''Phyllanthus'' is the largest

genus 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 nom ...

in the plant 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. Davis. 2009. "Malpighiales phylogenetics: Gaining ground on one of the m ...

. Estimates of the number of

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

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 Plant life-form schemes constitute a way of classifying plants alternatively to the ordinary species-genus-family scientific classification. In colloquial speech, plants may be classified as trees, shrubs, herbs (forbs and graminoids), etc. The sc ...

including

annual Annual may refer to: *Annual publication, periodical publications appearing regularly once per year ** Yearbook ** Literary annual *Annual plant *Annual report *Annual giving *Annual, Morocco, a settlement in northeastern Morocco *Annuals (band), ...

and

perennial A perennial plant or simply perennial is a plant that lives more than two years. The term ('' per-'' + '' -ennial'', "through the years") is often used to differentiate a plant from shorter-lived annuals and biennials. The term is also wid ...

herbs, shrubs, climbers, floating aquatics, and pachycaulous

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 ...

. Some have flattened leaflike stems called cladodes. It has a wide variety of floral morphologies and

chromosome number Ploidy () is the number of complete sets of chromosomes in a cell, and hence the number of possible alleles for autosomal and pseudoautosomal genes. Sets of chromosomes refer to the number of maternal and paternal chromosome copies, respectivel ...

s and has one of the widest range of pollen types of any

seed plant 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 ...

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 leaves, usually in the autumn; to the shedding of petals, ...

, 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 * Axess (disambiguation) *Axxess (disambiguation) Axxess may refer to ...

are reduced to scales called " cataphylls", while leaves on the other axes develop normally. ''Phyllanthus'' is distributed in all

tropical 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 ...

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

Carl Linnaeus Carl Linnaeus (; 23 May 1707 – 10 January 1778), also known after his Nobility#Ennoblement, ennoblement in 1761 as Carl von Linné#Blunt, Blunt (2004), p. 171. (), was a Swedish botanist, zoologist, taxonomist, and physician who formalise ...

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

Species

The

circumscription Circumscription may refer to: *Circumscribed circle * Circumscription (logic) *Circumscription (taxonomy) *Circumscription theory, a theory about the origins of the political state in the history of human evolution proposed by the American anthrop ...

of this genus has been a cause of much confusion and disagreement. Molecular phylogenetic studies have shown that ''Phyllanthus'' is paraphyletic over ''Reverchonia'', ''

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 ...

'', '' Sauropus'', and '' Breynia.'' A recent 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, also known as the Indochinese Peninsula or Indochina, 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 the west an ...

) 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 ''Phyllanthus emblica'', also known as emblic, emblic myrobalan, myrobalan, Indian gooseberry, Malacca tree, or amla, from the Sanskrit आमलकी (āmalakī), is a deciduous tree of the family Phyllanthaceae. Its native range is tropical a ...

'' L. –

Indian gooseberry ''Phyllanthus emblica'', also known as emblic, emblic myrobalan, myrobalan, Indian gooseberry, Malacca tree, or amla, from the Sanskrit आमलकी (āmalakī), is a deciduous tree of the family Phyllanthaceae. Its native range is tropical a ...

, also known as amla or

amalaki ''Phyllanthus emblica'', also known as emblic, emblic myrobalan, myrobalan, Indian gooseberry, Malacca tree, or amla, from the Sanskrit आमलकी (āmalakī), is a deciduous tree of the family Phyllanthaceae. Its native range is tropical a ...

. * '' 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'' Müll.Arg. – one of the four succulent species of this genus * '' Phyllanthus montanus'' (Sw.) Sw. * '' Phyllanthus myrtifolius'' (Wight.) Muell.Arg. * '' Phyllanthus muellerianus'' (Kuntze) Exell * ''

Phyllanthus niruri ''Phyllanthus niruri'' is a widespread tropical plant commonly found in coastal areas, known by the common names gale of the wind, stonebreaker, shatter stone , seed-under-leaf, quebra pedra and chance pierre. It is in the genus ''Phyllanthus'' ...

'' 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 preserved ...

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 petrol ...

samples of the Middle Miocene fresh water deposits in Nowy Sacz Basin,

West Carpathians The Western Carpathians are a mountain range and geomorphological province that forms the western part of the Carpathian Mountains. The mountain belt stretches from the Low Beskids range of the Eastern Carpathians along the border of Poland wit ...

Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populou ...

. The seeds are similar to seeds of the fossil species †''Phyllanthus triquetra'' and †''Phyllanthus compassica'' from the Oligocene 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 in the geologic time scale that extends from 5.333 million to 2.58Poland Poland, officially the Republic of Poland, is a country in Central Europe. It is divided into 16 administrative provinces called voivodeships, covering an area of . Poland has a population of over 38 million and is the fifth-most populou ...

Pollination biology

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

'' 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 precipitate proteins and various other organic compounds including amino acids and alkaloids. The term ''tannin'' (from Anglo-Norman ''tanner'' ...

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

traditional medicine Traditional medicine (also known as indigenous medicine or folk medicine) comprises medical aspects of traditional knowledge that developed over generations within the folk beliefs of various societies, including indigenous peoples, before the ...

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 role ...

s and other phytochemicals.

References

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