Phyllanthaceae
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Phyllanthaceae is a
family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...
of
flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...
s in the eudicot order
Malpighiales The Malpighiales comprise one of the largest orders of flowering plants, containing about 36 families and more than species, about 7.8% of the eudicots. The order is very diverse, containing plants as different as the willow, violet, poinsett ...
. It is most closely related to the family
Picrodendraceae Picrodendraceae is a family of flowering plants, consisting of 80 species in 24 genera.Stephens, P.F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008. http://www.mobot.org/MOBOT/Research/APweb/ These are subtropical to tropical ...
.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 The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...
, with many in the south temperate zone, and a few ranging as far north as the middle of the
north temperate zone In geography, the temperate climates of Earth occur in the middle latitudes (23.5° to 66.5° N/S of Equator), which span between the tropics and the polar regions of Earth. These zones generally have wider temperature ranges throughout t ...
.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 ''Antidesma'' is a genus of tropical plant in the family Phyllanthaceae formally described by Linnaeus in 1753. It is native to tropical Africa, S + E + SE Asia, Australia, and various oceanic islands. The greatest diversity occurs in Southe ...
'', ''
Margaritaria ''Margaritaria'' is a plant genus of the family Phyllanthaceae first published as a genus in 1782. It is the smallest pantropical genus of the Phyllanthaceae and, formerly, of the Euphorbiaceae Euphorbiaceae, the spurge family, is a large fam ...
'', and ''
Phyllanthus ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University P ...
'' 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 ''Antidesma'' is a genus of tropical plant in the family Phyllanthaceae formally described by Linnaeus in 1753. It is native to tropical Africa, S + E + SE Asia, Australia, and various oceanic islands. The greatest diversity occurs in Southe ...
'', ''
Baccaurea ''Baccaurea'Flora Cochinchinensis'' 2: 661. 1790. is a genus of flowering plant belonging to the family Phyllanthaceae. The genus comprises over 100 species, distributed from Malesia to the West Pacific. It is dioecious Dioecy (; ; adj ...
'', ''
Phyllanthus ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University P ...
'', and ''
Uapaca ''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 f ...
'' bear edible fruit. Phyllanthaceae comprises about 2000 species. Depending on the author, these are grouped into 54 to 60
genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...
. 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 ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University P ...
'', 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 ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University P ...
'' (1270), '' Cleistanthus'' (140), ''
Antidesma ''Antidesma'' is a genus of tropical plant in the family Phyllanthaceae formally described by Linnaeus in 1753. It is native to tropical Africa, S + E + SE Asia, Australia, and various oceanic islands. The greatest diversity occurs in Southe ...
'' (100), ''
Aporosa ''Aporosa'' is a genus of flowering plant belonging to the family Phyllanthaceae, first described as a genus in 1825. It is native to China, the Indian Subcontinent, Southeast Asia, Papuasia, and Queensland. These plants are mostly dioecious t ...
'' (90), ''
Uapaca ''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 f ...
'' (60), ''
Baccaurea ''Baccaurea'Flora Cochinchinensis'' 2: 661. 1790. is a genus of flowering plant belonging to the family Phyllanthaceae. The genus comprises over 100 species, distributed from Malesia to the West Pacific. It is dioecious Dioecy (; ; adj ...
'' (50), and ''
Bridelia ''Bridelia'' is a plant genus of the family Phyllanthaceae first described as a genus in 1806."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 ''Heterosavia'' is a genus of the family of Phyllanthaceae first described as a genus in 2008. It is native to the Caribbean and southern Florida. It is found in Bahamas, Cayman Islands, Cuba, Dominican Republic, Florida, Haiti, Jamaica, southwes ...
'' from '' Savia''. Another has separated '' Notoleptopus'' from ''
Leptopus ''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 flo ...
'', and segregated ''
Pseudophyllanthus ''Pseudophyllanthus'' is a monotypic genus of flowering plants in the family Phyllanthaceae. It is one of the eight genera in the tribe Poranthereae. Its closest relative is ''Poranthera''. Its sole species is ''Pseudophyllanthus ovalis''. It i ...
'' and ''
Phyllanthopsis ''Phyllanthopsis'' is a genus of flowering plants in the family Phyllanthaceae. It is one of the eight genera in the tribe Poranthereae. It has two species. The type species is '' Phyllanthopsis phyllanthoides''. ''Phyllanthopsis'' is native to ...
'' from '' Andrachne''. Also, ''
Oreoporanthera ''Poranthera'' is a genus of flowering plant belonging to the family Phyllanthaceae first described as a genus in 1811.
'' has been subsumed into ''
Poranthera ''Poranthera'' is a genus of flowering plant belonging to the family Phyllanthaceae first described as a genus in 1811.
'', while ''
Zimmermannia ''Meineckia'' is a genus of flowering plants in the family Phyllanthaceae first described as a genus in 1858. ''Meineckia'' is one of eight genera in the tribe Poranthereae.Maria S. Vorontsova and Petra Hoffmann. 2008. "A phylogenetic classifica ...
'' and ''
Zimmermanniopsis ''Meineckia'' is a genus of flowering plants in the family Phyllanthaceae first described as a genus in 1858. ''Meineckia'' is one of eight genera in the tribe Poranthereae.Maria S. Vorontsova and Petra Hoffmann. 2008. "A phylogenetic classifica ...
'' have been sunk into ''
Meineckia ''Meineckia'' is a genus of flowering plants in the family Phyllanthaceae first described as a genus in 1858. ''Meineckia'' is one of eight genera in the tribe Poranthereae.Maria S. Vorontsova and Petra Hoffmann. 2008. "A phylogenetic classifica ...
''. The large genus '' Cleistanthus'' is known to be
polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
, 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 Hutchinson John 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
ovule In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...
s in each
locule A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...
of the
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
, a trait that clearly distinguishes it from
Euphorbiaceae Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of a genus in the family. Most spurges, such as '' Euphorbia paralias'', are herbs, but some, ...
. The Phyllanthaceae are nearly all
tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...
s,
shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...
s, or
herbs In general use, herbs are a widely distributed and widespread group of plants, excluding vegetables and other plants consumed for macronutrients, with savory or aromatic properties that are used for flavoring and garnishing food, for medicinal ...
. A few are
climbers Climber may refer to: *Climber, a participant in the activity of climbing *Climber, general name for a vine *Climber, or climbing specialist, a road bicycle racer who can ride especially well on highly inclined roads * Climber (BEAM), a robot that ...
, or
succulents In botany, succulent plants, also known as succulents, are plants with parts that are thickened, fleshy, and engorged, usually to retain water in arid climates or soil conditions. The word ''succulent'' comes from the Latin word ''sucus'', meani ...
, and one species, ''
Phyllanthus fluitans ''Phyllanthus fluitans'', also known as the red root floater, floating spurge, or apple duckweed is a species of free floating aquatic plant and herbaceous perennial in the family Phyllanthaceae. This species is one of the only three non-terre ...
'', is aquatic. Unlike many of the
Euphorbiaceae Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of a genus in the family. Most spurges, such as '' Euphorbia paralias'', are herbs, but some, ...
, none has
latex Latex is an emulsion (stable dispersion) of polymer microparticles in water. Latexes are found in nature, but synthetic latexes are common as well. In nature, latex is found as a milky fluid found in 10% of all flowering plants (angiosperms ...
, and only a very few produce a
resin In polymer chemistry and materials science, resin is a solid or highly viscous substance of plant or synthetic origin that is typically convertible into polymers. Resins are usually mixtures of organic compounds. This article focuses on natu ...
ous exudate. Any
hairs Hair is a protein filament that grows from follicles found in the dermis. Hair is one of the defining characteristics of mammals. The human body, apart from areas of glabrous skin, is covered in follicles which produce thick terminal and f ...
, if present, are almost always simple. Rarely are they branched or scale-like. Thorns and other armament are rare.
Stipule In botany, a stipule is an outgrowth typically borne on both sides (sometimes on just one side) of the base of a leafstalk (the petiole). Stipules are considered part of the anatomy of the leaf of a typical flowering plant, although in many speci ...
s are produced with each
leaf A leaf ( : leaves) is any of the principal appendages of a vascular plant stem, usually borne laterally aboveground and specialized for photosynthesis. Leaves are collectively called foliage, as in "autumn foliage", while the leaves, ste ...
, but in some, these fall before the leaf is fully mature. Leaves are present, except for a few species of ''
Phyllanthus ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University P ...
'' that have flattened, leaflike stems called cladodes that bear flowers along their edges. The leaves are
compound Compound may refer to: Architecture and built environments * Compound (enclosure), a cluster of buildings having a shared purpose, usually inside a fence or wall ** Compound (fortification), a version of the above fortified with defensive struc ...
in '' Bischofia'', but otherwise simple and usually
alternate Alternative or alternate may refer to: Arts, entertainment and media * Alternative (''Kamen Rider''), a character in the Japanese TV series ''Kamen Rider Ryuki'' * ''The Alternative'' (film), a 1978 Australian television film * ''The Alternative ...
. Rarely are they opposite, in
fascicle Fascicle or ''fasciculus'' may refer to: Anatomy and histology * Muscle fascicle, a bundle of skeletal muscle fibers * Nerve fascicle, a bundle of axons (nerve fibers) ** Superior longitudinal fasciculus *** Arcuate fasciculus ** Gracile fas ...
s, or in
whorls A whorl ( or ) is an individual circle, oval, volution or equivalent in a whorled pattern, which consists of a spiral or multiple concentric objects (including circles, ovals and arcs). Whorls in nature File:Photograph and axial plane floral ...
around the stem. The leaf margin is almost always
entire Entire may refer to: * Entire function, a function that is holomorphic on the whole complex plane * Entire (animal), an indication that an animal is not neutered * Entire (botany) This glossary of botanical terms is a list of definitions of ...
, rarely toothed. A petiole is nearly always present, often with a
pulvinus A pulvinus (pl. ''pulvini'') is a joint-like thickening at the base of a plant leaf or leaflet that facilitates growth-independent movement. Pulvini are common, for example, in members of the bean family Fabaceae (Leguminosae) and the prayer plant ...
at its base. The
inflorescence An inflorescence is a group or cluster of flowers arranged on a stem that is composed of a main branch or a complicated arrangement of branches. Morphologically, it is the modified part of the shoot of seed plants where flowers are formed o ...
s are usually in the axils of leaves, rarely below the leaves or at the ends of stems. In ''
Uapaca ''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 f ...
'', the flowers are in a
pseudanthium A pseudanthium (Greek for "false flower"; ) is an inflorescence that resembles a flower. The word is sometimes used for other structures that are neither a true flower nor a true inflorescence. Examples of pseudanthia include flower heads, compos ...
, a tight bundle of flowers that resembles a single flower. Except for four species of ''
Aporosa ''Aporosa'' is a genus of flowering plant belonging to the family Phyllanthaceae, first described as a genus in 1825. It is native to China, the Indian Subcontinent, Southeast Asia, Papuasia, and Queensland. These plants are mostly dioecious t ...
'', the flowers are unisexual, the plants being either monoecious or
dioecious Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproductio ...
. The flowers are
actinomorphic Floral symmetry describes whether, and how, a flower, in particular its perianth, can be divided into two or more identical or mirror-image parts. Uncommonly, flowers may have no axis of symmetry at all, typically because their parts are spirall ...
in form. Detailed illustrations have been published for some of these. The
sepal A sepal () is a part of the flower of angiosperms (flowering plants). Usually green, sepals typically function as protection for the flower in bud, and often as support for the petals when in bloom., p. 106 The term ''sepalum'' was coined b ...
s are three to eight in number, usually free from each other.
Petal Petals are modified Leaf, leaves that surround the reproductive parts of flowers. They are often advertising coloration, brightly colored or unusually shaped to attract pollinators. All of the petals of a flower are collectively known as the ''c ...
s 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
nectar Nectar is a sugar-rich liquid produced by plants in glands called nectaries or nectarines, either within the flowers with which it attracts pollinating animals, or by extrafloral nectaries, which provide a nutrient source to animal mutualists ...
y disk is often present. It may be in the form of a ring, or divided into segments. The
stamen The stamen (plural ''stamina'' or ''stamens'') is the pollen-producing reproductive organ of a flower. Collectively the stamens form the androecium., p. 10 Morphology and terminology A stamen typically consists of a stalk called the filame ...
s are three to ten in number, or rarely more, free or variously fused. The
ovary The ovary is an organ in the female reproductive system that produces an ovum. When released, this travels down the fallopian tube into the uterus, where it may become fertilized by a sperm. There is an ovary () found on each side of the body. ...
is superior. The number of
locule A locule (plural locules) or loculus (plural loculi) (meaning "little place" in Latin) is a small cavity or compartment within an organ or part of an organism (animal, plant, or fungus). In angiosperms (flowering plants), the term ''locule'' usu ...
s in the ovary is highly variable, usually from two to five, but sometimes as many as fifteen. The
placentation Placentation refers to the formation, type and structure, or arrangement of the placenta. The function of placentation is to transfer nutrients, respiratory gases, and water from maternal tissue to a growing embryo, and in some instances to remov ...
is
apical Apical means "pertaining to an apex". It may refer to: *Apical ancestor, refers to the last common ancestor of an entire group, such as a species (biology) or a clan (anthropology) *Apical (anatomy), an anatomical term of location for features loc ...
, with a pair of
ovules In seed plants, the ovule is the structure that gives rise to and contains the female reproductive cells. It consists of three parts: the ''integument'', forming its outer layer, the ''nucellus'' (or remnant of the megasporangium), and the fe ...
hanging by their funicles from the top of each locule. Often, only one of the ovules will develop into a
seed A seed is an embryonic plant enclosed in a protective outer covering, along with a food reserve. The formation of the seed is a part of the process of reproduction in seed plants, the spermatophytes, including the gymnosperm and angiospe ...
. 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 Style is a manner of doing or presenting things and may refer to: * Architectural style, the features that make a building or structure historically identifiable * Design, the process of creating something * Fashion, a prevailing mode of clothing ...
is usually 2-lobed or
bifid Bifid refers to something that is split or cleft into two parts. It may refer to: * Bifid, a variation in the P wave, R wave, or T wave in an echocardiogram in which a wave which usually has a single peak instead has two separate peaks * Bifid ci ...
, sometimes
entire Entire may refer to: * Entire function, a function that is holomorphic on the whole complex plane * Entire (animal), an indication that an animal is not neutered * Entire (botany) This glossary of botanical terms is a list of definitions of ...
, or rarely multifid. The fruit is a
schizocarp A schizocarp is a dry fruit that, when mature, splits up into mericarps. There are different definitions: * Any dry fruit composed of multiple carpels that separate. : Under this definition the mericarps can contain one or more seeds (the m ...
,
drupe In botany, a drupe (or stone fruit) is an indehiscent fruit in which an outer fleshy part (exocarp, or skin, and mesocarp, or flesh) surrounds a single shell (the ''pit'', ''stone'', or '' pyrena'') of hardened endocarp with a seed (''kernel'') ...
, or berry. In some, the
schizocarp A schizocarp is a dry fruit that, when mature, splits up into mericarps. There are different definitions: * Any dry fruit composed of multiple carpels that separate. : Under this definition the mericarps can contain one or more seeds (the m ...
breaks up explosively.


Taxonomy


History

The name "Phyllanthaceae" was first validly published by Ivan Ivanovich Martynov in 1820 in a
Russian Russian(s) refers to anything related to Russia, including: *Russians (, ''russkiye''), an ethnic group of the East Slavic peoples, primarily living in Russia and neighboring countries *Rossiyane (), Russian language term for all citizens and peo ...
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 Euphorbiaceae, the spurge family, is a large family of flowering plants. In English, they are also commonly called euphorbias, which is also the name of a genus in the family. Most spurges, such as '' Euphorbia paralias'', are herbs, but some, ...
. The
monophyly In cladistics for a group of organisms, monophyly is the condition of being a clade—that is, a group of taxa composed only of a common ancestor (or more precisely an ancestral population) and all of its lineal descendants. Monophyletic grou ...
of Euphorbiaceae had long been held in doubt by some, but the first strong evidence of its
polyphyly A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...
came in 1993 with the first
maximum parsimony In phylogenetics, maximum parsimony is an optimality criterion under which the phylogenetic tree that minimizes the total number of character-state changes (or miminizes the cost of differentially weighted character-state changes) is preferred. ...
analysis of
DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...
s of the
gene In biology, the word gene (from , ; "...Wilhelm Johannsen coined the word gene to describe the Mendelian units of heredity..." meaning ''generation'' or ''birth'' or ''gender'') can have several different meanings. The Mendelian gene is a ba ...
rbcL Ribulose-1,5-bisphosphate carboxylase-oxygenase, commonly known by the abbreviations RuBisCo, rubisco, RuBPCase, or RuBPco, is an enzyme () involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is con ...
from a large number of
seed plants A spermatophyte (; ), also known as phanerogam (taxon Phanerogamae) or phaenogam (taxon Phaenogamae), is any plant that produces seeds, hence the alternative name seed plant. Spermatophytes are a subset of the embryophytes or land plants. They inc ...
.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 In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...
analyses have shown that the old concept of Euphorbiaceae consisted of several lineages that did not together form a
clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...
in the order
Malpighiales The Malpighiales comprise one of the largest orders of flowering plants, containing about 36 families and more than species, about 7.8% of the eudicots. The order is very diverse, containing plants as different as the willow, violet, poinsett ...
. 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 The family Pandaceae consists of three genera that were formerly recognized in the Euphorbiaceae. Those are: *''Galearia'' (from tribe Galearieae, subfamily Acalyphoideae, family Euphorbiaceae) *''Microdesmis'' (from tribe Galearieae, subfamily ...
, Phyllanthaceae,
Picrodendraceae Picrodendraceae is a family of flowering plants, consisting of 80 species in 24 genera.Stephens, P.F. (2001 onwards). Angiosperm Phylogeny Website. Version 9, June 2008. http://www.mobot.org/MOBOT/Research/APweb/ These are subtropical to tropical ...
,
Putranjivaceae Putranjivaceae is a rosid family that is composed of 218 species in 2 genera of evergreen tropical trees that are found mainly in the Old World tropics, but with a few species in tropical America. Members of this family have 2-ranked coriaceo ...
,
Peraceae Peraceae Klotzsch is a family of flowering plants in the eudicot order Malpighiales. The family was segregated from the Euphorbiaceae by Johann Friedrich Klotzsch in 1859, and its uniqueness was affirmed by the Royal Botanic Gardens, Kew's Eupho ...
, and
Centroplacaceae Centroplacaceae is a family of flowering plants in the order Malpighiales and is recognized by the APG III system of classification. The family comprises two genera: ''Bhesa'', which was formerly recognized in the Celastraceae, and ''Centroplacus ...
have been removed from it. The obsolete, older concept of Euphorbiaceae, known as Euphorbiaceae
sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular c ...
, 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 ''Centroplacus'' is a genus of the family Centroplacaceae. It was formerly classified in the Phyllanthaceae and given its own tribe, the Centroplaceae. It contains a single species, ''Centroplacus glaucinus''. General information ''C. glaucin ...
'', ''
Paradrypetes ''Paradrypetes'' is a plant genus under the family Rhizophoraceae. It is sometimes included in the Picrodendraceae and was formerly included in the family Euphorbiaceae Euphorbiaceae, the spurge family, is a large family of flowering plants. ...
'', and ''Phyllanoa'' had been placed in Phyllanthaceae, but these are now excluded from the family. ''Centroplacus'' is now in the family
Centroplacaceae Centroplacaceae is a family of flowering plants in the order Malpighiales and is recognized by the APG III system of classification. The family comprises two genera: ''Bhesa'', which was formerly recognized in the Celastraceae, and ''Centroplacus ...
. ''Paradrypetes'' is in
Rhizophoraceae The Rhizophoraceae is a family of tropical or subtropical flowering plants. It includes around 147 species distributed in 15 genera.Setoguchi, H., Kosuge, K., & Tobe, H. (1999). Molecular Phylogeny of Rhizophoraceae Based on rbcL Gene Sequences. ...
. ''Phyllanoa'' is known only from a single specimen. In 1996, this was examined and found to be a species of ''
Rinorea ''Rinorea'' is a genus of plant in family Violaceae. Species include: * '' Rinorea abbreviata'' G. Achoundong & J.J. Bos * '' Rinorea acommanthera'' Gagnep. * '' Rinorea antioquiensis'' Smith & Fernández * '' Rinorea bicornuta'' Hekking * '' R ...
'' (
Violaceae Violaceae is a family of flowering plants established in 1802, consisting of about 1000 species in about 25 genera. It takes its name from the genus ''Viola'', the violets and pansies. Older classifications such as the Cronquist system placed t ...
). 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 ''Chonocentrum'' is a genus of the family Phyllanthaceae described as a genus in 1922. It contains only known species, ''Chonocentrum cyathophorum'', native to the State of Amazonas in northwestern Brazil. The genus is still not well understoo ...
'' Subfamily Antidesmatoideae 6 tribes : Tribe Bischofieae 1 genus ::: '' Bischofia'' : Tribe Uapaceae 1 genus ::: ''
Uapaca ''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 f ...
'' : Tribe Spondiantheae 1 genus ::: '' Spondianthus'' : Tribe Scepeae 8 genera ::: ''
Aporosa ''Aporosa'' is a genus of flowering plant belonging to the family Phyllanthaceae, first described as a genus in 1825. It is native to China, the Indian Subcontinent, Southeast Asia, Papuasia, and Queensland. These plants are mostly dioecious t ...
'' ::: '' Ashtonia'' ::: ''
Baccaurea ''Baccaurea'Flora Cochinchinensis'' 2: 661. 1790. is a genus of flowering plant belonging to the family Phyllanthaceae. The genus comprises over 100 species, distributed from Malesia to the West Pacific. It is dioecious Dioecy (; ; adj ...
'' ::: '' Distichirhops'' ::: '' Maesobotrya'' ::: '' Nothobaccaurea'' ::: '' Protomegabaria'' ::: ''
Richeria ''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 ...
'' : Tribe Jablonskieae 2 genera ::: '' Jablonskia'' ::: '' Celianella'' : Tribe Antidesmateae 5 subtribes :: Subtribe Hieronyminae 1 genus ::: ''
Hieronyma ''Hieronyma'' is a genus in the plant family Phyllanthaceae. It was first described as a genus in 1848. This family was formerly united with spurges, crotons, copperleaves, etc. (Euphorbiaceae), but have turned out to be well distinct. The genu ...
'' :: Subtribe Leptonematinae 1 genus ::: '' Leptonema'' :: Subtribe Martretiinae 2 genera ::: '' Martretia'' ::: '' Apodiscus'' :: Subtribe Hymenocardiinae 2 genera ::: '' Hymenocardia'' ::: '' Didymocistus'' :: Subtribe Antidesmatinae 2 genera ::: ''
Antidesma ''Antidesma'' is a genus of tropical plant in the family Phyllanthaceae formally described by Linnaeus in 1753. It is native to tropical Africa, S + E + SE Asia, Australia, and various oceanic islands. The greatest diversity occurs in Southe ...
'' ::: '' Thecacoris'' Subfamily Phyllanthoideae 4 tribes : Tribe Bridelieae 5 subtribes :: Subtribe Securineginae 2 genera ::: ''
Securinega ''Securinega'' is a genus of plants in the family Phyllanthaceae, first described as a genus in 1789. As presently conceived, the genus is native to Madagascar and the Mascarene Islands in the Indian Ocean. In the past, it was considered to be m ...
'' ::: '' Lachnostylis'' :: Subtribe Saviinae 5 genera ::: '' Savia'' ::: '' Croizatia'' ::: '' Discocarpus'' ::: ''
Gonatogyne ''Gonatogyne'' is a genus of plants in the family Phyllanthaceae Phyllanthaceae 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. Davi ...
'' ::: '' Tacarcuna'' :: Subtribe Pseudolachnostylidinae 4 genera ::: '' Pseudolachnostylis'' ::: ''
Bridelia ''Bridelia'' is a plant genus of the family Phyllanthaceae first described as a genus in 1806.Cleistanthus'' ::: '' Pentabrachion'' :: Subtribe Keayodendrinae 1 genus ::: '' Keayodendron'' :: Subtribe Amanoinae 1 genus ::: '' Amanoa'' : Tribe Phyllantheae 5 genera ::: ''
Phyllanthus ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University P ...
'' (including ''
Breynia ''Breynia'' is a plant genus in the family Phyllanthaceae, first described in 1776. It is native to Southeast Asia, China, the Indian Subcontinent, Papuasia, Australia, and the island of Réunion. The name ''Breynia'' is a conserved name, it ...
'', ''
Glochidion ''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 P ...
'', '' Reverchonia'', and ''
Sauropus The genus ''Sauropus'', of the family Phyllanthaceae, comprises about 40 species of herbs, shrubs or subshrubs, sometimes with woody bases. These plants can be monoecious or dioecious. They are distributed in Southeast Asia, Malesia and Au ...
'') ::: ''
Flueggea ''Flueggea'', the bushweeds, is a genus of shrubs and trees in the family Phyllanthaceae first described as a genus in 1806. It is widespread across much of Asia, Africa, and various oceanic islands, with a few species in South America and on th ...
'' ::: ''
Lingelsheimia ''Lingelsheimia'' is a plant genus in the family (biology), families Phyllanthaceae (and previously placed in the family Putranjivaceae), first described as a genus in 1909. It is native to central Africa and Madagascar.Govaerts, R., Frodin, D.G ...
'' ::: ''
Margaritaria ''Margaritaria'' is a plant genus of the family Phyllanthaceae first published as a genus in 1782. It is the smallest pantropical genus of the Phyllanthaceae and, formerly, of the Euphorbiaceae Euphorbiaceae, the spurge family, is a large fam ...
'' ::: ''
Plagiocladus ''Plagiocladus'' is a genus of the family Phyllanthaceae, first described as a genus in 1987. It contains only one known species, ''Plagiocladus diandrus'', native to central Africa (Cameroon, Gabon Gabon (; ; snq, Ngabu), officially the ...
'' : Tribe Wielandieae 2 subtribes :: Subtribe Astrocasiinae 3 genera ::: '' Astrocasia'' ::: '' Chascotheca'' ::: ''
Heywoodia ''Heywoodia'' is a genus of plants in the Phyllanthaceae first described as a genus in 1907. It contains only one known species, ''Heywoodia lucens'', native to eastern, southeastern, and southern Africa (Kenya, Uganda, Tanzania, Mozambique, Kwa ...
'' :: Subtribe Wielandiinae 3 genera ::: '' Wielandia'' ::: '' Chorisandrachne'' ::: '' Dicoelia'' : Tribe
Poranthereae 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 gener ...
8 genera ::: ''
Poranthera ''Poranthera'' is a genus of flowering plant belonging to the family Phyllanthaceae first described as a genus in 1811.
'' ::: ''
Actephila ''Actephila'' is a genus of plants in the family Phyllanthaceae, first described as a genus in 1826. It is one of 8 genera in the tribe Poranthereae, and is most closely related to '' Leptopus''. The name of the genus is derived from two Greek ...
'' ::: '' Andrachne'' ::: ''
Leptopus ''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 flo ...
'' ::: ''
Meineckia ''Meineckia'' is a genus of flowering plants in the family Phyllanthaceae first described as a genus in 1858. ''Meineckia'' is one of eight genera in the tribe Poranthereae.Maria S. Vorontsova and Petra Hoffmann. 2008. "A phylogenetic classifica ...
'' ::: ''
Oreoporanthera ''Poranthera'' is a genus of flowering plant belonging to the family Phyllanthaceae first described as a genus in 1811.
'' ::: ''
Zimmermannia ''Meineckia'' is a genus of flowering plants in the family Phyllanthaceae first described as a genus in 1858. ''Meineckia'' is one of eight genera in the tribe Poranthereae.Maria S. Vorontsova and Petra Hoffmann. 2008. "A phylogenetic classifica ...
'' ::: ''
Zimmermanniopsis ''Meineckia'' is a genus of flowering plants in the family Phyllanthaceae first described as a genus in 1858. ''Meineckia'' is one of eight genera in the tribe Poranthereae.Maria S. Vorontsova and Petra Hoffmann. 2008. "A phylogenetic classifica ...
''


Genera

The list of 54 genera below is from the 2006 revision of Phyllanthaceae by Petra Hoffmann and co-workers. In their treatment, '' Blotia'' and ''
Petalodiscus ''Wielandia'' is a genus of flowering plant, of the family Phyllanthaceae first described as a genus in 1858. The plants are native to Kenya, Madagascar, and to various other islands in the Indian Ocean The Indian Ocean is the third-largest ...
'' were sunk into '' Wielandia'' and ''
Richeriella ''Flueggea'', the bushweeds, is a genus of shrubs and trees in the family Phyllanthaceae first described as a genus in 1806. It is widespread across much of Asia, Africa, and various oceanic islands, with a few species in South America and on th ...
'' into ''
Flueggea ''Flueggea'', the bushweeds, is a genus of shrubs and trees in the family Phyllanthaceae first described as a genus in 1806. It is widespread across much of Asia, Africa, and various oceanic islands, with a few species in South America and on th ...
''. ''
Breynia ''Breynia'' is a plant genus in the family Phyllanthaceae, first described in 1776. It is native to Southeast Asia, China, the Indian Subcontinent, Papuasia, Australia, and the island of Réunion. The name ''Breynia'' is a conserved name, it ...
'', ''
Glochidion ''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 P ...
'', '' Reverchonia'', and ''
Sauropus The genus ''Sauropus'', of the family Phyllanthaceae, comprises about 40 species of herbs, shrubs or subshrubs, sometimes with woody bases. These plants can be monoecious or dioecious. They are distributed in Southeast Asia, Malesia and Au ...
'' were recommended to be subsumed into ''
Phyllanthus ''Phyllanthus'' is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University P ...
'', 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 Putranjivaceae is a rosid family that is composed of 218 species in 2 genera of evergreen tropical trees that are found mainly in the Old World tropics, but with a few species in tropical America. Members of this family have 2-ranked coriaceo ...
.


Phylogeny

The revision of Phyllanthaceae by Hoffmann and co-authors was based on two molecular phylogenetic studies that were published in 2005. Since the revision, phylogenetic studies have been done on some of the tribes. The phylogenetic tree shown below is based on the results of several studies. Fifty-one genera are represented. ''Chonocentrum''(Phyllanthaceae,
incertae sedis ' () or ''problematica'' is a term used for a taxonomic group where its broader relationships are unknown or undefined. Alternatively, such groups are frequently referred to as "enigmatic taxa". In the system of open nomenclature, uncertain ...
), 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. In the phylogeny shown below, statistical support for the clades was measured by bootstrap percentage. All branches shown below have
maximum parsimony In phylogenetics, maximum parsimony is an optimality criterion under which the phylogenetic tree that minimizes the total number of character-state changes (or miminizes the cost of differentially weighted character-state changes) is preferred. ...
bootstrap support of at least 70%.


References


External links

* * *
Phyllanthaceae
At

At
Missouri Botanical Garden Website
* *
Key to Euphorbiaceae sensu lato
At
Index of /herbarium/keys
{{Taxonbar, from=Q133206 Malpighiales families