Phyllanthaceae
Encyclopedia
Phyllanthaceae is a family
of flowering plant
s in the eudicot order
Malpighiales
. It is most closely related to the family Picrodendraceae
.
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.
Some species of Andrachne
, Antidesma
, Margaritaria
, and Phyllanthus
are in cultivation. A few species of Antidesma
, Baccaurea
, Phyllanthus
, and Uapaca
provide 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 "absorbed" by others, while other genera have recently been divided. 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).
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 were sunk into Wielandia and Richeriella into Flueggea, while Breynia, Glochidion, Reverchonia, and Sauropus were subsumed into Phyllanthus. For older changes in the circumscription
of genera, see the synonymy
at Taxonomy of the Phyllanthaceae
.
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.
s in each locule
of the ovary
, a trait that clearly distinguishes it from Euphorbiaceae
.
The Phyllanthaceae are nearly all tree
s, shrub
s, or herbs
. A few are climbers
, or succulents
, and one species, Phyllanthus fluitans, is aquatic
. Unlike many of the Euphorbiaceae
, none have latex
, and only a very few produce a resin
ous exudate. Any hairs
, if present, are almost always simple. Rarely are they branched or scale-like. Thorns and other armament are rare.
Stipule
s 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 fascicle
s, 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 inflorescence
s 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 sepal
s are three to eight in number, usually free from each other. Petal
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 nectary disk is often present. It may be in the form of a ring, or divided into segments. The stamen
s are three to ten in number, or rarely more, free or variously fused.
The ovary
is superior. The number of locule
s 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.
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 sequence
s of the gene
rbcL
from a large number of seed plants
. 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. 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. 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.
, 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
Subfamily Phyllanthoideae 4 tribes
The phylogenetic tree shown below is based on the results of several studies. 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.
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%.
Family (biology)
In biological classification, family is* a taxonomic rank. Other well-known ranks are life, domain, kingdom, phylum, class, order, genus, and species, with family fitting between order and genus. As for the other well-known ranks, there is the option of an immediately lower rank, indicated by the...
of flowering plant
Flowering plant
The flowering plants , also known as Angiospermae or Magnoliophyta, are the most diverse group of land plants. Angiosperms are seed-producing plants like the gymnosperms and can be distinguished from the gymnosperms by a series of synapomorphies...
s in the eudicot order
Order (biology)
In scientific classification used in biology, the order is# a taxonomic rank used in the classification of organisms. Other well-known ranks are life, domain, kingdom, phylum, class, family, genus, and species, with order fitting in between class and family...
Malpighiales
Malpighiales
Malpighiales is one of the largest orders of flowering plants, containing about 16000 species, approximately 7.8% of the eudicots. The order is very diverse and hard to recognize except with molecular phylogenetic evidence. It is not part of any of the classification systems that are based only on...
. It is most closely related to the family Picrodendraceae
Picrodendraceae
Picrodendraceae is a family of flowering plants, consisting of 80 species in 24 genera. These are subtropical to tropical and found in New Guinea, Australia, New Caledonia, Madagascar, continental Africa as well as tropical America....
.
The Phyllanthaceae are most numerous in the tropics
Tropics
The tropics is a region of the Earth surrounding the Equator. It is limited in latitude by the Tropic of Cancer in the northern hemisphere at approximately N and the Tropic of Capricorn in the southern hemisphere at S; these latitudes correspond to the axial tilt of the Earth...
, with many in the south temperate zone, and a few ranging as far north as the middle of the north temperate zone.
Some species of Andrachne
Andrachne
Andrachne is a genus of flowering plants in the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. Andrachne comprises 22 species. They are monoecious herbs or subshrubs, native to semideserts and desert margins of the Americas, southern Europe, North Africa, and South...
, Antidesma
Antidesma
Antidesma is a species of tropical plant in the Phyllanthaceae. This is a variable plant which may be short and shrubby or tall and erect, approaching 30 meters in height.It has large oval shaped leathery evergreen leaves up to about 20 centimeters long and seven wide. The flowers have a strong,...
, Margaritaria
Margaritaria
Margaritaria is a plant genus of the Phyllanthaceae family. With 14 species, this is the smallest pantropical genus of the Phyllanthaceae and, formerly, of the Euphorbiaceae.-Synonymy:This genus is known under several other names:...
, and Phyllanthus
Phyllanthus
Phyllanthus is the largest genus in the flowering plant family Phyllanthaceae. Estimates of the number 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...
are in cultivation. A few species of Antidesma
Antidesma
Antidesma is a species of tropical plant in the Phyllanthaceae. This is a variable plant which may be short and shrubby or tall and erect, approaching 30 meters in height.It has large oval shaped leathery evergreen leaves up to about 20 centimeters long and seven wide. The flowers have a strong,...
, Baccaurea
Baccaurea
Baccaurea is a genus of flowering plant belonging to the family Phyllanthaceae. The genus comprises over 100 species, distributed from Indomalesia to the West Pacific.- Selected species :...
, Phyllanthus
Phyllanthus
Phyllanthus is the largest genus in the flowering plant family Phyllanthaceae. Estimates of the number 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...
, and Uapaca
Uapaca
Uapaca is a genus of plant, in the family Phyllanthaceae and is the only genus comprised in the subtribe Uapacinae. It comprises about 60 species, of which 49 are confined to continental Africa.-Synonyms:This genus is also known as:...
provide edible fruit.
Phyllanthaceae comprises about 2000 species. Depending on the author, these are grouped into 54 to 60 genera
Genus
In biology, a genus is a low-level taxonomic rank used in the biological classification of living and fossil organisms, which is an example of definition by genus and differentia...
. 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 "absorbed" by others, while other genera have recently been divided. 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).
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 were sunk into Wielandia and Richeriella into Flueggea, while Breynia, Glochidion, Reverchonia, and Sauropus were subsumed into Phyllanthus. For older changes in the circumscription
Circumscription (taxonomy)
In taxonomy, circumscription is the definition of the limits of a taxonomic group of organisms. One goal of taxonomy is to achieve a stable circumscription for every taxonomic group. Achieving stability can be simple or difficult....
of genera, see the synonymy
Synonym (taxonomy)
In scientific nomenclature, a synonym is a scientific name that is or was used for a taxon of organisms that also goes by a different scientific name. For example, Linnaeus was the first to give a scientific name to the Norway spruce, which he called Pinus abies...
at Taxonomy of the Phyllanthaceae
Taxonomy of the Phyllanthaceae
Here is a full taxonomy of the family Phyllanthaceae. Its circumscription is nearly coincident to that of ex-subfamily Phyllanthoideae of the Euphorbiaceae, where this family was split from...
.
Since Phyllanthaceae was revised in 2006, one paper has removed Heterosavia from Savia. Another has separated Notoleptopus
Notoleptopus
Notoleptopus is a genus of flowering plants in the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. It has only one species, Notoleptopus decaisnei...
from Leptopus, and segregated Pseudophyllanthus
Pseudophyllanthus
Pseudophyllanthus is a genus of flowering plants in the family Phyllanthaceae. It is one of the eight genera in the tribe Poranthereae. Its closest relative is Poranthera....
and Phyllanthopsis
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....
from Andrachne
Andrachne
Andrachne is a genus of flowering plants in the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. Andrachne comprises 22 species. They are monoecious herbs or subshrubs, native to semideserts and desert margins of the Americas, southern Europe, North Africa, and South...
. Also, Oreoporanthera has been subsumed into Poranthera
Poranthera
Poranthera is a genus of flowering plant belonging to the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. Its closest relative is Pseudophyllanthus....
, while Zimmermannia and Zimmermanniopsis have been sunk into Meineckia
Meineckia
Meineckia is a genus of flowering plants in the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. It comprises about 30 species. They are native to the Americas, South Asia, Africa, and Madagascar. The genus is particularly well represented in Madagascar. Species of...
. The large genus Cleistanthus is known to be polyphyletic, but further studies will be needed before it can be revised.
Genera
|
Breynia Breynia is a plant genus in the family Phyllanthaceae. It has 35 species, distributed from India to Australia.-Synonymy:This genus is also known under several names:*Foersteria Scop.*Forsteria Steud.*Melanthes Blume... Bridelia Bridelia is a plant genus of the family Phyllanthaceae. This genus comprises approximately between 60 and 70 species, found from Africa, Australia and Asia.... Celianella Celianella is a genus of flowering plant belonging to the family Phyllanthaceae. Its isotype is Celianella montana.... Chonocentrum Chonocentrum is a genus of the family Phyllanthaceae. The genus is still not well understood; W. John Hayden has observed that the plant seems have been collected just once, in upper Rio Negro of Brazil in the 1850s, and suggests that it has been misplaced taxonomically.- References :* Hayden, W.... Cleistanthus Cleistanthus is a plant genus of the family Phyllanthaceae. The genus comprises 140 species, found from Africa to the Pacific Islands. Cleistanthus collinus is known for being toxic and frequently used for homicidal or suicidal purposes.-Synonymy:... Didymocistus Didymocistus chrysadenius is a species of the family Phyllanthaceae. Its genus, Didymocistus is monotypic and it is recently described from Amazonia.... Discocarpus Discocarpus is a genus of the plant family Phyllanthaceae. It is a small genus with 3 species, which are found in Amazonia.... Glochidion Glochidion are a taxon of plants in the family Phyllanthaceae. It comprises about 300 species, distributed from Madagascar to the Pacific Islands... |
Heywoodia Heywoodia lucens is a plant species of the family Phyllanthaceae. Its genus, Heywoodia, is monotypic. The species is native to tropical East Africa.... Hieronyma Hieronyma is a genus belonging to the plant family Phyllanthaceae. This family was formerly united with spurges, crotons, copperleaves, etc , but have turned out to be well distinct.... Hymenocardia Hymenocardia is a genus of the family Phyllanthaceae. The genus has 7 or 8 species, native mainly to tropical Africa.... Lachnostylis Lachnostylis is a genus of the family Phyllanthaceae. It comprises 2 species, native to the Eastern Cape and the Western Cape of South Africa. It is often included in Savia.... Leptonema Leptonema is a genus of flowering plant belonging to the family Phyllanthaceae.... Maesobotrya Maesobotrya is a genus of flowering plant belonging to the family Phyllanthaceae.-Synonymy:The genus is also known as:*Staphysora Pierre... Margaritaria Margaritaria is a plant genus of the Phyllanthaceae family. With 14 species, this is the smallest pantropical genus of the Phyllanthaceae and, formerly, of the Euphorbiaceae.-Synonymy:This genus is known under several other names:... |
Meineckia Meineckia is a genus of flowering plants in the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. It comprises about 30 species. They are native to the Americas, South Asia, Africa, and Madagascar. The genus is particularly well represented in Madagascar. Species of... Nothobaccaurea Nothobaccaurea is a genus of flowering plant belonging to the family Phyllanthaceae, comprising 2 species found in the Pacific. The genus is named for its false resemblance with Baccaurea.... Oreoporanthera Oreoporanthera is a genus of flowering plant belonging to the family Phyllanthaceae.It contains at least two species, Oreoporanthera alpina and Oreoporanthera petalifera.... Pentabrachion Pentabrachion is a plant genus from the family Phyllanthaceae.... Petalodiscus Petalodiscus is a genus of the family Phyllanthaceae. The genus is sometimes included in Savia.... Phyllanthus Phyllanthus is the largest genus in the flowering plant family Phyllanthaceae. Estimates of the number 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... Poranthera Poranthera is a genus of flowering plant belonging to the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. Its closest relative is Pseudophyllanthus.... Protomegabaria Protomegabaria is a genus of flowering plant belonging to the family Phyllanthaceae.... |
Reverchonia Reverchonia arenaria is a plant species of the Phyllanthaceae family and the sole species in the genus Reverchonia. It is a sand dune annual and confined to the Southwestern United States and adjacent Mexico.... Richeria Richeria is a genus of flowering plant belonging to the family Phyllanthaceae.-Synonyms:*Bellevalia Roem. & Schult.*Guarania Wedd. ex Baill.... Richeriella Richeriella is a plant genus of the Phyllanthaceae family. It comprises 2 species, found from Southeast China to Central Malesia.... Sauropus The genus Sauropus, of the Phyllanthaceae family, 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 Australia.... Securinega Securinega was a genus of plant under the family Phyllanthaceae and the sole genus in the subtribe Securineginae. It consisted of several species, all of which are now synonyms for species under the genus Flueggea:... Spondianthus Spondianthus is a genus of plant, in the family Phyllanthaceae and is the only genus comprised in the subtribe Spondianthinae. The genus is found in tropical Africa.... Thecacoris Thecacoris is a genus of flowering plant belonging to the family Phyllanthaceae.-Synonyms:*Baccaureopsis Pax*Cyathogyne Müll.Arg.*Henribaillonia Kuntze... Uapaca Uapaca is a genus of plant, in the family Phyllanthaceae and is the only genus comprised in the subtribe Uapacinae. It comprises about 60 species, of which 49 are confined to continental Africa.-Synonyms:This genus is also known as:... Wielandia Wielandia is a genus of flowering plant, of the family Phyllanthaceae. Members of this genus are shrubs found in the Seychelles Islands.... Zimmermannia Zimmermannia is a genus of plant under the family Phyllanthaceae. The seven species appear similar, but careful examination of the polen of each has shown that they are distinct species.... Zimmermanniopsis Zimmermanniopsis is a genus of plant under the family Phyllanthaceae. The genus resembles Zimmermannia, hence its name.... |
Description
The description here is from Hoffmann, except for a few additions from Webster and Hutchinson 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 ovuleOvule
Ovule means "small egg". 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 , and the megaspore-derived female gametophyte in its center...
s in each locule
Locule
A locule is a small cavity or compartment within an organ or part of an organism ....
of the ovary
Ovary (plants)
In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule and is located above or below or at the point of connection with the base of the petals and sepals...
, a trait that clearly distinguishes it from Euphorbiaceae
Euphorbiaceae
Euphorbiaceae, the Spurge family are a large family of flowering plants with 300 genera and around 7,500 species. Most are herbs, but some, especially in the tropics, are also shrubs or trees. Some are succulent and resemble cacti....
.
The Phyllanthaceae are nearly all tree
Tree
A tree is a perennial woody plant. It is most often defined as a woody plant that has many secondary branches supported clear of the ground on a single main stem or trunk with clear apical dominance. A minimum height specification at maturity is cited by some authors, varying from 3 m to...
s, shrub
Shrub
A shrub or bush is distinguished from a tree by its multiple stems and shorter height, usually under 5–6 m tall. A large number of plants may become either shrubs or trees, depending on the growing conditions they experience...
s, or herbs
Herbaceous plant
A herbaceous plant is a plant that has leaves and stems that die down at the end of the growing season to the soil level. They have no persistent woody stem above ground...
. A few are climbers
Vine
A vine in the narrowest sense is the grapevine , but more generally it can refer to any plant with a growth habit of trailing or scandent, that is to say climbing, stems or runners...
, or succulents
Succulent plant
Succulent plants, also known as succulents or fat plants, are water-retaining plants adapted to arid climates or soil conditions. Succulent plants store water in their leaves, stems, and also in roots...
, and one species, Phyllanthus fluitans, is aquatic
Aquatic plant
Aquatic plants are plants that have adapted to living in aquatic environments. They are also referred to as hydrophytes or aquatic macrophytes. These plants require special adaptations for living submerged in water, or at the water's surface. Aquatic plants can only grow in water or in soil that is...
. Unlike many of the Euphorbiaceae
Euphorbiaceae
Euphorbiaceae, the Spurge family are a large family of flowering plants with 300 genera and around 7,500 species. Most are herbs, but some, especially in the tropics, are also shrubs or trees. Some are succulent and resemble cacti....
, none have latex
Latex
Latex is the stable dispersion of polymer microparticles in an aqueous medium. Latexes may be natural or synthetic.Latex as found in nature is a milky fluid found in 10% of all flowering plants . It is a complex emulsion consisting of proteins, alkaloids, starches, sugars, oils, tannins, resins,...
, and only a very few produce a resin
Resin
Resin in the most specific use of the term is a hydrocarbon secretion of many plants, particularly coniferous trees. Resins are valued for their chemical properties and associated uses, such as the production of varnishes, adhesives, and food glazing agents; as an important source of raw materials...
ous exudate. Any hairs
Indumentum
The indumentum is a covering of fine hairs or bristles on a plant or insect.In plants, the indumentum types are:*pubescent*hirsute*pilose*villous*tomentose*stellate*scabrous*scurfy...
, if present, are almost always simple. Rarely are they branched or scale-like. Thorns and other armament are rare.
Stipule
Stipule
In botany, stipule is a term coined by Linnaeus which refers to outgrowths borne on either side of the base of a leafstalk...
s are produced with each leaf
Leaf
A leaf is an organ of a vascular plant, as defined in botanical terms, and in particular in plant morphology. Foliage is a mass noun that refers to leaves as a feature of plants....
, 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
Phyllotaxis
In botany, phyllotaxis or phyllotaxy is the arrangement of leaves on a plant stem .- Pattern structure :...
, in fascicle
Fascicle (botany)
In botany, a fascicle is a bundle of leaves or flowers, or of the vascular tissues that supply these organs with nutrients...
s, or in whorls
Phyllotaxis
In botany, phyllotaxis or phyllotaxy is the arrangement of leaves on a plant stem .- Pattern structure :...
around the stem. The leaf margin is almost always entire
Leaf shape
In botany, leaf shape is characterised with the following terms :* Acicular : Slender and pointed, needle-like* Acuminate : Tapering to a long point...
, rarely toothed. A petiole
Petiole (botany)
In botany, the petiole is the stalk attaching the leaf blade to the stem. The petiole usually has the same internal structure as the stem. Outgrowths appearing on each side of the petiole are called stipules. Leaves lacking a petiole are called sessile, or clasping when they partly surround the...
is nearly always present, often with a pulvinus
Pulvinus
A pulvinus is a joint-like thickening at the base of a plant leaf or leaflet that facilitates growth-independent movement. It consists of a core of vascular tissue within a flexible, bulky cylinder of thin-walled parenchyma cells...
at its base.
The inflorescence
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. Strictly, it is the part of the shoot of seed plants where flowers are formed and which is accordingly modified...
s 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
Pseudanthium
A pseudanthium or flower head is a special type of inflorescence, in which several flowers are grouped together to form a flower-like structure. The real flowers are generally small and greatly reduced, but can sometimes be quite large...
, 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 sepal
Sepal
A sepal is a part of the flower of angiosperms . Collectively the sepals form the calyx, which is the outermost whorl of parts that form a flower. Usually green, sepals have the typical function of protecting the petals when the flower is in bud...
s are three to eight in number, usually free from each other. Petal
Petal
Petals are modified leaves that surround the reproductive parts of flowers. They often are brightly colored or unusually shaped to attract pollinators. Together, all of the petals of a flower are called a corolla. Petals are usually accompanied by another set of special leaves called sepals lying...
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 nectary disk is often present. It may be in the form of a ring, or divided into segments. The stamen
Stamen
The stamen is the pollen producing reproductive organ of a flower...
s are three to ten in number, or rarely more, free or variously fused.
The ovary
Ovary (plants)
In the flowering plants, an ovary is a part of the female reproductive organ of the flower or gynoecium. Specifically, it is the part of the pistil which holds the ovule and is located above or below or at the point of connection with the base of the petals and sepals...
is superior. The number of locule
Locule
A locule is a small cavity or compartment within an organ or part of an organism ....
s 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
Ovule
Ovule means "small egg". 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 , and the megaspore-derived female gametophyte in its center...
from the top of each locule. Often, only one of the ovules will develop into a seed
Seed
A seed is a small embryonic plant enclosed in a covering called the seed coat, usually with some stored food. It is the product of the ripened ovule of gymnosperm and angiosperm plants which occurs after fertilization and some growth within the mother plant...
. A single, massive obturator
Obturator
Obturator may refer to:In medicine:* Obturator artery* Obturator nerve* Obturator vein* Obturator externus muscle* Obturator internus muscle* Obturator membrane* Obturator canal* Obturator foramen...
may cover the micropyles
Ovule
Ovule means "small egg". 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 , and the megaspore-derived female gametophyte in its center...
of both ovules, or each ovule may have its own thin obturator. The megagametophyte
Ovule
Ovule means "small egg". 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 , and the megaspore-derived female gametophyte in its center...
is of the Polygonum type. The style is usually 2-lobed or bifid, sometimes entire, or rarely multifid.
The fruit is a schizocarp
Schizocarp
A schizocarp is a dry fruit that develops from multiple carpels. When mature it splits up into mericarps. Mericarps are often 1-seeded as in, for example, Malva, Malvastrum, and Sida...
, drupe
Drupe
In botany, a drupe is a fruit in which an outer fleshy part surrounds a shell of hardened endocarp with a seed inside. These fruits develop from a single carpel, and mostly from flowers with superior ovaries...
, or berry
Berry
The botanical definition of a berry is a fleshy fruit produced from a single ovary. Grapes are an example. The berry is the most common type of fleshy fruit in which the entire ovary wall ripens into an edible pericarp. They may have one or more carpels with a thin covering and fleshy interiors....
. In some, the schizocarp breaks up explosively.
History
The name "Phyllanthaceae" was first validly published by Ivan Ivanovich Martynov in 1820 in a RussianRussian language
Russian is a Slavic language used primarily in Russia, Belarus, Uzbekistan, Kazakhstan, Tajikistan and Kyrgyzstan. It is an unofficial but widely spoken language in Ukraine, Moldova, Latvia, Turkmenistan and Estonia and, to a lesser extent, the other countries that were once constituent republics...
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
Euphorbiaceae, the Spurge family are a large family of flowering plants with 300 genera and around 7,500 species. Most are herbs, but some, especially in the tropics, are also shrubs or trees. Some are succulent and resemble cacti....
. The monophyly
Monophyly
In common cladistic usage, a monophyletic group is a taxon which forms a clade, meaning that it contains all the descendants of the possibly hypothetical closest common ancestor of the members of the group. The term is synonymous with the uncommon term holophyly...
of Euphorbiaceae had long been held in doubt by some, but the first strong evidence of its polyphyly
Polyphyly
A polyphyletic group is one whose members' last common ancestor is not a member of the group.For example, the group consisting of warm-blooded animals is polyphyletic, because it contains both mammals and birds, but the most recent common ancestor of mammals and birds was cold-blooded...
came in 1993 with the first maximum parsimony
Maximum parsimony
Parsimony is a non-parametric statistical method commonly used in computational phylogenetics for estimating phylogenies. Under parsimony, the preferred phylogenetic tree is the tree that requires the least evolutionary change to explain some observed data....
analysis of DNA sequence
DNA sequence
The sequence or primary structure of a nucleic acid is the composition of atoms that make up the nucleic acid and the chemical bonds that bond those atoms. Because nucleic acids, such as DNA and RNA, are unbranched polymers, this specification is equivalent to specifying the sequence of...
s of the gene
Gene
A gene is a molecular unit of heredity of a living organism. It is a name given to some stretches of DNA and RNA that code for a type of protein or for an RNA chain that has a function in the organism. Living beings depend on genes, as they specify all proteins and functional RNA chains...
rbcL
RuBisCO
Ribulose-1,5-bisphosphate carboxylase oxygenase, commonly known by the shorter name RuBisCO, is an enzyme involved in the first major step of carbon fixation, a process by which atmospheric carbon dioxide is converted by plants to energy-rich molecules such as glucose. RuBisCo is an abbreviation...
from a large number of seed plants
Spermatophyte
The spermatophytes comprise those plants that produce seeds. They are a subset of the embryophytes or land plants...
. Since the 1993 study, all subsequent phylogenetic
Phylogenetics
In biology, phylogenetics is the study of evolutionary relatedness among groups of organisms , which is discovered through molecular sequencing data and morphological data matrices...
analyses have shown that the old concept of Euphorbiaceae consisted of several lineages
Lineage (evolution)
An evolutionary lineage is a sequence of species, that form a line of descent, each new species the direct result of speciation from an immediate ancestral species. Lineages are subsets of the evolutionary tree of life. Lineages are often determined by the techniques of molecular systematics.-...
that did not together form a clade
Clade
A clade is a group consisting of a species and all its descendants. In the terms of biological systematics, a clade is a single "branch" on the "tree of life". The idea that such a "natural group" of organisms should be grouped together and given a taxonomic name is central to biological...
in the order Malpighiales
Malpighiales
Malpighiales is one of the largest orders of flowering plants, containing about 16000 species, approximately 7.8% of the eudicots. The order is very diverse and hard to recognize except with molecular phylogenetic evidence. It is not part of any of the classification systems that are based only on...
. Euphorbiaceae is now defined as a much smaller family than it had been in the twentieth century. Pandaceae
Pandaceae
The family Pandaceae consists of three genera that were formerly recognized in the Euphorbiaceae. Those are:*Galearia...
, Phyllanthaceae, Picrodendraceae
Picrodendraceae
Picrodendraceae is a family of flowering plants, consisting of 80 species in 24 genera. These are subtropical to tropical and found in New Guinea, Australia, New Caledonia, Madagascar, continental Africa as well as tropical America....
, Putranjivaceae
Putranjivaceae
The rosid family Putranjivaceae is composed of about 210 species of evergreen tropical trees distributed into 4 genera. Members of this family have 2-ranked coriaceous leaves, which, if fresh, typically have a radish-like or peppery taste. The flowers are fasciculate and usually small, and the...
, Peraceae, and Centroplacaceae
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, which was formerly recognized in the Euphorbiaceae...
have been removed from it.
The obsolete, older concept of Euphorbiaceae, known as Euphorbiaceae sensu lato, is sometimes still used for continuity and convenience. 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.
Classification
In the past, the genera CentroplacusCentroplacus
Centroplacus is a genus of the family Centroplacaceae. It was formerly classified in the Phyllanthaceae and given its own tribe, the Centroplaceae....
, Paradrypetes
Paradrypetes
Paradrypetes is a plant genus under the family Picrodendraceae. It is sometimes included in the Rhizophoraceae and was formerly included in the Euphorbiaceae family....
, and Phyllanoa had been placed in Phyllanthaceae, but these are now excluded from the family. Centroplacus is now in the family Centroplacaceae
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, which was formerly recognized in the Euphorbiaceae...
. Paradrypetes is in Rhizophoraceae
Rhizophoraceae
Rhizophoraceae is a family constituted by tropical or subtropical flowering plants. Among the better known members are mangrove trees of the genus Rhizophora...
. Phyllanoa is known only from a single specimen. In 1996, this was examined and found to be a species of Rinorea
Rinorea
Rinorea is a genus of plant in family Violaceae.Species include:* Rinorea antioquiensis, Smith & Fernández* Rinorea bicornuta, Hekking* Rinorea brachythrix, Blake* Rinorea cordata, Smith & Fernández* Rinorea crenata, Blake...
(Violaceae
Violaceae
Violaceae are a family of flowering plants consisting of about 800 species in 21 genera. It takes its name from the genus Viola, the violets and pansies.Older classifications such as the Cronquist system placed Violaceae in an order named after it, the Violales...
).
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
-
- UapacaUapacaUapaca is a genus of plant, in the family Phyllanthaceae and is the only genus comprised in the subtribe Uapacinae. It comprises about 60 species, of which 49 are confined to continental Africa.-Synonyms:This genus is also known as:...
- Uapaca
-
- Tribe Spondiantheae 1 genus
-
- SpondianthusSpondianthusSpondianthus is a genus of plant, in the family Phyllanthaceae and is the only genus comprised in the subtribe Spondianthinae. The genus is found in tropical Africa....
- Spondianthus
-
- Tribe Scepeae 8 genera
-
- AporosaAporosaAporosa is a genus of flowering plant belonging to the family Phyllanthaceae. The genus comprises about 75 species, distributed from Indomalaya to the Solomon Islands.-Synonymy:The genus is also known as:*Aporusa Blume, orth...
- AshtoniaAshtoniaAshtonia is a genus of flowering plant belonging to the family Phyllanthaceae. It comprises 2 species, found in the Malay Peninsula and Borneo....
- BaccaureaBaccaureaBaccaurea is a genus of flowering plant belonging to the family Phyllanthaceae. The genus comprises over 100 species, distributed from Indomalesia to the West Pacific.- Selected species :...
- Distichirrhops
- MaesobotryaMaesobotryaMaesobotrya is a genus of flowering plant belonging to the family Phyllanthaceae.-Synonymy:The genus is also known as:*Staphysora Pierre...
- NothobaccaureaNothobaccaureaNothobaccaurea is a genus of flowering plant belonging to the family Phyllanthaceae, comprising 2 species found in the Pacific. The genus is named for its false resemblance with Baccaurea....
- ProtomegabariaProtomegabariaProtomegabaria is a genus of flowering plant belonging to the family Phyllanthaceae....
- RicheriaRicheriaRicheria is a genus of flowering plant belonging to the family Phyllanthaceae.-Synonyms:*Bellevalia Roem. & Schult.*Guarania Wedd. ex Baill....
- Aporosa
-
- Tribe Jablonskieae 2 genera
-
- Jablonskia
- CelianellaCelianellaCelianella is a genus of flowering plant belonging to the family Phyllanthaceae. Its isotype is Celianella montana....
-
- Tribe Antidesmateae 5 subtribes
- Subtribe Hieronyminae 1 genus
- HieronymaHieronymaHieronyma is a genus belonging to the plant family Phyllanthaceae. This family was formerly united with spurges, crotons, copperleaves, etc , but have turned out to be well distinct....
- Hieronyma
- Subtribe Leptonematinae 1 genus
- LeptonemaLeptonemaLeptonema is a genus of flowering plant belonging to the family Phyllanthaceae....
- Leptonema
- Subtribe Martretiinae 2 genera
- Subtribe Hymenocardiinae 2 genera
- HymenocardiaHymenocardiaHymenocardia is a genus of the family Phyllanthaceae. The genus has 7 or 8 species, native mainly to tropical Africa....
- DidymocistusDidymocistusDidymocistus chrysadenius is a species of the family Phyllanthaceae. Its genus, Didymocistus is monotypic and it is recently described from Amazonia....
- Hymenocardia
- Subtribe Antidesmatinae 2 genera
- AntidesmaAntidesmaAntidesma is a species of tropical plant in the Phyllanthaceae. This is a variable plant which may be short and shrubby or tall and erect, approaching 30 meters in height.It has large oval shaped leathery evergreen leaves up to about 20 centimeters long and seven wide. The flowers have a strong,...
- ThecacorisThecacorisThecacoris is a genus of flowering plant belonging to the family Phyllanthaceae.-Synonyms:*Baccaureopsis Pax*Cyathogyne Müll.Arg.*Henribaillonia Kuntze...
- Antidesma
- Subtribe Hieronyminae 1 genus
Subfamily Phyllanthoideae 4 tribes
- Tribe BridelieaeBridelieaeBridelieae is a tribe of the family Phyllanthaceae. It comprises 2 genera....
5 subtribes- Subtribe Securineginae 2 genera
- SecurinegaSecurinegaSecurinega was a genus of plant under the family Phyllanthaceae and the sole genus in the subtribe Securineginae. It consisted of several species, all of which are now synonyms for species under the genus Flueggea:...
- LachnostylisLachnostylisLachnostylis is a genus of the family Phyllanthaceae. It comprises 2 species, native to the Eastern Cape and the Western Cape of South Africa. It is often included in Savia....
- Securinega
- Subtribe Saviinae 5 genera
- Savia
- Croizatia
- DiscocarpusDiscocarpusDiscocarpus is a genus of the plant family Phyllanthaceae. It is a small genus with 3 species, which are found in Amazonia....
- Gonatogyne
- Tacarcuna
- Subtribe Pseudolachnostylidinae 4 genera
- Pseudolachnostylis
- BrideliaBrideliaBridelia is a plant genus of the family Phyllanthaceae. This genus comprises approximately between 60 and 70 species, found from Africa, Australia and Asia....
- CleistanthusCleistanthusCleistanthus is a plant genus of the family Phyllanthaceae. The genus comprises 140 species, found from Africa to the Pacific Islands. Cleistanthus collinus is known for being toxic and frequently used for homicidal or suicidal purposes.-Synonymy:...
- PentabrachionPentabrachionPentabrachion is a plant genus from the family Phyllanthaceae....
- Subtribe Keayodendrinae 1 genus
- Keayodendron
- Subtribe Amanoinae 1 genus
- AmanoaAmanoaAmanoa is a neotropical genus from the family Phyllanthaceae. It consists of trees or shrubs, occurring in the tropical rainforests of South America and Africa.-Synonymy:This genus is equally known as:*Micropetalum Poit. ex Baill....
- Amanoa
- Subtribe Securineginae 2 genera
- Tribe PhyllantheaePhyllantheaePhyllantheae is a tribe of the family Phyllanthaceae. It comprises 6 subtribes and 18 genera.-External links:*...
5 genera-
- PhyllanthusPhyllanthusPhyllanthus is the largest genus in the flowering plant family Phyllanthaceae. Estimates of the number 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...
- Flueggea
- LingelsheimiaLingelsheimiaLingelsheimia is a plant genus of the family Putranjivaceae.-Synonymy:The genus is equally known as:*Aerisilvaea Radcl.-Sm.*Danguyodrypetes LeandriSpecies include:*Lingelsheimia anitrata*Lingelsheimia longipedicellata...
- MargaritariaMargaritariaMargaritaria is a plant genus of the Phyllanthaceae family. With 14 species, this is the smallest pantropical genus of the Phyllanthaceae and, formerly, of the Euphorbiaceae.-Synonymy:This genus is known under several other names:...
- Plagiocladus
- Phyllanthus
-
- Tribe WielandieaeWielandieaeWielandieae is a tribe of the family Phyllanthaceae. It comprises 10 genera....
2 subtribes- Subtribe Astrocasiinae 3 genera
- AstrocasiaAstrocasiaAstrocasia is a plant genus of the family Phyllanthaceae and the sole genus comprised in the subtribe Astrocasiinae....
- Chascotheca
- HeywoodiaHeywoodiaHeywoodia lucens is a plant species of the family Phyllanthaceae. Its genus, Heywoodia, is monotypic. The species is native to tropical East Africa....
- Astrocasia
- Subtribe Wielandiinae 3 genera
- WielandiaWielandiaWielandia is a genus of flowering plant, of the family Phyllanthaceae. Members of this genus are shrubs found in the Seychelles Islands....
- Chorisandrachne
- DicoeliaDicoeliaDicoelia is a plant genus of the family Euphorbiaceae and the only member of its tribe . It comprises 3 species, found in West Malesia. They are shrubs or small trees....
- Wielandia
- Subtribe Astrocasiinae 3 genera
- Tribe PoranthereaePoranthereaePoranthereae is a tribe in the flowering 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...
8 genera-
- PorantheraPorantheraPoranthera is a genus of flowering plant belonging to the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. Its closest relative is Pseudophyllanthus....
- ActephilaActephilaActephila is a genus of the family Phyllanthaceae. It is one of 8 genera in the tribe Poranthereae, and is most closely related to Leptopus. It comprises 31 species, found in Australasia. The type species is Actephila javanica. The name of the genus is derived from two Greek words, akte, "the...
- AndrachneAndrachneAndrachne is a genus of flowering plants in the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. Andrachne comprises 22 species. They are monoecious herbs or subshrubs, native to semideserts and desert margins of the Americas, southern Europe, North Africa, and South...
- Leptopus
- MeineckiaMeineckiaMeineckia is a genus of flowering plants in the family Phyllanthaceae. It is one of eight genera in the tribe Poranthereae. It comprises about 30 species. They are native to the Americas, South Asia, Africa, and Madagascar. The genus is particularly well represented in Madagascar. Species of...
- OreoporantheraOreoporantheraOreoporanthera is a genus of flowering plant belonging to the family Phyllanthaceae.It contains at least two species, Oreoporanthera alpina and Oreoporanthera petalifera....
- ZimmermanniaZimmermanniaZimmermannia is a genus of plant under the family Phyllanthaceae. The seven species appear similar, but careful examination of the polen of each has shown that they are distinct species....
- ZimmermanniopsisZimmermanniopsisZimmermanniopsis is a genus of plant under the family Phyllanthaceae. The genus resembles Zimmermannia, hence its name....
- Poranthera
-
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
Incertae sedis
, is a term used to define a taxonomic group where its broader relationships are unknown or undefined. Uncertainty at specific taxonomic levels is attributed by , , and similar terms.-Examples:*The fossil plant Paradinandra suecica could not be assigned to any...
), 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
Bootstrapping (statistics)
In statistics, bootstrapping is a computer-based method for assigning measures of accuracy to sample estimates . This technique allows estimation of the sample distribution of almost any statistic using only very simple methods...
. All branches shown below have maximum parsimony
Maximum parsimony
Parsimony is a non-parametric statistical method commonly used in computational phylogenetics for estimating phylogenies. Under parsimony, the preferred phylogenetic tree is the tree that requires the least evolutionary change to explain some observed data....
bootstrap support of at least 70%.
External links
- Phyllanthaceae NCBI Taxonomy browser
- Wurdack and Davis on Malpighiales PMID 21628300
- Molecular Phylogenetics of Phyllanthaceae
- Pruesapan on tribe Phyllantheae
- Phyllanthaceae At: Angiosperm Phylogeny Website At: Missouri Botanical Garden Website
- Tokuoka on Euphorbiaceae PMID 17530165
- Supporting Online Material for Davis on Rafflesiaceae
- Key to Euphorbiaceae sensu lato At: Index of /herbarium/keys