Eudromaeosauria

Eudromaeosauria ("true dromaeosaurs") is a subgroup of terrestrial dromaeosaurid theropod dinosaurs. They were relatively large-bodied, feathered hypercarnivores (with diets consisting almost entirely of other terrestrial vertebrates) that flourished in the Cretaceous Period.

Eudromaeosaur fossils are known almost exclusively from the northern hemisphere. They first appeared in the early Cretaceous Period (early Aptian stage, about 124 million years ago) and survived until the end of the Cretaceous (Maastrichtian stage, 66 Ma). The earliest known definitive eudromaeosaur is the dromaeosaurine Utahraptor ostrommaysorum, from the Cedar Mountain Formation, dated to 124 million years ago.[1] However, the earlier (143-million-year-old) fossils such as those of Nuthetes destructor and several indeterminate teeth dating to the Kimmeridgian stage may represent eudromaeosaurs.[2][3]

Eudromaeosauria
Temporal range: Early CretaceousLate Cretaceous, 143–66 Ma
Likely Kimmeridgian record
Velociraptor Wyoming Dinosaur Center
Mounted cast of a Velociraptor mongoliensis skeleton, Wyoming Dinosaur Center
Scientific classification
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Order: Saurischia
Suborder: Theropoda
Family: Dromaeosauridae
Clade: Eudromaeosauria
Longrich & Currie, 2009
Subgroups

See text.

Description

While other dromaeosaurids filled a variety of specialized ecological niches, mainly those of small predators or larger fish-eating forms, eudromaeosaurs functioned as large-bodied predators of often medium to large-sized prey. Aside from their generally larger size, eudromaeosaurs are characterized by several features of the foot. First, there were differences in the positions of the grooves which anchored blood vessels and keratin sheathes of the toe claws. In primitive dromaeosaurids like Hesperonychus, these grooves ran parallel to each other on either side of the claw along its length. In eudromaeosaurs, the grooves were asymmetrical, with the inner one split into two distinct grooves and elevated toward the top of the claw, while the single outer groove remained positioned at the midline.[4]

The second distinguishing characteristic of eudromaeosaurs is an expanded and enlarged "heel" on the last bone in the second toe (phalange), which bore the enlarged, sickle-like toe claw. Finally, the first bone of the second toe also possessed an enlarged expansion at the joint, another adaptation relating to the unusually enlarged claw, and which helped the animal hold the claw high off the ground. Also unlike their more basal relatives, the sickle-claw of eudromaeosaurs was sharper and more blade-like. In unenlagiines and microraptorines, the claw is broader at its base.[4]

Classification

Eudromaeosauria was first defined as a node-based clade by Nick Longrich and Philip J. Currie in 2009, as the most inclusive natural group containing Dromaeosaurus, Velociraptor, Deinonychus, and Saurornitholestes, their most recent common ancestor and all of its other descendants. The various "subfamilies" have also been re-defined as clades, usually defined as all species closer to the groups namesake than to Dromaeosaurus or any namesakes of other sub-clades.[4]

The subgroups of Eudromaeosauria frequently shift in content based on new analysis, but typically consist of the following groups. For example, the subfamily Velociraptorinae has traditionally included Velociraptor, Deinonychus, and Saurornitholestes, and while the discovery of Tsaagan lent support to this grouping, the inclusion of Saurornitholestes is still uncertain. The Dromaeosaurinae is usually found to consist of medium to giant-sized species, with generally box-shaped skulls (the other subfamilies generally have narrower snouts). A number of eudromaeosaurs have not been assigned to any particular subfamily, because they are too poorly preserved to be placed confidently in phylogenetic analysis (see section Phylogeny below).[5]

The following classification of the various genera of eudromaeosaurs follows the table provided in Holtz, 2010 unless otherwise noted.[6]

Relationships

The below cladogram follows an analysis by Evans et al. in 2013. Their analysis used an updated version of the dataset originally compiled by Nick Longrich and Phil Currie to study dromaeosaurid relationships, and found a relatively traditional arrangement of eudromaeosaurian relationships.[7]

Eudromaeosauria
Saurornitholestinae

Bambiraptor feinbergi

Saurornitholestes langstoni

Atrociraptor marshalli

Deinonychus antirrhopus

 Dromaeosaurinae 

Achillobator giganticus

Balaur bondoc

Dromaeosaurus albertensis

Utahraptor ostrommaysorum

 Velociraptorinae 

IGN 100/23

Acheroraptor temertyorum

Velociraptor mongoliensis

Adasaurus mongoliensis

Tsaagan mangas

Velociraptor osmolskae

The cladogram below follows a 2015 analysis by paleontologists Robert DePalma, David Burnham, Larry Martin, Peter Larson, and Robert Bakker, using updated data from the Theropod Working Group. This analysis concluded that several members of traditional eudromaeosaurian subgroups are actually outside of that group, and that traditional groupings like Velociraptorinae are paraphyletic.[8]

Dromaeosauridae

Unenlagiinae

Microraptoria

Bambiraptor

Tianyuraptor

Adasaurus

Tsaagan

Eudromaeosauria

Saurornitholestes

Velociraptor

Dromaeosaurinae

Deinonychus

Atrociraptor

Achillobator

Utahraptor

Dakotaraptor

Dromaeosaurus

See also

References

  1. ^ McDonald AT, Kirkland JI, DeBlieux DD, Madsen SK, Cavin J, et al. (2010). "New Basal Iguanodonts from the Cedar Mountain Formation of Utah and the Evolution of Thumb-Spiked Dinosaurs". PLoS ONE. 5 (11): e14075. doi:10.1371/journal.pone.0014075. PMC 2989904. PMID 21124919.
  2. ^ Sweetman S.C. (2004). "The first record of velociraptorine dinosaurs (Saurischia, Theropoda) from the Wealden (Early Cretaceous, Barremian) of southern England". Cretaceous Research. 25 (3): 353–364. doi:10.1016/j.cretres.2004.01.004.
  3. ^ Van der Lubbe, T.; Richter, U.; Knotschke, N. (2009). "Velociraptorine dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany" (PDF). Acta Palaeontologica Polonica. 54 (3): 401–408. doi:10.4202/app.2008.0007.
  4. ^ a b c Longrich, N.R.; Currie, P.J. (2009). "A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America". PNAS. 106 (13): 5002–7. doi:10.1073/pnas.0811664106. PMC 2664043. PMID 19289829.
  5. ^ Turner, A.S.; Hwang, S.H.; Norell, M.A. (2007). "A small derived theropod from Öösh, Early Cretaceous, Baykhangor Mongolia" (PDF). American Museum Novitates. 3557: 1–27. doi:10.1206/0003-0082(2007)3557[1:ASDTFS]2.0.CO;2. hdl:2246/5845. Retrieved 2007-03-29.
  6. ^ Holtz, Thomas R. Jr. (2010) Dinosaurs: The Most Complete, Up-to-Date Encyclopedia for Dinosaur Lovers of All Ages, Winter 2010 Appendix.
  7. ^ a b c d e Evans, D. C.; Larson, D. W.; Currie, P. J. (2013). "A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America". Naturwissenschaften. 100 (11): 1041–9. doi:10.1007/s00114-013-1107-5. PMID 24248432.
  8. ^ a b DePalma, Robert A.; Burnham, David A.; Martin, Larry D.; Larson, Peter L.; Bakker, Robert T. (2015). "The First Giant Raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation". Paleontological Contributions (14).
  9. ^ Senter, P.; Kirkland, J. I.; Deblieux, D. D.; Madsen, S.; Toth, N. (2012). Dodson, Peter (ed.). "New Dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the Evolution of the Dromaeosaurid Tail". PLoS ONE. 7 (5): e36790. doi:10.1371/journal.pone.0036790. PMC 3352940. PMID 22615813.
  10. ^ Bell, P. R., and P. J. Currie. 2015. A high-latitude dromaeosaurid, Boreonykus certekorum, gen. et sp. nov. (Theropoda), from the upper Campanian Wapiti Formation, west-central Alberta. Journal of Vertebrate Paleontology. doi:10.1080/02724634.2015.1034359
Acheroraptor

Acheroraptor is an extinct genus of dromaeosaurid theropod dinosaur known from the latest Maastrichtian Hell Creek Formation of Montana, United States. It contains a single species, Acheroraptor temertyorum. A. temertyorum is one of the two geologically youngest known species of dromaeosaurids, the other being Dakotaraptor, which is also known from Hell Creek.

Achillobator

Achillobator ( ə-KIL-ə-BAY-tor) is a dromaeosaurid theropod dinosaur that lived roughly 93 to 80 million years ago during the Late Cretaceous in what is now Mongolia, in Asia. It was among the largest dromaeosaurs; the holotype and only known individual of Achillobator is estimated at 5 to 6 m (16.4 to 19.7 ft) long. Achillobator was a moderately-built, ground-dwelling, bipedal carnivore. It would have been an active predator, hunting with the enlarged, sickle-shaped claw on the second toe.

Dromaeosaurinae

Dromaeosaurinae is a subfamily of Dromaeosauridae. Most dromaeosaurines lived in what is now the United States and Canada, as well as Mongolia, and possibly Denmark as well. Isolated teeth that may belong to African dromaeosaurines have also been discovered in Ethiopia. These teeth date to the Tithonian stage, of the Late Jurassic Period.All North American and Asian dromaeosaurine dinosaurs from the Late Cretaceous were generally small, no more than 1.8 metres (5.9 ft) long, in Dromaeosaurus and Adasaurus. However, among the dromaeosaurines were the largest dromaeosaurs ever; Dakotaraptor was ~5.5 metres (18 ft) long, Achillobator 6 metres (20 ft), and Utahraptor up to ~7 metres (23 ft).

Dromaeosaurus

Dromaeosaurus (, "swift running lizard") is a genus of theropod dinosaur which lived during the Late Cretaceous period (middle late Campanian), sometime between 76.5 and 74.8 million years ago, in the western United States and Alberta, Canada. The type species is Dromaeosaurus albertensis, which was described by William Diller Matthew and Barnum Brown in 1922.

Halszkaraptorinae

Halszkaraptorinae is a basal ("primitive") subfamily of Dromaeosauridae that includes the enigmatic genera Halszkaraptor, Mahakala, and Hulsanpes. A comparison of the fossils of Halszkaraptor with the bones of extant crocodilians and aquatic birds revealed evidence of a semiaquatic lifestyle. The group is named after Polish paleontologist Halszka Osmólska.

Hesperonychus

Hesperonychus (meaning "western claw") was a genus of small, carnivorous dromaeosaurid dinosaur. There is one described species, Hesperonychus elizabethae; the type species was named in honor of the woman who collected it in 1982. It is known from fossils recovered from the lowermost strata of the Dinosaur Park Formation of Alberta, dating to the late Cretaceous Period (Campanian stage) about 76.5 million years ago).

Hesperonychus is known from one partial pelvic girdle, holotype specimen UALVP 48778, collected by Dr. Elizabeth Nicholls in Dinosaur Provincial Park in 1982. The fossil remained undescribed, however, until Nick Longrich and Phil Currie published on it in 2009. A number of very small toe bones, including "sickle claws", in the collection of the Royal Tyrrell Museum may also belong to Hesperonychus. The gracile appearance of these toe bones makes it unlikely that they belonged to a member of Eudromaeosauria. Despite their small size, the pubic bones were fused, a characteristic of adult dinosaurs, indicating that the specimen does not represent a juvenile of a known species.Though known from only partial remains, Longrich and Currie estimated its total length at under one meter and weight at about 1.9 kilograms, making it one of the smallest known carnivorous dinosaurs from North America. The alvarezsaurid Albertonykus was smaller.

Hulsanpes

Hulsanpes is a genus of dromaeosaurid theropod dinosaur from Mongolia that lived during the Late Cretaceous.

Itemirus

Itemirus is a genus of theropod dinosaur from the Turonian age of the Late Cretaceous period of Uzbekistan.

Linheraptor

Linheraptor is a genus of dromaeosaurid dinosaur which lived in what is now China in the Late Cretaceous. It was named by Xu Xing and colleagues in 2010, and contains the species Linheraptor exquisitus. This bird-like dinosaur was less than 2 m (6.5 ft) long and was found in Inner Mongolia. It is known from a single, nearly complete skeleton.

Luanchuanraptor

Luanchuanraptor (meaning "Luanchuan thief") is a genus of dromaeosaurid theropod dinosaur from the Late Cretaceous of China. It is based on a partial skeleton from the Qiupa Formation in Luanchuan, Henan. A medium-sized dromaeosaurid, it is the first Asian dromaeosaurid described from outside the Gobi Desert or northeastern China. The fossil material is cataloged as 4HIII-0100 in the Henan Geological Museum and includes four teeth, one frontal, a neck vertebra, one or two back vertebrae, seventeen tail vertebrae, ribs, chevrons, a humerus (upper arm bone), claw and finger bones, partial shoulder and pelvic girdles, and other fragmentary bones from a moderately sized dromaeosaurid. The type species is L. henanensis, described by Lü et al. in 2007.

Pyroraptor

Pyroraptor (meaning "fire thief") is a genus of dromaeosaurid dinosaur from the Late Cretaceous of what is now southern France, it lived during the late Campanian and early Maastrichtian stages, approximately 70.6 million years ago. It is known from a single partial specimen that was found in Provence in 1992. The animal was named Pyroraptor olympius by Allain and Taquet in 2000.

Saurornitholestinae

Saurornitholestinae is a subfamily of dromaeosaurid dinosaurs. The saurornitholestines currently include three monotypic genera: Atrociraptor marshalli, Bambiraptor feinbergorum, and Saurornitholestes langstoni. All are medium-sized dromaeosaurs from the Late Cretaceous of western North America. The group was originally recognized by Longrich and Currie as the sister taxon to a clade formed by the Dromaeosaurinae and Velociraptorinae. However, not all phylogenetic analyses recover this group.

Shanag

Shanag is a genus of dromaeosaurid theropod dinosaur from the Early Cretaceous Period of Mongolia.

The type species of Shanag is S. ashile. It was named and described by Alan Turner, Sunny Hai-Ching Hwang and Mark Norell in 2007. The generic name refers to the black-hatted dancers in the Buddhist Cham dance. The specific name refers to the Ashile Formation, the old name for the layers where Shanag was found, used by Henry Fairfield Osborn.The holotype of Shanag, IGM 100/1119, was discovered in the Öösh Formation, the stratification of which is uncertain but probably dating to the Berriasian-Barremian. Shanag bears a strong resemblance to basal Chinese dromaeosaurids such as Microraptor and Sinornithosaurus, suggesting a close similarity between the fauna of the Öösh deposits, dated tentatively to 130 million years ago, and the Jehol Biota of China (such as the animals found in the roughly contemporary Yixian Formation), during the Early Cretaceous. The holotype specimen, about six centimetres long, is composed of an associated uncompressed upper and lower jaw fragment, containing a nearly complete right maxilla with teeth, a partial right dentary with teeth and an attached partial splenial.Shanag was a small predator. In 2010 Gregory S. Paul estimated its length at 1.5 metres, the weight at five kilogrammes. Shanag shows a mixture of dromaeosaurid, troodontid and basal avialan traits.Turner et alii assigned Shanag to the Dromaeosauridae. Their cladistic analysis indicated that it was a basal dromaeosaurid but higher in the tree than the Unenlagiinae. Later analyses recovered it in the Microraptorinae.

Unenlagiinae

Unenlagiinae is a subfamily of dromaeosaurid theropods. Unenlagiines are known from South America and Antarctica.

Unquillosaurus

Unquillosaurus (meaning "Unquillo river lizard") is a genus of maniraptoran dinosaur from the Late Cretaceous Period, discovered in Argentina. Known only from a single fossilized pubis (a pelvic bone), its total body length may have reached 2 to 3 metres (6.6 to 9.8 ft).

Utahraptor

Utahraptor (meaning "Utah's predator" or "Utah's thief") is a genus of theropod dinosaurs. It contains a single species, Utahraptor ostrommaysorum, which is the largest-known member of the family Dromaeosauridae. Fossil specimens date to the upper Barremian stage of the early Cretaceous period (in rock strata dated to 126 ± 2.5 million years ago).In 2018, it was proposed that Utahraptor be the Utah state dinosaur, an act that was approved by the Senate. Initially Utahraptor would have replaced another dinosaur, Allosaurus, as the state's official fossil, but it was decided that Utahraptor would be another symbol of the state.

Variraptor

Variraptor ( VARR-i-rap-tor; "Var thief") is a genus of coelurosaurian theropod dinosaur from the Late Cretaceous of France.

Velociraptorinae

Velociraptorinae is a subfamily of the theropod group Dromaeosauridae. The earliest velociraptorines are probably Nuthetes from the United Kingdom, and possibly Deinonychus from North America. However, several indeterminate velociraptorines have also been discovered, dating to the Kimmeridgian stage, in the Late Jurassic Period. These fossils were discovered in the Langenberg quarry, Oker near Goslar, Germany.In 2007 paleontologists studied front limb bones of Velociraptor and discovered small bumps on the surface, known as quill knobs. The same feature is present in some bird bones, and represents the attachment point for strong secondary wing feathers. This finding provided the first direct evidence that velociraptorines, like all other maniraptorans, had feathers.While most velociraptorines were generally small animals, at least one species may have achieved gigantic sizes comparable to those found among the dromaeosaurines. So far, this unnamed giant velociraptorine is known only from isolated teeth found on the Isle of Wight, England. The teeth belong to an animal the size of dromaeosaurines of the genus Utahraptor, but they appear to belong to a velociraptorine, judging by the shape of the teeth and the anatomy of their serrations.

Yurgovuchia

Yurgovuchia is an extinct genus of dromaeosaurid theropod dinosaur known from the Early Cretaceous (probably Barremian stage) of Utah. It contains a single species, Yurgovuchia doellingi. According to a phylogenetic analysis performed by its describers, it represents an advanced dromaeosaurine, closely related to Achillobator, Dromaeosaurus and Utahraptor.

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