Phyllanthus

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Plagiotropic shoots of Phyllanthus pulcher
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Fruit of Phyllanthus acidus
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Male and female flowers of Phyllanthus acidus
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Flattened stems and flowers of Phyllanthus angustifolius
File:Phyllanthus urinaria leaves.jpg
Leaves of Phyllanthus urinaria

Phyllanthus is the largest genus in the plant family Phyllanthaceae. Estimates of the number of species in this genus vary widely, from 750[1] to 1200.[2] 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.

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.[3] Phyllanthus is distributed in all tropical and subtropical regions on Earth.

Phyllanthus was first described by Carl Linnaeus in 1753,[4][5] but the type was not designated.[4]

Species

Script error: No such module "Labelled list hatnote". 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 2006 revision of the family Phyllanthaceae has subsumed all four of these genera into Phyllanthus.[6] This enlarged version of Phyllanthus might eventually be divided into smaller genera,[7][8] including 32 Chinese (and northern Indochinese) species.[9] A complete overhaul of the genus, including a new classification is currently underway, following a recent indepth molecular treatment of major groups included.[10]

Selected species

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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]

References

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  1. ↑ David J. Mabberley. 2008. Mabberley's Plant-Book. third edition (2008). Cambridge University Press.Script error: No such module "Unsubst".
  2. ↑ Script error: No such module "Citation/CS1".
  3. ↑ Script error: No such module "Citation/CS1".
  4. ↑ a b Template:APNI2
  5. ↑ Linnaeus, C. (1753) Species Plantarum 2: 981.
  6. ↑ Script error: No such module "Citation/CS1".
  7. ↑ Script error: No such module "Citation/CS1".
  8. ↑ 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
  9. ↑ Script error: No such module "citation/CS1".
  10. ↑ Script error: No such module "Citation/CS1".
  11. ↑ "Phyllanthus phialanthoides (Phyllanthaceae), a new species from northeastern Cuba" (JSTOR)
  12. ↑ a b 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. ↑ Script error: No such module "Citation/CS1".
  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. ↑ Script error: No such module "Citation/CS1".
  23. ↑ Script error: No such module "Citation/CS1".
  24. ↑ Script error: No such module "Citation/CS1".
  25. ↑ Script error: No such module "Citation/CS1".
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