Phyllanthaceae comprises about 2000 species.[6][7] 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.[8]
Some of the genera have recently been sunk into others, while other genera have recently been divided.[9][10] 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).[11]
The description here is from Hoffmann,[4][14] except for a few additions from Webster[15] and Hutchinson [16] 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.
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.[13]
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.[16] 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.[15] The style is usually 2-lobed or bifid, sometimes entire, or rarely multifid.
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 generbcL from a large number of seed plants.[18] 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.[19][20]Pandaceae, Phyllanthaceae, Picrodendraceae, Putranjivaceae, Peraceae, and Centroplacaceae have been removed from it.[11]
The obsolete, older concept of Euphorbiaceae, known as Euphorbiaceae sensu lato, is sometimes still used for continuity and convenience.[21] 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.[15][22][23]
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.[11]Paradrypetes is in Rhizophoraceae.[3]Phyllanoa is known only from a single specimen. In 1996, this was examined and found to be a species of Rinorea (Violaceae).[24]
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.
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.[14] Plants of the World Online still accepts Breynia and Glochidion,[25] and subsumes Sauropus into Breynia.[26]
The revision of Phyllanthaceae by Hoffmann and co-authors was based on two molecular phylogenetic studies that were published in 2005.[6][7] Since the revision, phylogenetic studies have been done on some of the tribes.[8][27]
The phylogenetic tree shown below is based on the results of several studies.[8][12][13][14] Fifty-one genera are represented. Chonocentrum(Phyllanthaceae, incertae sedis), and three members of the tribe Scepeae (Ashtonia, Distichirrhops, and Nothobaccaurea) have not yet been sampled for DNA. Chonocentrum is known from only a single specimen collected in the 1850s.[24]
In the phylogeny shown below, statistical support for the clades was measured by bootstrap percentage. All branches shown below have maximum parsimony bootstrap support of at least 70%.
Phyllanthus is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750 to 1200. Phyllanthus has a remarkable diversity of growth forms including annual and perennial herbs, shrubs, climbers, floating aquatics, and pachycaulous succulents. Some have flattened leaflike stems called cladodes. It has a wide variety of floral morphologies and chromosome numbers and has one of the widest range of pollen types of any seed plant genus.
The family Pandaceae consists of three genera that were formerly recognized in the Euphorbiaceae. Those are:
Leptopus, the maidenbushes, are a genus of plants in the family Phyllanthaceae native to southern Asia from the Caucasus east to China and Maluku. The plants are monoecious herbs and shrubs with simple, entire leaves and small, green flowers.
Bischofia is a genus of plants in the family Phyllanthaceae first described as a genus in 1827. It is native to southern China, the Indian Subcontinent, Queensland, New Guinea, and various islands of the Pacific. It is the only member of the tribe Bischofieae. They are dioecious, with male and female flowers on separate plants, but may rarely be monoecious.
Martretia is a genus of plants in the family Phyllanthaceae first described as a genus in 1907. It contains only one known species, Martretia quadricornis, native to western and central Africa from Sierra Leone to Zaire. It is dioecious, with male and female flowers on separate plants.
Leptonema is a genus of flowering plant belonging to the family Phyllanthaceae first described in 1824. The entire genus is endemic to Madagascar. It is dioecious, with male and female flowers on separate plants.
Leptonema glabrum(Leandri) Leandri
Leptonema venosum(Poir.) A.Juss.
Thecacoris is a genus of flowering plant belonging to the family Phyllanthaceae first described as a genus in 1821. It is native to tropical Africa and Madagascar. It is dioecious, with male and female flowers on separate plants, although it may rarely be monoecious.
Ashtonia is a genus of flowering plant belonging to the family Phyllanthaceae first described as a genus in 1968. It is native to the Malay Peninsula and Borneo. It is dioecious, with male and female flowers on separate plants.
Ashtonia excelsa Airy Shaw - Borneo
Ashtonia praeterita Airy Shaw - S Thailand, W Malaysia
Maesobotrya is a genus of flowering plants belonging to the family Phyllanthaceae. It was first described as a genus in 1879 and is native to sub-Saharan Africa. It is dioecious, with male and female flowers on separate plants.
Richeria is a genus of flowering plant belonging to the family Phyllanthaceae first described as a genus in 1797. It is native to Central America, South America, and the West Indies. Richeria is dioecious, with male and female flowers on separate plants.
Richeria australis - São Paulo, Mato Grosso
Richeria dressleri - Costa Rica, Nicaragua, Panama, Ecuador
Richeria grandis - Panama, N South America, E West Indies
Richeria obovata - Costa Rica to Bolivia
Richeria tomentosa - Colombia, Ecuador
Uapaca is a genus of plant, in the family Phyllanthaceae first described as a genus in 1858. It is the only genus comprised in the tribe Uapaceae. The genus is native to Africa and Madagascar. Uapaca is dioecious, with male and female flowers on separate plants.
Didymocistus is a genus of plants in the family Phyllanthaceae first described as a genus in 1940. It contains only one known species, Didymocistus chrysadenius, native to the Loreto region of northeastern Peru, the Amazonas Department of southeastern Colombia, and the State of Amazonas in northwestern Brazil. It is dioecious, with male and female flowers on separate plants.
Hymenocardia is a genus of trees in the family Phyllanthaceae first described as a genus in 1836. Most of the species are native to Africa, with one in Southeast Asia.
Breynia is a genus in the flowering plant family Phyllanthaceae, first described in 1776. It is native to Southeast Asia, China, Réunion, the Indian Subcontinent, Papuasia and Australia.
Glochidion is a genus of flowering plants, of the family Phyllanthaceae, known as cheese trees or buttonwood in Australia, and leafflower trees in the scientific literature. It comprises about 300 species, distributed from Madagascar to the Pacific Islands. Glochidion species are used as food plants by the larvae of some Lepidoptera species including Aenetus eximia and Endoclita damor. The Nicobarese people have attested to the medicinal properties found in G. calocarpum, saying that its bark and seed are most effective in curing abdominal disorders associated with amoebiasis.
Keayodendron is a genus of plants in the family Phyllanthaceae first described as a genus in 1959. It contains only one known species, Keayodendron bridelioides, native to tropical western and Central Africa. It is dioecious, with male and female flowers on separate plants.
Gonatogyne is a genus of plants in the family Phyllanthaceae first described as a genus in 1873. It contains only one known species, Gonatogyne brasiliensis, endemic to southeastern Brazil. It is dioecious, with male and female flowers on separate plants.
Richeria grandis is a tree species in the family Phyllanthaceae which ranges from the Lesser Antilles to South America. The species is reputed to have aphrodisiac properties.
Poranthereae is a tribe in the plant family Phyllanthaceae. It is one of ten tribes in the family, and one of four tribes in the subfamily Phyllanthoideae. Poranthereae comprises about 111 species, distributed into eight genera. The largest genera and the number of species in each are Actephila (31), Meineckia (30), and Andrachne (22).
1 2 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 Botany96(8):1551-1570. (see External links below)
1 2 3 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. ISBN978-1-55407-206-4.
↑ 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. ISBN978-0-333-47494-5 (set)
1 2 3 Hashendra S. Kathriarachchi; Petra Hoffmann; Rosabelle Samuel; Kenneth J. Wurdack & Mark W. Chase (2005). "Molecular phylogenetics of Phyllanthaceae inferred from five genes (plastid atpB, matK, 3'ndhF, rbcL, and nuclear PHYC)". Molecular Phylogenetics and Evolution. 36 (1): 112–134. doi:10.1016/j.ympev.2004.12.002. PMID15904861.
↑ Hoffmann, Petra; McPherson, Gordon (2007). "Revision of Wielandia, including Blotia and Petalodiscus (Phyllanthaceae)". Annals of the Missouri Botanical Garden. 94 (3): 519–553. doi:10.3417/0026-6493(2007)94[519:ROWIBA]2.0.CO;2.
↑ 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 Botany102(6):1007-1018. (see External links below)
1 2 3 "Phyllanthaceae" In: Peter F. Stevens (2001 onwards). Angiosperm Phylogeny Website. In: Missouri Botanical Garden Website. (see external links below)
1 2 Hoffmann, Petra (2008). "Revision of Heterosavia, status novus, with notes on Gonatogyne and Savia (Phyllanthaceae)". Brittonia. 60 (2): 136–166. doi:10.1007/s12228-008-9012-5. S2CID34814559.
1 2 John Hutchinson. "Euphorbiaceae" pages 329-330. In: The Families of Flowering Plants, Third Edition (1973). Oxford University Press: London.
↑ Reveal, James L.; Hoffmann, Petra; Doweld, Alexander; Wurdack, Kenneth J. (2007). "(1765) Proposal to conserve the name Phyllanthaceae.". Taxon. 56 (1): 266.
↑ 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 Garden80(3):528-580.
↑ 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 Research120(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." Science315(5820):1812. (see External links below).
↑ 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).
↑ Alan Radcliffe-Smith. 2001. Genera Euphorbiacearum. Kew Publishing, Royal Botanic Gardens, Kew: Richmond, England.
1 2 W. John Hayden & Sheila M. Hayden (1996). "Two enigmatic biovulate Euphorbiaceae from the Neotropics: relationships of Chonocentron and the identity of Phyllanoa". American Journal of Botany. 83 (6): 162. doi:10.2307/2445447. JSTOR2445447.
Wurdack, KJ; Davis, CC (August 2009). "Malpighiales phylogenetics: Gaining ground on one of the most recalcitrant clades in the angiosperm tree of life". Am. J. Bot. 96 (8): 1551–70. doi:10.3732/ajb.0800207. PMID21628300.
Kathriarachchi, H; Hoffmann, P; Samuel, R; Wurdack, KJ; Chase, MW (July 2005). "Molecular phylogenetics of Phyllanthaceae inferred from five genes (plastid atpB, matK, 3'ndhF, rbcL, and nuclear PHYC)". Mol. Phylogenet. Evol. 36 (1): 112–34. doi:10.1016/j.ympev.2004.12.002. PMID15904861.
Tokuoka, T (July 2007). "Molecular phylogenetic analysis of Euphorbiaceae sensu stricto based on plastid and nuclear DNA sequences and ovule and seed character evolution". J. Plant Res. 120 (4): 511–22. doi:10.1007/s10265-007-0090-3. PMID17530165. S2CID19614584.
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