other is to split Phyllanthus into smaller, morphologically infrageneric structure of Phyllanthus was improved with the recognizable, monophyletic groups (Pruesapan et al. 2012, Van creation of several subgenera in a monographic work on the Welzen et al. 2014, Telford et al. 2016, followed by Chakrabarty Phyllanthus species of the West Indies by Webster (1956, 1957, & Balakrishnan 2012). A more exhaustive phylogenetic study 1958). Subsequent revisionary work followed Webster’s outline with higher sampling presented the case to maintain Breynia of subgenera and sections to illustrate the relations among (including Sauropus), Synostemon F.Muell. and Glochidion as groups within Phyllanthus (e.g., Bancilhon 1971, Webster & monophyletic and morphologically recognizable genera (Prue- Airy Shaw 1971, Punt 1972, Airy Shaw 1975, 1980a, Brunel sapan et al. 2008, 2012, Van Welzen et al. 2014), still leaving 1987, Rossignol et al. 1987, Santiago et al. 2006, Ralimanana the rest of Phyllanthus in its current state, a paraphyletic genus. & Hoffmann 2011, 2014, Ralimanana et al. 2013). Regional If Phyllanthus would be split, a larger phylogenetic study, which work on Phyllanthus (Merrill 1920, 1926, Pax & Hoffmann includes all subgenera and the majority of sections, is needed 1922, Beille 1925, 1927, Croizat 1942, 1943, Leandri 1958, Airy to prove which groups are monophyletic. Shaw 1963, 1969, 1972, 1975, 1976, 1980a, b, 1982, Webster 1986, Chantaranothai 2005, Silva & Sales 2006, 2008) and Phyllanthus is currently classified in about 18 subgenera with morphological studies (Punt 1967, 1972, 1980, 1986, Punt & numerous sections by past revision work. The most notable revi- Rentrop 1973, Lobreau-Callen et al. 1988, Stuppy 1995, Chen sions of Phyllanthus are those for the neotropics (Webster 2001b, 2002a, b, 2004), Asia (Airy Shaw 1960, 1975, 1977, 1980a, 1981, Webster & Airy Shaw 1971, McPherson & Schmid 1991) 1 Hortus botanicus Leiden, Leiden University, P.O. Box 9500, 2300 RA Leiden, and tropical Africa and Madagascar (Leandri 1958, Radcliffe- The Netherlands; corresponding author e-mail: [email protected]. Smith 1974, 1996, Brunel & Roux 1975, 1976, 1977, 1981, 2 Naturalis Biodiversity Center, P.O. Box 9517, 2300 RA Leiden, The Neth- 1984, 1985, Brunel 1987, Ralimanana & Hoffmann 2011, 2014, erlands. Ralimanana et al. 2013). There is some discussion regarding 3 Botany and N.C.W. Beadle Herbarium, School of Environmental and Rural the validity as publication of Brunel’s thesis (1987). The thesis Science, University of New England, Armidale, NSW 2351, Australia. 4 Institute of Biology Leiden, Leiden University, P.O. Box 9505, 2300 RA covers a large amount of work on the Phyllanthus species of Leiden, The Netherlands. Madagascar and Africa with many notes on subgenera and

© 2018 Naturalis Biodiversity Center You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 168 Blumea – Volume 63 / 2, 2018 sections. Because it is a thesis, this work was treated as not (1956). Unfortunately, the remaining 43 species could not be validly published based on article 32 of the International Code of assigned due to either incomplete descriptions, destroyed Botanical Nomenclature (McNeill et al. 2012) by Kathriarachchi type specimens, or lack of collections. We have opted to place et al. (2006). However, the thesis contains the name of a printing these species incertae sedis as their true relations need further company and numbered copies have been distributed to several detailed study. institutes, which is all in agreement with article 30.8, making it The classification of several subgenera from Webster’s origi- a validly published book. As such it is used in this publication. nal monographs (1956, 1957, 1958) has changed drastically. Several of the decisions in Brunel’s thesis were accepted in Subsequent palynological (e.g., Punt 1967, 1972, 1973, 1980, recent revisions of Phyllanthus in Madagascar (Ralimanana & 1986, Lobreau-Callen et al. 1988) and phylogenetic studies Hoffmann 2011, 2014, Ralimanana et al. 2013). (Kathriarachchi et al. 2006) have led to many new combinations The checklist by Govaerts et al. (2000) is often used to estimate and necessary transfers, some of which are discussed below. the number of species within Phyllanthus, but it does not con- Subgenus Isocladus was created to include about 60 species tain an infrageneric division. An attempted synopsis of all the with non-phyllanthoid branching (leaves on main stem not subgenera and sections was published by Kathriarachchi et al. reduced to scales and lateral axes not deciduous) and con- (2006). However, only the species included in the phylogenetic sisted of originally four sections, Loxopodium G.L.Webster, study were mentioned and a complete taxonomic treatment of Anisolobium Müll.Arg., Macraea (Wight) Baill. and Paraphyl­ the genus is still wanting. We hope that this list may serve as a lanthus Müll.Arg. (Webster 1956). However, subsequent studies framework for future studies. If Phyllanthus should ever be split (Brunel 1987, Webster 2002b) have reduced the size of this into various genera, this list can serve as a recommendation subgenus considerably. The sections Macraea and Cera­ for the species to include. manthus (Hassk.) Baill. (the latter with section Anisolobium merged with it; Punt 1972) were raised to subgeneric level by METHODS Brunel (1987). Section Loxopodium has been transferred to subgenus Phyllanthus on the basis of pollen characteristics In this study, we record 880 species, which are divided into and section Paraphyllanthus was placed in the synonymy of 18 subgenera, 70 sections and 14 subsections (Appendix 1). section Isocladus (Brunel 1987). Webster did create a new Govaerts et al. (2000) recorded 833 species and the difference section in subgenus Isocladus, Antipodanthus G.L.Webster, is mainly caused by the acceptance of Brunel (1987) and the which contained several neotropical and Australian species addition of newly published species after their work. Based on (Webster 2002a), but the Australian species appear to be better a combination of morphological descriptions, classifications in placed in section Lysiandra (F.Muell.) G.L.Webster of subgenus literature and published phylogenetic work (e.g., Samuel et al. Phyllanthus (Bouman, unpubl. data). Phylogenetic studies 2005, Kathriarachchi et al. 2006, Pruesapan et al. 2008, 2012, have confirmed the distinctness of subgenera Macraea and Manissorn et al. 2010, Challen et al. 2011, Luo et al. 2011), Ceramanthus from Loxopodium (Kathriararchchi et al. 2006). we propose the current list for the subgeneric classification of For section Antipodanthus, only one Australian species, P. caly­ Phyllanthus, in which we assign as many species as possible cinus Labill., and no neotropical species were included in the to subgenera and sections. Some placements are adopted phylogeny by Kathriararchchi et al. (2006), in which the group from and are now validly published from Webster’s unfinished appeared to be distinct from subgenus Isocladus. However, manuscripts, which are available online (http://herbarium.uc- to elucidate the relationship between sections Antipodanthus davis.edu/webster_manuscripts.html). For those species that and Lysiandra it is necessary to include more species in a were unplaced, we studied the distribution and morphological phylogenetic study. Therefore, section Antipodanthus is here descriptions (mainly branching type and the morphology of the maintained with no formal subgeneric placement. Ralimanana staminate flower), which allowed us to place them at least in & Hoffmann (2011) made the remainder of subgenus Isocladus subgenera. A synoptic key is provided by which most species (including former section Paraphyllanthus) monotypic, to only can be placed in the appropriate subgenera and/or sections. include P. maderaspatensis L., leaving some species unplaced However, sections and the species included have often not and in need of revision. been the subject of recent taxonomic revisions or are based All small shrubs and herbaceous Phyllanthus species were solely on palynological differences. This complicates the crea- originally placed in subgenus Phyllanthus. The subgenus tion of a key that can accommodate all species of Phyllanthus. was shown to be polyphyletic (Kathriarachchi et al. 2006) and The most important literature is cited after each species, which several subgenera are now recognized separately: subgenus either provides a direct placement or a morphological descrip- Swartziani (G.L.Webster) Ralim. & Petra Hoffm., containing tion. Hybrid species and infraspecific taxa were not included. the neotropical herbaceous species of subsection Swartziani; Some combinations, partly required by changes in level, are subgenus Afroswartziani Ralim. & Petra Hoffm., comprising published here, but only to solve nomenclatural anomalies the palaeotropical species of former subsection Swartziani (e.g., subsections that cannot be classified anymore in a sec- (largely comparable with section Anthophyllus Jean F.Brunel tion due to splitting of sections and changes in the taxonomic (Brunel 1987)), subgenus Tenellanthus Jean F.Brunel and sub- level of the taxa). genus Phyllanthus. Subgenus Phyllanthus now only contains sections Almadenses G.L.Webster, Choretropsis Müll.Arg., TAXONOMY LISTING OF PHYLLANTHUS Loxopodium G.L.Webster, Lysiandra, Phyllanthus and Salvini­ opsis Holm-Niels. ex Jean F.Brunel. Section Praephyllanthus We could assign 837 of the 880 species to a particular subge- Jean F.Brunel was found to be closely related to the species nus or (sub)section (Appendix 2), with some only listed as for- of subgenus Afroswartziani (Kathriarachchi et al. 2006) and is merly in subgenus Isocladus or the synonymized section transferred here to subgenus Afroswartziani. The type of sec- Para­phyllanthus Müll.Arg. One species of subgenus Isocladus tion Anthophyllus was placed in subgenus Swartziani, but all G.L.Webster, P. maderaspatensis L., was designated as the other palaeotropical species, including subsections Callidisci lectotype of the whole genus Phyllanthus by Ralimanana & Jean F.Brunel, Fluitantoides Jean F.Brunel and Odontadenii Hoffmann (2011). However, Phyllanthus niruri L. was already Jean F.Brunel & Jacq.Roux (here raised to section level) are designated as the lectotype of the genus Phyllanthus by better placed in subgenus Afroswartziani. These two subgenera Small (1913) and later independently confirmed by Webster are closely related (see Kathriarachchi et al. 2006) and mostly R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 169 distinguished by the inflorescences (unisexual in Afroswart­ Phyllanthus (Li 1987a), but is placed here in subgenus Eriococ­ ziani, bisexual in Swartziani) (Ralimanana et al. 2013). The cus (Hassk.) Croizat & Metcalf based on its pollen morphology species in sections Odontadenii, Fluitantoides and Callidisci (see Webster 1958, Brunel 1987, Webster & Carpenter 2008). have unisexual inflorescences and are tentatively placed in Species of subgenus Eriococcus occur in Asia and Australia subgenus Afroswartziani. and are characterized by the staminate flower with four sepals Subgenus Kirganelia (A.Juss.) Kurz is polyphyletic (Kathira­ and two or four stamens. Section Physoglochidion Müll.Arg. rachchi et al. 2006) and currently consists of eight sections: is placed here as a subsection under section Gomphidium Anisonema (A.Juss.) Griseb., Brazzeani Jean F.Brunel & Roux, Baill. based on the treatment of McPherson & Schmid (1991), Chorisandra (Wight) Müll.Arg., Cicca (L.) Müll.Arg., Hemicicca which is discussed below. Subgenus Gomphidium is a diverse (Baill.) Müll.Arg., Omphacodopsis Jean F.Brunel, Polyanthi group, with its main centres of diversity in New Guinea and Jean F.Brunel and Pseudomenarda Müll.Arg. As noted by New Caledonia. The monotypic section Hemicicca Baill. is Ralimanana & Hoffmann (2011), the type species for subgenus here placed in subgenus Kirganelia based on its similarity in Kirganelia is P. casticum P.Willemet, but P. reticulatus Poir. pollen (see Brunel 1987) and baccate fruits. The remaining is the type species for the type section Anisonema. Some previously un-assigned sections are here treated as synonyms: African and Madagascan species, originally attributed to this section Heteroglochidion Müll.Arg. is a synonym of subsection subgenus, were shown to be phylogenetically separate and Eleutherogynium (Müll.Arg.) G.L.Webster ex R.W.Bouman (see placed in subgenus Anesonemoides (Jean F.Brunel) Ralim. below for new combinations based on Webster 1986); sections & Petra Hoffm. (Ralimanana & Hoffmann 2014). Subgenus Meiandroglochidion S.Moore and Polyandroglochidion S.Moore Anesonemoides differs from subgenus Kirganelia in fruit mor- are synonyms of section Adenoglochidion (Müll.Arg.) Müll. phology (dehiscent in subgenus Anesonemoides vs baccate Arg. (McPherson & Schmid 1991); section Pentaglochidion in subgenus Kirganelia), a lack of brachyblasts in some spe- Müll.Arg.is a homeotypic synonym of section Leptonema Baill. cies of subgenus Anesonemoides, pollen with colpi bordered (see Baillon 1862, Müller 1863). The type species of section by parallel muri and the androecium (free or centrally fused Hedycarpidium Müll.Arg. has been re-identified as Baccaurea stamens in subgenus Anesonemoides vs two sets of stamens javanica (Blume) Müll.Arg. (see Müller 1866, Haegens 2000) (one fused, one free) in subgenus Kirganelia) (Ralimanana and even though the name is sometimes still used (Thin 2007), & Hoffmann 2014). Subgenus Kirganelia sections Cicca and it is invalid and the other species assigned to this section need Chorisandra were also shown to be in a clade separate from to be re-evaluated. A small number of Phyllanthus species section Anisonema (Kathriarachchi et al. 2006), but no nomen­- from Vietnam was placed in subgenus Eriococcus subsection clatural changes have yet been published. The sections Integra Thin (see Thin 2007), which is not included in our list. Omphacodopsis, Polyanthi and Brazzeani have not yet been No description was provided and it is quite possibly a synonym included in any phylogenetic study. Section Brazzeani was of subgenus Eriococcus subsection Spiciferens Jean F.Brunel originally placed in subgenus Conami (Aubl.) G.L.Webster as they include some of the same species, but we were not based on pollen characters (Brunel & Roux 1977), but these able to see the original publication. seem to have arisen through convergence and Brazzeani is Some nomenclatural issues are still present within Phyllanthus, better placed in subgenus Kirganelia (Meeuwis & Punt 1983). particularly when looking at the names of subdivisions of certain The stamen in the staminate flowers of section Brazzeanii are subgenera. Recommendation 22A of the International Code of arranged in two sets, similarly to subgenus Kirganelia section Nomenclature (McNeill et al. 2012) states that if there are no Anisonema. Though still used in Kathriarachchi et al. (2006) problems any subdivision of a subgenus that bears the type, and Ralimanana & Hoffmann (2011), section Floribundi Pax should be given the same epithet. However, in a few subgenera, & K.Hoffm. was reorganized by Brunel (1987) into two new this is currently not the case (Webster 1960). The type section sections Polyanthi and Omphacodopsis, while the type spe- of subgenus Conami is section Nothoclema G.L.Webster and cies of section Floribundi (P. muellerianus (Kuntze) Exell) was the type section of subgenus Kirganelia is Anisonema. In sub- transferred to section Anisonema, and the two sections were genus Kirganelia section Cicca, the type species, P. acidus (L.) combined. Sections Polyanthi and Omphacodopsis, though dis- Skeels, is in subsection Cheramela Kuntze (Webster 2001b). tinguished by pollen and fruit (in)dehiscence by Brunel (1987), can possibly be combined (see Breteler 2012). The staminate TAXONOMIC CHANGES flowers of these sections are similar to species in subgenus Anesonemoides, but the indehiscent fruit is more like subgenus Phyllanthus L. subgenus Afroswartziani Ralim. & Petra Hoffm. Kirganelia section Anisonema or Cicca. section Callidisci (Jean F.Brunel) R.W.Bouman, stat. nov.

Subgenus Emblica, sections Microglochidion (Müll.Arg.) Müll. Phyllanthus L. subgenus Afroswartziani Ralim. & Petra Hoffm. section Calli­ Arg., Pityrocladus G.L.Webster (subg. Emblica) and subgenus disci (Jean F.Brunel) R.W.Bouman. — Phyllanthus L. subsect. Callidisci Cyclanthera G.L.Webster were not yet included in any phylo- Jean F.Brunel, Gen. Phyllanthus Afr. Intertrop. Madag. (1987) 334. — Type: genetic research and their relationships within Phyllanthus are Phyllanthus callidiscus Jean F.Brunel. not well known. Webster chose to include section Microglo­ Note — Species of section Callidisci were originally placed chidion and Pityrocladus in the Asiatic subgenus Emblica on by Brunel (1987) in subgenus Phyllanthus section Anthophyllus account of their similarity in pollen (Webster 2002b, Webster & together with other palaeotropical subsections and recognized Carpenter 2002, 2008). A possible relationship between sub- by the fringed disc in the pistillate flowers. As this group has genus Cyclan­thera and subgenus Xylophylla was suggested recently been shown to be polyphyletic (Kathriarachchi et al. by Brunel (1987), but not incorporated in the latest revision by 2006) and after revision were split into a few new subgenera Webster (2002b). (Ralimanana et al. 2013), it seems necessary also to transfer Kathriarachchi et al. (2006) listed several sections as ‘not as- Brunel’s subsections. All other palaeotropical species of sub- signed to subgenus’, which are either already placed by other genus Phyllanthus were placed in subgenus Afroswartziani and authors, placed here, or treated as synonyms. Sections Bivia were distinguished from the neotropical subgenus Swartziani Jean F.Brunel & Jacq.Roux, Ceramanthus (Hassk.) Baill. and by their unisexual inflorescences (Ralimanana et al. 2013). This Cluytopsis Müll.Arg. are all placed in subgenus Ceramanthus is in agreement with species of subsection Callidisci, which is (Punt 1972, Brunel & Roux 1985, Brunel 1987). Section Nym­ transferred here and raised to sectional level to accommodate phanthus (Lour.) Müll.Arg. has often been treated in subgenus the separation from section Anthophyllus. 170 Blumea – Volume 63 / 2, 2018

Phyllanthus L. subgenus Afroswartziani Ralim. & Petra Physoglochidion to a subsection level and place it with section Hoffm. section Odontadenii (Jean F.Brunel & Jacq.Roux) Gomphidium. R.W.Bouman, stat. nov.

Phyllanthus L. subgenus Afroswartziani Ralim. & Petra Hoffm. section Transfer of Phyllanthus oxycarpus to Glochidion: Odontadenii (Jean F.Brunel & Jacq.Roux) R.W.Bouman. — Phyllanthus L. subsect. Odontadenii Jean F.Brunel & Jacq.Roux, Willdenowia 11 (1981) 70; Glochidion oxycarpum (Müll.Arg.) R.W.Bouman, comb. nov. Jean F.Brunel, Gen. Phyllanthus Afr. Intertrop. Madag. (1987) 339. — Type: Phyllanthus odontadensis Müll.Arg. Phyllanthus oxycarpus Müll.Arg., Prodr. 15, 2 (1866) 1270. — Diasperus oxycarpus (Müll.Arg.) Kuntze (1891) 600. — Type: Teijsmann s.n. (holo Note — Species in the palaeotropical section Odontadenii GDC), Indonesia, Sumatra. also have unisexual inflorescences and are therefore more Note — In his treatment of the genus Phyllanthus for De suited to be placed in subgenus Afroswartziani than the neo- Candolle, Müller (1866) reduced the genus Glochidion to a tropical subgenus Swartziani. The species are distinguished few sections within Phyllanthus. Phyllanthus oxycarpus Müll. form other sections by their winged plagiotropic branchlets Arg. was first described by Müller (1866) and placed in section (Brunel & Roux 1981). Euglochidion Müll.Arg. as it closely resembled P. subscandens (Zoll. & Moritzi) Müll.Arg. (a synonym of G. zeylanicum (Gaertn.) Phyllanthus L. subgenus Gomphidium (Baill.) G.L.Webster A.Juss.). Other species first published in section Euglochidion section Adenoglochidion (Müll.Arg.) Müll.Arg. subsection by Müller were all transferred to the genus by other authors Eleutherogynium (Müll.Arg.) G.L.Webster ex R.W.Bouman, (e.g., Boerlage 1900, Smith 1910), but we were unable to find a stat. nov. transfer for G. oxycarpum. The description lists no nectar disc, Phyllanthus L. subgenus Gomphidium (Baill.) G.L.Webster section Adeno­ a 5- or 6-locular ovary with columnar style, which are all typical glochidion (Müll.Arg.) Müll.Arg. subsection Eleutherogynium (Müll.Arg.) features for the genus Glochidion and therefore this species is G.L.Webster ex R.W.Bouman. — Phyllanthus L. sect. Eleutherogynium transferred here. Müll.Arg., Linnaea 32 (1863) 4, 14. — Type: Phyllanthus loranthoides Baill. Glochidion J.R.Forst. & G.Forst. sect. Chorizogynium Müll.Arg. (1863) 58, 59. — Lecto­type (designated by Webster 1986): Phyllanthus macrocho­ KEY TO THE SUBGENERA AND (SUB)SECTIONS OF rion Baill. PHYLLANTHUS Phyllanthus L. sect. Heteroglochidion Müll.Arg. (1866) 319. — Type: Phyl­ lanthus baladensis Baill. A provisional key is here provided based on characters men- Phyllanthus L. sect. Scleroglochidion Müll.Arg. (1866) 317. — Type: Phyl­ tioned in the literature. A key for full identification purposes, lanthus myrianthus Müll.Arg. using morphology only (not pollen) is difficult due to the absence of recent complete treatments for several groups and the fact Note — Section Scleroglochidion was previously placed in that some characters have evolved multiple times within Phyl­ synonymy by Webster (1986) who expanded the description lanthus. The key is not completely dichotomous (trichotomous of Eleuterhogynium to include also Phyllanthus species with 3 questions are marked with *). Authors of the various subgenera, free filaments. Section Heteroglochidion was defined by Mül- sections and subsections are listed in Appendix 1 and all spe- ler on its biseriate sepals, which is a common character for cies within a particular group are listed in Appendix 2. subgenus Gomphidium. All of these sections are character- ized by a rudimentary to absent nectar disc (see Müller 1866). 1. Branching non-phyllanthoid (laminate leaves and flowers on Lobreau-Callen et al. (1988) in a palynological study, showed all axes, branchlets not deciduous) ...... 2 that the pollen of these groups showed a continuous variation in 1. Branching phyllanthoid (leaves on main stem reduced to pollen characters and were difficult to differentiate. The lack of scales, cataphylls, laminate leaves and flowers on lateral distinguishing floral and vegetative characters and the overlap axes, lateral branchlets deciduous) or sub-phyllanthoid in palynological characters leads us to the decision to combine (leaves at base of branchlets not reduced to scales (often the above sections in one subsection Eleutherogynium, with in juveniles), lateral branchlets deciduous) . . . . . 15 as main character the absent nectar disc to distinguish it from other species within section Adenoglochidion. 2. Aquatic herbs. — Americas ...... Subgenus Phyllanthus section Salviniopsis 2. Herbs, shrubs or trees, but not aquatic ...... 3 Phyllanthus L. subgenus Gomphidium (Baill.) G.L.Webster section Gomphidium Baill. subsection Physoglochidion 3. Palm-like (monocaul) shrubs to trees; stigma petaloid. — (Müll.Arg.) R.W.Bouman, stat. nov. South America . . Subgenus Xylophylla section Asterandra 3. Herbs, shrubs to small trees, rarely climbers; stigmas vari- Phyllanthus L. subgenus Gomphidium (Baill.) G.L.Webster section Gomphi­ ously bifid to multifid, not petaloid ...... 4 dium Baill. subsection Physoglochidion (Müll.Arg.) R.W.Bouman. — Glo­ chidion J.R.Forst. & G.Forst. sect. Physoglochidion Müll.Arg., Linnaea 32 4. Leaves on all axes spirally arranged ...... 5 (1863) 58. — Phyllanthus L. sect. Physoglochidion (Müll.Arg.) Müll.Arg., 4. Leaves on all axes distichous ...... 9 Prodr. 15,2 (1866) 318. — Type: Phyllanthus faguetii Baill. 5. Sepals 4 in staminate flowers, 6 in pistillate flowers; stami- Phyllanthus L. sect. Phyllocalyx Baill. (1862) 236 (nom. illeg., non Phyllocalyx nate disc entire, H-shaped around filaments; stamens 2, A.Rich., 1847). — Glochidion J.R.Forst. & G.Forst. sect. Physoglochidion Müll.Arg. (1863) 58, 71. — Lectotype (designated here by R.W.Bouman, filaments free. — North America ...... but see Webster (2001) manuscript synopsis of Gomphidium): Phyllanthus ...... Subgenus Swartziani section Reverchonia faguetii Baill. 5. Sepals 5 or 6 in both sexes; staminate disc segmented; stamens 3 or 5, filaments free or connate ...... 6 Note — Phyllanthus section Physoglochidion (Müll.Arg.) Müll.Arg. is characterized by 3 free stamens, 6 sepals in two 6. Sepals 5; stamens 5, filaments free. — Africa . . . . . whorls and a calyx that becomes saccate in fruit. Apart from . . . . .Subgenus Kirganelia section Pseudomenarda the saccate calyx, these characters also occur in section 6. Sepals 5 or 6; stamens usually 3, filaments connate (free in Gomphidium and within section Physoglochidion and the P. rosmarinifolius Müll.Arg.) ...... 7 saccate calyx shows a continuous variation between species 7. Inflorescences axillary cymules with 1–4 flowers; pistillate (Lobreau-Callen et al. 1988). Since these groups can also not disc consisting of free glands. — Africa and Asia, introduced be distinguished on palynological data we opt to reduce section in North America ...... Subgenus Isocladus R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 171

7. Inflorescences axillary cymules or thyrses; pistillate disc 19. Anther connective often enlarged, thecae not appearing as entire ...... 8 stipitate; seeds scalariform with slight transverse striations 8. Inflorescences axillary glomerules; pollen 3- or 4-colpo- or smooth. — Australia and Central America (?) . . . rate, subglobose. — South America and Australia? ...... Subgenus Phyllanthus section Lysiandra ...... Section Antipodanthus incertae sedis 19. Anther connective variable, sometimes deeply emarginate 8. Inflorescences axillary glomerules or thyrses (sometimes with the two thecae appearing to be stipitate; seeds striate paniculate at end of branch); pollen areolate. — Central or linearly verrucate. — South America ...... and South America . . .Subgenus Xylophylla section . . . . . Subgenus Phyllanthus section Phyllanthus ...... Elutanthos ...... subsection Clausseniani 9. Staminate disc segmented, pistillate disc entire or seg­ 20. Herbs or subshrubs ...... 21 ment­ed; filaments free ...... 10 20. Shrubs to trees, rarely climbers ...... 36 9. Staminate disc segmented, or entire and urceolate, pistil- 21. Herbs; each branchlet bearing just one pair of (sub)oppo- late disc often massive and urceolate; filaments connate. site leaves and terminating in a raceme; anther connective Pollen with macroreticulate exine ...... enlarged. — South America ...... 12 (Subgenus Ceramanthus) . . . . . Subgenus Phyllanthus section Almadanses 10. Anthers dehiscing with horizontal slits; pollen 4-colpo- 21. Herbs or subshrubs; branchlet with more than 2 alternate rate. — Americas ...... leaves and flowers in leaf axils; anther connective (not) . . . . . Subgenus Phyllanthus section Loxopodium enlarged ...... 22 10. Anthers often deflexed, but dehiscing with vertical slits; pollen clypeate or perisyncolporate ...... 11 22. Flowers 5-merous; stamens 5, filaments free (except 3 stamens in P. cocumbiensis Jean F.Brunel). Pollen sub- 11. Leaves distichous; pollen clypeate; seeds verrucate. — globose, 3- or 4-colporate. — Pantropical, but origin Africa Africa, Asia, Australia and Pacific . Subgenus Macraea ...... 23 (Subgenus Tenellanthus) 11. Leaves spiral at basal nodes, distichous at upper nodes; 22. Flowers 5- or 6-merous; stamens 2 or 3, filaments free or pollen grains perisyncolporate with median pores, colpi connate ...... 24 bordered by parallel muri; seeds smooth or striate. — Madagascar ...... Subgenus Betsileani 23*. Shrubs or hemicryptophytes; stamens 5, filaments basally 12. Staminate flowers with sepals 4 (6 in pistillate ones); stami- united; pollen 3-colporate, with macroreticulate exine. nate disc consisting of 4 massive segments; stamens 2, Seeds with fine punctuation. — Africa ...... filaments connate and thecae on an enlarged connective...... Subgenus Tenellanthus section Loandani Pollen stephanoporate. — Africa ...... 23*. Herbs; stamens 5 (3 in P. cocumbiensis), filaments connate ...... Subgenus Ceramanthus section Bivia or free; pollen 3-colporate, with tectate, microperforate 12. Sepals 6 in both sexes; staminate disc entire or 6 seg- exine. — Africa ...... ments; stamens 3 with connate filaments, thecae not on . . . . . Subgenus Tenellanthus section Pentandra an enlarged connective ...... 13 23*. Herbs or subshrubs; stamens 5, filaments completely free; 13. Staminate disc entire and cup-shaped. — Asia . . . . pollen 4-colporate with sponge-like exine. — Pantropical, . . . . Subgenus Ceramanthus section Ceramanthus but origin Africa ...... 13. Staminate disc segmented or only slightly fused into a . . . . Subgenus Tenellanthus section Tenellanthus ring ...... 14 24. Inflorescences unisexual ...... 25 14. Sepals in two dimorphic whorls; staminate disc segmented; 24. Inflorescences bisexual ...... 35 pollen peribrevicolporate. — Africa and Asia . . . . . 25. Cataphyllary stipules (unilaterally) auriculate . . . 26 . . . . Subgenus Ceramanthus section Anisolobium 25. Cataphyllary stipules not auriculate ...... 32 14. Sepals in two equal whorls; staminate disc segmented to 26. Leaf base symmetric; plagiotropic branches carinate (wing- slightly fused into a ring; pollen pantoporate. — Asia . ed) (Brunel & Roux 1981). Pollen exine tectate; seeds with . . . . . Subgenus Ceramanthus section Cluytopsis longitudinal striae or smooth. — Africa ...... 15. Branching sub-phyllanthoid ...... 16 . . . . Subgenus Afroswartziani section Odontadenii 15. Branching phyllanthoid (sometimes sub-phyllanthoid in 26. Leaf base asymmetric; plagiotropic branches not cari- very young plants, check mature plants) . . . . . 20 nate ...... 27 16. Branchlets short, with only 5–10 leaves (Webster 2001b); 27. Pistillate disc entire with delicate fringes. — Africa . . staminate disc entire. — South and Central America ...... Subgenus Afroswartziani section Callidisci . . . . . Subgenus Xylophylla section Brachycladus 27. Pistillate disc entire, but not fringed ...... 28 16. Branchlet length variable, usually bearing more than 10 leaves; staminate disc segmented ...... 17 28. Sepals 5 in staminate flowers ...... 29 28. Sepals 6 in staminate flowers ...... 30 17. Anther connective not enlarged; fruit an indehiscent capsule; seeds smooth with fleshy sarcotesta. — South 29. Cataphyllary stipules usually black and indurate; stamen America and West Indies ...... 2 or 3, filaments partially or wholly connate, anthers ...... Subgenus Conami section Hylaeanthus sometimes deeply emarginate, dehiscing mostly horizon- 17. Anther connective variable, often enlarged; fruit a dehis­ tal; seeds longitudinally striate or banded, possibly with cent capsule; seeds ornamented, without a fleshy sarco­ transverse striae. Pollen 3-colporate. — West Indies . testa ...... 18 . . . . . Subgenus Phyllanthus section Phyllanthus 18. Filaments connate, stamens mostly 3 ...... subsection Pentaphyllus ...... 26 (subgenus Afroswartziani) 29. Cataphyllary stipules thin and membranous, not indu- 18. Filaments free (filaments connate in P. allemii G.L.Webster rate or black; stamen 3, filaments connate, anthers not and P. fastigiatus Mart ex Müll.Arg., but then only 2 sta- emarginate, dehiscing horizontally to vertically; seeds mens) ...... 19 longitudinally striate ...... 30 172 Blumea – Volume 63 / 2, 2018

30. Pistillate inflorescences on proximal position and stami- 38. Main axes often flat, branching monopodial, leaves disti­ nate inflorescences on distal position of plagiotropic chous; inflorescences usually bisexual, stamens 3 (rarely 4). branchlets; seeds transversely striate. Ovary often cov- — South America ...... Subgenus Phyllanthus ered with tubercles. — Pantropical, but origin Asia ...... section Choretropsis subsection Applanata ...... Subgenus Emblica section Urinaria 38. Main axes rounded, branching monopodial or sympodial, 30. Pistillate inflorescences on distal position and staminate leaves spiral; inflorescences mostly unisexual; stamens inflorescences on proximal position of plagiotropic branch- 2 or 3. — South America . . . Subgenus Phyllanthus lets; seeds longitudinally striate ...... section Choretropsis subsection Choretropsis ...... 31 (Subgenus Afroswartziani) 39. Fruits indehiscent, berries or drupes ...... 40 31*. Pollen 3- or 4-colporate, exine bireticulate. — Africa . 39. Fruits dehiscent, capsules (or absent) ...... 48 . . .Subgenus Afroswartziani section Praephyllanthus 40. Fruits drupaceous ...... 41 31*. Pollen 3-sulcate, exine macroreticulate. Often found in 40. Fruits baccate ...... 43 water. — Africa ...... Subgenus Afroswartziani section Fluitantoides 41. Sepals 6; stamens 3, filaments connate. — Asia . . . 31*. Pollen perihexabrevisulcate, exine macro-rugulose (Brunel ...... Subgenus Emblica section Emblica 1987). Ovary on gynophore. — Africa ...... 41. Sepals 4–6; stamens 3 or 4 (rarely 2 or 5), filaments free . . . Subgenus Afroswartziani section Microdendron ...... 42 (Subgenus Kirganelia section Cicca) 32. Branchlets and flowers not purplish; stamens 3, filaments 42. Plants dioecious; disc absent in both sexes; staminodes mostly free or united to 2/3 of length; pollen 4-colporate, absent; fruits spongy (Webster 1957). — West Indies and exine (hetero-)reticulate; pistillate sepals 5; pistillate disc South America) . . Subgenus Kirganelia section Cicca entire; stigmas free, bifid, tips sometimes subcapitate 33 ...... subsection Aporosella 32. Branchlets and flowers often purplish; stamens 2 or 3, 42. Plants monoecious; disc present in both sexes; sometimes filaments connate; pollen pantoporate, exine shields elon- staminodes present; fruits hard. — Commonly cultivated, gated or if round with only 1 pila; pistillate sepals 6; pistillate origin possibly African? . . . . .Subgenus Kirganelia disc dissected or lobed; stigmas free or connate, bifid to ...... section Cicca subsection Cheramella emarginate, tips not capitate 34 (Subgenus Cyclanthera) 43. Stamen 2, filaments connate; ovary 2-locular. — Africa, 33. Anther connective not enlarged, thecae not stipitate; Madagascar, Mainland Asia ...... style branches sub-capitate; seeds verrucate. — South ...... Subgenus Kirganelia section Chorisandra America, pantropically invasive Subgenus Phyllanthus 43. Stamen 3‒6, filaments free or connate; ovary 3-locular ...... section Phyllanthus subsection Niruri ...... 44 33. Anther connective variable, deeply emarginate with the 44. Branchlets subtended by reduced leaves, but not cata- two thecae appearing stipitate; style branches not capitate; phylls, flowers on brachyblasts; stamens 3, filaments free seeds striate or linearly verrucate. — South America . or connate. — South America ...... Subgenus Phyllanthus section Phyllanthus ...... Subgenus Conami section Hylaeanthus ...... subsection Clausseniani 44. Branchlets subtended by (spinescent) cataphylls, stamens 34. Branchlets unramified, rooting at nodes; leaves crisply suc- 4‒6, filaments free ...... 45 culent; stamens 2, filaments free; pollen shields elongated 45. Branchlets subtended by spinescent cataphylls; stamens (banded) (Webster & Carpenter 2002). — West Indies . 5 in 2 sets, one free and the other basally fused. — Africa . . . . Subgenus Cyclanthera section Callitrichoides and Asia . . .Subgenus Kirganelia section Anisonema 34. Branchlets often with 1 or 2 lateral branches (bipinnati- 45. Branchlets subtended by scale or stipule like cataphylls; form), not rooting at nodes; leaves not succulent; stamens stamens 4–6, filaments free ...... 46 3, filaments completely connate into a circular synandrium; 46. Staminate inflorescences on separate (leafless) plagio- pollen shields isodiametric, each with a central pila sur- tropic branches, pistillate flowers axillary. Seeds globular, rounded by a murus (Webster & Carpenter 2002). — West smooth. — Africa Subgenus Kirganelia section Polyanthi Indies . . .Subgenus Cyclanthera section Cyclanthera 46. Inflorescences axillary, on all plagiotropic branches . 47 35. Stamens 3, filaments free, anthers dehiscing horizontally; 47. Stamens 5. — Asia ...... pistillate disc dissected; pollen grains brevicolporate and ...... Subgenus Kirganelia section Hemicicca diorate or porate, exine pilate; seeds verruculose. — South 47. Stamens 6. — Africa, Madagascar, Mainland Asia . . America . . . . Subgenus Conami section Apolepsis ...... Subgenus Kirganelia section Chorisandra 35. Stamens 2 or 3, filaments entirely or partially connate (free 48. Anthers apiculate ...... 49 in P. warnockii G.L.Webster), anthers dehiscing oblique to 48. Anthers non-apiculate ...... 56 horizontally (vertically in P. warnockii); pistillate disc entire; pollen 3-colporate, exine reticulate; seeds smooth or longi­ 49. Sepals often caudate-acuminate; filaments connate, sta- tudinally striate. — North America, pantropical invasive minate disc consisting of linear spathulate segments; pistil- ...... Subgenus Swartziani late disc entire. Pollen 4-colporate, exine reticulate. — Asia ...... 50 (Subgenus Phyllanthodendron) 36. Leaves reduced and branchlets transformed to phyllo- 49. Sepals often acuminate, but not caudate; filaments free clades (at least in older branches) ...... 37 or connate, staminate disc segmented, globular; pistillate 36. Leaves not reduced and branchlets not transformed to disc entire (or absent) ...... 54 phylloclades ...... 39 50. Shoots not differentiated, all leaves similar in size, flowers 37. All stems rounded or flat; stipules unilaterally auriculate, sta- on lateral shoots ...... 51 mens free or connate; pollen 3- or 4-colporate, exine reti­ 50. Shoots differentiated into sterile leaf bearing shoots with culate 38 (Subgenus Phyllanthus section Choretropsis) larger leaves and fertile shoots with smaller leaves . 53 37. Lateral stems flattened with wide phylloclades, (bi-)pinnati­ form; stipules not auriculate; stamens usually united at base; 51. Sepals 4 in staminate flowers; stamens 4. — Asia . . pollen clypeate, exine areolate. — West Indies ...... Subgenus Phyllanthodendron section Tetrandrum ...... Subgenus Xylophylla section Xylophylla 51. Sepals 5 or 6 in staminate flowers; stamens 3 . . . 52 R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 173

52. Shrubs; sepals 5 in staminate flowers. Fruit reminiscent of 65. Flowers in all leaf axils. — Asia ...... Actephila (Croizat 1942). — Asia ...... Subgenus Eriococcus section Eriococcus .Subgenus Phyllanthodendron section Pseudoactephila 65. Pistillate flowers on leafy panicles at end of branchlets and 52. Twining shrubs; Sepals 6 sepals in staminate flowers. — staminate flowers closer to the base of branchlets without Asia Subgenus Phyllanthodendron section Arachnodes leaves (see Brunel 1987). — Asia ...... 53. Trunk often succulent and enlarged at base; leaf blades ...... Subgenus Eriococcus section Eriococcus > 6 cm long. — Asia ...... subsection Spiciferens Subgenus Phyllanthodendron section Phyllanthodendron 66. Branchlets (bi-)pinnatiform; sepals often biseriate; stami- 53. Trunk not succulent or enlarged at base; leaf blades < 6 cm nate nectar disc often 3 massive emarginate (or 6 sepa- long. — Asia ...... rate) segments to absent, stamens may be inserted on a . . Subgenus Phyllanthodendron section Calophyllum wide receptaculum. Pollen 3-(syn)colporate . . . . . 54. Filaments connate. — South America ...... 67 (Subgenus Gomphidium) ...... Subgenus Xylophylla section Ciccastrum 66. Branchlets pinnatiform; sepal whorls indistinct; staminate 54. Filaments free ...... 55 nectar disc entire or segmented ...... 79 55. Leaves with or without laminar glands; sepals in two in- 67. Branchlets bipinnatiform ...... 68 67. Branchlets pinnatiform ...... 70 distinct whorls; pollen 4–8-colporate or diorate. — South America . .Subgenus Emblica section Microglochidion 68. Axes incrustate or hirsutulous with red hairs; stamens 2‒6; 55. Leaves without laminar glands; sepals in two distinct whorls; pollen clypeate. — West Indies ...... pollen 3-syncolporate . . 67 (Subgenus Gomphidium) . . . . Subgenus Xylophylla section Hemiphyllanthus 68. Axes not incrustate or hirsutulous, hairs usually white; 56. Leaves opposite or subopposite ...... 57 sta­mens mainly 3‒5 (up to 20); pollen 3-(syn)colporate 56. Leaves alternate ...... 59 ...... 69 57. Bark lenticellate; filaments connate. — Cuba . . . . 69. Inflorescences glomerules; pollen diverse, often 3-colpo- . . Subgenus Xylophylla section Williamia subsection rate or porate with diorate colpi (see Webster & Carpenter ...... Mirifici 2002), exine vermiculate to pilate; fruit conspicuously veined. 57. Bark smooth; filaments free ...... 58 — South America ...... 58. Branchlets sometimes opposite bipinnatiform; staminate ...... Subgenus Conami section Nothoclema sepals 5, not distinctly biseriate; staminate disc consisting 69. Inflorescences glomerules or panicles; pollen 3- or 4-syn- of 5 free segments; stamens 5. — Madagascar and Middle colporate with vermiculate/rugulate exine (Lobreau-Callen East (?) ...... Subgenus Menarda et al. 1988); fruit smooth. — Southeast Asia, mostly New 58. Branchlets pinnatiform, not opposite; staminate sepals Guinea . . . Subgenus Gomphidium section Nymania 5 or 6, in both sexes often distinctly biseriate; staminate 70. Disc absent or rudimentary in both sexes . . . . . 71 disc entire, 3 emarginate segments or 5 or 6 massive 70. Disc entire or segmented in both sexes . . . . . 74 segments; stamens mostly 3 or 5 (up to 20) ...... 67 (Subgenus Gomphidium) 71. Sepals 6, biseriate, inner whorl petal-like, pistillate sepals leafy; stamens 3; ovary 3-locula. Calyx in fruit saccate. 59. Sepals 4 in staminate flowers; stamen 2, filaments con- — New Caledonia . . . . . Subgenus Gomphidium nate. Pollen pantoporate or clypeate ...... 60 . . .section Gomphidium subsection Physoglochidion 59. Sepals 5 or 6 in staminate flowers; stamens 3–15, fila­ 71. Sepals 5 or 6, not distinctly biseriate; stamens (3–)5(–15), ments free or connate ...... 66 ovary 3–5-locular ...... 72 60. Leaf margins very thick, conspicuously revolute; staminate 72. Sepals 5; stamens 5; ovary 4- or 5-locular. — New Cale­ disc massive, entire; pollen clypeate. — Cuba . . . . donia . . .Subgenus Gomphidium section Leptonema . . . . Subgenus Xylophylla section Glyptothamnus 72. Sepals 5 sometimes 6; stamens mostly (3–)5(–15); ovary 60. Leaf margins not thickened, sometimes slightly revolute; 3-locular ...... 73 staminate disc segmented; pollen pantoporate or clype­ ate ...... 61 73. Disc rudimentary in both sexes; filaments shorter than anthers, inserted on a wide receptaculum. — Southeast 61. Anthers dehiscing vertically; sepal margins entire . . . 62 Asia, New Caledonia ...... 61. Anthers dehiscing horizontally/transversely; sepal margins . . . Subgenus Gomphidium section Adenoglochidion entire to dentate or lacerate ...... 64 73. Disc rudimentary or absent in both sexes; filaments longer 62. Inflorescences usually bisexual, appearing with the ex- than anthers, diverging from centre of receptaculum. — panding leaves (Webster 1958); pollen clypeate; style New Caledonia ...... Subgenus Gomphidium connate in a tube and stigmas often reduced to acute tips. . section Adenoglochidion subsection Eleutherogynium — West Indies . . . . .Subgenus Xylophylla section 74. Stamens connate ...... 75 Thamnocharis 74. Stamens free ...... 76 62. Inflorescences mostly unisexual, appearing after the 75. Inflorescences glomerules; pollen diverse, often 3-colpo- leaves; pollen pantoporate; style connate or free . . 63 rate or porate with diorate colpi (see Webster & Carpenter 63. Ovary papillose or verruculose, 3-locular. — Asia . . . 2002), exine vermiculate to pilate; fruit conspicuously . . . . . Subgenus Eriococcus section Eriococcodes veined. — South America ...... 63. Ovary smooth, 6-locular. — Asia ...... Subgenus Conami section Nothoclema . . . . .Subgenus Eriococcus section Nymphanthus 75. Inflorescences glomerules or paniculate; pollen 3- or 4- 64*. Stigmas entire, connate — filaments thickened at top. — syncolporate, exine vermiculate/rugulate (Lobreau-Callen Asia to Australia ...... et al. 1988); fruit smooth. — Southeast Asia, mainly New . . . . .Subgenus Eriococcus section Emblicastrum Guinea . . . Subgenus Gomphidium section Nymania 64*. Stigmas entire or emarginate. — Asia ...... 76. Sepals 5, not distinctly biseriate; stamens mostly (3–) ...... Subgenus Eriococcus section Scepasma 5(‒15), filaments free; disc consisting of 3 emarginate seg- 64*. Stigmas free, bifid ...... ments or absent. — Southeast Asia, New Caledonia . . . . . 65 (Subgenus Eriococcus section Eriococcus) . . . Subgenus Gomphidium section Adenoglochidion 174 Blumea – Volume 63 / 2, 2018

76. Sepals 5 or 6, often biseriate (except in P. tuerckheimii 89. Stamens fused in several whorls or sets . . . . . 90 G.L.Webster); stamens 3, filaments free or connate; disc 89. Stamens fused in a central column ...... 93 consisting of 3 emarginate segments or 6 free segments 90. Brachyblasts present; stamens fused in two sets with one ...... 77 central column and two separate free stamens . . 91 77. Sepals 5; pollen grains not syncolpate, colpi without dis- 90. Brachyblasts absent; stamens in 2 or 3 whorls, fused in tinct borders; exine reticulate. — South America ...... various ways. — West Indies ...... Subgenus Gomphidium section Calodictyon . . . . . 94 (Subgenus Xylophylla section Willamia) 77. Sepals 6; pollen grains with marginate colpi, often meeting 91. Pollen 3-colporate, exine pilate or reticulate . . . 92 at poles; exine reticulate or ± vermiculate . . . . . 78 91. Pollen clypeate, exine areolate. — West Indies . . . 78*. Inflorescences axillary cymules; pollen 3-syncolporate with . . . . . 94 (Subgenus Xylophylla section Willamia) fine to course reticulate exine. — Southeast Asia, New 92. Exine pilate. — Africa ...... Caledonia Subgenus Gomphidium section Gomphidium ...... Subgenus Kirganelia section Brazzeani 78*. Inflorescences glomerules or panicles; pollen 3-syncolpo- 92. Exine reticulate. — Africa and Asia ...... rate with vermiculate/rugulate exine. — Southeast Asia, ...... Subgenus Kirganelia section Anisonema mostly New Guinea ...... Subgenus Gomphidium section Nymania 93. Stems and branchlets incrustate with dark platelets of 78*. Inflorescences glomerules; pollen diverse, often 3-col- bark or lenticellate ...... porate with diorate colpi (see Webster & Carpenter . . . . . 94 (Subgenus Xylophylla section Willamia) 2002), exine vermiculate to pilate. Fruit conspicuously 93. Stems smooth ...... 95 veined. — South America . .Subgenus Conami section 94*. Stems smooth; leaves alternate; stamens 3–15 in 3 whorls, ...... Nothoclema connate in various ways; stigmas erect but not lacerate. 79. Staminate disc entire. — South America ...... Sepals 5 or 6. — West Indies . . Subgenus Xylophylla . . . . . Subgenus Xylophylla section Adianthoides ...... section Williamia subsection Discolores 79. Staminate disc segmented ...... 80 94*. Stems and branchlets incrustate with small dark platelets on the fissured bark; leaves alternate; stamens (2–)3–6 with 80. Filaments free or only fused at base ...... 81 filaments connate, usually in 2 whorls; stigmas erect, con- 80. Filaments fused at least partially to completely, sometimes spicuously lacerate (see Webster 1958). — West Indies fused in separate sets or whorls ...... 89 ...... Subgenus Xylophylla section Williamia 81. Leaves often with glands; anthers apiculate. Leaves thick; ...... subsection Incrustati style entire. — South America ...... 94*. Stems smooth but prominently lenticellate; leaves op- . . . . . Subgenus Emblica section Microglochidion posite; stamens 5 with filaments connate, but 2 anthers 81. Leaves without glands; anthers not apiculate . . . 82 inserted lower than the other 3; stigmas reflexed and cov- 82. Stamens 3 ...... 83 ering the ovary, apex sometimes blunt. — West Indies 82. Stamens 4 or 5 ...... 85 ...... Subgenus Xylophylla section Williamia 83. Brachyblasts often present; inflorescences cauliflorous; ...... subsection Mirifici sepals 6. — South America ...... 95. Pollen 3–5-colporate or 5-brevicolporate (Webster & ...... Subgenus Kirganelia section Ciccopsis Carpenter 2008), exine reticulate, microperforate or 83. Brachyblasts absent; inflorescences axillary; sepals 5 84 scabrous. — Africa and Asia ...... 96 84. Leaf blades < 8 cm long; anther connective enlarged; pollen 95. Pollen clypeate, exine areolate. — Americas . . . . 4-colporate. — South America . . Subgenus Phyllanthus ...... 99 (Subgenus Xylophylla) . . . . .section Phyllanthus subsection Clausseniani 96. Anthers dehiscing obliquely to horizontally; exine micro- 84. Leaf blades > 8 cm long; anther connectives not enlarged; perforate or scabrous. — Pantropical, mostly African . pollen perisyncolporate. Pistillate pedicel quite massive ...... Subgenus Afroswartziani (up to 3 cm wide (Brunel 1987)), fruit ornamented. — Af- 96. Anthers dehiscing vertically; exine reticulate . . . . rica . . . .Subgenus Ceramanthus section Ebolowani ...... 97 (Subgenus Emblica) 85. Pistillate sepals 8–10. — South America ...... 97. Sepals mostly 5; staminate disc consisting of 5 segments; ...... Subgenus Xylophylla section Diplocicca stamens 2‒5(‒7); anthers dehiscing horizontally; pollen 85. Pistillate sepals 5 or 6 ...... 86 3‒5-colporate; pistillate disc entire or segmented. — 86. Brachyblasts present ...... 87 South America Subgenus Emblica section Pityrocladus 86. Brachyblasts absent ...... 88 97. Sepals 6; staminate disc consisting of 6 segments; stamens 3; anthers dehiscing vertically; pollen 4- or 87. Inflorescences (stalked) fascicles; stamens 5, filaments 5-colporate or 5-brevisulcate; pistillate disc entire . 98 completely free; fruits 3-locular, dehiscent; seeds kidney- shaped, smooth with mottled patterns (similar to seeds of 98. Inflorescences at terminal end of branchlet and secondary P. juglandifolius Willd.). — Africa ...... ramification on more basal end; pollen 5-brevisulcate. — . . . . Subgenus Kirganelia section Omphacodopsis Asia . . . . .Subgenus Emblica section Botryoides 87. Inflorescences panicles; stamens 4 or 5, filaments free or 98. Inflorescences found along entire branchlet; pollen 4- or sometimes basally fused; fruits 3–5-locular, indehiscent; 5-colporate. — Asia Subgenus Emblica section Emblica seeds globular, smooth. — Africa ...... 99. Inflorescences cauliflorous thyrses; stigmas petaloid. — ...... Subgenus Kirganelia section Polyanthi West Indies . . Subgenus Xylophylla section Epistylium 88. Stamens 3–5, free or slightly fused at base; anthers de- 99. Inflorescences axillary cymules; stigmas tapering, not hiscing vertically; fruits capsular; seeds smooth or faintly petaloid, sometimes fused into a tube . . . . . 100 longitudinally striate. — Africa, Madagascar, Asia . . . 100. Leaf blades mostly 1–2 cm long, with mesophyllar scle- ...... Subgenus Anesonemoides reids; stamens 3‒7. — West Indies ...... 88. Stamen 2‒5(‒7); anthers dehiscing horizontally; fruits ...... Subgenus Xylophylla section Orbicularia capsular; seeds smooth. — South America ...... 100. Leaf blades > 2 cm long, sometimes with mesophyllar ...... Subgenus Emblica section Pityrocladus sclereids; stamens 2‒7(‒8) ...... 101 R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 175

101. Brachyblasts often present; sepals 5; staminate disc con- Acknowledgements This work was done as part of the PhD research of sisting of 5 segments; stamens 3 (rarely 4); fruit a large the main author, funded by the Hortus botanicus Leiden and Leiden Univer- fleshy capsule (usually > 2 cm diam). — West Indies . sity. Help with preparing the list for the manuscript by Peter Hovenkamp is . . . . .Subgenus Xylophylla section Omphacodes gratefully acknowledged. The second author thanks the Leiden University Fund (LUF) for their support of the chair Botanical Gardens and Botany of 101. Brachyblasts absent; sepals 4–6; staminate disc usually Southeast Asia. The last author thanks the Treub Maatschappij, the Society consisting of 6 segments; stamens 2–7(–8); fruit small for the Advancement of Research in the Tropics, for their support of the dry capsule (< 1 cm diam) ...... 102 Ornstein chair in Tropical Plant Biogeography. 102. Staminate sepals 5, pistillate sepals 6; inflorescences mostly unisexual cymules appearing after the leaves, REFERENCES several pistillate flowers per node; stamens 3‒7, thecae dehiscing horizontally; style present, elongated and ex- Airy Shaw HK. 1960. Notes on Malaysian Euphorbiaceae. Kew Bulletin 14: erted from calyx, stigmas dilated, bifid to multifid. — South 469‒471. America . . .Subgenus Xylophylla section Oxalistylis Airy Shaw HK. 1963. Notes on Malaysian and other Asiatic Euphorbiaceae. 102. Sepals in both sexes 4‒6; inflorescences bisexual cymules Kew Bulletin 16: 341‒372. appearing with the expanding leaves on new branch- Airy Shaw HK. 1969. New or noteworthy species of Phyllanthus L. Kew Bulletin 23: 26‒40. lets, usually only 1 or 2 pistillate flowers among several Airy Shaw HK. 1971. CXXV. New or Noteworthy species of Phyllanthus L. Kew staminate flowers; stamens 2‒6 (or 8), thecae dehiscing Bulletin 25: 493–495. vertically; style like an erect tube, stigma branches nar- Airy Shaw HK. 1972. The Euphorbiaceae of Siam. Kew Bulletin 26: 191‒363. rowed to acute tips. — West Indies ...... Airy Shaw HK. 1974. Notes on Malesian and other Asiatic Euphorbiaceae. . . . . Subgenus Xylophylla section Thamnocharis Kew Bulletin 29: 281‒331. Airy Shaw HK. 1975. The Euphorbiaceae of Borneo. Kew Bulletin, Additional Series 4: 1‒224. DISCUSSION AND CONCLUSION Airy Shaw HK. 1976. New or noteworthy Australian Euphorbiaceae. Kew Bulletin 31: 341‒398. Taxonomic discussions on the circumscription of genus Phyl­ Airy Shaw HK. 1977. Additions and corrections to the Euphorbiaceae of lanthus are still ongoing, mainly with the question whether the Siam. Kew Bulletin 32: 69‒83. genera nested within should be subsumed (Hoffmann et al. Airy Shaw HK. 1980a. The Euphorbiaceae of New Guinea. Kew Bulletin, 2006) or remain separate (Van Welzen et al. 2014). However, Additional Series 8: 3‒230. a good understanding and clear structure within the genus Airy Shaw HK. 1980b. Notes on Euphorbiaceae from Indomalesia, Australia and the Pacific. Kew Bulletin 35: 383‒399. Phyllanthus in its current circumscription is necessary. Here Airy Shaw HK. 1981. The Euphorbiaceae of Sumatra. Kew Bulletin 36: an attempt was made to summarize over 200 years of taxo- 239‒374. nomic work on this immense group. Several issues that still Airy Shaw HK. 1982. The Euphorbiaceae of Central Malesia (Celebes, exist will hopefully be resolved in new systematic studies. The Moluccas. Lesser Sunda Is.). Kew Bulletin 37: 1‒40. provisional key to the subgenera and (sub)sections provided Alston AHG. 1931. Part VI. Supplement. In: Trimen H, A hand-book to the here works with most typical examples of Phyllanthus species. flora of Ceylon: 1‒350. Dulae & Co., London. Future research and revision work should focus on treatments Backer CA. 1945. Notes on the Flora of Java, II. Blumea 5: 490‒524. of the individual subgenera and/or sections within the genus. Baillon HE. 1858. Etude générale du groupe des Euphorbiacées. Libraire De Victor Masson, Paris. Unfortunately, not all species could be fitted in this subgeneric Baillon HE. 1862. Euphorbiaceae Neo-Caledonicae. Adansonia 2: 211‒242. classification due to exceptional characters or incomplete de- Balakrishnan NP, Chakrabarty T. 2007. Phyllanthus. In: Gangopadhyay M, scriptions (see Appendix 2). These will need further study or Chakrabarty T, Balakrishnan NP (eds), The family Euphorbiaceae in India: a more new collections to elucidate their place within the genus. synopsis of its profile, taxonomy and bibliography: 361‒391. Bishen Singh Often these are species of which only the type specimen is Mahendra Pal Singh, Dehradun, Uttarakhand. Balakrishnan NP, Nair N. 1982. New taxa and record from Saddle Peak, known and which were not collected since, and some might be Andaman Islands. Nelumbo 24: 28‒36. extinct (e.g., P. aoraiensis Nadeaud; Wagner & Lorence 2011), Bancilhon L. 1971. Contribution à l’étude taxonomique du genre Phyllanthus or they might be exceptional forms, which should be united (Euphorbiacées). Boissiera 18: 9‒81. with other species. For some we could only assign them to Barrett RL, Telford IRH. 2015. Two new species of Phyllanthus from northern subgenus level and further revision work should place them in Australia and notes on Phyllanthus, Sauropus and Synostemon (Phyllan- their appropriate sections. The placement of some species may thaceae) in Western Australia. Nuytsia 26: 149‒166. change with new research and we welcome these changes as Beille L. 1925. Sur quelques Euphorbiacées nouvelles de la flore indo- chinoise. Bulletin de la Société Botanique de France 72: 156‒163. they will lead to a better understanding of the genus Phyllanthus Beille L. 1927. Phyllanthus. In: Lecomte MH (ed), Flore Générale de l’Indo- and we hope this article inspires discussion. Chine 5: 571‒608. Masson & Cie, Éditeurs, Paris. Several issues are still unresolved and will require further at- Bentham G. 1873. Flora Australiensis 6. Reeve & Co., London. tention. Subgenus Phyllanthus, which previously spanned all Boerlage JG. 1900. Handleiding tot de kennis der flora van Nederlands Indië herbaceous species, remains difficult and more species need 3: 185–296. Brill, Leiden. to be included in new phylogenetic studies. Several groups in Bouman RW, Van Welzen PC, Sumail S, et al. 2018. Phyllanthus rufus- chaneyi: a new nickel hyperaccumulator from Sabah (Borneo Island) with our list have not had formal taxonomic treatment for some time potential for tropical agromining. Botanical studies 59 (9): 1‒12. and new revision work may identify new species and better Brenan JPM. 1967. Tropical African Plants: XXVIII. Kew Bulletin 21: 241–262. characters to differentiate them within Phyllanthus. Another Breteler FJ. 2012. Flore du Gabon 43: 1‒107. Muséum National d’Histoire taxonomic problem was created by the discussions on the va- Naturelle, Paris. lidity of Brunel’s thesis (1987), which has led to many species Britton NL. 1920. Description of Cuban plants new to science. Memoirs of being published twice under different names (see Radcliffe- the Torrey Botanical Club 16: 57‒118. Smith 1996). This will require close scrutiny in determining how Brunel JF. 1987. Sur le genre Phyllanthus L. et quelques genres voisins de la tribu des Phyllantheae Dumort. (Euphorbiaceae, Phyllantheae) en Afrique many should be synonymized. Finally, a decision should be intertropicale et à Madagascar. Institut de Botanique, Strasbourg, France. made on how to treat the paraphyly of the genus Phyllanthus. Brunel JF, Roux J. 1975. Notes sur les Phyllanthoideae (Euphorbiaceae) Whether the genus will be split or whether the clades will be ouest-africaines. I. Phyllanthus niruri L., une espèce à exclure des flores subsumed within Phyllanthus, we hope that this treatment will de l’Afrique occidentale. Bulletin de la Société Botanique de France 122: provide structure to this diverse genus. 153‒162. 176 Blumea – Volume 63 / 2, 2018

Brunel JF, Roux J. 1976. Notes sur les Phyllanthoideae (Euphorbiaceae) Hemsley W. 1898. Phyllanthodendron mirabilis Hemsl. Hooker’s Icones ouest-africaines: II. A propos du polymorphisme du Phyllanthus sublanatus Plantarum 26: 2563‒2564. Schum. et Thonn. Bulletin de la Société Botanique de France 123: 365‒376. Hidalgo BF, Hechavarría JLG, Bazan SF. 2017. Phyllanthus phialanthoides Brunel JF, Roux J. 1977. Notes sur les Phyllanthoideae (Euphorbiaceae) (Phyllanthaceae), a new species from northeastern Cuba. Revista del ouest-africaines III. A propos de la position systématique du Phyllanthus Jardín Botánico Nacional 38: 1–6. dinklagei Pax. Bulletin de la Société Botanique de France 124: 217‒225. Hoffmann P, Baker AJ, Madulid DA, et al. 2003. Phyllanthus balgooyi (Eu- Brunel JF, Roux J. 1981. Phyllanthus subsect. Odontadenii (Euphorbiaceae) phorbiaceae s.l.), a new nickel-hyperaccumulating species from Palawan au nord du fleuve Congo (Afrique de l’Ouest). Willdenowia 11: 69‒90. and Sabah. Blumea 48: 193‒199. Brunel JF, Roux J. 1984. South-east Asian Phyllantheae. II. Some Phyllan- Hoffmann P, Cheek M. 2003. Two new species of Phyllanthus (Euphorbia- thus of subsect. Swartziani. Nordic Journal of Botany 4: 469‒473. ceae) from Southwest Cameroon. Kew Bulletin 58: 347–446. Brunel JF, Roux J. 1985. Les Phyllanthus (Euphorbiaceae) affines de la Hoffmann P, Kathriarachchi H, Wurdack KJ. 2006. A phylogenetic classifi­ section Anisoliobum en Afrique intertropicale. Beiträge zur Flora von West- cation of Phyllanthaceae (Malpighiales; Euphorbiaceae sensu lato). Kew Afrika 18: 235‒254. Bulletin 61: 37‒53. Chakrabarty T, Balakrishnan NP. 2009. Transfer of Indian species of Breynia, Hooker JD. 1890. The Flora of British India 5. Reeve, London. Glochidion and Sauropus to Phyllanthus (Phyllanthaceae) – new combi- Hunter JT, Bruhl JJ. 1997a. Three new species of Phyllanthus (Euphorbia­ nations and new names. Journal of Economic and Taxonomic Botany 33: ceae: Phyllantheae) for the Northern Territory, one new species for West- 712‒716. ern Australia, and notes on other Phyllanthus species occurring in these Chakrabarty T, Balakrishnan NP. 2012. Nineteen new combinations and a regions. Nuytsia 11: 147–163. new name in Breynia J.R. Forst. & G. Forst. (Phyllanthaceae) from Indian Hunter JT, Bruhl JJ. 1997b. Two new species of Phyllanthus and notes on subcontinent. Bangladesh Journal of Plant Taxonomy 19: 119‒122. Phyllanthus and Sauropus (Euphorbiaceae: Phyllantheae) in New South Chakrabarty T, Gangopadhyay M. 1993. A new Phyllanthus L. (Euphorbia- Wales. Telopea 7 (2): 149–165. ceae) from North Andaman island. The Journal of the Bombay Natural Jablonski E. 1967. Euphorbiaceae. In: Maguire B and collaborators (eds), History Society 90: 69‒70. The botany of the Guayana Highland – part VII. Memoirs of the New York Challen G, Vorontsova MS, Schneider H, et al. 2011. Phylogenetically distinct Botanical Garden 17: 80‒190. and critically endangered new tree species of Phyllanthus from Cameroon Jangid PP, Gupta S. 2016. Systematic wood anatomy of the tribe Phyl- (Phyllanthaceae, Euphorbiaceae s.l.). Systematic Botany 36: 933‒938. lantheae (Phyllanthaceae, Euphorbiaceae s.l.) from India: implication in Chantaranothai P. 2005. Taxonomic notes on the genus Phyllanthus L. reinstatement of Phyllanthus, Glochidion and allies. Nordic Journal of (Euphorbiaceae) in Thailand. Thai Forest Bulletin (Botany) 33: 16‒20. Botany 34: 496‒512. Chen Y, Chen S, Huang T, et al. 2009. Pollen morphology of Philippine species Johnston MC. 1986. Phyllanthus barbarae (Euphorbiaceae), new species of Phyllanthus (Phyllanthaceae, Euphorbiaceae s.l.). Blumea 54: 47‒58. from southwestern Tamaulipas, Mexico. Systematic Botany 11: 35‒38. Colla L. 1831. Herbarium pedemontanum 5. Augustae Taurinorum, ex Typis Kathriarachchi H, Samuel R, Hoffmann P, et al. 2006. Phylogenetics of tribe Regis, Piedmont. Phyllantheae (Phyllanthaceae; Euphorbiaceae sensu lato) based on nrITS Collett H, Hemsley WB. 1890. On a collection of plants from Upper Burma and plastid matK DNA sequence data. American Journal of Botany 93 (4): and the Shan States. Botanical Journal of the Linnean Society 28: 1‒150. 637‒655. doi: https://doi.org/10.3732/ajb.93.4.637. Cordeiro I, Carneiro-Torres DS. 2004. A new species of Phyllanthus (Phyl- Kurosawa T. 2016. Transfer of Japanese Breynia and Glochidion to Phyl- lanthaceae) from Chapada Diamantina, Bahia, Brazil. Botanical Journal lanthus (Phyllanthaceae). Journal of Japanese Botany 91: 118‒121. of the Linnean Society 146: 247‒250. Kurz S. 1873. New Burmese plants, part III. The Journal of the Asiatic Society Croizat L. 1942. On certain Euphorbiaceae from the tropical Far East. Journal of Bengal 42: 227‒254. of the Arnold Arboretum 23: 29‒54. Leandri J. 1958. Euphorbiacées. In: Humbert H (ed), Flore de Madagascar Croizat L. 1943. Notes on American Euphorbiaceae, with descriptions of et des Comores, famille 111, part I. Didot, Paris. eleven new species. Journal of the Washington Academy of Sciences Li PT. 1987a. A revision of Chinese Phyllanthus (Euphorbiaceae). Acta 33: 11‒20. Phytotaxonomica Sinica 25: 371‒383. Croizat LCM. 1945. New or critical Euphorbiaceae from the Americas. Journal Li PT. 1987b. A revision of the Chinese Phyllanthodendron (Euphorbiaceae). of the Arnold Arboretum 26 (2): 181–196. Bulletin of Botanical Research 7: 1‒9. De Jussieu AL (ed). 1824. De Euphorbiacearum generibus medicisque Lobreau-Callen D, Punt W, Schmid M. 1988. Pollen morphology and taxon- earumdem viribus tentamen. Didot, Paris. omy of the Phyllanthus species (Euphorbiaceae) native to New Caledonia. Funez LA, Ferreira JPR, Hassemer G. 2018. Phyllanthus timboënsis (Phyl- Review of Palaeobotany and Palynology 53: 283‒304. lanthaceae), a new species from Santa Catarina, southern Brazil. Webbia Luo S, Esser H-J, Zhang D, et al. 2011. Nuclear ITS sequences help disen- 73 (1): 63–69. doi: https://doi.org/10.1080/00837792.2018.1452370. tangle Phyllanthus reticulatus (Phyllanthaceae), an Asian species not oc- Funez LA, Hassemer G. 2017. Phyllanthus eremitus (Phyllanthaceae), a curring in Africa, but introduced to Jamaica. Systematic Botany 36: 99‒104. narrowly endemic new species from Santa Catarina, southern Brazil, and Manissorn J, Sukrong S, Ruangrungsi N, et al. 2010. Molecular phyloge- lectotypification and range extension of P. hyssopifolioides. Phytotaxa netic analysis of Phyllanthus species in Thailand and the application of 319: 149‒158. polymerase chain reaction-restriction fragment length polymorphism for Gage AT. 1914. New Euphorbiaceae from India and Malaya. Bulletin of Phyllanthus amarus identification. Biological and Pharmaceutical Bulletin Miscellaneous Information Kew: 236‒241. 33: 1723‒1727. Gandoger MM. 1919. Sertum plantarum novarum. Pars secunda (Suite). McNeill J, Barrie FR, Buck WR, et al. 2012. International Code of Nomen- Bulletin de la Société Botanique de France 66: 286‒307. clature for algae, fungi and plants (Melbourne Code). Koeltz Scientific Ganesan R. 2003. Identification, distribution and conservation of Phyllan- Books, Oberreifenberg. thus indofischeri, another source of Indian gooseberry. Current Science McPherson G, Schmid M. 1991. Flore de la Nouvelle Calédonie et Dépen- 84: 1515‒1518. dances 17: Euphorbiaceae. Muséum National d’Histoire Naturelle, Paris. Gilbert MG. 1987. New and interesting species of Euphorbiaceae from Ethio-­ Meeuwis B, Punt W. 1983. Pollen morphology and taxonomy of the subgenus pia. Kew Bulletin 42: 351‒368. Kirganelia (Jussieu) Webster (genus Phyllanthus, Euphorbiaceae) from Gilbert MG, Thulin M. 1993. A new species of Phyllanthus (Euphorbiaceae) Africa. Review of Palaeobotany and Palynology 39: 131–160. from Somalia. Nordic Journal of Botany 13: 171‒173. Merrill ED. 1912. Notes on Philippine Euphorbiaceae. Philippine Journal of Govaerts R, Frodin DG, Radcliffe-Smith A. 2000. World checklist and biblio­ Science 7: 379‒410. graphy of Euphorbiaceae 1-4. Royal Botanic Gardens, Kew. Merrill ED. 1914. An enumeration of the plants of Guam. The Philippine Greenman JM. 1898. Diagnosis of new and critical phanerograms. Proceed- Journal of Science ser. C, 9: 97‒155. ings of the American Academy of Arts and Sciences 33: 453‒490. Merrill ED. 1920. Notes on Philippine Euphorbiaceae, III. Philippine Journal Guillaumin A. 1927. Contributions à la flore de la Nouvelle Calédonie. Bulletin of Science 16: 539‒580. du Museum National d’Histoire Naturelle 33: 272‒276. Merrill ED. 1926. Additions to our knowledge of the Philippine flora III. Philip- Guillaumin A. 1937. Matériaux pour la flore de la Nouvelle-Calédonie. XL, XLI pine Journal of Science 30: 389‒430. et XLII. Bulletin de la Société Botanique de France 2: 294‒315. Merrill ED. 1935. A sixth supplementary list of Hainan plants. Lingnan Sci- Haegens RMAP. 2000. Taxonomy, phylogeny and biogeography of Bac- ence Journal 14: 1‒62. caurea, Distichirhops, and Nothobaccaurea (Euphorbiaceae). Blumea Mitra D, Bennet SSR. 1967. Phyllanthus mukerjeeanus (fam. Euphorbia- Supplement 12: 1–218. ceae) ‒ A new species from west Bengal. Bulletin of the Botanical Society Hayata B. 1904. Revisio Euphorbiacearum et Buxacearum Japonicarum. of Bengal 19: 145‒146. The Journal of the College of Science, Imperial University Tokyo, Japan Moore S. 1920. A contribution to the flora of Australia. Journal of the Linnean 20: 1‒92. Society of London, Botany 45: 159‒220. R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 177

Müller J. 1863. Euphorbiaceae. Vorläufige Mittheilungen aus dem für De Rossignol L, Rossignol M, Haicour R. 1987. A systematic revision of Phyl- Candolle’s Prodromus bestimmten Manuscript über diese Familie. Lin- lanthus subsection Urinaria (Euphorbiaceae). American Journal of Botany naea 32: 1‒126. 74: 1853‒1862. Müller J. 1865. Euphorbiaceae. Vorläufige Mittheilungen aus dem für De Rusby HH. 1912. New species from Bolivia, collected by R.S. William – II. Candolle’s Prodromus bestimmten Manuscript über diese Familie. Pro- Bulletin of the New York Botanical Garden 8: 89‒136. dromus Systematis Naturalis Regni Vegetabilis 34: 1‒224. Rusby HH. 1927. Description of new genera and species of plants collected Müller J. 1866. Euphorbiaceae. In: De Candolle AP (ed), Prodromus systema- on the Mulford Biological Exploration of the Amazon Valley. Memoires of tis naturalis regni vegetabilis: 15, 2: 189‒1286. Masson, Paris. the New York Botanical Garden 7: 205‒387. Müller J. 1873. Euphorbiaceae, Phyllantheae. In: Von Martius CFP (ed), Flora Samuel R, Kathriarachchi H, Hoffmann P, et al. 2005. Molecular phyloge- Brasiliensis 11: 7‒80. Oldenbourg, Munich & Leipzig. netics of Phyllanthaceae: evidence from plastid MATK and nuclear PHYC Pagare RS, Naik SS, Krishnan S, et al. 2016. Lectotypification of Pseudo- sequences. American Journal of Botany 92: 132‒141. glochidion anamalayanum Gamble and its taxonomic position under the Santiago LJM, Louro RP, Emmerich M. 2006. Phyllanthus section Chore­ genus Phyllanthus (Phyllanthaceae). Phytotaxa 286: 61‒74. tropsis (Euphorbiaceae) in Brazil. Botanical Journal of the Linnean Society Parodi D. 1881. Duez nuevas especies pertenecientes á la Familias de las 150: 131‒164. Euphorbiáceas. Anales de la Sociedad Científica Argentina 11: 49‒56. Schweinfurth G. 1899. Sammlung arabisch-aethiopischer Pflanzen. Ergeb- Pax F, Hoffmann K. 1922. Euphorbiaceae - Phyllanthoideae - Phyllantheae. nisse von Reisen in den Jahren 1881, 88, 91 und 92. Bulletin de l’Herbier In: Engler A (ed), Das Pflanzenreich IV.147.xv: 1‒349. Engelmann, Berlin. Boissier 7 (Appendix 2): 299‒340. Pornpongrungrueng P, Parnell JA, Hodkinson TR, et al. 2017. Phyllanthus Secco RdS, De Rosário AS. 2015. A new species of Phyllanthus (Phyllantha- kaweesakii (Phyllanthaceae), a new species from Thailand. Botany 95: ceae) endemic to Amazonas State, Brazil. Novon 24: 209‒211. 567‒577. Silva MJD. 2009. Two new Brazilian Species of Phyllanthus (Phyllanthaceae). Pruesapan K, Telford IRH, Bruhl JJ, et al. 2008. Delimitation of Sauropus Novon 19 (2): 229–233. (Phyllanthaceae) based on plastid matK and nuclear ribosomal ITS DNA Silva MJD, Sales MFD. 2006. Phyllanthus L. (Phyllanthaceae) em Pernam- sequence data. Annals of Botany 102: 1007‒1018. buco, Brasil. Acta Botanica Brasilica 21 (1): 79–98. Pruesapan K, Telford IRH, Bruhl JJ, et al. 2012. Phylogeny and proposed cir- Silva MJD, Sales MFD. 2008. Sinopse do gênero Phyllanthus (Phyllantha- cumscription of Breynia, Sauropus and Synostemon (Phyllanthaceae), ceae) no nordeste do Brasil. Rodriguésia 59: 407‒422. based on chloroplast and nuclear DNA sequences. Australian Systematic Small JK. 1913. Famlily 70. Euphorbiaceae. In: Britton NL, Brown A (eds), Botany 25: 313‒330. An illustrated flora of the northern United States, Canada and the British Punt W. 1967. Pollen morphology of the genus Phyllanthus (Euphorbiaceae). possessions, from Newfoundland to the parallel of the southern boundary Review of Palaeobotany and Palynology 3: 141‒150. of Virginia, and from the Atlantic Ocean westward to the 102d meridian 2: Punt W. 1972. Pollen morphology and taxonomy of section Ceramanthus 452‒476. Charles Scribner’s Sons, New York. Baillon s.l. of the genus Phyllanthus (Euphorbiaceae). Review of Palaeo- Smith JJ. 1910. Euphorbiaceae. In: Koorders SH, Valeton T (eds), Bijdrage no. 12 tot de kennis der boomsoorten op Java. Mededeelingen uitgaande botany and Palynology 13: 213‒228. van het Departement van Landbouw 10: 9‒638. Kolff & Co., Batavia. Punt W. 1980. Pollen morphology of the Phyllanthus species (Euphorbia- Smith JJ. 1912. Euphorbiaceae. In: Lorentz MHA (ed), Nova Guinea: résul- ceae) occurring in New Guinea. Review of Palaeobotany and Palynology tats de l’expédition scientifique néerlandaise à la Nouvelle-Guinée: 8 (2): 31: 155‒177. 779‒796. Brill, Leiden. Punt W. 1986. Convergence in some interesting pollen types of Phyllanthus Smith JJ. 1920. Plantae novae vel criticae ex Herbario et Horto Bogoriensi: (Euphorbiaceae). Canadian Journal of Botany 64: 3127‒3129. I. Bulletin du Jardin Botanique de Buitenzorg ser. 3, 1: 390‒410. Punt W. 1987. A survey of pollen morphology in Euphorbiaceae with special Smith WW. 1921. Diagnosem specierum novarum in herbario Horti Regii reference to Phyllanthus. Botanical Journal of the Linnean Society 94: Botanici Edinburgensis cognitarum. Notes from the Royal Botanic Garden, 127‒142. Edinburgh 13: 149‒187. Punt W, Rentrop J. 1973. Pollen morphology of the Phyllanthus species Stuppy W. 1995. Systematische Morphologie und Anatomie der Samen der occurring in the continental United States. Review of Palaeobotany and biovulaten Euphorbiaceen. PhD Thesis Fachbereich Biologie, Universität Palynology 16: 243‒261. Kaiserslautern, Kaiserslautern, Germany. Radcliffe-Smith A. 1974. Notes on African Euphorbiaceae: V. Kew Bulletin Sunil CN, Prabhukumar KM, Kumar VVN, et al. 2016. Phyllanthus balakrish- 29: 435‒441. nanii sp. nov. (Phyllanthaceae), an interesting discovery from the southern Radcliffe-Smith A. 1982. Notes on African Euphorbiaceae: XII. Kew Bulletin Western Ghats, India. Phytotaxa 273: 65‒71. 37: 421‒428. Telford IR, Pruesapan K, Van Welzen PC, et al. 2016. Morphological and Radcliffe-Smith A. 1992. Notes on African Euphorbiaceae XXVIII. Kew Bul- molecular data show Synostemon trachyspermus (Phyllanthaceae, Phyl- letin 47: 677‒683. lantheae) to be a heterogeneous species assemblage. Australian Syste­ Radcliffe-Smith A. 1996. Notes on African Euphorbiaceae XXX: Phyllanthus matic Botany 29: 218‒234. (V) & c. Kew Bulletin 51: 301‒331. Thin NN. 2007. Taxonomy of Euphorbiaceae in Vietnam. National University Radcliffe-Smith A, Hoffmann P. 2006. New records, names and combinations Publishers, Hanoi. in African Euphorbiaceae sensu lato. Kew Bulletin 61: 609‒611. Thulin M. 2003. Phyllanthus xylorrhizus (Phyllanthaceae), a new species Ralimanana H, Hoffmann P. 2011. Taxonomic revision of Phyllanthus (Phyl- from Somalia. Nordic Journal of Botany 23: 385‒387. lanthaceae) in Madagascar and the Comoro Islands I: Synopsis and sub- Van Welzen PC, Pruesapan K, Telford IRH, et al. 2014. Phylogenetic re- genera Isocladus, Betsileani, Kirganelia and Tenellanthus. Kew Bulletin construction prompts taxonomic changes in Sauropus, Synostemon and 66: 331‒365. Breynia (Phyllanthaceae tribe Phyllantheae). Blumea 59: 77‒94. Ralimanana H, Hoffmann P. 2014. Taxonomic revision of Phyllanthus L. (Phyl- Visiani RD. 1858. Recensio Altera Plantarum Minus Cognitarum quas Hortus lanthaceae) in Madagascar and the Comoro Islands II: subgenera Ani- Patavinus colit. Atti Del Reale Istituto Veneto di Scienze, Lettere ed Arti. sonemoides (Jean F.Brunel) Ralim. & Petra Hoffm., stat. nov. and Menarda 3: 133‒142. (Müll.Arg.) Ralim. & Petra Hoffm., stat. nov. Adansonia 36: 265‒301. Von Mueller FJH. 1859. Fragmenta Phytographiae Australiae 1. Joannis Ralimanana H, Hoffmann P, Rajeriarison C. 2013. Taxonomic revision of Phyl- Ferris, Melbourne. lanthus (Phyllanthaceae) in Madagascar and the Comoro Islands III: sub- Vorontsova MS, Hoffmann P, Kathriarachchi H, et al. 2007. Andrachne cu- genera Swartziani, Afroswartziani and Emblica. Kew Bulletin 68: 535‒558. neifolia (Phyllanthaceae; Euphorbiaceae s.l.) is a Phyllanthus. Botanical Rendle AB, Baker EG, Moore S. 1921. A systematic account of the plants Journal of the Linnean Society 155: 519‒525. collected in New Caledonia and the Isle of Pines by Prof. RH Compton, MA, Wagner WL, Lorence DH. 2011. A nomenclator of Pacific oceanic island in 1914. – Part I. Flowering Plants (Angiosperms). Journal of the Linnean Phyllanthus (Phyllanthaceae), including Glochidion. Phytokeys 4: 67‒94. Society of London, Botany 45: 245‒417. Warburg O. 1891. Beiträge zur Kenntnis der papuanischen Flora (Schluss). Ribeiro RTM, Neto RLS, Loiola MIB. 2017. Phyllanthus carmenluciae, a Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzen- supreme species of Phyllanthus (Phyllanthaceae) from Brazil. Phytotaxa geographie 13: 273–455. 305: 35‒40. Webster GL. 1955. Studies of the Euphorbiaceae, Phyllanthoideae I. Taxo- Ridley HN. 1909. The Flora of the Telom and Batang Padang Valleys. Journal nomic notes on the West Indian species of Phyllanthus. Contributions from of the Federated Malay States Museums 4: 1‒98. the Gray Herbarium of Harvard University 176: 45–63. Robinson CB. 1909. Philippine Phyllanthinae. Philippine Journal of Science Webster GL. 1956. A monographic study of the West Indian species of Phyl- 4: 71‒106. lanthus. Journal of the Arnold Arboretum 37: 91–122, 217–268, 340–359. Robinson CB. 1911. Alabastra Philippensia, III. Philippine Journal of Science, Webster GL. 1957. A monographic study of the West Indian species of Phyl- C, Botany 6: 319‒358. lanthus. Journal of the Arnold Arboretum 38: 51–80,170–198, 295–373. 178 Blumea – Volume 63 / 2, 2018

Webster GL. 1958. A monographic study of the West Indian species of Phyl- Manuscripts lanthus. Journal of the Arnold Arboretum 39: 49–100, 111–212. Available online at the website of the University of California: Webster GL. 1960. Supplement to a monographic study of the West Indian http://herbarium.ucdavis.edu/webster_manuscripts.html (last accessed March species of Phyllanthus. Journal of the Arnold Arboretum 41: 279–286. 2017). Webster GL. 1967a. A remarkable new Phyllanthus (Euphorbiaceae) from Central America. Annals of the Missouri Botanical Garden 54: 194–198. Webster GL. 2001 Synopsis of Gomphidium Webster GL. 1967b. The genera of Euphorbiaceae in the southeastern United http://herbarium.ucdavis.edu/manuscripts/webster/New%20World/2001%20 States. Journal of the Arnold Arboretum 48: 363‒430. Synopsis%20of%20Gomphidium.pdf. Webster GL. 1970. A revision of Phyllanthus (Euphorbiaceae) in the conti- Webster GL. 2002 Outline of the Neotropical infrageneric taxa of Phyllanthus nental United States. Brittonia 22: 44‒76. (Euphorbiaceae) Webster GL. 1978. A new Mexican species of Phyllanthus (Euphorbiaceae) http://herbarium.ucdavis.edu/manuscripts/webster/New%20World/2002%20 with Southern Hemisphere affinities. Rhodora 80: 570‒574. Outline%20neotropical%20Phyllanthus%202.pdf. Webster GL. 1979. A revision of Margaritaria (Euphorbiaceae). Journal of Webster GL. 2002 Synopsis of Phyllanthus subgenus Emblica the Arnold Arboretum 60: 403‒444. http://herbarium.ucdavis.edu/manuscripts/webster/New%20World/2002%20 Webster GL. 1986. A revision of Phyllanthus (Euphorbiaceae) in eastern Synopsis%20of%20Phyllanthus%20subgenus%20Emblica.pdf. Melanesia. Pacific Science 40: 88‒105. Webster GL. 1995. A new species of Phyllanthus (Euphorbiaceae) from Webster GL. 2002 Synopsis of Phyllanthus subgenus Eriococcus Ceylon. Kew Bulletin 50: 266. http://herbarium.ucdavis.edu/manuscripts/webster/New%20World/2002%20 Webster GL. 1997. Phyllanthus. In: Dassanayake MD, Clatyon WD (eds), A Synopsis%20of%20Phyllanthus%20subgenus%20Eriococcus.pdf. revised handbook to the Flora of Ceylon 11: 206–235. Balkema, Rotterdam. Webster GL. 2002 Synopsis of Phyllanthus subgenus Kirganelia Webster GL. 2001a. An unusual new species of Phyllanthus (Euphorbiaceae) http://herbarium.ucdavis.edu/manuscripts/webster/New%20World/2002%20 from Colombia. Lundellia 4: 64‒68. Synopsis%20of%20Phyllanthus%20subgenus%20Kirganelia.pdf. Webster GL. 2001b. Synopsis of Croton and Phyllanthus (Euphorbiaceae) Webster GL. 2002 Synopsis of Phyllanthus subgenus Phyllanthus in western tropical Mexico. Contributions from the University of Michigan http://herbarium.ucdavis.edu/manuscripts/webster/New%20World/2002%20 Herbarium 23: 353‒388. Synopsis%20of%20Phyllanthus%20subgenus%20Phyllanthus.pdf. Webster GL. 2002a. A synopsis of the Brazilian taxa of Phyllanthus section Phyllanthus (Euphorbiaceae). Lundellia 5: 1‒26. Webster GL. 2002 Synopsis of Phyllanthus subgenus Xylophylla Webster GL. 2002b. Three new sections and a new subgenus of Phyllanthus http://herbarium.ucdavis.edu/manuscripts/webster/New%20World/2002%20 (Euphorbiaceae). Novon 12: 290‒298. Synopsis%20of%20Phyllanthus%20subgenus%20Xylophylla.pdf. Webster GL. 2003. A synopsis of Phyllanthus section Nothoclema (Euphor- Webster GL. Outline of Australian Phyllanthus (No date) biaceae). Lundellia 6: 19–36. http://herbarium.ucdavis.edu/manuscripts/webster/Austral%20and%20 Webster GL. 2004. A revision of Phyllanthus section Hylaeanthus (Euphor- the%20Pacific/Outline%20of%20Australian%20Phyllanthus.pdf. biaceae). Lundellia 7: 11‒27. Webster GL. 2007. Taxonomic and nomenclatural changes in American Euphorbiaceae sensu lato. Contributions from the University of Michigan Herbarium 25: 235‒239. Webster GL, Airy Shaw HK. 1971. A provisional synopsis of the New Guinea taxa of Phyllanthus (Euphorbiaceae). Kew Bulletin 26: 85‒109. Webster GL, Carpenter KJ. 2002. Pollen morphology and phylogenetic re- lationships in neotropical Phyllanthus (Euphorbiaceae). Botanical Journal of the Linnean Society 138: 325‒338. Webster GL, Carpenter KJ. 2008. Pollen morphology and systematics of pa­laeotropical Phyllanthus and related genera of subtribe Phyllanthinae (Euphorbiaceae). Botanical Journal of the Linnean Society 157: 591‒608. Wilson PG. 1962. Tabula 3589. Phyllanthus petaloideus. Hooker’s Icones Plantarum, ser. 5: 1‒3. Wu M-J, Ferreras U, Chen Y-J. 2017. Phyllanthus coi (Phyllanthaceae), a new herbaceous species from the Philippines. Taiwania 62: 375–380. Wu M-J, Huang T-C, Liu C-C, et al. 2016. Pollen morphology and taxonomy in Malesian Phyllanthus (Phyllanthaceae). The Journal of Japanese Botany 91, Supplement: 257‒292. R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 179 Africa Australia Asia Tropical America) Indies America, West Australia, Pacific Asia, Malesia, America Australia Australia Australia Asia, Malesia, Asia Asia Asia Asia Asia Asia America America Indies America, West America, Central America America America Asia Asia, Malesia Asia, Malesia Asia, Malesia, Asia, Malesia, Asia, Malesia, ?Australia Asia Asia, Malesia Asia, Malesia, Africa Africa Africa Africa Africa, Africa, Africa Africa, Africa Africa Africa, Mainland Mainland Africa, Madagascar, America’, Malesia, Pantropical Mainland Madagascar, Tropical Tropical Tropical Tropical Tropical Madagascar Madagascar Tropical Tropical Tropical Asia Tropical South Tropical South Tropical South Tropical South Tropical Indies West Indies West Indies West Pantropical, origin Tropical Mainland Tropical Mainland Pantropical, origin Pantropical, origin Mainland Pantropical South Tropical South Tropical Mainland Mainland Mainland Mainland Australia, Malesia, Pacific Pantropical (Introduced in North Mainland New Caledonia New Caledonia Mainland ‘Central Malesia, New Caledonia New Caledonia Tropical Distribution Pantropical: origin South South Tropical Central America Malesia, Pacific

Congo

P. niruri P. lokohensis P. P. callidiscus P. P. felicis P. P. microdendron P. P. odontadensis P. P. arvensis P. P. bojerianus P. albidiscus P. P. betsileanus P. P. welwitschianus P. P. petraeus P. cochinchinensis P. P. albidiscus P. P. letouzeyanus P. P. brasiliensis P. orbiculatus P. P. acuminatus P. P. attenuatus P. P. carnosulus P. lindenianus P. emblica P. P. lindenianus P. baeobotryoides P. dinklagei P. P. reticulatus P. P. emblica P. casticum P. P. urinaria P. acutissimus P. P. vaciniifolius P. ruscifolius P. urinaria P. gracilipes P. lamprophyllus P. gracilipes P. maderaspatensis P. P. clamboides P. P. nhatrangensis P. ruber P. kanalensis P. P. faguettii P. P. buxifolius P. chamaecerasus P. aeneus P. loranthoides P. P. pinnatus P. species Type acidus P. chacoensis P. P. chamaecerasus P. P. tuerckheimii P. Appendix 2. Countries in parentheses indicate unlikely disjunct distributions that require further study. L. Ralim. & Petra Hoffm. R.W.Bouman (Jean F.Brunel) Jean F.Brunel Jean F.Brunel & Jacq.Roux) R.W.Bouman (Jean F.Brunel Jean F.Brunel Ralim. & Petra Hoffm. (Jean F.Brunel) (Hassk.) Jean F.Brunel Ralim. & Petra Hoffm. (Jean F.Brunel) Müll.Arg. & Jacq.Roux Jean F.Brunel Müll.Arg. (Hassk.) Baill. Jean F.Brunel (Aubl.) G.L.Webster G.L.Webster G.L.Webster G.L.Webster G.L.Webster G.L.Webster (Gaertn.) Kurz G.L.Webster Jean F.Brunel & Jacq.Roux Jean F.Brunel (A.Juss.) Griseb. (Gaertn.) Baill. (A.Juss.) Kurz (Gaertn.) G.L.Webster Müll.Arg. (Müll.Arg.) Müll.Arg. G.L.Webster Haicour & Rossignol (Hassk.) Croizat & Metcalf Müll.Arg. (Hassk.) Müll.Arg. G.L.Webster (K.Schum.) J.J.Sm. Jean F.Brunel Müll.Arg. (Lour.) Baill. (Blume) Müll.Arg. (Baill.) G.L.Webster (Müll.Arg.) Müll.Arg. (Wight) Müll.Arg. Author (L.) Müll.Arg. (Chodat) G.L.Webster Baill. G.L.Webster ex R.W.Bouman (Müll.Arg.) G.L.Webster (Müll.Arg.) R.W.Bouman Author of type species can be found in

Urinaria

Spiciferens Eleutherogynium Subsection Aporosella Physoglochidion

strum

Callidisci Fluitantoides Microdendron Odontadenii Praephyllanthus Anisolobium

Bivia Cluytopsis Ebolowani Ceramanthus Apolepsis Hylaeanthus Nothoclema Callitrichoides Cyclanthera Botryoides Brazzeani Chorisandra Emblica Anisomena Microglochidion Pityrocladus Urinaria Urinaria Emblica Eriococcodes Eriococcus Eriococcus Nymphanthus Leptonema Nymania Scepasma Adenoglochidion Adenoglochidion Calodictyon Section Cicca Cicca Gomphidium Gomphidium

Afroswartziani

Afroswartziani Afroswartziani Afroswartziani Afroswartziani Afroswartziani Anisonemoides Ceramanthus Betsileani Ceramanthus Ceramanthus Ceramanthus Ceramanthus Ceramanthus Conami Conami Conami Conami Cyclanthera Cyclanthera Emblica Cyclanthera Emblica Kirganelia Kirganelia Emblica Kirganelia Kirganelia Emblica Emblica Emblica Emblica Eriococcus Eriococcus Eriococcus Eriococcus Isocladus Eriococcus Eriococcus Gomphidium Gomphidium Eriococcus Gomphidium Gomphidium Gomphidium Gomphidium Subgenus Kirganelia Kirganelia Gomphidium Gomphidium

Phyllanthus Appendix 1 Synopsis of the infrageneric classification genus Phyllanthus . Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Genus Phyllanthus Phyllanthus Phyllanthus Phyllanthus 180 Blumea – Volume 63 / 2, 2018 Australia Africa America America America? America Africa Africa America’ America America America America America America America America, ‘Australia’ America America America America America Indies America, West Asia Asia Asia, Malesia, Pacific, Asia Asia Asia Asia Africa Africa Africa Africa Africa South Tropical China, Japan Tropical Tropical Pantropical, origin possibly Middle East? Madagascar, Mainland Mainland Tropical Indies West Mainland Mainland Mainland Mainland Mainland South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical Indies West Distribution South Tropical Indies West Australia, ‘North Pantropical South Tropical Pantropical, origin North North America Pantropical, origin Tropical Tropical Pantropical, origin South Tropical South Tropical South and Middle Tropical South and Middle Tropical South Tropical South Tropical Central and South Indies West Indies West Indies West Indies West Indies West South Tropical Indies West Indies West Indies West Indies West

Americas

P. flexuosus P. physocarpus P. P. pseudocicca P. polyanthus P. P. acidus P. P. cryptophilus P. mirabilis P. P. arachnodes P. P. purpureus P. mirificus P. P. virgatus P. P. anthopotamicus P. P. mirabilis P. P. roseus P. P. moi P. P. niruri P. choretroides P. klotschianus P. choretroides P. P. claussenii P. niruri P. P. pentaphyllus P. species Type P. dictyospermus P. P. epiphyllanthus P. P. caroliniensis P. P. subcrenulatus P. niruri P. almadensis P. P. fluitans Americas P. amarus P. P. tenellus P. P. warnockii P. P. loandensis P. pentandrus P. P. tenellus P. P. epiphyllanthus P. P. adianthoides P. P. juglandifolius P. P. riedelianus P. P. rupestris P. P. glaucescens P. axillaris P. chryseus P. ovatus P. subcarnosus P. orbicularis P. salviifolius P. cinctus P. P. octomerus P. P. discolor P. P. discolor P. P. williamioides P. A.Juss.) Ralim. & Petra Hoffm. Airy Shaw (Baill.) Müll.Arg. Jean F.Brunel G.L.Webster Jean F.Brunel Kuntze (Hemsl.) G.L.Webster (Gagnep.) Müll.Arg. G.L.Webster (Wight) Jean F.Brunel (Comm. ex Croizat Croizat Croizat P.T.Li – Müll.Arg. L.J.M.Santiago – G.L.Webster G.L.Webster Author G.L.Webster (L.) Baill. G.L.Webster G.L.Webster (F.Muell.) – G.L.Webster Holm-Niels. ex Jean F.Brunel Ralim. & Petra Hoffm. (G.L.Webster) Jean F.Brunel (A.Gray) G.L.Webster Jean F.Brunel G.L.Webster Jean F.Brunel (L.) Pers. Jabl. ex Jean F.Brunel (Klotzsch) Müll.Arg. Müll.Arg. G.L.Webster Croizat Griseb. (Sw.) G.L.Webster (Müll.Arg.) Müll.Arg. G.L.Webster (Baill.) Griseb Baill. G.L.Webster Müll.Arg. (Baill.) Müll.Arg. G.L.Webster G.L.Webster G.L.Webster

Cheramela

Mirifici

Applanata Choretropsis Niruri Pentaphylli Subsection

Almadanses Clausseniani

Discolores Incrustati

Hemicicca Omphacodopsis Ciccopsis Polyanthi Pseudomenarda Cicca Williamia Arachnodes Calophyllum Xylophylla

Phylllanthodendron Pseudoactephila Tetrandrum Choretropsis Choretropsis Choretropsis Loxopodium Phyllanthus Phyllanthus Salviniopsis Section Antipodanthus Lysiandra Phyllanthus Phyllanthus Phyllanthus

Reverchonia

Loandani Pentandra Tenellanthus Adianthoides Asterandra Cicca strum Diplocicca Brachycladus Elutanthos Epistylium Glyptothamnus Hemiphyllanthus Omphacodes Orbicularia Oxalistylis Thamnocharis Williamia Williamia Williamia

Kirganelia Kirganelia Kirganelia Kirganelia Kirganelia Kirganelia Phyllanthodendron Phyllanthodendron Phyllanthodendron Xylophylla Macraea Menarda Phyllanthodendron Phyllanthodendron Phyllanthodendron Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Subgenus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Swartziani Swartziani Tenellanthus Tenellanthus Tenellanthus Tenellanthus Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla

Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Appendix 1 (cont.) Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Genus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 181 - subg. Phyllanthus subg. Phyllanthus Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 D x x x Brunel & Roux 1984 Balakrishnan & Chakrabarty 2007 x x x x Ralimanana et al. 2013 Ralimanana et al. 2013 x x x x Brunel 1987 Ralimanana et al. 2013 x x Brunel 1987 x x 1957 Webster x x x Brunel 1987 Balakrishnan & Chakrabarty 2007 x x Brunel 1987 x x & Cheek 2003 Hoffmann x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x 2002 synopsis of Webster x x Ralimanana et al. 2013 x x Brunel 1987 x x x Ralimanana et al. 2013 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x x x Ralimanana et al. 2013 Balakrishnan & Chakrabarty 2007 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Ralimanana et al. 2013 x x x x Brunel 1987 Ralimanana et al. 2013 x x Ralimanana et al. 2013 x x x x Brunel 1987 2002 synopsis of Webster x x Brunel 1987 x x x Brunel 1987 Balakrishnan & Chakrabarty 2007 x x Brunel 1987 x x Brunel 1987 x x 2006 & Hoffmann Radcliffe-Smith x x x x x Brunel 1987 Ralimanana et al. 2013 x x Brunel 1987 x x x Ralimanana et al. 2013 Asia Asia Asia Asia Asia Asia Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa, Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Global area Mainland Tropical Madagascar Madagascar Madagascar Tropical Tropical Tropical Mainland Tropical Tropical Tropical Tropical Tropical Tropical Indian Ocean Madagascar Tropical Tropical Mainland Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Mainland Tropical Tropical Tropical Tropical Tropical Tropical Tropical Madagascar Tropical Tropical Indian Ocean Madagascar Tropical Mainland Tropical Tropical Tropical Tropical Madagascar Tropical Madagascar Thailand Mainland Tanzania Zimbabwe Tanzania, Tanzania Zambia Tanzania, Zambia Tanzania, Zambia Tanzania, Tanzania Tanzania Tanzania Zambia, Zimbabwe Tanzania, Africa Africa African Republic African Republic Asia, widespread Africa Major area Zambia Madagascar Northern Angola Ivory Coast India Congo Cameroon Cameroon Democratic Republic of the Congo, Zambia Central Chad, Kenya, Sudan, United Republic of Mascarenes Madagascar Democratic Republic of the Congo, Malawi, United Republic of Africa,

Angola Tropical Uganda Cameroon Togo Congo Cameroon United Republic of Equatorial Guinea, Gabon Uganda Democratic Republic of the Congo, Kenya, Zambia, Zimbabwe Tropical Angola, Democratic Republic of the Congo, Zambia Bhutan, India, Nepal Kenya, United Republic of South Ethiopia, Somalia Angola, Democratic Republic of the Congo, Zambia Angola Mozambique, United Republic of United Republic of Madagascar Kenya, United Republic of Cameroon Mascarenes Madagascar Democratic Republic of the Congo Bhutan, India Central Kenya, Mozambique, Somalia, Uganda, United Republic of United Republic of Kenya Madagascar Angola, Democratic Republic of the Congo, United Republic of Madagascar Arg. for location based placements, a question mark and/or a ~ symbol when morphology does not completely comply with the group. Müll.Arg. Jean F.Brunel Leandri & Jacq.Roux Jean F.Brunel Madagascar S.Moore Müll.Arg. Jean F.Brunel Jean F.Brunel Jean F.Brunel Ralim. & Petra Hoffm. Madagascar Radcl.-Sm. ex Müll.Arg. E.Mey. Müll Radcl.-Sm. Jean F.Brunel Lisowski Jean F.Brunel Jean F.Brunel Jean F.Brunel Müll.Arg. Pax Hutch. Jean F.Brunel Hutch. Jean F.Brunel Jean F.Brunel Poir. Hutch. L.B.Chaudhary & R.R.Rao India & Cheek Petra Hoffm. & Jacq.Roux Jean F.Brunel Republic of Bangladesh, People’s India, Myanmar, Jean F.Brunel Leandri Hutch. Beille Leandri Jean F.Brunel Jean F.Brunel J.G.Klein ex Willd. P. airy - shawii P. Species P. ajmerianus P. ampandrandavae P. P. andranovatensis P. angustatus Hutch. P. P. ankazobensis P. asperulatus P. P. benguelensis P. P. bonnardii P. P. brevipes Hook.f. P. brynaertii Jean F.Brunel P. P. caesiifolius P. P. camerunensis P. P. carunculatus P. P. cerastotemon Brenan P. P. chevalieri P. P. consanguineus P. P. coursii Leandri P. crassinervius P. debilis P.

P. dekindtianus P. P. delagoensis P. P. denticulatus P. P. discolaciniatus P. & Jacq.Roux) eliae (Jean F.Brunel P. Jean F.Brunel epiphylliferens P. P. fotii Jean F.Brunel P. P. friesii Hutch. P. P. gabonensis P. P. geniculatostemon P. P. gillettianus P. P. gossweileri P. P. griffithii Müll.Arg. P. harrisii Radcl.-Sm. P. P. heterophyllus P. P. hildebrandtii P. P. holostylus Milne-Redh. P. P. humpatanus P. P. hutchinsonianus P. P. irriguus Radcl.-Sm. P. P. ivohibeus P. kaessneri Hutch. P. P. kelleanus P. P. lanceolatus P. P. leandrianus P. leonardianus P. P. leschenaultii P. letestui Jean F.Brunel P. P. leucocalyx P. leucochlamys P. P. leucosepalus P. P. lokohensis P. macranthus Pax P. madagascariensis P.

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literature references or phylo Appendix 2 Species checklist of Phyllanthus based on the current infrageneric classification. Each species denotation contains information whether classification was morphology, genetic evidence. Unsure placements are noted with ‘loc’ Subg. Afroswartziani Subgeneric placement Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani

Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani 182 Blumea – Volume 63 / 2, 2018

Jean F.Brunel P. xiphophorus P. subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus Phyllanthus Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 D x x 1996 Radcliffe-Smith x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x x x Ralimanana et al. 2013 x Ralimanana et al. 2013 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x x x Brunel 1987 Ralimanana et al. 2013 Mitra & Bennet 1967 x x Müller 1863, 1866 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x & Cheek 2003 Hoffmann x x 1996 Radcliffe-Smith x x x x Brunel 1987 2002 synopsis of Webster x Balakrishnan & Chakrabarty 2007 x x Brunel 1987 x Balakrishnan & Chakrabarty 2007 x x 2002 synopsis of Webster x x x x Brunel 1987 2002 synopsis of Webster x x Brunel 1987 x x x x x Ralimanana et al. 2013 Ralimanana et al. 2013 x x carunculatus 1996, but see P. Radcliffe-Smith x x Brunel 1987 x x Brunel 1987 x x x x x Brunel 1987 Ralimanana et al. 2013 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x x x Brunel 1987 Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x 1996 Radcliffe-Smith x x x x Brunel 1987 Brunel 1987 x x 1996 Radcliffe-Smith x x Brunel 1987 x x Brunel 1987 x x x x x x Brunel 1987 x Ralimanana et al. 2013 x Ralimanana et al. 2013 Brunel 1987 x x 1996, but see Brunel 1987 Radcliffe-Smith Jean F.Brunel x x 1996 Radcliffe-Smith x x 1996 Radcliffe-Smith x x Brunel 1987 Asia Asia Asia Asia Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Global area Tropical Tropical Tropical Tropical Tropical Indian Ocean Madagascar Tropical Tropical Tropical Tropical Tropical Madagascar Mainland Tropical Tropical Tropical Tropical Tropical Tropical Tropical Indian Ocean Mainland Tropical Mainland Indian Ocean Tropical Indian Ocean Tropical Tropical Madagascar Mainland Tropical Tropical Tropical Tropical Tropical Madagascar Tropical Tropical Tropical Tropical Madagascar Tropical Tropical Tropical Tropical Madagascar Madagascar Madagascar Tropical Tropical Tropical Tropical Togo Tropical Tanzania Tropical Tanzania Tropical Tanzania Tanzania, Tanzania Tanzania Tanzania Zambia Tanzania, Tanzania Tanzania Tanzania Africa to Sri Lanka and India Togo Major area Malawi Cameroon United Republic of Congo Cameroon, Nigeria Aldabra, Seychelles Madagascar Malawi, United Republic of Congo United Republic of Democratic Republic of the Congo Liberia Madagascar India Zimbabwe Cameroon Nigeria Guinea, Kenya, Uganda, United Republic of Zaire, Zimbabwe Cameroon Malawi Angola, Botswana, Namibia, Zambia, Zimbabwe Mascarenes Nepal Bhutan, India, Myanmar, Angola, Democratic Republic of the Congo, Malawi, United Republic of India Mascarenes Togo Mascarenes Angola, Zimbabwe Democratic Republic of the Congo, Zambia Madagascar India, Sri Lanka, Myanmar United Republic of Tropical Ghana, Ethiopia, Kenya, Sudan, Uganda, United Republic of Guinea Democratic Republic of the Congo Benin, Burkina Faso, Ghana, Guinea, Ivory Coast, Mali, Ethiopia, Kenya, Somalia, Uganda, United Republic of Madagascar Kenya Cameroon to Ethiopia and Zimbabwe Zambia United Republic of Madagascar Malawi, Zambia, Zimbabwe United Republic of Congo Democratic Republic of the Congo Madagascar Madagascar Madagascar Zambia Zambia Malawi, Zambia, Zimbabwe Ethiopia, Kenya, Uganda, United Republic of Thonn. Radcl.-Sm. Jean F.Brunel Jean F.Brunel Jean F.Brunel (Leandri) Leandri D.Mitra & Bennet Leandri Jean F.Brunel Leandri Jean F.Brunel Dinter & Pax Baill. ex Müll.Arg. Poir. Radcl.-Sm. J.G.Klein ex Willd. Radcl.-Sm. Leandri De Wild. Coode Jean F.Brunel Müll.Arg. Radcl.-Sm. Schumach. & Jean F.Brunel Jean F.Brunel Radcl.-Sm. Buch.-Ham. ex D.Don Müll.Arg. Roxb. Radcl.-Sm. Brenan Jean F.Brunel Wight Müll.Arg. Jean F.Brunel Jean F.Brunel Radcl.-Sm. Species mafingenis P. P. magdemeanus P. P. mahengeaensis P. P. makitae Jean F.Brunel P. P. mannianus P. P. mckenziei Fosberg P. melleri Müll.Arg. P. micromeris Radcl.-Sm. P. P. mindouliensis P. P. mkurirae Jean F.Brunel P. P. moeroensis P. P. monroviae P. P. moramangicus P. mukerjeeanus P. myrtaceus Sond. P. P. ndikinimekianus P. P. nigericus P. P. niruroides P. & Cheek nyale Petra Hoffm. P. P. nyikae Radcl.-Sm. P. P. omahekensis P. P. oppositifolius P. P. parvifolius P. parvus Hutch. P. pendulus P. phillyreifolius P. P. pierlotii P. P. pileostigma P. P. prostratus Müll.Arg. P. P. pseudocarunculatus P. P. rangoloakesis P. rheedei P. rhizomatosus Radcl.-Sm. P. P. rotundifolius P. P. rouxii Jean F.Brunel P. P. sepialis P. serandii P. P. shabaensis P. P. sublanatus P. P. suffrutescens Pax P. P. tampinensis P. tanaensis Jean F.Brunel P. P. taylorianus P. P. tenuis Radcl.-Sm. P. P. thulinii P. P. tukuyuanus Jean F.Brunel P. udoricola P. P. ukagurensis P. P. upembaensis P. P. vanderystii Hutch. & De Wild. P. P. vatovaviensis P. venustulus Leandri P. viquieri P. xiphephorus P. zambicus Radcl.-Sm. P. P. zornioides P. P. fischeri Pax P.

Callidisci

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sect. Appendix 2 (cont.) Subgeneric placement Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 183 subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 D x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x 2002 synopsis of Webster x x 2002 synopsis of Webster x x Brunel & Roux 1981 x x Brunel & Roux 1981 x x 2002 synopsis of Webster x x 2002 synopsis of Webster x x 2002 synopsis of Webster x x 2002 synopsis of Webster x x Brunel & Roux 1981 x x Brunel & Roux 1981 x x Brunel & Roux 1981 x x Brunel & Roux 1981 x x 2002 synopsis of Webster x x 2002 synopsis of Webster x x 2002 synopsis of Webster x x Brunel & Roux 1981 x x Brunel & Roux 1981 x x 2002 synopsis of Webster x x Brunel & Roux 1981 x x Brunel & Roux 1981 x x 2002 synopsis of Webster x x 2002 synopsis of Webster x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987

x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x 1992 Radcliffe-Smith x Gilbert 1987 Brenan 1967 x Gilbert 1987 x x x x 2014 Ralimanana & Hoffmann 2014 Ralimanana & Hoffmann 1992 Radcliffe-Smith x x x x 2014 Ralimanana & Hoffmann

Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Global area Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Madagascar Madagascar Tropical Madagascar Tropical Madagascar Zaire, Zambia, Tanzania, Zambia Tanzania, Tanzania Tanzania Togo Tanzania Tanzania Zaire Tanzania, Zambia, Zimbabwe Tanzania, Mozambique, Zambia Tanzania, Tanzania Tanzania Tanzania Zambia Tanzania, Africa Africa Tanzani Africa Tropical Major area Democratic Republic of the Congo, Ethiopia United Republic of Democratic Republic of the Congo Angola Guinea Angola, Zambia Tropical Burundi Nigeria Nigeria, Sierra Leone, Sierra Leone Congo Democratic Republic of the Congo, Rwanda Ethiopia, Malawi, Mozambique, Zaire, Zambia, Zimbabwe Tropical Ghana United Republic of Angola, Burundi, Kenya, Rwanda, Sierra Leone, Uganda, Tropical United Republic of Botswana, Burundi, Cameroon, Ethiopia, Kenya, Rwanda, Tropical Zimbabwe Cameroon, Gabon Sudan, Uganda, United Republic of Congo, Rwanda Kenya, Democratic Republic of the Congo Zambia Angola, Democratic Republic of the Congo, Malawi, United Republic of Democratic Republic of the Congo, Ethiopia, Kenya, Malawi, Tropical Uganda, United Republic of Malawi Kenya South Angola Kenya, United Republic of Ehtiopia, Kenya, Zaire Angola, United Republic of Angola, Burundi, Democratic Republic of the Congo, Malawi, Tropical United Republic of Cameroon United Republic of Angola, Democratic Republic of the Congo, Malawi, Zambia Tropical United Republic of Ethiopia Malawi, Zambia Ethiopia United Republic of Madagascar Madagascar Ethiopia Madagascar Democratic Republic of the Congo, Malawi, Madagascar United Republic of Jean F.Brunel Ralim. Jean F.Brunel Radcl.-Sm. Müll.Arg. Radcl.-Sm. Leandri Hutch. Müll.Arg. Arg. & Jacq.Roux Jean F.Brunel Ivory Coast Müll.Arg. Jean F.Brunel Brenan M.G.Gilbert M.G.Gilbert Müll.Arg. Jean F.Brunel & Jacq.Roux Jean F.Brunel West Jean F.Brunel Radcl.-Sm. Radcl.-Sm. Leandri Müll.Arg. Cufod. M.G.Gilbert Pax & Jacq. Roux Jean F.Brunel Democratic Republic of the Congo, Ghana, Ivory Coast, Robyns & Lawalrée Radcl.-Sm. Leandri Engl. Jean F.Brunel Pax Robyns & Lawalrée Jean F.Brunel Hutch. & Jacq.Roux Jean F.Brunel & Jacq.Roux Jean F.Brunel Species P. limmuensis P. P. wingfieldii P. P. yangambiensis P. P. youngii P. P. felicis Jean F.Brunel P. P. microdendron P. & Jacq.Roux aspersus Jean F.Brunel P. Cameroon P. bancilhonae P. P. bequaertii P. P. burundiensis P. & Jacq.Roux caligatus Jean F.Brunel P. Cameroon P. dusenii P. P. gagnioeva P. jaegeri P. P. kivuensis Jean F.Brunel P. P. lebrunii P. P. leucanthus P. P. magnificens P. & Jacq.Roux mieschii Jean F.Brunel P. Democratic Republic of the Congo Congo, P. nozeranianus P. P. odontadenioides P. P. odontadenius P. pseudoniruri P. raynalii P.

P. trichotepalus P. P. volkensii P. P. wittei Robyns & Lawalrée P. P. xiphophorus P. P. arvensis Müll P. boehmii P. confusus Brenan P. P. fluminis - athi P. P. meyerianus P. P. microphyllinus P. P. mittenianus Hutch. P. P. oblongiglans P. P. oxycoccifolius P. P. paxii Hutch. P. schaulsii P. P. tanzanianus P. P. virgulatus Müll.Arg. P. P. austroparensis P. P. borenensis P. P. caespitosus Brenan P. P. dewildeorum P. P. bemangidiensis P. P. ankarana P. P. dictyophlebsis P. P. analamerae P. P. mooneyi M.G.Gilbert P. P. ambatovolana P. P. retinervis Hutch. P.

Callidisci Callidisci Callidisci Callidisci Fluitantoides Microdendron Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Odontadenii Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus Praephyllanthus

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. loc loc loc loc loc loc loc Appendix 2 (cont.) Subgeneric placement Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani

Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Afroswartziani Subg. Anisonemoides Subg. Afroswartziani Subg. Anisonemoides Subg. Afroswartziani Subg. Anisonemoides Subg. Anisonemoides 184 Blumea – Volume 63 / 2, 2018 Most important literature 3 L 2 M 1 x Thin 2007 Airy Shaw 1969 x x x x x x x x 2014 Ralimanana & Hoffmann x 2011 Ralimanana & Hoffmann x 2011 Ralimanana & Hoffmann ~ 2011 Ralimanana & Hoffmann Thin 2007 x x & Carpenter 2008 Webster x x x x 2014 Ralimanana & Hoffmann 2014 Ralimanana & Hoffmann x x Brunel & Roux 1985, 1987 x x Alston 1931 x x 2014 Ralimanana & Hoffmann x x Airy Shaw 1974 x x 2014 Ralimanana & Hoffmann x x x Brunel & Roux 1985, 1987 x x x x 2014 Ralimanana & Hoffmann x x x x Brunel & Roux 1985, 1987 Brunel & Roux 1985, 1987 x x Balakrishnan & Nair 1982 x x 2014 Ralimanana & Hoffmann x x x Punt 1972, Brunel & Roux 1985, 1987 x x 2014 Ralimanana & Hoffmann x x x Brunel & Roux 1985, 1987 Wilson 1962 x x 2004 Webster x x 2003 Webster x x x x x x 2002b Webster x x 1957 Webster 1957 Webster x Pagare et al. 2016 x x 2014 Ralimanana & Hoffmann x x x x 2002b Webster 1957 Webster x x 2014 Ralimanana & Hoffmann x x 2014 Ralimanana & Hoffmann D x x 2014 Ralimanana & Hoffmann America America America x America America x x America x America & Carpenter 2002 Punt 1987, Webster x x America x x 2004 Webster x x America 2004 Webster x x 2004 Webster x x America 2004 Webster x America 2004 Webster x x 2004 Webster America 2004 Webster x x America x x 2004 Webster x America x x 2003 Webster x America x 2003 Webster x x 2003 Webster 2001b Webster x x 2003 Webster x 2003 Webster Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Africa Africa America x x America x America 2001b Webster x x x 2003 Webster x 2001b Webster America Mainland Mainland Madagascar Madagascar Madagascar Madagascar Mainland Tropical Malesia Madagascar Madagascar Indian Ocean Madagascar Tropical Mainland Madagascar Tropical Mainland Madagascar Mainland Mainland Madagascar Tropical Madagascar North South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical Indies West South Tropical South Tropical Indies West South Tropical Central South Tropical Central South Tropical Indies West Indies West Indies West Mainland Madagascar Indies West Indies West Madagascar Tropical Madagascar Global area Togo Thailand, Vietnam Zambia Tanzania, Thailand Venezuela Trinidad, Vietnam Thailand Madagascar Madagascar Madagascar Madagascar Vietnam Benin, Ghana, Guinea, Ivory Coast, Sierra Leone, Indonesia (Java) Madagascar Madagascar Sri Lanka Angola, Democratic Republic of the Congo, Malawi, Democratic Republic, Lao People’s Madagascar United Republic of Sierra Leone, Liberia, Guinea Cambodia Madagascar Indonesia (Java), Madagascar Democratic Republic, Cambodia, China, India, Lao People’s Vietnam Cameroon Mainland Madagascar Mexico Peru, Bolivia, Brazil, Colombia, French Guiana, Paraguay, Surinam, French Guyana, Peru, Venezuela Brazil, Colombia, Ecuador, Ecuador South Tropical Venezuela South Tropical Colombia Brazil, Colombia Brazil Brazil Costa Rica Colombia Argentina, Brazil, Caribean, French Guiana, Mexico, Venezuela Nicaragua, Panama, Peru Colombia, Costa Rica, Ecuador, Central Brazil, Guyanas, Peru, Surinam, Venezuela, Mexico, Peru, Venezuela Brazil, Colombia, Costa Rica, Ecuador, South Tropical Venezuela South Tropical Costa Rica, Mexico Venezuela Mexico Costa Rica, El Salvador, Peru Cuba Haiti Dominican Republic, Haiti Madagascar Cuba, Dominican Republic, Haiti Cuba, Haiti Madagascar Mozambique Madagascar, Madagascar Major area Airy Shaw Leandri Spreng. Müll.Arg. (Gamble) G.L.Webster India Leandri Leandri Müll.Arg. Leandri & N.G.Nair N.P.Balakr. India Ralim. & Petra Hoffm. Madagascar Jean F.Brunel W.A.Rodrigues & Proctor G.L.Webster Islands Cayman Leandri Leandri (Baill.) Müll.Arg. Croizat Müll.Arg. Baill. Paul G.Wilson Baill. Tagane Leandri Baill. Baill. (Aubl.) Poir Kunth

Rich. Müll.Arg. G.L.Webster Müll.Arg. Thin Müll.Arg. (Baill.) Müll.Arg. Alston

G.L.Webster (Ridl.) Leandri

Croizat G.L.Webster & Beille ex Beille A.Chev. Craib Jabl. G.L.Webster Ralim. & Petra Hoffm. G.L.Webster Thin Airy Shaw

P. pacoensis P. P. kerrii P. P. obdeltophyllus P. vakinankaratrae P. P. bathianus P. betsileanus P. philippioides P. binhii P. kerstingii Jean F.Brunel P. P. hasskarlianus P. P. multiflorus Poir. P. P. welwitschianus P. P. dealbatus P. P. mantadiensis P. P. collinsae P. P. mananarensis P. petraeus P. P. bokorensis P. P. isomonensis P. albidiscus P. cochinchinensis P. P. andamanicus P. P. iratsiensis Leandri P. letouzeyanus P.

P. anamalayanus P. P. humbertianus P. P. petaloideus P. orbiculatus P. attenuatus Miq. P. awaensis P. bernardii P. callejasii P. madeirensis P. manausensis P. puntii G.L.Webster P. skutchii Standl. P. valleanus P. acuminatus Vahl P. anisolobus P. brasiliensis P. caymanensis P. graveolens P. liesneri P. mcvaughii P. meridensis P. mocinianus P. pavonianus P. carnosulus P. abditus G.L.Webster P. berteroanus P. lindenianus P. P. goudotianus P. P. tenuicaulis P. P. gordonii P. P. bojerianus P. P. bernieranus P. Species

Anisolobium Anisolobium Bivia Ceramanthus Cluytopsis Ebolowani Callitrichoides Cyclanthera Cyclanthera Cyclanthera Cyclanthera

Apolepsis Hylaeanthus Hylaeanthus Hylaeanthus Hylaeanthus Hylaeanthus Hylaeanthus Hylaeanthus Hylaeanthus Hylaeanthus Nothoclema Nothoclema Nothoclema Nothoclema Nothoclema Nothoclema Nothoclema Nothoclema Nothoclema Nothoclema ? sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

? sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Appendix 2 (cont.) Subg. Emblica Subg. Betsileani Subg. Betsileani Subg. Betsileani Subg. Ceramanthus Subg. Ceramanthus Subg. Emblica Subg. Anisonemoides Subg. Anisonemoides Subg. Ceramanthus Subg. Emblica Subg. Anisonemoides Subg. Emblica Subg. Anisonemoides Subg. Ceramanthus Subg. Emblica Subg. Anisonemoides Subg. Ceramanthus Subg. Ceramanthus Subg. Emblica Subg. Anisonemoides Subg. Ceramanthus Subg. Anisonemoides Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Conami Subg. Cyclanthera Subg. Cyclanthera Subg. Cyclanthera Subg. Cyclanthera Subg. Emblica Subg. Anisonemoides Subg. Cyclanthera Subg. Emblica Subg. Anisonemoides Subg. Anisonemoides Subg. Anisonemoides Subgeneric placement Subg. Anisonemoides R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 185

Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus P. major Steyerm. subg. Emblica subg. Emblica subg. Emblica subg. Emblica subg. Emblica subg. Emblica subg. Emblica Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus

Most important literature 3 L 2 M 1 ~ x x Hooker 1890 Brunel 1987 subg. Phyllanthus subg. Phyllanthus x x 2002 Synopsis Webster ~ x x 2002 Synopsis Ralimanana et al. 2013, Webster x x x Chen et al. 2009 Robinson 1911, 2002 Synopsis Ralimanana et al. 2013, Webster x x x x x x x x x 2002 Synopsis Webster Müller 1866 x Müller 1866 2002 Synopsis Webster ~ Collett & Hemsley 1890 subg. Phyllanthus subg. Phyllanthus x x x x x x x x 2002 Synopsis Webster x Ganesan 2003 x x x Müller 1866 x Hayata 1904 x x Müller 1866 x x 2002 Synopsis Webster x x x x 2002 Synopsis Webster Beille 1927 x Müller 1866 2002 Synopsis Webster

x x 2002 Synopsis Ralimanana et al. 2013, Webster x x 2002 Synopsis Ralimanana et al. 2013, Webster D subg. Phyllanthus x x 2002 Synopsis Ralimanana et al. 2013, Webster America America America x America x x x Secco & De Rosário 2015 x x x 1957 Webster x 1957 Webster 1957 Webster America America America America x America x America x x America x x America 1957 Webster x x America 1957 Webster x x America 1957 Webster x x America 1957 Webster x x America & Carpenter 2008 Webster x x America 1957, sometimes referenced as Webster x x America 1957 Webster x x America 1957 Webster x x America 1957 Webster x x America 1957 Webster x x America 1957 Webster x x America 1957 Webster x x America 1957 Webster x x America & Carpenter 2008 Webster x x America 1957 Webster x x 1957 Webster x x 1957 Webster x x & Carpenter 2002 Webster x x 2002b Webster x 2002b Webster x 2002b Webster 2002b Webster America x x 2002b Webster Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Africa America x x 2002b Webster Mainland Mainland

Mainland Tropical Mainland Mainland Malesia Malesia Mainland Mainland Mainland Tropical Mainland

Mainland Mainland Mainland Mainland Mainland Mainland Mainland Mainland South Tropical South Tropical South Tropical South Tropical Mainland Malesia Mainland South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical Global area South Tropical Central Mainland Thailand Thailand, Vietnam Thailand, Vietnam Thailand Thailand Asia, Malesia Republic of Bangladesh People’s India, Myanmar, India Thailand, Vietnam

India, Myanmar Cambodia China Indonesia (Borneo incl. Sabah), Peninsular Malaysia, Philippines Marianas Myanmar, Mainland Democratic Republic, Lao People’s Cambodia Myanmar

India India Taiwan Peninsular Malaysia, Indonesia (Sumatra), Democratic China (Hainan), Indonesia (Borneo), Lao People’s Peninsular Malaysia, Republic, Myanmar, Peninsular Malaysia Mainland Vietnam India, Sri Lanka, Myanmar Venezuela Venezuela Venezuela Bhutan, China, India, Myanmar Philippines Venezuela Venezuela Venezuela Venezuela Venezuela Venezuela Venezuela Colombia, Guyana, Venezuela Venezuela Venezuela Venezuela Guyana, Venezuela Venezuela Venezuela Venezuela Guyana, Venezuela Venezuela Brazil Colombia to Peru Colombia Colombia Colombia, Ecuador Major area Colombia Costa Rica Vietnam Kunth G.L.Webster Wall. Ridl. & Kantachot Chantar. Thailand Beille Pax Steyerm. & Luteyn Jabl. C.B.Rob. Steyerm. Müll.Arg. Müll.Arg. M.G.Gilbert & P.T.Li (Müll.Arg.) Müll.Arg. Beille Jabl. (Kurz) Hook. Bennet Hook.f. Jabl. Willd. Jabl. Müll.Arg. Müll.Arg. ex Secco & G.L.Webster Brazil Miq. Jabl. Collett & Hemsl. Beille Jabl. Beille Steyerm. Jabl. Jabl. Hayata Gleason P. baeobotryoides P. albizzioides P. P. scabrifolius P.

P. angkorensis P. P. rheophyticus P. P. chamaepeuce P. P. erythrotrichus C.B.Rob. P. marianus Müll.Arg. P. P. columnaris P. emblica L. P. geoffrayi Beille P. harmandii P. P. prainianus P.

P. indofischeri P. lawii J.Graham P. niinamii P. oxyphyllus P. pachyphyllus P. pectinatus Hook.f. P. phuquocensis P. polyphyllus P. pomiferus Hook.f. P. aracaensis P. A. Rosário carrenoi Steyerm. P. chimantae Jabl. P. duidae P. jablonskianus P. P. praetervisus Müll.Arg. P. P. benguetensis P. P. chayamaritiae P. P. jauaensis P. lediformis Jabl. P. longistylus P. maguirei P. majus Steyerm. P. minutifolius P. myrsinites Kunth P. neblinae P. obfalcatus Lasser & Maguire P. paraqueensis P. pycnophyllus P. strobilaceus P. subapicalis P. tepuicola P. vacciniifolius P. ventuarii Jabl. P. websterianus P. cuatrecasasianus P. popayanensis P. ruscifolius Müll.Arg. P. sponiifolius P. symphoricarpoides P.

Species P. valerii Standl. P. arenarius P.

Urinaria Botryoides Emblica Emblica Urinaria Urinaria Urinaria Emblica Emblica Emblica Emblica Emblica Emblica Emblica Emblica Emblica Emblica Emblica Emblica Emblica Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Urinaria Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Microglochidion Pityrocladus Pityrocladus Pityrocladus Pityrocladus Pityrocladus Pityrocladus Urinaria sect. sect. sect. sect. ? sect. sect. sect. sect. sect. sect. sect.

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. ? sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. ? sect. sect. Appendix 2 (cont.) Subg. Emblica Subg. Emblica Subg. Emblica

Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica

Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica

Subgeneric placement Subg. Emblica Subg. Emblica Subg. Emblica 186 Blumea – Volume 63 / 2, 2018

Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus subg. Eriococcus subg. Eriococcus subg. Eriococcus subg. Eriococcus Phyllanthus Phyllanthus Phyllanthus Phyllanthus

Most important literature 3 L 2 M 1 x x x x x Balakrishnan & Chakrabarty 2007 x et al. 2003 Hoffmann Robinson 1909 x x Merrill 1912 x x Balakrishnan & Chakrabarty 2007 x x x x x x Airy Shaw 1971 x Airy Shaw 1969 x Li 1987a Li 1987a x x x x Ridley 1909 x Balakrishnan & Chakrabarty 2007 x x x x x Thin 2007 x Li 1987a 2002 Synopsis Webster ~ x Müller 1866 x Robinson 1911 x x x x x x Müller 1863 x x Beille 1927 x x Balakrishnan & Chakrabarty 2007 x Müller 1863 x Müller 1863 2002 Synopsis Webster x x 2002 Synopsis Webster x x x x x x x Müller 1866 x x Beille 1927 x x Beille 1927 x Gage 1914 x Müller 1863 2002 Synopsis Webster x x Balakrishnan & Chakrabarty 2007 x x Müller 1863 x x Li 1987a x x Li 1987a x x x Balakrishnan & Chakrabarty 2007 x x x x Müller 1866 Sunil et al. 2016 subg. Phyllanthus x x x 2002 Synopsis Ralimanana et al. 2013, Webster subg. Phyllanthus ~ x x x Thin 2007 x x Thin 2007 Smith 1921 et al. 2017 Wu x 2002 Synopsis Ralimanana et al. 2013, Webster subg. Phyllanthus x x 2002 Synopsis Ralimanana et al. 2013, Webster subg. Phyllanthus x x 2002 Synopsis Ralimanana et al. 2013, Webster subg. Phyllanthus D x x x Thin 2007 2002 Synopsis Ralimanana et al. 2013, Webster Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Malesia Malesia Mainland Australia, Malesia Malesia x x Müller 1866 Mainland Mainland Mainland Mainland Mainland Tropical Mainland Mainland Mainland Mainland Mainland Mainland ?Australia Malesia Malesia Mainland Mainland Mainland Malesia Mainland Mainland Malesia Mainland Mainland Mainland Mainland Mainland Malesia Mainland Mainland Mainland Mainland Mainland Malesia Mainland

Pantropical

Mainland Mainland Mainland Malesia Mainland

Australia

Malesia

Global area Mainland Pacific Thailand Taiwan Territory) Asia Thailand, Vietnam Philippines Republic of Bangladesh India, People’s Australia, Indonesia (Java) Philippines India Peninsular Malaysia Indonesia (Java, Sumatra), China China Cambodia, Vietnam India Peninsular Malaysia Hainan Democratic Republic Lao People’s Malaysia Australia (Northern Philippines Sri Lanka Vietnam India Myanmar Philippines China China Sri Lanka Myanmar, Vietnam Peninsular Malaysia India China Lesser Sunda Islands India China China Vietnam Sri Lanka

Pantropical: origin

Vietnam Vietnam China Philippines and China to Vietnam

Australia (Queensland)

Indonesia, Flores

Major area Vietnam Cook Islands, French Polynesia A.N.Henry) India A.J.M.Baker Malaysia (Borneo), Philippines Y.J.Chen P.T.Li (Sebast. & P.T.Li Airy Shaw M.Schmid Müll.Arg. Müll.Arg.

& Sunil, K.M.P.Kumar India Murugan, Kabeer & Govaerts & Radcl.-Sm. India W.W.Sm. (Miq.) Müll.Arg. Müll.Arg. C.B.Rob. Müll.Arg. Müll.Arg. Haicour & Rossignol (Gamble) M.Mohanan (Gamble) M.Mohanan André Merr. Beille Gage Airy Shaw Rehder (H.Lév.) & Petra Hoffm. P.T.Li Wall. L. Thin Thin P. balgooyi P. curranii C.B.Rob. P. lamprophyllus P. P. tetrandrus Roxb. P. P. robinsonii P. watsonii P. P. rangachariarii P. singampattianus P. talbotii Sedgw. P. P. acutissimus Miq. P. fangchengensis P. guangdongensis P. kampotensis Beille P. G.V.S.Murthy Kumari & Chandrab. P. muscosus Ridl. P. P. nanellus P. spirei Beille P. ex Müll.Arg. acutus Wall. P. armstrongii Benth. P. baillonianus P. P. macgregorii P. P. balansae P. beddomei P. birmanicus P. blancoanus P. bodinieri P. P. chekiangensis P. P. cinereus Müll.Arg. P. elegans P. evrardii Beille P. filicifolius P. fimbriatus (Wight) Müll.Arg. P. forrestii W.W.Sm. P. P. glabrescens P. P. gageanus P. P. franchetianus H.Lév. P. P. chantrieri P. fimbricalyx P.T.Li P. P. balakrishnanii P. chandrabosei P. P. anabaptizatus P. Naveen Kum.

P. urinaria P.

P. thaii P. trungii P. tsarongensis P. Ferreras & coi M.J.Wu, P. embergeri P.

& J.J.Bruhl sulcatus J.T.Hunter P.

P. submarginalis P.

P. societatis Müll.Arg. P. Species petelotii Croizat P.

subsect. Urinaria subsect. subsect. Urinaria

Emblicastrum Emblicastrum Emblicastrum Emblicastrum Emblicastrum Eriococcodes Eriococcodes Eriococcodes Eriococcodes Eriococcodes Eriococcodes Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Urinaria Urinaria Urinaria Urinaria Urinaria Urinaria Urinaria Urinaria Urinaria Urinaria sect. sect. sect. sect. sect.

sect. sect. sect. sect.

sect. sect. sect. sect. sect.

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. ? sect.

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Appendix 2 (cont.) Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus

Subg. Emblica

Subg. Emblica Subg. Emblica Subg. Emblica Subg. Emblica Benguetensis Subg. Emblica

Subg. Emblica

Subg. Emblica

Subg. Emblica Subgeneric placement Subg. Emblica R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 187

subg. Eriococcus subg. Eriococcus subg. Eriococcus subg. Eriococcus subg. Eriococcus subg. Eriococcus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 x x x x 2011 Ralimanana & Hoffmann D x x x x x Balakrishnan & Chakrabarty 2007 Airy Shaw 1975 x 2002 Synopsis Webster x x x x x x Merril 1935 x x Müller 1863 x Beille 1927 x Airy Shaw 1974, 1975 2002 Synopsis Webster x x x x Müller 1863 2002 Synopsis Webster x x x x Müller 1863 2002 Synopsis Webster x x x x x Chen et al. 2009 x Chen et al. 2009 2002 Synopsis Webster x x x x x x x x x Müller 1863 Beille 1927 x Airy Shaw 1975 x x x Beille 1927 x Beille 1927 2002 Synopsis Webster x x x x x Chen et al. 2009 x Müller 1866 x x x x Müller 1863 x Beille 1927 x Robinson 1909 Beille 1927 x x Airy Shaw 1975 x x x x x x McPherson & Schmid 1991 x x Merrill 1920 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x Guillaumin 1927 x x x Robinson 1909 x x McPherson & Schmid 1991 x x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x x x McPherson & Schmid 1991 McPherson & Schmid 1991 x 2011 Ralimanana & Hoffmann McPherson & Schmid 1991 x x x x x x Backer 1945 x Robinson 1909 x x Müller 1863 x Brunel 1987 x x Beille 1927 Beille 1927 x x x x 2011 Ralimanana & Hoffmann x x McPherson & Schmid 1991 McPherson & Schmid 1991 Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Malesia Madagascar Global area Mainland Malesia Mainland Mainland Mainland Mainland Malesia Malesia Mainland Malesia Mainland Mainland Malesia Malesia Malesia Malesia Mainland Mainland Mainland Mainland Mainland Malesia Mainland Malesia Mainland Malesia Mainland Malesia Malesia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Malesia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Madagascar New Caledonia New Caledonia Malesia Malesia Mainland Mainland Mainland Mainland New Caledonia New Caledonia Madagascar Thailand, Vietnam Thailand, Vietnam Thailand Caroline Islands Major area Democratic Republic, China, Indonesia, Lao People’s Malaysia, Philippines Peninsular Malaysia India, Myanmar, China India Vietnam Borneo Philippines China Philippines India Philippines Philippines Indonesia (Java) Sri Lanka Vietnam Democratic Republic, China, Indonesia, Lao People’s Malaysia, Cambodia, Vietnam Vietnam Philippines Nepal, Peninsular Malaysia, Bhutan, India, Myanmar, Republic of Bangladesh People’s Indonesia (Sumatra) China Philippines Cambodia, China, Indonesia (Borneo, Java) Philippines New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Philippines New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Indonesia (Celebes, Java) Philippines India Vietnam China, Vietnam Vietnam New Caledonia Madagascar Hook.f. Beille Guillaumin ex M.Schmid New Caledonia M.Schmid M.Schmid M.Schmid Guillaumin Elmer Airy Shaw C.B.Rob. M.Schmid Adelb.

(Gamble) Kumari & Chandrab. India C.B.Rob. Govaerts & Radcl.-Sm. Vietnam (Leandri ex Humbert) Merr. Craib Beille Beille Beille Benth. (Miq.) Müll.Arg. Müll.Arg. Hosok. M.Schmid Müll.Arg. Müll.Arg. C.B.Rob. M.Schmid Merr. Beille (Miq.) Müll.Arg. C.B.Rob. (Blume) Müll.Arg. Elmer M.Schmid P. palauensis P. & ankaratrae (Leandri) Petra Hoffm. P. McPherson aoupinieensis P. Madagascar P. greenei P. P. gracilipes P. Species gomphocarpus P. P. hainanensis P. heyneanus P. insulensis P. kinabaluicus P. laciniatus P. P. leptoclados P. leytensis Elmer P. P. macrocalyx Müll.Arg. P. megacarpus P. P. megalanthus P. mindorensis P. muriculatus J.J.Sm. P. P. pulchroides P. P. oreophilus P. pireyi Beille P. ex Müll.Arg. pulcher Wall. P. rubescens Beille P. rubristipulus P. P. buxifolius P. P. sibuyanensis P. sikkimensis Müll.Arg. P. singalensis P. sootepensis P. stipularis Merr. P. taxodiifolius P. trichosporus P. apiculatus P. baraouaensis P. boguenensis P. carlottae M.Schmid P. cherrieri M.Schmid P. ciliaris Baill. P. cordatulus C.B.Rob. P. dorotheae P. dumbeaensis P. favieri M.Schmid P. fractiflexus M.Schmid P. golonensis P. & McPherson humbertii (Leandri) Petra Hoffm. P. Madagascar koumacensis P. lichenisilvae P. P. triphlebius P. wightianus P. nhatrangensis P. Spreng. ruber (Lour.) P. touranensis P. & McPherson Petra Hoffm. longeramosus P. luciliae P.

subsect.

Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Eriococcus Scepasma Eriococcus Eriococcus Eriococcus Nymphanthus Nymphanthus Scepasma

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

Appendix 2 (cont.) Subg. Gomphidium Subg. Gomphidium Subg. Eriococcus Subg. Eriococcus Subgeneric placement Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Spiciferens Subg. Eriococcus Subg. Eriococcus Subg. Eriococcus Subg. Gomphidium Subg. Gomphidium 188 Blumea – Volume 63 / 2, 2018 Gomphidium Gomphidium Gomphidium Airy Shaw 1971 Most important literature 3 L 2 M 1 D x x McPherson & Schmid 1991 x x x x x 2011 Ralimanana & Hoffmann x x x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x 2011 Ralimanana & Hoffmann x x Smith 1920 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x 2011 Ralimanana & Hoffmann McPherson & Schmid 1991 x x x x ~ x McPherson & Schmid 1991 x x McPherson & Schmid 1991 2011 Ralimanana & Hoffmann x 1891 Warburg Airy Shaw 1980a x x 2001 Synopsis of Webster x x x x x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x x x McPherson & Schmid 1991 x x x McPherson & Schmid 1991 x 2001 Synopsis of Webster x x x McPherson & Schmid 1991 x Guillaumin 1927 x x McPherson & Schmid 1991 x x 2001 Synopsis of Webster x x McPherson & Schmid 1991 x x & Webster x x Chen et al. 2009 pollen x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x McPherson & Schmid 1991 Airy Shaw 1980a x x x Müller 1866, McPherson & Schmid 1991 x x Rendle et al. 1921, McPherson & Schmid 1991 x x Rendle et al. 1921, McPherson & Schmid 1991 x McPherson & Schmid 1991 x x x x x x x Rendle et al. 1921, McPherson & Schmid 1991 Rendle et al. 1921, McPherson & Schmid 1991 x x Lobreau-Callen et al. 1988 x McPherson & Schmid 1991 Global area New Caledonia Madagascar New Caledonia New Caledonia New Caledonia Madagascar Malesia New Caledonia New Caledonia New Caledonia Madagascar New Caledonia New Caledonia Pacific Madagascar Malesia Malesia Malesia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Australia New Caledonia New Caledonia New Caledonia Australia New Caledonia Malesia Malesia New Caledonia New Caledonia New Caledonia Malesia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Archipel, Papua New Guinea Major area New Caledonia New Caledonia New Caledonia New Caledonia Madagascar Moluccas New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Madagascar New Guinea Irian Jaya Papua New Guinea New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Australia (Queensland) New Caledonia New Caledonia New Caledonia Australia (Queensland) New Caledonia Irian Jaya Philippines New Caledonia New Caledonia New Caledonia Bismarck New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia M.Schmid M.Schmid Baill. M.Schmid M.Schmid Leandri M.Schmid Baill. Däniker & (Leandri) Petra Hoffm. Madagascar Airy Shaw Baill. Däniker F.Muell. S.Moore

Baill. S.Moore Guillaumin S.Moore M.Schmid Däniker Guillaumin M.Schmid Baill. S.Moore M.Schmid Guillaumin S.Moore M.Schmid Airy Shaw Airy Shaw M.Schmid Species mandjeliaensis P. P. marojejiensis P. McPherson memaoyaensis P. meuieensis P. nitens M.Schmid P. oreichtitus Leandri P. ovatifolius J.J.Sm. P. parainduratus P. parangoyensis P. peltatus Guillaumin P. & McPherson perrieri (Leandri) Petra Hoffm. P. Madagascar pronyensis P. pseudotrichopodus P. P. rozennae P. sambiranensis P. sessilis Warb. P. stultitiae P. tabularis P. P. tixieri M.Schmid P. trichopodus P. umbricola P. unifoliatus P. valeriae P. virgultiramus P. vulcani Guillaumin P. yvettae M.Schmid P. baladensis P. aeneus Baill. P. artensis M.Schmid P. brassii C.T.White P. deciduiramus P. francii Guillaumin P. guillauminii P. hypospodius P. ligustrifolius P. maritimus J.J.Sm. P. ramosii Quisumb. & Merr. P. serpentinus P. sylvincola P. tireliae M.Schmid P. verrucicaulis P. bupleuroides P. P. casearioides P. comptonii S.Moore P. conjugatus P. dracunculoides P. P. gneissicus P. loranthoides P. macrochorion P.

Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Appendix 2 (cont.) Subgeneric placement Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 189 Most important literature 3 L 2 M 1 D x x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x Rendle et al. 1921 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x Müller 1866, McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x x x Airy Shaw 1980a x McPherson & Schmid 1991 Airy Shaw 1980a x x McPherson & Schmid 1991 x x x x x x x x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x Rendle et al. 1921, McPherson & Schmid 1991 x McPherson & Schmid 1991 x McPherson & Schmid 1991 x McPherson & Schmid 1991 McPherson & Schmid 1991 x x Airy Shaw 1980a x x x x x Chen et al. 2009 pollen x McPherson & Schmid 1991 1986 Webster x x x x x x x McPherson & Schmid 1991 McPherson & Schmid 1991 McPherson & Schmid 1991 x x x x x x x Airy Shaw 1980a x x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x x Müller 1866 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x McPherson & Schmid 1991 Rendle et al. 1921, McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x x x 1986 Webster McPherson & Schmid 1991 America x x & Carpenter 2008 1967a. Webster Webster Global area New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Central Malesia New Caledonia Malesia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Malesia Malesia Malesia New Caledonia Pacific New Caledonia New Caledonia New Caledonia Malesia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Malesia Australia Major area New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Guatemala New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Irian Jaya Papua New Guinea Philippines New Caledonia Fiji New Caledonia New Caledonia New Caledonia Irian Jaya New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Caroline Isl. Australia (Queensland) Baill. M.Schmid Airy Shaw M.Schmid

M.Schmid Guillaumin M.Schmid M.Schmid Baill. M.Schmid M.Schmid Airy Shaw Elmer Airy Shaw & G.L.Webster Papua New Guinea M.Schmid Hosok. Airy Shaw & G.L.Webster Irian Jaya M.Schmid Guillaumin Müll.Arg. S.Moore

G.L.Webster S.Moore Schltr. Guillaumin M.Schmid M.Schmid Baill. Guillaumin M.Schmid Baill. Baill. M.Schmid Däniker Guillaumin M.Schmid Guillaumin M.Schmid Airy Shaw P. montrouzieri P. moratii M.Schmid P. ouveanus P. rhodocladus P. salicifolius P. sarasinii P. torrentium Müll.Arg. P. vespertilio P. yaouhensis P. tuerckheimii P. acinacifolius P. amieuensis P. ardisianthus P. balansaeanus P. Species P. margaretae M.Schmid P. P. bourgeoisii P. P. buxoides P. calcicola P. castus S.Moore P. caudatus Müll.Arg. P. chamaecerasus P. cornutus Baill. P. dzumacensis P. frodinii P. P. glochidioides P. P. helenae P. heterodoxus P. houailouensis P. P. jaffrei M.Schmid P. koniamboensis P. kostermansii P. P. kouaouaensis P. macphersonii P. mangenotii P. montis - fontium M.Schmid P. moorei M.Schmid P. mouensis M.Schmid P. ningaensis P. pancherianus P. paucitepalus P. petchikaraensis P. pinjenensis P. polygynus P. poueboensis P. poumensis P. pterocladus P. quintuplinervis P. P. rupi - insularis P. P. sauropodoides P.

Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Adenoglochidion Calodictyon Gomphidium Gomphidium Gomphidium Gomphidium Adenoglochidion Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Appendix 2 (cont.) subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium subsect. Eleutherogynium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subgeneric placement Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium 190 Blumea – Volume 63 / 2, 2018 Airy Shaw 1971 Airy Shaw 1971 Most important literature 3 L 2 M 1 D x x x x x x Chen et al. 2009 pollen x 1986 Webster x McPherson & Schmid 1991 McPherson & Schmid 1991 x x x x x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x McPherson & Schmid 1991 x x McPherson & Schmid 1991 McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x McPherson & Schmid 1991 x x x McPherson & Schmid 1991 x x x x 1986 Guillaumin 1937, but see Webster x x & Carpenter 2008 Webster x Airy Shaw 1980a x 1986 Webster x Airy Shaw 1980a x x x x Airy Shaw 1980a x Airy Shaw 1980a Airy Shaw 1980a x x x x x x Smith 1912 x x Airy Shaw 1980a x x Airy Shaw 1980a x Airy Shaw 1980a x x 1986 Webster x Airy Shaw 1980a x x x x Airy Shaw 1980a x x Airy Shaw 1980a x x Airy Shaw 1980a x x Smith 1912 x x Bouman et al. 2018 x Airy Shaw 1980a x Airy Shaw 1980a Airy Shaw 1980a x Merril 1926 x x x x x Airy Shaw 1980a x Airy Shaw 1980a 1986 Webster Asia x x x 2011 Ralimanana & Hoffmann Africa, Global area Malesia Malesia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Pacific New Caledonia Malesia Pacific Malesia Australia, Malesia Australia, Malesia Malesia x Malesia Malesia x x x x Smith 1912 & Webster New Caledonia Malesia Malesia Malesia Malesia Malesia Australia, Malesia Malesia Malesia Malesia Malesia x Malesia Malesia x Malesia & Webster Tropical Malesia Malesia Malesia Malesia Malesia Malesia Major area Philippines Fiji New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia New Caledonia Vanuatu New Caledonia Irian Jaya Tonga Australia (Queensland), Papua New Guinea Australia (Queensland), Papua New Guinea Papua New Guinea Papua New Guinea Papua New Guinea New Caledonia Papua New Guinea Papua New Guinea Irian Jaya Fiji Irian Jaya Papua New Guinea Papua New Guinea Papua New Guinea Papua New Guinea Papua New Guinea Papua New Guinea Palaeotropics Papua New Guinea Philippines Papua New Guinea Fiji L. M.Schmid Merr. Guillaumin Diels Airy Shaw Airy Shaw & G.L.Webster Papua New Guinea J.J.Sm. Airy Shaw & G.L.Webster Australia (Queensland), Papua New Guinea

M.Schmid M.Schmid

& Ent R.W.Bouman Welzen, Malaysia (Borneo) Airy Shaw Guillaumin Merr. J.J.Sm.

Airy Shaw & G.L.Webster Papua New Guinea Müll.Arg. Diels (F.Muell.) M.Schmid

M.Schmid Airy Shaw Guillaumin Baill. Airy Shaw

Gillespie

Airy Shaw & G.L.Webster Papua New Guinea Airy Shaw Gage Airy Shaw & G.L.Webster M.Schmid

J.J.Sm. Airy Shaw & G.L.Webster ex Guillaumin Vieill. Airy Shaw & G.L.Webster M.Schmid

Species P. securinegoides P. P. smithianus G.L.Webster P. stenophyllus P. tangoensis P. tenuipedicellatus P. P. tiebaghiensis P. tritepalus M.Schmid P. unioensis P. veillonii P. avanguiensis P. faguetii Baill. P. jaubertii P. koghiensis P. natoensis M.Schmid P. nothisii M.Schmid P. pilifer M.Schmid P. pindaiensis P. platycalyx Müll.Arg. P. stipitatus M.Schmid P. fimbriatitepalus P. kanalensis P. actephilifolius P. amicorum G.L.Webster P. aphanostylus P. clamboides P. cuscutiflorus S.Moore P. effusus S.Moore P. finschii K.Schum. P. flaviflorus P. gjellerupi P. P. maderaspatensis P. P. insulae - japen P. merinthopodus P. papuanus P. pergracilis P. poli - borealis P. praelongipes P. pullenii P. rheophilus P. rosselensis P. rubriflorus J.J.Sm. P. rufuschaneyi P. salomonis P. tagulae P. tenuirhachis P. utricularis P. P. caudatifolius P. P. warburgii P. wilkesianus P.

Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Gomphidium Leptonema Leptonema Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania Nymania sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. ?

Appendix 2 (cont.) Subgeneric placement Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium subsect. Physoglochidion Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Gomphidium Subg. Kirganelia Subg. Gomphidium Subg. Gomphidium Subg. Isocladus R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 191

sect. , : probably

Kirganelia , subg. Kirganelia subg. Kirganelia subg. Kirganelia subg. Kirganelia subg. Kirganelia subg. Kirganelia subg. Kirganelia

subg.

2011: Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus

P. wildemannii P. Hoffmann

&

Airy Shaw 1971 Airy Shaw 1971 Airy Shaw 1971 Airy Shaw 1971 2015 Telford Anisonema Flueggeopsis Most important literature 3 L 2 M 1 D x x 1891 Warburg x x x Brunel 1987 x x 2002 Synopsis Webster x x x x x x 1996 Radcliffe-Smith x x Li 1987 x x Brunel 1987a & Webster Barrett & x x x x x x x x x & Webster Müller 1866 x x x x Müller 1863 x 2002 Synopsis Webster x 2002 Synopsis Webster 2002 Synopsis Webster x x Brunel 1987 x x Meeuwis & Punt 1983 x x 2002 Synopsis Webster x x x Brunel 1987 x x Brunel 1987 x x Challen et al. 2011 x x Brunel 1987 x x Brunel 1987 ~ x Airy Shaw 1971 x x x Ralimanana x x Brunel 1987 x x Brunel 1987, but see Breteler 2012 x x x 1957 Webster x x Brunel 1987, but check x x Brunel 1987 x x Müller 1866 x x x x 2001b Webster x x x 2002 Synopsis Webster 2002 Synopsis Webster but see Brunel 1987 Tropical x x 2001b Webster America x x Rusby 1927 America x x Rusby 1927 America x x x 2001b Webster Asia, Malesia x x x Airy Shaw 1975, see Luo et al. 2011 Asia, Malesia x x x Luo et al. 2011 Asia Asia Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Africa Indies America, West America, Tropical Global area Malesia Malesia Tropical Tropical Madagascar Tropical Malesia Madagascar Malesia Australia Malesia Madagascar Australia, Malesia New Caledonia Madagascar Malesia Madagascar Mainland x x & Webster Madagascar Tropical Australia, Malesia x x & Webster Tropical Tropical Mainland Madagascar Tropical Tropical Tropical South Tropical Tropical South Tropical South Tropical North South Pantropical Indies West Mainland Africa, Tanzania, Tropical Africa, Zambia Tobago Africa, Uganda, Yemen & Trinidad Togo Thailand Zambia Tanzania, African Republic, Congo, Zambia, Zimbabwe Tanzania, Zambia Tanzania, Tanzania, Tanzania Tanzania Thailand Africa Cameroon, Congo, Gabon, Major area Philippines Papua New Guinea Zambia Papua New Guinea, Bhutan, China, India, Indonesia, Myanmar, Mainland Ghana, Kenya, Mozambique, South Republic of Bangladesh, Peninsular Malaysia, People’s Philippines, Madagascar Cameroon, Democratic Republic of the Congo, Gabon, Taiwan Madagascar Australia Papua New Guinea Madagascar Australia, Papua New Guinea New Caledonia Madagascar Indonesia (Sumatra) Madagascar India, Indonesia, Myanmar, United Republic of Republic of Equatorial Guinea Angola, Benin, Cameroon, Democratic Republic of the Congo, Tropical Benin, Cameroon, Congo, Democratic Republic of the Tropical Madagascar Cameroon Gabon, Guinea, Ivory Coast, Liberia, Nigeria, South United Republic of Australia (Queensland), Papua New Guinea Angola, Cameroon,Democratic Republic of the Congo, Ethiopia, Tropical Gabon, United Republic of South Kenya, Malawi, Mozambique, Rwanda, Uganda, United Republic of United Republic of Thailand Comoros, Madagascar Cameroon, Central Kenya, Malawi, Mozambique, United Republic of Democratic Republic of the Congo, South Cameroon, Ghana, Ivory Coast, Liberia Zimbabwe United Republic of Bolivia Angola, Namibia Bolivia Bolivia Argentina, Brazil, Paraguay, Colombia, Mexico, Guyana,

Pantropical Cuba Müll.Arg. Airy Shaw Airy Shaw & G.L.Webster Papua New Guinea

Rusby Hayata Schumach. (Kuntze) Exell Müll.Arg. Griseb. Müll.Arg. (Baill.) Müll.Arg. Verdc. Leandri Baill. Morong Müll.Arg. Pax Mansf. ex Radcl.-Sm. Müll.Arg. Forssk. (Miq.) Müll.Arg. Rusby (Craib) Pax Airy Shaw & G.L.Webster Pax P. polyspermus P. P. keyensis Warb. P. martinii Radcl.-Sm. P. Species dumetosus Poir. P. P. reticulatus Poir. P. P. inflatus Hutch. P.

P. seyrigii Leandri P. dinklagei P. P. oligospermus P. angavensis P. archboldianus P. ex Benth. baccatus F.Muell. P. P. caesius P. casticum P.Willemet P. ciccoides Müll.Arg. P. deplanchei P. fuscoluridus P. glaucinus P. matitanensis Leandri P. microcarpus (Benth.) Müll.Arg. P. P. muellerianus P.

P. physocarpus P. P. kidna Challen & Petra Hoffm. P. P. coluteoides P. novae - hollandiae P. ovalifolius P. cedrelifolius P. pervilleanus P. P. engleri P. P. orientalis P. P. polyanthus P. P. pinnatus (Wight) G.L.Webster P. profusus N.E.Br. P. schliebenii P.

P. ibonensis P. P. pseudonobilis P. P. purpureus P. P. chacoensis P. P. elsiae Urb. P.

P. acidus Skeels P. P. pseudocicca P. flexuosus (Siebold & Zucc.) Müll.Arg. P. China, Japan

Anisonema Omphacodopsis Anisonema Anisonema Brazzeani Anisonema Anisonema Anisonema Anisonema Anisonema Anisonema Anisonema Anisonema Anisonema Anisonema Anisonema Anisonema Omphacodopsis Chorisandra Anisonema Anisonema Polyanthi Chorisandra Anisonema Polyanthi Chorisandra Polyanthi Polyanthi Polyanthi Cicca Cicca Pseudomenarda Cicca subsect. Cicca subsect. Cicca subsect. Ciccopsis Hemicicca sect. Chorisandra sect. sect.

? sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

Appendix 2 (cont.) Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subgeneric placement Subg. Kirganelia Subg. Kirganelia

Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Subg. Kirganelia Aporosella Subg. Kirganelia Subg. Kirganelia Aporosella Cheramela Subg. Kirganelia Subg. Kirganelia 192 Blumea – Volume 63 / 2, 2018 Airy Shaw 1971 2015 Telford Most important literature 3 L 2 M 1 x x x x x x & Webster x Ralimanana et al. 2014 x Ralimanana et al. 2014 Schweinfurth 1899 x x x 1995 Webster x x x x x x Müller 1866 Li 1987b x Li 1987b Li 1987b x x x Lobreau-Callen et al. 1988 x x x x Li 1987b Beille 1927 x x x x x x Airy Shaw 1960 x Croizat 1942 x Croizat 1942 Croizat 1942 x x x Brunel 1987 x x & Carpenter 2008 Croizat 1942, but see Webster x x Brunel 1987 x Thin 2007 x et al. 2016 Robinson 1909, Wu x x x x x x Croizat 1942 x Pornpongrungrueng et al. 2017 x Hemsley 1898 x x Croizat 1942 x x Croizat 1942 x x & Carpenter 2008 Croizat 1942, but see Webster x x Croizat 1942 x Li 1987b x Chakrabarty & Gangopadhyay 1993 x x Chen et al. 2009 x x Hunter & Bruhl 1997a et al. 2016 Merrill 1920, Wu x x Punt 1980 x x x Brunel 1987 Balakrishnan & Chakrabarty 2007 x x Barrett & x x Müller 1866 x x Chen et al. 2009 x x x Robinson 1909 Robinson 1909 x x 1986 Webster x x x Luo et al. 2011 x x x Lobreau-Callen et al. 1988 D x & Lorence 2011 Wagner x x Brunel 1987 x x Brunel 1987 America America x x 1858 Visiani Rusby 1912 Asia Asia Asia Asia, Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Malesia Madagascar Madagascar Middle east Malesia Madagascar Mainland Mainland Mainland Mainland Mainland Tropical Australia, Pacific Mainland Mainland Mainland Mainland Mainland Indies West Mainland Malesia Mainland Mainland Mainland Mainland Mainland Mainland Mainland South Tropical South Tropical Mainland Malesia Australia Malesia Pacific Mainland Malesia Australia Mainland Malesia Australia Malesia Malesia Pacific Mainland New Caledonia Global area Pacific Tropical Thailand Thailand, Vietnam Mainland Thailand, Vietnam Thailand, Vietnam Thailand Thailand, Vietnam Territory) Tahiti Madagascar Yemen Sri Lanka Madagascar China New Caledonia, Democratic Republic, Myanmar, Lao People’s Mainland Vietnam Cambodia Australia, China, Fiji, India, Indonesia (Borneo), Papua New Guinea, Democratic Republic, Lao People’s Vietnam Peninsular Malaysia, China, Japan, Korea, Russia South China Moorea, Raiatea, Vietnam Philippines Democratic Republic, China, Lao People’s Vietnam Democratic Republic, China, Lao People’s China Venezuela Bolivia India Philippines Australia Philippines Marquesas India Sri Lanka Australia India Philippines Australia (Northern Philippines Philippines Hawaii Nepal, China, India, Myanmar, New Caledonia Major area Tahiti Benin, Ivory Coast, Liberia, Republic of Equatorial Guinea, Tropical Kenya, Somalia Sierra Leone A.Juss.) Müll.Arg. Madagascar Arn. Hand.-Mazz. Gamble Müll.Arg. Govaerts & Radcl.-Sm. (P.T.Li) China Thin

Govaerts & Radcl.-Sm. China & J.J.Bruhl J.T.Hunter (Croizat) Govaerts & Radcl.-Sm. China (Comm. ex Govaerts & Radcl.-Sm. Airy Shaw & G.L.Webster Papua New Guinea Govaerts & Radcl.-Sm. (P.T.Li) China Merr. Schweinf.

& Maxim. Rupr. Hutch. Merr. Beille & J.Parn. Pornp., Chantar. Thailand Airy Shaw Rusby Hand.-Mazz. ex Rehder (H.Lév.) China Chakrab. & M.Gangop.

Nadeaud Beille (Croizat) Govaerts & Radcl.-Sm. China G.L.Webster Müll.Arg. Beille P. womersleyi P. P. coodei Ralim. & Petra Hoffm. P. cryptophilus P. hodjelensis P. zygophylloides P. P. wheeleri P. breynioides P. guangxiensis P. lii Govaerts & Radcl.-Sm. P. orbicularifolius P. rubicundus P. arachnodes P. P. virgatus G.Forst. P. P. lingulatus P. poilanei P. ridleyanus P. P. ussuriensis P. P. anthopotamicus P. P. urceolatus Baill. P. P. dongmoensis P. P. tenuipes C.B.Rob. P. P. dunnianus P. P. kaweesakii P. mirabilis P. carinatus Beille P. lativenius P. roseus (Craib & Hutch.) Beille P. yunnanensis P. moi P.T.Li P. bicolor Vis. P. cassioides P. P. sanjappae P. P. samarensis Müll.Arg. P. P. prominulatus P. P. panayensis P. P. narayanswamii P. pacificus Müll.Arg. P. P. myrtifolius (Wight) Müll.Arg. P. ex Müll.Arg. minutiflorus F.Muell. P. P. macraei Müll.Arg. P. P. lanceifolius P. P. exilis S.Moore P. P. everettii C.B.Rob. P. P. dumosus C.B.Rob. P. P. distichus Hook. & P. P. clarkei Hook.f. P. P. chrysanthus Baill. P. Species P. aoraiensis P. P. alpestris Beille P. P. somalensis P.

Arachnodes Arachnodes Arachnodes Arachnodes Calophyllum Calophyllum Calophyllum Tetrandrum

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Pseudomenarda

sect. ? ?

?

Appendix 2 (cont.) Subg. Menarda Subg. Menarda Subg. Menarda Subg. Menarda Subg. Macraea Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Macraea Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Macraea Subg. Phyllanthodendron Subg. Macraea Subg. Phyllanthodendron Subg. Macraea Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Phyllanthodendron Phyllanthodendron Phyllanthodendron Pseudoactephila Pseudoactephila Pseudoactephila Pseudoactephila Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthodendron Subg. Phyllanthodendron Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subg. Macraea Subgeneric placement Subg. Kirganelia Subg. Macraea Subg. Macraea Subg. Macraea R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 193 Lysiandra subg. Phyllanthus subg. Phyllanthus Phyllanthus Phyllanthus Phyllanthus ) Phyllanthus Phyllanthus Australian Australian Australian ( Euphorbiaceae 2015 Telford 2015 Telford 2015 Telford 2015 Telford Lysiandra Phyllanthus Lysiandra Lysiandra Most important literature 3 L 2 M 1 x x Barrett & x x 1978 Webster x x x x x Hunter & Bruhl 1997b x x 2002 Synopsis Webster 2002b Webster x x Barrett & x x Outline of Webster x x Barrett & x x x x Manuscript enumeration australian taxa: Webster x x x x x Airy Shaw 1980b x Airy Shaw 1980b Bentham 1873 x x x x 1978 Webster Barrett & x x x Moore 1920 Outline of Webster x x Outline of Webster D

x x 1955 Webster ~ Greenman 1898 x x x x 1970 Webster 1956 Webster x x x 2002b Webster x x x 1956 Webster America x America America America x x Müller 1873 Parodi 1881 Müller 1873 America America ~ America x America x x Müller 1863, 1866 America Santiago et al. 2006 x x America x Santiago et al. 2006 x x America x Santiago et al. 2006 x America x Santiago et al. 2006 x America Santiago et al. 2006 x x America Santiago et al. 2006 x x America Santiago et al. 2006 x x America Santiago et al. 2006 x x 2002 Synopsis Webster x x Santiago et al. 2006 x 2002 Outline of the Neotropical infrageneric taxa Webster America America America America America Australia North Australia Australia Australia Australia Australia Australia Australia Australia Australia Australia Australia Australia Australia Australia Australia Australia Global area Central to North South Tropical South Tropical South Tropical

North South Tropical North South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical North Indies West Australia Australia Indies West Australia Territory) Territory) Queensland) Territory, Territory) Australia) Australia) Australia) Australia (coastal areas of the Carnarvon bioregion) Mexico Australia (Queensland) Australia (Queensland) Australia Australia (Queensland, New South Wales) Australia Australia (East and Southeast) Australia (Western Australia (Queensland) Australia Australia (Queensland) Australia (Southeast Queensland and East New South Wales) Australia Australia Australia (Queensland) Australia (Western Australia (Northern Australia (Northern Australia (Queensland) Major area Brazil, Costa Rica, Guatemala, Honduras, Mexico, Nicaragua, Panama, Peru, Venezuela Brazil Paraguay Brazil

Mexico Brazil Mexico Brazil Brazil Brazil Brazil Brazil, Guyana Brazil Brazil Brazil Brazil Brazil Argentina, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, French Guiana, Guyana, Mexico, Nicaragua, El Salvador, North, Central and South Tropical x x x 1955 Webster Peru, Surinam, United States, Venezuela Panama, Paraguay, Mexico Cuba Australia (Northern Australia (Northern Colombia, Costa Rica, Dominican Republic, Surinam, Venezuela Australia (Western Govaerts & Radcl.-Sm. Brazil Kunth. Parodi Arg. Müll.Arg. Benth. Müll.Arg. Benth. Müll.Arg. & J.J.Bruhl J.T.Hunter Müll.Arg. Müll.Arg. Müll.Arg. Walter Müll.Arg. Brandegee Domin & J.J.Bruhl J.T.Hunter S.Moore Müll.Arg. ex Griseb. C.Wright Pax & K.Hoffm. L.J.M.Santiago Airy Shaw Airy Shaw F.Muell.

Millsp. G.L.Webster L.J.M.Santiago Müll.Arg. Benth. Müll Müll.Arg. R.L.Barrett & I.Telford Benth. Domin P. hamelinii P. P. harrimanii P. indigoferoides P. P. lacerosus P. occidentalis P. savannicola P. saxosus F.Muell. P. similis Müll.Arg. P. P. fuernrohrii P. P. gunni Hook.f. P. & J.J.Bruhl involutus J.T.Hunter P. P. lacunellus P. mitchellii P. microcladus P. P. dallachyanus P. eremicus R.L.Barrett & I.Telford P. P. eutaxioides P. flagellaris P. P. collinus P. leptocaulos P. paraguayensis P. pohlianus P. simplicicaulis P. Species compressus Kunth P.

P. brandegei P. caroliniensis P. P. subcuneatus Greenm. P. angustissimus P. edmundoi P. flagelliformis P. gladiatus P. klotzschianus P. scoparius Müll.Arg. P. choretroides P. goianensis P. sarothamnoides P. spartioides P. avicularis P.

evanescens P. heliotropus P. hyssopifolioides P. P. carpentariae P. & J.J.Bruhl cauticola J.T.Hunter P. baeckeoides P.

? ?

? ?

Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Lysiandra Loxopodium Loxopodium Choretropsis Choretropsis Choretropsis Choretropsis Choretropsis Choretropsis Choretropsis Choretropsis Choretropsis Choretropsis Loxopodium Loxopodium Loxopodium Loxopodium Lysiandra Lysiandra Lysiandra Lysiandra

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Appendix 2 (cont.) Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subgeneric placement Subg. Phyllanthus

Subg. Phyllanthus Subg. Phyllanthus Subg. phyllanthus sect. Subg. Phyllanthus Subg. Phyllanthus subsect. Applanata Subg. Phyllanthus subsect. Applanata Subg. Phyllanthus subsect. Applanata Subg. Phyllanthus subsect. Applanata Subg. Phyllanthus subsect. Applanata Subg. Phyllanthus subsect. Applanata Subg. Phyllanthus subsect. Choretropsis Subg. Phyllanthus subsect. Choretropsis Subg. Phyllanthus subsect. Choretropsis Subg. Phyllanthus subsect. Choretropsis Subg. Phyllanthus

Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus 194 Blumea – Volume 63 / 2, 2018 Most important literature 3 L 2 M 1 D x x Mueller 1859 Von x x 1957 Webster America America America America America x ~ America x x America Ribeiro et al. 2017 x x x Funez & Hassemer 2017 America Funez et al. 2018 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America x 2002a Webster x x America x 2002a Webster x America x 2002a Webster x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America 2004 Cordeiro & Carneiro-Torres x x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America 2002a Webster x x America Silva & Sales 2008 x x America 2002a Webster x x America 2002a Webster x x 2002a Webster x x 2002 outline neotropical taxa Webster x Silva 2009 South Tropical Global area South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical Indies West North, Central and x x x 2002a Webster Australia America Tobago

Major area Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Brazil Southeast Brazil Bolivia Brazil & Trinidad North and South Australia (Queensland, New South Wales) M.J.Silva Müll.Arg. F.Muell. Müll.Arg. G.L.Webster & Loiola R.T.M.Ribeiro Brazil Müll.Arg. Cordeiro & Carn.-Torres Brazil Müll.Arg. Müll.Arg. G.L.Webster G.L.Webster G.L.Webster Brade Pax & K.Hoffm. Müll.Arg. Müll.Arg. Baill. Casar. G.L.Webster ex Müll.Arg. Wall. G.L.Webster G.L.Webster

P. carmenluciae P. eremitus Funez & Hassemer P. & Hassemer timboensis Funez, J.P.R.Ferreira P. Brazil almadensis P. ex Spreng. acutifolius Poir. P. allemii P. arenicola P. atalaiensis P. blanchetianus P. bradeanus P. caparaoensis P. carvalhoi P. claussenii P. fastigiatus Mart. ex Müll.Arg. P. glaziovii P. gongyloides P. heteradenius P. hypoleucus P. itatiaiensis P. lagoensis P. mocotensis G.L.Webster P. piranii P. retroflexus Brade P. sincorensis P. subemarginatus P. augustinii P. bolivianus P. longipedicellatus P. mimicus G.L.Webster P. niruri L. P. Species subcrenulatus P. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect.

Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Lysiandra sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

Appendix 2 (cont.) Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Almadanses Subg. Phyllanthus Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Clausseniani Niruri Subg. Phyllanthus Niruri Subg. Phyllanthus Niruri Subg. Phyllanthus Niruri Subg. Phyllanthus Niruri Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subg. Phyllanthus Subgeneric placement Subg. Phyllanthus R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 195 subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus subg. Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 D x x x 1955, 1957 Webster x x 1957 Webster x x 1957 Webster x x 1957 Webster x x 1957 Webster x x 1955, 1957 Webster x x 1955, 1957 Webster x x 1957 Webster x x 1957 Webster x x 1957 Webster x 1955, 1957 Webster x x x x 1957 Webster x x x 1957 Webster x x x x x Ralimanana et al. 2013 x x Ralimanana et al. 2013 x x 2002 Synopsis Webster x 1957 Webster x 2002 Synopsis Webster 2002 Synopsis Webster x x x x x 2002 Synopsis Webster x 2002 Synopsis Webster x 2007 Webster x 1982 Radcliffe-Smith x 1996 Radcliffe-Smith x Gandoger 1919 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 x x Brunel 1987 ~ x x x Brunel 1987 2002 Synopsis Webster x x Brunel 1987 America x x 2002a Webster America America x x 2002 Synopsis Webster America America America x x America x x x America 2002 Synopsis Webster x 2002 Synopsis Webster x 2002 Synopsis Webster x x 2002 Synopsis Webster x Croizat 1945 Asia Africa Africa Africa Africa Africa Africa Africa Africa Africa America America America America America Global area South Tropical Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West North, Central and South Tropical Indies West Indies West x North x 1955, 1957 Webster Pantropical Indies West Indies West North South Tropical North South Tropical South Tropical South Tropical Indies West North South Tropical South Tropical North Tropical Tropical Tropical Tropical Tropical Tropical Tropical Tropical Madagascar Tropical Africa, Venezuela Tobago, Asia, naturalised elsewhere Mainland Africa & Trinidad Africa to Tanzania Zambia, Zimbabwe Tanzania, Tropical Major area Brazil Dominican Republic Haiti Dominican Republic, Haiti Cuba Cuba Dominican Republic, Haiti Cuba Cuba Dominican Republic, Haiti Cuba Cuba Caribbean, Florida, Venezuela Cuba Cuba Mexico, Paraguay, Argentina, Brazil, Colombia, Ecuador, Peru, United States, Venezuela Pantropical South to Tropical Brazil, Colombia, Mexico, Jamaica Widespread in Mexico Brazil x x Brunel 1987 United States Brazil, Colombia, Surinam, Venezuela Bolivia, Brazil, Colombia, Venezuela Brazil, Colombia, Venezuela Cuba Mexico Guyana, Bolivia, Brazil, Colombia, Costa Rica, Ecuador, Nicaragua, Venezuela Colombia United States United Republic of Mozambique Angola, Botswana, Malawi, Mozambique, South United Republic of Mozambique, Zimbabwe Angola, Mozambique, South Mozambique Angola Madagascar Angola, Botswana, Namibia, Zambia, Zimbabwe Thonn. Gand. C.Wright Urb. Jean F.Brunel Urb. McVaugh ex Griseb. C.Wright ex Radcl.-Sm. Jean F.Brunel Mozambique Kunth Müll.Arg. Urb. Croizat A.Rich. Hutch. Baill. Müll.Arg. Welw. Britton & P.Wilson Urb. G.L.Webster G.L.Webster Müll.Arg. Jean F.Brunel Species P. perpusillus P. P. amnicola P. brachyphyllus P. buchii Urb. P. dimorphus P. echinospermus P. fuertesii Urb. P. imbricatus G.L.Webster P. junceus Müll.Arg. P. leptoneurus P. maestrensis Urb. P. micranthus P. pentaphyllus P. pulverulentus P. selbyi Britton & P.Wilson P. fluitans Benth. ex Müll.Arg. P. amarus Schumach. & P. caribaeus P. fadyenii Urb. P. fraternus G.L.Webster P. hexadactylus P. leptophyllus P. lindbergii P. P. abnormis Baill. P. P. microphyllus P. minutulus Müll.Arg. P. procerus C.Wright P. standleyi McVaugh P. stipulatus (Raf.) G.L.Webster P. vichadensis P. warnockii P. frazieri Radcl.-Sm. P. P. manicaensis P. P. mozambicensis P. P. angolensis P.

P. graminicola P. P. loandensis P. P. tsetserrae Jean F.Brunel P. P. cocumbiensis P. P. geayi Leandri ex Humbert P. mendesii P.

subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect. subsect.

Loandani Loandani Loandani Loandani Pentandra Pentandra Pentandra Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Phyllanthus Salviniopsis Reverchonia ? ? ? sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

? sect.

Appendix 2 (cont.) Subgeneric placement Subg. Phyllanthus Niruri Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Pentaphylli Subg. Phyllanthus Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Swartziani Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus 196 Blumea – Volume 63 / 2, 2018 subg. Xylophylla subg. Xylophylla subg. Xylophylla subg. Phyllanthus ) ) Phyllanthus Phyllanthus Phyllanthus Phyllanthus

Synopsis

( Euphorbiaceae ( Euphorbiaceae 2002

Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 D

x x x x x x x Müller 1866 x x x Müller 1866 x Müller 1866 x 1958 Webster x x 2002 Synopsis of Webster 2002 Synopsis of Webster x 2002 Synopsis of Webster x x x 2011 Ralimanana & Hoffmann x x Brunel 1987 x x Webster x x x 2011 Ralimanana & Hoffmann x x x 2011 Ralimanana & Hoffman x x 2001b Webster x x 1958 Webster x x x 1958 Webster x x x x x Croizat 1943 Croizat 1943 x 1958 Webster x x x x x x x 1958 Webster Croizat 1943 1958 Webster x x x 1958 Webster x 1958 Webster America America 2002 Outline of the Neotropical infrageneric taxa Webster x x Müller 1873 America America America x 1958 Webster America x x Brunel 1987 America America ~ x x Jablonski 1967 1958 Webster America America x America x America x America x x x 1958 Webster x 2001b Webster x x x 2001b Webster x 2001b Webster 2001b Webster America America x America x x x 1958 Webster x Müller 1873 x 1958 Webster America America x x x 2001b Webster x Silva & Sales 2006 America America x x x Croizat 1943 x 2001a Webster America America America America x ~ America x x x America Müller 1866 x America x x 1958 Webster 1958 Webster x x x 1958 Webster x x 1967b Webster 2002 Outline of the Neotropical infrageneric taxa Webster Africa Africa Africa

America America x ~ America America America America America America America America Global area

South Tropical Indies West Indies West Indies West Indies West Indies West South Tropical Indies West Indies West Indian Ocean South Tropical South Tropical Central Central Pantropical South Tropical South Tropical South Tropical South Tropical South Tropical South Tropical North South Tropical South Tropical South Tropical North South Tropical South Tropical North Indies West South Tropical South Tropical North North South Tropical North South Tropical Central and North America South Tropical Indies West North Indies West x x Croizat 1943 South Tropical South Tropical South Tropical South Tropical North South Tropical Indies West South Tropical South Tropical Africa Tropical Africa, Tropical Tanzania Tanzania America Benin, Botswana, Burkina Faso, Namibia, Nigeria, South Major area

Venezuela Jamaica Jamaica Cuba Tobago Dominican Republic, Haiti Brazil Dominica, Jamaica, Martinique Jamaica United Republic of Botswana, Democratic Republic of the Congo, Namibia, South Comoros Mozambique, Gabon, Ghana, Guinea, Liberia, Madagascar, Tropical Republic of Equatorial Guinea, United Brazil Venezuela Guatemala Guatemala Pantropical Uruguay Argentina, Brazil, Paraguay, Guyana, Surinam Colombia, Venezuela Panama Tropical Brazil, Colombia, Guyana, Venezuela Mexico Venezuela Brazil, Colombia, Venezuela Brazil Mexico Brazil Brazil Mexico: Veracruz Windward Islands Brazil Colombia, Mexico, Peru, Venezuela Mexico Mexico Brazil Mexico Peru Guatemala, Mexico Colombia Cuba, Jamaica Mexico Cuba Brazil, Peru Bolivia Peru, Venezuela Brazil Mexico Brazil Haiti Peru Peru Thonn. Arg Arg. Poir. Croizat Steyerm. Jabl. (Müll.Arg.) Müll.Arg. Willd. Arg. Klotzsch Müll. Standl. ex Croizat Müll.Arg. G.L.Webster (Klotzsch) Müll.Arg. Müll.Arg. Müll.Arg. L. Croizat Mansf. Müll.Arg. Lundell Müll.Arg. Brandegee Urb. Sprague B.L.Rob.& Greenm. Urb. & Ekman Jabl. Rusby Griseb. (Sw.) Jabl. Mart. ex Colla Sond. Brandegee Müll P. pentandrus Schumach. & P. P. parvulus P. Species

P. zanthoxyloides P. axillaris P. cauliflorus P. cladanthus P. chryseus R.A.Howard P. acacioides P. maleolens P. martii Müll.Arg. P. megapodus P.

P. comorensis Leandri P. P. nummulariifolius P.

P. eurisladro P. larensis Steyerm. P. minarum Standl. & Steyerm. P. petenensis P. sellowianus P. P. tenellus Roxb. P. P. adianthoides P. P. borjaensis P. gentryi G.L.Webster P. juglandifolius P. P. atabapoensis P. mickelii McVaugh P. paezensis P. rupestris Kunth P. spruceanus P. purpusii P. riedelianus P. P. octomerus Müll.Arg. P. adenodiscus P. anderssonii P. biantherifer P. botryanthus Müll P. P. chiapensis P. coalcomanensis P. gradyi M.J.Silva & M.F.Sales P. grandifolius P. huallagensis P. P. laxiflorus Benth. P. mutisianus G.L.Webster P. nutans Sw. P. oaxacanus P. pachystylus Urb. P. poeppigianus P. P. prunifolius P. racemigerus P. ramosus Vell. P. tequilensis P. umbratus Müll.Arg. P. urbanianus P. ventricosus G.L.Webster P. vincentae J.F.Macbr. P.

Pentandra Pentandra Tenellanthus Tenellanthus Tenellanthus Elutanthos Epistylium Epistylium Epistylium Glyptothammus Hemiphyllanthus Hemiphyllanthus Hemiphyllanthus Hemiphyllanthus Adianthoides Adianthoides Asterandra Asterandra Brachycladus Brachycladus Brachycladus Brachycladus Brachycladus Ciccastrum Ciccastrum Diplocicca Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos Elutanthos sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. ? ? ? ? ? sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Appendix 2 (cont.) Subg. Tenellanthus Subgeneric placement Subg. Tenellanthus

Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Tenellanthus Subg. Tenellanthus Subg. Tenellanthus

Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla R.W. Bouman et al.: Subgeneric delimitation of Phyllanthus (Phyllanthaceae) 197 subg. Xylophylla subg. Xylophylla subg. Xylophylla Phyllanthus Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 D x x 2002 Synopsis of Webster x x x x x x 2002 Synopsis of Webster x x 2002 Synopsis of Webster x 1958 Webster x x 1958 Webster x 1958 Webster x x 1958 Webster 1958 Webster x x x x 1958 Webster 1958 Webster x x x x x Hidalgo et al. 2017 x x 1958 Webster x 1958 Britton 1920, Webster x x x 1955,1958 Webster x 1955,1958 Webster 1955,1958 Webster ~ x 1958 Webster x x x Müller 1866 x x x 1958 Webster x x 1958 Webster x x 1958 Webster x x 1958 Webster x x x x x x 1958 Webster x 1958 Webster 1958 Webster x x x Müller 1866 x x x x x Brunel 1987 x x x x 1996 Radcliffe-Smith x 1996 Radcliffe-Smith x 2002b Webster 2002b Webster 2002b Webster x x x x x x 2002b Webster x 2002b Webster x 2002b Webster 2002b Webster x x x x x 1960 Webster x x 1958 Webster 1958 Webster x 1958 Webster America America America Müller 1865 America x x x x x America 2002b Webster x America 2002b Webster 2002b Webster x x x x 2002b Webster 2002b Webster America America x x Müller 1866 x Colla 1831 Africa Africa Africa America x x et al. 2007 Vorontsova America x x Müller 1866 Global area Indies West Indies West Malesia Indies West Indies West Indies West Central Indies West Indies West Indies West Indies West Indies West Indies West Indies West Central Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Indies West Tropical Tropical Tropical South Tropical Australia Australia Australia South Tropical South Tropical South Tropical Australia Australia Australia Australia South Tropical South Tropical Indies West Indies West Indies West South Tropical Indies West South Tropical Tobago & Trinidad Trinidad Africa Major area Leeuward Isles, Haiti Martinique Cuba, Haiti Cuba Cuba Puerto Rico Cuba Cuba Cuba to Puerto Rico, Surinam Dominican Republic Cuba Cuba Cuba Colombia, Costa Rica, Peru, Venezuela Cuba Cuba Cuba Cuba Cuba Cuba Cuba Cuba Cuba Cuba Cuba, Jamaica, Nicaragua Jamaica Tropical Zimbabwe Zambia Haiti, Puerto Rico, Brazil Australia Australia (Tasmania) Australia Argentina Brazil Brazil Australia Australia Australia Brazil Argentina, Brazil Anguilla, Bahamas, Cuba, Dominican Republic, Guadeloupe, Indies West Jamaica Jamaica Jamaica Brazil Jamaica Brazil Urb. Müll.Arg. Falcón & J.L.Gómez L. Radcl.-Sm. & J.J.Bruhl J.T.Hunter Australia Urb. ex Griseb. C.Wright Griseb. Urb. Sw. (Sw.) (Kuntze) K.Schum. Sw. (Britton) Urb. Griseb. (Britton) Croizat Kunth Kunth F.Muell. Müll.Arg. J.F.Gmel. (Sw.) Thunb. Steyerm. Baill. & J.J.Bruhl J.T.Hunter Species P. mimosoides P. P. subcarnosus P. P. myriophyllus P. ovatus Poir. P. chamaecristoides P. comosus Urb. P. cuneifolius P. formosus Urb. P. myrtilloides P. nummularioides P. P. orbicularis P. phialanthoides P. P. phlebocarpus P. scopulorum P. salviifolius P. cinctus Urb. P. comptus G.L.Webster P. ekmanii G.L.Webster P. P. cristalensis Urb. P. P. discolor Poepp. P. P. microdictyus Urb. P. P. excisus Urb. P. P. incrustatus Urb. P. P. williamioides P. P. incurvus P. serpentinicola P. tener Radcl.-Sm. P. P. angustifolius P. arbuscula P. epiphyllanthus P. P. mirificus G.L.Webster P. P. fallax Müll.Arg. P. aridus Benth. P. australis Hook.f. P. calycinus Labill. P. cordobensis P. dawsonii P. dictyospermus Müll.Arg. P. erwinii P. ex Müll.Arg. hirtellus F.Muell. P. lacunarius P. oblanceolatus P. pinifolius P. ramillosus P. & Proctor eximius G.L.Webster P. latifolius (L.) Sw. P. Sw. montanus (Sw.) P. obtusatus (Billb.) Müll.Arg. P. proctoris G.L.Webster P. robustus Mart. ex Colla P.

subsect. subsect. subsect. subsect. subsect. subsect. subsect. Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Antipodanthus Hemiphyllanthus Hemiphyllanthus Hemiphyllanthus Omphacodes Orbicularia Orbicularia Orbicularia Orbicularia Orbicularia Orbicularia Orbicularia Orbicularia Orbicularia Orbicularia Oxalistylis Thamnocharis Thamnocharis Thamnocharis Williamia Williamia Williamia Williamia Williamia Williamia Williamia Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla Xylophylla sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect.

Appendix 2 (cont.) Subgeneric placement Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Discolores Subg. Xylophylla Discolores Subg. Xylophylla Discolores Subg. Xylophylla Incrustati Subg. Xylophylla Incrustati Subg. Xylophylla Incrustati Subg. Xylophylla Mirifici Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Subg. Xylophylla Former Isocladus 198 Blumea – Volume 63 / 2, 2018 Phyllanthus , but species not seen , but species not seen , but species not seen ) ) Eriococcus Eriococcus Eriococcus , but species not seen , but species not seen , but species not seen ( Euphorbiaceae ( Euphorbiaceae Eriococcus Eriococcus Eriococcus Phyllanthus Phyllanthus Most important literature 3 L 2 M 1 Thin 2007, possible Thin 2007, possible Thin 2007, possible D x x x x 2002b Webster x 2002b Webster Brunel 1987 x Brunel 1987 x Brunel 1987 x Brunel 1987 x Brunel 1987 x 2001b Webster x Brunel 1987 x Brunel 1987 x x Johnston 1986 x 2001b Webster 2002 Outline of the Neotropical infrageneric taxa Webster

x x 2001b Webster 2001b Webster 1997 Webster x 2002 Outline of the Neotropical infrageneric taxa Webster x 2001b Webster Merrill 1914 Possible Specimen not seen or described Male flowers not known Possible

Male flowers not known 2011 See Ralimanana & Hoffmann Possible Thulin 2003 America America x x x x 2002b Webster Silva 2009 America America x Müller 1863, 1866, 1873 America x Müller 1866, 1873 America Manuscript outline neotropical See Webster America America Sterile Asia Asia Asia Asia Asia Asia Asia Asia Africa Africa Africa Africa Africa Africa Africa Africa Africa America America America America America America America America Tropical Tropical Tropical Tropical Tropical Tropical Tropical Mainland Mainland Mainland Global area South Tropical Australia Australia North, Central and South Tropical North x 2001b Webster South Tropical North North North South Tropical

North North Malesia South Tropical Malesia Malesia South Tropical Malesia North North Tropical Malesia South Tropical Mainland South Tropical Mainland Mainland Malesia Mainland Madagascar Mainland Tropical Malesia Africa is invalid. Angola Angola Namibia, South Ethiopia, Mozambique Angola, Democratic Republic of the Congo, Namibia Mascarenes, Mozambique Somalia Vietnam Vietnam Vietnam Major area Brazil Australia Australia Mexico, United States: Florida Mexico, United States Brazil Mexico Mexico Mexico Brazil

Mexico Mexico Sri Lanka Paraguay Guam Philippines Venezuela Celebes Mexico Mexico Unknown Somalia Celebes Venezuela India Uruguay India Philippines Vietnam Madagascar China Somalia Lesser Sunda Isl. Thin (2007) for Phyllanthus Trimen Thulin Herter & Mansf. and its use by R.L.Mitra & Sanjappa Nepal Bhutan, India, Myanmar, Merr. Müll.Arg. Müll.Arg. Thin Nutt. ex Spreng. Croizat

Jean F.Brunel Stokes (Lour.) Thwaites ex Thin Baill. Koord. Brandegee Jean F.Brunel

Steyerm. Jean F.Brunel

Coode Jean F.Brunel Steyerm. Thulin Müll.Arg. Müll.Arg. Torr. Müll.Arg. McVaugh Koord. Dinter M.C.Johnst.

Roxb. M.G.Gilbert & M.J.Silva Elmer Baccaurea Thin P. caraculiensis P. cunenensis P. karibibensis P. liebmannianus P. mendoncae P. paxianus P. polygonoides P. revaughanii P. spinosus Chiov. P. P. daclacensis P. songboiensis P. tui P. P. salesiae P. scaber Klotzsch P. & J.J.Bruhl striaticaulis . J.T.Hunter P. bahiensis P. barbarae P. ericoides P. fraguensis M.C.Johnst. P. Species P. rosmarinifolius P.

P. galeottianus P. gypsicola P. hakgalensis P. lasiogynus P. neoleonensis P. peninsularis P. P. saffordii Merr. P. anisophyllioides P. bolivarensis P. celebicus P. P. hortensis Govaerts & Radcl.-Sm. P. lunifolius P. P. minahassae P. P. orinocensis P. petiolaris P. pseudoguyanensis P. pseudoparvifolius P. P. rotundatus Poir. P. rupicola P. squamifolius P. vergens Baill. P. villosus Poir. P. P. xylorrhizus P. P. zippelianus P.

sensu sensu sensu sensu sensu sensu sensu sensu sensu is currently placed in the genus Antipodanthus Antipodanthus Antipodanthus Isocladus Isocladus Isocladus Isocladus Isocladus Isocladus Isocladus Isocladus Isocladus Paraphyllanthus Paraphyllanthus Paraphyllanthus Paraphyllanthus Antipodanthus Paraphyllanthus Paraphyllanthus Paraphyllanthus Paraphyllanthus Paraphyllanthus Paraphyllanthus Paraphyllanthus

4 4 4 sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. sect. Placement based on DNA. Placement based on morphology. Placement based on a literature reference. Section Hedycarpidium Appendix 2 (cont.) sect. Hedycarpidium sect. Hedycarpidium Former Isocladus Former Isocladus Former Isocladus Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Brunel 1987 Former Isocladus Former Isocladus Former Isocladus Former Isocladus Subgeneric placement Former Isocladus 1 2 3 4 Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Former Isocladus Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis Incertae sedis sect. Hedycarpidium