Hydrochoerus Hesperotiganites
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''Hydrochoerus hesperotiganites'' is an extinct species of
capybara The capybaraAlso called capivara (in Brazil), capiguara (in Bolivia), chigüire, chigüiro, or fercho (in Colombia and Venezuela), carpincho (in Argentina, Paraguay and Uruguay) and ronsoco (in Peru). or greater capybara (''Hydrochoerus hydro ...
that lived in
San Diego County San Diego County (), officially the County of San Diego, is a county in the southwestern corner of the U.S. state of California. As of the 2020 census, the population was 3,298,634, making it California's second-most populous county and the fi ...
,
California California is a U.S. state, state in the Western United States, located along the West Coast of the United States, Pacific Coast. With nearly 39.2million residents across a total area of approximately , it is the List of states and territori ...
, during the
Rancholabrean The Rancholabrean North American Land Mammal Age on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from less than 240,000 years to 11,000 years BP, a p ...
stage of the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
(between 130,000 and 80,000 years ago). It is currently the only known capybara of the genus ''
Hydrochoerus The genus ''Hydrochoerus'' contains two living and two extinct species of rodents from South America, the Caribbean island of Grenada, California and Panama. Capybaras are the largest living rodents in the world. The genus name is derived from th ...
'' found in North America. It was closely related to the modern Greater and
Lesser Capybara The lesser capybara (''Hydrochoerus isthmius'')
University of Michigan M ...
.


History and naming

The holotype specimen of ''H. hesperotiganites'' was collected by the San Diego Society of Natural History by Bradford O. Riney during mitigation monitoring at the Town Center North shopping center, City of
Oceanside, California Oceanside is a city on the South Coast (California), South Coast of California, located in San Diego County, California, San Diego County. The city had a population of 167,086 at the 2010 United States Census, 2010 census. The city is a popular ...
. The fossil was recovered from the lower of two stratigraphic units, consisting of
sandstone Sandstone is a clastic sedimentary rock composed mainly of sand-sized (0.0625 to 2 mm) silicate grains. Sandstones comprise about 20–25% of all sedimentary rocks. Most sandstone is composed of quartz or feldspar (both silicates) ...
,
mudstone Mudstone, a type of mudrock, is a fine-grained sedimentary rock whose original constituents were clays or muds. Mudstone is distinguished from '' shale'' by its lack of fissility (parallel layering).Blatt, H., and R.J. Tracy, 1996, ''Petrology. ...
and
siltstone Siltstone, also known as aleurolite, is a clastic sedimentary rock that is composed mostly of silt. It is a form of mudrock with a low clay mineral content, which can be distinguished from shale by its lack of fissility.Blatt ''et al.'' 1980, p ...
dating to the
Pleistocene The Pleistocene ( , often referred to as the ''Ice age'') is the geological Epoch (geology), epoch that lasted from about 2,580,000 to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fina ...
. The sediments more specifically most likely date to the
Rancholabrean The Rancholabrean North American Land Mammal Age on the geologic timescale is the North American faunal stage according to the North American Land Mammal Ages chronology (NALMA), typically set from less than 240,000 years to 11,000 years BP, a p ...
based on typical mammal fauna of that age and the presence of ''
Bison latifrons ''Bison latifrons'', also known as the giant bison or long-horned bison, is an extinct species of bison that lived in North America during the Pleistocene epoch ranging from Alaska to Mexico. It was the largest and heaviest bovid ever to live in ...
'' from possibly correlative strata upriver from the locality. The species name "hesperotiganites" translates to "western pancake" from the Greek words "hesperos" (western) and "tiganites" (pancake), chosen to reflect both the geographic location of the holotype skull as well as its crushed state.


Evolution

Capybaras are the only
caviomorph Caviomorpha is the rodent infraorder or parvorder that unites all New World hystricognaths. It is supported by both fossil and molecular evidence. The Caviomorpha was for a time considered to be a separate order outside the Rodentia, but is now a ...
rodents besides the new world porcupines that managed to colonize temperate North America during the Pliocene and Pleistocene stages of the
Great American Interchange The Great American Biotic Interchange (commonly abbreviated as GABI), also known as the Great American Interchange and the Great American Faunal Interchange, was an important late Cenozoic paleozoogeographic biotic interchange event in which lan ...
. Capybaras are present in North America in the form of three genera, ''
Neochoerus ''Neochoerus'' ("new hog") is an extinct genus of rodent closely related to the living capybara. Fossil remains of ''Neochoerus'' have been found through North America (México and United States) and South America in Boyacá, Colombia C ...
'' from Pliocene to Pleistocene
Mexico Mexico (Spanish: México), officially the United Mexican States, is a country in the southern portion of North America. It is bordered to the north by the United States; to the south and west by the Pacific Ocean; to the southeast by Guatema ...
,
Texas Texas (, ; Spanish language, Spanish: ''Texas'', ''Tejas'') is a state in the South Central United States, South Central region of the United States. At 268,596 square miles (695,662 km2), and with more than 29.1 million residents in 2 ...
,
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 to ...
and
South Carolina )''Animis opibusque parati'' ( for, , Latin, Prepared in mind and resources, links=no) , anthem = " Carolina";" South Carolina On My Mind" , Former = Province of South Carolina , seat = Columbia , LargestCity = Charleston , LargestMetro = ...
, '' Phugatherium'' from late Blancan Florida and
Arizona Arizona ( ; nv, Hoozdo Hahoodzo ; ood, Alĭ ṣonak ) is a state in the Southwestern United States. It is the 6th largest and the 14th most populous of the 50 states. Its capital and largest city is Phoenix. Arizona is part of the Fou ...
and ''Hydrochoerus hesperotiganites'' from the Rancholabrean of California. This makes ''H. hesperotiganites'' the only member of the extant genus ''Hydrochoerus'' to have made it into the United States, with any species previously referred to ''Hydrochoerus'' subsequently being considered to be part of ''Neochoerus''. This however presents a problem regarding the dispersal of ''Hydrochoerus'' into North America, as there is no indication for their presence between San Diego County and the northernmost extent of their current range (the
Lesser Capybara The lesser capybara (''Hydrochoerus isthmius'')
University of Michigan M ...
in
Panama Panama ( , ; es, link=no, Panamá ), officially the Republic of Panama ( es, República de Panamá), is a transcontinental country spanning the southern part of North America and the northern part of South America. It is bordered by Cos ...
) and the only known fossil remains of capybaras in Central America belong to ''Neochoerus''. Tracing the route ''Hydrochoerus'' could have taken to reach California is subsequently difficult, with White ''et al.'' proposing two different paths. Both begin with the ancestors of ''H. hesperotiganites'' traveling along the Pacific coastline of
Sinaloa Sinaloa (), officially the Estado Libre y Soberano de Sinaloa ( en, Free and Sovereign State of Sinaloa), is one of the 31 states which, along with Mexico City, comprise the Administrative divisions of Mexico, Federal Entities of Mexico. It is d ...
and
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 Administrative divisions of Mexico, Federal Entities of Mexico. The state is d ...
, before two different paths become available. The first would have led them further along the coast and up the
Colorado River The Colorado River ( es, Río Colorado) is one of the principal rivers (along with the Rio Grande) in the Southwestern United States and northern Mexico. The river drains an expansive, arid drainage basin, watershed that encompasses parts of ...
. However, to reach the San Luis Rey River overland travel would have been necessary with permanent water sources being rare, thusly this route is deemed the less likely of the two by the authors. The second route would have seen capybaras travel up some of the other rivers draining into the Gulf of California, traveling further north and eventually dispersing southwards. A similar route along the Rio Yaqui is known to have been taken by the other two capybara genera as they spread throughout Northern Mexico. White and colleagues further suggest that increased sampling of Pleistocene localities in Mexico might help resolve the matter.


Paleoenvironment

The sediments ''H. hesperotiganites'' was discovered in where fluvial-lacustrine in nature and have been interpreted to represent what was once a freshwater pond or
oxbow lake An oxbow lake is a U-shaped lake or pool that forms when a wide meander of a river is cut off, creating a free-standing body of water. In South Texas, oxbows left by the Rio Grande are called '' resacas''. In Australia, oxbow lakes are call ...
. The lake was formed atop of eroded
Eocene The Eocene ( ) Epoch is a geological epoch (geology), epoch that lasted from about 56 to 33.9 million years ago (mya). It is the second epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), Era. The name ''Eocene' ...
strata, likely as the ancestral San Luis Rey River deposited sediments across its floodplains. The presence of changing layers of silt- and sandstone indicates that sediment input changed across the seasons. The upper layers indicate that the river was eventually meandering south again, forming yet another oxbow lake while the floodplains were aggrading. As most mammal remains are isolated, it's been proposed that mammalian corpses washed ashore, where parts decomposed and fossilized while others were transported further downriver. The lower strata preserve little fauna, only consisting of the capybara remains as well as those of ''
Stockoceros ''Stockoceros'' is an extinct genus of the North American artiodactyl family Antilocapridae (pronghorns), known from Mexico and the southwestern United States. Its horns are each divided near their base into two prongs of roughly equal length. T ...
'', a relative of the pronghorn. The slightly younger units however show a diverse fauna including
pond turtles Emydidae (Latin ''emys'' (freshwater tortoise) + Ancient Greek εἶδος (''eîdos'', “appearance, resemblance”)) is a family of testudines (turtles) that includes close to 50 species in 10 genera. Members of this family are commonly calle ...
,
grebes Grebes () are aquatic diving birds in the order (biology), order Podicipediformes . Grebes are widely distributed freshwater birds, with some species also found in sea, marine habitats during Bird migration, migration and winter. Some flightless ...
,
pelicans Pelicans (genus ''Pelecanus'') are a genus of large water birds that make up the family Pelecanidae. They are characterized by a long beak and a large throat pouch used for catching prey and draining water from the scooped-up contents before s ...
,
ducks Duck is the common name for numerous species of waterfowl in the family Anatidae. Ducks are generally smaller and shorter-necked than swans and geese, which are members of the same family. Divided among several subfamilies, they are a form t ...
coots Coots are medium-sized water birds that are members of the rail family, Rallidae. They constitute the genus ''Fulica'', the name being the Latin term for "coot". Coots have predominantly black plumage, and—unlike many rails—they are usually ...
,
rails Rail or rails may refer to: Rail transport *Rail transport and related matters *Rail (rail transport) or railway lines, the running surface of a railway Arts and media Film * ''Rails'' (film), a 1929 Italian film by Mario Camerini * ''Rail'' ( ...
as well as various other shore- and songbirds. Mammals are likewise diverse and abundant, with scientists having recovered the fossils of
grey fox The gray fox (''Urocyon cinereoargenteus''), or grey fox, is an omnivorous mammal of the family Canidae, widespread throughout North America and Central America. This species and its only congener (biology), congener, the diminutive island fox ...
es,
tapirs Tapirs ( ) are large, herbivorous mammals belonging to the family Tapiridae. They are similar in shape to a pig, with a short, prehensile nose trunk. Tapirs inhabit jungle and forest regions of South and Central America, with one species inhabit ...
,
horses The horse (''Equus ferus caballus'') is a domesticated, one-toed, hoofed mammal. It belongs to the taxonomic family Equidae and is one of two extant subspecies of ''Equus ferus''. The horse has evolved over the past 45 to 55 million yea ...
, the
American Mastodon American(s) may refer to: * American, something of, from, or related to the United States of America, commonly known as the "United States" or "America" ** Americans, citizens and nationals of the United States of America ** American ancestry, pe ...
as well as the ground sloths ''
Nothrotheriops ''Nothrotheriops'' is a genus of Pleistocene ground sloth found in North America, from what is now central Mexico to the southern United States. This genus of bear-sized xenarthran was related to the much larger, and far more famous ''Megatherium ...
'' and ''
Megalonyx ''Megalonyx'' (Greek, "large claw") is an extinct genus of ground sloths of the family Megalonychidae, native to North America during the Pliocene and Pleistocene epochs. It became extinct during the Quaternary extinction event at the end of the ...
''. The presence of abundant waterfowl and shorebird all match the environment these strata represented during the Pleistocene and also matches the presence of Capybaras, known to inhabit areas close to bodies of water in modern South America.


References

{{Taxonbar, from=Q111110016 Glires Prehistoric mammals Cavies Fossil taxa described in 2022