Mayaheros

{{Short description|Genus of fishes}} {{Automatic taxobox | image = Mayan cichlid.jpg | image_caption = ''Mayaheros urophthalmus'' | taxon = Mayaheros | authority = Říčan & Piálek, 2016<ref name=Rican2016/> | type_species = ''Heros urophthalmus'' | type_species_authority = Günther, 1862 }}

'''''Mayaheros''''' is a genus of cichlid fish that is native to Mexico and northern Central America. This genus has a disjunct distribution, with the ''M. urophthalmus'' group being found in the Atlantic drainages of southeastern Mexico (southern Veracruz east to the Yucatán Peninsula and southwards), Belize, eastern Guatemala, northern Honduras, and northeastern Nicaragua, while ''M. beani'' is the northernmost cichlid in the Pacific drainage, ranging from Jalisco to Sonora in northwestern Mexico.<ref name=Rican2016>{{cite journal|author=Říčan, O.|author2=Piálek, L.|author3=Dragová, K.|author4=Novák, J.|name-list-style=amp|year=2016|title=Diversity and evolution of the Middle American cichlid fishes (Teleostei: Cichlidae) with revised classification|journal=Vertebrate Zoology|volume=66|issue=1|pages=1–102|doi=10.3897/vz.66.e31534 |s2cid=257203657 |doi-access=free}}</ref><ref name=IUCNbeani>{{cite iucn |author=Lyons, T.J. |date=2019 |title=''Mayaheros beani'' |volume=2019 |article-number=e.T191844A2007699 |doi=10.2305/IUCN.UK.2019-2.RLTS.T191844A2007699.en |access-date=29 December 2021}}</ref><ref name=IUCNurophthalmus>{{cite iucn |author=Schmitter-Soto, J. & Vega-Cendejas, M. |date=2019 |title=''Mayaheros urophthalmus'' |volume=2019 |article-number=e.T62217A3110044 |doi=10.2305/IUCN.UK.2019-2.RLTS.T62217A3110044.en |access-date=29 December 2021}}</ref> Both inhabit a wide range of habitats, such as freshwater rivers, streams, lakes, ponds, pools and marshes, as well as brackish waters including estuaries, lagoons, and mangroves, with members of the ''M. urophthalmus'' group sometimes even occurring in caves or coastal marine waters.<ref name=IUCNbeani/><ref name=IUCNurophthalmus/><ref name=AqGl28Dec>{{cite web| title=Mayaheros beani (formerly: Cichlasoma b.) | url=https://www.aquariumglaser.de/en/fish-archives/mayaheros-beani-formerly-cichlasoma-b/ | date=28 December 2021 | publisher=AquariumGlaser | access-date=29 December 2021 }}</ref> They are medium-sized to large cichlids that are omnivorous, feeding mostly on small animals, but also plants and detritus.<ref name=Bergmanna2005>{{cite journal| last1=Bergmann | first1=G.T. | last2=Motta | first2=P.J. | year=2005 | title=Diet and morphology through ontogeny of the nonindigenous Mayan cichlid 'Cichlasoma (Nandopsis)' urophthalmus (Günther 1862) in southern Florida | journal=Environmental Biology of Fishes | volume=72 | issue=2 | pages=205–211 | doi=10.1007/s10641-004-1480-1 | bibcode=2005EnvBF..72..205B | s2cid=7750210 }}</ref><ref>{{cite web| title=Mayaheros urophthalmus – Maya Cichlide | url=https://aquainfo.nl/artikel/mayaheros-urophthalmum-maya-cichlide/ | publisher=AquaInfo | access-date=30 December 2021 }}</ref><ref>{{cite web| title=Mayaheros beani | url=https://aquainfo.nl/artikel/mayaheros-beani/ | publisher=AquaInfo | access-date=30 December 2021 }}</ref>

''Mayaheros'' cichlids are sometimes kept in aquaria and commonly caught for food. Broadstock of the ''M. urophthalmus'' group that is better suited for farming is being developed by Mexican authorities;<ref>{{cite web| title=Researchers aim to boost production of two native fish species in Mexico | url=https://aquafishcrsp.oregonstate.edu/researchers-aim-boost-production-two-native-fish-species-mexico | date=27 June 2021 | publisher=AquaFish, Oregon State University | access-date=29 December 2021 }}</ref> ''M. beani'' also has potential in farming, but preliminary tests have been complicated by its high levels of aggression.<ref>{{cite journal| last1=Martinez-Cardenas | first1=L. | last2=Valdez-Hernández | first2=E.F. | last3=González-Díaz | first3=A.A. | last4=Soria-Barreto | first4=M. | last5=Castañeda-Chavez | first5=M.R. | last6=Ruiz-Velazco | first6=J.M. | last7=Peña-Messina | first7=E. | last8=Robles-Bermudez | first8=A. | year=2014 | title=Preliminary observations on Cichlasoma beani in culture conditions | journal=Lat. Am. J. Aquat. Res. | volume=43 | issue=3 | pages=639–643 | doi=10.3856/vol42-issue3-fulltext-20 | doi-access=free }}</ref> Both ''M. beani'' and ''M. urophthalmus'' are overall common and regarded as least concern by the IUCN, although certain local populations are under pressure from habitat loss, pollution, overfishing and introduced species.<ref name=IUCNbeani/><ref name=IUCNurophthalmus/> A captive breeding program exists for a few populations of ''M. beani''.<ref name=AqGl28Dec/> If many species are recognized within the ''M. urophthalmus'' group (''see #Species''), most are restricted to a single location in the Yucatán Peninsula region, such as a lagoon, cenote, cave, island, or bay, and some are seriously threatened. The most threatened –if not already extinct– are likely ''M. conchitae'' of the Conchita Cenote and ''M. ericymba'' of the Sambulá Cave (''M. ericymba'' has enlarged eyes and sensory pores, likely an adaption to its low-light habitat<ref>{{cite journal| author=Hubbs, C.L. | author-link=Carl Leavitt Hubbs | year=1938 | title=Fishes from the caves of Yucatan | journal=Carnegie Institution of Washington Publications | volume=491 | pages=261–295 }}</ref>), which both are located at the city of Mérida. The Conchita Cenote appears to have disappeared entirely, and while the Sambulá Cave still exists, ''M. ericymba'' has not been seen since the 1930s.<ref name=BarrientosMedina2005>{{citation| last=Barrientos-Medina | first=R.C. | year=2005 | title=Estado taxonómico de la mojarra rayada "Cichlasoma" urophthalmus Günther, 1862 (Teleostei: Cichlidae) | publisher=El Colegio de la Frontera Sur y la Universidad Autónoma de Yucatán, Chetumal, Quintana Roo }}</ref><ref>{{cite web| last=Barrientos-Medina | first=R.C. | date=8 March 2003 | title=The Mayan cichlid, 'Cichlasoma' urophthalmus: An Overview | url=https://cichlidae.com/article.php?id=181 | publisher=CichlidRoom | access-date=30 December 2021 }}</ref> If recognized as valid species or subspecies of ''M. urophthalmus'', these two are the only likely extinct cichlids of the Americas.<ref>{{cite journal| last=Burkhead | first=N.M. | year=2012 | title=Extinction Rates in North American Freshwater Fishes, 1900–2010 | journal=BioScience | volume=62 | issue=9 | pages=798–808 | doi=10.1525/bio.2012.62.9.5 | s2cid=86188840 | doi-access=free }}</ref><ref>{{cite web| title=Neopterygii (New-Finned Fish) | url=https://recentlyextinctspecies.com/databases/neopterygii-new-finned-fish | publisher=recentlyextinctspecies.com | access-date=30 January 2021 }}</ref> Conversely, ''M. urophthalmus'' has been introduced outside its native range in Oaxaca (Mexico), Florida (United States), the Philippines, Thailand and Singapore.<ref name=IUCNurophthalmus/><ref>{{cite web| title=Cichlasoma urophthalmum (Mayan cichlid) | url=https://www.cabi.org/isc/datasheet/88479 | publisher=CABI - Invasive Species Compendium | access-date=30 December 2021 }}</ref><ref>{{cite journal| last1=Ordoñez | first1=J.F.F. | last2=Asis | first2=A.M.J.M. | last3=Catacutan | first3=B.J. | last4=Pena | first4=J.d. | last5=Santos | first5=M.D. | year=2015 | title=First report on the occurrence of invasive black-chin tilapia Sarotherodon melanotheron (Ruppell, 1852) in Manila Bay and of Mayan cichlid Cichlasoma urophthalmus (Gunther, 1892) in the Philippines | journal=BioInvasions Records | volume=4 | issue=2 | pages=115–124 | doi= 10.3391/bir.2015.4.2.08 | doi-access=free }}</ref> It is unclear which member of the ''M. urophthalmus'' group is involved in these introductions, except in Florida where the population was based on individuals from several different locations in the ''M. urophthalmus'' group's native range.<ref name=Harrison2014>{{cite journal| last1=Harrison | first1=E. | last2=Trexler | first2=J.C. | last3=Collins | first3=T.M. | last4=Vazquez-Domınguez | first4=E. | last5=Razo-Mendivil | first5=U. | last6=Matamoros | first6=W.A. | last7=Barrientos | first7=C. | year=2014 | title=Genetic Evidence for Multiple Sources of the Non-Native Fish Cichlasoma urophthalmus (Günther; Mayan Cichlids) in Southern Florida | journal=PLOS ONE | volume=9 | issue=9 | pages=1–12 | doi=10.1371/journal.pone.0104173 | pmid=25184569 | pmc=4153574 | bibcode=2014PLoSO...9j4173H | doi-access=free }}</ref>

==Taxonomy== Both the species ''M. beani'' and ''M. urophthalmus'' were first scientifically described as members of the genus ''Heros'' and then moved to ''Cichlasoma'' despite clearly belonging elsewhere.<ref name=CoF>{{Catalog of Fishes |genus=Mayaheros |list=genus |access-date=29 December 2021}}</ref><ref name=Kullander2003>{{cite book| author=Kullander, S.O. | author-link=Sven O. Kullander | year= 2003 | chapter=Family Cichlidae (Cichlids) | pages=605–654 | editor-last=Reis | editor-first=R.E. | editor-last2=Kullander | editor-first2=S.O. | editor-last3=Ferraris | editor-first3=C.J. Jr. | title=Check List of the Freshwater Fishes of South and Central America | publisher=Edipucrs, Porto Alegre | isbn=978-85-7430-361-1 }}</ref> ''Mayaheros'' was only established for them in 2016.<ref name=Rican2016/>

===Species=== Traditionally, only two species have been recognized in this group: the northwest Mexican ''M. beani'', and the southeast Mexican and northern Central American ''M. urophthalmus''. The latter had a large number of subspecies that had been described in 1935–38 by Carl Leavitt Hubbs based on exact colour patterns (notably of their body stripes and tail base spots) and morphometrics. In a taxonomic review of 2003, those that had been described in 1936–38 by Hubbs and ''M. troschelii'' described in 1867 by Franz Steindachner were preliminarily elevated to species status, thus increasing the number of recognized species in the ''urophthalmus'' group to 11.<ref name=Rican2016/><ref name=Kullander2003/> A 2005 thesis proposed that all those, ''M. trispilus'' described in 1935 by Hubbs, and eight additional currently undescribed species (names were proposed, but the thesis does not comply with article 8 of the ICZN Code<ref>{{cite web| editor=Ride, W.D.L. | editor2=Cogger, H.G. | editor3=Dupuis, C. | editor4=Kraus, O. | editor5=Minelli, A. | editor6=Thompson, F.C. | editor7=Tubbs, P.K. | date=31 December 1999 | title=Article 8. What constitutes published work | url=https://code.iczn.org/criteria-of-publication/article-8-what-constitutes-published-work/?frame=1 | work=International Code of Zoological Nomenclature | edition=4 | access-date=29 December 2021 }}</ref>) should be recognized in the ''urophthalmus'' group based on colour patterns and morphometrics.<ref name=BarrientosMedina2005/> This would effectively limit "true" ''M. urophthalmus'' to the Lake Petén system in Guatemala.<ref name=Rican2016/><ref name=BarrientosMedina2005/> Other reviews of many of the populations in the ''urophthalmus'' group found that genetic differences are very small without clear geographic structure, that there likely has been widespread translocations of the cichlids by humans in the pre-Columbian era, and that a large amount of individual variations in pattern and morphometrics exists, mostly depending on environmental factors.<ref name=Harrison2014/><ref name=Miller2003>{{cite book| last=Miller | first=R.R. | year=2003 | title=Freshwater fishes of México | publisher=University of Chicago Press, Chicago | isbn=978-0-226-52604-1 }}</ref><ref>{{cite journal | last=Morgenstern | first=P. | year=2018 | title=Fishes collected by Emanuel Ritter von Friedrichsthal in Central America between 1838–1841 | journal=Vertebrate Zoology | volume=68 | issue=3 | pages=253–267 | doi=10.3897/vz.68.e31614 | s2cid=54074416 | doi-access=free }}</ref><ref name=BarrientosVillalobos2018>{{cite journal | last1=Barrientos-Villalobos | first1=J. | last2=Schmitter-Soto | first2=J.J. | last3=Espinosa de los Monteros | first3=A.J. | year=2018 | title=Several Subspecies or Phenotypic Plasticity? A Geometric Morphometric and Molecular Analysis of Variability of the Mayan Cichlid Mayaheros urophthalmus in the Yucatan | journal=Copeia | volume=106 | issue=2 | pages=268–278 | doi=10.1643/CI-17-657 | s2cid=90809586 | doi-access=free }}</ref> This points to a single variable species with a high degree of phenotypic plasticity rather than several different species in the ''urophthalmus'' group.<ref name=Harrison2014/><ref name=Miller2003/><ref name=BarrientosVillalobos2018/> As a consequence, all proposed species that were directly checked in those studies have been synonymized with ''M. urophthalmus'' by Catalog of Fishes, leaving only the unchecked ''M. aguadae'', ''M. amarus'', ''M. ericymba'' and ''M. trispilus'' as still recognized in this group.<ref name=CoF/> In the IUCN's evaluation of 2019, ''M. urophthalmus'' had been returned to its original single-species form without other recognized species in the group.<ref name=IUCNurophthalmus/> In contrast to the complicated taxonomic status of ''M. urophthalmus'', the single-species taxonomy of ''M. beani'' is agreed upon by all major authorities.<ref name=Rican2016/><ref name=IUCNbeani/><ref name=CoF/><ref name=FishBase>{{FishBase genus| genus=Mayaheros | year=2021 | month=December }}</ref>

Unlike Catalog of Fishes and the IUCN, FishBase continues to recognize 13 species in this genus, including ''M. urophthalmus'' itself and 11 others in its group (indented in list):<ref name=FishBase/>

* ''Mayaheros beani'' (D. S. Jordan, 1889) (Green guapote) * ''Mayaheros urophthalmus'' (Günther, 1862) (Mayan cichlid) ** ''Mayaheros aguadae'' (C. L. Hubbs, 1936) ** ''Mayaheros alborus'' (C. L. Hubbs, 1936) ** ''Mayaheros amarus'' (C. L. Hubbs, 1936) ** ''Mayaheros cienagae'' (C. L. Hubbs, 1936) ** ''Mayaheros conchitae'' (C. L. Hubbs, 1936) ** ''Mayaheros ericymba'' (C. L. Hubbs, 1938) ** ''Mayaheros mayorum'' (C. L. Hubbs, 1936) ** ''Mayaheros stenozonus'' (C. L. Hubbs, 1936) ** ''Mayaheros trispilus'' (C. L. Hubbs, 1935) ** ''Mayaheros troschelii'' (Steindachner, 1867) ** ''Mayaheros zebra'' (C. L. Hubbs, 1936)

The species ''Amphilophus istlanus'' (still included in ''Cichlasoma'' by some authorities) is a special case, as nDNA places it in ''Amphilophus'', but mDNA places it in ''Mayaheros''. ''M. istlanus'' likely is the result of hybrid speciation involving ''M. beani'' and ''A. trimaculatus''.<ref name=Rican2016/>

==References== {{Reflist}}

Category:Heroini