''Phyllanthus'' is the largest

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

in the plant family Phyllanthaceae. Estimates of the number of

species A species () is often defined as the largest group of organisms in which any two individuals of the appropriate sexes or mating types can produce fertile offspring, typically by sexual reproduction. It is the basic unit of Taxonomy (biology), ...

in this genus vary widely, from 750David J. Mabberley. 2008. ''Mabberley's Plant-Book.'' third edition (2008). Cambridge University Press. to 1200. ''Phyllanthus'' has a remarkable diversity of growth forms including annual and

perennial In horticulture, the term perennial ('' per-'' + '' -ennial'', "through the year") is used to differentiate a plant from shorter-lived annuals and biennials. It has thus been defined as a plant that lives more than 2 years. The term is also ...

herbs,

shrub A shrub or 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 by their multiple ...

s, climbers, floating aquatics, and pachycaulous succulents. Some have flattened leaflike stems called cladodes. It has a wide variety of floral morphologies and chromosome numbers and has one of the widest range of

pollen Pollen is a powdery substance produced by most types of flowers of seed plants for the purpose of sexual reproduction. It consists of pollen grains (highly reduced Gametophyte#Heterospory, microgametophytes), which produce male gametes (sperm ...

types of any seed plant genus. Despite their variety, almost all ''Phyllanthus'' species express a specific type of growth called "phyllanthoid branching" in which the vertical stems bear

deciduous In the fields of horticulture and botany, the term deciduous () means "falling off at maturity" and "tending to fall off", in reference to trees and shrubs that seasonally shed Leaf, leaves, usually in the autumn; to the shedding of petals, aft ...

, floriferous (flower-bearing), plagiotropic (horizontal or oblique) stems. The leaves on the main (vertical)

axes Axes, plural of ''axe'' and of ''axis'', may refer to * ''Axes'' (album), a 2005 rock album by the British band Electrelane * a possibly still empty plot (graphics) See also * Axis (disambiguation) An axis (: axes) may refer to: Mathematics ...

are reduced to scales called "

cataphyll In plant morphology, a cataphyll (sometimes also called a ''cataphyllum'' or cataphyll leafJackson, Benjamin, Daydon; A Glossary of Botanic Terms with their Derivation and Accent; Published by Gerald Duckworth & Co. London, 4th ed 1928) is a reduce ...

s", while leaves on the other axes develop normally. ''Phyllanthus'' is distributed in all

tropical The tropics are the regions of Earth surrounding the equator, where the sun may shine directly overhead. This contrasts with the temperate or polar regions of Earth, where the Sun can never be directly overhead. This is because of Earth's ax ...

and

subtropical The subtropical zones or subtropics are geographical zone, geographical and Köppen climate classification, climate zones immediately to the Northern Hemisphere, north and Southern Hemisphere, south of the tropics. Geographically part of the Ge ...

regions on Earth. ''Phyllanthus'' was first described by

Carl Linnaeus Carl Linnaeus (23 May 1707 – 10 January 1778), also known after ennoblement in 1761 as Carl von Linné,#Blunt, Blunt (2004), p. 171. was a Swedish biologist and physician who formalised binomial nomenclature, the modern system of naming o ...

in 1753,Linnaeus, C. (1753) Species Plantarum 2: 981. but the type was not designated.

Species

The circumscription of this genus has been a cause of much confusion and disagreement.

Molecular phylogenetic Molecular phylogenetics () is the branch of phylogeny that analyzes genetic, hereditary molecular differences, predominantly in DNA sequences, to gain information on an organism's evolutionary relationships. From these analyses, it is possible to ...

studies have shown that ''Phyllanthus'' is

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

over ''Reverchonia'', '' Glochidion'', '' Sauropus'', and '' Breynia.'' A 2006 revision of the family Phyllanthaceae has subsumed all four of these genera into ''Phyllanthus''. This enlarged version of ''Phyllanthus'' might eventually be divided into smaller genera,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 including 32 Chinese (and northern

Indochinese Mainland Southeast Asia (historically known as Indochina and the Indochinese Peninsula) is the continental portion of Southeast Asia. It lies east of the Indian subcontinent and south of Mainland China and is bordered by the Indian Ocean to th ...

) species. A complete overhaul of the genus, including a new classification is currently underway, following a recent indepth molecular treatment of major groups included.

Selected species

* '' Phyllanthus abnormis'' Baill. – Drummond's leafflower * '' Phyllanthus acidus'' (L.) Skeels – Otaheite gooseberry * '' Phyllanthus acuminatus'' Vahl – Jamaican gooseberry tree * '' Phyllanthus amarus'' Schumacher * '' Phyllanthus anamalayanus'' (Gamble) G.L.Webster * '' Phyllanthus angustifolius'' (Sw.) Sw. * '' Phyllanthus arbuscula'' (Sw.) J.F.Gmel. * '' Phyllanthus axillaris'' (Sw.) Müll.Arg. * '' Phyllanthus brasiliensis'' (Aubl.) Poir. * '' Phyllanthus caesiifolius'' Petra Hoffm. & Cheek * '' Phyllanthus caroliniensis'' Walt. – native to the Americas * '' Phyllanthus cauliflorus'' (Sw.) Griseb. * '' Phyllanthus cladanthus'' Müll.Arg. * '' Phyllanthus cochinchinensis'' (Lour.) Spreng. * '' Phyllanthus coluteoides'' Baill. ex Müll.Arg. * '' Phyllanthus cuneifolius'' (Britt.) Croizat * '' Phyllanthus debilis'' Klein ex Willd. * '' Phyllanthus distichus'' Hook. & Arn. * '' Phyllanthus emblica'' L. – Indian gooseberry, also known as amla or amalaki. * '' Phyllanthus engleri'' Pax * '' Phyllanthus epiphyllanthus'' L. * '' Phyllanthus ericoides'' Torr. * '' Phyllanthus eximius'' G.L.Webster & Proctor * '' Phyllanthus fadyenii'' Urb. * '' Phyllanthus fluitans'' Benth. ex Müll.Arg. – red root floater, sometimes sold in aquarium shops * '' Phyllanthus fraternus'' G.L.Webster * '' Phyllanthus gentryi'' Webster * '' Phyllanthus grandifolius'' L. * '' Phyllanthus gunnii'' Hook.f. * '' Phyllanthus hakgalensis'' * '' Phyllanthus hirtellus'' F.Muell. ex Mull.Arg. * '' Phyllanthus juglandifolius'' Willd. * '' Phyllanthus lacunarius'' F.Muell. * '' Phyllanthus latifolius'' (L.) Sw. * '' Phyllanthus liebmannianus'' Muell.Arg. * '' Phyllanthus maderaspatensis'' L. * '' Phyllanthus microcladus'' Muell.Arg. * '' Phyllanthus millei'' Standl. * ''

Phyllanthus mirabilis ''Phyllanthus mirabilis'' is a plant species of family (biology), family Phyllanthaceae and is native to Thailand, Laos and Myanmar. It is one of the only four ''Phyllanthus'' to be caudex, caudiciform and one of the only two caudiciform ''Phylla ...

'' Müll.Arg. – one of the four

succulent 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 ...

species of this genus * '' Phyllanthus montanus'' (Sw.) Sw. * '' Phyllanthus myrtifolius'' (Wight.) Muell.Arg. * ''

Phyllanthus muellerianus ''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 ...

'' (Kuntze) Exell * '' Phyllanthus niruri'' L. – Chanca piedra (Also includes ''P. amarus'' and ''P. debilis'') * '' Phyllanthus nyale'' Petra Hoffm. & Cheek * '' Phyllanthus parvifolius'' Buch.-Ham. ex D.Don * '' Phyllanthus pavonianus'' Baill. * '' Phyllanthus pentaphyllus'' C.Wright ex Griseb. * '' Phyllanthus phialanthoides'' Falcón & J.L.Gómez * '' Phyllanthus polygonoides'' Nutt. ex Spreng. – Smartweed leafflower * '' Phyllanthus polyspermus'' Shumach. & Thonn. – often misidentified as ''P. reticulatus'' * '' Phyllanthus profusus'' N.E.Br. * '' Phyllanthus pulcher'' Wallich ex Muell.Arg. * '' Phyllanthus reticulatus'' Poir. – Asian sp. similar in appearance to ''P. polyspermus''Luo, S.X., H.-J. Esser, D. Zhang, and S. S. Renner. 2011. Nuclear ITS sequences help disentangle ''Phyllanthus reticulatus'' (Phyllanthaceae), an Asian species not occurring in Africa, but introduced to Jamaica. Systematic Botany 36(1): 99-104. * '' Phyllanthus revaughanii'' Coode * '' Phyllanthus rufuschaneyi'' Welzen, R.W.Bouman & Ent * '' Phyllanthus saffordii'' Merr. * '' Phyllanthus salviifolius'' Kunth * '' Phyllanthus sepialis'' Müll.Arg. * '' Phyllanthus societatis'' Müll.Arg. * '' Phyllanthus sponiifolius'' Müll.Arg. * '' Phyllanthus stipulatus'' (Raf.) G.L. Webster * '' Phyllanthus taxodiifolius'' Beille * '' Phyllanthus tenellus'' Roxb. * '' Phyllanthus urinaria'' L. – chamberbitter * '' Phyllanthus virgatus'' G.Forst. * '' Phyllanthus warnockii'' G.L.Webster * '' Phyllanthus watsonii'' Airy Shaw * '' Phyllanthus welwitschianus'' Müll.Arg.

Fossil record

Two

fossil A fossil (from Classical Latin , ) is any preserved remains, impression, or trace of any once-living thing from a past geological age. Examples include bones, shells, exoskeletons, stone imprints of animals or microbes, objects preserve ...

seeds of a ''Phyllanthus'' species have been extracted from

borehole A borehole is a narrow shaft bored in the ground, either vertically or horizontally. A borehole may be constructed for many different purposes, including the extraction of water ( drilled water well and tube well), other liquids (such as petr ...

samples of the

Middle Miocene The Middle Miocene is a sub-epoch of the Miocene epoch (geology), epoch made up of two Stage (stratigraphy), stages: the Langhian and Serravallian stages. The Middle Miocene is preceded by the Early Miocene. The sub-epoch lasted from 15.97 ± 0. ...

fresh water deposits in Nowy Sacz Basin, West Carpathians,

Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...

. The seeds are similar to seeds of the fossil species †''Phyllanthus triquetra'' and †''Phyllanthus compassica'' from the

Oligocene The Oligocene ( ) is a geologic epoch (geology), epoch of the Paleogene Geologic time scale, Period that extends from about 33.9 million to 23 million years before the present ( to ). As with other older geologic periods, the rock beds that defin ...

and

Miocene The Miocene ( ) is the first epoch (geology), geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and mea ...

of West Siberia. ''Phyllanthus'' fossils are known from several

and

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Poland Poland, officially the Republic of Poland, is a country in Central Europe. It extends from the Baltic Sea in the north to the Sudetes and Carpathian Mountains in the south, bordered by Lithuania and Russia to the northeast, Belarus and Ukrai ...

Pollination biology

''Phyllanthus'' are of note in the fields of pollination biology and

coevolution In biology, coevolution occurs when two or more species reciprocally affect each other's evolution through the process of natural selection. The term sometimes is used for two traits in the same species affecting each other's evolution, as well a ...

because some but not all species in the genus have a specialized mutualism with moths in the genus '' Epicephala'' (leafflower moths), in which the moths actively pollinate the flowers. While ensuring that the tree may produce viable seeds, the moths also lay eggs in the flowers' ovaries where their larvae consume a subset of the developing seeds as nourishment. Other species of ''Epicephala'' are pollinators of certain species of plants in the genera '' Glochidion'' and '' Breynia'', both of which are phylogenetically nested within ''Phyllanthus''.

Research and traditional medicine

Particularly for its content of

tannin Tannins (or tannoids) are a class of astringent, polyphenolic biomolecules that bind to and Precipitation (chemistry), precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' is widel ...

s, ''P. emblica'' fruit has a history of use in

traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) refers to the knowledge, skills, and practices rooted in the cultural beliefs of various societies, especially Indigenous groups, used for maintaining health and treatin ...

and is under study for its potential biological properties. Leaves, roots, stem, bark and berries of this genus contain

lignan The lignans are a large group of low molecular weight polyphenols found in plants, particularly seeds, whole grains, and vegetables. The name derives from the Latin word for "wood". Lignans are precursors to phytoestrogens. They may play a rol ...

s and other

phytochemical Phytochemicals are naturally-occurring chemicals present in or extracted from plants. Some phytochemicals are nutrients for the plant, while others are metabolites produced to enhance plant survivability and reproduction. The fields of ext ...

References

{{Taxonbar, from=Q310927 Phyllanthaceae genera Taxa named by Carl Linnaeus Taxa described in 1753