Androdioecy
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Androdioecy is a
reproductive system The reproductive system of an organism, also known as the genital system, is the biological system made up of all the anatomical organs involved in sexual reproduction. Many non-living substances such as fluids, hormones, and pheromones are ...
characterized by the coexistence of
males Male (symbol: ♂) is the sex of an organism that produces the gamete (sex cell) known as sperm, which fuses with the larger female gamete, or ovum, in the process of fertilization. A male organism cannot reproduce sexually without access to ...
and hermaphrodites. Androdioecy is rare in comparison with the other major reproductive systems:
dioecy Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproducti ...
,
gynodioecy Gynodioecy is a rare breeding system that is found in certain flowering plant species in which female and hermaphroditic plants coexist within a population. Gynodioecy is the evolutionary intermediate between hermaphroditism (exhibiting both fem ...
and
hermaphroditism In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have s ...
. In animals, androdioecy has been considered a stepping stone in the transition from dioecy to hermaphroditism, and vice versa. Androdioecy is sometimes referred to as a mixed breeding system with trioecy and
gynodioecy Gynodioecy is a rare breeding system that is found in certain flowering plant species in which female and hermaphroditic plants coexist within a population. Gynodioecy is the evolutionary intermediate between hermaphroditism (exhibiting both fem ...
. It is a dimorphic sexual system in plants alongside
gynodioecy Gynodioecy is a rare breeding system that is found in certain flowering plant species in which female and hermaphroditic plants coexist within a population. Gynodioecy is the evolutionary intermediate between hermaphroditism (exhibiting both fem ...
and
dioecy Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproducti ...
.


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. 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. Androdioecy can evolve either from hermaphroditic ancestors through the invasion of males or from dioecious ancestors through the invasion of hermaphrodites. 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 Out-crossing or out-breeding is the technique of crossing between different breeds. This is the practice of introducing distantly related genetic material into a breeding line, thereby increasing genetic diversity. Outcrossing can be a usefu ...
. Hermaphrodites may be beneficial in colonizing new habitats, because a single hermaphrodite can generate many other individuals. In the well-studied roundworm ''
Caenorhabditis elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' (r ...
'', males are very rare and only occur in populations that are in bad condition or stressed. In ''Caenorhabditis elegans'' androdioecy is thought to have evolved from dioecy, through a trioecous intermediate.


Androdioecy with hermaphroditic ancestry

In barnacles androdioecy evolved from hermaphroditism. Many plants self-fertilize, and males may be sustained in a population when
inbreeding depression Inbreeding depression is the reduced biological fitness which has the potential to result from inbreeding (the breeding of related individuals). Biological fitness refers to an organism's ability to survive and perpetuate its genetic material. ...
is severe because males guarantee outcrossing.


Types of Androdioecy

The most common form of androdioecy in animals involves hermaphrodites that can reproduce by autogamy or allogamy through ovum with males. However, this type doesn’t involve outcrossing with sperm. This type of androdioecy generally occurs in predominantly gonochoric taxonomy groups. One type of androdioecy contains outcrossing hermaphrodites which is present in some
angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
. Another type of androdioecy has males and simultaneous hermaphrodites in a population due to developmental or conditional sex allocation. Like in some fish species small individuals are hermaphrodites and under circumstances of high density, large individuals become male.


Androdioecious species

Despite their unlikely evolution, 115 androdioecious animal and about 50 androdioecious plant species are known. These species include


Anthozoa Anthozoa is a subphylum of marine invertebrates which includes the sea anemones, stony corals and soft corals. Adult anthozoans are almost all attached to the seabed, while their larvae can disperse as part of the plankton. The basic unit of ...
(Corals)

*'' Goniastra australensis'' *'' Stylophora pistillata''


Nematoda The nematodes ( or grc-gre, Νηματώδη; la, Nematoda) or roundworms constitute the phylum Nematoda (also called Nemathelminthes), with plant-parasitic nematodes also known as eelworms. They are a diverse animal phylum inhabiting a broa ...
(Roundworms)

Rhabditidae (Order
Rhabditida Rhabditida is an order of free-living, zooparasitic, and phytoparasitic microbivorous nematodes living in soil. The Cephalobidae, Panagrolaimidae, Steinernematidae, and Strongyloididae seem to be closer to the Tylenchia, regardless of whet ...
) *'' Caenorhabditis briggsae'' *''
Caenorhabditis elegans ''Caenorhabditis elegans'' () is a free-living transparent nematode about 1 mm in length that lives in temperate soil environments. It is the type species of its genus. The name is a blend of the Greek ''caeno-'' (recent), ''rhabditis'' (r ...
'' *'' Caenorhabditis sp. 11'' *''
Oscheius myriophila ''Oscheius'' is a genus of nematode. '' O. tipulae'' is a satellite developmental genetic model organism used to study vulva formation. In phylogenetic studies, based on the analysis of sequences of three nuclear genes, ''Oscheius'' groups wit ...
'' *''
Oscheius dolchura ''Oscheius'' is a genus of nematode. '' O. tipulae'' is a satellite developmental genetic model organism used to study vulva formation. In phylogenetic studies, based on the analysis of sequences of three nuclear genes, ''Oscheius'' groups wit ...
'' *'' Oscheius tipulae'' *'' Oscheius guentheri'' *'' Rhabditis rainai'' *'' Rhabditis sp. (AF5)'' *''
Rhabdias nigrovenosum ''Rhabdias'' is a genus of nematodes belonging to the family Rhabdiasidae. The genus has cosmopolitan distribution. Species The genus includes the following species: * ''Rhabdias aegyptiaca'' * ''Rhabdias africanus'' * ''Rhabdias alabialis'' * ...
'' *'' Rhabdias rubrovenosa'' *'' Rhabdias ranae'' *'' Entomelas entomelas''
Diplogastridae Diplogastridae, formerly Diplogasteridae, are a family of nematodes (roundworms) known from a wide range of habitats, often in commensal or parasitic associations with insects. Description Diplogastrid nematodes are characterized by a distinct ...
(Order Rhabditida) *'' Allodiplogaster sudhausi'' *'' Diplogasteroides magnus'' *''
Levipalatum texanum ''Levipalatum texanum'' is a free-living nematode (roundworm) in the family Diplogastridae. The species is androdioecious, consisting of self-fertile hermaphrodites which are morphologically females, and males. It is known from the south-eastern ...
'' *'' Pristionchus boliviae'' *'' Pristionchus fissidentatus'' *'' Pristionchus maupasi'' *'' Pristionchus mayeri'' *''
Pristionchus pacificus ''Pristionchus pacificus'' is a species of free-living nematodes (roundworms) in the family Diplogastridae. The species has been established as a satellite model organism to ''Caenorhabditis elegans'', with which it shared a common ancestor 200 ...
'' *'' Pristionchus triformis'' *''
Sudhausia aristotokia ''Sudhausia'' is a genus of nematodes (roundworms) of the family Diplogastridae. They live in association with dung beetles and are primarily known from Africa. Species of ''Sudhausia'' show a suite of biological features that, together, are unus ...
'' *''
Sudhausia crassa ''Sudhausia'' is a genus of nematodes (roundworms) of the family Diplogastridae. They live in association with dung beetles and are primarily known from Africa. Species of ''Sudhausia'' show a suite of biological features that, together, are unus ...
'' Steinernematidae (Order Rhabditida) *''
Steinernema hermaphroditum ''Steinernema hermaphroditum'' is a species of nematodes in the Steinernematidae family. It was the first steinernematid reported to be able to self fertilize. The species has been classified as androdioecious Androdioecy is a reproductive sy ...
'' Allanotnematidae (Order Rhabditida) *'' Allantonema mirabile'' *'' Bradynema rigidum'' Dorylaimida *'' Dorylaimus liratus''


Nemertea Nemertea is a phylum of animals also known as ribbon worms or proboscis worms, consisting of 1300 known species. Most ribbon worms are very slim, usually only a few millimeters wide, although a few have relatively short but wide bodies. Many h ...
(Ribbon worms)

*'' Prostoma eilhardi''


Arthropoda Arthropods (, (gen. ποδός)) are invertebrate animals with an exoskeleton, a segmented body, and paired jointed appendages. Arthropods form the phylum Arthropoda. They are distinguished by their jointed limbs and cuticle made of chitin ...

Clam shrimp *'' Eulimnadia texana'' *''
Eulimnadia africana ''Eulimnadia'' is a genus of branchiopods in the family Limnadiidae. There are about 13 described species in ''Eulimnadia''. Species * '' Eulimnadia agassizii'' Packard, 1874 * '' Eulimnadia antillarum'' (Baird, 1852) * '' Eulimnadia antlei'' ...
'' *''
Eulimnadia agassizii ''Eulimnadia'' is a genus of branchiopods in the family Limnadiidae Limnadiidae is a family of crustaceans in the order Spinicaudata that live in seasonal wetlands, inland saline pools and lakes. They are found on all the world's continents ...
'' *''
Eulimnadia antlei ''Eulimnadia antlei'' is a species of branchiopod in the family Limnadiidae Limnadiidae is a family of crustaceans in the order Spinicaudata that live in seasonal wetlands, inland saline pools and lakes. They are found on all the world's cont ...
'' *'' Eulimnadia braueriana'' *'' Eulimnadia brasiliensis'' *''
Eulimnadia colombiensis ''Eulimnadia'' is a genus of branchiopods in the family Limnadiidae. There are about 13 described species in ''Eulimnadia''. Species * ''Eulimnadia agassizii'' Packard, 1874 * ''Eulimnadia antillarum'' (Baird, 1852) * ''Eulimnadia antlei'' Mack ...
'' *'' Eulimnadia cylondrova'' *'' Eulimnadia dahli'' *''
Eulimnadia diversa ''Eulimnadia'' is a genus of branchiopods in the family Limnadiidae. There are about 13 described species in ''Eulimnadia''. Species * ''Eulimnadia agassizii'' Packard, 1874 * ''Eulimnadia antillarum'' (Baird, 1852) * ''Eulimnadia antlei'' Mack ...
'' *'' Eulimnadia feriensis'' *'' Eulimnadia follisimilis'' *'' Eulimnadia thompsoni'' *'' Eulimnadia sp. A'' *''
Eulimnadia sp. B ''Eulimnadia'' is a genus of branchiopods in the family Limnadiidae. There are about 13 described species in ''Eulimnadia''. Species * ''Eulimnadia agassizii'' Packard, 1874 * ''Eulimnadia antillarum'' (Baird, 1852) * ''Eulimnadia antlei'' Mack ...
'' *'' Eulimnadia sp. C''
Tadpole shrimp The order Notostraca, containing the single family Triopsidae, is a group of crustaceans known as tadpole shrimp or shield shrimp. The two genera, ''Triops'' and '' Lepidurus'', are considered living fossils, with similar forms having existed sinc ...
*''
Triops cancriformis ''Triops cancriformis'', European tadpole shrimp or tadpole shrimp, is a species of tadpole shrimp found in Europe to the Middle East and India. Due to habitat destruction, many populations have recently been lost across its European range, ...
'' *''
Triops newberryi ''Triops newberryi'' is a species of ''Triops'' found on the western coast of North America, commonly in valleys throughout the states of Washington, Oregon, California, and small areas of Nevada, Utah, New Mexico, and Mexico, with at least one ...
'' *''
Triops longicaudatus ''Triops longicaudatus'' (commonly called American tadpole shrimp or longtail tadpole shrimp) is a freshwater crustacean of the order Notostraca, resembling a miniature horseshoe crab. It is characterized by an elongated, segmented body, a flatten ...
''
Barnacles A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallow and tidal waters, typically in erosiv ...
*''
Paralepas klepalae Paralepas is a genus of goose barnacles in the family Heteralepadidae. Species The World Register of Marine Species includes the following species In biology, a species is the basic unit of Taxonomy (biology), classification and a taxo ...
'' *'' Paralepas xenophorae'' *''
Koleolepas avis ''Koleolepas'' is a genus of crustaceans belonging to the family Heteralepadidae Heteralepadidae is a family of goose barnacles. Genera The World Register of Marine Species The World Register of Marine Species (WoRMS) is a taxonomic datab ...
'' *''
Koleolepas tinkeri ''Koleolepas'' is a genus of crustaceans belonging to the family Heteralepadidae. The species of this genus are found in Southwestern Asia. Species: *''Koleolepas avis ''Koleolepas'' is a genus of crustaceans belonging to the family Heteral ...
'' *''
Ibla quadrivalvis ''Ibla quadrivalvis'' is a species of barnacle in the Iblidae family. The common name for this species is hairy stalked barnacle. The species was studied by Charles Darwin. He first described that this specie is androdioecious Androdioecy is a ...
'' *'' Ibla cumingii'' *'' Ibla idiotica'' *''
Ibla segmentata Ragusa (; scn, Rausa ; la, Ragusia) is a city and '' comune'' in southern Italy. It is the capital of the province of Ragusa, on the island of Sicily, with 73,288 inhabitants in 2016. It is built on a wide limestone hill between two deep va ...
'' *''
Calantica studeri ''Calantica'' is a genus of barnacle A barnacle is a type of arthropod constituting the subclass Cirripedia in the subphylum Crustacea, and is hence related to crabs and lobsters. Barnacles are exclusively marine, and tend to live in shallo ...
'' *''
Calantica siemensi ''Calantica'' is a genus of barnacles in the family Calanticidae, containing the following species: *'' Calantica affinis'' Broch, 1922 *'' Calantica darwini'' Jones & Hosie, 2009 *'' Calantica eos'' (Pilsbry, 1907) *'' Calantica flagellata'' ...
'' *''
Calantica spinosa ''Calantica'' is a genus of barnacles in the family Calanticidae, containing the following species: *'' Calantica affinis'' Broch, 1922 *'' Calantica darwini'' Jones & Hosie, 2009 *'' Calantica eos'' (Pilsbry, 1907) *'' Calantica flagellata'' ...
'' *'' 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 ''Chelonibia'' is a genus of acorn barnacles in the family Chelonibiidae of the subphylum Crustacea. Its members are epizootic and live attached to manatees, turtles, marine molluscs, crabs and horseshoe crabs in all tropical and subtropical o ...
'' *''
Chelonibia testudinaria ''Chelonibia testudinaria'' is a species of barnacle in the family Chelonibiidae. It is native to the Atlantic Ocean, Mediterranean Sea and Gulf of Mexico where it lives as a symbiont on sea turtles, being particularly abundant on the loggerhead ...
'' *''
Bathylasma alearum ''Bathylasma alearum'' is a species of barnacles in the Pachylasmatidae family. The species is androdioecious Androdioecy is a reproductive system characterized by the coexistence of males and hermaphrodites. Androdioecy is rare in comparison wi ...
'' *''
Bathylasma corolliforme ''Bathylasma'' is a genus of barnacles. Species Species in this genus include. * '' Bathylasma alearum'' * '' Bathylasma aucklandicum'' * '' Bathylasma chilense'' * '' Bathylasma corolliforme'' * '' Bathylasma hirsutum'' * '' Bathylasma ranga ...
'' *'' Conopea galeata'' *'' Conopea calceola'' *'' Conopea merrilli'' *'' Solidobalanus masignotus'' *'' Tetrapachylasma trigonum'' *'' Megalasma striatum'' *'' Octolasmis warwickii'' Lysmata *'' Lysmata wurdemanni'' *''
Lysmata amboinensis ''Lysmata amboinensis'' is an omnivorous shrimp species known by several common names including the Pacific cleaner shrimp. It is considered a cleaner shrimp as eating parasites and dead tissue from fish makes up a large part of its diet. The s ...
'' *''
Lysmata californica ''Lysmata californica'', known generally as the red rock shrimp or lined shrimp, is a species of caridean shrimp in the family Hippolytidae Hippolytidae is a family of cleaner shrimp, also known as broken-back shrimp or anemone shrimp. The te ...
'' *'' Lysmata bahia'' *'' Lysmata intermedia'' *''
Lysmata grabhami ''Lysmata grabhami'' is a species of saltwater shrimp in the family Hippolytidae. It was first described by Gordon in 1935. It occurs in the tropical and subtropical Atlantic Ocean and is a cleaner shrimp, operating a cleaning station to which ...
'' *'' Lysmata seticaudata'' *'' Lysmata nilita'' *'' Lysmata hochi'' *'' Lysmata nayaritensis'' *''
Lysmata rafa ''Lysmata'' is a genus of shrimp in the infraorder Caridea, the caridean shrimp. The genus belongs to the family Lysmatidae. ''Lysmata'' are popular ornamental shrimp in the marine aquarium trade for their bright color patterns, interesting be ...
'' *'' Lysmata boggessi'' *'' Lysmata ankeri'' *''
Lysmata pederseni ''Lysmata pederseni'' is a species of saltwater shrimp first classified as ''Lysmata wurdemanni ''Lysmata wurdemanni'', commonly known as the peppermint shrimp, is a species of shrimp. Description It reaches in length, and is named for the ...
'' *''
Lysmata debelius ''Lysmata debelius'' is a species of cleaner shrimp indigenous to the Indo-Pacific. It is popular in the aquarium trade, where it is known as the fire shrimp, blood shrimp or scarlet cleaner shrimp. Taxonomy ''Lysmata debelius'' was first desc ...
'' *'' Lysmata galapaguensis'' *''
Lysmata cf. trisetacea ''Lysmata'' is a genus of shrimp in the infraorder Caridea, the caridean shrimp. The genus belongs to the family Lysmatidae. ''Lysmata'' are popular ornamental shrimp in the marine aquarium trade for their bright color patterns, interesting be ...
''
Insects Insects (from Latin ') are pancrustacean hexapod invertebrates of the class Insecta. They are the largest group within the arthropod phylum. Insects have a chitinous exoskeleton, a three-part body (head, thorax and abdomen), three pairs of j ...
*'' Icerya bimaculata'' *'' Icerya purchasi'' *'' Crypticerya zeteki''


Annelida (Ringed worms)

*'' Salvatoria clavata'' *''
Ophryotrocha gracilis ''Ophryotrocha'' is a genus of marine polychaete worms in the family Dorvilleidae. Species The World Register of Marine Species lists the following species in the genus : *'' Ophryotrocha adherens'' Paavo, Bailey-Brock & Akesson, 2000 *'' Op ...
'' *'' Ophryotrocha hartmanni'' *''
Ophryotrocha diadema ''Ophryotrocha'' is a genus of marine polychaete worms in the family Dorvilleidae. Species The World Register of Marine Species lists the following species in the genus : *'' Ophryotrocha adherens'' Paavo, Bailey-Brock & Akesson, 2000 *'' Op ...
'' *'' Ophryotrocha bacci'' *''
Ophryotrocha maculata ''Ophryotrocha'' is a genus of marine polychaete worms in the family Dorvilleidae Dorvilleidae is a family of polychaetes Polychaeta () is a paraphyletic class of generally marine annelid worms, commonly called bristle worms or polychae ...
'' *'' Ophryotrocha socialis''


Chordata A chordate () is an animal of the phylum Chordata (). All chordates possess, at some point during their larval or adult stages, five synapomorphies, or primary physical characteristics, that distinguish them from all the other taxa. These fi ...

*'' Kryptolebias marmoratus'' *'' Serranus fasciatus'' *'' Serranus baldwini''


Angiosperms Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants ...
(Flowering plants)

*Some '' Acer'' (maple) species *'' Castilla elastica'' *''
Culcita macrocarpa ''Culcita macrocarpa'', the woolly tree fern, is a species of fern in the family Culcitaceae native to Macaronesia and parts of the Iberian Peninsula, where it might have been introduced. Description ''Culcita macrocarpa'' is a large plant ...
'' *'' Datisca cannabina'' (false hemp) *'' Datisca glomerata'' (Durango root) *'' Fraxinus lanuginosa'' (Japanese ash) *''
Fraxinus ornus ''Fraxinus'' (), commonly called ash, is a genus of flowering plants in the olive and lilac family, Oleaceae. It contains 45–65 species of usually medium to large trees, mostly deciduous, though a number of subtropical species are evergr ...
'' *'' Fuchsia microphylla'' *''
Gagea serotina ''Gagea serotina'', synonym ''Lloydia serotina'', is an Arctic–alpine flowering plant of the lily family. It is widespread across the mountainous parts of western North America, from Alaska to New Mexico, and in Europe is found in the Alps an ...
'' *'' Mercurialis annua'' (Annual mercury) *'' Neobuxbaumia mezcalaensis'' *'' Nephelium lappaceum'' (Rambutan) *'' Panax trifolius'' ( Ginseng) *'' Oxalis suksdorfii'' *''
Phillyrea angustifolia ''Phillyrea angustifolia'', the narrow-leaved mock privet,RHS Plants. Available at: https://www.rhs.org.uk/Plants/24488/i-Phillyrea-angustifolia-i/Details ccessed 22/02/21/ref> is a species of flowering plant in the olive family Oleaceae, native ...
'' *'' Phillyrea latifolia'' *'' Ricinocarpus pinifolius'' *'' Sagittaria lancifolia'' (sub-androdioecy) *'' Saxifraga cernua'' *'' Schizopepon bryoniaefolius'' *'' Spinifex littoreus'' *'' Ulmus minor''


See also

*
Gynodioecy Gynodioecy is a rare breeding system that is found in certain flowering plant species in which female and hermaphroditic plants coexist within a population. Gynodioecy is the evolutionary intermediate between hermaphroditism (exhibiting both fem ...
*
Plant sexuality Plant reproductive morphology is the study of the physical form and structure (the morphology) of those parts of plants directly or indirectly concerned with sexual reproduction. Among all living organisms, flowers, which are the reproductive s ...
*
Dioecy Dioecy (; ; adj. dioecious , ) is a characteristic of a species, meaning that it has distinct individual organisms (unisexual) that produce male or female gametes, either directly (in animals) or indirectly (in seed plants). Dioecious reproducti ...
* Trioecy *
Hermaphrodite In reproductive biology, a hermaphrodite () is an organism that has both kinds of reproductive organs and can produce both gametes associated with male and female sexes. Many taxonomic groups of animals (mostly invertebrates) do not have ...
* Monoicy


References


External links

* *{{cite journal , last1 = Pennisi , first1 = Elizabeth , author-link = Elizabeth Pennisi , s2cid = 84857451 , year = 2006, title = Sex and the Single Killifish , url = http://www3.uakron.edu/biology/science06.pdf , journal = Science , volume = 313 , issue = 5792, page = 2006 , doi = 10.1126/science.313.5792.1381 , pmid = 16959986 , access-date = 2008-01-25 , archive-url = https://web.archive.org/web/20081001223411/http://www3.uakron.edu/biology/science06.pdf , archive-date = 2008-10-01 , url-status = dead *Diana Wolf
'Breeding systems: Evolution of androdioecy'
Sex Mating systems Sexual system