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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 them into the own family Orestiidae. They are found in lakes, rivers and springs in the
Andean The Andes, Andes Mountains or Andean Mountains (; ) are the List of mountain ranges#Mountain ranges by length, longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range i ...
highlands of South America, and several species are considered
threatened Threatened species are any species (including animals, plants and fungi) which are vulnerable to endangerment in the near future. Species that are threatened are sometimes characterised by the population dynamics measure of ''critical depensat ...
. 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 surface of fish. The paired pelvic fins are homologous to the hindlimbs of tetrapods. Structure and function Structure In actinopterygians, the pelvic fin consists of two ...
, 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 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 people t ...
. 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 (; grc-gre, Ὀρέστης ) was the son of Clytemnestra and Agamemnon, and the brother of Electra. He is the subject of several Ancient Greek plays and of various myths connected with his madness and ...
, a
Greek mythological A major branch of classical mythology, Greek mythology is the body of myths originally told by the ancient Greeks, and a genre of Ancient Greek folklore. These stories concern the origin and nature of the world, the lives and activities of d ...
character who
Valenciennes Valenciennes (, also , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
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 Mountains (; ) are the longest continental mountain range in the world, forming a continuous highland along the western edge of South America. The range is long, wide (widest between 18°S – 20°S ...
of central and southern
Peru , image_flag = Flag of Peru.svg , image_coat = Escudo nacional del Perú.svg , other_symbol = Great Seal of the State , other_symbol_type = Seal (emblem), National seal , national_motto = "Fi ...
, western
Bolivia , image_flag = Bandera de Bolivia (Estado).svg , flag_alt = Horizontal tricolor (red, yellow, and green from top to bottom) with the coat of arms of Bolivia in the center , flag_alt2 = 7 × 7 square p ...
, and northeastern
Chile Chile, officially the Republic of Chile, is a country in the western part of South America. It is the southernmost country in the world, and the closest to Antarctica, occupying a long and narrow strip of land between the Andes to the east a ...
.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 sm ...
'' 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 (; es, Lago Titicaca ; qu, Titiqaqa Qucha) 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. By volume of water and by surface area, ...
(including 23
endemics Endemism is the state of a species being found 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 found else ...
to its basin), with the remaining from other
Altiplano The Altiplano (Spanish for "high plain"), Collao (Quechua and Aymara: Qullaw, meaning "place of the Qulla") or Andean Plateau, in west-central South America, is the most extensive high plateau on Earth outside Tibet. The plateau is located at the ...
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 macrophytes, 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 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 9,900 amphipod species so far descri ...
and
cladocera The Diplostraca or Cladocera, commonly known as water fleas, are a superorder of small crustaceans that feed on microscopic chunks of organic matter (excluding some predatory forms). Over 1000 species have been recognised so far, with many more ...
), aquatic insects and larvae (such as
chironomid The Chironomidae (informally known as chironomids, nonbiting midges, or lake flies) comprise a family of nematoceran flies with a global distribution. They are closely related to the Ceratopogonidae, Simuliidae, and Thaumaleidae. Many specie ...
s), snails, small fish, fish eggs, algae, macrophyte seeds and plant
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 commun ...
. There are significant variations depending on exact species, especially in Lake Titicaca where there is a degree of
niche differentiation In ecology, niche differentiation (also known as niche segregation, niche separation and niche partitioning) refers to the process by which competing species use the environment differently in a way that helps them to coexist. The competitive excl ...
, including some that mainly feed on
zooplankton Zooplankton are the animal component of the planktonic community ("zoo" comes from the Greek word for ''animal''). Plankton are aquatic organisms that are unable to swim effectively against currents, and consequently drift or are carried along by ...
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 tributaries of the Pacific Ocean in Asia and North America. The steelhead (sometimes called "steelhead trout") is an anadromous (sea-run) form of the coasta ...
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 Aqua-Lung, open-circuit SCUBA (self-contained underwater breathing apparatus). T ...
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 their anatomies. These data are used in many areas of ichthyology, including taxonomy and fisheries biology. Overall length * Standard length (SL) is the length of a fish m ...
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; 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 ...
in 2009, but in 2014 the species was regarded as critically endangered 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 allelic drift or the Wright effect, is the change in the frequency of an existing gene variant (allele) in a population due to random chance. Genetic drift may cause gene variants to disappear completely and there ...
,
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. In ...
, 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 ''Lago Junín'', named after the nearby town of Junin) or Chinchaycocha (possibly from Quechua ''chincha, chinchay'' north, northern, ''chinchay'' ocelot, ''qucha'' lake, lagoon, "northern lake" or "ocelot lake") is the lar ...
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 pan (geology), 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 end ...
and ''O. chungarensis'' found only in Lake Chungará. 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 (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...
of several species including
mtDNA Mitochondrial DNA (mtDNA or mDNA) is the DNA located in mitochondria, cellular organelles within eukaryotic cells that convert chemical energy from food into a form that cells can use, such as adenosine triphosphate (ATP). Mitochondrial DNA ...
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 calle ...
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 their anatomies. These data are used in many areas of ichthyology, including taxonomy and fisheries biology. Overall length * Standard length (SL) is the length of a fish m ...
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 , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
, 1846
(Carache Negro) * '' Orestias albus''
Valenciennes Valenciennes (, also , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
, 1846
(Carache Amarillo) * '' Orestias ascotanensis'' 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 The cope (known in Latin as ''pluviale'' 'rain coat' or ''cappa'' 'cape') is a liturgical vestment, more precisely a long mantle or cloak, open in front and fastened at the breast with a band or clasp. It may be of any liturgical colours, litu ...
, 1876
* '' Orestias gloriae'' Vila, S. Scott,
Méndez Méndez is a common Spanish surname, originally a patronymic, meaning ''Son of Mendo'', ''Menendo'', or ''Mem''. A longer form sharing the same root is Menéndez, while the Portuguese form is Mendes. Méndez may refer to: General * Ana G. Mén ...
,
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 n ...
, 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 Orestias ( el, Ὀρεστιάς) was an ancient Greek settlement next to the Maritsa (or Evros) river, near or at the site of present-day Edirne, and close to the current border between Turkey and Greece. Legends claim that Orestias was founded ...
''
Valenciennes Valenciennes (, also , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
, 1846
(Carache Amarillo) * '' Orestias lastarriae'' Philippi , 1876 * '' Orestias laucaensis'' Arratia, 1982 * '' Orestias luteus''
Valenciennes Valenciennes (, also , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
, 1846
(Carache Amarillo) * '' Orestias multiporis'' 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 Orestias ( el, Ὀρεστιάς) was an ancient Greek settlement next to the Maritsa (or Evros) river, near or at the site of present-day Edirne, and close to the current border between Turkey and Greece. Legends claim that Orestias was founded ...
'' Vila, 2006 * '' Orestias polonorum'' Tchernavin, 1944 * '' Orestias puni'' Tchernavin, 1944 — likely a
synonym A synonym is a word, morpheme, or phrase that means exactly 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 all ...
of ''O. jussiei'' * '' Orestias richersoni'' Parenti, 1984 * '' Orestias silustani'' 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 , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
, 1846
(Titicaca Orestias, Amanto; likely
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 ...
(mid-20th century)) * '' Orestias forgeti'' Lauzanne, 1981 (Ispi) * '' Orestias ispi'' Lauzanne, 1981 (Ispi) * '' Orestias pentlandii''
Valenciennes Valenciennes (, also , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
, 1846
(Boga)


Gilsoni species complex

All restricted to the Lake Titicaca basin. * ''
Orestias gilsoni Orestias ( el, Ὀρεστιάς) was an ancient Greek settlement next to the Maritsa (or Evros) river, near or at the site of present-day Edirne, and close to the current border between Turkey and Greece. Legends claim that Orestias was founded ...
'' 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 exactly 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 all ...
of ''O. minimus'' * '' Orestias mooni'' Tchernavin, 1944 * '' Orestias robustus'' Parenti, 1984 * '' Orestias taquiri'' Tchernavin, 1944 * '' Orestias tchernavini'' Lauzanne, 1981 * ''
Orestias tomcooni Orestias ( el, Ὀρεστιάς) was an ancient Greek settlement next to the Maritsa (or Evros) river, near or at the site of present-day Edirne, and close to the current border between Turkey and Greece. Legends claim that Orestias was founded ...
'' Parenti, 1984 * '' Orestias uruni'' Tchernavin, 1944


Mulleri species complex

All restricted to the Lake Titicaca basin. * '' Orestias crawfordi'' 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 , , ; nl, label=also Dutch, Valencijn; pcd, Valincyinnes or ; la, Valentianae) is a commune in the Nord department, Hauts-de-France, France. It lies on the Scheldt () river. Although the city and region experienced a s ...
, 1846
* ''
Orestias tutini Orestias ( el, Ὀρεστιάς) was an ancient Greek settlement next to the Maritsa (or Evros) river, near or at the site of present-day Edirne, and close to the current border between Turkey and Greece. Legends claim that Orestias was founded ...
'' 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 in 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