Contents 1 Evolution of androdioecy 1.1 Androdioecy with dioecious ancestry 1.2 Androdioecy with hermaphroditic ancestry 2 Androdioecious species 2.1 Anthozoa (Corals) 2.2 Nematoda (Roundworms) 2.3 Nemertea (Ribbon worms) 2.4 Arthropoda 2.5 Annelida (Ringed worms) 2.6 Chordata 2.7 Plants 3 External links 4 See also 5 References

Evolution of androdioecy[edit] The fitness requirements for androdioecy to arise and sustain itself are theoretically so improbable that it was long considered that such systems do not exist.[3][4] Particularly, males and hermaphrodites have to have the same fitness, in other words the same number of offspring, in order to be maintained. However, males only have offspring by fertilizing eggs or ovules of hermaphrodites, while hermaphrodites have offspring both through fertilizing eggs or ovules of other hermaphrodites and their own ovules. This means that all else being equal, males have to fertilize twice as many eggs or ovules as hermaphrodites to make up for the lack of female reproduction.[5][6] Androdioecy can evolve either from dioecious ancestors through the invasion of hermaphrodites or from hermaphroditic ancestors through the invasion of males. The ancestral state is important because conditions under which androdioecy can evolve differ significantly. Androdioecy with dioecious ancestry[edit] In roundworms, clam shrimp, tadpole shrimp and cancrid shrimps, androdioecy has evolved from dioecy. In these systems, hermaphrodites can only fertilize their own eggs (self-fertilize) and do not mate with other hermaphrodites. Males are the only means of outcrossing. Hermaphrodites may be beneficial in colonizing new habitats, because a single hermaphrodite can generate many other individuals.[7] In the well-studied roundworm Caenorhabditis elegans, males are very rare and only occur in populations that are in bad condition or stressed.[8] Androdioecy with hermaphroditic ancestry[edit] In animals, corals (phyla Cnidaria) and barnacles (Subphyla Crustacea), androdioecy has evolved from hermaphroditism. Many plants self-fertilize, and males may be sustained in a population when inbreeding depression is severe because males guarantee outcrossing.

Androdioecious species[edit] Despite their unlikely evolution, 115 androdioecious animal and about 50 androdioecious plant species are known.[2][9] These species include Anthozoa (Corals)[edit] Goniastra australensis Stylophora pistillata Nematoda (Roundworms)[edit] Rhabditidae (Order Rhabditida) Caenorhabditis briggsae Caenorhabditis elegans[8] Caenorhabditis sp. 11 Oscheius myriophila Oscheius dolchura Oscheius tipulae Oscheius guentheri Rhabditis rainai Rhabditis sp. (AF5) Rhabdias nigrovenosum Rhabdias rubrovenosa Rhabdias ranae Entomelas entomelas Diplogastridae (Order Rhabditida) Allodiplogaster sudhausi[10] Diplogasteroides magnus[11] Levipalatum texanum[12] Pristionchus boliviae[13] Pristionchus fissidentatus[14] Pristionchus maupasi[15] Pristionchus mayeri[13] Pristionchus pacificus Pristionchus triformis[16] Sudhausia aristotokia[17] Sudhausia crassa[17] Steinernematidae (Order Rhabditida) Steinernema hermaphroditum Allanotnematidae (Order Rhabditida) Allantonema mirabile Bradynema rigidum Dorylaimida Dorylaimus liratus Nemertea (Ribbon worms)[edit] Prostoma eilhardi Arthropoda[edit] Clam shrimp Eulimnadia texana[18] Eulimnadia africana Eulimnadia agassizii Eulimnadia antlei Eulimnadia braueriana Eulimnadia brasiliensis Eulimnadia colombiensis Eulimnadia cylondrova Eulimnadia dahli Eulimnadia diversa Eulimnadia feriensis Eulimnadia follisimilis Eulimnadia thompsoni Eulimnadia sp. A Eulimnadia sp. B Eulimnadia sp. C Tadpole shrimp Triops cancriformis[19] Triops newberryi Triops longicaudatus Barnacles Paralepas klepalae Paralepas xenophorae Koleolepas avis Koleolepas tinkeri Ibla quadrivalvis Ibla cumingii Ibla idiotica Ibla segmentata Calantica studeri Calantica siemensi Calantica spinosa Calantica villosa Arcoscalpellum sp. Euscalpellum squamuliferum Scalpellum peronii Scalpellum scalpellum Scalpellum vulgare Scillaelepas arnaudi Scillaelepas bocquetae Scillaelepas calyculacilla Scillaelepas falcate Scillaelepas fosteri Smilium hastatum Smilium peronii Chelonibia patula[20] Chelonibia testudinaria[21] Bathylasma alearum[22] Bathylasma corolliforme Conopea galeata[23] Conopea calceola[23] Conopea merrilli[23] Solidobalanus masignotus[24] Tetrapachylasma trigonum Megalasma striatum Octolasmis warwickii[25] Lysmata Lysmata wurdemanni Lysmata amboinensis Lysmata californica Lysmata bahia Lysmata intermedia Lysmata grabhami Lysmata seticaudata Lysmata nilita Lysmata hochi Lysmata nayaritensis Lysmata rafa Lysmata boggessi Lysmata ankeri Lysmata pederseni Lysmata debelius Lysmata galapaguensis Lysmata cf. trisetacea Insects Icerya bimaculata Icerya purchasi Crypticerya zeteki Annelida (Ringed worms)[edit] Salvatoria clavata Ophryotrocha gracilis Ophryotrocha hartmanni Ophryotrocha diadema Ophryotrocha bacci Ophryotrocha maculata Ophryotrocha socialis Chordata[edit] Kryptolebias marmoratus[26] Serranus fasciatus Serranus baldwini Plants[edit] Acer sp. (Maple)[27] Castilla elastica[28] Culcita macrocarpa Datisca glomerata (Durango root) Fraxinus lanuginosa (Japanese Ash) Fraxinus ornus Fuchsia microphylla Gagea serotina Mercurialis annua (Annual mercury)[29] Neobuxbaumia mezcalaensis[30] Nephelium lappaceum (Rambutan) Panax trifolius (Ginseng) Oxalis suksdorfii Phillyrea angustifolia Phillyrea latifolia Ricinocarpus pinifolius[31] Sagittaria lancifolia (sub-androdioecy)[32] Saxifraga cernua Schizopepon bryoniaefolius Spinifex littoreus Ulmus minor[33]

External links[edit] Ishida, Kiyoshi; Hiura, Tsutom (1998). "Pollen Fertility and Flowering Phenology in an Androdioecious Tree, Fraxinus lanuginosa (Oleaceae), in Hokkaido, Japan". International Journal of Plant Sciences. 159: 941–947. doi:10.1086/314088.  Pennisi, Elizabeth. "'Sex and the Single Killifish'" (PDF). Science. 313: 2006. doi:10.1126/science.313.5792.1381.  Diana Wolf. 'Breeding systems: Evolution of androdioecy'

See also[edit] Gynodioecy Plant sexuality

References[edit] ^ Pannell, JR. (2002). "The evolution and maintenance of androdioecy". Annual Review of Ecology and Systematics. 33: 397–425. doi:10.1146/annurev.ecolsys.33.010802.150419.  ^ a b Weeks, SC (2012). "The role of androdioecy and gynodioecy in mediating evolutionary transitions between dioecy and hermaphroditism in the Animalia". Evolution. 66 (12): 3670–3686. doi:10.1111/j.1558-5646.2012.01714.x. PMID 23206127.  ^ Charlesworth, D (1984). "Androdioecy and the evolution of dioecy". Biological Journal of the Linnean Society. 22 (4): 333–348. doi:10.1111/j.1095-8312.1984.tb01683.x.  ^ Darwin C. 1877. The different forms of flowers and plants of the same species. New York: Appleton. ^ Lloyd, DG (1975). "The maintenance of gynodioecy and androdioecy in angiosperms". Genetica. 45: 325–339. doi:10.1007/bf01508307.  ^ Charlesworth, B; Charlesworth, D (1978). "A Model for the Evolution of Dioecy and Gynodioecy". The American Naturalist. 112 (988): 975–997. doi:10.1086/283342.  ^ Pannell, J (2000). "A hypothesis for the evolution of androdioecy: the joint influence of reproductive assurance and local mate competition in a metapopulation". Evolutionary Ecology. 14 (3): 195–211. doi:10.1023/A:1011082827809.  ^ a b Stewart, AD; Phillips, PC (2002). "Selection and maintenance of androdioecy in Caenorhabditis elegans". Genetics. 160 (3): 975–982.  ^ Weeks, SC; Benvenuto, C; Reed, SK (2006). "When males and hermaphrodites coexist: a review of androdioecy in animals". Integrative and Comparative Biology. 46 (4): 449–464. doi:10.1093/icb/icj048.  ^ Fürst von Lieven A (2008). "Koerneria sudhausi n. sp. (Nematoda: Diplogastridae); a hermaphroditic diplogastrid with an egg shell formed by zygote and uterine components". Nematology. 10 (1): 27–45. doi:10.1163/156854108783360087.  ^ Kiontke K, Manegold A, Sudhaus W (2001). "Redescription of Diplogasteroides nasuensis Takaki, 1941 and D. magnus Völk, 1950 (Nematoda: Diplogastrina) associated with Scarabaeidae (Coleoptera)". Nematology. 3 (8): 817–832. doi:10.1163/156854101753625317.  ^ Ragsdale EJ, Kanzaki N, Sommer RJ (2014). "Levipalatum texanum n. gen., n. sp. (Nematoda: Diplogastridae), an androdioecious species from the south-eastern USA". Nematology. 16: 695–709. doi:10.1163/15685411-00002798.  ^ a b Kanzaki N, Ragsdale EJ, Herrmann M, Susoy V, Sommer RJ (2013). "Two androdioecious and one dioecious new species of Pristionchus (Nematoda: Diplogastridae): new reference points for the evolution of reproductive mode". Journal of Nematology. 45 (3): 172–194. PMC 3792836 . PMID 24115783.  ^ Kanzaki N, Ragsdale EJ, Herrmann M, Sommer RJ (2012). "Two new species of Pristionchus (Rhabditida: Diplogastridae): P. fissidentatus n. sp. from Nepal and La Réunion Island and P. elegans n. sp. from Japan". Journal of Nematology. 44 (1): 80–91. PMC 3593256 . PMID 23483847.  ^ Potts FA (1908). "Sexual phenomena in the free-living nematodes". Proceedings of the Cambridge Philosophical Society. 14: 373–375.  ^ Ragsdale EJ, Kanzaki N, Röseler W, Herrmann M, Sommer RJ (2013). "Three new species of Pristionchus (Nematoda: Diplogastridae) show morphological divergence through evolutionary intermediates of a novel feeding-structure polymorphism". Zoological Journal of the Linnean Society. 168: 671–698. doi:10.1111/zoj.12041.  ^ a b Hermmann M, Ragsdale EJ, Kanzaki N, Sommer RJ (2013). "Sudhausia aristotokia n. gen., n. sp. and S. crassa n. gen., n. sp. (Nematoda: Diplogastridae): viviparous new species with precocious gonad development". Nematology. 15: 1001–1020.  ^ Vicky G. Hollenbeck; Stephen C. Weeks; William R. Gould; Naida Zucker (2002). "Maintenance of androdioecy in the freshwater shrimp Eulimnadia texana: sexual encounter rates and outcrossing success". Behavioral Ecology. 13 (4): 561–570. doi:10.1093/beheco/13.4.561.  ^ Zierold, T; Hanfling, B; Gómez, A (2007). "Recent evolution of alternative reproductive modes in the'living fossil'Triops cancriformis". BMC Evolutionary Biology. 7 (1): 161. doi:10.1186/1471-2148-7-161. PMC 2075510 . PMID 17854482.  ^ Crisp, DJ (1983). "Chelonobia patula (Ranzani), a pointer to the evolution of the complemental male". Marine Biology Letters. 4: 281–294.  ^ Zardus, JD; Hadfield, MG (2004). "Larval Development and Complemental Males in Chelonibia testudinaria, a Barnacle Commensal with Sea Turtles". Journal of Crustacean Biology. 24 (3): 409–421. doi:10.1651/c-2476.  ^ Foster, BA (1983). "Complemental males in the barnacle Bathylasma alearum (cirripedia, pachylasmatidae)". Mem. Aus. Mus. 18: 133–140. doi:10.3853/j.0067-1967.18.1984.379.  ^ a b c McLaughlin, PA; Henry, DP (1972). "Comparative Morphology of Complemental Males in Four Species of Balanus (Cirripedia Thoracica)". Crustaceana. 22 (1): 13–30. doi:10.1163/156854072x00642.  ^ Henry, DP; McLaughlin, PA (1967). "A Revision of the Subgenus Solidobalanus Hoek (Cirripedia Thoracica) including a Description of a New Species with Complemental Males". Crustaceana. 12 (1): 43–58. doi:10.1163/156854067x00693.  ^ Yusa, Y; Takemura, M; Miyazaki, K; Watanabe, T; Yamato, S (2010). "Dwarf Males of Octolasmis warwickii (Cirripedia: Thoracica): The First Example of Coexistence of Males and Hermaphrodites in the Suborder Lepadomorpha". The Biological Bulletin. 218 (3): 259–265.  ^ Mackiewicz; Tatarenkov, A; Taylor, DS; Turner, BJ; Avise, JC; et al. (2006). "Extensive outcrossing and androdioecy in a vertebrate species that otherwise reproduces as a self-fertilizing hermaphrodite". Proc Natl Acad Sci USA. 103 (26): 9924–9928. doi:10.1073/pnas.0603847103. PMC 1502555 . PMID 16785430.  ^ Gleiser G, Verdú M. 2005. Repeated evolution of dioecy from androdioecy in Acer" New Phytologist 165(2):633-640. doi=10.1111/j.1469-8137.2004.01242.x ^ Sakai, S (2001). "Thrips pollination of androdioecious Castilla elastica (Moraceae) in a seasonal tropical forest". American Journal of Botany. 88 (9): 1527–1534. doi:10.2307/3558396. PMID 21669685.  ^ Pannell J (1997). "Widespread functional androdioecy in Mercurialis annua L. (Euphorbiaceae)". Biological Journal of the Linnean Society. 61: 95–116. doi:10.1111/j.1095-8312.1997.tb01779.x.  ^ Valiente-Banuet, A; Rojas-Martínez, A; Del Coro, Arizmendi M; Dávila, P (1997). "Pollination biology of two columnar Cacti (Neobuxbaumia mezcalaensis and Neobuxbaumia macrocephala) in the Tehuacan Valley, central Mexico". American Journal of Botany. 84 (4): 452. doi:10.2307/2446020.  ^ Thomson JD, Shivanna KR, Kenrick J and Knox RB. 1989" American Journal of Botany 76 (7):1048-1059 ^ Muenchow, G (1998). "Subandrodioecy and male fitness in Sagittaria lancifolia subsp. lancifolia (Alismataceae)". American Journal of Botany. 85 (4): 513. doi:10.2307/2446435.  ^ López-Almansa, JC; Pannell, JR; Gil, L (2003). "Female sterility in Ulmus minor (Ulmaceae): a hypothesis invoking the cost of sex in a clonal plant". American Journal of Botany. 90 (4): 603–609. doi:10.3732/ajb.90.4.603. PMID 21659155.  Retrieved from "" Categories: Biology of genderMating systemsHidden categories: Wikipedia references cleanup from December 2013All articles needing references cleanupArticles covered by WikiProject Wikify from December 2013All articles covered by WikiProject Wikify

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