Unenlagiines
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Unenlagiinae is a subfamily of long-snouted
paravian 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 av ...
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 ...
s. They are traditionally considered to be members of
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 ...
, though some authors place them into their own family,
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 ...
, sometimes alongside the subfamily
Halszkaraptorinae Halszkaraptorinae is an extinct basal ("primitive") subfamily of Dromaeosauridae (or possibly Unenlagiidae) that includes the enigmatic genera '' Halszkaraptor'', '' Natovenator'', ''Mahakala'', and '' Hulsanpes''. Halszkaraptorines are definit ...
. Definitive members are known from 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 ...
of
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 ...
, though some researchers include taxa from other continents within this subfamily based on phylogenetic analyses. Two probable unenlagiine specimens (NMV P257601, NMV P180889) from the upper
Strzelecki Group The Halibut Field is an oil field, within the Gippsland Basin offshore of the Australian state of Victoria (Australia), Victoria. The oil field is located approximately 64 km offshore of southeastern Australia. The total area of this field ...
(
Aptian The Aptian is an age (geology), age in the geologic timescale or a stage (stratigraphy), stage in the stratigraphic column. It is a subdivision of the Early Cretaceous, Early or Lower Cretaceous epoch (geology), Epoch or series (stratigraphy), S ...
) and
Eumeralla Formation The Eumeralla Formation is a geological formation in Victoria, Australia whose strata date back to the Early Cretaceous. It is Aptian to Albian in age. Dinosaur remains are among the fossils that have been recovered from the formation, particul ...
(lower
Albian The Albian is both an age (geology), age of the geologic timescale and a stage (stratigraphy), stage in the stratigraphic column. It is the youngest or uppermost subdivision of the Early Cretaceous, Early/Lower Cretaceous epoch (geology), Epoch/s ...
) of
Australia Australia, officially the Commonwealth of Australia, is a country comprising mainland Australia, the mainland of the Australia (continent), Australian continent, the island of Tasmania and list of islands of Australia, numerous smaller isl ...
might potentially extend their known fossil range to the
Early Cretaceous The Early Cretaceous (geochronology, geochronological name) or the Lower Cretaceous (chronostratigraphy, chronostratigraphic name) is the earlier or lower of the two major divisions of the Cretaceous. It is usually considered to stretch from 143.1 ...
, and ''
Kakuru ''Kakuru'' is a Nomen dubium, dubious genus of theropod dinosaur from the early Cretaceous Period. Discovery The only described species, ''Kakuru kujani'', is known primarily from evidence of a single tibia, which had been fossilised through a ...
,'' which is considered a maniraptoran, and might be an unenlagiine as well. ''
Imperobator ''Imperobator'' ("powerful warrior") is a genus of probable unenlagiid Paraves, paravian theropod dinosaurs, that lived during the Maastrichtian age of the Late Cretaceous in what is now James Ross Island in Antarctica. ''Imperobator'' is one of ...
'' from the Late Cretaceous of Antarctica, previously considered enigmatic, has also been recently interpreted as an unenlagiine.


Description

Most unenlagiines have been discovered 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 ...
. The largest was ''
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, ...
'', which measured up to 5–6 m (16.4–19.7 ft) in length, making it also one of the largest dromaeosaurids. The subfamily is distinguished from other dromaeosaurids by a tail stiffened by lengthy chevrons and superior processes, a reduced second pedal ungual, a posteriorly oriented pubis and very elongated snouts. Unenlagiines also had elongated, slender hindlimbs with a subarctometatarsalian metatarsus, which is characterized by the pinched metatarsal III at the upper end. Their distinct anatomy from Laurasian dromaeosaurids was likely a consequence of the breakup of
Pangaea Pangaea or Pangea ( ) was a supercontinent that existed during the late Paleozoic and early Mesozoic eras. It assembled from the earlier continental units of Gondwana, Euramerica and Siberia during the Carboniferous period approximately 335 mi ...
into
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 ...
and Laurasia, where the geological isolation of unenlagiines from their relatives resulted in
allopatric speciation Allopatric speciation () – also referred to as geographic speciation, vicariant speciation, or its earlier name the dumbbell model – is a mode of speciation that occurs when biological populations become geographically isolated from ...
.


Classification

During the description of '' Halszkaraptor'' in 2017, Cau et al. published a phylogenetic analysis 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 ...
, in which, members of the Unenlagiinae are classified as: In 2019, during the description of ''
Hesperornithoides ''Hesperornithoides'' (meaning "western bird form"; nicknamed "Lori") is a genus of troodontid theropod dinosaur that lived in North America during the Late Jurassic period. Discovery Although several Troodontid teeth were found, with the trood ...
'', many
Paravian 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 av ...
groups were examined for the inclusion of this new
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 ...
, including the Unenlagiinae. The analysis ended in the inclusion of ''
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 ...
'', ''
Pyroraptor ''Pyroraptor'' (meaning "fire thief") 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 earl ...
'' and ''
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 ...
'' to the Unenlagiinae.


Paleobiology

A study performed by Gianechini and colleagues in 2020 indicates that unenlagiine dromaeosaurids of Gondwana possessed different hunting specializations than the eudromaeosaurs from Laurasia. The shorter second
phalanx The phalanx (: phalanxes or phalanges) was a rectangular mass military formation, usually composed entirely of heavy infantry armed with spears, pikes, sarissas, or similar polearms tightly packed together. The term is particularly used t ...
in the second digit of the foot of eudromaeosaurs allowed for increased force to be generated by that digit, which, combined with a shorter and wider
metatarsus The metatarsal bones or metatarsus (: metatarsi) are a group of five long bones in the midfoot, located between the tarsal bones (which form the heel and the ankle) and the phalanges ( toes). Lacking individual names, the metatarsal bones are ...
, and a noticeable marked hinge‐like morphology of the articular surfaces of metatarsals and phalanges, possibly allowed eudromaeosaurs to exert a greater gripping strength than unenlagiines, allowing for more efficient subduing and killing of large prey. In comparison, the unenlagiine dromaeosaurids possess a longer and slender sub
arctometatarsus An arctometatarsalian organism is one in which the proximal part of the middle metatarsal is pinched between the surrounding metatarsals. The trait appears to be highly homoplastic, common in certain sorts of dinosaurs accustomed to running (among ...
, and less well‐marked hinge joints, a trait that possibly gave them greater
cursorial A cursorial organism is one that is adapted specifically to run. An animal can be considered cursorial if it has the ability to run fast (e.g. cheetah) or if it can keep a constant speed for a long distance (high endurance). "Cursorial" is often ...
capacities and allowed for greater speed than eudromaeosaurs. Additionally, the longer second phalanx of the second digit allowed unenlagiines fast movements of their feet's second digits to hunt smaller, faster types of prey. These differences in locomotor and predatory specializations may have been a key feature that influenced the evolutionary paths that shaped both groups of dromaeosaurs in the northern and southern hemispheres, respectively.


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 ...
*
Halszkaraptorinae Halszkaraptorinae is an extinct basal ("primitive") subfamily of Dromaeosauridae (or possibly Unenlagiidae) that includes the enigmatic genera '' Halszkaraptor'', '' Natovenator'', ''Mahakala'', and '' Hulsanpes''. Halszkaraptorines are definit ...


References

{{Taxonbar, from=Q1258292 Dinosaur subfamilies Late Cretaceous dinosaurs