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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]

Anatomy[edit]

Eudromaeosaurs were all bipedal and had relatively long arms in comparison to other theropods, like most other maniraptorans. 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. However unlike many other groups of coelurosaurs, dromaeosaurs possessed tibiae that were relatively short in relation to their metatarsals. 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. Eudromaeosaurs also generally possessed long and stiff tails, which are believed to have been used for balance.

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 all other paravians, oviraptorosaurs, and possibly ornithomimosaurs.

Size[edit]

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

Eudroameosaurs likely evolved from small ancestors, similar in size to microraptorians. Eudromaeosaurs were generally all larger than this, with most being between 2–3 metres (6.6–9.8 ft) long. Some were likely smaller than this, and the largest species reached sizes comparable to modern large terrestrial carnivores.

Most North American and Asian dromaeosaurines from the Late Cretaceous were generally medium to large-sized animals, with an average length of around 1.8 metres (5.9 ft); i. e., Dromaeosaurus and Yurgovuchia.[8] However, among the dromaeosaurines were the largest dromaeosaurs ever, with the feathered Dakotaraptor measuring 5.5 metres (18 ft) long,[9] Achillobator 6 metres (20 ft),[10][8] and Utahraptor up to 7 metres (23 ft).[11]

While most velociraptorines were generally small animals, at least one 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.[12]

Skull and tooth morphology[edit]

Skull diagram of Dromaeosaurus

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][8]

Integument[edit]

See also: Feathered dinosaur and List of non-avian dinosaur species preserved with evidence of feathers

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.[13] Today, it is generally believed that all paravians and oviraptorosaurs (and possibly ornithomimosaurs) had pennaceous wing feathers.

Arms and wings[edit]

See also: Wing-assisted incline running
The ulna of Dineobellator showing the position of the quill knobs

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

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]

Eyes and senses[edit]

Nervous system[edit]

Respiration and pneumaticity[edit]

Preening[edit]

Reproduction[edit]

A life reconstruction of Deinonychus brooding with the aid of its wings

Growth and ontogeny[edit]

Ontogenetic size of Utahraptor

Pathologies[edit]

Classification[edit]

Relationships[edit]

Paraves


Hartman et al., 2019
Paraves
Motta et al., 2020
Paraves
Cau et al., 2017
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 et al. 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.[17] 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.[18] 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.

Novas and Pol (2005) found a distinct velociraptorine clade close to the traditional view, which included Velociraptor, Deinonychus, and material that was later named Tsaagan. A cladistic analysis conducted by Turner et al. (2012) 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.[19] 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.[20]

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.[21] However, not all phylogenetic analyses recover this group and/or with the same proposed genera.[22][23][24]

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.[25] 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.[26] A similar result to Jasinski and colleagues was recovered by Hartman and colleagues a year earlier.[27] None of these aberrant clades have received support from other analyses, however they serve to illustrate the lack of a robust scientific consensus regarding the classification of the various genera of eudromaeosaurs.

Taxonomic uncertainty[edit]

Choiniere et al. (2014)[29]

Eudrom.

Dromaeosaurinae

Saurornitholestinae

Velociraptorinae

Turner et al. (2021)[30]

Eudrom.

Saurornitholestinae

Dromaeosaurinae

Velociraptorinae

Czepiński (2022)[31]

Eudrom.

Velociraptorinae

Dromaeosaurinae

Saurornitholestinae

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

Classification of Eudromaeosaur Genera

Unambiguous Eudromaeosaurs

Genus Possible classifications
Acheroraptor Originally described as a velociraptorine,[25] subsequently recovered as a saurornitholestine[32]
Achillobator
Adasaurus
Atrociraptor
Bambiraptor
Boreonykus
Deinonychus
Dineobellator Originally described as a velociraptorine,[33] may be outside of all three major subfamilies[34]
Dromaeosaurus Belongs to Dromaeosaurinae by definition
Itemirus
Kansaignathus Early-diverging velociraptorine[35][36]
Kuru
Luanchuanraptor
Linheraptor
Saurornitholestes Belongs to Saurornitholestinae by definition
Shri
Tsaagan
Utahraptor
Velociraptor Belongs to Velociraptorinae by definition
Yurgovuchia

Possible Eudromaeosaurs

Genus Possible classifications
Balaur
Dakotaraptor Described as a dromaeosaurinae, subsequently found as a possible unenlagiine, possibly a chimera
Dromaeosauroides Potentially dubious, only known from teeth
Nuthetes Potentially dubious, only known from teeth
Pyroraptor
Richardoestesia Potentially dubious, only known from teeth
Tianyuraptor
Ulughbegsaurus Originally described as a carcharodontosaur, but may be a large eudromaeosaur[37]
Variraptor
Vectiraptor
Zapsalis Potentially dubious, only known from teeth, possible junior synonym of Saurornitholestes

Phylogenetic hypotheses[edit]

Evolutionary history[edit]

Origin and dispersal[edit]

Diversification[edit]

Extinction[edit]

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.[38]

See also[edit]

Source gathering[edit]

Gallery[edit]

References[edit]

  1. ^ Senter, P.; Kirkland, J. I.; Deblieux, D. D.; Madsen, S.; Toth, N. (2012). "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.
  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.
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  6. ^ 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. Bibcode:2009PNAS..106.5002L. doi:10.1073/pnas.0811664106. PMC 2664043. PMID 19289829.
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  12. ^ Naish, D. Hutt, and Martill, D.M. (2001). "Saurischian dinosaurs: theropods." in Martill, D.M. and Naish, D. (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association, Field Guides to Fossils. 10, 242–309.
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  14. ^ Rauhut, Oliver WM; Tischlinger, Helmut; Foth, Christian (2019). "A non-archaeopterygid avialan theropod from the Late Jurassic of southern Germany". eLife. 8. doi:10.7554/eLife.43789. PMC 6516837. PMID 31084702.
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  16. ^ Hartman, Scott; Mortimer, Mickey; Wahl, William R.; Lomax, Dean R.; Lippincott, Jessica; Lovelace, David M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi:10.7717/peerj.7247. PMC 6626525. PMID 31333906.
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  18. ^ Sereno, P. C. (1998). "A rationale for phylogenetic definitions, with application to the higher-level taxonomy of Dinosauria". Neues Jahrbuch für Geologie und Paläontologie - Abhandlungen. 210 (1): 41–83. doi:10.1127/njgpa/210/1998/41.
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  20. ^ Godefroit, Pascal; Cau, Andrea; Hu, Dong-Yu; Escuillié, François; Wu, Wenhao; Dyke, Gareth (2013). "A Jurassic avialan dinosaur from China resolves the early phylogenetic history of birds". Nature. 498 (7454): 359–362. Bibcode:2013Natur.498..359G. doi:10.1038/nature12168. PMID 23719374. S2CID 4364892.
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  22. ^ 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.
  23. ^ Senter, P.; Kirkland, J. I.; Deblieux, D. D.; Madsen, S.; Toth, N. (2012). "New Dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the Evolution of the Dromaeosaurid Tail". PLOS ONE. 7 (5): e36790. Bibcode:2012PLoSO...736790S. doi:10.1371/journal.pone.0036790. PMC 3352940. PMID 22615813.
  24. ^ Hartman, S.; Mortimer, M.; Wahl, W.R.; Lomax, D.R.; Lippincott, J.; Lovelace, D.M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi:10.7717/peerj.7247. PMC 6626525. PMID 31333906.
  25. ^ a b 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. Bibcode:2013NW....100.1041E. doi:10.1007/s00114-013-1107-5. PMID 24248432. S2CID 14978813.
  26. ^ Jasinski, Steven E.; Sullivan, Robert M.; Dodson, Peter (2020). "New Dromaeosaurid Dinosaur (Theropoda, Dromaeosauridae) from New Mexico and Biodiversity of Dromaeosaurids at the end of the Cretaceous". Scientific Reports. 10: 5105. Bibcode:2020NatSR..10.5105J. doi:10.1038/s41598-020-61480-7.
  27. ^ Hartman, S.; Mortimer, M.; Wahl, W. R.; Lomax, D. R.; Lippincott, J.; Lovelace, D. M. (2019). "A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight". PeerJ. 7: e7247. doi:10.7717/peerj.7247. PMC 6626525. PMID 31333906.
  28. ^ Zanno, Lindsay E. (2010). "A taxonomic and phylogenetic re-evaluation of Therizinosauria (Dinosauria: Maniraptora)". Journal of Systematic Palaeontology. 8 (4): 503–543. Bibcode:2010JSPal...8..503Z. doi:10.1080/14772019.2010.488045.
  29. ^ Choiniere, Jonah N.; Clark, James M.; Norell, Mark A.; Xu, Xing (2014). "Cranial Osteology of Haplocheirus sollers Choiniere et al., 2010 (Theropoda: Alvarezsauroidea)". American Museum Novitates (3816): 1–44. doi:10.1206/3816.1. S2CID 4667472.
  30. ^ Turner, A.H.; Montanari, S.; Norell, M.A. (2021). "A new dromaeosaurid from the Late Cretaceous Khulsan locality of Mongolia". American Museum Novitates (3965): 1–46. doi:10.1206/3965.1.
  31. ^ Czepiński, Łukasz (2023). "Skull of a dromaeosaurid Shri devi from the Upper Cretaceous of the Gobi Desert suggests convergence to the North American forms". Acta Palaeontologica Polonica. 68. doi:10.4202/app.01065.2023. S2CID 259441055.
  32. ^ Powers, Mark J.; Fabbri, Matteo; Doschak, Michael R.; Bhullar, Bhart-Anjan S.; Evans, David C.; Norell, Mark A.; Currie, Philip J. (2021). "A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters". Journal of Vertebrate Paleontology. 41 (5). Bibcode:2021JVPal..41E0087P. doi:10.1080/02724634.2021.2010087.
  33. ^ Jasinski, Steven E.; Sullivan, Robert M.; Dodson, Peter (December 2020). "New Dromaeosaurid Dinosaur (Theropoda, Dromaeosauridae) from New Mexico and Biodiversity of Dromaeosaurids at the end of the Cretaceous". Scientific Reports. 10 (1): 5105. Bibcode:2020NatSR..10.5105J. doi:10.1038/s41598-020-61480-7. PMC 7099077. PMID 32218481.
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  37. ^ Sues, Hans-Dieter; Averianov, Alexander; Britt, Brooks B. (2022-12-22). "A giant dromaeosaurid theropod from the Upper Cretaceous (Turonian) Bissekty Formation of Uzbekistan and the status of Ulughbegsaurus uzbekistanensis". Geological Magazine. 160 (2): 355–360. doi:10.1017/S0016756822000954. ISSN 0016-7568.
  38. ^ Goodwin, M. B.; Clemens, W. A.; Hutchison, J. H.; Wood, C. B.; Zavada, M. S.; Kemp, A.; Duffin, C. J.; Schaff, C. R. (1999). "Mesozoic continental vertebrates with associated palynostratigraphic dates from the northwestern Ethiopian plateau". Journal of Vertebrate Paleontology. 19 (4): 728–741. doi:10.1080/02724634.1999.10011185.
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