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''Cyonasua'' (meaning "dog-coati" in
Greek Greek may refer to: Greece Anything of, from, or related to Greece, a country in Southern Europe: *Greeks, an ethnic group. *Greek language, a branch of the Indo-European language family. **Proto-Greek language, the assumed last common ancestor ...
) is an extinct
genus Genus ( plural genera ) is a taxonomic rank used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In the hierarchy of biological classification, genus com ...
of
procyonid Procyonidae is a New World family of the order Carnivora. It comprises the raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of environments and are generally omnivorous. Characteri ...
from the
Late Miocene The Late Miocene (also known as Upper Miocene) is a sub-epoch of the Miocene epoch (geology), Epoch made up of two faunal stage, stages. The Tortonian and Messinian stages comprise the Late Miocene sub-epoch, which lasted from 11.63 Ma (million ye ...
to
Middle Pleistocene The Chibanian, widely known by its previous designation of Middle Pleistocene, is an age in the international geologic timescale or a stage in chronostratigraphy, being a division of the Pleistocene Epoch within the ongoing Quaternary Period. The ...
of South America. Fossils of ''Cyonasua'' have been found in
Argentina Argentina (), officially the Argentine Republic ( es, link=no, República Argentina), is a country in the southern half of South America. Argentina covers an area of , making it the second-largest country in South America after Brazil, th ...
( Ituzaingó, Epecuén, Huayquerías, Monte Hermoso, Chapadmalal, Maimará, Ensenada, La Playa, Chiquimil, Andalhuala, and
Cerro Azul Formation The Cerro Azul Formation ( es, Formación Cerro Azul), in the Buenos Aires Province also described as Epecuén Formation, is a geological formation of Late Miocene (Tortonian, or Huayquerian in the SALMA classification) age in the Colorado Basin ...
s),
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 ...
( Tariquía Formation), Uruguay ( Camacho Formation), and
Venezuela Venezuela (; ), officially the Bolivarian Republic of Venezuela ( es, link=no, República Bolivariana de Venezuela), is a country on the northern coast of South America, consisting of a continental landmass and many islands and islets in th ...
( San Gregorio Formation). The oldest well-dated fossils of ''Cyonasua'' are approximately 7.3 million years old. Most fossils of ''Cyonasua'' are late
Miocene The Miocene ( ) is the first geological epoch of the Neogene Period and extends from about (Ma). The Miocene was named by Scottish geologist Charles Lyell; the name comes from the Greek words (', "less") and (', "new") and means "less recen ...
to early late
Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58Huayquerian The Huayquerian ( es, Huayqueriense) age is a period of geologic time (9.0–6.8 Ma) within the Late Miocene epoch of the Neogene, used more specifically within the SALMA classification. It follows the Mayoan and precedes the Montehermosan age. ...
to
Chapadmalalan The Chapadmalalan age is a period of geologic time (4.0–3.0 Ma) within the Pliocene epoch of the Neogene used more specifically with South American Land Mammal Ages. It follows the Montehermosan and precedes the Uquian age. Fossil content F ...
SALMAs, 7.3-3 million years old) in age, but a single early
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 ...
specimen (the
holotype A holotype is a single physical example (or illustration) of an organism, known to have been used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of several ...
and only known specimen of ''Cyonasua meranii'') indicates that members of this genus survived until at least 0.99 million years ago (the fossil layer where this specimen was collected dating to the Jaramillo Chron). ''Cyonasua'' is the oldest terrestrial
carnivoran Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
known from South America, and represents the earliest undisputed southward mammalian migrants 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 ...
. ''Cyonasua'' appears in the fossil record much earlier than other
North American North America is a continent in the Northern Hemisphere and almost entirely within the Western Hemisphere. It is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Ca ...
immigrant groups, most of which did not appear until 3 million years ago, including other
carnivorans Carnivora is a monophyletic order of placental mammals consisting of the most recent common ancestor of all cat-like and dog-like animals, and all descendants of that ancestor. Members of this group are formally referred to as carnivorans, ...
, many of which did not appear in South America until the early
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 ...
(about 1.2 million years ago). The next oldest remains of carnivorans in South America are rare specimens of ''
Lycalopex The South American foxes (''Lycalopex''), commonly called ''raposa'' in Portuguese, or ''zorro'' in Spanish, are a genus from South America of the subfamily Caninae. Despite their name, they are not true foxes, but are a unique canid genus more ...
'' and ''
Galictis A grison , , also known as a South American wolverine, is any mustelid in the genus ''Galictis''. Native to Central and South America, the genus contains two extant species: the greater grison (''Galictis vittata''), which is found widely in So ...
'' from the Barrancalobian (~2.9 million years old), nearly 4.4 million years after the first appearance of ''Cyonasua'' in South America. The ancestors of ''Cyonasua'' are thought to have arrived from
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. ...
by
island hopping Leapfrogging, also known as island hopping, was a military strategy employed by the Allies in the Pacific War against the Empire of Japan during World War II. The key idea is to bypass heavily fortified enemy islands instead of trying to captu ...
before the formation of the
Isthmus of Panama The Isthmus of Panama ( es, Istmo de Panamá), also historically known as the Isthmus of Darien (), is the narrow strip of land that lies between the Caribbean Sea and the Pacific Ocean, linking North and South America. It contains the country ...
. ''Cyonasua'' arrived in South America when ecosystems there were still dominated by
metatherian Metatheria is a mammalian clade that includes all mammals more closely related to marsupials than to placentals. First proposed by Thomas Henry Huxley in 1880, it is a more inclusive group than the marsupials; it contains all marsupials as well ...
carnivores, namely
sparassodonts Sparassodonta (from Greek to tear, rend; and , gen. , ' tooth) is an extinct order of carnivorous metatherian mammals native to South America, related to modern marsupials. They were once considered to be true marsupials, but are now thoug ...
and carnivorous
opossums Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 93 species in 18 genera. Opossums originated in South America and entered North A ...
. Unlike most sparassodonts, which were
hypercarnivore A hypercarnivore is an animal which has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant examp ...
s, ''Cyonasua'' was an omnivore and it is thought that it did not directly
compete Competition is a rivalry where two or more parties strive for a common goal which cannot be shared: where one's gain is the other's loss (an example of which is a zero-sum game). Competition can arise between entities such as organisms, indivi ...
with sparassodonts for food. This is thought to be one possible reason why ''Cyonasua'' and its close relatives were able to colonize South America many millions of years before other carnivorans. Because sparassodonts and other native predators like
terror birds Phorusrhacids, colloquially known as terror birds, are an extinct clade of large carnivorous flightless birds that were one of the largest species of apex predators in South America during the Cenozoic era; their conventionally accepted temporal ...
and
teratorn Teratornithidae is an extinct family of very large birds of prey that lived in North and South America from the Late Oligocene to the Late Pleistocene. They include some of the largest known flying birds. Taxonomy Teratornithidae are related ...
s were
hypercarnivore A hypercarnivore is an animal which has a diet that is more than 70% meat, either via active predation or by scavenging. The remaining non-meat diet may consist of non-animal foods such as fungi, fruits or other plant material. Some extant examp ...
s, and other predators such as
opossums Opossums () are members of the marsupial order Didelphimorphia () endemic to the Americas. The largest order of marsupials in the Western Hemisphere, it comprises 93 species in 18 genera. Opossums originated in South America and entered North A ...
were much smaller than most
procyonid Procyonidae is a New World family of the order Carnivora. It comprises the raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of environments and are generally omnivorous. Characteri ...
s, the large, omnivorous ''Cyonasua'' could occupy an otherwise unoccupied
niche Niche may refer to: Science *Developmental niche, a concept for understanding the cultural context of child development *Ecological niche, a term describing the relational position of an organism's species *Niche differentiation, in ecology, the ...
in South American ecosystems. ''Cyonasua'' was much larger than any extant procyond, weighing about 15–25 kg, about the same size as a medium-sized dog. However, it was much smaller than its later relative ''
Chapalmalania ''Chapalmalania'' is an extinct genus of procyonid from the Pliocene (Chapadmalalan to Uquian) of Argentina and Colombia (Ware Formation, Cocinetas Basin, La Guajira). Description Though related to raccoons and coatis, ''Chapalmalania'' was a ...
'', which was the size of a small bear. ''
Chapalmalania ''Chapalmalania'' is an extinct genus of procyonid from the Pliocene (Chapadmalalan to Uquian) of Argentina and Colombia (Ware Formation, Cocinetas Basin, La Guajira). Description Though related to raccoons and coatis, ''Chapalmalania'' was a ...
'' is thought to have been closely related to ''Cyonasua'', with the two genera frequently referred to as "''Cyonasua''-group procyonids". ''Chapalmalania'' is thought to have evolved from a species of ''Cyonasua'' in
South America South America is a continent entirely in the Western Hemisphere and mostly in the Southern Hemisphere, with a relatively small portion in the Northern Hemisphere at the northern tip of the continent. It can also be described as the southe ...
, with ''Cyonasua'' as traditionally defined potentially being
paraphyletic In taxonomy (general), taxonomy, a group is paraphyletic if it consists of the group's most recent common ancestor, last common ancestor and most of its descendants, excluding a few Monophyly, monophyletic subgroups. The group is said to be pa ...
. Limb bones of ''Cyonasua'' suggest that this genus was a generalized terrestrial mammal with some capability to dig and climb, similar to
raccoons The raccoon ( or , ''Procyon lotor''), sometimes called the common raccoon to distinguish it from other species, is a mammal native to North America. It is the largest of the procyonid family, having a body length of , and a body weight of . ...
(''Procyon'') and
coatis Coatis, also known as coatimundis (), are members of the family Procyonidae in the genera '' Nasua'' and '' Nasuella''. They are diurnal mammals native to South America, Central America, Mexico, and the southwestern United States. The name ...
(''Nasua''). Fossils of ''Cyonasua'' have been found inside crotovines (fossil burrows), but these are thought to have been made by the large
armadillo Armadillos (meaning "little armored ones" in Spanish) are New World placental mammals in the order Cingulata. The Chlamyphoridae and Dasypodidae are the only surviving families in the order, which is part of the superorder Xenarthra, along wi ...
''Ringueletia'' and later occupied by ''Cyonasua''. The teeth of ''Cyonasua'' suggest it was omnivorous, more carnivorous than most living procyonids, but less carnivorous than the modern ringtails ( ''Bassariscus'' spp.)


References

{{Taxonbar, from=Q16941543 Prehistoric procyonids Prehistoric carnivoran genera Miocene musteloids Miocene mammals of South America Pliocene mammals of South America Chapadmalalan Montehermosan Huayquerian Neogene Argentina Fossils of Argentina Cerro Azul Formation Ituzaingó Formation Neogene Venezuela Fossils of Venezuela Fossil taxa described in 1885 Taxa named by Florentino Ameghino