Mayaheros
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''Mayaheros'' is a
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nom ...
of
cichlid Cichlids are fish from the family Cichlidae in the order Cichliformes. Cichlids were traditionally classed in a suborder, the Labroidei, along with the wrasses ( Labridae), in the order Perciformes, but molecular studies have contradicted th ...
fish that is native to
Mexico Mexico (Spanish language, Spanish: México), officially the United Mexican States, is a List of sovereign states, country in the southern portion of North America. It is borders of Mexico, bordered to the north by the United States; to the so ...
and northern
Central America Central America ( es, América Central or ) is a subregion of the Americas. Its boundaries are defined as bordering the United States to the north, Colombia to the south, the Caribbean Sea to the east, and the Pacific Ocean to the west. ...
. This genus has a
disjunct distribution In biology, a taxon with a disjunct distribution is one that has two or more groups that are related but considerably separated from each other geographically. The causes are varied and might demonstrate either the expansion or contraction of a s ...
, with the ''M. urophthalmus'' group being found in the Atlantic drainages of southeastern Mexico (southern
Veracruz Veracruz (), formally Veracruz de Ignacio de la Llave (), officially the Free and Sovereign State of Veracruz de Ignacio de la Llave ( es, Estado Libre y Soberano de Veracruz de Ignacio de la Llave), is one of the 31 states which, along with Me ...
east to the
Yucatán Peninsula The Yucatán Peninsula (, also , ; es, Península de Yucatán ) is a large peninsula in southeastern Mexico and adjacent portions of Belize and Guatemala. The peninsula extends towards the northeast, separating the Gulf of Mexico to the north ...
and southwards),
Belize Belize (; bzj, Bileez) is a Caribbean and Central American country on the northeastern coast of Central America. It is bordered by Mexico to the north, the Caribbean Sea to the east, and Guatemala to the west and south. It also shares a wa ...
, eastern
Guatemala Guatemala ( ; ), officially the Republic of Guatemala ( es, República de Guatemala, links=no), is a country in Central America. It is bordered to the north and west by Mexico; to the northeast by Belize and the Caribbean; to the east by Hon ...
, northern
Honduras Honduras, officially the Republic of Honduras, is a country in Central America. The republic of Honduras is bordered to the west by Guatemala, to the southwest by El Salvador, to the southeast by Nicaragua, to the south by the Pacific Oce ...
and northeastern
Nicaragua Nicaragua (; ), officially the Republic of Nicaragua (), is the largest country in Central America, bordered by Honduras to the north, the Caribbean to the east, Costa Rica to the south, and the Pacific Ocean to the west. Managua is the coun ...
, while ''M. beani'' is the northernmost cichlid in the Pacific drainage, ranging from
Jalisco Jalisco (, , ; Nahuatl: Xalixco), officially the Free and Sovereign State of Jalisco ( es, Estado Libre y Soberano de Jalisco ; Nahuatl: Tlahtohcayotl Xalixco), is one of the 31 states which, along with Mexico City, comprise the 32 Federal ...
to
Sonora Sonora (), officially Estado Libre y Soberano de Sonora ( en, Free and Sovereign State of Sonora), is one of the 31 states which, along with Mexico City, comprise the Federal Entities of Mexico. The state is divided into 72 municipalities; the ...
in northwestern Mexico. Both inhabit a wide range of habitats such as freshwater rivers, streams, lakes, ponds, pools and marshes, as well as
brackish Brackish water, sometimes termed brack water, is water occurring in a natural environment that has more salinity than freshwater, but not as much as seawater. It may result from mixing seawater (salt water) and fresh water together, as in estua ...
waters like
estuaries An estuary is a partially enclosed coastal body of brackish water with one or more rivers or streams flowing into it, and with a free connection to the open sea. Estuaries form a transition zone between river environments and maritime environme ...
, lagoons and
mangrove A mangrove is a shrub or tree that grows in coastal saline or brackish water. The term is also used for tropical coastal vegetation consisting of such species. Mangroves are taxonomically diverse, as a result of convergent evolution in severa ...
, with members of the ''M. urophthalmus'' group sometimes even occurring in caves or coastal marine waters. They are medium-sized to large cichlids that are
omnivorous An omnivore () is an animal that has the ability to eat and survive on both plant and animal matter. Obtaining energy and nutrients from plant and animal matter, omnivores digest carbohydrates, protein, fat, and fiber, and metabolize the nut ...
, feeding mostly on small animals, but also plants and
detritus In biology, detritus () is dead particulate organic material, as distinguished from dissolved organic material. Detritus typically includes the bodies or fragments of bodies of dead organisms, and fecal material. Detritus typically hosts comm ...
. ''Mayaheros'' cichlids are sometimes kept in aquariums and commonly caught for food. Broadstock of the ''M. urophthalmus'' group that is better suited for
farming Agriculture or farming is the practice of cultivating plants and livestock. Agriculture was the key development in the rise of sedentary human civilization, whereby farming of domesticated species created food surpluses that enabled peopl ...
is being developed by Mexican authorities; ''M. beani'' also has potential in farming, but preliminary tests have been complicated by its high levels of aggression. Both ''M. beani'' and ''M. urophthalmus'' are overall common and regarded as
least concern A least-concern species is a species that has been categorized by the International Union for Conservation of Nature (IUCN) as evaluated as not being a focus of species conservation because the specific species is still plentiful in the wild. ...
by the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
, although certain local populations are under pressure from habitat loss, pollution,
overfishing Overfishing is the removal of a species of fish (i.e. fishing) from a body of water at a rate greater than that the species can replenish its population naturally (i.e. the overexploitation of the fishery's existing fish stock), resulting in t ...
and
introduced species An introduced species, alien species, exotic species, adventive species, immigrant species, foreign species, non-indigenous species, or non-native species is a species living outside its native distributional range, but which has arrived there ...
. A captive breeding program exists for a few populations of ''M. beani''. 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 A cenote ( or ; ) is a natural pit, or sinkhole, resulting from the collapse of limestone bedrock that exposes groundwater. The regional term is specifically associated with the Yucatán Peninsula of Mexico, where cenotes were commonly used ...
, cave, island or bay, and some are seriously threatened. The most threatened –if not already
extinct Extinction is the termination of a kind of organism or of a group of kinds (taxon), usually a species. The moment of extinction is generally considered to be the death of the last individual of the species, although the capacity to breed and ...
– 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), 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. If recognized as valid species or
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
of ''M. urophthalmus'', these two are the only likely extinct cichlids of the Americas. Conversely, ''M. urophthalmus'' has been introduced outside its native range in
Oaxaca Oaxaca ( , also , , from nci, Huāxyacac ), officially the Free and Sovereign State of Oaxaca ( es, Estado Libre y Soberano de Oaxaca), is one of the 32 states that compose the Federative Entities of Mexico. It is divided into 570 municipaliti ...
(Mexico),
Florida Florida is a state located in the Southeastern region of the United States. Florida is bordered to the west by the Gulf of Mexico, to the northwest by Alabama, to the north by Georgia, to the east by the Bahamas and Atlantic Ocean, and ...
(United States), the Philippines, Thailand and Singapore. 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.


Taxonomy

Both the species ''beani'' and ''urophthalmus'' were first
scientifically described A species description is a formal description of a newly discovered species, usually in the form of a scientific paper. Its purpose is to give a clear description of a new species of organism and explain how it differs from species that have be ...
as members of the genus '' Heros'' and then moved to ''
Cichlasoma ''Cichlasoma'' is a genus of freshwater fish in the cichlid family. The genus was previously very large (a wastebasket taxon), including cichlids from North America, including Central America, and South America. Reclassification and subsequent ...
'' despite clearly belonging elsewhere. ''Mayaheros'' was only established for them in 2016.


Species

Traditionally, only two species have been recognized in this group: the northwest Mexican ''beani'', and the southeast Mexican and northern Central American ''urophthalmus''. The latter had a large number of
subspecies In biological classification, subspecies is a rank below species, used for populations that live in different areas and vary in size, shape, or other physical characteristics ( morphology), but that can successfully interbreed. Not all specie ...
that had been described in 1935–38 by
Carl Leavitt Hubbs Carl Leavitt Hubbs (October 19, 1894 – June 30, 1979) was an American ichthyologist. Biography Youth He was born in Williams, Arizona. He was the son of Charles Leavitt and Elizabeth (née Goss) Hubbs. His father had a wide variety of jobs (f ...
based on exact colour patterns (notably of their body stripes and tail base spot) and
morphometrics Morphometrics (from Greek μορϕή ''morphe'', "shape, form", and -μετρία ''metria'', "measurement") or morphometry refers to the quantitative analysis of ''form'', a concept that encompasses size and shape. Morphometric analyses are co ...
. In a
taxonomic Taxonomy is the practice and science of categorization or classification. A taxonomy (or taxonomical classification) is a scheme of classification, especially a hierarchical classification, in which things are organized into groups or types. ...
review of 2003, those that had been described in 1936–38 by Hubbs and ''troschelii'' described in 1867 by
Franz Steindachner Franz Steindachner (11 November 1834 in Vienna – 10 December 1919 in Vienna) was an Austrian Zoology, zoologist, ichthyologist, and herpetologist. He published over 200 papers on fishes and over 50 papers on reptiles and amphibians. Steindachne ...
were preliminarily elevated to species status, thus increasing the number of recognized species in the ''urophthalmus'' group to eleven. In a 2005 thesis, it was proposed that all those, ''trispilus'' described in 1935 by Hubbs, and eight additional currently
undescribed species In taxonomy, an undescribed taxon is a taxon (for example, a species) that has been discovered, but not yet formally described and named. The various Nomenclature Codes specify the requirements for a new taxon to be validly described and named. U ...
(names were proposed, but the thesis does not comply with article 8 of the
ICZN Code The International Code of Zoological Nomenclature (ICZN) is a widely accepted convention in zoology that rules the formal scientific naming of organisms treated as animals. It is also informally known as the ICZN Code, for its publisher, the ...
) should be recognized in the ''urophthalmus'' group based on colour patterns and morphometrics. This would effectively limit "true" ''M. urophthalmus'' to the Lake Petén system in Guatemala. 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 In the history of the Americas, the pre-Columbian era spans from the Migration to the New World, original settlement of North and South America in the Upper Paleolithic period through European colonization of the Americas, European colonization, w ...
, and that there is a large amount of individual variations in pattern and morphometrics, mostly depending on environmental factors. This points to a single variable species with a high degree of
phenotypic plasticity Phenotypic plasticity refers to some of the changes in an organism's behavior, morphology and physiology in response to a unique environment. Fundamental to the way in which organisms cope with environmental variation, phenotypic plasticity encompa ...
rather than several different species in the ''urophthalmus'' group. As a consequence, all proposed species that were directly checked in those studies have been synonymized with ''M. urophthalmus'' by
Catalog of Fishes Catalog of Fishes is a comprehensive on-line database and reference work on the scientific names of fish species and genera. It is global in its scope and is hosted by the California Academy of Sciences. It has been compiled and is continuously up ...
, leaving only the unchecked ''M. aguadae'', ''M. amarus'', ''M. ericymba'' and ''M. trispilus'' as still recognized in this group. In the
IUCN The International Union for Conservation of Nature (IUCN; officially International Union for Conservation of Nature and Natural Resources) is an international organization working in the field of nature conservation and sustainable use of natu ...
's evaluation of 2019, ''M. urophthalmus'' had been returned to its original single-species form without other recognized species in the group. In contrast to the complicated taxonomic status of ''M. urophthalmus'', the single-species taxonomy of ''M. beani'' is agreed upon by all major authorities. Unlike Catalog of Fishes and the IUCN, FishBase continues to recognize thirteen species in this genus, including ''M. urophthalmus'' itself and eleven others in its group (indented in list): * '' 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 ''Amphilophus'' is a genus of cichlid fishes from Central America, ranging from southern Mexico to Panama. The genus currently contains 23 species, including several that are well-known from the aquarium trade. However, studies led by Oldřich ...
istlanus'' (still included in ''Cichlasoma'' by some authorities) is a special case, as
nDNA Nuclear DNA (nDNA), or nuclear deoxyribonucleic acid, is the DNA contained within each cell nucleus of a eukaryotic organism. It encodes for the majority of the genome in eukaryotes, with mitochondrial DNA and plastid DNA coding for the rest. I ...
places it in ''Amphilophus'', but mDNA places it in ''Mayaheros''. It is likely that ''istlanus'' is the result of
hybrid speciation Hybrid speciation is a form of speciation where hybridization between two different species leads to a new species, reproductively isolated from the parent species. Previously, reproductive isolation between two species and their parents was th ...
involving ''M. beani'' and '' A. trimaculatus''.


References

Heroini {{Cichlidae-stub