Phyllanthus

Phyllanthus is the largest genus in the flowering plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750[2] to 1200.[3] Phyllanthus has a remarkable diversity of growth forms including annual and perennial herbs, shrubs, 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 types of any seed plant genus.

For the bird genus of the same name, see Capuchin babbler.

Not to be confused with Philanthus.

Phyllanthus
Phyllanthus mirabilis
Scientific classification
Kingdom:	Plantae
Clade:	Tracheophytes
Clade:	Angiosperms
Clade:	Eudicots
Clade:	Rosids
Order:	Malpighiales
Family:	Phyllanthaceae
Subfamily:	Phyllanthoideae
Tribe:	Phyllantheae
Genus:	Phyllanthus L.
Diversity
About 800 species
Synonyms[1]
List Agyneia L., nom. rej. Anisonema A.Juss. Aporosella Chodat Arachnodes Gagnep. Ardinghalia Comm. ex A.Juss. Asterandra Klotzsch Bradleia Banks ex Gaertn. Cathetus Lour. Ceramanthus Hassk. Chlorolepis Nutt. Chorisandra Wight, nom. illeg. Chorizonema Jean F.Brunel Cicca L. Clambus Miers Coccoglochidion K.Schum. Conami Aubl. Cycca Batsch Dendrophyllanthus S.Moore Diasperus L. ex Kuntze, nom. illeg. Dichelactina Hance Dichrophyllum Klotzsch & Garcke Dimorphocladium Britton Emblica Gaertn. Episteira Raf. Epistylium Sw. Eriococcus Hassk. Flueggeopsis K.Schum. Frankia Steud. Geminaria Raf. Genesiphyla Raf. Genesiphylla L'Hér. Glochidionopsis Blume Glochisandra Wight Gynoon A.Juss. Hemicicca Baill. Hemiglochidion (Müll.Arg.) K.Schum., nom. illeg. Hexadena Raf. Hexaspermum Domin Kirganelia Juss. Leichhardtia F.Muell. Lobocarpus Wight & Arn. Lomanthes Raf. Macraea Wight Maschalanthus Nutt., nom. illeg. Meborea Aubl. Menarda Comm. ex A.Juss. Moeroris Raf. Nellica Raf. Niruri Adans. Niruris Raf. Nymania K.Schum. Nymphanthus Lour. Orbicularia Baill. Oxalistylis Baill. Phyllanthodendron Hemsl. Pseudoglochidion Gamble Ramsdenia Britton Reidia Wight Reverchonia A.Gray Rhopium Schreb. Roigia Britton Scepasma Blume Staurothyrax Griff. Synexemia Raf. Tetraglochidion K.Schum. Tricarium Lour. Uranthera Pax & K.Hoffm. Urinaria Medik. Williamia Baill. Xylophylla L. Zarcoa Llanos

Plagiotropic shoots of Phyllanthus pulcher

Fruit of Phyllanthus acidus

Male and female flowers of Phyllanthus acidus

Flattened stems and flowers of Phyllanthus angustifolius

Despite their variety, almost all Phyllanthus species express a specific type of growth called "phyllanthoid branching" in which the vertical stems bear deciduous, floriferous (flower-bearing), plagiotropic (horizontal or oblique) stems. The leaves on the main (vertical) axes are reduced to scales called "cataphylls", while leaves on the other axes develop normally.[4] Phyllanthus is distributed in all tropical and subtropical regions on Earth.

Phyllanthus was first described by Carl Linnaeus in 1753,[5][6] but the type was not designated.[5]

Species

Main article: List of Phyllanthus species

The circumscription of this genus has been a cause of much confusion and disagreement. Molecular phylogenetic studies have shown that Phyllanthus is paraphyletic over Reverchonia, Glochidion, Sauropus, and Breynia. A recent 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) species.[10]

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 - 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. - the only succulent species of this genus
• Phyllanthus montanus (Sw.) Sw.
• Phyllanthus myrtifolius - Wight. Muell. Arg.
• Phyllanthus muellerianus (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[11]
• Phyllanthus polygonoides Nutt. ex Spreng. - Smartweed leafflower
• Phyllanthus polyspermus Shumach. & Thonn. - often misidentified as P. reticulatus[12]
• Phyllanthus profusus N.E.Br.
• Phyllanthus pulcher Wallich ex Muell.-Arg.
• Phyllanthus reticulatus Poir. - Asian sp. similar in appearance to P. polyspermus[12]
• Phyllanthus revaughanii Coode
• Phyllanthus saffordii Merr.
• Phyllanthus salviifolius Kunth
• Phyllanthus sepialis Müll.Arg.[13]
• 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 A. Shaw
• Phyllanthus welwitschianus Müll.Arg.

Fossil record

Two fossil seeds of a Phyllanthus species have been extracted from borehole samples of the Middle Miocene fresh water deposits in Nowy Sacz Basin, West Carpathians, Poland. The seeds are similar to seeds of the fossil species †Phyllanthus triquetra and †Phyllanthus compassica from the Oligocene and Miocene of West Siberia. Phyllanthus fossils are known from several Miocene and Pliocene sites in Poland.[14]

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.[15][16] Other species of Epicephala are pollinators of certain species of plants in the genera Glochidion[17][18] and Breynia,[19][20] both of which are phylogenetically nested within Phyllanthus.[21]

Research and traditional medicine

Particularly for its content of tannins, P. emblica fruit has a history of use in traditional medicine and is under study for its potential biological properties.[22] Leaves, roots, stem, bark and berries of this genus contain lignans and other phytochemicals.[23][24][25]

Phyllanthus species have been assessed for use in people with chronic hepatitis B virus infection, but the poor quality of clinical trials has prevented any conclusion about efficacy, as of 2013.[26]

References

1. "World Checklist of Selected Plant Families".
2. David J. Mabberley. 2008. Mabberley's Plant-Book. third edition (2008). Cambridge University Press.
3. 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. doi:10.1016/j.ympev.2004.12.002. PMID 15904861.
4. Webster, Grady L. (1994). "Classification of the Euphorbiaceae". Annals of the Missouri Botanical Garden. 81 (1): 3–32. doi:10.2307/2399908. JSTOR 2399908.
5. "Phyllanthus". Australian Plant Name Index (APNI), IBIS database. Centre for Plant Biodiversity Research, Australian Government.
6. Linnaeus, C. (1753) Species Plantarum 2: 981.
7. Hoffmann, Petra; Kathriarachchi, Hashendra S.; Wurdack, Kenneth J. (2006). "A Phylogenetic Classification of Phyllanthaceae". Kew Bulletin. 61 (1): 37–53.
8. 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. PMID 21646224.
9. 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
10. "Flora of China". eFlora. Retrieved 18 April 2017.
11. "Phyllanthus phialanthoides (Phyllanthaceae), a new species from northeastern Cuba" (JSTOR)
12. 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.
13. 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. PMC 1475560. PMID 16674830.
14. 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.
15. Kawakita, A.; Kato, M. 2004. "Evolution of obligate pollination mutualism in New Caledonian Phyllanthus (Euphorbiaceae)." American Journal of Botany 91: 410–415.
16. 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.
17. Hembry, D. H.; Okamoto, T.; Gillespie, R. G. (2012) Repeated colonization of remote islands by specialized mutualists. Biology Letters. 8: 258–261.
18. 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
19. 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.
20. 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.
21. 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.
22. 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. PMID 24369850.
23. 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. PMID 17418855.
24. Srivastava V, Singh M, Malasoni R, et al. (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. PMID 18064620.
25. 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. PMID 17331318.
26. Xia, Y; Luo, H; Liu, J. P.; Gluud, C (2013). "Phyllanthus species versus antiviral drugs for chronic hepatitis B virus infection". The Cochrane Database of Systematic Reviews (4): CD009004. doi:10.1002/14651858.CD009004.pub2. PMID 23633363.