''Devincenzia'' is an

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 ...

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 giant flightless

predator Predation is a biological interaction where one organism, the predator, kills and eats another organism, its prey. It is one of a family of common feeding behaviours that includes parasitism and micropredation (which usually do not kill th ...

bird Birds are a group of warm-blooded vertebrates constituting the class Aves (), characterised by feathers, toothless beaked jaws, the laying of hard-shelled eggs, a high metabolic rate, a four-chambered heart, and a strong yet lightweigh ...

s in the

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

Phorusrhacidae 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 ...

or "terror birds" that lived during the Early Miocene (

Deseadan The Deseadan ( es, Deseadense) age is a period of geologic time (29.0–21.0 Ma) within the Oligocene epoch of the Paleogene to the Early Miocene epoch of the Neogene, used more specifically within the SALMA classification of South America. It fo ...

) Fray Bentos Formation of

Uruguay Uruguay (; ), officially the Oriental Republic of Uruguay ( es, República Oriental del Uruguay), is a country in South America. It shares borders with Argentina to its west and southwest and Brazil to its north and northeast; while bordering ...

and

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 ...

(

Huayquerian 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. ...

)

Ituzaingó Formation The Ituzaingó Formation ( es, Formación Ituzaingó), in older literature also described as Entre Ríos or Entrerriana Formation, is an extensive geological formation of Late Miocene (Tortonian, or Huayquerian in the SALMA classification) age in ...

Early Pliocene Early may refer to: History * The beginning or oldest part of a defined historical period, as opposed to middle or late periods, e.g.: ** Early Christianity ** Early modern Europe Places in the United States * Early, Iowa * Early, Texas * Early ...

(

Montehermosan The Montehermosan age is a period of geologic time (6.8–4.0 Ma) within the Miocene and Pliocene epochs of the Neogene used more specifically with South American Land Mammal Ages. It follows the Huayquerian and precedes the Chapadmalalan The C ...

) of

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 ...

. The

type species In zoological nomenclature, a type species (''species typica'') is the species name with which the name of a genus or subgenus is considered to be permanently taxonomically associated, i.e., the species that contains the biological type specimen ...

''D. pozzi'' was formerly known as ''Onactornis pozzi''.''Devincenzia''
at

Fossilworks Fossilworks is a portal which provides query, download, and analysis tools to facilitate access to the Paleobiology Database The Paleobiology Database is an online resource for information on the distribution and classification of fossil animals ...

.org It stood about tall, making it one of the largest Phorusrhacids and carnivorous birds known.

Etymology

The generic name ''Devincenzia'' comes from Uruguayan museum director and zoologist Garibaldi Devincenzi (1882-1943) and the specific name of ''D. gallinali'' comes from Alejandro Gallinal, another Uruguayan scientist. The specific name of ''D. pozzi'' was after the lead taxidermist at the museum, Antonio Pozzi.

History and taxonomy

In 1931, a very large distal right tarsometatarsus associated with a ungual phalanx from digit II, was described by the

an Paleontologist Lucas Kraglievich as a new species of ''Phororhacos'' (a misspelling of ''Phorusrhacos''), ''P.'' ''pozzi'' and deposited at the Museo Argentino de Ciencias Naturales Bernardino Rivadavia in

Buenos Aires, Argentina Buenos Aires ( or ; ), officially the Autonomous City of Buenos Aires ( es, link=no, Ciudad Autónoma de Buenos Aires), is the capital and primate city of Argentina. The city is located on the western shore of the Río de la Plata, on South ...

under specimen numbers MACN-6554 and 6681.Kraglievich, L. (1931). Contribución al conocimiento de las aves fósiles de la época araucoentrerriana. ''Physis'', ''10'', 304-315. The fossils had been found in Lower Pliocene rock layers at El Brete in Cordoba, Argentina, specifically from the Mesopotamian. Later in the same paper, Kraglievich named a subspecies of ''Phororhacos'' (''Phorusrhacos'') ''longissimus mendocinus'' based on a partial proximal right femur from the

Huayquerías Formation The Huayquerías Formation ( es, Formación Huayquerías) is a Late Miocene fossiliferous geological formation of the Frontal Cordillera and Cuyo Basin of Argentina. The formation crops out in the central Mendoza Province. The formation, with a m ...

Mendoza, Argentina Mendoza (, ), officially the City of Mendoza ( es, Ciudad de Mendoza) is the capital of the province of Mendoza in Argentina. It is located in the northern-central part of the province, in a region of foothills and high plains, on the eastern si ...

.Brodkorb, P. (1967). ''Catalogue of fossil birds: part 3 (Ralliformes, Ichthyornithiformes, Charadriiformes)''. University of Florida. The subspecies has since been synonymized with ''Devincenzia pozzi''. Kraglievich also referred a synphysis fragment to ''Phororhacos'' (''Phorusrhacos'') ''platygnathus,'' but the fossil has since been referred to ''Devincenzia pozzi.'' The next year, Kraglievich named a new genus and species of Phorusrhacid from Argentina based on a partial right tarsometatarsus of a juvenile individual (MNHN-M-189), naming it ''Devincenzia gallinali.'' The origin of the fossil is unknown, with Kraglievich initially speculating that it was from Uruguay, but the coloration corroborates with that of Patagonian fossils from the Miocene. Although they are sometimes considered distinct, Herculano Alvarenga and Elizabeth Höfling synonymized the two species in 2013 in their reassesment of Phorusrhacidae and moved ''P. pozzi'' to ''Devincenzia'' in the same paper. In August 1936 in a lagoon at Campo de Robilotte on Lake Epecuén, around 600 km southwest of the city of

Buenos Aires Buenos Aires ( or ; ), officially the Autonomous City of Buenos Aires ( es, link=no, Ciudad Autónoma de Buenos Aires), is the capital and primate city of Argentina. The city is located on the western shore of the Río de la Plata, on South ...

, Antonio Castro collected a partial skull and two pedal phalanges of a large Phorusrhacid, as well as fossils of other Cenozoic fossil taxa like

Xenarthrans Xenarthra (; from Ancient Greek wikt:ξένος, ξένος, xénos, "foreign, alien" + wikt:ἄρθρον, ἄρθρον, árthron, "joint") is a major clade of Placentalia, placental mammals native to the Americas. There are 31 living species: ...

and Macraucheniids. The fossils were sent to the Museo de la Plata, where they were described by Angelo Cabrera in 1939, naming the Phorusrhacid specimens (MLP 37-III-7-8) ''Onactornis depressus.'' Parts of the ''Onactornis'' skull are missing, so they were reconstructed with plaster to be put on display at the MLP, but this has caused some of the preserved parts to be confused with the plaster ones. ''P. pozzi'' and ''P. longissimus mendocinus'' were moved to ''Onactornis'' by Pierce Brodkorb, who also believed that ''Devincenzia gallinali'' was a synonym of ''

Brontornis ''Brontornis'' is an extinct genus of giant bird that inhabited Argentina during the Early to Middle Miocene. Its taxonomic position is highly controversial, with authors alternatively considering it to be a cariamiform, typically a phorusrhaci ...

.'' The Ituzaingo Formation also bears ''Devincenzia'' fossils, including a pedal phalange, tibiotarsus fragment, tarsometatarsus fragment, a cervical vertebra, and a dorsal vertebra that were referred to ''D. gallinali'' based on size and morphology.Alvarenga, H., Jones, W., & Rinderknecht, A. (2010)
The youngest record of phorusrhacid birds (Aves, Phorusrhacidae) from the late Pleistocene of Uruguay.
''Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen'', ''256''(2), 229-234. Two other fossil remains include a right tarsometatarsus, about 40 cm long, lacking the inner knuckle, probably from the Arroyo Roman river basin in the

Río Negro Department Río Negro Department ( es, Departamento de Río Negro, ) is a department of the northwestern region of Uruguay. It has an area of and a population of 54,765. Its capital is Fray Bentos. It borders Paysandú Department to the north, Tacuarembó D ...

, and the lower articular end of another tarsometatarsus. These specimens come from the

upper Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58lower Pleistocene, extending the potential range of the taxon.

Description

''Devincenzia'' was one of the largest representatives of

. It surpassed the type genus '' Phorusrhacos'', whose total height is given at about 2.4 m, making it potentially the largest phorusrhacid. The probable body weight of ''Devincenzia'' is given as about 161 kg. It probably resembled its relatives ''Phorusrhacos'' and ''

Kelenken ''Kelenken'' is a genus of phorusrhacid ("terror bird"), an extinct group of large, predatory birds, which lived in what is now Argentina in the middle Miocene about 15 million years ago. The only known specimen was discovered by high school stu ...

'', like them, it was equipped with a narrow body, conspicuously elongated walking legs, and retracted wings. The skull was preserved with the rear part and areas of the upper jaw. Reconstructed, it was probably about 64.5 cm long, which roughly corresponded to the known skull of ''Phorusrhacos,'' but was slightly smaller than that of ''Kelenken''. When viewed from above, it had a wedge shape, reaching a width of 32.3 cm at the

occipital bone The occipital bone () is a neurocranium, cranial dermal bone and the main bone of the occiput (back and lower part of the skull). It is trapezoidal in shape and curved on itself like a shallow dish. The occipital bone overlies the occipital lobe ...

and a height of 12.7 cm. The height of the skull at the back of the head corresponded to about 39% of the width, which is less than in ''Phorusrhacos'' with 47% or '' Psilopterus'' with 48%. In this regard, the skull of ''Devincenzia'' more closely resembled that of ''Kelenken''. It narrowed towards the front, with the width at the frontal bone being 24.4 cm, and at the temporal fossa it drew in significantly and measured 19.1 cm. The

frontal bone The frontal bone is a bone in the human skull. The bone consists of two portions.''Gray's Anatomy'' (1918) These are the vertically oriented squamous part, and the horizontally oriented orbital part, making up the bony part of the forehead, par ...

was very wide, and the two processes occurring there, the processus postorbitalis and the processus supraorbitalis, were separated by a deep recess. The

quadrate Quadrate may refer to: * Quadrate bone * Quadrate (heraldry) * Quadrate lobe of liver * Quadrate tubercle The quadrate tubercle is a small tubercle found upon the upper part of the femur. It serves as a point of insertion of the quadratus femori ...

appeared comparatively large, differing from that of ''Psilopterus'' in addition to the different design of the three extensions, among other things, by the comparatively smaller part of the main bone, which was connected to the cheekbone. Likewise, the os quadratojugale showed a more robust structure and was comparatively higher. Since the upper jaw is incomplete, and only the middle section is available, the dimensions of the beak can only be inferred. Using dimensions from other phorusrhacids, it may have been 36 cm long and 17 cm high. The symphysis of the lower jaw, which has also been handed down in fragments, had a much narrower and lower shape than in ''Brontornis''. It was only preserved to a length of 11 cm, but it would have reached a length of around 16 cm. At the rear end it was 6.2 cm wide and 4.7 cm thick, and towards the front it became significantly lower, measuring about 2.8 cm at half length. The underside had a slight curvature, more pronounced than that of ''

Physornis ''Physornis'' is an extinct genus of giant flightless predatory birds of the family Phorusrhacidae or "terror birds", most closely related to '' Paraphysornis'', that lived in Argentina. The type species is ''P. fortis''. It lived during the Mi ...

,'' which had a nearly flat surface. The externally visible foramina were wide and deep.Cabrera, Á. (1939)
Sobre vertebrados fósiles del Plioceno de Adolfo Alsina.
''Revista del Museo de La Plata'', ''2''(6), 3-35. Of the few surviving elements of the body skeleton, a complete tibiotarsus remained, and it was 72 cm long and 10.4 cm wide at the lower end of the joint, resembling that of ''Phorusrhacos'' in its long and slender build, although it was larger. At the lower end of the joint it had a bony bridge (pons supratendineus) that was conspicuous for the phorusrhacids. In addition, the lower inner joint roller protruded further forward than that of

Galliformes Galliformes is an order of heavy-bodied ground-feeding birds that includes turkeys, chickens, quail, and other landfowl. Gallinaceous birds, as they are called, are important in their ecosystems as seed dispersers and predators, and are often ...

.Tambussi, C., Ubilla, M., & Perea, D. (1999). The youngest large carnassial bird (Phorusrhacidae, Phorusrhacinae) from South America (Pliocene–Early Pleistocene of Uruguay). ''Journal of Vertebrate Paleontology'', ''19''(2), 404-406. An almost complete specimen, missing only the inner jointed roller, reached a length of 40 cm and a width at the top of 11 cm. In the middle of the shaft, the side edges drew in significantly further than in ''Phorusrhacos'', which showed a relatively even course of the diaphysis there. With a width of 4.3 cm, the middle joint roll at the bottom was significantly more voluminous than the other two and indicates that ''Devincenzia'''s middle toe was also more massive. The first toe of the third (middle) ray measured 12.5 cm in length. It was long and narrow with a width of 5.3 cm and a height of 5.7 cm. The surviving end phalange of the second toe measured around 9 cm, had an oval cross-section, was strongly curved in side view, and was strongly pressed laterally. The width at the joint end facing the body was 2.4 cm and the height was 3.9 cm.

Paleobiology

Phorusrhacids are though to have been ground predators or scavengers, and have often been considered apex predators that dominated Cenozoic South America in the absence of mammalian predators, though they did co-exist with some large, carnivorous

borhyaenid Borhyaenidae is an extinct metatherian family of low-slung, heavily built predatory mammals in the order Sparassodonta. Borhyaenids are not true marsupials, but members of a sister taxon, Sparassodonta. Like most metatherians, borhyaenids and o ...

mammals. Earlier hypotheses of phorusrhacid feeding ecology were mainly based on them possessing large skulls with hooked beaks rather than through detailed hypotheses and biomechanical studies, and such studies of their running and predatory adaptations were only tested from the beginning of the 21st century. Alvarenga and Elizabeth Höfling made some general remarks about phorusrhacid habits in a 2003 article. They were flightless, as evidenced by the proportional size of their wings and body mass, and wing-size was more reduced in larger members of the group. They pointed out that the narrowing of the pelvis, upper maxilla, and

thorax The thorax or chest is a part of the anatomy of humans, mammals, and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main divisions of the cre ...

may have been adaptations for hunting in regions with high vegetation, which would permit greater agility when moving between vertical obstacles. The narrow upper maxilla would also help catching small animals hidden among tree trunks or stones. The large expansions above the eyes formed by the lacrimal bones (similar to what is seen in modern

hawks Hawks are birds of prey of the family Accipitridae. They are widely distributed and are found on all continents except Antarctica. * The subfamily Accipitrinae includes goshawks, sparrowhawks, sharp-shinned hawks and others. This subfamily a ...

) would have protected the eyes against the sun, and enabled keen eyesight, which indicates they hunted by sight in open, sunlit areas, and not shaded forests. A 2010 study by Degrange and colleagues of ''

Andalgalornis ''Andalgalornis'' is a genus of flightless predatory birds of the extinct family Phorusrhacidae (often called "terror birds") that lived in Argentina. The type and only species is ''A. steulleti''. Taxonomy ''Andalgalornis'' is known from an in ...

'', based on

Finite Element Analysis The finite element method (FEM) is a popular method for numerically solving differential equations arising in engineering and mathematical modeling. Typical problem areas of interest include the traditional fields of structural analysis, heat ...

using

CT scans A computed tomography scan (CT scan; formerly called computed axial tomography scan or CAT scan) is a medical imaging technique used to obtain detailed internal images of the body. The personnel that perform CT scans are called radiographers ...

, estimated its

bite force Bite force quotient (BFQ) is a numerical value commonly used to represent the bite force of an animal, while also taking factors like the animal's size into account. The BFQ is calculated as the regression of the quotient of an animal's bite f ...

and stress distribution in its skull, and showed that it had lost a large degree of intracranial immobility (mobility of skull bones in relation to each other), as was also the case for other large phorusrhacids. These researchers interpreted this loss as an adaptation for enhanced rigidity of the skull, and compared to the modern

red-legged seriema The red-legged seriema (''Cariama cristata''), also known as the crested cariama and crested seriema, is a mostly predatory terrestrial bird in the seriema family ( Cariamidae), included in the Gruiformes in the old paraphyletic circumscription ...

and

white-tailed eagle The white-tailed eagle (''Haliaeetus albicilla'') is a very large species of sea eagle widely distributed across temperate Palearctic, Eurasia. Like all eagles, it is a member of the family Accipitridae (or accipitrids) which includes other diur ...

, the skull of the phorusrhacid showed relatively high stress under sideways loadings, but low stress where force was applied up and down, and in simulations of “pullback”. Due to the relative weakness of the skull at the sides and middle, these researchers considered it unlikely that ''Andalgalornis'' engaged in potentially risky behavior that involved using its beak to subdue large, struggling prey. Instead, they suggested that it either fed on smaller prey that could be killed and consumed more safely, by for example swallowing it whole, or that when targeting large prey, it used a series of well-targeted repetitive strikes with the beak, in a "attack-and-retreat" strategy. Struggling prey could also be retained with the feet, despite the lack of sharp talons. A 2012 follow up study by Tambussi and colleagues analyzed the flexion abilities of the neck of ''Andalgalornis'', based on the morphology of its neck vertebrae, finding the neck to be divided into three sections. They concluded that the neck musculature and skeleton of ''Andalgalornis'' was adapted to carrying a large head, and for helping it rise after a maximum downwards strike, and the researchers assumed the same would be true for other large, big-headed phorusrhacids. A 2020 study of phorusrhacid skull morphology by Degrange found that there were two main morphotypes within the group, derived from a seriema-like ancestor; the "Psilopterine Skull Type", which was

plesiomorphic In phylogenetics, a plesiomorphy ("near form") and symplesiomorphy are synonyms for an ancestral character shared by all members of a clade, which does not distinguish the clade from other clades. Plesiomorphy, symplesiomorphy, apomorphy, and ...

(more similar to the ancestral type), and the "Terror Bird Skull Type", which included ''Andalgalornis'' and other large members, that was more specialized, with more rigid and stiff skulls. Despite the differences, studies have shown the two types handled prey similarly, while the more rigid skulls and resulting larger bite force of the "Terror Bird" type would have been an adaptation to handling larger prey.

Phylogeny

Due to the fragmentary nature of ''Devincenzia'' fossils and the internal taxonomy of the genus being in a flux, its classification within Phorusrhacidae frequently changes. Despite this, ''Devincenzia'' is frequently found to be a member of the Phorusrhacinae along with '' Phorusrhacos,

Titanis ''Titanis'' was an extinct genus of giant flightless terror birds that inhabited North America during the early Pliocene to early Pleistocene epochs. The generic name, ''Titanis'', refers to the titans, Ancient Greek gods that preceded the Twelv ...

,'' and possibly the Patagornithines as well. The following phylogenetic tree shows the internal relationships of Phorusrhacidae under the exclusion of ''Brontornis'' as published by Degrange and colleagues in 2015, which recovers ''Devincenzia'' as a member of a large clade that includes ''Physornis'', ''Phorusrhacos'' and ''Andalgalornis'', among others. After the extinction of the non-bird dinosaurs, during the early

Cenozoic The Cenozoic ( ; ) is Earth's current geological era, representing the last 66million years of Earth's history. It is characterised by the dominance of mammals, birds and flowering plants, a cooling and drying climate, and the current configura ...

, mammals underwent an evolutionary diversification, and birds around the world developed a tendency towards

gigantism Gigantism ( el, γίγας, ''gígas'', "giant", plural γίγαντες, ''gígantes''), also known as giantism, is a condition characterized by excessive growth and height significantly above average. In humans, this condition is caused by ove ...

; this includes the

Gastornithidae ''Gastornis'' is an extinct genus of large flightless birds that lived during the mid Paleocene to mid Eocene epochs of the Paleogene period. Fossils have been found in Europe, Asia and North America, with the remains from North America origina ...

, the

Dromornithidae Dromornithidae, known as mihirungs and informally as thunder birds or demon ducks, were a clade of large, flightless Australian birds of the Oligocene through Pleistocene Epochs. All are now extinct. They were long classified in Struthioniform ...

, the

Palaeognathae Palaeognathae (; ) is a infraclass of birds, called paleognaths, within the class Aves of the clade Archosauria. It is one of the two extant infraclasses of birds, the other being Neognathae, both of which form Neornithes. Palaeognathae contai ...

, and the Phorusrhacidae. Phorusrhacids are an extinct group within

Cariamiformes Cariamiformes (or Cariamae) is an order of primarily flightless birds that has existed for over 60 million years. The group includes the family Cariamidae (seriemas) and the extinct families Phorusrhacidae, Bathornithidae, Idiornithidae and Ame ...

, the only living members of which are the two species of

seriemas The seriemas are the sole living members of the small bird family Cariamidae, which is also the only surviving lineage of the order Cariamiformes. Once believed to be related to cranes, they have been placed near the falcons, parrots and passeri ...

in the family Cariamidae. While they are the most speciose group within Cariamiformes, the interrelationships between phorusrhacids are unclear due to the incompleteness of their remains. Phorusrhacinae skulls

Phorusrhacids were present in South America from the

Paleocene The Paleocene, ( ) or Palaeocene, is a geological epoch (geology), epoch that lasted from about 66 to 56 mya (unit), million years ago (mya). It is the first epoch of the Paleogene Period (geology), Period in the modern Cenozoic Era (geology), E ...

(when the continent was an isolated island) and survived until 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 ...

. They also appeared in North America probably due to the Great American Biotic Interchange, and while there are records from Europe, these are disputed. It is unclear where the group originated; both cariamids and phorusrhacids may have arisen in South America, or arrived from elsewhere when southern continents were closer together or when sea levels were lower, and they may also have made reverse movements. Since phorusrhacids survived until the Pleistocene, they appear to have been more successful than for example the South American

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 ...

thylacosmilid predators (which disappeared in the

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58placental predators that invaded from North America in the Pleistocene.

Paleoenvironment

''Devincenzia'' fossils usually have poor locality records, but fossils are primarily known from the Upper Miocene,

Pliocene The Pliocene ( ; also Pleiocene) is the epoch in the geologic time scale that extends from 5.333 million to 2.58lower Pleistocene of

, though the holotype was originally speculated to be from Uruguay. Some fossils have been unearthed from the Ituzaingo Formation of

Mesopotamia, Argentina La Mesopotamia or Región Mesopotámica is the humid and verdant area of northeast Argentina, comprising the provinces of Misiones, Entre Ríos, and Corrientes Corrientes (; Guaraní language, Guaraní: Taragüí, literally: "Currents") is t ...

, which preserves vast tidal flats similar to those in the modern day

Amazon Amazon most often refers to: * Amazons, a tribe of female warriors in Greek mythology * Amazon rainforest, a rainforest covering most of the Amazon basin * Amazon River, in South America * Amazon (company), an American multinational technology c ...

and a warm climate. Large, herbivorous notoungulate mammals in the Ituzaingo Formation were widespread, including the toxodontids ''

Xotodon ''Xotodon'' is an extinct genus of toxodontid mammal that lived during the Late Miocene (Huayquerian in the SALMA classification) in Argentina, South America. Fossils of ''Xotodon'' have been found in the Ituzaingó, Maimará and Chiquimil F ...

'' and ''

Adinotherium ''Adinotherium'' (meaning "not terrible beast")Palmer (1904) p. 80. is an extinct genus of Toxodontidae, large bodied hoofed ungulates which inhabited South America during the Middle to Late Miocene, from 17.5 to 6.8 Ma and existed for approxi ...

'', smaller notoungulates such as ''

Protypotherium ''Protypotherium'' is an extinct genus of notoungulate mammals native to South America during the Oligocene and Miocene epochs. A number of closely related animals date back further, to the Eocene. Fossils of ''Protypotherium'' have been found in ...

'', smaller litopterns such as ''

Brachytherium ''Brachytherium'' is an extinct genus of proterotheriid mammal from the Late Miocene to Late Pliocene of Argentina. It is represented by the type and only species ''B. cuspidatum'', a taxon named in 1883 by Ameghino for a partial mandible with t ...

'', ''

Cullinia ''Cullinia'' is an extinct genus of litoptern, an order of South American native ungulates that included horse-like and camel-like animals such as ''Macrauchenia''. It is only known from fragmentary remains. ''Cullinia levis'' is known from Cha ...

'', ''

Diadiaphorus ''Diadiaphorus'' is an extinct genus of litoptern mammal from the Miocene of Argentina ( Ituzaingó, Pinturas, Chiquimil and Santa Cruz Formations) and Bolivia ( Nazareno Formation), South America. Description ''Diadiaphorus'' closely rese ...

Neobrachytherium ''Neobrachytherium'' is an extinct genus of proterotheriid mammal from the Late Miocene of Argentina and Uruguay. It is represented by multiple species, including the type ''N. intermedium'', originally named in 1891 by Moreno and Mercerat as a ...

, Oxyodontherium,

Paranauchenia ''Paranauchenia'' is an extinct genus of South American litopterns belonging to the family Macraucheniidae. It is known only from fossil finds in Argentina. It possessed three toes and long limbs. The species ''Paranauchenia denticulata'' lived i ...

Promacrauchenia ''Promacrauchenia'' is an extinct genus of macraucheniids that lived during the Late Miocene to Late Pliocene epochs of what is now Argentina and Bolivia. It belongs to the subfamily Macraucheniinae, which also includes ''Huayqueriana'', ''Macrau ...

Proterotherium ''Proterotherium'' (meaning "first beast") is an extinct genus of litoptern mammal of the family Proterotheriidae that lived during the Late Miocene of Argentina and Chile. Fossils of this genus have been found in the Ituzaingó Formation of Arge ...

'' and ''

Scalabrinitherium ''Scalabrinitherium'' is an extinct genus of mammals of the family Macraucheniidae. Fossils of this animal were found among the fossils of prehistoric xenarthrans in the Ituzaingó Formation of Argentina. Carnivores included the other phorusrhacid ''

'' and

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 ...

, with giant crocodilians like ''

Gryposuchus ''Gryposuchus'' is an extinct genus of gavialid crocodilian. Fossils have been found from Argentina, Colombia, Venezuela, Brazil and the Peruvian Amazon. The genus existed during the Miocene epoch (Colhuehuapian to Huayquerian). One recently d ...

'' and '' Mourasuchus'' in the freshwater.

Bamboos Bamboos are a diverse group of evergreen perennial flowering plants making up the subfamily Bambusoideae of the grass family Poaceae. Giant bamboos are the largest members of the grass family. The origin of the word "bamboo" is uncertain, but ...

coconut palms The coconut tree (''Cocos nucifera'') is a member of the palm tree family (Arecaceae) and the only living species of the genus ''Cocos''. The term "coconut" (or the archaic "cocoanut") can refer to the whole coconut palm, the seed, or the f ...

, and other palms were prevalent. The Huayquerias Formation also preserves fossils, but has much fewer preserved taxa and has had less exploration. Its paleoenvironment was hypothesized to be a gravely fluvial environment near large mountains and small lakes. The strata dates to the Upper Miocene.Garrido, A. C., Bonini, R., & Barbeau, D. L. (2017). Paleoambiente, edad y vertebrados de la Formación Huayquerías (Mioceno tardío), Provincia de Mendoza, República Argentina. XX Congreso Geológico Argentino. In ''IV Simposio del Mioceno-Plioceno del Centro y Norte de Argentina. Actas Simposio'' (Vol. 1, pp. 50-55). Fossils of litopterns and

ground sloths Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. The term is used to refer to all extinct sloths because of the large size of the earliest forms discovered, compared to existing tree sloths. The Carib ...

are the most common. The unusual carnivorous armadillo ''

Macroeuphractus ''Macroeuphractus'' is a genus of extinct armadillos from the Late Miocene to Late Pliocene of South America. The genus is noted for its large size, with ''Macroeuphractus outesi'' being the largest non- pampathere or glyptodont armadillo disc ...

'' is also known from the formation''.''Cione, A. L., Azpelicueta, M. D. L. M., Bond, M., Carlini, A. A., Casciotta, J. R., Cozzuol, M. A., ... & Vucetich, M. G. (2000). Miocene vertebrates from Entre Ríos province, eastern Argentina. ''El Neógeno de Argentina. Serie Correlación Geológica'', ''14'', 191-237. At the Campo de Robilotte locality on Lake Epecuén near

, in the

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 ...

, several fossil mammals were uncovered alongside the holotype of ''Onactornis depressus'', a synonym of ''Devincenzia'', in 1936 and many were described by Angelo Cabrera as new fossil species and even some genera, though few remain valid. Fossils of several kinds of litopterns have been found, such as the macraucheniid '' Huayqueriania,'' the proterothere ''

Eoauchenia ''Eoauchenia'' is a genus of extinct proterotheriidae, proterotheriid from the Late Miocene and Early Pliocene of Argentina. The genus was named by Ameghino in 1887 for the type species ''E. primitiva'', which was originally known from the early ...

,'' and several mesotheriids like ''

Pseudotypotherium ''Pseudotypotherium'' is an extinct genus of Notoungulates, belonging to the suborder Typotheria. It lived from the Late Miocene to the Late Pliocene, and its fossilized remains were discovered in South America. Description This animal, like a ...

'' and ''

Typotheriopsis ''Typotheriopsis'' is an extinct genus of Notoungulate, belonging to the family Mesotheriidae, which included several small sized Meridiungulates specialized in digging. It is considered as the sister taxon of the clade including ''Mesotherium'' ...

.'' Many

xenarthrans Xenarthra (; from Ancient Greek wikt:ξένος, ξένος, xénos, "foreign, alien" + wikt:ἄρθρον, ἄρθρον, árthron, "joint") is a major clade of Placentalia, placental mammals native to the Americas. There are 31 living species: ...

fossils were unearthed and described by Cabrera including the type specimen of the glyptodont '' Coscinocercus'' and the type specimen of the

ground sloth Ground sloths are a diverse group of extinct sloths in the mammalian superorder Xenarthra. The term is used to refer to all extinct sloths because of the large size of the earliest forms discovered, compared to existing tree sloths. The Caribbe ...

s, ''

Proscelidodon ''Proscelidodon'' is an extinct genus of ground sloths in the family Scelidotheriidae. It lived during the Miocene and Pliocene of what is now Argentina and Bolivia. The genus was described in 1935. Taxonomy During the Quaternary the taxonomic ...

'' and ''

Mcdonaldocnus ''Mcdonaldocnus'' is an extinct genus of nothrotheriid ground sloths that lived during the Middle Miocene and Early Pliocene of what is now Bolivia and Argentina. It was originally placed in the genus '' Xyophorus'' but was subsequently recognize ...

''.

References

{{Taxonbar, from=Q4895333 Phorusrhacinae Extinct flightless birds Miocene birds of South America Pliocene birds of South America Deseadan Montehermosan Neogene Argentina Fossils of Argentina Ituzaingó Formation Neogene Uruguay Fossils of Uruguay Paraná Basin Fossil taxa described in 1932