''Unenlagia'' (meaning "half-bird" in
Latin
Latin ( or ) is a classical language belonging to the Italic languages, Italic branch of the Indo-European languages. Latin was originally spoken by the Latins (Italic tribe), Latins in Latium (now known as Lazio), the lower Tiber area aroun ...
ized
Mapudungun
Mapuche ( , ; from 'land' and 'people', meaning 'the people of the land') or Mapudungun (from 'land' and 'speak, speech', meaning 'the speech of the land'; also spelled Mapuzugun and Mapudungu) is either a language isolate or member of the s ...
) is a
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 ...
of
dromaeosaurid
Dromaeosauridae () is a family (biology), family of feathered coelurosaurian Theropoda, theropod dinosaurs. They were generally small to medium-sized feathered carnivores that flourished in the Cretaceous period (geology), Period. The name Drom ...
theropod
Theropoda (; from ancient Greek , (''therion'') "wild beast"; , (''pous, podos'') "foot"">wiktionary:ποδός"> (''pous, podos'') "foot" is one of the three major groups (clades) of dinosaurs, alongside Ornithischia and Sauropodom ...
dinosaur
Dinosaurs are a diverse group of reptiles of the clade Dinosauria. They first appeared during the Triassic Geological period, period, between 243 and 233.23 million years ago (mya), although the exact origin and timing of the #Evolutio ...
that lived 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 ...
during the
Late Cretaceous
The Late Cretaceous (100.5–66 Ma) is the more recent of two epochs into which the Cretaceous Period is divided in the geologic time scale. Rock strata from this epoch form the Upper Cretaceous Series. The Cretaceous is named after ''cre ...
period.
The genus ''Unenlagia'' has been assigned two species: ''U. comahuensis'', the
type species
In International_Code_of_Zoological_Nomenclature, 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 spe ...
described by Novas and Puerta in 1997,
and ''U. paynemili'', described by Calvo ''et al.'' in 2004.
Discovery and naming

In 1996 in the
Neuquén
Neuquén (; ) is the capital city of the Argentine province of Neuquén and of the Confluencia Department, located in the east of the province. It occupies a strip of land west of the confluence of the Limay and Neuquén rivers which form t ...
province of Argentina a skeleton of a theropod was discovered in the
Sierra del Portezuelo and reported the same year. In 1997
Fernando Emilio Novas
Fernando Emilio Novas (born 1960) is an Argentina, Argentine Paleontology, paleontologist working for the Comparative Anatomy Department of the Bernardino Rivadavia Natural Sciences Museum in Buenos Aires, Argentina. and
Pablo Puerta named and described ''Unenlagia comahuensis''. The generic name is derived from
Mapuche
The Mapuche ( , ) also known as Araucanians are a group of Indigenous peoples of the Americas, Indigenous inhabitants of south-central Chile and southwestern Argentina, including parts of Patagonia. The collective term refers to a wide-ranging e ...
''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 Specific name may refer to:
* in Database management systems, a system-assigned name that is unique within a particular database
In taxonomy, either of these two meanings, each with its own set of rules:
* Specific name (botany), the two-part (bino ...
refers to the
Comahue
Comahue is a sub-region of Argentina slightly to the south of the country's centre that covers the northern part of Argentine Patagonia and includes the provinces of Argentina, provinces of Neuquén Province, Neuquén and Río Negro Province, Rí ...
, the region the find was made.
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 ...
specimen, MCF PVPH 78, was uncovered in layers of the
Portezuelo Formation
The Portezuelo Formation is a geologic formation of Late Cretaceous (Turonian, Late Turonian to Coniacian, Early Coniacian) age, outcropping in the Mendoza Province, Mendoza, Río Negro Province, Río Negro and Neuquén Province, Neuquén provinc ...
dating to the
Coniacian
The Coniacian is an age or stage in the geologic timescale. It is a subdivision of the Late Cretaceous Epoch or Upper Cretaceous Series and spans the time between 89.8 ± 1 Ma and 86.3 ± 0.7 Ma (million years ago). The Coniacian is preceded by ...
. 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.
In 2002 near the
Lago Barreales a second skeleton was uncovered and reported in 2003. In 2004 it was named and described by
Jorge Calvo,
Juan Porfiri and
Alexander Kellner
Alexander Wilhelm Armin Kellner (born September 26, 1961) is a Brazilian geologist and paleontologist who is a leading expert in the field of studying pterosaurs. His research has focused mainly on fossil reptiles from the Cretaceous Period, i ...
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
paratype
In zoology and botany, a paratype is a specimen of an organism that helps define what the scientific name of a species and other taxon actually represents, but it is not the holotype (and in botany is also neither an isotype (biology), isotype ...
s were also assigned: MUCPv-343, a claw; MUCPv-409, a partial ilium; MUCPv-415, a phalanx and MUCPv-416, a vertebra.
Some researchers consider ''
Neuquenraptor
''Neuquenraptor'' (meaning Neuquén thief) 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 ...
'' to be a
junior subjective synonym
In taxonomy, the scientific classification of living organisms, a synonym is an alternative scientific name for the accepted scientific name of a taxon. The botanical and zoological codes of nomenclature treat the concept of synonymy differently.
...
of ''Unenlagia''.
Description

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.
Estimates have thus varied between a length of and a weight of on the one hand, and a length of just on the other.
However, Thomas Holtz has estimated ''Unenlagia'' at long and 9.1-22.7 kg (20-50 lbs) in weight. The describers of ''Austroraptor'' estimated that ''Unenlagia'' reached over long and weighed about . 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
''Buitreraptor'' (meaning "La Buitrera seizer") 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 g ...
'' and ''
Austroraptor
''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.
''Austroraptor'' was a large, moderately-built, ground-dwelling, ...
''.

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
''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird'') is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaîos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
''.
The shoulder girdle of ''Unenlagia'' was originally interpreted as if it was adapted for flapping, with a flat
scapula
The scapula (: scapulae or scapulas), also known as the shoulder blade, is the bone that connects the humerus (upper arm bone) with the clavicle (collar bone). Like their connected bones, the scapulae are paired, with each scapula on either side ...
positioned on top of the ribcage, making the shoulder joint point more laterally.
However, in 2002
Kenneth Carpenter
Kenneth Carpenter (born 21 September 1949) is an American paleontologist. He is the former director of the USU Eastern Prehistoric Museum and author or co-author of books on dinosaurs and Mesozoic life. His main research interests are armore ...
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. 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
''Archaeopteryx'' (; ), sometimes referred to by its German name, "" ( ''Primeval Bird'') is a genus of bird-like dinosaurs. The name derives from the ancient Greek (''archaîos''), meaning "ancient", and (''ptéryx''), meaning "feather" ...
''.
[Senter, P. (2006). "Scapular orientation in theropods and basal birds, and the origin of flapping flight." ''Acta Palaeontologica Polonica'', 51(2): 305–31]
PDF fulltext
/ref> South American workers have remained unconvinced though, countering that a laterally positioned scapula would make the coracoid
A coracoid is a paired bone which is part of the shoulder assembly in all vertebrates except therian mammals (marsupials and placentals). In therian mammals (including humans), a coracoid process is present as part of the scapula, but this is n ...
of ''Unenlagia'' jut into its ribcage, which seems anatomically implausible.
Classification
Novas and Puerta in the original description considered ''Unenlagia'' a sister taxon
In phylogenetics, a sister group or sister taxon, also called an adelphotaxon, comprises the closest relative(s) of another given unit in an evolutionary tree.
Definition
The expression is most easily illustrated by a cladogram:
Taxon A and ...
of the birds, which would make it a basal member of the Avialae
Avialae ("bird wings") is a clade containing the only living dinosaurs, the birds, and their closest relatives. It is usually defined as all theropod dinosaurs more closely related to birds (Aves) than to Deinonychosauria, deinonychosaurs, though ...
''sensu'' Gauthier 1986.
However, in 1999 Mark Norell
Mark Allen Norell (born July 26, 1957) is an American vertebrate paleontologist. He is currently the chairman of paleontology and a research associate at the American Museum of Natural History. He is best known as the discoverer of the first ...
and Peter Makovicky
Peter may refer to:
People
* List of people named Peter, a list of people and fictional characters with the given name
* Peter (given name)
** Saint Peter (died 60s), apostle of Jesus, leader of the early Christian Church
* Peter (surname), a sur ...
stated that ''Unenlagia'' was a member of the Dromaeosauridae
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 ...
. ''Unenlagia'' would have belonged to the extremely bird-like Gondwana
Gondwana ( ; ) was a large landmass, sometimes referred to as a supercontinent. The remnants of Gondwana make up around two-thirds of today's continental area, including South America, Africa, Antarctica, Australia (continent), Australia, Zea ...
n clade
In biology, a clade (), also known as a Monophyly, monophyletic group or natural group, is a group of organisms that is composed of a common ancestor and all of its descendants. Clades are the fundamental unit of cladistics, a modern approach t ...
of dromaeosaurids called the Unenlagiinae
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 Halszkar ...
, and be closely related to species such as ''Buitreraptor
''Buitreraptor'' (meaning "La Buitrera seizer") 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 g ...
'' and ''Neuquenraptor
''Neuquenraptor'' (meaning Neuquén thief) 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 ...
'' (which might be the same species as ''Unenlagia''). Makovicky ''et al.'' in 2005 also recovered the 'flying raptor' ''Rahonavis
''Rahonavis'' is a genus of bird-like theropod from the Late Cretaceous (Maastrichtian, from about 72.1 to 66 mya) of what is now northwestern Madagascar. It is known from a partial skeleton ( UA 8656) found by Catherine Forster and colleagues ...
'' 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. Norell ''et al.'' in 2006 also found ''Rahonavis'' to be the sister taxon of ''Unenlagia''.
This position has not been unanimously accepted. In 2011 Novas and Federico Agnolín
Federico (; ) is a given name and surname. It is a form of Frederick, most commonly found in Spanish, Portuguese and Italian.
People with the given name Federico
Arts and language
* Federico Ágreda, Venezuelan composer and DJ
* Federico Aguil ...
published a study in which a Unenlagiidae
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 g ...
was positioned within Avialae.
Below is a cladogram conducted by Hartman et al. 2019, in which ''Unenlagia'' is found to be a more derived member alongside ''Dakotaraptor
''Dakotaraptor'' (meaning “thief from Dakota”) is a possible chimaeric genus of maniraptoriform theropod dinosaur that lived in western North America during the Late Cretaceous period. The remains have been found in the Maastrichtian-aged ...
'':
This cladogram is from Motta ''et al.'', 2020, showing ''Unenlagia'' being more derived than traditional dromaeosaurids:
See also
* Timeline of dromaeosaurid research
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 ...
* Origin of birds
The scientific question of which larger group of animals birds evolved within has traditionally been called the "origin of birds". The present scientific consensus is that birds are a group of maniraptoran theropod dinosaurs that originated du ...
''Unenlagia'' in the Dino Directory
References
{{Taxonbar, from=Q134159
Unenlagiinae
Dinosaur genera
Coniacian dinosaurs
Portezuelo Formation
Taxa named by Fernando Novas
Fossil taxa described in 1997
Dinosaurs of Argentina