Several groups within tribe Phyllantheae (Phyllanthaceae) formed, independently, an (obligate) pollination mutualism with Epicephala moths, which originally had been parasitic. In this pollination... Show moreSeveral groups within tribe Phyllantheae (Phyllanthaceae) formed, independently, an (obligate) pollination mutualism with Epicephala moths, which originally had been parasitic. In this pollination system, female moths actively collect pollen from staminate flowers and deposit it on the stigma of pistillate flowers, after which they place at least one egg in or against the ovary. The high pollination rate makes the system beneficial for the plants, whereas the larvae are provided with food (part of the developing seeds) and some protection against predation. Qualitative comparisons are made between non-moth-pollinated lineages, used as outgroups and various, independently moth-pollinated Phyllantheae clades, used as ingroups, thereby looking for parallel developments. The flowers of both sexes of various groups display similar, convergent morphological adaptations to the pollination system, likely to secure the obligate relationship and to improve efficiency. Sepals in both sexes, free or partly to highly connate, are commonly upright and form a narrow tube. The staminate flowers often have united, vertical stamens with the anthers along the androphore or on top of the androphore. Pistillate flowers generally reduce the stigmatic surface, either by making the stigmas shorter or by uniting them into a cone with a small opening at the top for pollen deposition. Less obvious is the reduction of the stigmatic papillae; these are often present in non-moth-pollinated taxa, but absent in the moth-pollinated species. The most diverging, parallel adaptations to moth pollination are currently found in the Palaeotropics, whereas in the Neotropics, some groups continue to also be pollinated by other insect groups and are morphologically less changed. Show less
The largest genus within the Phyllanthaceae family is a group called Phyllanthus L. Recent studies have shown, that Phyllanthus is paraphyletic with the genera Glochidion, Breynia and Synostemon... Show moreThe largest genus within the Phyllanthaceae family is a group called Phyllanthus L. Recent studies have shown, that Phyllanthus is paraphyletic with the genera Glochidion, Breynia and Synostemon nested within it. In this thesis, I study the evolution and systematics of Phyllanthus and how to solve the apparent paraphyly. First we explore past morphological subgroups to come to a subdivision, which was tested using molecular phylogenetics. Several species of Phyllanthus are also used in traditional medicine and by utilizing the living collections of the Hortus botanicus Leiden, we tested for antibacterial and antifungal bioactivity. The phylogeny resulting from this study is used to study the historical biogeography and diversification of the tribe and to propose a new classification where Phyllanthus is separated into several smaller genera. Show less
The majority of tribe Phyllantheae (Phyllanthaceae) is currently placed in the paraphyletic genus Phyllanthus and discussions have persisted on how to resolve this issue. Here, we split Phyllanthus... Show moreThe majority of tribe Phyllantheae (Phyllanthaceae) is currently placed in the paraphyletic genus Phyllanthus and discussions have persisted on how to resolve this issue. Here, we split Phyllanthus into ten monophyletic genera, which are all reinstatements of former genera, but with changes made to the circumscription and constituent species of each group. The genera Breynia, Glochidion and Synostemon were recently found to be nested within Phyllanthus and discussions ensued whether or not to subsume everything into Phyllanthus s.l. Instead of combining all these genera, we here implement the solution of splitting Phyllanthus into strictly monophyletic genera to ensure that the classification is consistent with the latest phylogenetic results. The new classification is based on a phylogenetic framework combined with differences in habit, branching type, floral, fruit and pollen morphology. With this new division of the genus Phyllanthus, tribe Phyllantheae will consist of the following 18 genera: Breynia, Cathetus, Cicca, Dendrophyllanthus, Emblica, Flueggea, Glochidion, Heterosavia, Kirganelia, Lingelsheimia, Lysiandra, Margaritaria, Moeroris, Nellica, Nymphanthus, Phyllanthus, Plagiocladus and Synostemon. As a result of the reinstated genera, five new names for illegitimate combinations or previous overlooked nomenclatural anomalies and 645 new combinations are proposed. Several keys are provided to distinguish the reinstated genera. Full species lists are given for the reinstated genera treated here except for Breynia, Synostemon and Glochidion. Show less
The genus Phyllanthus is paraphyletic as currently circumscribed, with the genera Breynia, Glochidion and Synostemon nested within it. A phylogeny based on nuclear (ITS, PHYC) and chloroplast (matK, .Show moreThe genus Phyllanthus is paraphyletic as currently circumscribed, with the genera Breynia, Glochidion and Synostemon nested within it. A phylogeny based on nuclear (ITS, PHYC) and chloroplast (matK, accD-psaI, trnS-trnG) markers is presented, including 18/18 subgenera and 53/70 sections. Differences in habit, branching type, floral and fruit characters are discussed, and we find indications for shifts in pollination and dispersal strategies possibly underlying the convergent evolution of these characters in multiple clades. Several taxonomic issues were found in the subgeneric classification of Phyllanthus that will require new transfers and rank changes. Phyllanthus subg. Anesonemoides, subg. Conami, subg. Emblica, subg. Gomphidium, subg. Kirganelia and subg. Phyllanthus are polyphyletic, and several sections appear to be paraphyletic (e.g., P. sect. Anisonema, sect. Emblicastrum, sect. Pseudoactephila, sect. Swartziani, and sect. Xylophylla); P. subg. Phyllanthodendron is furthermore paraphyletic with the genus Glochidion nested within. To create a classification of tribe Phyllantheae that comprises exclusively monophyletic taxa, it is necessary to treat several clades at the same taxonomic rank as the genera Breynia, Glochidion and Synostemon. Since combining all genera would lead to one giant heterogeneous genus that is difficult to define, we recommend dividing Phyllanthus into several monophyletic genera, which have previously been recognized and often possess diagnostic (combinations of) morphological characters. This new classification is forthcoming. Show less
During the environmental impact study for a proposed nickel mine near Weda Bay on Halmahera in North Moluccas (Maluku Utara Province), Indonesia, two unknown Euphorbiaceae were discovered.... Show moreDuring the environmental impact study for a proposed nickel mine near Weda Bay on Halmahera in North Moluccas (Maluku Utara Province), Indonesia, two unknown Euphorbiaceae were discovered. Morphological comparisons and molecular phylogenetic analyses using four markers (plastid trnL-F and rbcL, and nuclear ribosomal internal transcribed spacer and external transcribed spacer) indicated that they should be recognized as constituting a new, distinct genus of two species, which are described and illustrated here as Weda fragarioides and Weda lutea. The new taxa are members of the Australasian tribe Ricinocarpeae in subfamily Crotonoideae, and they are most closely related to Alphandia. In contrast with the otherwise mostly sclerophyllous Ricinocarpeae, Weda possesses stellate to dendritic hairs, large, long-petiolate, glandular leaves, and inflorescences with a pair of large, leafy, subopposite bracts. The two narrowly distributed species are distinguished from each other by vegetative and floral features, molecular data, and elevational preferences. Leaf elemental analysis of Weda indicated manganese, but not nickel, accumulation. Newly resolved generic relationships and potential morphological synapomorphies within Crotonoideae are discussed, and the circumscription of Ricinocarpeae is expanded from 7 to 11 genera. Show less
Bouman, R.W.; Kessler, P.J.A.; Welzen, P.C. van 2019
Two species of Phyllanthus collected and described by Koorders during his travels on the island of Sulawesi(Indonesia) are lectotypified, descriptions amended and their taxonomic affinity is... Show moreTwo species of Phyllanthus collected and described by Koorders during his travels on the island of Sulawesi(Indonesia) are lectotypified, descriptions amended and their taxonomic affinity is discussed. Phyllanthus mindorensis was found to be too similar to P. celebicus and is placed in the synonymy of the latter. A key is provided to the species of Phyllanthus on Sulawesi. Show less
Over two centuries of taxonomic studies on the species rich genus Phyllanthus have culminated in a broadand complicated classification with many subgenera and (sub)sections. Past taxonomic work has... Show moreOver two centuries of taxonomic studies on the species rich genus Phyllanthus have culminated in a broadand complicated classification with many subgenera and (sub)sections. Past taxonomic work has only focused on local revisions, mostly because of the size of the genus. In this study we aim to summarize most of the taxonomic work in a list containing the infrageneric delimitations of Phyllanthus. This work will serve as a reference, placing most currently recognized species in subgenera and if possible, in sections for further study. Here we recognize 880 species of Phyllanthus, classified in 18 subgenera, 70 sections and 14 subsections. A few taxonomic changes 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. Show less
Bouman, R.W.; Welzen, P.C. van; Sumail, S.; Echevarria, G.; Erskine, P.D.; Ent, A. van der 2018
