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Eudromaeosaurs
Temporal range: Early Cretaceous – Late Cretaceous, 143–66 Ma Possible Kimmeridgian record
Eudromaeosauria diversity, featuring from top left to lower right: Utahraptor, Deinonychus, Velociraptor and Bambiraptor
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Chordata
Clade: Dinosauria
Clade: Saurischia
Clade: Theropoda
Family: †Dromaeosauridae
Clade: †Eudromaeosauria
Longrich & Currie, 2009
Type species
†Dromaeosaurus albertensis
Matthew & Brown, 1922
Subgroups
  • †Dromaeosaurinae
    Matthew & Brown, 1922
  • †Saurornitholestinae
    Longrich & Currie, 2009
  • †Velociraptorinae
    Barsbold, 1983

For classification of genera, see text

Eudromaeosauria ("true dromaeosaurs") is a subgroup of terrestrial dromaeosaurid theropod dinosaurs. They were small to large-sized, 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 probable dromaeosaurine Yurgovuchia, from the Cedar Mountain Formation, dated to 139 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]

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.

History of study[edit]

The Dromaeosaurinae was first erected in 1922 by Matthew and Brown as a part of the "Deinodontidae" (now named Tyrannosauridae).[4] Today, Dromaeosaurinae is defined as a monophyletic group including Dromaeosaurus and all the other dromaeosaurs closer to it than to Velociraptor, Microraptor, Passer and Unenlagia.[5]

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 redefined as clades, usually defined as all species closer to Velociraptor, Dromaeosaurus, or Saurornitholestes, respectively.[6]

Dromaeosaurines are usually found to consist of medium- to giant-sized species, with generally box-shaped skulls while 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. Velociraptorinae has traditionally included the more slender-snouted species which are found primarily in Asia, although this group may also include North American genera like Dineobellator and Deinonychus. Saurornitholestinae, the most recently-named subfamily, typically consists of smaller species with shortened snouts.[7]

Most eudromaeosaur genera are known from only 1-2 specimens. The major exceptions to this are Deinonychus, known from over 30 specimens belonging to an indeterminate number of individuals,[citation needed] Utahraptor, which is known from several specimens that have yet to receive a full description,[citation needed] Saurornitholestes, which is known from at least four individuals,[citation needed] and Velociraptor, which is known from a dozen almost complete skeletons.[citation needed]

Anatomy[edit]

Eudromaeosaurs were all bipedal and had relatively long arms in comparison to other theropods, like most other maniraptorans.[citation needed] Their wrists exhibited the typical maniraptoran condition in the semi-lunate carpal, which allowed them to fold their arms against their body in the same way that modern birds fold their wings.[citation needed] However unlike many other groups of coelurosaurs, dromaeosaurs possessed tibiae that were relatively short in relation to their metatarsals.[citation needed] Their second toe possessed the archetypal sickle-claw that all known dromaeosaurids bore which was held off the ground so that only the third and fourth toes touched the ground when walking.[citation needed] Eudromaeosaurs also generally possessed long and stiff tails, which are believed to have been used for balance.[citation needed]

There is some direct evidence of eudromaeosaurs such as Velociraptor being feathered. Today, it is believed that all eudromaeosaurs were fully-feathered and possessed wings, along with most, if not all, other maniraptorans.[citation needed]

Diagnostic traits[edit]

The original definition of Eudromaeosauria was a node-based definition. The apomorphic features of the group were left unresolved at the time of its naming.

Size[edit]

The size of three of the largest eudromaeosaurs compared to a human

Eudroameosaurs likely evolved from small ancestors, only around 1 kilogram (2.2 lb) in mass.[8] Later eudromaeosaurs were generally larger than this, with most being less than 2–3 metres (6.6–9.8 ft) long[9] and having masses estimated at around 15–40 kilograms (33–88 lb).[10]

Eudromaeosaurs are also known to have reached relatively large sizes. Among these were the dromaeosaurines Achillobator, at around 6 metres (20 ft),[11][9] and Utahraptor at up to 7 metres (23 ft).[12] The largest eudromaeosaurs are estimated to have been more than 200 kilograms (440 lb) in mass.[10] At least one velociraptorine taxon 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 appear to have belonged to an animal similar in size to the North American genus Utahraptor, but the morphology of the teeth suggests that the large size may only be homoplastic.[13] Remains from giant eudromaeosaurs are also reported from the Bissekty[14] and Bayan Shireh formations.[15]

Skull and tooth morphology[edit]

Skull diagram of Dromaeosaurus

The main difference in the skull morphology of eudromaeosaur species that has been observed is that those known from Asia have typically narrower skulls than those in North America. This is generally attributed to a phylogenetic difference (most Asian eudromaeosaurs are considered to be velociraptorines), but an analysis by Mark Powers and colleagues in 2020 demonstrated that dromaeosaur snouts in general increased in length during the Cretaceous. The reason for this is not fully understood, but it has been suggested that this reflects a change in the preferred prey of dromaeosaurs that existed from the Early Cretaceous to the Late Cretaceous.[15]

Eudromaeosaur skulls are also relatively solid in comparison to their primitive coelurosaur ancestors (i.e. they had smaller paranasal sinuses). In particular, the skull pneumaticity of oviraptorosaurs, which share a common ancestor with both birds and eudromaeosaurs is much higher than in any eudromaeosaurs. A 2021 survey of the premaxillae, maxillae, nasals, lacrimals, and jugals of several eudromaeosaurs was conducted in an attempt to reconstruct the ancestral condition of facial pneumaticity for coelurosaurs. The pneumatic elements of all five bones show a marked decline from basal coelurosaurs to derived paravians, with eudromaeosaurs completely lacking pneumatic spaces in their premaxillae. The reason for this evolutionary trend is unclear.[16]

  • Skull ecology[17]
  • Velociraptor endocranium[18]
  • Cranial osteology of Saurornitholestes[19]

Dromaeosaurines were a group of eudromaeosaurs that can be recognised in having stouter, box-shaped skulls, as opposed to the other subfamilies, which generally have narrower snouts, also, dromaeosaurines are generally more heavily built than the other members of their family, with thick, heavy-set legs, which were designed more for strength, rather than for speed. They differ from velociraptorines, in having a low DSDI ratio; i. e., their teeth have equal-sized serrations, on both the posterior and on the anterior edges. By contrast, velociraptorines often have larger serrations on the posterior side of the tooth, than the anterior, or no serrations on the anterior side at all.[5][9]

Integument[edit]

See also: Feathered dinosaur and List of non-avian dinosaur species preserved with evidence of feathers
The ulna of Dineobellator showing the position of the quill knobs

In 2007 paleontologists studied the ulna of a specimen 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 eudromaeosaurs had feathers.[20] Today, it is generally believed that all paravians and oviraptorosaurs (and possibly ornithomimosaurs) had pennaceous wing feathers.[citation needed]

Arms and wings[edit]

Diagram of the arms of several dinosaurs including maniraptorans (left); semilunate carpal of Linheraptor (right)

Feet and claws[edit]

A reconstruction of the bones of the foot of Deinonychus, emblematic of the condition in eudromaeosaurs

Aside from their generally larger size when compared to earlier-diverging dromaeosaurids, eudromaeosaurs are characterized by several features of the foot. First, differences existed in the positions of the grooves that 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.[6]

The second distinguishing characteristic of eudromaeosaurs is an expanded and enlarged "heel" on the last bone in the second toe (phalanx), 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.[6]

Tail[edit]

The tail of Dromaeosaurus, showing the ossified tendons
  • Yurgovuchia description[1]

Paleobiology and behavior[edit]

Claw function[edit]

A diagram showing possible functions of the sickle-claw in eudromaeosaurs

Teeth and diet[edit]

Metabolism[edit]

Locomotion[edit]

See also: Wing-assisted incline running

Eyes and senses[edit]

Nervous system and cognition[edit]

Respiration and pneumaticity[edit]

Reproduction[edit]

A life reconstruction of Deinonychus brooding with the aid of its wings
  • Description of a dromaeosaur ootaxon[22]

Growth and ontogeny[edit]

Ontogenetic size of Utahraptor

Pathologies[edit]

Classification[edit]

Relationships[edit]

  • Turner et al. (2012)[5]
  • Rauhut and Pol[23]
  • Angolin at al. (2019)[24]

One of the primary phylogenetic matrices in the scientific literature is the so-called "TWiG Matrix" from the Theropod Working Group. This matrix was first published by Steven Brusatte and colleagues in 2014.[25]

Paraves


Cau et al., 2017[edit]

Paraves

Hartman et al., 2019[edit]

Paraves

Motta et al., 2020[edit]

Paraves

Subgroups[edit]

Dromaeosaurus albertensis (cast)
Saurornitholestes langstoni (MOR 660R)
Velociraptor mongoliensis (MPC-D 100/54)

Eudromaeosauria is divided into three major subgroups. The composition of these groups is not universally agreed upon, but there is consensus on the classification of a few key taxa. Most phylogenetic analyses recover these three groups with varying members and varying degrees of confidence.

According to Turner and colleagues in 2012, technical diagnoses for the first subfamily of eudromaeosaurs, the Dromaeosaurinae, can be established based on several synapomorphies. These include fully serrated teeth; vertically oriented pubis; pubic boot (or end) projecting anteriorly and posteriorly; the jugal process of the maxilla, in a ventral view to the external antorbital fenestra, is dorsoventrally wide. This subfamily includes the eponymous Dromaeosaurus and all of its closest relatives.[5]

When erected by Barsbold in 1983, the second subfamily of eudromaeosaurs — Velociraptorinae, was conceived as a group containing Velociraptor and supposed closely related species.[27] It was not until 1998 that this group was defined as a clade by Paul Sereno. Sereno defined the group as all dromaeosaurids more closely related to Velociraptor than to Dromaeosaurus.[28] While several studies have since recovered a group of dromaeosaurids closely related to Velociraptor, they vary widely regarding which species are actually velociraptorines and which are either more basal or closer to Dromaeosaurus.

In 2005, Novas and Pol found a distinct velociraptorine clade close to the traditional view, which included Velociraptor, Deinonychus, and material that was later named Tsaagan. The analysis conducted by Turner and colleagues also supported a traditional, monophyletic of Velociraptorinae.[5] However, some studies found a very different group of dromaeosaurids in velociraptorinae, such as Longrich and Currie (2009), which found Deinonychus to be a non-velociraptorine, non-dromaeosaurine eudromaeosaur, and Saurornitholestes to be a member of a more basal group they named Saurornitholestinae.[6] A larger analysis in 2013 found some traditional velociraptorines, such as Tsaagan, to be more basal than Velociraptor, while others to be more closely related to Dromaeosaurus, making them dromaeosaurines. This study found Balaur, previously found to be a velociraptorine by most analyses, to be an avialan instead.[29]

Saurornitholestinae is the third, and most recently named, subfamily of Eudromaeosauria. The saurornitholestines currently include three monotypic genera: Atrociraptor marshalli, Bambiraptor feinbergi, 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.[6] However, not all phylogenetic analyses recover this group and/or with the same proposed genera.[5][1][26]

Occasionally, phylogenetic analyses will produce results that do not conform to the traditional topology that includes only three sub-clades. In their description of Acheroraptor in 2013, Evans and colleagues recovered the genera Atrociraptor and Deinonychus in a clade more derived than Saurornitholestinae, but more basal than either Dromaeosaurinae or Velociraptorinae.[30] In the 2020 description of Dineobellator, Jasinski and colleagues recovered the genera Utahraptor, Achillobator, and Adasaurus in a clade that was the sister-taxon of Velociraptorinae.[31] A similar result to Jasinski and colleagues was recovered by Hartman and colleagues a year earlier.[26] None of these aberrant clades have received strong support from subsequent analyses.

Taxonomic uncertainty[edit]

Choiniere et al. (2014)[33]

Eudrom.

Dromaeosaurinae

Saurornitholestinae

Velociraptorinae

Turner et al. (2021)[34]

Eudrom.

Saurornitholestinae

Dromaeosaurinae

Velociraptorinae

Czepiński (2022)[35]

Eudrom.

Velociraptorinae

Dromaeosaurinae

Saurornitholestinae

Table[edit]

The list below includes a comprehensive summary of all the possible classifications of each dromaeosaur species.

Possible Affinities of Eudromaeosaur Genera

Unambiguous Eudromaeosaurs

Genus Suggested classifications
Acheroraptor Originally described as a velociraptorine,[30] subsequently recovered as a saurornitholestine[36]
Achillobator Generally believed to be a dromaeosaurine,[37][38] has been recovered as being closely related to, but outside of, velociraptorines[31]
Adasaurus Originally described as a dromaeosaurine,[27] generally believed to be a velociraptorine,[37][39][31] but has been subsequently recovered as a dromaeosaurine[40] or as a basal member of eudromaeosauria[31][35][41]
Atrociraptor Originally described as a velociraptorine,[42] subsequently recovered as a dromaeosaurine,[43] a saurornitholestine,[44][36] or as being outside all three subfamilies[30]
Bambiraptor Either a saurornitholestine,[6] a junior synonym of Saurornitholestes,[45] or a basal member of eudromaeosauria,[31][44][37][19] has also been found as a velociraptorine[46]
Boreonykus Originally described as a velociraptorine,[47] has also been recovered as a dromaeosaurine[31]
Deinonychus Variously found as a dromaeosaurine,[43][36] a velociraptorine,[37][39][15] a saurornitholestine,[44][35] or being outside of all three groups[30][44]
Dineobellator Originally described as a velociraptorine,[31] which has been supported,[36][39] may be outside of all three major subfamilies[44]
Dromaeosaurus Must belong to Dromaeosaurinae by definition[5]
Itemirus Originally assigned to its own family ("Itemiridae") outside of dromaeosauridae,[48] later recovered as a dromaeosaurine[49] or possibly a velociraptorine[6]
Kansaignathus Early-diverging velociraptorine[39][50]
Kuru Velociraptorine very closely related to Velociraptor[37][39][35]
Luanchuanraptor Described as a velociraptorine[51]
Linheraptor Originally described as a basal eudromaeosaur closely related to Tsaagan,[52] has also been recovered as a velociraptorine[37][15][39][35] and a dromaeosaurine[26]
Saurornitholestes Must belong to Saurornitholestinae by definition[5]
Shri Generally accepted to be a velociraptorine[37][35]
Tsaagan Originally described as a velociraptorine,[53] which has been supported by subsequent analyses,[15][37][39] has also been recovered as a dromaeosaurine[26] or as being outside all three subfamilies[43]
Utahraptor Generally believed to be a dromaeosaurine,[43][37][39][1][15][5] has been suggested to be a velociraptorine[19] or to be outside all three subfamilies[30]
Velociraptor Must belong to Velociraptorinae by definition[5]
Yurgovuchia Originally described as a dromaeosaurine,[1] may be a basal eudromaeosaur[26]

Possible Eudromaeosaurs

Genus Suggested classifications
Balaur Described as a velociraptorine,[37] has been recovered as a dromaeosaurine, may be a stem-avialan[29][26]
Dakotaraptor Believed by some to be a chimera,[54][31][46] originally described as a dromaeosaurine,[43] subsequently found as a possible unenlagiine[26]
Dromaeosauroides Potentially dubious genus, only known from teeth, originally described as a dromaeosaurine,[55] subsequently recovered as an indeterminate dromaeosaurid[56]
Hesperonychus Originally described as a microraptorian,[6] but may be an avialan,[26] a eudromaeosaur, or some other kind of dromaeosaurid[31]
Nuthetes Potentially dubious genus, only known from teeth, may or may not be a velociraptorine[57][58]
Ornithodesmus Potentially dubious genus, may be an unenlagiine[26] or a eudromaeosaur[59]
Pyroraptor May be a dromaeosaurine or an indeterminate eudromaeosaur,[60] has been recovered as an unenlagiine[26]
Tianyuraptor Generally believed to be a microraptorian,[41] sometimes recovered as a eudroameosaur[41] or an unenlagiine[15]
Ulughbegsaurus Originally described as a carcharodontosaur, but may be a large eudromaeosaur[14]
Variraptor May be a dromaeosaurine, an indeterminate eudromaeosaur, an unenlagiine,[60] or a microraptorian[26]
Vectiraptor Tentatively described as a eudromaeosaur[60]
Yixianosaurus Originally described as an indeterminate maniraptoran,[61] but has since been recovered as an anchiornithid,[38] a scansoriopterygid,[62] as a basal eudromaeosaur,[26] or some other kind of stem-paravian[63]
Zapsalis Potentially dubious genus, only known from teeth, possible junior synonym of Saurornitholestes[19]

Phylogenetic hypotheses[edit]

  • Powers matrix,[35] TWiG matrix,[37] Currie/Evans matrix,[19] Hartman matrix,[26] Jasinski 2020 (TWiG + Currie/Evans matrices),[31] Jasinski 2022 (three combined matrices)[44]
  • Powers matrix analysis (Czepinski 2022)
Eudromaeosauria

Adasaurus

Velociraptorinae

Kuru

"Velociraptor" osmolskae

Shri

Velociraptor sp. (MPC-D 100/982)

Velociraptor mongoliensis

  • Hartman 2019
Eudromaeosauria
  • Napoli 2021 (TWiG matrix)
Eudromaeosauria
  • Zanno matrix
  • Dineobellator description (TWiG + Currie and Evans matrix) and re-description (TWiG + Currie/Evans + Powers matrices)
Eudromaeosauria
Eudromaeosauria

Evolutionary history[edit]

Mesozoic paravian distribution

Origin and dispersal[edit]

Diversification[edit]

Extinction[edit]

  • Terminal Cretaceous localities: Hell Creek, Udurchukan, Qiupa, Nanxiong, Khouribga

Paleoecology[edit]

Predation[edit]

See also: Fighting Dinosaurs

Social behavior[edit]

A skeletal mount of several Deinonychus attacking a Tenontosaurus in Philadelphia

Contemporary fauna[edit]

Geographic distribution[edit]

See also: Biogeography of paravian dinosaurs

Most eudromaeosaurs lived in what is now Asia and North America during the Cretaceous period, from the Berriasian to the Maastrichtian stages. Two possible eudromaeosaurs, Balaur and Dromaeosauroides, lived in Europe during that same time.[5] However, isolated teeth that may belong to African eudromaeosaurs have also been discovered in Ethiopia. These teeth date to the Tithonian stage, of the Late Jurassic period.[64]

See also[edit]

Source gathering[edit]

  • Pittman, Michael; Xu, Xing (2020). "Pennaraptoran Theropod Dinosaurs Past Progress and New Frontiers". Bulletin of the American Museum of Natural History. 440: 1. doi:10.1206/0003-0090.440.1.1.
  • Yang, Li; Li, Xiaobo; Chen, Jun; Reisz, Robert R. (2024). "First discovery of large-bodied dromaeosaurid fossil materials (Dinosauria: Theropoda) from the Upper Cretaceous Quantou Formation, Songliao Basin, Northeast China". Cretaceous Research. 153. Bibcode:2024CrRes.15305711Y. doi:10.1016/j.cretres.2023.105711.
  • Wick, Steven L.; Lehman, Thomas M.; Brink, Alyson A. (2015). "A theropod tooth assemblage from the lower Aguja Formation (Early Campanian) of West Texas, and the roles of small theropod and varanoid lizard mesopredators in a tropical predator guild". Palaeogeography, Palaeoclimatology, Palaeoecology. 418: 229–244. Bibcode:2015PPP...418..229W. doi:10.1016/j.palaeo.2014.11.018.
  • Manafzadeh, Armita R.; Gatesy, Stephen M.; Bhullar, Bhart-Anjan S. (2024). "Articular surface interactions distinguish dinosaurian locomotor joint poses". Nature Communications. 15: 854. Bibcode:2024NatCo..15..854M. doi:10.1038/s41467-024-44832-z. PMID 38365765.
  • Tada, Seishiro; Tsuihiji, Takanobu; Matsumoto, Ryoko; Hanai, Tomoya; Iwami, Yasuko; Tomita, Naoki; Sato, Hideaki; Tsogtbaatar, Khishigjav (2023). "Evolutionary process toward avian-like cephalic thermoregulation system in Theropoda elucidated based on nasal structures". Royal Society Open Science. 10 (4). Bibcode:2023RSOS...1020997T. doi:10.1098/rsos.220997. PMID 37063996.
  • Gianechini, Federico A.; Ercoli, Marcos D.; Díaz-Martínez, Ignacio (2020). "Differential locomotor and predatory strategies of Gondwanan and derived Laurasian dromaeosaurids (Dinosauria, Theropoda, Paraves): Inferences from morphometric and comparative anatomical studies". Journal of Anatomy. 236 (5): 772–797. doi:10.1111/joa.13153. PMC 7163733. PMID 32023660.
  • Frederickson, J.A.; Engel, M.H.; Cifelli, R.L. (2020). "Ontogenetic dietary shifts in Deinonychus antirrhopus (Theropoda; Dromaeosauridae): Insights into the ecology and social behavior of raptorial dinosaurs through stable isotope analysis". Palaeogeography, Palaeoclimatology, Palaeoecology. 552. Bibcode:2020PPP...55209780F. doi:10.1016/j.palaeo.2020.109780.
  • Bishop, Peter J. (2019). "Testing the function of dromaeosaurid (Dinosauria, Theropoda) 'sickle claws' through musculoskeletal modelling and optimization". PeerJ. 7: e7577. doi:10.7717/peerj.7577. PMID 31523517.

Gallery[edit]

References[edit]

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  2. ^ Sweetman S.C. (2004). "The first record of velociraptorine dinosaurs (Saurischia, Theropoda) from the Wealden (Early Cretaceous, Barremian) of southern England" (PDF). 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. ^ Matthew, W. D.; Brown, B. (1922). "The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta". Bulletin of the American Museum of Natural History. 46: 367–385. hdl:2246/1300.
  5. ^ a b c d e f g h i j k Turner, A.H.; Makovicky, P.J.; Norell, M.A. (2012). "A Review of Dromaeosaurid Systematics and Paravian Phylogeny". Bulletin of the American Museum of Natural History. 2012 (371): 1–206. doi:10.1206/748.1. hdl:2246/6352. S2CID 83572446.
  6. ^ a b c d e f g h Longrich, N.R.; Currie, P.J. (2009). "A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America". PNAS. 106 (13): 5002–7. Bibcode:2009PNAS..106.5002L. doi:10.1073/pnas.0811664106. PMC 2664043. PMID 19289829.
  7. ^ 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. S2CID 31096081. Archived from the original (PDF) on 2009-03-26. Retrieved 2007-03-29.
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  10. ^ a b Molina-Pérez, Rubén; Larramendi, Asier (2019). Dinosaur Facts and Figures: The Theropods and Other Dinosauriformes. Translated by Connolly, David; Ramírez Cruz, Gonzalo Ángel. Illustrated by Andrey Atuchin and Sante Mazzei. Princeton University Press. ISBN 978-0691180311.
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