''Cyonasua'' (meaning "dog-coati" in

Greek Greek may refer to: 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 of all kno ...

) is an extinct

genus Genus (; : genera ) is a taxonomic rank above species and below family (taxonomy), family as used in the biological classification of extant taxon, living and fossil organisms as well as Virus classification#ICTV classification, viruses. In bino ...

procyonid Procyonidae ( ) is a New World family (biology), family of the order (biology), order Carnivora. It includes the Procyon (genus), raccoons, ringtails, cacomistles, coatis, kinkajous, olingos, and olinguitos. Procyonids inhabit a wide range of e ...

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 ...

Middle Pleistocene The Chibanian, more widely known as the Middle Pleistocene (its previous informal name), is an Age (geology), age in the international geologic timescale or a Stage (stratigraphy), stage in chronostratigraphy, being a division of the Pleistocen ...

of South America. Fossils of ''Cyonasua'' have been found in

Argentina Argentina, officially the Argentine Republic, is a country in the southern half of South America. It covers an area of , making it the List of South American countries by area, second-largest country in South America after Brazil, the fourt ...

( Ituzaingó, Epecuén, Huayquerías, Monte Hermoso, Chapadmalal, Maimará, Ensenada, La Playa, Chiquimil, Andalhuala, and

Cerro Azul Formation The Cerro Azul Formation (), also described as Epecuén Formation, is a formation (geology), geological formation of Late Miocene (Tortonian, or Huayquerian in the South American land mammal age, SALMA classification) age in the Colorado Basin, Arg ...

s),

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 ...

( Tariquía Formation), Uruguay ( Camacho Formation), and

Venezuela Venezuela, officially the Bolivarian Republic of Venezuela, is a country on the northern coast of South America, consisting of a continental landmass and many Federal Dependencies of Venezuela, islands and islets in the Caribbean Sea. It com ...

( 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 epoch (geology), 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 mea ...

to early late

Pliocene The Pliocene ( ; also Pleiocene) is the epoch (geology), epoch in the geologic time scale that extends from 5.33 to 2.58Huayquerian The Huayquerian () 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 Chasicoan and precedes the Montehermosan age. Etymology T ...

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 ...

SALMAs, 7.3-3 million years old) in age, but a single early

Pleistocene The Pleistocene ( ; referred to colloquially as the ''ice age, Ice Age'') is the geological epoch (geology), epoch that lasted from to 11,700 years ago, spanning the Earth's most recent period of repeated glaciations. Before a change was fin ...

specimen (the

holotype A holotype (Latin: ''holotypus'') is a single physical example (or illustration) of an organism used when the species (or lower-ranked taxon) was formally described. It is either the single such physical example (or illustration) or one of s ...

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 an order of placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at least 279 species. Carnivor ...

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 land ...

. ''Cyonasua'' appears in the fossil record much earlier than other

North American North America is a continent in the Northern and Western hemispheres. North America is bordered to the north by the Arctic Ocean, to the east by the Atlantic Ocean, to the southeast by South America and the Caribbean Sea, and to the sou ...

immigrant groups, most of which did not appear until 3 million years ago, including other

carnivorans Carnivora ( ) is an Order (biology), order of Placentalia, placental mammals specialized primarily in eating flesh, whose members are formally referred to as carnivorans. The order Carnivora is the sixth largest order of mammals, comprising at ...

, many of which did not appear in South America until the early

(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 inhabiting South America. Despite their name, they are not true foxes, but are a unique canid genus more closely related t ...

'' and ''

Galictis A grison (, ) 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 South America, through Central America to s ...

'' 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 is a subregion of North America. Its political boundaries are defined as bordering Mexico to the north, Colombia to the southeast, the Caribbean to the east, and the Pacific Ocean to the southwest. Central America is usually ...

island hopping Leapfrogging was an amphibious military strategy employed by the Allies in the Pacific War against the Empire of Japan during World War II. The key idea was to bypass heavily fortified enemy islands instead of trying to capture every island i ...

before the formation of the

Isthmus of Panama The Isthmus of Panama, 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 America, North and South America. The country of Panama is located on the i ...

. ''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 though ...

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 126 species in 18 genera. Opossums originated in South America and entered North A ...

. Unlike most sparassodonts, which were

hypercarnivore A hypercarnivore is an animal that 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 exampl ...

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, individ ...

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 family of large carnivorous, mostly flightless birds that were among the largest apex predators in South America during the Cenozoic era. Their definitive fossil records range from ...

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 Late Pleistocene. They include some of the largest known flying birds. Its members are known as teratorns. Taxonom ...

s were

s, and other predators such as

were much smaller than most

s, the large, omnivorous ''Cyonasua'' could occupy an otherwise unoccupied niche in South American ecosystems. ''Cyonasua'' was much larger than any extant procyonid, weighing about 15–25 kg, about the same size as a medium-sized dog. However, it was much smaller than its later relative '' Chapalmalania'', which was the size of a small bear. '' Chapalmalania'' 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 considerably smaller portion in the Northern Hemisphere. It can also be described as the southern Subregion#Americas, subregion o ...

, with ''Cyonasua'' as traditionally defined potentially being

paraphyletic Paraphyly is a taxonomic term describing a grouping that consists of the grouping's last common ancestor and some but not all of its descendant lineages. The grouping is said to be paraphyletic ''with respect to'' the excluded subgroups. In co ...

. 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 North American, northern or common raccoon (also spelled racoon) to distinguish it from Procyonina, other species of raccoon, is a mammal native to North America. It is the largest ...

(''Procyon'') and

coatis Coatis (from Tupí), also known as coatimundis (), are members of the family Procyonidae in the genera '' Nasua'' and '' Nasuella'' (comprising the subtribe Nasuina). They are diurnal mammals native to South America, Central America, Mexico, ...

(''Nasua''). Fossils of ''Cyonasua'' have been found inside crotovines (fossil burrows), but these are thought to have been made by the large

armadillo Armadillos () are New World placental mammals in the order (biology), order Cingulata. They form part of the superorder Xenarthra, along with the anteaters and sloths. 21 extant species of armadillo have been described, some of which are dis ...

'' 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 Camacho Formation Neogene Venezuela Fossils of Venezuela Fossil taxa described in 1885 Taxa named by Florentino Ameghino