Androdioecy
Androdioecy is a reproductive system characterized by the coexistence of males and hermaphrodites. Androdioecy is rare in comparison to the other major reproductive systems: dioecy, gynodioecy and hermaphroditism.[1] In animals, androdioecy has been considered an important stepping stone in the transition from dioecy to hermaphroditism, and vice versa.[2]
Evolution of androdioecy
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
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
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
Despite their unlikely evolution, 115 androdioecious animal and about 50 androdioecious plant species are known.[2][9] These species include
Anthozoa (Corals)
Nematoda (Roundworms)
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)
Allanotnematidae (Order Rhabditida)
Nemertea (Ribbon worms)
- 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
- 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 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
Annelida (Ringed worms)
- Salvatoria clavata
- Ophryotrocha gracilis
- Ophryotrocha hartmanni
- Ophryotrocha diadema
- Ophryotrocha bacci
- Ophryotrocha maculata
- Ophryotrocha socialis
Plants
- 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]
See also
References
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- ^ 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 (8): 1001–1020. doi:10.1163/15685411-00002738.
- ^ 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.
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{{cite journal}}
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External links
- 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 (6): 941–947. doi:10.1086/314088.
- Pennisi, Elizabeth (2006). "Sex and the Single Killifish" (PDF). Science. 313 (5792): 2006. doi:10.1126/science.313.5792.1381. PMID 16959986. Archived from the original (PDF) on 2008-10-01. Retrieved 2008-01-25.
- Diana Wolf. 'Breeding systems: Evolution of androdioecy'