Phyllanthaceae, commonly known as the leaf-flower family, is a family of flowering plants in the eudicot order Malpighiales. It is most closely related to the family Picrodendraceae.Kenneth J. Wurdack and Charles C. Davis. 2009. "Malpighiales phylogenetics: Gaining ground on one of the most recalcitrant clades in the angiosperm tree of life." American Journal of Botany 96(8):1551-1570. (see External links below)
The Phyllanthaceae are most numerous in the tropics, with many in the south temperate zone, and a few ranging as far north as the middle of the north temperate zone.Petra Hoffman. 2007. "Phyllanthaceae" pages 250-252. In: Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alastair Culham. Flowering Plant Families of the World. Firefly Books: Ontario, Canada. .
Some species of Andrachne, Antidesma, Margaritaria, and Phyllanthus are in cultivation.Anthony J. Huxley, Mark Griffiths, and Margot Levy (editors). 1992. The New Royal Horticultural Society Dictionary of Gardening. The Macmillan Press Limited, London; The Stockton Press, New York. (set) A few species of Antidesma, Baccaurea, Phyllanthus, and Uapaca bear edible fruit.
Phyllanthaceae comprises about 2000 species. Depending on the author, these are grouped into 54 to 60 genera. Some of the genera are poorly defined, and the number of genera in the family is likely to change as the classification is further refined. The genus Phyllanthus, one of the largest genera of flowering plants with over 1200 species, has more than half of the species in the family.
Some of the genera have recently been sunk into others, while other genera have recently been divided.Kanchana Pruesapan, Ian R.H. Telford, Jeremy J. Bruhl, Stefano G.A. Draisma, and Peter C. Van Welzen. 2008. "Delimitation of Sauropus (Phyllanthaceae) Based on Plastid matK and Nuclear Ribosomal ITS DNA Sequence Data." Annals of Botany 102(6):1007-1018. (see External links below) The largest genera and the approximate number of species in each are: Phyllanthus (1270), Cleistanthus (140), Antidesma (100), Aporosa (90), Uapaca (60), Baccaurea (50), and Bridelia (50)."Phyllanthaceae" In: Peter F. Stevens (2001 onwards). Angiosperm Phylogeny Website. In: Missouri Botanical Garden Website. (see external links below)
Since Phyllanthaceae was revised in 2006, one paper has removed Heterosavia from Savia. Another has separated Notoleptopus from Leptopus, and segregated Pseudophyllanthus and Phyllanthopsis from Andrachne. Also, Oreoporanthera has been subsumed into Poranthera, while Zimmermannia and Zimmermanniopsis have been sunk into Meineckia. The large genus Cleistanthus is known to be polyphyletic, but further studies will be needed before it can be revised.
Description The description here is from Hoffmann, except for a few additions from Webster and HutchinsonJohn Hutchinson. "Euphorbiaceae" pages 329-330. In: The Families of Flowering Plants, Third Edition (1973). Oxford University Press: London. where cited. Phyllanthaceae is an unusually diverse family for its moderate size. It can be recognized only by a combination of characters because there are a few exceptions to almost everything that is generally true of the family. It is most notable for having two ovules in each locule of the ovary, a trait that clearly distinguishes it from Euphorbiaceae.
The Phyllanthaceae are nearly all trees, shrubs, or herbs. A few are climbers, or succulents, and one species, Phyllanthus fluitans, is aquatic. Unlike many of the Euphorbiaceae, none has latex, and only a very few produce a resinous exudate. Any hairs, if present, are almost always simple. Rarely are they branched or scale-like. Thorns and other armament are rare.
Stipules are produced with each leaf, but in some, these fall before the leaf is fully mature. Leaves are present, except for a few species of Phyllanthus that have flattened, leaflike stems called cladodes that bear flowers along their edges. The leaves are compound in Bischofia, but otherwise simple and usually alternate. Rarely are they opposite, in fascicles, or in whorls around the stem. The leaf margin is almost always entire, rarely toothed. A petiole is nearly always present, often with a pulvinus at its base.
The inflorescences are usually in the axils of leaves, rarely below the leaves or at the ends of stems. In Uapaca, the flowers are in a pseudanthium, a tight bundle of flowers that resembles a single flower.
Except for four species of Aporosa, the flowers are unisexual, the plants being either monoecious or dioecious. The flowers are actinomorphic in form. Detailed illustrations have been published for some of these.
The sepals are three to eight in number, usually free from each other. Petals may be absent or present. If present, there are usually four to six, and their color is yellow to green, or rarely, pink or maroon.
A nectary disk is often present. It may be in the form of a ring, or divided into segments. The stamens are three to ten in number, or rarely more, free or variously fused.
The ovary is superior. The number of locules in the ovary is highly variable, usually from two to five, but sometimes as many as fifteen. The placentation is apical, with a pair of ovules hanging by their funicles from the top of each locule. Often, only one of the ovules will develop into a seed. A single, massive obturator may cover the micropyles of both ovules, or each ovule may have its own thin obturator. The megagametophyte is of the Polygonum type. The style is usually 2-lobed or bifid, sometimes entire, or rarely multifid.
The fruit is a schizocarp, drupe, or berry. In some, the schizocarp breaks up explosively.
Taxonomy
History The name "Phyllanthaceae" was first validly published by Ivan Ivanovich Martynov in 1820 in a Russian book entitled Tekhno-botanico Slovar. A proposal to conserve this name was published in 2007.
Martynov's name was rarely used in the 180 years after he published it. During that time, the plants that are now in Phyllanthaceae were placed in the large and heterogeneous family Euphorbiaceae. The monophyly of Euphorbiaceae had long been held in doubt by some, but the first strong evidence of its polyphyly came in 1993 with the first maximum parsimony analysis of DNA sequences of the gene rbcL from a large number of seed plants.Mark W. Chase et alii (42 authors). 1993. "Phylogenetics of seed plants: an analysis of nucleotide sequences from the plastid gene rbcL." Annals of the Missouri Botanical Garden 80(3):528-580. Since the 1993 study, all subsequent phylogenetic analyses have shown that the old concept of Euphorbiaceae consisted of several lineages that did not together form a clade in the order Malpighiales. Euphorbiaceae is now defined as a much smaller family than it had been in the twentieth century.Toru Tokuoka. 2007. "Molecular phylogenetic analysis of Euphorbiaceae sensu stricto based on plastid and nuclear DNA sequences and ovule and seed character evolution." Journal of Plant Research 120(4):511-522. (see External links below).Charles C. Davis, Maribeth Latvis, Daniel L. Nickrent, Kenneth J. Wurdack, and David A. Baum. 2007. "Floral Gigantism in Rafflesiaceae." Science 315(5820):1812. (see External links below). Pandaceae, Phyllanthaceae, Picrodendraceae, Putranjivaceae, Peraceae, and Centroplacaceae have been removed from it.
The obsolete, older concept of Euphorbiaceae, known as Euphorbiaceae sensu lato, is sometimes still used for continuity and convenience.Petra Hoffmann, Don Kirkup, Aimee Galster, Gill Challen, and Alan Radcliffe-Smith. 2005 onward. Interactive Key to the Genera of Euphorbiaceae sensu lato. In: Index of /herbarium/keys. (see External links below). It was the subject of a book and two papers which stood as the standard works on Phyllanthaceae until that family was revised by Hoffmann and co-authors in 2006.Alan Radcliffe-Smith. 2001. Genera Euphorbiacearum. Kew Publishing, Royal Botanic Gardens, Kew: Richmond, England.
Classification
In the past, the genera Centroplacus, Paradrypetes, and Phyllanoa had been placed in Phyllanthaceae, but these are now excluded from the family. Centroplacus is now in the family Centroplacaceae. Paradrypetes is in Rhizophoraceae. Phyllanoa is known only from a single specimen. In 1996, this was examined and found to be a species of Rinorea (Violaceae).
The family Phyllanthaceae is divided into two subfamilies: Antidesmatoideae and Phyllanthoideae. Antidesmatoideae is divided into six tribes and Phyllanthoideae is divided into four. The tribe Antidesmateae of Antidesmatoideae, and the tribes Bridelieae and Wielandieae of Phyllanthoideae are further divided into subtribes. The following classification table is from the 2006 revision of Phyllanthaceae.
Incertae sedis: Chonocentrum
Subfamily Antidesmatoideae 6 tribes Tribe Bischofieae 1 genus Bischofia Tribe Uapaceae 1 genus Uapaca Tribe Spondiantheae 1 genus Spondianthus Tribe Scepeae 8 genera Aporosa Ashtonia Baccaurea Distichirhops Maesobotrya Nothobaccaurea Protomegabaria Richeria Tribe Jablonskieae 2 genera Jablonskia Celianella Tribe Antidesmateae 5 subtribes Subtribe Hieronyminae 1 genus Hieronyma Subtribe Leptonematinae 1 genus Leptonema Subtribe Martretiinae 2 genera Martretia Apodiscus Subtribe Hymenocardiinae 2 genera Hymenocardia Didymocistus Subtribe Antidesmatinae 2 genera Antidesma Thecacoris Subfamily Phyllanthoideae 4 tribes Tribe Bridelieae 5 subtribes Subtribe Securineginae 2 genera Securinega Lachnostylis Subtribe Saviinae 5 genera Savia Croizatia Discocarpus Gonatogyne Tacarcuna Subtribe Pseudolachnostylidinae 4 genera Pseudolachnostylis Bridelia Cleistanthus Pentabrachion Subtribe Keayodendrinae 1 genus Keayodendron Subtribe Amanoinae 1 genus Amanoa Tribe Phyllantheae 5 genera Phyllanthus (including Breynia, Glochidion, Reverchonia, and Sauropus) Flueggea Lingelsheimia Margaritaria Plagiocladus Tribe Wielandieae 2 subtribes Subtribe Astrocasiinae 3 genera Astrocasia Chascotheca Heywoodia Subtribe Wielandiinae 3 genera Wielandia Chorisandrachne Dicoelia Tribe Poranthereae 8 genera Poranthera Actephila Andrachne Leptopus Meineckia Oreoporanthera Zimmermannia Zimmermanniopsis
Genera , Plants of the World Online accepted 58 genera.Phyllanthaceae Martinov. Plants of the World Online. Retrieved 2 January 2026.
A 2006 revision of Phyllanthaceae by Petra Hoffmann and co-authors recognized 54 genera. In their treatment, Blotia and Petalodiscus were sunk into Wielandia and Richeriella into Flueggea. Breynia, Glochidion, Reverchonia, and Sauropus were recommended to be subsumed into Phyllanthus, but many new species combinations must be published to effect this change. Genera previously considered as the tribe Drypeteae are now placed in the separate family Putranjivaceae. Plants of the World Online still accepts Breynia and Glochidion, and subsumes Sauropus into Breynia.Sauropus Blume. Plants of the World Online. Retrieved 17 March 2024.
Actephila Amanoa Andrachne Antidesma Apodiscus Aporosa Ashtonia Astrocasia Baccaurea Bischofia Breynia Bridelia Celianella Chascotheca Chonocentrum Chorisandrachne Cleistanthus Croizatia Dicoelia Didymocistus Discocarpus Distichirhops Flueggea Glochidion Gonatogyne Heterosavia Heywoodia Hieronyma Hymenocardia Jablonskia Keayodendron Lachnostylis Leptonema Leptopus Lingelsheimia Maesobotrya Margaritaria Martretia Meineckia Nothobaccaurea Notoleptopus Pentabrachion Phyllanthopsis Phyllanthus Plagiocladus Poranthera Protomegabaria Pseudolachnostylis Pseudophyllanthus Richeria Savia Securinega Spondianthus Synostemon Tacarcuna Thecacoris Uapaca Wielandia
Phylogeny