Unenlagia Temporal range: Late Cretaceous ~ | |
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Illustration of known material of U. comahuensis | |
Scientific classification ![]() | |
Domain: | Eukaryota |
Kingdom: | Animalia |
Phylum: | Chordata |
Clade: | Dinosauria |
Clade: | Saurischia |
Clade: | Theropoda |
Family: | † Dromaeosauridae |
Subfamily: | † Unenlagiinae |
Genus: | † Unenlagia Novas & Puerta, 1997 |
Species | |
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Unenlagia (meaning "half-bird" in Latinized Mapudungun) is a genus of dromaeosaurid theropod dinosaur that lived in South America during the Late Cretaceous period. [1] The genus Unenlagia has been assigned two species: U. comahuensis, the type species described by Novas and Puerta in 1997, [1] and U. paynemili, described by Calvo et al. in 2004. [2]
In 1996 in the Neuquén province of Argentina a skeleton of a theropod was discovered in the Sierra del Portezuelo and reported the same year. [3] In 1997 Fernando Emilio Novas and Pablo Puerta named and described Unenlagia comahuensis. The generic name is derived from Mapuche uñùm, 'bird', and llag, 'half', in reference to the fact that the describers considered the species to be a link between birds and more basal theropods. The specific name refers to the Comahue, the region the find was made. [1]
The holotype specimen, MCF PVPH 78, was uncovered in layers of the Portezuelo Formation dating to the Coniacian. It consists of a partial skeleton lacking the skull but including vertebrae, a sacrum, ribs, chevrons, a scapula, a humerus, a partial pelvis, a femur and a tibia. [1]
In 2002 near the Lago Barreales a second skeleton was uncovered and reported in 2003. [4] In 2004 it was named and described by Jorge Calvo, Juan Porfiri and Alexander Kellner as a second species: Unenlagia paynemili. The specific name honours Maximino Paynemil, the chief of the Paynemil community. The holotype is MUCPv-349, a partial skeleton consisting of a humerus and two pubes. Several paratypes were also assigned: MUCPv-343, a claw; MUCPv-409, a partial ilium; MUCPv-415, a phalanx and MUCPv-416, a vertebra. [2]
Some researchers consider Neuquenraptor to be a junior subjective synonym of Unenlagia. [5]
The body length of Unenlagia has been disputed, due to the fact that only the leg length is well known and it is uncertain whether this should be extrapolated using the proportions of the low-slung Dromaeosauridae or the long-legged basal birds. [1] Estimates have thus varied between a length of 3.5 m (11 ft) and a weight of 75 kg (165 lb) on the one hand, [6] and a length of just 2 m (6.6 ft) on the other. [7] However, Thomas Holtz has estimated Unenlagia at 2.3 m (7.5 ft) long and 9.1-22.7 kg (20-50 lbs) in weight. [8] The describers of Austroraptor estimated that Unenlagia reached over 3.2 metres (10 ft) long and weighed about 95 kilograms (209 lb). [9] Likewise, the interpretation of the head form has changed from a shorter-snouted dromaeosaurid condition to the elongated shape known from the later discovered related genera Buitreraptor and Austroraptor .
Novas and Puerta found that the pelvic region of Unenlagia, especially the form of the ilium, was very similar to that of the early bird Archaeopteryx . [1] The shoulder girdle of Unenlagia was originally interpreted as if it was adapted for flapping, with a flat scapula positioned on top of the ribcage, making the shoulder joint point more laterally. [1] However, in 2002 Kenneth Carpenter pointed out that this would imply that the shoulder-blade was dorsoventrally flattened instead of laterally as with other theropods and that it thus were more likely the scapula was located on the side of the ribcage. [10] This conformed to a later hypothesis by Philip Senter that non-avian theropods like Unenlagia were unable to lift their forelimbs above their back, as even would still have been the case for the basalmost bird Archaeopteryx . [11] South American workers have remained unconvinced though, countering that a laterally positioned scapula would make the coracoid of Unenlagia jut into its ribcage, which seems anatomically implausible. [7]
Novas and Puerta in the original description considered Unenlagia a sister taxon of the birds, which would make it a basal member of the Avialae sensu Gauthier 1986.
However, in 1999 Mark Norell and Peter Makovicky stated that Unenlagia was a member of the Dromaeosauridae. [12] [13] Unenlagia would have belonged to the extremely bird-like Gondwanan clade of dromaeosaurids called the Unenlagiinae, and be closely related to species such as Buitreraptor and Neuquenraptor (which might be the same species as Unenlagia). Makovicky et al. in 2005 also recovered the 'flying raptor' Rahonavis as a member of this group, which would mean that either Unenlagia is secondarily flightless, having evolved from flying, Rahonavis-like ancestors, or that bird-like flight evolved at least twice. [14] Norell et al. in 2006 also found Rahonavis to be the sister taxon of Unenlagia. [15]
This position has not been unanimously accepted. In 2011 Novas and Federico Agnolín published a study in which a Unenlagiidae was positioned within Avialae. [16]
Below is a cladogram conducted by Hartman et al. 2019, in which Unenlagia is found to be a more derived member alongside Dakotaraptor : [17]
This cladogram is from Motta et al., 2020, showing Unenlagia being more derived than traditional dromaeosaurids: [18]
Dromaeosauridae is a family of feathered coelurosaurian theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous Period. The name Dromaeosauridae means 'running lizards', from Greek δρομαῖος (dromaîos), meaning 'running at full speed', 'swift', and σαῦρος (saûros), meaning 'lizard'. In informal usage, they are often called raptors, a term popularized by the film Jurassic Park; several genera include the term "raptor" directly in their name, and popular culture has come to emphasize their bird-like appearance and speculated bird-like behavior.
Neuquenraptor is a genus of dromaeosaurid theropod dinosaurs that lived in South America during the Late Cretaceous in what is now the Portezuelo Formation of Argentina. It is one of the first dromaeosaurids found in the Southern Hemisphere.
Buitreraptor is a genus of dromaeosaurid dinosaurs that lived during the Late Cretaceous of Argentina at the Candeleros Formation. Buitreraptor was described in 2005 and the type species is Buitreraptor gonzalezorum. It was rooster-sized and had a very elongated head with many small teeth.
Adasaurus is a genus of dromaeosaurid dinosaur that lived in Asia during the Late Cretaceous period about 70 million years ago. The genus is known from two partial specimens found in the Nemegt Formation of Mongolia that were partially described in 1983 by the paleontologist Rinchen Barsbold.
Rahonavis is a genus of bird-like theropod from the Late Cretaceous of what is now northwestern Madagascar. It is known from a partial skeleton found by Catherine Forster and colleagues in Maevarano Formation rocks at a quarry near Berivotra, Mahajanga Province. Rahonavis was a small predator, at about 70 centimetres (2.3 ft) long and 0.45-2.27 kg, with the typical dromaesaurid-like raised sickle claw on the second toe. It was originally the first African coelurosaur until the discovery of Nqwebasaurus in 2000.
Megaraptor is a genus of large theropod dinosaur that lived in the ages of the Late Cretaceous. Its fossils have been discovered in the Patagonian Portezuelo Formation of Argentina, South America. Initially thought to have been a giant dromaeosaur-like coelurosaur, it was classified as a neovenatorid allosauroid in previous phylogenies, but more recent phylogeny and discoveries of related megaraptoran genera has placed it as either a basal tyrannosauroid or a basal coelurosaur with some studies still considering it a neovenatorid.
Pyroraptor is an extinct genus of paravian dinosaur, probably a dromaeosaurid or unenlagiid, from the Late Cretaceous Ibero-Armorican island, of what is now southern France. It lived during the late Campanian and early Maastrichtian stages, approximately 72 million years ago. It is known from a single partial specimen that was found in Provence in 1992, after a forest fire. The animal was named Pyroraptor olympius by Allain and Taquet in 2000.
Patagonykus is a genus of theropod dinosaur from the Upper Cretaceous of Argentina. This alvarezsauroid was discovered in exposures of the Portezuelo Formation (Turonian-Coniacian) of the Rio Neuquén Subgroup in the Neuquén Basin, Neuquen Province of Patagonia, Argentina. The holotype consists of an incomplete but well-preserved skeleton, lacking a skull, but including many vertebrae, the coracoids, a partial forelimb, pelvic girdle, and hindlimbs. Patagonykus has been classed with the Alvarezsauridae, a family which includes such taxa as the Mongolian Mononykus and the Argentinian Alvarezsaurus. In 2010 Gregory S. Paul estimated its length at 1 meter and its weight at 3.5 kg.
The Huincul Formation is a geologic formation of Late Cretaceous age of the Neuquén Basin that outcrops in the Mendoza, Río Negro and Neuquén Provinces of northern Patagonia, Argentina. It is the second formation in the Río Limay Subgroup, the oldest subgroup within the Neuquén Group. Formerly that subgroup was treated as a formation, and the Huincul Formation was known as the Huincul Member.
The Portezuelo Formation is a geologic formation of Late Cretaceous age, outcropping in the Mendoza, Río Negro and Neuquén provinces of Argentina. It is the fourth-oldest formation in the Neuquén Group and the older of the two formations in the Río Neuquén Subgroup. Formerly, that subgroup was treated as a formation, and the Portezuelo Formation was known as the Portezuelo Member.
Unenlagiinae is a subfamily of long-snouted paravian theropods. They are traditionally considered to be members of Dromaeosauridae, though some authors place them into their own family, Unenlagiidae, sometimes alongside the subfamily Halszkaraptorinae. Definitive members are known from South America, though some researchers include taxa from other continents within this subfamily based on phylogenetic analyses.
Velociraptorinae is a subfamily of the theropod group Dromaeosauridae. Definitive fossils attributed to the subfamily have only been found in the Late Cretaceous deposits of Asia, with Kansaignathus as the basalmost member. While numerous taxa from North America are previously assigned to the velociraptorines, they are now reclassified within different lineages of Eudromaeosauria. Several teeth that may belong to indeterminate velociraptorines have also been discovered in Germany, dating to the Kimmeridgian stage of the Late Jurassic.
Shanag is a genus of paravian theropod dinosaur from the Early Cretaceous Period of Mongolia. It may be a dromaeosaurid, but some researchers are skeptical of this classification. The type species is S. ashile.
Paraves are a widespread group of theropod dinosaurs that originated in the Middle Jurassic period. In addition to the extinct dromaeosaurids, troodontids, anchiornithids, and possibly the scansoriopterygids, the group also contains the avialans, which include diverse extinct taxa as well as the over 10,000 species of living birds. Basal members of Paraves are well known for the possession of an enlarged claw on the second digit of the foot, which was held off the ground when walking in some species. A number of differing scientific interpretations of the relationships between paravian taxa exist. New fossil discoveries and analyses make the classification of Paraves an active subject of research.
Austroraptor is a genus of dromaeosaurid theropod dinosaur that lived during the Campanian and Maastrichtian ages of the Late Cretaceous period in what is now Argentina.
Tianyuraptor is a genus of short-armed dromaeosaurid dinosaur that lived during the Early Cretaceous, about 122 million years ago. Its remains have been found in western Liaoning, China. It was similar to other dromaeosaurids found in Liaoning, with the exception of being somewhat more primitive. The type specimen, formally named in 2009, shows features not seen in previously known Northern Hemisphere (Laurasian) dromaeosaurids, but present in Southern Hemisphere (Gondwanan) species and early birds. Because of this, the scientists who first studied Tianyuraptor described it as a "transitional species", bridging the gap between northern and southern types of dromaeosaurid. Tianyuraptor also differs from previously known dromaeosaurids in that it possesses a relatively small furcula ("wishbone"), and unusually short forelimbs.
Pamparaptor is an extinct genus of paravian theropod dinosaur from the Late Cretaceous Portezuelo Formation of the Neuquén province in Argentine Patagonia. Its precise classification is uncertain, but the authors who described this taxon have argued that it is a dromaeosaurid. The genus contains a single species, P. micros, which is known from a single specimen consisting of a mostly complete and fully-articulated left foot, which preserves the iconic dromaeosaur-like "killing claw".
This timeline of dromaeosaurid research is a chronological listing of events in the history of paleontology focused on the dromaeosaurids, a group of sickle-clawed, bird-like theropod dinosaurs including animals like Velociraptor. Since the Native Americans of Montana used the sediments of the Cloverly Formation to produce pigments, they may have encountered remains of the dromaeosaurid Deinonychus hundreds of years before these fossils came to the attention of formally trained scientists.
Unenlagiidae is a proposed family of eumaniraptoran paravians that includes the subfamilies Unenlagiinae and possibly Halszkaraptorinae. Fossils of both subfamilies have been found in both Gondwanan and Laurasian deposits. The biology of the group suggests that some members were semiaquatic specialists.
Overoraptor is an extinct genus of paravian theropod of uncertain affinities from the Late Cretaceous Huincul Formation of Argentinian Patagonia. The genus contains a single species, O. chimentoi, known from several bones of the hands, feet, and hips alongside some vertebrae.