# Phyllanthus

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"Reidia" redirects here. Not to be confused with [Reedia](/source/Reedia).

"Conami" redirects here. Not to be confused with [Konami](/source/Konami) or [CONAMI](/source/CONAMI).

Not to be confused with [Philanthus](/source/Philanthus).

***Phyllanthus*** is the largest [genus](/source/Genus) in the plant family [Phyllanthaceae](/source/Phyllanthaceae). Estimates of the number of [species](/source/Species) in this genus vary widely, from 750[1] to 1200.[2] *Phyllanthus* has a remarkable diversity of [growth forms](/source/Habit_(biology)) including [annual](/source/Annual_plant) and [perennial](/source/Perennial_plant) [herbs](/source/Herbaceous_plant), [shrubs](/source/Shrub), [climbers](/source/Vine), [floating aquatics](/source/Aquatic_plant), and [pachycaulous](/source/Pachycaul) [succulents](/source/Succulent_plant). Some have flattened leaflike stems called [cladodes](https://en.wiktionary.org/wiki/cladode). It has a wide variety of [floral morphologies](/source/Flower#Morphology) and [chromosome numbers](/source/Chromosome_number) and has one of the widest range of [pollen](/source/Pollen) types of any [seed plant](/source/Spermatophyte) genus.

Despite their variety, almost all *Phyllanthus* species express a specific type of growth called "phyllanthoid branching" in which the vertical stems bear [deciduous](/source/Deciduous), [floriferous](/source/Floriferous) (flower-bearing), [plagiotropic](/source/Glossary_of_botanical_terms#P) (horizontal or oblique) stems. The leaves on the main (vertical) [axes](/source/Axis_of_symmetry) are reduced to scales called "[cataphylls](/source/Cataphyll)", while leaves on the other axes develop normally.[3] *Phyllanthus* is distributed in all [tropical](/source/Tropical) and [subtropical](/source/Subtropical) regions on Earth.

*Phyllanthus* was first described by [Carl Linnaeus](/source/Carl_Linnaeus) in 1753;[4][5] the [type species](/source/Type_species) has subsequently been designated as *[Phyllanthus niruri](/source/Phyllanthus_niruri)*.[6]

## Species

Main article: [List of Phyllanthus species](/source/List_of_Phyllanthus_species)

The [circumscription](/source/Circumscription_(taxonomy)) of this genus has been a cause of much confusion and disagreement. [Molecular phylogenetic](/source/Molecular_phylogenetic) studies have shown that *Phyllanthus* is [paraphyletic](/source/Paraphyletic) over *Reverchonia*, *[Glochidion](/source/Glochidion)*, *[Sauropus](/source/Sauropus)*, and *[Breynia](/source/Breynia).* A 2006 revision of the family Phyllanthaceae has subsumed all four of these genera into *Phyllanthus*.[7] This enlarged version of *Phyllanthus* might eventually be divided into smaller genera,[8][9] including 32 Chinese (and northern [Indochinese](/source/Indochinese)) species.[10] A complete overhaul of the genus, including a new classification is currently underway, following a recent indepth molecular treatment of major groups included.[11]

### Selected species

- *[Phyllanthus abnormis](/source/Phyllanthus_abnormis)* Baill. – Drummond's leafflower
- *[Phyllanthus acidus](/source/Phyllanthus_acidus)* (L.) Skeels – Otaheite gooseberry
- *[Phyllanthus acuminatus](/source/Phyllanthus_acuminatus)* Vahl – Jamaican gooseberry tree
- *[Phyllanthus amarus](/source/Phyllanthus_amarus)* Schumacher
- *[Phyllanthus anamalayanus](/source/Phyllanthus_anamalayanus)* (Gamble) G.L.Webster
- *[Phyllanthus angustifolius](/source/Phyllanthus_angustifolius)* (Sw.) Sw.
- *[Phyllanthus arbuscula](/source/Phyllanthus_arbuscula)* (Sw.) J.F.Gmel.
- *[Phyllanthus axillaris](/source/Phyllanthus_axillaris)* (Sw.) Müll.Arg.
- *[Phyllanthus brasiliensis](/source/Phyllanthus_brasiliensis)* (Aubl.) Poir.
- *[Phyllanthus caesiifolius](/source/Phyllanthus_caesiifolius)* Petra Hoffm. & Cheek
- *[Phyllanthus caroliniensis](/source/Phyllanthus_caroliniensis)* Walt. – native to the Americas
- *[Phyllanthus cauliflorus](/source/Phyllanthus_cauliflorus)* (Sw.) Griseb.
- *[Phyllanthus cladanthus](/source/Phyllanthus_cladanthus)* Müll.Arg.
- *[Phyllanthus cochinchinensis](/source/Phyllanthus_cochinchinensis)* (Lour.) Spreng.
- *[Phyllanthus coluteoides](/source/Phyllanthus_coluteoides)* Baill. ex Müll.Arg.
- *[Phyllanthus cuneifolius](/source/Phyllanthus_cuneifolius)* (Britt.) Croizat
- *[Phyllanthus debilis](/source/Phyllanthus_debilis)* Klein ex Willd.
- *[Phyllanthus distichus](/source/Phyllanthus_distichus)* Hook. & Arn.
- *[Phyllanthus emblica](/source/Phyllanthus_emblica)* L. – [Indian gooseberry](/source/Indian_gooseberry), also known as [amla](/source/Indian_gooseberry) or [amalaki](/source/Amalaki).
- *[Phyllanthus engleri](/source/Phyllanthus_engleri)* Pax
- *[Phyllanthus epiphyllanthus](/source/Phyllanthus_epiphyllanthus)* L.
- *[Phyllanthus ericoides](/source/Phyllanthus_ericoides)* Torr.
- *[Phyllanthus eximius](/source/Phyllanthus_eximius)* G.L.Webster & Proctor
- *[Phyllanthus fadyenii](/source/Phyllanthus_fadyenii)* Urb.
- *[Phyllanthus fluitans](/source/Phyllanthus_fluitans)* Benth. ex Müll.Arg. – red root floater, sometimes sold in aquarium shops
- *[Phyllanthus fraternus](/source/Phyllanthus_fraternus)* G.L.Webster
- *[Phyllanthus gentryi](/source/Phyllanthus_gentryi)* Webster
- *[Phyllanthus grandifolius](/source/Phyllanthus_grandifolius)* L.
- *[Phyllanthus gunnii](/source/Phyllanthus_gunnii)* Hook.f.
- *[Phyllanthus hakgalensis](/source/Phyllanthus_hakgalensis)*
- *[Phyllanthus hirtellus](/source/Phyllanthus_hirtellus)* F.Muell. ex Mull.Arg.
- *[Phyllanthus juglandifolius](/source/Phyllanthus_juglandifolius)* Willd.
- *[Phyllanthus lacunarius](/source/Phyllanthus_lacunarius)* F.Muell.
- *[Phyllanthus latifolius](/source/Phyllanthus_latifolius)* (L.) Sw.
- *[Phyllanthus liebmannianus](/source/Phyllanthus_liebmannianus)* Muell.Arg.
- *[Phyllanthus maderaspatensis](/source/Phyllanthus_maderaspatensis)* L.
- *[Phyllanthus microcladus](/source/Phyllanthus_microcladus)* Muell.Arg.
- *[Phyllanthus millei](/source/Phyllanthus_millei)* Standl.
- *[Phyllanthus mirabilis](/source/Phyllanthus_mirabilis)* Müll.Arg. – one of the four [succulent](/source/Succulent) species of this genus
- *[Phyllanthus montanus](/source/Phyllanthus_montanus)* (Sw.) Sw.
- *[Phyllanthus myrtifolius](/source/Phyllanthus_myrtifolius)* (Wight.) Muell.Arg.
- *[Phyllanthus muellerianus](/source/Phyllanthus_muellerianus)* (Kuntze) Exell
- *[Phyllanthus niruri](/source/Phyllanthus_niruri)* L. – Chanca piedra (Also includes *P. amarus* and *P. debilis*)
- *[Phyllanthus nyale](/source/Phyllanthus_nyale)* Petra Hoffm. & Cheek
- *[Phyllanthus parvifolius](/source/Phyllanthus_parvifolius)* Buch.-Ham. ex D.Don
- *[Phyllanthus pavonianus](/source/Phyllanthus_pavonianus)* Baill.
- *[Phyllanthus pentaphyllus](/source/Phyllanthus_pentaphyllus)* C.Wright ex Griseb.
- *[Phyllanthus phialanthoides](/source/Phyllanthus_phialanthoides)* Falcón & J.L.Gómez[12]
- *[Phyllanthus polygonoides](/source/Phyllanthus_polygonoides)* Nutt. ex Spreng. – Smartweed leafflower
- *[Phyllanthus polyspermus](/source/Phyllanthus_polyspermus)* [Shumach.](/source/Heinrich_Christian_Friedrich_Schumacher) & [Thonn.](/source/Peter_Thonning) – often misidentified as *P. reticulatus*[13]
- *[Phyllanthus profusus](/source/Phyllanthus_profusus)* N.E.Br.
- *[Phyllanthus pulcher](/source/Phyllanthus_pulcher)* Wallich ex Muell.Arg.
- *[Phyllanthus reticulatus](/source/Phyllanthus_reticulatus)* Poir. – Asian sp. similar in appearance to *P. polyspermus*[13]
- *[Phyllanthus revaughanii](/source/Phyllanthus_revaughanii)* Coode
- *[Phyllanthus rufuschaneyi](/source/Phyllanthus_rufuschaneyi)* Welzen, R.W.Bouman & Ent
- *[Phyllanthus saffordii](/source/Phyllanthus_saffordii)* Merr.
- *[Phyllanthus salviifolius](/source/Phyllanthus_salviifolius)* Kunth
- *[Phyllanthus sepialis](/source/Phyllanthus_sepialis)* Müll.Arg.[14]
- *[Phyllanthus societatis](/source/Phyllanthus_societatis)* Müll.Arg.
- *[Phyllanthus sponiifolius](/source/Phyllanthus_sponiifolius)* Müll.Arg.
- *[Phyllanthus stipulatus](/source/Phyllanthus_stipulatus)* (Raf.) G.L. Webster
- *[Phyllanthus taxodiifolius](/source/Phyllanthus_taxodiifolius)* Beille
- *[Phyllanthus tenellus](/source/Phyllanthus_tenellus)* Roxb.
- *[Phyllanthus urinaria](/source/Phyllanthus_urinaria)* L. – chamberbitter
- *[Phyllanthus virgatus](/source/Phyllanthus_virgatus)* G.Forst.
- *[Phyllanthus warnockii](/source/Phyllanthus_warnockii)* G.L.Webster
- *[Phyllanthus watsonii](/source/Phyllanthus_watsonii)* Airy Shaw
- *[Phyllanthus welwitschianus](/source/Phyllanthus_welwitschianus)* Müll.Arg.

## Fossil record

Two [fossil](/source/Fossil) seeds of a *Phyllanthus* species have been extracted from [borehole](/source/Borehole) samples of the [Middle Miocene](/source/Middle_Miocene) fresh water deposits in [Nowy Sacz](/source/Nowy_Sacz) Basin, [West Carpathians](/source/West_Carpathians), [Poland](/source/Poland). The seeds are similar to seeds of the fossil species †*[Phyllanthus triquetra](/source/Phyllanthus_triquetra)* and †*[Phyllanthus compassica](/source/Phyllanthus_compassica)* from the [Oligocene](/source/Oligocene) and [Miocene](/source/Miocene) of [West Siberia](/source/West_Siberia). *Phyllanthus* fossils are known from several [Miocene](/source/Miocene) and [Pliocene](/source/Pliocene) sites in [Poland](/source/Poland).[15]

## Pollination biology

*Phyllanthus* are of note in the fields of [pollination biology](/source/Anthecology) and [coevolution](/source/Coevolution) because some but not all species in the genus have a specialized [mutualism](/source/Mutualism_(biology)) with moths in the genus *[Epicephala](/source/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.[16][17] Other species of *Epicephala* are pollinators of certain species of plants in the genera *[Glochidion](/source/Glochidion)*[18][19] and *[Breynia](/source/Breynia)*,[20][21] both of which are phylogenetically nested within *Phyllanthus*.[22]

## Research and traditional medicine

Particularly for its content of [tannins](/source/Tannin), *P. emblica* fruit has a history of use in [traditional medicine](/source/Traditional_medicine) and is under study for its potential biological properties.[23] Leaves, roots, stem, bark and berries of this genus contain [lignans](/source/Lignan) and other [phytochemicals](/source/Phytochemical).[24][25][26]

## References

1. David J. Mabberley. 2008. *Mabberley's Plant-Book.* third edition (2008). Cambridge University Press.[page needed]

1. Kathriarachchi H, Hoffmann P, Samuel R, Wurdack KJ, Chase MW (July 2005). "Molecular phylogenetics of Phyllanthaceae inferred from five genes (plastid atpB, matK, 3'ndhF, rbcL, and nuclear PHYC)". *Molecular Phylogenetics and Evolution*. **36** (1): 112–34. [Bibcode:2005MolPE..36..112K](https://ui.adsabs.harvard.edu/abs/2005MolPE..36..112K). [doi:10.1016/j.ympev.2004.12.002](https://doi.org/10.1016/j.ympev.2004.12.002). [PMID 15904861](https://pubmed.ncbi.nlm.nih.gov/15904861)

1. Webster, Grady L. (1994). ["Classification of the Euphorbiaceae"](https://www.biodiversitylibrary.org/part/30297). *Annals of the Missouri Botanical Garden*. **81** (1): 3–32. [doi:10.2307/2399908](https://doi.org/10.2307/2399908). [JSTOR 2399908](https://www.jstor.org/stable/2399908)

1. Linnaeus, C. (1753) Species Plantarum 2: 981.

1. ["Phyllanthus"](https://www.ipni.org/n/327609-2). *International Plant Names Index*. 1913-06-07. Retrieved 2026-01-08.

1. Hoffmann, Petra; Kathriarachchi, Hashendra S.; Wurdack, Kenneth J. (2006). "A Phylogenetic Classification of Phyllanthaceae". *Kew Bulletin*. **61** (1): 37–53.

1. Kathriarachchi, Hashendra S.; Samuel, Rosabelle; Hoffmann, Petra; Mlinarec, Jelena; Wurdack, Kenneth J.; Ralimanana, Hélène; Stuessy, Tod F.; Chase, Mark W. (2006). "Phylogenetics of tribe Phyllantheae (Phyllanthaceae) based on nrITS and plastid matK DNA sequence data". *American Journal of Botany*. **93** (4): 637–655. [doi:10.3732/ajb.93.4.637](https://doi.org/10.3732/ajb.93.4.637). [PMID 21646224](https://pubmed.ncbi.nlm.nih.gov/21646224)

1. 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

1. ["Flora of China"](http://www.efloras.org/florataxon.aspx?flora_id=2&taxon_id=125179). eFlora. Retrieved 18 April 2017.

1. Bouman, Roderick W.; Keßler, Paul J. A.; Telford, Ian R. H.; Bruhl, Jeremy J.; Strijk, Joeri S.; Saunders, Richard M. K.; Welzen, Peter C. van (2021). "Molecular phylogenetics of Phyllanthus sensu lato (Phyllanthaceae): Towards coherent monophyletic taxa". *Taxon*. **70** (1): 72–98. [doi:10.1002/tax.12424](https://doi.org/10.1002/tax.12424). [hdl:1887/3248720](https://hdl.handle.net/1887/3248720). [ISSN 1996-8175](https://www.worldcat.org/issn/1996-8175)

1. ["*Phyllanthus phialanthoides* (Phyllanthaceae), a new species from northeastern Cuba"](https://www.jstor.org/stable/26296180?seq=1#page_scan_tab_contents) ([JSTOR](/source/JSTOR))

1. 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.

1. Bussmann, R. W.; Gilbreath, GG; Solio, J; Lutura, M; Lutuluo, R; Kunguru, K; Wood, N; Mathenge, SG (2006). "Plant use of the Maasai of Sekenani Valley, Maasai Mara, Kenya". *J Ethnobiol Ethnomed*. **2**: 22. [doi:10.1186/1746-4269-2-22](https://doi.org/10.1186/1746-4269-2-22). [PMC 1475560](https://www.ncbi.nlm.nih.gov/pmc/articles/PMC1475560). [PMID 16674830](https://pubmed.ncbi.nlm.nih.gov/16674830)

1. Macroscopic plant remains from the freshwater Miocene of the Nowy Sącz Basin (West Carpathians, Poland) by Łańcucka-Środoniowa M, Acta Palaeobotanica 1979 20 (1): 3-117.

1. Kawakita, A.; Kato, M. 2004. "Evolution of obligate pollination mutualism in New Caledonian *Phyllanthus* (Euphorbiaceae)." *American Journal of Botany* **91**: 410–415.

1. Kawakita, A.; Kato, M. 2009. "Repeated independent evolution of obligate pollination mutualism in the Phyllantheae-*Epicephala* association." *Proceedings of the Royal Society B.* **276**: 417–426.

1. Hembry, D. H.; Okamoto, T.; Gillespie, R. G. (2012) Repeated colonization of remote islands by specialized mutualists. *Biology Letters*. **8**: 258–261.

1. Luo, S.-X.; Yao, G.; Wang, Z.; Zhang, D.; Hembry, D. H. (2017) "A novel, enigmatic basal leafflower moth lineage pollinating a derived leafflower host illustrates the dynamics of host shifts, partner replacement, and apparent co-adaptation in intimate mutualisms." *The American Naturalist*. **189**: 422–435

1. Kawakita, A.; Kato, M. 2004. Obligate pollination mutualism in *Breynia* (Phyllanthaceae): further documentation of pollination mutualism involving *Epicephala* moths (Gracillariidae). *American Journal of Botany.* 91: 1319–1325.

1. Zhang, J.; Wang, S.; Li, H.; Hu, B.; Yang, X.; Wang, Z. 2012. "Diffuse coevolution between two *Epicephala* species (Gracillariidae) and two *Breynia* species (Phyllanthaceae). *PLOS ONE.* **7**: e41657.

1. Kathriarachchi, H.; Samuel, R.; Hoffmann, P.; Mlinarec, J.; Wurdack, K. J.; Ralimanana, H.; Stuessy, T. F.; Chase, M. W. 2006. "Phylogenetics of tribe Phyllantheae (Phyllanthaceae: Euphorbiaceae sensu lato) based on nrITS and plastid matK DNA sequence data." *American Journal of Botany.* **93**: 637–655.

1. Yang, B & Liu, P (2014). "Composition and biological activities of hydrolyzable tannins of fruits of Phyllanthus emblica". *Journal of Agricultural and Food Chemistry*. **62** (3): 529–41. [doi:10.1021/jf404703k](https://doi.org/10.1021/jf404703k). [PMID 24369850](https://pubmed.ncbi.nlm.nih.gov/24369850)

1. Murugaiyah V, Chan KL (June 2007). "Determination of four lignans in *Phyllanthus niruri* L. by a simple high-performance liquid chromatography method with fluorescence detection". *Journal of Chromatography A*. **1154** (1–2): 198–204. [doi:10.1016/j.chroma.2007.03.079](https://doi.org/10.1016/j.chroma.2007.03.079). [PMID 17418855](https://pubmed.ncbi.nlm.nih.gov/17418855)

1. Srivastava V, Singh M, Malasoni R, etal (January 2008). "Separation and quantification of lignans in *Phyllanthus* species by a simple chiral densitometric method". *Journal of Separation Science*. **31** (1): 47–55. [doi:10.1002/jssc.200700282](https://doi.org/10.1002/jssc.200700282). [PMID 18064620](https://pubmed.ncbi.nlm.nih.gov/18064620)

1. Bagalkotkar G, Sagineedu SR, Saad MS, Stanslas J (December 2006). "Phytochemicals from *Phyllanthus niruri* Linn. and their pharmacological properties: a review". *The Journal of Pharmacy and Pharmacology*. **58** (12): 1559–70. [doi:10.1211/jpp.58.12.0001](https://doi.org/10.1211/jpp.58.12.0001). [PMID 17331318](https://pubmed.ncbi.nlm.nih.gov/17331318)

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