Orestias (Cyprinodontidae)
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''Orestias'' is a genus of
pupfish Pupfish are a group of small killifish belonging to ten genera of the family Cyprinodontidae of ray-finned fish. Pupfish are especially noted for being found in extreme and isolated situations. They are primarily found in North America, South Am ...
. Older systematics
classified Classified may refer to: General *Classified information, material that a government body deems to be sensitive *Classified advertising or "classifieds" Music *Classified (rapper) (born 1977), Canadian rapper * The Classified, a 1980s American ro ...
them into the own family Orestiidae. They are found in lakes, rivers and springs in the
Andean The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long and wide (widest between 18°S ...
highlands of South America, and several species are considered
threatened A threatened species is any species (including animals, plants and fungi) which is vulnerable to extinction in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensatio ...
. They are egg-laying fish that feed on small animals and plant matter. The largest species can reach a total length of , but most remain far smaller. Their most characteristic feature is the absence of the
ventral fin Pelvic fins or ventral fins are paired fins located on the ventral (belly) surface of fish, and are the lower of the only two sets of paired fins (the other being the laterally positioned pectoral fins). The pelvic fins are homologous to the hind ...
, although this is shared by a few other pupfish. Despite their moderate to small size, they are important to local fisheries and a few species are farmed. Several species are locally and colloquially known as ''carache''. The name of the genus is a reference to
Orestes In Greek mythology, Orestes or Orestis (; ) was the son of Agamemnon and Clytemnestra, and the brother of Electra and Iphigenia. He was also known by the patronymic Agamemnonides (), meaning "son of Agamemnon." He is the subject of several ...
, a
Greek mythological Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of ancient Greek folklore, today absorbed alongside Roman mythology into the broader designation of classical mythology. These stories concern the ancient ...
character who
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
described as the "nymph of the mountains".


Range and habitat

''Orestias'' is restricted to freshwater habitats at an altitude of in the
Andes The Andes ( ), Andes Mountains or Andean Mountain Range (; ) are the List of longest mountain chains on Earth, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range ...
of central and southern
Peru Peru, officially the Republic of Peru, is a country in western South America. It is bordered in the north by Ecuador and Colombia, in the east by Brazil, in the southeast by Bolivia, in the south by Chile, and in the south and west by the Pac ...
, western
Bolivia Bolivia, officially the Plurinational State of Bolivia, is a landlocked country located in central South America. The country features diverse geography, including vast Amazonian plains, tropical lowlands, mountains, the Gran Chaco Province, w ...
, and northeastern
Chile Chile, officially the Republic of Chile, is a country in western South America. It is the southernmost country in the world and the closest to Antarctica, stretching along a narrow strip of land between the Andes, Andes Mountains and the Paci ...
.Vila, Morales, Scott, Poulin, Veliz, Harrod and Mendez (2013). Phylogenetic and phylogeographic analysis of the genus Orestias (Teleostei: Cyprinodontidae) in the southern Chilean Altiplano: the relevance of ancient and recent divergence processes in speciation. Journal of Fish Biology 82, 927–943. Although freshwater habitats at low and middle altitudes in the Andes are relatively rich in fish, few are found in the highest, with ''Orestias'' and the catfish ''
Astroblepus ''Astroblepus'' is a genus of fish in the family Astroblepidae found in South America and Panama. This genus is the only member of its family. These catfishes are primarily found in torrential streams in the Andean area. ''Astroblepus pholeter'' ...
'' and ''
Trichomycterus ''Trichomycterus'' is a genus of fish in the family Trichomycteridae, the largest genus of its family with over 170 species currently described. This genus is native to freshwater habitats in Central and South America. These fish are generally s ...
'' being the primary—in many places only—native genera. Of the 45 recognized species of ''Orestias'', the majority are found in
Lake Titicaca Lake Titicaca (; ; ) is a large freshwater lake in the Andes mountains on the border of Bolivia and Peru. It is often called the highest navigable lake in the world. Titicaca is the largest lake in South America, both in terms of the volume of ...
(including 23
endemics Endemism is the state of a species being found only in a single defined geographic location, such as an island, state, nation, country or other defined zone; organisms that are indigenous to a place are not endemic to it if they are also foun ...
to its basin), with the remaining from other
Altiplano The Altiplano (Spanish language, Spanish for "high plain"), Collao (Quechuan languages, Quechua and Aymara language, Aymara: Qullaw, meaning "place of the Qulla people, Qulla") or Andean Plateau, in west-central South America, is the most extens ...
lakes, rivers or springs. In addition to differences in feeding, the many species in Lake Titicaca segregate by habitat preference, with some living among totora reeds, some among submerged
macrophyte Aquatic plants, also referred to as hydrophytes, are vascular plants and non-vascular plants that have adapted to live in aquatic environments ( saltwater or freshwater). In lakes, rivers and wetlands, aquatic vegetations provide cover for aquat ...
s, some at the bottom in water too deep for macrophytes (deeper than ), and some pelagically in the open water. ''Orestias'' species found elsewhere than Lake Titicaca often have very small ranges.


Behavior

Relatively little is known about the behavior of ''Orestias''. They feed on small crustaceans (such as
amphipods Amphipoda () is an order (biology), order of malacostracan crustaceans with no carapace and generally with laterally compressed bodies. Amphipods () range in size from and are mostly detritivores or scavengers. There are more than 10,700 amphip ...
and
cladocera The Diplostraca or Cladocera, commonly known as water fleas, is a superorder (biology), superorder of small, mostly freshwater crustaceans, most of which feed on microscopic chunks of organic matter, though some forms are predatory. Over 1000 sp ...
), aquatic insects and larvae (such as
chironomid Chironomidae , commonly known as non-biting midges or chironomids , are a family of Nematoceran flies with a global distribution. They are closely related to the families Ceratopogonidae, Simuliidae, and Thaumaleidae. Although many chironomid s ...
s), snails, small fish, fish eggs, algae,
macrophyte Aquatic plants, also referred to as hydrophytes, are vascular plants and non-vascular plants that have adapted to live in aquatic environments ( saltwater or freshwater). In lakes, rivers and wetlands, aquatic vegetations provide cover for aquat ...
seeds and plant
detritus In biology, detritus ( or ) is organic matter made up of the decomposition, decomposing remains of organisms and plants, and also of feces. Detritus usually hosts communities of microorganisms that colonize and decomposition, decompose (Reminera ...
. There are significant variations depending on exact species, especially in Lake Titicaca where there is a degree of
niche differentiation In ecology, a niche is the match of a species to a specific environmental condition. Three variants of ecological niche are described by It describes how an organism or population responds to the distribution of resources and competitors (for e ...
, including some that mainly feed on
zooplankton Zooplankton are the heterotrophic component of the planktonic community (the " zoo-" prefix comes from ), having to consume other organisms to thrive. Plankton are aquatic organisms that are unable to swim effectively against currents. Consequent ...
ic organisms, two (''O. albus'' and ''O. cuvieri'') that are particularly willing to take other fish, primarily smaller ''Orestias'' (fish eggs are regularly consumed by a wider range of species), several that mainly feed on small bottom- or plant-living organisms, and some that are generalists that will feed on a wide range of things, sometimes varying depending on season. Females tend to grow larger than males. Although generally rather dull-coloured, breeding males may become partially yellow or orange. The eggs, up to a few hundred, are placed in shallow water among vegetation. ''Orestias'' appear to reach maturity when around one year old.


Conservation and relationship to humans

Many species of the genus became rare in recent decades due to predation by, and competition with introduced fish species (especially
rainbow trout The rainbow trout (''Oncorhynchus mykiss'') is a species of trout native to cold-water tributary, tributaries of the Pacific Ocean in North America and Asia. The steelhead (sometimes called steelhead trout) is an Fish migration#Classification, ...
and Argentinian silverside), pollution, overfishing and other human activities in their habitats. In the 1960s, an expedition to Lake Titicaca led by
Jacques Cousteau Jacques-Yves Cousteau, (, also , ; 11 June 191025 June 1997) was a French naval officer, oceanographer, filmmaker and author. He co-invented the first successful open-circuit self-contained underwater breathing apparatus (SCUBA), called the ...
reported seeing many dead ''Orestias'' and when studied they were found to be infected by a disease introduced with the trout. The two largest species in the genus, the Titicaca orestias (''O. cuvieri'') at up to in
standard length Fish measurement is the measuring of individual fish and various parts of fish anatomy, their anatomies, for data used in many areas of ichthyology, including Taxonomy (biology), taxonomy and fishery biology. Overall length Standard length (SL) is ...
and in total length, and ''O. pentlandii'' up to and respectively, have fared the worst. The Titicaca orestias was last seen in 1939 and is almost certainly extinct. ''O. pentlandii'' is from the same lake and it may have disappeared in its native form. It was still regarded as "only" vulnerable by the
IUCN The International Union for Conservation of Nature (IUCN) is an international organization working in the field of nature conservation and sustainable use of natural resources. Founded in 1948, IUCN has become the global authority on the status ...
in 2009, but in 2014 the species was regarded as
critically endangered An IUCN Red List critically endangered (CR or sometimes CE) species is one that has been categorized by the International Union for Conservation of Nature as facing an extremely high risk of extinction in the wild. As of December 2023, of t ...
in a review of Lake Titicaca fish. In the 1990s, some were introduced to a small highland lake in Peru in an attempt of safeguarding the species, but over time they have changed (possibly due to
genetic drift Genetic drift, also known as random genetic drift, allelic drift or the Wright effect, is the change in the Allele frequency, frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene va ...
,
inbreeding depression Inbreeding depression is the reduced biological fitness caused by loss of genetic diversity as a consequence of inbreeding, the breeding of individuals closely related genetically. This loss of genetic diversity results from small population siz ...
, or their new and different habitat) and now appear very different from the original ''O. pentlandii'' of Lake Titicaca. The remaining ''Orestias'' of Lake Titicaca are also threatened to various degrees, as are the three species in
Lake Junin Lake Junin (; Spanish language, Spanish ''Lago Junín'', named after the nearby town of Junín, Peru, Junin) or Chinchaycocha (possibly from Quechua language, Quechua ''chincha, chinchay'' north, northern, ''chinchay'' ocelot, ''qucha'' lake, lag ...
and its vicinity, which in turn may threaten the Lake Junin giant frog that feeds on the fish and invertebrates. Many ''Orestias'' found elsewhere are vulnerable because of their highly restricted ranges,Parenti, Lynne R. (1984): A taxonomic revision of the Andean Killifish Genus ''Orestias'' (Cyprinodontiformes, Cyprinodontidae). ''Bulletin of the American Museum of Natural History'' 178: 107-214
PDF fulltext
/ref> like ''O. ascotanensis'' where the entire habitat covers of
Salar de Ascotán Salar de Ascotán, also known as Salar de Cebollar, is a salt flat in northern Chile. Its drainage basin is and is shared with Bolivia. The basin is bordered on the north by the Salar de Carcote basin, on the east by small endorheic basins, inc ...
and ''O. chungarensis'' found only in
Lake Chungará A lake is often a naturally occurring, relatively large and fixed body of water on or near the Earth's surface. It is localized in a depression (geology), basin or interconnected basins surrounded by dry land. Lakes lie completely on land an ...
. Despite their moderate to small size, some species are important to local fisheries, but they are overfished, there are few regulations and limited enforcement of the rules that do exist. In Lake Titicaca, the moderately sized ''O. agassizii'' and ''O. luteus'' ( complexes) represent more than 90% of the total catches of native fish species, but others like the small open-water ispi (''O. forgeti'' and ''O. ispi'') are also frequently caught. The natives have now been greatly surpassed by fisheries for the non-native introduced trout and Argentinian silverside. Because of pollution, studies have revealed levels of metals in ''Orestias'' of Lake Titicaca that exceed the internationally recommended safety thresholds for human consumption. ''O. agassizii'' and ''O. luteus'' are also farmed and spawned in captivity; both to supply the food market and for release to support their vulnerable wild populations.


Species and taxonomy

There are currently 45 recognized species in this genus, divided depending on their relationships into four
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
es. The genus is estimated to be 5 million years old, with a significant diversification occurring within the last one million years. Lüssen (2003) researched the
phylogeny A phylogenetic tree or phylogeny is a graphical representation which shows the evolutionary history between a set of species or Taxon, taxa during a specific time.Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, M ...
of several species including
mtDNA Mitochondrial DNA (mtDNA and mDNA) is the DNA located in the mitochondria organelles in a eukaryotic cell that converts chemical energy from food into adenosine triphosphate (ATP). Mitochondrial DNA is a small portion of the DNA contained in ...
sequence In mathematics, a sequence is an enumerated collection of objects in which repetitions are allowed and order matters. Like a set, it contains members (also called ''elements'', or ''terms''). The number of elements (possibly infinite) is cal ...
data; because hybridization is known to occur (Villwock 1964, Parenti 1984), mtDNA data cannot be relied upon on its own to resolve the evolution of this genus. For example, ''O. agassizii'' and ''O. luteus'' reach a similar maximum
standard length Fish measurement is the measuring of individual fish and various parts of fish anatomy, their anatomies, for data used in many areas of ichthyology, including Taxonomy (biology), taxonomy and fishery biology. Overall length Standard length (SL) is ...
of , but otherwise they are quite different. Despite this, hybrids between them are known.


Agassizii species complex

Some species restricted to the Lake Titicaca basin (which includes the inflowing rivers and connected smaller lakes like Arapa, Lagunillas and Saracocha), and some restricted to other lakes, rivers and springs elsewhere in the Andean highlands. ''O. agassizii'' (often spelled ''agassii'' instead) is the only member of the genus found both in the Lake Titicaca basin and elsewhere. * '' Orestias agassizii''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1846
(Carache Negro) * '' Orestias albus''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1846
(Carache Amarillo) * ''
Orestias ascotanensis ''Orestias ascotanensis'' is a species of freshwater ray-finned fish within the family Cyprinodontidae, endemic to 12 springs in the Ascotán Salt Flat in Chile Chile, officially the Republic of Chile, is a country in western South Americ ...
'' Parenti, 1984 * '' Orestias chungarensis'' Vila & M. Pinto, 1987 * '' Orestias ctenolepis'' Parenti, 1984 * '' Orestias elegans''
Garman Garman is a surname or first name. Notable people with the name include: Sports * Ann Garman, All-American Girls Professional Baseball League player * Judi Garman (born 1954), American softball coach * Mike Garman (born 1949), American baseball pla ...
, 1895
* '' Orestias empyraeus'' W. R. Allen, 1942 * '' Orestias frontosus''
Cope A cope ( ("rain coat") or ("cape")) is a liturgical long mantle or cloak, open at the front and fastened at the breast with a band or clasp. It may be of any liturgical colour. A cope may be worn by any rank of the Catholic or Anglican clerg ...
, 1876
* ''
Orestias gloriae Orestias (), later refounded by Hadrian as Adrianople (Greek: Άδριανούπολις), was an ancient Greek settlement next to the Evros river in Thrace, near or at the site of present-day Edirne, and close to the current border between Turk ...
'' Vila, S. Scott,
Méndez Méndez is a common Spanish language, Spanish surname to all of Germanic origin, originally a patronymic, meaning ''Son of Mendo'', ''Menendo'', or ''Mem_(given_name), Mem''. A longer form sharing the same root is Melendez, Menéndez, while the Por ...
,
Valenzuela Valenzuela may refer to: Places * Valenzuela, Paraguay * Valenzuela, Metro Manila, Philippines * Valenzuela, Spain * Valenzuela de Calatrava, Spain * Valenzuela, Louisiana Other uses * Valenzuela (surname), including a list of people with the ...
, Iturra & Poulin, 2012
Vila, I., Scott, S., Mendez, M.A., Valenzuela, F., Iturra, P. & Poulin, E. (2012)
Orestias gloriae, a new species of cyprinodontid fish from saltpan spring of the southern high Andes (Teleostei: Cyprinodontidae).
''Ichthyological Exploration of Freshwaters, 22 (4) 011 345-353.''
* '' Orestias gymnotus'' Parenti, 1984 * '' Orestias hardini'' Parenti, 1984 * '' Orestias jussiei''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1846
(Carache Amarillo) * '' Orestias lastarriae'' Philippi , 1876 * ''
Orestias laucaensis Orestias (), later refounded by Hadrian as Adrianople (Greek: Άδριανούπολις), was an ancient Greek settlement next to the Evros river in Thrace, near or at the site of present-day Edirne, and close to the current border between Turk ...
'' Arratia, 1982 * '' Orestias luteus''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1846
(Carache Amarillo) * ''
Orestias multiporis Orestias (), later refounded by Hadrian as Adrianople (Greek: Άδριανούπολις), was an ancient Greek settlement next to the Evros river in Thrace, near or at the site of present-day Edirne, and close to the current border between Turk ...
'' Parenti, 1984 * '' Orestias mundus'' Parenti, 1984 * '' Orestias olivaceus''
Garman Garman is a surname or first name. Notable people with the name include: Sports * Ann Garman, All-American Girls Professional Baseball League player * Judi Garman (born 1954), American softball coach * Mike Garman (born 1949), American baseball pla ...
, 1895
* '' Orestias parinacotensis'' Arratia, 1982 * '' Orestias piacotensis'' Vila, 2006 * ''
Orestias polonorum Orestias (), later refounded by Hadrian as Adrianople (Greek: Άδριανούπολις), was an ancient Greek settlement next to the Evros river in Thrace, near or at the site of present-day Edirne, and close to the current border between Turk ...
'' Tchernavin, 1944 * '' Orestias puni'' Tchernavin, 1944 — likely a
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
of ''O. jussiei'' * '' Orestias richersoni'' Parenti, 1984 * ''
Orestias silustani Orestias (), later refounded by Hadrian as Adrianople (Greek: Άδριανούπολις), was an ancient Greek settlement next to the Evros river in Thrace, near or at the site of present-day Edirne, and close to the current border between Turk ...
'' W. R. Allen, 1942 * '' Orestias tschudii'' Castelnau, 1855 * '' Orestias ututo'' Parenti, 1984


Cuvieri species complex

All restricted to the Lake Titicaca basin. * '' Orestias cuvieri''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1846
(Titicaca Orestias, Amanto; likely
extinct Extinction is the termination of an organism by the death of its Endling, last member. A taxon may become Functional extinction, functionally extinct before the death of its last member if it loses the capacity to Reproduction, reproduce and ...
(mid-20th century)) * '' Orestias forgeti'' Lauzanne, 1981 (Ispi) * '' Orestias ispi'' Lauzanne, 1981 (Ispi) * '' Orestias pentlandii''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1846
(Boga)


Gilsoni species complex

All restricted to the Lake Titicaca basin. * '' Orestias gilsoni'' Tchernavin, 1944 * '' Orestias imarpe'' Parenti, 1984 * '' Orestias minimus'' Tchernavin, 1944 * '' Orestias minutus'' Tchernavin, 1944 — likely a
synonym A synonym is a word, morpheme, or phrase that means precisely or nearly the same as another word, morpheme, or phrase in a given language. For example, in the English language, the words ''begin'', ''start'', ''commence'', and ''initiate'' are a ...
of ''O. minimus'' * '' Orestias mooni'' Tchernavin, 1944 * '' Orestias robustus'' Parenti, 1984 * '' Orestias taquiri'' Tchernavin, 1944 * '' Orestias tchernavini'' Lauzanne, 1981 * '' Orestias tomcooni'' Parenti, 1984 * '' Orestias uruni'' Tchernavin, 1944


Mulleri species complex

All restricted to the Lake Titicaca basin. * ''
Orestias crawfordi Orestias (), later refounded by Hadrian as Adrianople (Greek: Άδριανούπολις), was an ancient Greek settlement next to the Evros river in Thrace, near or at the site of present-day Edirne, and close to the current border between Turk ...
'' Tchernavin, 1944 * '' Orestias gracilis'' Parenti, 1984 * '' Orestias incae''
Garman Garman is a surname or first name. Notable people with the name include: Sports * Ann Garman, All-American Girls Professional Baseball League player * Judi Garman (born 1954), American softball coach * Mike Garman (born 1949), American baseball pla ...
, 1895
* '' Orestias mulleri''
Valenciennes Valenciennes (, also , , ; ; or ; ) is a communes of France, commune in the Nord (French department), Nord Departments of France, department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced ...
, 1846
* '' Orestias tutini'' Tchernavin, 1944


References

* Lüssen, Arne (2003): Zur Systematik, Phylogenie und Biogeographie chilenischer Arten der Gattung ''Orestias'' VALENCIENNES, 1839. Doctoral thesis, University of Hamburg, Germany.
n German N, or n, is the fourteenth letter of the Latin alphabet, used in the modern English alphabet, the alphabets of other western European languages, and others worldwide. Its name in English is ''en'' (pronounced ), plural ''ens''. History ...
br>PDF fulltext
* Villwock, W. (1964): Vermeintliche Artbastarde in der Gattung ''Orestias'' (Pisces, Cyprinodontidae). ''Mitteilungen aus dem Hamburgischen Zoologischen Museum und Institut'', Kosswig-Festschrift: 285-291. rticle in German* Villwock, W. (1972): Gefahren für die endemische Fischfauna durch Einbürgerungsversuche und Akklimatisation von Fremdfischen am Beispiel des Titicacas-Sees (Peru/Bolivien) und des Lanao-Sees (Mindanao/Philippinen). ''Verhandlungen des Internationalen Vereins für Limnologie'' 18: 1227-1234. rticle in German {{Taxonbar, from=Q490401 Cyprinodontidae * Taxa named by Achille Valenciennes Freshwater fish genera