Ningyuansaurus Ji, Lu, Wei and Wang, 2012
N. wangi Ji, Lu, Wei and Wang, 2012
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (Xingcheng Confuciusornis Museum coll.) skull, mandibles, nine cervical vertebrae, cervical ribs, ten dorsal vertebrae, dorsal ribs, gastralia, ~22 caudal vertebrae, chevrons, scapula (73 mm), coracoid, (?)sternal plates, humerus (75 mm), radii (55 mm), ulnae (58 mm), scapholunare, semilunate carpal, distal carpal III, metacarpals I (12 mm), phalanx I-1 (25 mm), manual ungual I, metacarpals II (28 mm), phalanx II-1, phalanx II-2, metacarpal III (29 mm), two manual phalanges, three manual unguals, ilia (70 mm), pubes (109 mm), ischia (41 mm), femora (135 mm), tibiae (172 mm), fibula (172 mm), astragali (17 mm wide), metatarsals II (88 mm), metatarsals III (105 mm), phalanx III-1 (24 mm), metatarsals IV (102 mm), pedal phalanges, pedal unguals, metatarsal V, body feathers, seeds
Diagnosis- (after Ji et al., 2012) at least 10 upper and 14 lower jaw teeth on each side; femur much longer than ilium; distal caudal vertebrae much elongated.
Comments- Ji et al. state the holotype, discovered prior to April 2012, was offered for study by a private individual. The locality was listed as "Lamadong, Jianchang County, Liaoning Province; Yixian Formation", but Li et al. (2014) state "recent fieldwork confirmed the outcrops of fossil beds exposed in Lamadong Village mainly belong to the Jiufotang Formation (X. L. Wang, personal commun.)." The description is brief and the illustration and photo quality poor, so that not much information is determinable. While the authors believe it is an oviraptorosaur more basal than Caudipteryx or Incisivosaurus, the low iliofemoral ratio, low ischiopubic ratio and enlarged pedal ungual ?II suggest possible paravian relations. Funston and Currie (2016) added Ningyuansaurus to Osmolska et al.'s oviraptorosaur matrix lineage and recovered it in a polytomy with Caudipteryx, Similicaudipteryx and Luoyanggia, but other than oviraptorosaurs only include Protarchaeopteryx, Archaeopteryx, Velociraptor and Herrerasaurus, not even recovering a paravian Velociraptor. Hartman et al. (2019) used a full taxon sample and recovered the taxon as a halszkaraptorine paravian, requiring 4 steps to move to Oviraptorosauria.
References- Ji, Lu, Wei and Wang, 2012. A new oviraptorosaur from the Yixian Formation of Jianchang, western Liaoning Province, China. Geological Bulletin of China. 31(12), 2102-2107.
Li, Zhou, Wang and Clarke, 2014. A new specimen of large-bodied basal enantiornithine Bohaiornis from the Early Cretaceous of China and the inference of feeding ecology in Mesozoic birds. Journal of Paleontology. 88(1), 99-108.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4), e1160910. 36(4), e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 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

Hulsanpes Osmolska, 1982
H. perlei Osmolska, 1982
Late Campanian, Late Cretaceous
Khulsan, Baruungoyot Formation, Mongolia
Holotype- (ZPAL MgD-I/173) (~560 mm; juvenile) prootic-opisthotic fragment, incomplete metatarsal II (34 mm), phalanx II-1 (6.5 mm), proximal phalanx II-2, incomplete metatarsal III (~39 mm), proximal phalanx III-1, incomplete metatarsal IV (~36 mm)
Diagnosis- (after Cau and Madzia, 2018) distal third of metatarsal II shaft extensively overlapped by metatarsal III in extensor view; proximal shaft of metatarsal III unconstricted and wider than adjacent metatarsals; metatarsal III-IV contact straight in extensor view; distal end of metatarsal IV diverges laterodistally.
Other diagnoses- Osmolska (1982) originally diagnosed Hulsanpes based on symplesiomorphies for Neotheropoda (functionally tridactyl pes; metatarsals II-IV subequally thick), Tyrannoraptora (metatarsus slender [width: length ratio 0.16]), Deinonychosauria (second pedal digit specialized), and Halszkarptorinae (metatarsal III not wedged in anterior view).
Comments- This specimen was discovered in Summer 1970 (Kielan-Jaworowska and Barsbold, 1972) and described twelve years later. Using Halszkaraptor as a guide, we can estimate Hulsanpes was about 560 mm long, but it was juvenile based on the rough bone texture and the badly abraded articular joints.
Osmolska allied this species with dromaeosaurids based on the ginglymoid second and third metatarsals and subequally developed metatarsals II and IV, but noted it resembled troodontids in the narrow metatarsus and weakly developed second pedal digit. She dismissed avian origins based on the lack of fusion. Chiappe and Norell (Norell pers. comm. to Currie, 2001) think Hulsanpes is not a dromaeosaurid, but from "another more speciose branch of the Maniraptora". Senter et al. (2004) was the first published analysis to include Hulsanpes, which emerged as a dromaeosaurid because they misscored the ginglymoid metatarsal II as absent in Sinovenator. Cau et al. (2017) described the more complete taxon Halszkaraptor and created Halszkaraptorinae for these and Mahakala finding them sister to unenlagiids plus dromaeosaurids using his Megamatrix.
References- Kielan-Jaworowska and Barsbold, 1972. Narrative of the Polish-Mongolian Palaeontological Expeditions 1967-1971. Palaeontologica Polonica. 27, 5-13.
Osmolska, 1982. Hulsanpes perlei n. g. n. sp. (Deinonychosauria, Saurischia, Dinosauria) from the Upper Cretaceous Barun Goyot Formation of Mongolia. Neues Jahrbuch fur Geologie und Palaeontologie, Monatshefte. 1982(7), 440-448.
Currie, 2001. Theropod dinosaurs from the Cretaceous of Mongolia. in Benton, Shishkin, Unwin and Kurochkin, eds. The Age of Dinosaurs in Russia and Mongolia. 434-455.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Natural History Museum. 8, 1-20.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399.
Cau and Madzia, 2018. Redescription and affinities of Hulsanpes perlei (Dinosauria, Theropoda) from the Upper Cretaceous of Mongolia. PeerJ. 6:e4868.

Mahakala Turner, Pol, Clarke, Ericson and Norell, 2007
M. omnogovae Turner, Pol, Clarke, Ericson and Norell, 2007
Late Campanian, Late Cretaceous
Tugriken Shire, Djadokhta Formation, Mongolia
Holotype- (IGM 100/1033) (~700 mm; adult) maxillary fragment, frontals (25.2 mm), partial quadrate, ectopterygoid, pterygoid fragment, posterior braincase, mandibular fragment, articular? fragment, dentary tooth, atlas, partial axis, third cervical prezygapophysis, incomplete fourth cervical vertebra (~16.6 mm), partial fifth cervical vertebra, two fragmentary posterior cervical vertebrae, parts of twelve dorsal vertebrae including first dorsal neural arch, second dorsal neural arch, third dorsal vertebra, anterior dorsal centrum (~9.5 mm), six posterior dorsal vertebrae (~9.2, ~9.8, ~10 mm), incomplete sacrum, first caudal vertebra, second caudal vertebra, third caudal vertebra, fourth caudal vertebra, fifth caudal vertebra, sixth caudal vertebra, seventh caudal vertebra, eighth caudal vertebra, ninth caudal vertebra, tenth caudal vertebra, eleventh caudal vertebra, twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra, fifteenth caudal vertebra, sixteenth caudal vertebra, seventeenth caudal vertebra, eighteenth caudal vertebra, nineteenth caudal vertebra, twentieth caudal vertebra, six chevrons, partial scapulae, incomplete humeri (~35-40 mm), radius (36 mm), partial ulnae (~40 mm), semilunate carpal, distal carpal III, metacarpal I (~6.8 mm), phalanx I-1 (~13.1 mm), metacarpals II (18 mm), phalanx II-1 (~9.5 mm), phalanx II-2 (~13.2 mm), manual ungual II (~12.5 mm), metacarpals III (16 mm), phalanx III-1, manual ungual III (~11.4 mm), manual ungual, incomplete ilium (52.5 mm), femur (76.2 mm), tibiae (110 mm; one fragmentary), proximal fibula, astragalocalcaneum, metatarsals I (one incomplete), distal phalanx I-1, pedal ungual I, metatarsal II (~74 mm), phalanx II-2 (9 mm), pedal ungual II (16 mm straight), metatarsals III (82 mm; one fragmentary), phalanges III-1 (19.1 mm), phalanx III-2 (16 mm), phalanges III-3 (13 mm), incomplete pedal ungual III (10.5 mm), partial metatarsals IV (~77 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, three phalanges IV-?, pedal unguals IV
Diagnosis- (after Turner et al., 2007) proximal caudal vertebrae with subhorizontal, laterally directed prezygapophyses (unknown in Hulsanpes); strongly compressed and anteroposteriorly broad ulna tapering posteriorly to a narrow edge; elongate lateral crest on the posterodistal part of the femur.
(after Turner et al., 2011) ledgelike depression at the confluence of metotic strut and posterior tympanic recess on the anterior face of the paroccipital process; posteriorly tapering scapula; short forelimb (humerus 50% femur length); prominent supratrochanteric process; absence of a cuppedicus fossa (unknown in Hulsanpes).
(after Cau et al., 2017) pendant paroccipital processes (unknown in Hulsanpes); short forelimb (humerus not longer than 50% of femur; (unknown in Hulsanpes); distal ends of metatarsals I and II
diverge medially (unknown in Hulsanpes); metatarsal III longer than femur (unknown in Hulsanpes); distal end of metatarsal III with symmetrical constriction proximal to trochlea.
Comments- Discovered in 1992 and first noted by Turner et al. (2006).
This is generally recovered as sister to unenlagiids plus dromaeosaurids using the TwiG matrix (as in Turner et al. 2007 and 2011), although Senter et al. (2012) recovered it as a microraptorian. Cau et al. (2017) described the more complete Halszkaraptor and created Halszkaraptorinae for these and Hulsanpes, finding them sister to unenlagiids plus dromaeosaurids using his Megamatrix and Brusatte's TWiG analysis.
References- Turner, Pol, Norell and Hwang, 2006. Resolving dromaeosaurid phylogeny: New information and additions to the tree. Journal of Vertebrate Paleontology. 26(3), 133A.
Turner, Pol, Clarke, Ericson and Norell, 2007. A basal dromaeosaurid and size evolution preceding avian flight. Science. 317, 1378-1381.
Turner, Pol, Conicet, Clarke and Norell, 2007. The basal-most dromaeosaurid: A new species from Tugrugyin Shireh, Mongolia. Journal of Vertebrate Paleontology. 27(3), 161A.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS ONE. 4(10), e7390.
Turner, Pol and Norell, 2011. Anatomy of Mahakala omnogovae (Theropoda: Dromaeosauridae), Tögrögiin Shiree, Mongolia. American Museum Novitates. 3722, 66 pp.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399.

Unenlagiinae Makovicky et al., 2005
Definition- (Unenlagia comahuensis <- Halszkaraptor escuilliei, Microraptor zhaoianus, Velociraptor mongoliensis, Dromaeosaurus albertensis, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019)
Other definitions- (Unenlagia comahuensis <- Velociraptor mongoliensis) (Makovicky, Apesteguia and Agnolin, 2005)
(Unenlagia comahuensis <- Microraptor zhaoianus, Velociraptor mongoliensis, Dromaeosaurus albertensis, Passer domesticus) ( Turner, Makovicky and Norell, 2012)
(Unenlagia comahuensis <- Microraptor zhaoianus, Dromaeosaurus albertensis, Vultur gryphus) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017)
Comments- Unenlagiinae was erected by Makovicky et al. (2005) to contain Unenlagia, Buitreraptor and Rahonavis at the base of Dromaeosauridae. It has been recovered in this position in most subsequent Theropod Working Group analyses, though sometimes in Avialae instead (Agnolin and Novas, 2011, 2013).
Turner et al.'s (2012) definition replaced Makovicky et al.'s, which only included Velociraptor as an external specifier, while Hartman et al. (2019) added Halszkaraptor so that halszkaraptorines could be a subfamiliy of unenlagiines.
References- Bonaparte, 1999. Tetrapod faunas from South America and India: A paleobiogeographic interpretation. PINSA. 65A(3), 427-437.
Makovicky, Apesteguía and Agnolín, 2005. The earliest dromaeosaurid theropod from South America. Nature. 437, 1007-1011.
Novas and Agnolin, 2010. Phylogenetic relationships of Unenlagiidae theropods: Are they members of Dromaeosauridae?
Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the Dromaeosauridae (Theropoda, Maniraptora)? Anais da Academia Brasileira de Ciências. 83(1), 117-162.
Gianechini and Apesteguia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods from South America. Anais da Academia Brasileira de Ciências. 83(1), 163-195.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. Springer. 96 pp.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 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

unnamed possible unenlagiine (Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton, Zhang, Wolfe, Yi, Tseng, Ran and Currie, 2016)
Early Cenomanian, Late Cretaceous
Angbamo, Myanmar
Material- (DIP-V-15103) (juvenile?) about nine mid to distal caudal vertebrae (2.97, 3.41, 3.41, 3.76, 4.01 mm), skin, feathers
Reference- Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton, Zhang, Wolfe, Yi, Tseng, Ran and Currie, 2016. A feathered dinosaur tail with primitive plumage trapped in Mid-Cretaceous amber. Current Biology. 26(24), 3352-3360.

unnamed unenlagiine (Langston, 1953)
Maastrichtian, Late Cretaceous
Ortega, Columbia
Material- (UCMP 39649b) incomplete lateral tooth
Comments- Initially identified as a carnosaur (Langston, 1953), this was described as an unenlagiine similar to Austroraptor by Ezcurra (2009).
References- Langston, 1953. Cretaceous terrestrial vertebrates from Colombia, South America. Bulletin of the Geological Society of America. 64, 1519.
Ezcurra, 2009. Theropod remains from the latest Cretaceous of Colombia and their implications on the palaeozoogeography of western Gondwana. Cretaceous Research. 30, 1339-1344.

unnamed unenlagiine (Novas, Agnolin, Rozadilla, Aranciaga-Rolando, Brissón-Eli, Motta, Cerroni, Ezcurra, Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento, García-Marsà, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez and Salgado, 2019)
Late Campanian-Early Maastrichtian, Late Cretaceous
Chorrillo Formation, Santa Cruz, Argentina
Material- (MPM 21548) partial pedal ungual II
?...(MPM 21549) pedal phalanx III-2 (45 mm)
Comments- Discovered between January and March 2019, Novas et al. assign MPM 21545 and 21546 to Unenlagiidae.
Reference- Novas, Agnolin, Rozadilla, Aranciaga-Rolando, Brissón-Eli, Motta, Cerroni, Ezcurra, Martinelli, D'Angelo, Álvarez-Herrera, Gentil, Bogan, Chimento, García-Marsà, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez and Salgado, 2019. Paleontological discoveries in the Chorrillo Formation (upper Campanian-lower Maastrichtian, Upper Cretaceous), Santa Cruz Province, Patagonia, Argentina. Revista del Museo Argentino de Ciencias Naturales. 21(2), 217-293.

unnamed unenlagiine (Candeiro, Cau, Fanti, Nava and Novas, 2012)
Turonian-Santonian, Late Cretaceous
Adamantina Formation of the Bauru Group, Brazil
Material- (MPM 011) (~1 m; adult) anterior-mid dorsal vertebra
Reference- Candeiro, Cau, Fanti, Nava and Novas, 2012. First evidence of an unenlagiid (Dinosauria, Theropoda, Maniraptora) from the Bauru Group, Brazil. Cretaceous Research. 37, 223-226.

unnamed unenlagiine (Benson, Rich, Vickers-Rich and Hall, 2012)
Late Aptian-Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P180889) proximal femur
Reference- Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from Southern Australia indicates high polar diversity and climate-driven dinosaur provinciality. PLoS ONE. 7(5), e37122.

Diuqin Porfiri, Baiano, dos Santos, Gianechini, Pittman and Lamanna, 2024
D. lechiguanae Porfiri, Baiano, dos Santos, Gianechini, Pittman and Lamanna, 2024
Etymology- "Genus name: Diuqin (from Mapuzungun, the language of the Mapuce people indigenous to the region where the fossil was found), bird of prey. Species name: lechiguanae, after Lechiguana, the witch in the 1975 film Nazareno Cruz y el Lobo (directed by eminent Argentinian filmmaker Leonardo Favio) who foresaw that the film's titular character would become a werewolf. Intended etymology: "Lechiguana's bird of prey.""
Santonian, Late Cretaceous
isthmus between Lago Barreales and Lago Mari Menuco, Bajo de la Carpa Formation, Neuquen, Argentina
Holotype- (MUCPv 1401) (subadult?) last sacral neural arch, incomplete proximal caudal neural arch, incomplete left humerus (~225 mm), four fragments
Diagnosis- (after Porfiri et al., 2024; autapomorphies) horizontal accessory lamina between spinopostzygapophyseal laminae on posteriormost sacral vertebra; pair of elliptical, bilateral, dorsolaterally-ventromedially oriented foramina immediately anterolateral to base of neural spine in (at least) posteriormost sacral and proximal caudal vertebrae; distolateral deltopectoral ridge of humerus arises on distal half of deltopectoral crest.
Comments- Of the four unidentified fragments in the holotype, Porfiri et al write "At least two of these may correspond to pieces of vertebrae (potentially transverse processes), but this identification is extremely tentative and as such the fragments are not described further herein". They further state "The neurocentral suture is open (Fig. 2A, B, D, E), and as such, the neural arch was unfused to its respective centrum at the time of death; consequently, the holotype of Diuqin lechiguanae is inferred to represent a somatically immature individual."
Porfiri et al. (2024) added Diuqin to a version of Brusatte's TWiG analysis and recovered it as a paravian in either basal Anchiornithinae, basal Dromaeosauridae or anywhere within Unenlagiinae.  When added to Mortimer's maniraptoromorph analysis it emerges sister to Coelurus, but only one step is needed to force it into Unenlagiinae, which seems more plausible considering its age and geographic location.
Reference- Porfiri, Baiano, dos Santos, Gianechini, Pittman and Lamanna, 2024. Diuqin lechiguanae gen. et sp. nov., a new unenlagiine (Theropoda: Paraves) from the Bajo de la Carpa Formation (Neuquén Group, Upper Cretaceous) of Neuquén Province, Patagonia, Argentina. BMC Ecology and Evolution. 24:77.

"Lopasaurus" Price vide Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021 online
Maastrichtian, Late Cretaceous
Ponto 1 do Price, Serra da Galga Formation, Brazil
Holotype- (DNPM coll.; lost) metatarsal II (~162 mm), distal metatarsal III, distal metatarsal IV
Comments- In 2021 the Serra da Galga and Ponte Alta Members of the Marília Formation were recognized as the Serra da Galga Formation. Collected in the 1950s and only surviving as an unfinished plate drawn by Price labeled 'Lopasaurus', this is presumably the "metatarsals (all in the DNPM, Rio de Jineiro) which Price believed included at least two new genera and families." Brum et al. (2021 online) stated it disappeared after Price's death and first published the figure and name, identifying it as an unenlagiine based on "the anterior surface of metatarsal III transversally expanded; the expansion of the posterolateral region of metatarsal II; and the expansion of the posteromedial region of metatarsal IV. In posterior view, the expansions observed in metatarsals II and IV overlap metatarsal III." While this could belong to the contemporaneous Ypupiara, Brum et al. note it was not associated with the type skull fragments, and multiple unenlagiines may have coexisted as in the Portezuelo Formation. "Lopasaurus" is a nomen nudum, as it was not used as valid when proposed (ICZN Article 11.5), "accompanied by a description or definition that states in words characters that are purported to differentiate the taxon" (Article 13.1.1), or "accompanied by the fixation of a type species in the original publication" (Article 13.3).
References- Bertini, Marshall, Gayet and Brito, 1993. Vertebrate faunas from the Adamantina and Marília formations (Upper Bauru Group, Late Cretaceous, Brazil) in their stratigraphic and paleobiogeographic context. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen. 188(1), 71-101.
Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021 online. A new unenlagiine (Theropoda, Dromaeosauridae) from the Upper Cretaceous of Brazil. Papers in Palaeontology. DOI: 10.1002/spp2.1375

Austroraptor Novas, Pol, Canale, Porfiri and Calvo, 2009
= "Austroraptor" Novas, Pol, Canale, Porfiri and Calvo, 2008 online
A. cabazai Novas, Pol, Canale, Porfiri and Calvo, 2009
= "Austroraptor cabazai" Novas, Pol, Canale, Porfiri and Calvo, 2008 online
Campanian-Maastrichtian, Late Cretaceous
Allen Formation, Rio Negro, Argentina
Holotype- (MML-195) (~4.9 m; 368 kg) (skull ~800 mm) incomplete maxillae, lacrimals, frontal, postorbital, incomplete dentaries, surangular, prearticular, third cervical vertebra (~111 mm), fifth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra (~79 mm), second dorsal vertebra, fourth dorsal vertebra (~68 mm), ribs, gastralia, humerus (262 mm), manual ungual III (53 mm), partial pubis, incomplete femur (~560 mm), tibia (565 mm), astragalus, calcaneum, pedal ungual I, metatarsal II (325 mm), phalanx II-2 (57 mm), metatarsal III (330 mm), pedal ungual III (67 mm), phalanx IV-2 (48 mm)
Referred- (MML-220) (adult) fragmentary maxilla, two teeth, incomplete mid-posterior dorsal vertebra (96 mm), twelve distal caudal centra (32-82 mm), incomplete humerus, radii (161 mm), ulnae (one incomplete, one partial; 188 mm), metacarpals I (39 mm), phalanx I-1 (90.5 mm), incomplete manual ungual I, manual unguals II (one partial; 67 mm), incomplete manual ungual III, tibia (~560 mm), phalanx I-1 (41 mm), proximal metatarsal II, distal metatarsal II, phalanx II-1 (61.2 mm), phalanx II-2 (58.1 mm), distal metatarsals III, phalanx III-1 (98.2 mm), phalanx III-2 (56.2 mm), phalanx III-3 (48.5 mm), phalanx IV-1 (49.3 mm), phalanx IV-2 (33.7 mm), pedal ungual IV (Carabajal et al., 2009; described in Currie and Paulina Carabajal, 2012)
Diagnosis- (modified from Novas et al., 2009) lacrimal highly pneumatized; ventral process strongly curved anteriorly; posterior process flaring out horizontally above orbit; postorbital lacking dorsomedial process for articulation with the frontal; squamosal process of postorbital extremely reduced; maxillary and dentary teeth fluted; humerus short, representing slightly less than 50 per cent of femur length; pedal phalanx II-2 transversely narrow, contrasting with the extremely robust phalanx IV-2.
Other diagnoses- Novas et al. (2009) also included the small tooth size and lack of serrations as apomorphies, but these are probably basal for paravians.
Comments- Note "Austroraptor" was earlier used as an unpublished name for Ozraptor (Pigdon, DML 1997). Austroraptor's name and description appeared online in December 2008 before being officially published in March 2009.
Novas et al. (2009) assigned the taxon to Unenlagiinae based on a TWiG analysis, which has been the result of all subsequent published analyses as well.
References- Pigdon, DML 1997. https://web.archive.org/web/20191030133231/http://dml.cmnh.org/1997Sep/msg00942.html
Novas, Canale and Isasi, 2004. Giant deinonychosaurian theropod from the Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 24(3), 26A-27A.
Carabajal, Currie, Garcia, Salgado, Cerda, Fernandez, Reichel, Sissons, Koppelhus and Cabaza, 2009. Un nuevo especimen de Austroraptor (Dinosauria: Theropoda: Dromaeosauridae) del Cretácico Tardío (Maastrichtiano) de Río Negro, Argentina. Ameghiniana. 46(S), 41R.
Novas, Pol, Canale, Porfiri and Calvo, 2009. A bizarre Cretaceous theropod dinosaur from Patagonia and the evolution of Gondwanan dromaeosaurids. Proceedings of the Royal Society B. 276(1659), 1101-1107.
Gianechini and Apesteguiia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods from South America. Anais da Academia Brasileira de Ciências. 83(1), 163-195.
Currie and Paulina Carabajal, 2012. A new specimen of Austroraptor cabazai Novas, Pol, Canale, Porfiri and Calvo, 2008 (Dinosauria, Theropoda, Unenlagiidae) from the Latest Cretaceous (Maastrichtian) of Rio Negro, Argentina. Ameghiniana. 49(4), 662-667.

Ypupiara Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021
Y. lopai Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021
Maastrichtian, Late Cretaceous
Ponto 1 do Price, Serra da Galga Formation, Brazil
Holotype- (DGM 921-R; destroyed) anterior maxillary fragment, dentary fragment
Diagnosis- (after Brum et al., 2021) preantorbital body of maxilla with only one neurovascular foramen (unknown in other unenlagiines); maxillary interdental plate rectangular and anteroposteriorly expanded (unknown in other unenlagiines); maxillary teeth with a CBR of between 0.6 and 1 (also in Austroraptor); teeth of maxilla widely spaced (also in Buitreraptor).
Comments- In 2021 the Serra da Galga and Ponte Alta Members of the Marília Formation were recognized as the Serra da Galga Formation. Originally collected in the 1950s, Holgado et al. (2018) announced this in an abstract as a new taxon of unenlagiine. Brum et al. state "After the great fire that destroyed the main building of the MN-UFRJ in September 2018 ... the specimen was not recovered and is here considered as lost." They added Ypupiara to Cau's matrix and recovered it as an unenlagiine sister to Austroraptor, which is also the result when added to the Hartman et al. matrix.
References- Holgado, Brum, Pegas, Bandeira, Souza, Kellner and Campos, 2018. A new Unenlagiinae (Theropoda: Dromaeosauridae) from the Maastrichtian of Brazil. Journal of Vertebrate Paleontology, Program and Abstracts, 2018. 148.
Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021 online. A new unenlagiine (Theropoda, Dromaeosauridae) from the Upper Cretaceous of Brazil. Papers in Palaeontology. DOI: 10.1002/spp2.1375

Alcmonavis Rauhut, Tischlinger and Foth, 2019
A. poeschli Rauhut, Tischlinger and Foth, 2019
Early Tithonian, Late Jurassic
Mörnsheim Formation, Germany
Holotype- (SNSB-BSPG 2017 I 133; Muhlheim specimen; 13th specimen) humerus (~90 mm), radius (~80-82 mm), ulna (82 mm), scapholunare, pisiform, semilunate carpal, distal carpal III, incomplete metacarpal I (~7.1 mm), phalanx I-1 (28.5 mm), manual ungual I (16 mm), metacarpal II (40.9 mm), phalanx II-1 (22.8 mm), phalanx II-2 (28 mm), manual ungual II (17.5 mm), metacarpal III (36.8 mm), phalanx III-1 (8.8 mm), phalanx III-2 (5.9 mm), phalanx III-3 (17.9 mm), manual ungual III (11 mm), manual claw sheaths
Diagnosis- (after Rauhut et al., 2019) humerus with large deltopectoral crest, with a maximal expansion that exceeds the width of the humeral shaft; proximal part of humerus strongly angled at approximately 38 degrees with respect to distal shaft; ulna with well-defined, single, oval, concave proximal cotyle and small lateral tubercle; distal end of ulna slightly asymmetrically expanded; large, crest-like biceps tubercle on proximal radius; longitudinal groove along medial side of radial shaft; metacarpal II considerably more robust than metacarpals I and III; manual phalanx I-1 with longitudinal groove; manual phalanx II-1 very robust, but with rounded, rather than flattened cross-section; manual phalanx II-1 slightly twisted; manual unguals with strongly developed and palmarly transversely expanded flexor tubercles.
Comments- This was discovered in 2017 and initially thought to be a new Archaeopteryx specimen. Paul (2024) used the informal term alcmonavians for Alcmonavis, seemingly entirely based on the "flattened" manual phalanx II-1 being developed early, admitting an uncertain placement within Eumaniraptora. This character is polymorphic for archaeopterygids and dromaeosaurids, but in any case the term remains informal as an 'Alcmonavia' has never been published.
Rauhut et al. (2019) added it to Foth et al.'s TWiG analysis and recovered it as an avialan more derived than archaeopterygids but outside Shemzhouraptor+Sapeornis+Pygostylia. When added to Hartman et al.'s TWiG analysis, it emerges as an unenlagiine, which can be avialans with the addition of a single step (albeit further from Aves than archaeopterygids).
References- Rauhut, Tischlinger and Foth, 2019. A non-archaeopterygid avialan theropod from the Late Jurassic of southern Germany. eLife. 8:e43789.
Paul, 2024. The Princeton Field Guide to Dinosaurs 3rd edition. Princeton University Press. 384 pp.

Buitreraptor Makovicky, Apesteguia and Agnolin, 2005
B. gonzalezorum Makovicky, Apesteguia and Agnolin, 2005
Cenomanian-Turonian, Late Cretaceous
Candeleros Formation of the Rio Neuquen Subgroup, Rio Negro, Argentina
Holotype- (MPCA 245) (>4 year old subadult) incomplete skull (~190 mm), incomplete mandibles (~166 mm), five teeth (2.5-4.6 mm), incomplete axis, partial third cervical vertebra, partial fourth cervical vertebra, fifth cervical vertebra, partial sixth cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth cervical vertebra, partial first dorsal vertebra, second dorsal vertebra, third dorsal vertebra, partial fourth dorsal vertebra, fifth dorsal vertebra, incomplete sixth dorsal vertebra, seventh dorsal vertebra, eighth dorsal vertebra, ninth dorsal vertebra, incomplete tenth dorsal vertebra, incomplete eleventh dorsal vertebra, partial twelfth dorsal vertebra, partial thirteenth dorsal vertebra, fifteen dorsal ribs, incomplete sacrum (57 mm), first caudal vertebra, second caudal vertebra, third caudal vertebra, fourth caudal vertebra, fifth caudal vertebra, sixth caudal vertebra, seventh caudal vertebra, eighth caudal vertebra, ninth caudal vertebra, tenth caudal vertebra, eleventh caudal vertebra, twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra, fifteenth caudal vertebra, nine chevrons, incomplete scapulae (~96 mm), coracoids (45 mm), furcula (~73 mm), humeri (136 mm), incomplete radius (95 mm), incomplete ulna (112.2 mm), semilunate carpal?, partial metacarpal I?, metacarpal II (20.1 mm), partial metacarpal III?, manual phalanges, partial ilia, ischium (54 mm), femora (149.5 mm), tibiae (one incomplete; >158 mm), fibulae (one incomplete; >127 mm), incomplete metatarsal II, phalanx II-1 (22.8 mm), phalanx II-2 (19.3 mm), phalanx IV-1 (23.9 mm), fragments
Paratype- (MPCA 238) (>4 year old subadult) incomplete sacrum, ilium, pubis, femur, tibiotarsus, metatarsal I (20 mm), phalanx I-1 (16.2 mm), incomplete metatarsal II (90.6 mm), phalanx II-1 (27.9 mm), phalanx II-2 (18.3 mm), pedal ungual II, incomplete metatarsal III (99.4 mm), incomplete metatarsal IV (90.6 mm)
Referred- (MPCA 471-C) material including partial phalanx I-1, partial metacarpal II, partial phalanx III-3, four manual fragments (Anonymous, 2005)
(MPCA 471-D) material including appendicular element (Cerda and Gianechini, 2015)
(MPCA? coll.) (Anonymous, 2005)
(MPCN-PV-598) (8+ year old subadult) three incomplete anterior cervical vertebrae with fused cervical ribs, sixth dorsal vertebra, partial seventh dorsal vertebra, eighth dorsal centrum, ninth dorsal centrum, tenth dorsal centrum, eleventh dorsal centrum, partial twelfth dorsal centrum, partial thirteenth dorsal vertebra, dorsal rib, fragmentary gastralia, partial synsacrum, fragmentary first caudal vertebra, fragmentary second caudal vertebra, partial third caudal vertebra, incomplete fourth caudal vertebra, incomplete fifth caudal vertebra, sixth caudal vertebra, seventh caudal vertebra, eighth caudal vertebra, ninth caudal vertebra, tenth caudal vertebra, incomplete eleventh caudal vertebra, twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal centrum, anterior fifteenth caudal centrum, two distal caudal vertebrae, chevrons, incomplete scapula, incomplete coracoid, incomplete furcula, incomplete humerus, incomplete radius, incomplete ulna, semilunate carpal, incomplete metacarpal I, phalanx I-1, manual ungual I, partial metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, partial metacarpal III, incomplete phalanges III-1, phalanx III-2, phalanx III-3, manual elements, incomplete ilia, incomplete pubes, fragmentary ischia, femur (148 mm), incomplete tibia (~185 mm), incomplete fibula, partial astragalus, metatarsal I, phalanx I-1, pedal ungual I, metatarsals II (one distal), phalanx II-1, phalanx II-2, pedal ungual II, phalanges II-?, metatarsals III (one distal; ~104 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV (one distal), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, phalanges IV-? (Novas et al., 2017)
Diagnosis- (after Turner et al., 2012) skull long, exceeding femoral length by 25%; large maxillary fenestra; continuous transition from frontal margin to postorbital process; quadrate with large lateral flange and pneumatic foramen; dentary bears a deep subalveolar groove; teeth small, unserrated, without root-crown constriction; posterior cervical centra with ventrolateral ridge; furcula pneumatic; flexor process present on humerus; brevis shelf expanded and lobate, projects laterally from posterior end of ilium.
(after Novas et al., 2017) dorsal vertebrae lacking pleurocoels; mid and distal caudals with lateral complex of ridges (transverse process with ridge that extends as feebly developed rims that reachs the prezygapophyses and postzygapophyses; anterior half of centrum with small ridge running towards the transverse process and a similar ridge is located at the posterior end of the centrum; sub-triangular fossa on anterior and posterior lateral margins, delimited by tiny ridges); scapular blade axially expanded at mid-length; extremely slender manual elements; manus longer than femur (117% of femoral length); penultimate pedal and manual phalanges with dorsally displaced collateral ligament pits that are medially displaced and nearly contact each other.
Comments- Briefly described by Makovicky et al. (2005) and Gianechini and Apesteguia (2011), Gianechini is studying this taxon for his thesis. While Novas et al. (2017) identified the most complete dorsaal vertebra as the seventh "based on comparisons with [the] holotype specimen", but given the two posteriormost dorsals (12 and 13) articulated with the sacrum and the five articulated posterior dorsals, it must be at least as anterior as the sixth.
Makovicky et al. found Buitreraptor to be an unenlagiine using the TWiG matrix, which has been recovered in all subsequent analyses as well.
References- Anonymous, 2005. Newly discovered birdlike dinosaur is oldest raptor ever found in South America.
Makovicky, Apesteguía and Agnolín, 2005. The earliest dromaeosaurid theropod from South America. Nature. 437, 1007-1011.
Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the Dromaeosauridae (Theropoda, Maniraptora)? Anais da Academia Brasileira de Ciências. 83(1), 117-162.
Gianechini and Apesteguiia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods from South America. Anais da Academia Brasileira de Ciências. 83(1), 163-195.
Gianechini, Makovicky and Apesteguia, 2011. The teeth of the unenlagiine theropod Buitreraptor from the Cretaceous of Patagonia, Argentina, and the unusual dentition of the Gondwanan dromaeosaurids. Acta Palaeontologica Polonica. 56(2), 279-290.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. Springer. 96 pp.
Cerda and Gianechini, 2015. Determinacion de estadios ontogeneticos en Buitreraptor gonzalezorum (Dinosauria: Theropoda) a partir de la microestructura osea. XXIX Jornadas Argentinas de Paleontología de Vertebrados, resumenes. Ameghiniana. 52(4) suplemento, 11-12.
Novas, Brissón Egli, Agnolin, Gianechini and Cerda, 2017. Postcranial osteology of a new specimen of Buitreraptor gonzalezorum (Theropoda, Coelurosauria). Cretaceous Research. doi: 10.1016/j.cretres.2017.06.003.

Pamparaptor Porfiri, Calvo and dos Santos, 2011
P. micros Porfiri, Calvo and dos Santos, 2011
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of the Rio Neuquen Subgroup, Neuquen, Argentina
Holotype- (MUCPv-1163) (~500-700 mm) metatarsal II (82.1 mm), phalanx II-1 (16.6 mm), phalanx II-2 (18.1 mm), proximal pedal ungual II, metatarsal III (93 mm), phalanx III-1 (26.5 mm), phalanx III-2 (17.4 mm), metatarsal IV (92.5 mm), distal phalanx IV-1, proximal phalanx IV-2
Diagnosis- (after Porfiri et al., 2011) slender metatarsus; metatarsal II approximately twice the proximal width of III or IV; metatarsal II distally overlapping metatarsal III; distal end of metatarsal II with small medially directed sulcus anteriorly; pedal phalanx II-2 longer than II-1; proximal half of metatarsal III narrow and with subparallel margins; metatarsal III not ginglymoid; metatarsals IV and III subequal in length; distal half of metatarsal IV strongly compressed transversely, acquiring a blade-like shape in posterior view.
Comments- This was discovered in 2005 and initially referred to Neuquenraptor by Porfiri et al. (2007), with the differences considered ontogenetic. Porfiri et al. (2011) later described it as a new taxon of dromaeosaurid, though they noted troodontid similarities and did not run a phylogenetic analysis. Novas et al. (2013) questioned the unenlagiine identification, stating the elongate metatarsus, "distally extended metatarsal IV, subequal in distal extension to metatarsal III", non-ginglymoid metatarsals II and IV, and more elongate and acute pedal ungual were more similar to birds. They regarded it as Paraves incertae sedis.
References- Porfiri, Calvo, dos Santos and Valieri, 2007. New record of Neuquenraptor (Theropoda, Dromaeosauridae) from the Late Cretaceous of Patagonia. Ameghiniana. 44(S), 34R.
Porfiri, Calvo and dos Santos, 2011. A new small deinonychosaur (Dinosauria: Theropoda) from the Late Cretaceous of Patagonia, Argentina. Anais da Academia Brasileira de Ciências. 83(1), 109-116.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research. 45, 174-215.

Overoraptor Motta, Agnolin, Brissón Egli and Novas, 2020
O. chimentoi Motta, Agnolin, Brissón Egli and Novas, 2020
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of Rio Limay Subgroup, Rio Negro, Argentina
Holotype- (MPCA-Pv 805) (~1.3 m) proximal caudal vertebra, two mid caudal vertebrae (20.5 mm), distal caudal centrum (16.6 mm), two chevrons, incomplete scapula, ulna (~116 mm), incomplete metacarpal I (~14.9 mm), partial phalanx I-1, manual ungual I (~25.1 mm), manual ungual II, ilial fragment, pubic shaft, phalanx I-1 (13.6 mm), incomplete pedal ungual I (~14.5 mm), metatarsals II (100.6 mm), phalanx II-2 (20.0 mm), incomplete pedal ungual II, metatarsals III (106.5 mm)
Paratype- (MPCA-Pv 818) (~1 m) manual phalanx I-1, phalanx III-2, ilial fragment, incomplete pubis, metatarsal II (~79 mm), phalanx II-1, phalanx III-1
Diagnosis- (after Motta et al., 2020) distal caudal centra with complex system of lateral longitudinal ridges and concavities (also in Buitreraptor and Rahonavis); scapula with medially deflected distal end; acromion reduced and ridge-like; robust ulna; radial cotyle of ulna saddle-shaped and prominent; metacarpal I with extensive medioventral crest; metatarsal II with longitudinal lateroventral crest on distal half, ending distally in a posterior tubercle; strongly dorsally displaced collateral pits on pedal phalanx II-2; metatarsal III distal end non-ginglymoid (also in Pamparaptor); metatarsal III distal end dorsoventrally deeper than transversely wide.
Comments- The mid and distal caudal vertebrae, metacarpal I and pedal ungual II of the holotype were originally described as ?Unenlagiidae gen. et sp. indet. by Motta et al. (2016). Note the measurements in Motta et al.'s (2020) Supplementary Table 1 are in cm, not mm as listed.
Motta et al. (2020) described the taxon and added it to Agnolin and Novas' TWiG analysis where it emerged sister to Rahonavis, one node closer to Ornithes than unenlagiids. Adding it to Hartman et al.'s maniraptoromorph analysis results in it being an unenlagiine close to Rahonavis, but making it sister to that henus takes 4 more steps.
References- Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson Egli and Novas, 2016. New theropod fauna from the Upper Cretaceous (Huincul Formtation) of northwestern Patagonia, Argentina. In Khosla and Lucas (eds.). Cretaceous period: Biotic diversity and biogeography. New Mexico Museum of Natural History and Science Bulletin. 71, 231-253.
Motta, Agnolin, Brissón Egli and Novas, 2020. New theropod dinosaur from the Upper Cretaceous of Patagonia sheds light on the paravian radiation in Gondwana. The Science of Nature. 107(3), 24.

Rahonaviformes Livezey and Zusi, 2007
"Rahonavidae" Miller, 2004
Comments- Miller (2004) proposed Rahonavidae for Rahonavis and Jeholornis, but did not indicate the family was new (e.g. on page 142 it lacks the asterisk he used to indicate which taxa were new), so is a nomen nudum (ICZN Article 16.1). Livezey and Zusi (2007) erected Rahonaviformes and Rahonavidae containing only Rahonavis itself. As these were only used in a table that lacked a diagnosis or definition, "Rahonavidae" was still a nomen nudum (ICZN Article 13.1.1).
References- Miller, 2004. A new phylogeny of the Dromaeosauridae. 2004 Student Showcase Journal. 20, 123-158.
Livezey and Zusi, 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society. 149 (1), 1-95.
Rahonavis Forster, Sampson, Chiappe and Krause, 1998b
= Rahona Forster, Sampson, Chiappe and Krause, 1998a (preoccupied Griveaud, 1975)
R. ostromi (Forster, Sampson, Chiappe and Krause, 1998) Forster, Sampson, Chiappe and Krause, 1998b
= Rahona ostromi Forster, Sampson, Chiappe and Krause, 1998a
Maastrichtian, Late Cretaceous
Anembalemba Member of Maevarano Formation, Madagascar
Holotype- (UA 8656) (adult) incomplete ~first dorsal vertebra (12.0 mm), seventh dorsal vertebra (10.8 mm), eighth dorsal vertebra (10.7 mm), partial ninth dorsal neural arch, partial tenth dorsal vertebra, incomplete eleventh dorsal vertebra (9.8 mm), incomplete twelfth dorsal vertebra (9.1 mm), incomplete synsacrum (42.3 mm), incomplete first caudal vertebra, second caudal vertebra (6.3 mm), third caudal vertebra (6.4 mm), fourth caudal vertebra (6.9 mm), fifth caudal vertebra (8.1 mm), sixth caudal vertebra (9.4 mm), seventh caudal vertebra (9.5 mm), eighth caudal vertebra (12.2 mm), ninth caudal vertebra (14.9 mm), tenth caudal vertebra (18.5 mm), eleventh caudal vertebra (22.6 mm), twelfth caudal vertebra (24.7 mm), thirteenth caudal vertebra (25.8 mm), twelve chevrons (2.8-11.6 mm dorsoventrally), scapula (82.2 mm), radius (126.5 mm), ulna (132.3 mm), ilia (66.5, 67.1 mm), pubes (one incomplete; 69.1 mm), incomplete ischia (27.4 mm), femora (87.6, 86.9 mm), tibiae (one incomplete; 118.1mm), incomplete fibulae, incomplete astragalocalcaneum (10.7 mm trans), distal tarsal IV, metatarsal I (8.6 mm), phalanx I-1 (10.2 mm), pedal ungual I (11.1 mm), metatarsals II (44.8, 43.9 mm), phalanx II-1 (12.8 mm), phalanx II-2 (11.3 mm), pedal ungual II (22.8 mm), metatarsals III (47.8, 47.6 mm), phalanx III-1 (19.2 mm), phalanx III-2 (12.9 mm), phalanx III-3 (10.5 mm), metatarsals IV (44.8, 43.1 mm), phalanx IV-1 (14.7 mm), phalanx IV-2 (9.6 mm), phalanx IV-3 (7.3 mm), pedal claw II sheath,
Referred- (FMNH PA 740) incomplete dentary (Forster et al., 2020)
(FMNH PA 746) distal humerus (Forster and O'Conner, 2000; described by O'Connor and Forster, 2010)
(FMNH PR 2821) incomplete ulna (Forster et al., 2020)
(UA 9604) distal humerus (Forster and O'Conner, 2000; described by O'Connor and Forster, 2010)
Diagnosis- (after Turner et al., 2012) neural canal at least 40% of the height of the dorsal vertebral centra; six sacral vertebrae; deeply concave glenoid fossa on scapula; elongate acromion process (length greater than length of glenoid); long muscle scar above glenoid on lateral face of scapula; proximally kinked scapular shaft; elongate ulna with ulnar papillae; ulnar distal condyle subtriangular in distal view and twisted more than 54 degrees with respect to the proximal end; undivided trochanteric crest, that is distally shifted on the femoral shaft; prominent and proximally placed fingerlike lateral ridge on femur; mediolaterally broad cnemial crest with short lateral process; proximal end of tibia of equal width and length; greatly reduced fibula lacking contact with the calcaneum; lack of a medial fossa on the fibula.
Comments- The genus Rahona is preoccupied by limantriid moths (Griveaud, 1975), so Forster et al. (1998b) renamed their genus Rahonavis. All elements of Rahonavis were found in 1995. The distal humeri were both found near to the holotype and articulate perfectly with the type ulna, so may belong to the same individual (O'Connor and Forster, 2010). O'Connor and Forster call them Humeral Taxon B. However, a second right ulna was also found in the same layer, so another individual of the same size was present who the humeri could also belong to.
Originally recovered as an avialan by Forster et al (1998a) and the TWiG matrix (Norell et al., 2001), Makovicky et al. (2005) and several other iterations of the TWiG matrix instead place it as a basal dromaeosaurid in Unenlagiinae. This was also recovered by Foth et al. (2014), and the most recent versions of the TWiG matrix using Turner's paravian updates (Brusatte et al., 2014). Agnolin and Novas (2011) recover a compromise phylogeny where unenlagiines include Rahonavis and are placed as basal avialans, while Xu et al. (2008) and Lee et al. (2014) again found it to be an avialan, apart from deinonychosaurian unenlagiines.
References- Griveaud, 1975. Descriptions préliminaires de nouveaux genres et espèces de Lymantriidae malgaches (Lep.). Bulletin de la Société entomologique de France. 80, 225-232.
Forster, Sampson, Chiappe and Krause, 1998a. The theropod ancestry of birds: New evidence from the Late Cretaceous of Madagascar. Science. 279, 1915-1919.
Forster, Sampson, Chiappe and Krause, 1998b. Genus correction. Science. 280(5361), 179.
Schweitzer, Watt, Avci, Forster, Krause, Knapp, Rogers, Beech and Marshall, 1999. Keratin immunoreactivity in the Late Cretaceous bird Rahonavis ostromi. Journal of Vertebrate Paleontology. 19(4), 712-722.
Forster and O'Conner, 2000. The avifauna of the Upper Cretaceous Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 20(3), 41A-42A.
Chinsamy and Elzanowski, 2001. Evolution of growth pattern in birds. Nature. 412, 402-403.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian theropods. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. 49-67.
Makovicky, Apesteguía and Agnolín, 2005. The earliest dromaeosaurid theropod from South America. Nature. 437, 1007-1011.
Xu, Zhao, Norell, Sullivan, Hone, Erickson, Wang, Han and Guo, 2008. A new feathered maniraptoran dinosaur fossil that fills a morphological gap in avian origin. Chinese Science Bulletin. 54(3), 430-435.
O'Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4), 1178-1201.
Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the Dromaeosauridae (Theropoda, Maniraptora)? Anais da Academia Brasileira de Ciências. 83(1), 117-162.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Forster, O'Connor, Chiappe and Turner, 2020. The osteology of the Late Cretaceous paravian Rahonavis ostromi from Madagascar. Palaeontologia Electronica. 23(2):a31.

Dakotaraptor DePalma, Burnham, Martin, Larson and Bakker, 2015
D. steini DePalma, Burnham, Martin, Larson and Bakker, 2015
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Holotype- (PBMNH.P.10.113.T) (robust adult) fragmentary dorsal centrum (~70 mm), mid caudal centrum (~75 mm), distal caudal vertebra (70 mm), four incomplete distal caudal vertebra, two distal caudal centra, partial distal caudal centrum, incomplete humeri (~320 mm), incomplete radii (~320 mm), ulnae (360 mm), metacarpal I (75 mm), distal phalanx I-1, metacarpal II (130 mm), distal phalanx II-2, distal phalanx III-2, femur (558 mm), tibiae (673 mm), astragalocalcaneum (~120 mm trans), metatarsal II (247 mm), pedal ungual II (160 mm straight), fragmentary metatarsal III (~320 mm), pedal ungual III (70 mm), metatarsals IV (one fragmentary; 290 mm)
Paratypes- ?(KUVP 156045) tooth
(PBMNH.P.10.115.T) (gracile adult) tibia (485 mm)
(PBMNH.P.10.118.T) (gracile) astragalocalcaneum (60 mm trans)
?(PBMNH.P.10.119.T) tooth
?(PBMNH.P.10.121.T) tooth
?(PBMNH.P.10.122.T) tooth
?(PBMNH.P.10.124.T) tooth
Diagnosis- (after DePalma et al., 2015) large size (~5.5 m long); teeth have 15-20 denticles/5 mm on distal carina and 20-27 denticles/5 mm on mesial carina; lateral and medial condyles of metacarpal II subequal in size; proximodorsal margin of metacarpal II straight in dorsal view; shallow lateral ligament fossa on metacarpal II; pedal ungual II (straight measurement) ~29% of femur length; fibular crest of tibia long and gracile (height does not exceed 9% of total length); proximal margin of fibular crest hooked; sharp ventral keel on pedal unguals II and III; reduced flexor tubercle on pedal ungual IV; lateral groove of pedal ungual IV fully enclosed in bone for about its distal half.
Comments- The holotype was discovered in 2005. Arbour et al. (2015) reidentified the three supposed furcula (one in the holotype, KUVP 152429 and NCSM 13170) as entoplastra of the trionychid turtle cf. Axestemys splendida.
DePalma et al. (2015) found it to be a dromaeosaurine sister to Dromaeosaurus using a version of the TWiG analysis. However, Cau (online, 2015) recovered it as sister to Velociraptorinae+Dromaeosaurinae in his larger unpublished analysis, and my own unpublished analysis agrees it is outside that clade.
References- Arbour, Zanno, Larson, Evans and Sues, 2015. The furculae of the dromaeosaurid dinosaur Dakotaraptor steini are trionychid turtle entoplastra. PeerJ PrePrints. https://doi.org/10.7287/peerj.preprints.1570v1
Cau, online 2015. http://theropoda.blogspot.com/2015/10/dakotaraptor-un-acheroraptor-gigante.html
DePalma, Burnham, Martin, Larson and Bakker, 2015. The first giant raptor (Theropoda: Dromaeosauridae) from the Hell Creek Formation. Paleontological Contributions. 14, 16 pp.

Dineobellator Jasinski, Sullivan and Dodson, 2020
= "Dineobellator" Jasinski, 2018
D. notohesperis Jasinski, Sullivan and Dodson, 2020
= "Dineobellator notohesperis" Jasinski, 2018
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Holotype- (SMP VP-2430) premaxillary fragment, maxillary fragment, ?maxillary tooth (11.3x7.78x4.15 mm), ?nasal fragment, lacrimal fragment, jugal fragment, partial basisphenoid, occipital condyle, several cranial fragments, partial dorsal rib, several rib fragments, first caudal vertebra (26.4 mm), ~tenth caudal vertebra (32.56 mm), several vertebral fragments, incomplete humerus (~215 mm), incomplete ulna (~140 mm), incomplete manual ungual ?II (~50 mm), proximal metacarpal III, incomplete manual ungual III (~55 mm), incomplete femur (~275 mm), partial ?astragalus, incomplete metatarsal I, incomplete metatarsal II, proximal metatarsal III, fragments
Diagnosis- (after Jasinski et al., 2020) short and robust neural spine on proximal caudal vertebrae; sharp angle of distal deltopectoral crest of humerus (also in Dakotaraptor); vertical offset of side grooves on manual ungual II; dorsodistally oriented groove just distal to articular surface on medial side of manual ungual II; tall and distally wide flexor tubercle on manual ungual II.
(suggested) accessory ventral side grooves on manual ungual III.
Other diagnoses- Jasinski et al. describe the first caudal as opisthocoelous but state "the anterior surface is flat, the posterior is concave", making it platycoelous as is common in dromaeosaurids (e.g. Deinonychus, Adasaurus). They list "proximal caudal vertebrae with curved ventral surface", but this is equally true of Bambiraptor, Velociraptor, Deinonychus, Adasaurus, IGM 100/22 and Yurgovuchia. Similarly, 'proximal caudal vertebrae with oval to subrectangular anterior and posterior centrum surfaces' describes all dromaeosaurids. The proximal caudal transverse processes are "gracile and subrectangular" in Bambiraptor, Deinonychus, Adasaurus and IGM 100/22 as well, but are actually broken distally in Dineobellator (Jasinski et al., 2011- fig. 10C-D) so that their shape is indeterminate. 'Distinct round concavities on anterior and posterior centrum surfaces in mid caudal vertebrae' are on rarely exposed surfaces in dromaeosaurid tails, but a similar structure is present at least posteriorly in Deinonychus, and anteriorly in Utahraptor. The supposed enlarged flexor tubercle on pedal ungual III is actually manual ungual III, and not larger than in other dromaeosaurids. This also affects their character of a secondary side groove on pedal ungual III, which is instead found in manual ungual III.
Comments- Discovered in 2008, Jasinski et al. (2011) briefly describe SMP VP-2430 as Dromaeosauridae indet., noting it "probably represents a new taxon and will be described in detail elsewhere." The dorsal vertebra they note and figure seems to be the centrum of the first caudal vertebra. Jasinoski et al. (2015) mention it as "a specimen with both cranial and postcranial material [that] is known from the Maastrichtian of New Mexico, probably representing a distinct taxon, and is by far the most complete dromaeosaurid material recovered from Late Cretaceous strata in southern Laramidia."Jasinski (2018) described the specimen in his thesis as Dineobellator notohesperis, which is a nomen nudum as theses are generally considered not "issued for the purpose of providing a public and permanent scientific record" (ICZN Areticle 8.1.1). He used Longrich and Currie's dromaeosaurid analysis to recover it as a eudromaeosaur closer to dromaeosaurines and velociraptorines than to saurornitholestiines. The taxon was officially named and described by Jasinski et al. (2020) who used the same analysis to recover it as a velociraptorine sister to Tsaagan sensu lato. Adding it to the Hartman et al. maniraptoromorph analysis results in it being an unenlagiid close to Dakotaraptor with 2 steps and 3 steps needed respectively to move it to the positions of Jasinski's thesis and eventual description.
The left lacrimal fragment is supp. fig. 2A would seem to be in medial view based on comparison with Saurornitholestes, making the labeled lacrimal recess either autapomorphic or taphonomic. Jasinoski et al. describe an articulated string of four vertebrae as fused distal caudals, but they are about half as long as tall (43-54%) unlike theropod distal caudals which are always much longer than tall. For a comparison, their centra are 75% as tall as the anterior face of the first caudal vertebra but 16-19% of its length, whereas in Deinonychus the caudal 75% as tall as the first (~caudal 15) is 143% its length. Instead, these small (4.12, 5.10 mm) and tall centra resemble actinopterygians, in both size and the presence of foramina and anteroposterior ridges which the authors attributed to prezygapophyseal rods. Multiple fish are present in the Alamo Wash local fauna including lepisosteids, although amiid and esocid vertebrae are also common in similar formations. The supposed pedal ungual III is not only far more curved than non-avialan pedal unguals (except deinonychosaur pedal unguals II), its dorsal margin arches far dorsal to the articular surface, which Senter described as distinguishing dromaeosaurid manual unguals from pedal unguals. Its articular surface is much smaller (59% as tall, 49% as wide) than the other ungual, suggesting it is from manual digit III.
References- Jasinski, Sullivan and Lucas, 2011. Taxonomic composition of the Alamo Wash local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann (eds.). Fossil Record 3. New Mexico Museum of Natural History and Science Bulletin. 53, 216-271.
Jasinski, Sullivan and Dodson, 2015. Late Cretaceous dromaeosaurid theropod dinosaurs (Dinosauria: Dromaeosauridae) from southern Laramidia and implications for dinosaur faunal provinciality in North America. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 150.
Jasinski, 2018. The integration of morphology, variation, and phylogenetics to better understand fossil taxa and their modern relatives. PhD thesis, University of Pennsylvania. 564 pp.
Jasinski, Sullivan and Dodson, 2020. New dromaeosaurid dinosaur (Theropoda, Dromaeosauridae) from New Mexico and biodiversity of dromaeosaurids at the end of the Cretaceous. Scientific Reports. 10:5105.

Neuquenraptor Novas and Pol, 2005
= "Araucanoraptor" Novas vide anonymous, 1997
N. argentinus Novas and Pol, 2005
= "Araucanoraptor argentinus" Novas vide anonymous, 1997
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of the Rio Neuquen Subgroup, Neuquen, Argentina
Holotype- (MCF PVPH 77) (~2 m) fragmentary cervical vertebra, dorsal ribs, chevrons, proximal radius, incomplete femur (~250 mm), distal tibia, distal fibula, incomplete astragalus, calcaneum, metatarsal I (30.4 mm), phalanx I-1 (27 mm), pedal ungual I (26.9 mm straight), incomplete metatarsal II (~147.9 mm), phalanx II-1 (39.1 mm), phalanx II-2 (36.2 mm), pedal ungual II (39.7 mm), incomplete metatarsal III (~172.7 mm), phalanx III-1 (57.4 mm), phalanx III-2 (36.6 mm), phalanx III-3 (34.4 mm), incomplete metatarsal IV (~149.6 mm), phalanx IV-2 (32.3 mm), phalanx IV-3 (25.4 mm), phalanx IV-4 (25.5 mm), pedal ungual IV (21.7 mm)
Other diagnoses- All of the characters listed in Novas and Pol's (2005) diagnosis are actually more widely distributed. A subarctometatarsus is present in most basal paravians. Metatarsal II laterally expands over metatarsal III posteriorly in Buitreraptor and Pamparaptor. An extensor sulcus on metatarsal III is also present in Buitreraptor, troodontids and microraptorians. Metatarsal III is also incipiently ginglymoid in Buitreraptor, Microraptor and several other paravians. Subequal pedal phalanges II-1 and II-2 are also present in several paravians including Adasaurus, Archaeopteryx, Microraptor, NGMC 91, Bambiraptor and Velociraptor. A strongly curved pedal ungual II is found in most dromaeosaurids and basal avialans.
Comments- This was discovered in 1996 and originally reported as a troodontid until its official description identified it as a dromaeosaurid. Novas (pers. comm., 2005) confirms that it is the same specimen as "Araucanoraptor". Gianechini and Apesteguia (2011) illustrate the proximal ungual originally placed as ungual I by Novas and Pol (2005) as ungual IV. The distal ungual originally placed as IV may be I or III, but is not illustrated by Gianechini and Apesteguia. Makovicky et al. (2005) suggested that Neuquenraptor and Unenlagia may be synonymous based on supposedly identical femur (U. comahuensis), pedal phalanx II-1 and pedal ungual II (U. paynemili). See the comments under Unenlagia for details, but Neuquenraptor seems distinct from U. comahuensis, but cannot be compared to U. paynemili, except for the possibly referrable phalanx II-1, which differ between taxa. Though usually referred to Unenlagiinae, without the addition of information from Unenlagia, Neuquenraptor can be placed in any of several basal paravian positions outside derived dromaeosaurids, derived troodontids and derived avialans.
Porfiri et al. (2007) reported another specimen of Neuquenraptor (MUCPv-1163), but this has since been made the holotype of Pamparaptor by Porfiri et al. (2011).
References- Novas, Cladera and Puerta, 1996. New theropods from the Late Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 16(3), 56A.
Anonymous, 1997. Clarin (Argentine newspaper). 7-23-97.
Novas, Apesteguia, Pol and Cambiaso, 1999. Un probable Troodontido (Theropoda - Coelurosauria) del Cretacico Tardio de Patagonia. XV Jornadas Argentinas de Paleontologia de Vertebrados. Ameghiniana. 36(S), 17R.
Novas and Pol, 2005. New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia. Nature. 433, 858-861.
Porfiri and Calvo, 2007. La validez taxonómica de Neuquenraptor argentinus (Theropoda, Dromaeosauridae) y la monofilia del nodo Unenlagiinae. Ameghiniana. 44(S), 34R.
Porfiri, Calvo, dos Santos and Valieri, 2007. New record of Neuquenraptor (Theropoda, Dromaeosauridae) from the Late Cretaceous of Patagonia. Ameghiniana. 44(S), 34R.
Gianechini and Apesteguiia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods from South America. Anais da Academia Brasileira de Ciências. 83(1), 163-195.
Porfiri, Calvo and dos Santos, 2011. A new small deinonychosaur (Dinosauria: Theropoda) from the Late Cretaceous of Patagonia, Argentina. Anais da Academia Brasileira de Ciências. 83(1), 109-116.

Dromaeosauridae Matthew and Brown, 1922 sensu Russell, 1969
Definition- (Dromaeosaurus albertensis <- Unenlagia comahuensis, Troodon formosus, Archaeopteryx lithographica, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019)
Other definitions- (Dromaeosaurus albertensis + Velociraptor mongoliensis) (modified from Padian et al., 1999)
(Microraptor zhaoianus + Sinornithosaurus millenii + Velociraptor mongoliensis) (Norell and Makovicky, 2004)
(Deinonychus antirrhopus <- Troodon formosus, Passer domesticus) (modified from Hu, Hou, Zhang and Xu, 2009)
(Dromaeosaurus albertensis <- Troodon formosus, Passer domesticus) (Turner et al., 2012)
(Dromaeosaurus albertensis <- Saurornithoides mongoliensis, Vultur gryphus) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017)
(Dromaeosaurus albertensis <- Ornithomimus edmontonicus, Troodon formosus, Passer domesticus) (Sereno, in press)
= Itemiridae Kurzanov, 1976
Definition- (Itemirus medullaris <- Dromaeosaurus albertensis, Stenonychosaurus inequalis, Tyrannosaurus rex) (Martyniuk, 2012)
= Dromaeosauridae sensu Sereno, 1998
Definition- (Velociraptor mongoliensis <- Troodon formosus) (modified from Sereno, 1998)
= Avenychia Miller, 2004
= Enantiavenychia Miller, 2004
= "Tetrapteryxidae" Miller, 2004
= Dromaeosauroidea Matthew and Brown, 1922 sensu Livezey and Zusi, 2007
= Dromaeosauridae sensu Hu, Hou, Zhang and Xu, 2009
Definition- (Deinonychus antirrhopus <- Troodon formosus, Passer domesticus)
= Dromaeosauridae sensu Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Dromaeosaurus albertensis <- Saurornithoides mongoliensis, Vultur gryphus)
= Serraraptoria Wang and Pei, 2024 online
Definition- (Microraptor zhaoianus, Velociraptor mongoliensis <- Mahakala omnogovae, Halszkaraptor escuilliei, Unenlagia comahuensis) (Wang and Pei, 2024 online)
= Dromaeosauridae sensu Sereno, in press
Definition- (Dromaeosaurus albertensis <- Ornithomimus edmontonicus, Troodon formosus, Passer domesticus)
Comments- Sereno's new (in press) definition modifies his earlier (1998) one by using the eponymous genus as an internal specifier, and adding Ornithomimus and Passer as external specifiers. This contrasts with the two published node-based definitions- (Dromaeosaurus albertensis + Velociraptor mongoliensis) by Padian et al. (1999) and (Microraptor zhaoianus + Sinornithosaurus millenii + Velociraptor mongoliensis) by Norell et al. (2004). Norell et al.'s is problematic for not including Dromaeosaurus, and for explicitly including Microraptor. The latter would make pygostylians dromaeosaurids in Mayr et al.'s (2005) and Agnolin and Novas' (2013) topology. Padian et al.'s is equivalent to the use of Dromaeosauridae prior to 1999, but microraptorians weren't known and 'unenlagiines' were considered avialans. Once microraptorians were discovered, they were assigned to Dromaeosauridae by most authors except Senter et al. (2004). So using a definition which won't probably exclude them does have some precedence, but Sereno's should have more external specifiers if it is adopted, namely Archaeopteryx (Paul, 1988; 2002), Tyrannosaurus (Matthew and Brown, 1922; Russell and Dong, 1994), Ornitholestes (Makovicky, 1995) and Oviraptor (Barsbold et al., 1990). I actually think this stem-based definition is better than Padian et al.'s node-based one for the additional reason that exactly which taxa fall into the (Dromaeosaurus + Velociraptor) crown is uncertain. With the recognition that velociraptorine sensu lato characters are symplesiomorphic for dromaeosaurs (e.g. found in Bambiraptor and microraptorians), taxa like Deinonychus or Saurornitholestes could easily fall just outside the defined node, not to mention more fragmentary taxa such as Variraptor or Pyroraptor. So I advocate the following redefinition of Sereno's- (Dromaeosaurus albertensis <- Tyrannosaurus rex, Ornithomimus velox, Oviraptor philoceratops, Troodon formosus, Archaeopteryx lithographica, Passer domesticus).
Kurzanov (1976) erected Itemiridae for Itemirus only, sister to tyrannosaurids than to dromaeosaurids. Martynuik (2012) defined Itemiridae as excluding Dromaeosaurus, but since the ICZN states families cannot contain other families, Itemiridae is listed as a synonym of Dromaeosauridae here.
Miller (2004) stated "I would ascribe Microraptor gui, and possibly Cryptovolans, to its own family, the Tetrapteryxidae", but Tetrapteryx is a gruid genus whose name cannot be used for a family of dromaeosaurs, so Miller's taxon is invalid. Miller also proposed Avenychia for "The most inclusive dromaeosaurid clade, including the Tetrapteryxidae* plus all later dromaeosaurid genera", which would make it a junior synonym of Dromaeosauridae here. Similarly, Miller proposes Enantiavenychia for the clade including Microraptor, Sinornithosaurus, Bambiraptor and eudromaeosaurs but not "tetrapteryxids", which is not recognized here as Cryptovolans and gui are synonymized with Microraptor zhaoianus.
Not dromaeosaurids- Teeth from the Albian Eumeralla Formation and Aptian Wonthoggi Formation of Victoria, Australia have been commonly referred to Dromaeosauridae (Rich and Vickers-Rich, 1994), but are retained as Coelurosauria incertae sedis here as the lack of mesial serrations is known in many other coelurosaurs (e.g. Orkoraptor, compsognathids, most troodontids).
A partial braincase referred to as possibly dromaeosaurid (CCMGE 466/12457) by Nessov (1995) was referred to Urbacodon sp. by Averianov and Sues (2007).
A partial pedal phalanx II-2 (IGM-7715) was tentatively referred to Dromaeosauridae by Rodriguez de la Rosa and Cevallos-Ferriz (1998) based on the dorsally placed collateral ligament pit. However, this character is also present in basal troodontids (e.g. IGM 100/44, Sinornithoides, Borogovia), Neuquenraptor and Rahonavis. It is here referred to Paraves incertae sedis.
Tavares et al. (2014) described two teeth from the Adamantina Formation of Brazil as dromaeosaurid, but these are here considered more likely to be abelisaurid.
References- Kurzanov, 1976. Braincase structure in the carnosaur Itemirus n. gen., and some aspects of the cranial anatomy of dinosaurs. Paleontological Journal. 1976, 361-369.
Rich and Vickers-Rich, 1994. Digs at Dinosaur Cove and Flat Rocks 1994. Excavation Report. Dinosaur Cove 1993 - 1994 & Inverloch 1994. 10-13.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Rodriguez de la Rosa and Cevallos-Ferriz, 1998. Vertebrates of the El Pelillal locality (Campanian, Cerro del Pueblo Formation), Southeastern Coahuila, Mexico. Journal of Vertebrate Paleontology. 18, 751-764.
Norell, Makovicky and Clark, 2000. A Review of the Dromaeosauridae. The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History 20.
Burnham, Senter, Barsbold and Britt, 2004. Phylogeny of the Dromaeosauridae. Journal of Vertebrate Paleontology. 24(3), 204A-205A.
Miller, 2004. A new phylogeny of the Dromaeosauridae. 2004 Student Showcase Journal. 20, 123-158.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the Cenomanian of Uzbekistan, with a review of troodontid records from the territories of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
Livezey and Zusi, 2007. Higher-order phylogeny of modern birds (Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological Journal of the Linnean Society. 149 (1), 1-95.
Manning, Ali, McDonald, Mummery and Sellers, 2007. Biomechanics of dromaeosaurid claws: Application of x-ray microtomography, nanoidentification and finite element analysis. Journal of Vertebrate Paleontology. 27(3), 111A-112A.
Hoffman and Hwang, 2009. Identifying diagnostic characters in the tooth enamel microstructure of dromaeosaurid dinosaurs. Journal of Vertebrate Paleontology. 29(3), 115A.
Longrich and Currie, 2009. A microraptorine (Dinosauria - Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Manning, Margetts, Johnson, Mustansar and Mummery, 2009. A finite approach to the biomechanics of dromaeosaurid dinosaur claws. Journal of Vertebrate Paleontology. 29(3), 141A.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Pan Aves. 189 pp.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. Springer. 96 pp.
Tavares, Branco and Santucci, 2014. Theropod teeth from the Adamantina Formation (Bauru Group, Upper Cretaceous), Monte Alto, São Paulo, Brazil. Cretaceous Research. 50, 59-71.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 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
Wang and Pei, 2024 online. The smallest known specimen of Microraptor (Dinosauria: Dromaeosauridae) from the Jiufotang Formation in northeastern China. Historical Biology. DOI: 10.1080/08912963.2024.2385604

"Velociraptor" osmolskae Godefroit, Currie, Li, Shang and Dong, 2008
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype- (IMM 99NM-BYM-3/3) incomplete maxillae, lacrimal
Diagnosis- (after Godefroit et al., 2008) long anterior ramus of maxilla, with elongation index (L/H ratio) 1.38; promaxillary fenestra subequal in size to maxillary fenestra and teardrop-shaped; long axis of promaxillary fenestra perpendicular to dorsal border of the maxilla; long axis of maxillary fenestra parallel to this border; ten maxillary teeth with short unserrated carina on the apical end of the mesial edge and with incipient serrations on the distal carina.
Comments- Evans et al. (2013) entered osmolskae into a version of Longrich and Currie's dromaeosaurid analysis and stated "Velociraptor is found to be paraphyletic in this analysis, with V. osmolskae being more closely related to Tsaagan than to Velociraptor mongolienesis from Mongolia. Although the elongate shape of the maxilla in V. oskmolskae is similar to that of V. mongoliensis, V. osmolskae shares a number of features with Linheraptor (Xu et al. 2010a) (here synonomized with T. mangas following Turner et al. (2012)) from the same host formation, including a large, teardrop-shaped maxillary fenestra positioned anteriorly within the antorbital fossa (115[1]) and directly above the promaxillary fenestra."
References- Godefroit, Currie, Li, Shang and Dong, 2008. A new species of Velociraptor (Dinosauria: Dromaeosauridae) from the Upper Cretaceous of northern China. Journal of Vertebrate Paleontology. 28(2), 432-438.
Evans, Larson and Currie, 2013. A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America. Naturwissenschaften. 100(11), 1041-1049.

Acheroraptor Evans, Larson and Currie, 2013
A. temertyorum Evans, Larson and Currie, 2013
Etymology- "The generic name is derived from Acheron, the River of Pain in the underworld of ancient Greek mythology, in reference to the Hell Creek Formation, and Latin raptor, robber, a suffix used in previously named dromaeosaurids. The name Acheron has been deliberately shortened in the combination for euphony. The specific epithet honors James and Louise Temerty for their outstanding service and contributions to the Royal Ontario Museum."
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, ~45 km southwest of of Jordan, Montana, US
Holotype- (ROM 63777) (skull ~224 mm) incomplete right maxilla, second maxillary tooth (12x7.125x3.25 mm)
Referred- ?...(ROM 63778) incomplete left dentary (Evans, Larson and Currie, 2013)
Diagnosis- (after Evans et al., 2013) relatively small anterior region of antorbital fossa (defined as the region between the anterior borders of the antorbital fenestra and the antorbital fossa); maxillary fenestra that almost reaches ventralmargin of antorbital fossa and is positioned directly posterior to promaxillary fenestra; extensive, posteriorly projected postantral wall that is visible laterally through antorbital fenestra (also in Austroraptor); maxillary dentition with unique configuration of prominent apicobasal ridges on both labial and lingual surfaces of tooth.
Comments- The holotype was found on August 28, 2009, while the referred dentary was collected several years later. Evans et al. stated "The referred dentary, ROM 63778, was found approximately 4 m from the holotype maxilla in the same sandstone-hosted bonebed" and "The corresponding size and close physical association of the holotype maxilla and the dentary in the same bonebed suggest that they may belong to the same individual animal." Note while they claim "Morphometric analyses ... of tooth measurements from the specimen (Tables S1 and S2) indicate that the teeth are similar to previously collected isolated dromaeosaur teeth from the Hell Creek and Lance formations and that they are more similar to these teeth than any other North American dromaeosaur", Figure S4 actually shows some Saurornitholestes fall out just as closely as Lancian teeth. Powers et al. (2022) described maxillary characters in detail using CT scans.
Evans et al. (2013) entered Acheroraptor into a version of Longrich and Currie's dromaeosaurid analysis and found it to be a basal velociraptorine.
References- Evans, Larson and Currie, 2013. A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America. Naturwissenschaften. 100(11), 1041-1049.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as 2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters. Journal of Vertebrate Paleontology. 41(5), 2010087.

Boreonykus Bell and Currie, 2015
B. certekorum Bell and Currie, 2015
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Holotype- (RTMP 89.55.47) incomplete frontal (52.4 mm)
Diagnosis- (after Bell and Currie, 2015) supratemporal ridge on frontals forming acute angle (55 degrees) anteriorly.
Combination of- frontal elongate and slender; low supratemporal ridge forming arc that is concave relative to interfrontal suture in dorsal view; area immediately posterior to supratemporal ridge smooth and posteroventrally sloping.
Comments- Ryan and Russell (2001) listed the holotype frontal as "Saurornitholestes undescribed n. sp. (Currie pers. comm.)". Bell and Currie (2015) described this and other material (distal caudal RTMP 88.55.129, manual ungual UALVP 53597, and pedal ungual II RTMP 86.55.184) from the Pipestone Creek Pachyrhinosaurus lakustai bonebed as Boreonykus certekorum, noting that preserved elements are not duplicated and are properly proportioned to belong to one individual. The fourteen referred teeth are shed so were admitted to at least belong to another individual. When entered in Longrich and Currie's dromaeosaurid analysis, it emerged as a velociraptorine if the teeth were included, and a member of Velociraptorinae+Dromaeosaurinae if they were excluded. However, Cau (online, 2015) noted that given the paucity of remains and number of small Judithian theropod taxa, it should not be assumed that the frontal belongs to the same individual as the dromaeosaurid postcrania. Indeed, he reported that when the frontal is entered into his theropod supermatrix, it emerges in several position within Maniraptoriformes but never as a dromaeosaurid. My own analysis finds it to be a coelurosaur excluded from ornithomimosaurs, alvarezsauroids, therizinosaurs, oviraptorosaurs, unenlagiines, microraptorians, dromaeosaurines, derived troodontids and birds. As it is clearly not a member of contemporaneous clades known from frontals, Boreonykus may belong to one of the problematic taxa such as Richardoestesia or Paronychodon which are both placed in Paraves here. If I'm correct and Richardoestesia is a microraptorian, Boreonykus may be Paronychodon.
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press. 279-297.
Bell and 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. e1034359. DOI: 10.1080/02724634.2015.1034359.
Cau, online 2015. http://theropoda.blogspot.com/2015/12/boreonykus-e-un-dromaeosauridae.html

"Julieraptor" Thompson vide Robbins, 2009
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Material- (ROM coll.; Sid Vicious) non-jaw cranial elements, cervical series, dorsal series, dorsal ribs, incomplete caudal series, distal chevrons, pectoral girdles, forelimbs (missing four manual elements), ilia, pubes, hindlimbs (missing two pedal elements)
Comments- As detailed by Bakker (online 2012) and Robbins (online 2009), material exposed on the surface (cranial elements, manual phalanges and unguals) were collected in 2002 and nicknamed "Julieraptor" by Thompson, while the rest of the specimen was excavated by 2006 and nicknamed Sid Vicious by Murphy of the Judith River Dinosaur Institute. Once reuninted and after various legal issues "the skeleton was returned to the land owner, who arranged to sell the specimen to the Royal Ontario Museum" (Bakker, online 2012).
Described in BHI's website as having "many differences from another significant small raptor, Bambiraptor; primarily in the ilia, scapula, arms, hands, feet and size (about 35% larger than Bambiraptor)." Whether these differences prove convincing and whether "Julieraptor" can be distinguished from other contemporaneous dromaeosaurids such as Saurornitholestes and Dromaeosaurus remains to be seen.
References- Robbins, 2009. Dinosaur fossil faces charges of theft. The New York Times. January 21.
Bakker, online 2012. http://blog.hmns.org/2012/06/bakker-blogs-the-kleptomania-continues-with-a-sid-vicious-julieraptor-dino-rustlers-part-ii/
BHI online, 2012. http://www.bhigr.com/store/product.php?productid=679

Kansaignathus Averianov and Lopatin, 2021a
K. sogdianus Averianov and Lopatin, 2021a
Early Santonian, Late Cretaceous
Kansai, Yalovach Formation, Tajikistan
Holotype- (PIN 2398/15) (~2 m) dentary (140 mm)
Referred- ?(PIN 2398/4) manual ungual II (Averianov and Lopatin, 2021)
?(PIN 3041/11) anterior dorsal vertebra (Alifanov and Averianov, 2006)
?(PIN 3041/45) anterior tooth (Averianov and Lopatin, 2021)
Early Santonian, Late Cretaceous
Kyzylpilyal, Yalovach Formation, Tajikistan
? teeth (Nessov, Kaznyshkina and Cherepanov, 1987)
Diagnosis- (after Averianov and Lopatin, 2021) twelve dentary teeth; differs from ?Itemirus ZIN PH 2338/16 in being straight in dorsal view and lacking vertical grooves in its interdental plates; differs from Dromaeosaurus and Deinonychus in being more ventrally convex and having chin; differs from Saurornitholestes and Linheraptor in lacking ventral curvature at the symphysis; differs from Atrociraptor in being more concave dorsally and lacking grooves between interdental plates; differs from Velociraptor in being taller, lacking ventral curvature at the symphysis, and lacking a paradental groove medially; differs from Acheroraptor in lesser development of lateral foramina.
(suggested) differs from Bambiraptor in lacking ventral curvature at symphysis.
Other diagnoses- Averianov and Lopatin (2021b) claim Kansaignathus differs from Bambiraptor "in the presence of the ventral row of vascular foramina in the middle of the dentary and a smaller difference in the depth between the anterior and posterior parts of the dentary", but Bambiraptor also has the ventral row of foramina in the middle and is not notably different in anterior taper (pers. obs.). They also claim it differs from Velociraptor in having the dorsal row of lateral foramina be closer to the alveolar edge, but this doesn't appear to be true. The authors also say it differs from Tsaagan "in the absence of a common groove for the dorsal row of vascular foramina on the labial side of the dentary", but the grooves in both taxa are not continuous.
Comments- The holotype dentary was discovered in 1963-1964 and described by Averianov and Lopatin (2021a) as a new taxon of dromaeosaurid. When they entered it into Currie and Longrich's dromaeosaurid matrix it emerged as a velociraptorine closer to Velociraptor than Deinonychus but further than Acheroraptor.
Nessov (1995) noted (translated) "dromaeosaurids (D. A. Russell, personal communication, 1988)" from Kansai. Alifanov and Averianov (2006) stated that at Kansai "The family Dromaeosauridae is probably represented by a corpus of a relatively short anterior thoracic vertebra with a stout hypapophysis (PIN, no. 3041/11)." Averianov and Loptain (2021b) stated the vertebra may belong to Kansaignathus, as well as "a tooth with considerably lingually displaced carina (specimen PIN, no. 3041/45), ... and an ungual phalanx of manual finger II (specimen PIN, no. 2398/4)." Nessov et al. (1987) had earlier reported deinonychosaurian teeth from a different locality. While these materials could easily belong to other dromaeosaurid taxa given the presence of multiple genera in well sampled formations, they are listed here for convenience.
References- Nessov, Kaznyshkina and Cherepanov, 1987. Dinosaurs, crocodiles, and other archosaurs of the late Mesozoic of Middle Asia and their place in ecosystems. Abstracts of the XXXII Session of the All-Union Paleontological Society. 46-47.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology and paleobiogeography. Scientific Research Institute of the Earth's Crust. 156 pp.
Alifanov and Averianov, 2006. On the finding of ornithomimid dinosaurs (Saurischia, Ornithomimosauria) in the Upper Cretaceous beds of Tajikistan. Paleontological Journal. 40(1),103-108.
Averianov and Lopatin, 2021a. A new carnivorous dinosaur (Theropoda, Dromaeosauridae) from the Late Cretaceous of Tajikistan. Reports of the Russian Academy of Sciences. Earth Sciences. 499(1), 49-53.
Averianov and Lopatin, 2021b. A new theropod dinosaur (Theropoda, Dromaeosauridae) from the Late Cretaceous of Tajikistan. Doklady Earth Sciences. 499(1), 570-574.

"Kitadanisaurus" Lambert, 1990
Middle-Late Aptian, Early Cretaceous
Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Material- (Kitadani-ryu) tooth (Dong et al., 1990)
?(FPDM-V96082619) dorsal vertebra (Azuma and Currie, 2000)
?(FPDM-V9812638, 96072901, 97082906, 980815181, 98092604) five teeth (Azuma and Currie, 2000)
?(FPDM-V98081115) humerus (144 mm) (Currie and Azuma, 2006)
?(FPDM-V980801141) manual ungual (Currie and Azuma, 2006)
? several unguals (Currie and Azuma, 2006)
Comments- The original tooth was informally called "Kitadani-ryu", as found in Azuma (1991). Lambert (1990) inappropriately made it into a genus name, listing it as "Kitadanisaurus" in a childrens' book. Dong et al. (1990) published photos of the tooth, labeling it Dromaeosauridae indet.. In 1991, jaw fragments were found in the quarry and identified as dromaeosaurid based on their fused interdental plates. This was followed by the discovery of a manual ungual I, astragalus and metatarsal III in 1993. Azuma and Currie (1995) described these remains in an abstract as those of a giant dromaeosaurid, which was associated in the paleontological community with "Kitadanisaurus" through the late 1990's. Azuma and Currie (2000) later described the skeletal remains as a new taxon of carnosaur, Fukuiraptor. Yet they also noted smaller teeth are known, similar in size and shape to dromaeosaurids and sharing Dromaeosaurus' mesial carina twist, as well as a dorsal vertebra "suggestive of dromaeosaurid affinities." Currie and Azuma (2006) list the teeth which are not referrable to Fukuiraptor, and describe and illustrate a humerus they refer to Dromaeosauridae indet.. In addition, they state several unguals seem to be dromaeosaurid, though these remain undescribed. One manual ungual is listed as having a proximodorsal lip, which is present in most dromaeosaurids as well as some other coelurosaurs. It is probably one of the several, though the only other listed ungual is a distal tip (FPDM-V9912141). Even if these remains are all dromaeosaurid, they do not necessarily belong to the same taxon. "Kitadanisaurus" remains a nomen nudum, as the original tooth has never been described.
References- Dong, Hasegawa and Azuma, 1990. The Age of Dinosaurs in Japan and China. Fukui, Japan: Fukui Prefectural Museum. 65 pp.
Lambert, 1990. The Dinosaur Data Book. New York: Avon Books, 66. ISBN 0-380-75896-3.
Azuma, 1991. Early Cretaceous Dinosaur Fauna from the Tetori Group, central Japan. Research on Dinosaurs from the Tetori Group (1). Professor S. Miura Memorial Volume, 55-69.
Azuma and Currie, 1995. A new giant dromaeosaurid from Japan. Journal of Vertebrate Paleontology. 15(3), 17A.
Azuma and Currie, 2000. A new carnosaur (Dinosauria: Theropoda) from the Lower Cretaceous of Japan. Canadian Journal of Earth Sciences. 37(12), 1735-1753.
Currie and Azuma, 2006. New specimens, including a growth series of Fukuiraptor (Dinosauria, Theropoda) from the Lower Cretaceous Kitadani Quarry of Japan. J. Paleont. Soc. Korea. 22(1), 173-193.

undescribed Dromaeosauridae (Chiba, Ryan, Braman, Eberth, Scott, Brown, Kobayashi and Evans, 2015)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
Material- (RTMP coll.) two teeth
Reference- Chiba, Ryan, Braman, Eberth, Scott, Brown, Kobayashi and Evans, 2015. Taphonomy of a monodominant Centrosaurus apertus (Dinosauria: Ceratopsia) bonebed from the upper Oldman Formation of southeastern Alberta. Palaios. 30, 655-667.

probable Dromaeosauridae indet. (Currie, Rigby and Sloan, 1990)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
Material- (AMNH 5387) manual ungual (Gilmore, 1920)
?(AMNH coll.) three manual unguals (Gilmore, 1920)
(CMN 12413) presacral vertebra (Russell, 1969)
(CMN 12415) presacral vertebra (Russell, 1969)
?(CMN 12438) sacrum (Russell, 1969)
?(CMN 12439) sacrum (Russell, 1969)
(RTMP 80.13.34) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.180) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.20.255) tooth (Currie, Rigby and Sloan, 1990)
(UA 5286) presacral vertebra
Comments- Gilmore (1920) noted four unguals from the Belly River Group of Canada in the AMNH (including AMNH 5387) were nearly identical to the holotype of Coelurus gracilis. Gilmore (1924) believed they belonged to Chirostenotes, but at least AMNH 5387 more closely resembles derived dromaeosaurids in being more curved, having no proximodorsal lip, a more proximally placed flexor tubercle and a proximally undivided longitudinal groove. It is likely Saurornitholestes or Dromaeosaurus, and the other unguals may be as well.
Russell (1969) listed the presacral vertebrae (CMN 12413 and 12414, UA 5286) as being comparable to Oviraptor in their short centra and pleurocoels. The sacra (CMN 12438 and 12439) were tentatively referred to the same taxon, and described as being strongly concave ventrally and containing five vertebrae. The latter resembles dromaeosaurids more closely (oviraptorids have at least six sacrals), many of which also share the presacral characters listed above. They probably belong to Richardoestesia, Dromaeosaurus or Saurornitholestes based on provenance. Another possibility is that they are from juvenile tyrannosaurids.
The three RTMP teeth have a chalky appearence and lack serrations due to being swallowed and digested. They are thus difficult to assign to particular genera, but are near certainly coelurosaurs, and probably dromaeosaurids based on provenence and elongate shape.
References- Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States National Museum with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. United States National Museum Bulletin. 110, l-154.
Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta. Canada Geological Survey, Bulletin 38, geological series 43, 1-13.
Russell, 1969. A new specimen of Stenonychosaurus from the Oldman Formation (Cretaceous) of Alberta. Canadian Journal of Earth Science. 6, 595-612.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.

undescribed Dromaeosauridae (Chapman, Deck, Varricchio and Jackson, 2004)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2240/45084; Morph 4) tooth (?x4x2.6 mm) (Krumenacker et al., 2016)
(IMNH 2251/49826; Morph 4) tooth (8.4x4.2x2.5 mm) (Krumenacker et al., 2016)
(IMNH 2251/50034; Morph 4) tooth (~9.4x~10.6x~5.2 mm) (Krumenacker et al., 2016)
(IMNH 2251/50832; Morph 4) tooth (8.9x5.3x~2.5 mm) (Krumenacker et al., 2016)
(IMNH 2251/50849; Morph 4) tooth (~3x~1.9x? mm) (Krumenacker et al., 2016)
(IMNH 2167/50104; Morph 4) tooth (4.1x2.1x1 mm) (Krumenacker et al., 2016)
caudal vertebrae (Krumenacker, 2005)
material (Chapman, Deck, Varricchio and Jackson, 2004)
Comments- Stated to have a low HCR and high amount of labiolingual compression.
References- Chapman, Deck, Varricchio and Jackson, 2004. Cretaceous Wayan Formation of Idaho: A preliminary report. 24(3), 151-152.
Krumenacker, 2005. Preliminary report on new vertebrate fossils from the Draney Limestone (Aptian) and Wayan Formation (Albian) of east Idaho. Journal of Vertebrate Paleontology. 25(3), 80A.
Krumenacker and Scofield, 2015. A diverse theropod tooth assemblage from the Mid-Cretaceous (Albian-Cenomanian) Wayan Formation of Idaho. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 158.
Krumenacker, Simon, Scofield and Varricchio, 2016. Theropod dinosaurs from the Albian-Cenomanian Wayan Formation of eastern Idaho. Historical Biology. DOI: 10.1080/08912963.2015.1137913

undescribed possible dromaeosaurid (Turner and Peterson, 1999)
Late Oxfordian-Tithonian, Late Jurassic
Mother's Day Quarry, Morrison Formation, Montana, US
Material- tooth
Comments- Turner and Patterson reference a dromaeosaur tooth from this undescribed quarry.
Reference- Turner and Peterson, 1999. Biostratigraphy of dinosaurs in the Upper Jurassic Morrison Formation of the Western Interior U.S.A. in Gillette (ed). Vertebrate Paleontology in Utah. Utah Geological Survey Miscellaneous Publication. 99-1, 77-114.

undescribed dromaeosaurid (Ullmann, Varricchio, Knell and Lacovara, 2010)
Albian, Early Cretaceous
Vaughn Member of the Blackleaf Formation, Montana, US
Reference- Ullmann, Varricchio, Knell and Lacovara, 2010. Taphonomy and taxonomy of a vertebrate microsite in the Cretaceous Blacklaef Formation in Southwest Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 179A.

undescribed Dromaeosauridae (MOR online)
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
Material- (MOR 721) partial skeleton
Comments- This probably belongs to Saurornitholestes and/or Bambiraptor.

undescribed Dromaeosauridae (MOR online)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Material- (MOR 029) teeth (MOR online)
(MOR 040) ungual fragment (MOR online)
(MOR 119) digit, two phalanges (MOR online)
(MOR 126) manual ungual (MOR online)
(MOR 156) tooth (MOR online)
(MOR 176) tooth (MOR online)
(UCMP 129723) maxillary fragment (UCMP online)
(UCMP 140582) tooth (UCMP online)
(UCMP 154576) ungual (UCMP online)
(UCMP 154577) phalanx (UCMP online)
(UCMP 154578) phalanx (UCMP online)
(YPM PU 22301) (YPM online)
(YPM PU 22302) (YPM online)
Comments- These remains likely belong to Richardoestesia, Paronychodon, Saurornitholestes and/or Dromaeosaurus.

unnamed Dromaeosauridae (Triebold and Russell, 1995)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (MOR 059) tooth (MOR online)
(MOR 061) ungual (MOR online)
(UCMP 64139) phalanx (UCMP online)
(UCMP 64140) phalanx (UCMP online)
(UCMP 119675) tooth (UCMP online)
(UCMP 119679) pedal ungual (UCMP online)
(UCMP 119751) tooth (UCMP online)
(UCMP 119924) three teeth (UCMP online)
(UCMP 119925) four teeth (UCMP online)
(UCMP 119926) tooth (UCMP online)
(UCMP 120077) two teeth (UCMP online)
(UCMP 120256) two teeth (UCMP online)
(UCMP 120304) ungual (UCMP online)
(UCMP 123338) over twenty teeth (UCMP online)
(UCMP 123339) twenty-four teeth (UCMP online)
(UCMP 123505) tooth (UCMP online)
(UCMP 123506) tooth (UCMP online)
(UCMP 123507) three teeth (UCMP online)
(UCMP 123525) two teeth (UCMP online)
(UCMP 123544) tooth (UCMP online)
(UCMP 123566) tooth (UCMP online)
(UCMP 124989) tooth (UCMP online)
(UCMP 129085) two teeth (UCMP online)
(UCMP 134801) tooth (UCMP online)
(UCMP 145737) tooth (UCMP online)
(UCMP 145877) two teeth (UCMP online)
(UCMP 145968) tooth (UCMP online)
(UCMP 174752) tooth (UCMP online)
(UCMP 174796) tooth (UCMP online)
(UCMP 186838) tooth (UCMP online)
(UCMP 186870) teeth (UCMP online)
(UCMP 186874) teeth (UCMP online)
(UCMP 186877) teeth (UCMP online)
(UCMP 186882) teeth (UCMP online)
(UCMP 186893) teeth (UCMP online)
(UCMP 186899) tooth (UCMP online)
(UCMP 186903) teeth (UCMP online)
(UCMP 186909) teeth (UCMP online)
(UCMP 186912) teeth (UCMP online)
(UCMP 186929) teeth (UCMP online)
(UCMP 186944) teeth (UCMP online)
(UCMP 186954) teeth (UCMP online)
(UCMP 186956-186962) seven teeth (UCMP online)
(UCMP 186963) teeth (UCMP online)
(UCMP 186964) tooth (UCMP online)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Material- (FMNH PR2896) tooth (6.5x3.1x1.3 mm) (Gates, Zanno and Makovicky, 2015)
material (Triebold and Russell, 1995)
material (Jacobsen and Sroka, 1998)
Comments- The above material has merely been identified as Dromaeosauridae. Acheroraptor and largely dental material referred to Saurornitholestes and Dromaeosaurus have been identified in the Hell Creek Formation, so these specimens may belong to those taxa.
References- Triebold and Russell, 1995. A new small dinosaur locality in the Hell Creek Formation: Journal of Vertebrate Paleontology. 15(3), 57A.
Jacobson and Sroka, 1998. Preliminary assement of a Hell Creek Dinosaurian Fauna from sites in Corson County, South Dakota. Journal of Vertebrate Paleontology. 18(3), 53A.
Gates, Zanno and Makovicky, 2015. Theropod teeth from the upper Maastrichtian Hell Creek Formation "Sue" Quarry: New morphotypes and faunal comparisons. Acta Palaeontologica Polonica. 60(1), 131-139.

undescribed Dromaeosauridae (AMNH online)
Cretaceous
Montana, US
Material- (AMNH 2363) six teeth
Comments- These are listed on the AMNH wbsite as cf. Velociraptor sp., but are more likely another basal dromaeosaurid taxon based on provenence.

undescribed Dromaeosauridae (UCMP online)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- (UCMP 120851) four teeth
(UCMP 120852) over thirty-five teeth

unnamed Dromaeosauridae (Ostrom, 1969)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- (UCMP 73079) phalanx (UCMP online)
(UCMP 186861) teeth (UCMP online)
(UCMP 186862) teeth (UCMP online)
(UCMP 186888) teeth (UCMP online)
(UCMP 186891) tooth (UCMP online)
(UCMP 186901) teeth (UCMP online)
(UCMP 186917) teeth (UCMP online)
(UCMP 186921) teeth (UCMP online)
(YPM PU 20589) proximal pedal phalanx II-2 (Ostrom, 1969)
(YPM PU 55525) (YPM online)
Comments- Ostrom (1969) notes PU 20589 is almost identical to the element in Deinonychus except that the proximoventral heel is a third larger. He considered it an indeterminate new dromaeosaurid taxon.
The UCMP specimens and YPM PU 55525 are merely listed as Dromaeosauridae.
References- Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin. 30, 1-165.

undescribed possible Dromaeosauridae (Chure, Madsen and Britt, 1993)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of Morrison Formation, Utah, US
Material- teeth
Comments- Chure et al. (1993) reported small teeth with large hooked distal serrations and small or absent mesial serrations.
References- Chure, Madsen and Britt, 1993. New data on theropod dinosaurs from the Late Jurassic Morrison FM. (MF). Journal of Vertebrate Paleontology. 13(3), 30A.
Chure, 1995. The teeth of small theropods from the Morrison Formation (Upper Jurassic: Kimmeridgian), UT. Journal of Vertebrate Paleontology. 15(3), 23A.

unnamed dromaeosaurid (Senter, Kirkland, Deblieux and Madsen, 2010)
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Material- (UMNH VP 20209) proximal caudal vertebra (45 mm), six distal caudal vertebrae, distal caudal vertebra (41 mm), distal caudal vertebra (~40 mm), distal caudal vertebra (~43 mm), distal caudal vertebra (~46 mm), distal chevrons
Comments- This was initially identified (Senter et al., 2010) as part of what would later be named Yurgovuchia by Senter et al. (2012).
References- Senter, Kirkland, Deblieux and Madsen, 2010. Three new theropods from the Cedar Mountain Formation (Lower Cretaceous) of Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 162A.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.

undescribed Dromaeosauridae (Kirkland, 1996)
Early Albian, Early Cretaceous
Dinosaur National Monument, Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- partial skeleton including mid dorsal vertebra, femur (~290 mm), astragalus and pes (Chure, Britt and Scheetz, 2007)
multiple elements (Chure, Britt and Scheetz, 2007)
Early Albian, Early Cretaceous
Hotel Mesa Site, Ruby Ranch Member of Cedar Mountain Formation, Utah, US
(OMNH coll.) teeth (Kirkland, 1996)
Aptian, Early Cretaceous
Long Walk Quarry, Lower Ruby Ranch Member of Cedar Mountain Formation, Emery County, Utah, US
(OMNH coll.) pedal ungual ?III (DeCourten, 1998)
Early Albian, Early Cretaceous
Lorrie's Site, Dinosaur National Monument, Ruby Ranch Member of Cedar Mountain Formation, Utah, US
(DMNH coll.) manual ungual (Kirkland, online 2007)
Comments- Kirkland (1996) initially reported dromaeosaurid teeth from the "middle fauna" of the Cedar Mountain Formation, seemingly referring to "teeth possibly referable to the dromaeosaurid Deinonychus" from Hotel Mesa mentioned by Kirkland et al. (1997).
DeCourten (1998) wrote "A large curving foot claw demonstrates the presence of dromaeosaurs. This dromaeosaur was much smaller than Utahraptor but probably represents a carnivore much like Deinonychus from the Cloverly Formation of Montana. Until more dromaeosaur material is found at the Long Walk Quarry scientists cannot be sure about the exact identity of this theropod, but Deinonychus is a good guess at this point because the claw is almost identical to those of that genus (Figure 7-12A) and the Cloverly Formation is very nearly the same age as the Ruby Ranch Member of the Cedar Mountain Formation." Unfortunately, Figure 7-12A is only a redrawing of Ostrom's illustration of Deinonychus' pedal ungual III, said to be "very similar to that found in the Long Walk Quarry." So perhaps the Long Walk specimen is also from digit III.
Chure et al. (2007) reported a non-Utahraptor taxon from Dinosaur National Monument "represented by a partial skeleton plus isolated elements of a second individual. Femoral length is ~290 mm. The single mid-dorsal bear a pneumatic foramen on its centrum. The Monument specimen differs from Utahraptor in the extreme height of the ascending process of the astragalus plus its symmetrical shape, with the apex at mid-width."
Carpenter (online 2007) stated "A single hand claw of Utahraptor is known from near the base of the Ruby Ranch at Lorrie's Site", and "The foot of Deinonychus is known from high in the Ruby Ranch at Dinosaur National Monument" which probably refers to the partial skeleton.
References- Kirkland, 1996. Biogeography of western North America's Mid-Cretaceous dinosaur faunas: Losing European ties and the first great Asian-North American interchange. Journal of Vertebrate Paleontology. 16(3), 45.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the central Colorado plateau: A key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young University Geology Studies. 42, 69-103.
DeCourten, 1998. Dinosaurs of Utah. University of Utah Press. 300 pp.
Chure, Britt and Scheetz, 2007. Sickle-claw theropod dinosaurs of the Lower Cretaceous Cedar Mountain Formation from the Dalton Wells Quarry and Dinosaur National Monument, Utah. Journal of Vertebrate Paleontology. 27(3), 59A.
Carpenter, online 2007. https://web.archive.org/web/20090517064058/http://scientists.dmns.org/kenCarpenter/cedar-mountain-project/dinosaurs-of-the-cedar-mountain-formation/

Dromaeosauridae indet. (Kirkland, Lucas and Estep, 1998)
Late Cenomanian, Late Cretaceous
Dakota Formation, Utah, US
Material- teeth
Comments- These remains were listed as Velociraptorinae indet. by Kirkland et al. (1998) and Eaton et al. (1999). They are presumably one of the two Dromaeosauridae indet. gen. et sp. teeth listed by Kirkland et al. (1997).
References- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Eaton, Cifelli, Hutchison, Kirkland and Parrish, 1999. Cretaceous vertebrate faunas from the Kaiparowits Plateau, south central Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 345-353.

unnamed Dromaeosauridae (Nelson and Crooks, 1987)
Cenomanian-Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71599) two teeth (Fiorillo, 1999)
(FHSU coll.) two teeth (~1.1x~1.2x? mm, ~2.3x~2.1x? mm) (Nelson and Crooks, 1987)
(NCSM 33267) lateral tooth (10.15x5.03x3.32 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
(NCSM 33275) lateral tooth (5.83x3.23x1.36 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
teeth (Kirkland et al., 1997)
partial skeleton (Cifelli et al., 1999)
Comments- Nelson and Crooks (1987) figure two teeth as "Theropoda, Troodontidae (= Saurornithoididae)" from the Rough Road Quarry. Currie et al. (1990) said these "are more likely from a velociraptorine because the denticles are too small (there are 12 posterior denticles per millimeter in their figures) and elongate."
Material was called cf. Deinonychus sp. nov. by Kirkland et al. (1997), and noted to be large. Fiorillo (1999) described two teeth which were strongly compressed, lack mesial serrations, and have distal serrations which are taller than wide and are straight with a slight apically oriented hook. He referred these to Velociraptorinae. Cifelli et al. (1999) state a partial skeleton is known, though do not say if it is of the "velociraptorine" or "dromaeosaurine" taxon. Cifelli et al. also list Velociraptorinae indet. teeth.
Avrahami et al. (2016) reported "four complete and several fragmentary dromaeosaur teeth from an as-yet indeterminate species" from the Cliffs of Insanity microsite, which by the time of the material's description (Avrahamai, 2018; Avrahami et al., 2018) included NCSM 33267 and 33275, and perhaps also Richardoestesia teeth NCSM 33274 and 33288.
References- Nelson and Crooks, 1987. Stratigraphy and paleontology of the Cedar Mountain Formation (Lower Cretaceous), eastern Emery County, Utah. In Averett (ed.). Paleontology and Geology of the Dinosaur Triangle: Guidebook for 1987 Field Trip. Museum of Western Colorado. 55-63.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press. 107-125.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial Cretaceous vertebrates from the Cedar Mountain Formation, Emery County, Utah: the Mussentuchit Local Fauna. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison Eggshell site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 259-268.
Avrahami, Humphrey, Heckert, Gates, Makovicky and Zanno, 2016. The early Late Cretaceous paleofaunal vertebrate assemblage of the Cliffs of Insanity microsite in the Mussentuchit Member of the Cedar Mountain Formation, Utah. Journal of Vertebrate Paleontology. Program and Abstracts, 90.
Avrahami, 2018. Paleobiodiversity of a new microvertebrate locality from the Upper Cretaceous Mussentuchit Member, Cedar Mountain Formation, Utah: Testing morphometric multivariate approaches for quantifying shape variation in microvertebrate specimens. Masters thesis, North Carolina State University. 181 pp.
Avrahami, Gates, Heckert, Makovicky and Zanno, 2018. A new microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain Formation: Insights into the paleobiodiversity and paleobiogeography of early Late Cretaceous ecosystems in western North America. PeerJ. 6:e5883.

Dromaeosauridae indet. (Kirkland, Lucas and Estep, 1998)
Middle-Late Turonian, Late Cretaceous
Smoky Hollow Member of the Straight Cliffs Formation, Utah, US
Material- (MNA 994) tooth (Parrish, 1999)
(OMNH 24439) tooth (Parrish, 1999)
(OMNH 24441) tooth (Parrish, 1999)
(OMNH 24442) tooth (Parrish, 1999)
Comments- These were listed as Velociraptorinae by Kirkland et al. (1998) and Parrish (1999).
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

Dromaeosauridae indet. (Kirkland, Lucas and Estep 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Material- (OMNH 21238) tooth (Parrish, 1999)
(OMNH 21673) tooth (Parrish, 1999)
? material (Eaton et al., 1999)
Comments- These were listed as Velociraptorinae by Kirkland et al. (1998) and Parrish (1999). In addition, Eaton et al. (1999) listed ?Dromaeosauridae indet. from a Santonian possible Straight Cliffs (or Wahweap?) locality.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Eaton, Diem, Archibald, Schierup and Munk, 1999. Vertebrate paleontology of the Upper Cretaceous rocks of the Markagunt Plateau, southwestern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 323-333.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

possible undescribed dromaeosaurid (Eaton, 1999)
Santonian-Campanian, Late Cretaceous
Iron Springs Formation, Utah, US
Material- tooth
Reference- Eaton, 1999. Vertebrate paleontology of the Iron Springs Formation, Upper Cretaceous, southwestern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 339-343.

Dromaeosauridae indet. (Kirkland, Lucas and Estep 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah, US
Material- (UCM 8613) tooth (Parrish, 1999)
? material (Eaton et al., 1999)
Comments- UCM 8613 was listed as Velociraptorinae by Kirkland et al. (1998) and Parrish (1999). In addition, Eaton et al. (1999) listed ?Dromaeosauridae indet. from an Early Campanian possible Wahweap locality.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Eaton, Diem, Archibald, Schierup and Munk, 1999. Vertebrate paleontology of the Upper Cretaceous rocks of the Markagunt Plateau, southwestern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 323-333.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

Dromaeosauridae indet. (Parrish, 1999)
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Material- (MNA HM-6) tooth (Parrish, 1999)
(OMNH 21240) tooth (Parrish, 1999)
(OMNH 24158) tooth (Parrish, 1999)
(UCM 8312, 83240 (in part), 8642 (in part), 8655, 8659 (in part), 8663) six teeth (Parrish, 1999)
Comments- The teeth were listed as Velociraptorinae by Parrish (1999). UCMP 149171 was listed in the UCMP online database as dromaeosaurid, but is a troodontid (Zanno et al., 2011).
References- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.
Zanno, Varricchio, O’Connor, Titus and Knell, 2011. A new troodontid theropod, Talos sampsoni gen. et sp. nov., from the Upper Cretaceous western interior basin of North America. PLoS ONE. 6(9), e24487.

unnamed Dromaeosauridae (Bonde, 2008)
Albian, Early Cretaceous
Willow Tank Formation, Nevada, US
Material- teeth (?x7x3.5, ?x6x3.5 mm), incomplete femur, partial tibia
Reference- Bonde, 2008. Paleoecology and taphonomy of the Willow Tank Formation (Albian), southern Nevada. Masters thesis, Montana State University. 96 pp.

Dromaeosauridae indet. (Ratkevich and Duffek, 1996)
Campanian, Late Cretaceous
Fort Crittenden Formation, Arizona, US
Material- two teeth
Comments- These were identified as cf. Richardoestesia and cf. Saurornitholestes by Ratkevich and Duffek (1996), but reidentified merely as Dromaeosauridae indet. by Sullivan and Lucas (2006).
References- Ratkevich and Duffek, 1996. Small macro-and large micro-vertebrate fauna of the Fort Crittenden Formation, Southeast Arizona. Proceedings of Southwest Paleontological Society and Mesa Southwest Museum, Mesa, Arizona. 4, 115-120.
Sullivan and Lucas, 2006. The Kirtlandian land-vertebrate "age" - faunal composition, temporal position and biostratigraphic correlation in the nonmarine Upper Cretaceous of western North America. New Mexico Museum of Natural History and Science Bulletin. 35, 7-29.

undescribed dromaeosaurid (Lucas et al., 1988)
Mid Santonian, Late Cretaceous
Point Lookout Sandstone, New Mexico, US
Material- (NMMNH P-27481) tooth (8.9x5.2x2.7 mm) (Williamson and Brusatte, 2014)
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.

Dromaeosauridae indet. (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-33147) tooth
References- Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.

Dromaeosauridae indet. (Lehman, 1981)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- ?(Lafon coll.) metatarsal I (Lehman, 1981)
(NMMNH P-32814) tooth (5.2x3.3x1.6 mm) (Williamson and Brusatte, 2014)
(SMP VP-2505) tooth (34 mm) (Jasinski, Sullivan and Lucas, 2011)
(SMP VP-2595) tooth (14 mm) (Jasinski, Sullivan and Lucas, 2011)
Comments- Williamson and Brusatte (2014) state NMMNH P-32814 is more similar to Acheroraptor than Saurornitholestes due to rounded distal serrations and lingual ridges.
References- Lehman, 1981. The Alamo Wash local fauna: A new look at the old Ojo Alamo fauna. In Lucas, Rigby and Kues (eds.). Advances in San Juan Basin paleontology. University of New Mexico Press. 189-221.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Jasinski, Sullivan and Lucas, 2011. Taxonomic composition of the Alamo Wash local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann (eds.). Fossil Record 3. New Mexico Museum of Natural History and Science Bulletin. 53, 216-271.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.

undescribed Dromaeosauridae (Fix, Darrough, Parris and Grandstaff, 2012)
Campanian, Late Cretaceous
Chronister site, Missouri, US
Reference- Fix, Darrough, Parris and Grandstaff, 2012. Western Appalachia Dinosauria and associated vertebrates of the Late Cretaceous of Southeast Missouri. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 94.

undescribed Dromaeosauridae (Martin and Brett-Surman, 1992 unpublished)
Late Aptian, Early Cretaceous
Dinosaur Park / Cherokee-Sanford Brick Clay Pit / Muirkirk Clay Pit USNM 41614, Arundel Formation, Prince George's County, Maryland, US
Material- (NHRD-AP 2011.v.267) tooth (Dinosaur Fund online)
(NHRD-AP 2012.v.601) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.104) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.105) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.131) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.132) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.133) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.134) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.135) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.136) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.137) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.138) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.139) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.140) tooth (Dinosaur Fund online)
(NHRD-AP 2014.s.141) tooth (Dinosaur Fund online)
(USNM 437630) tooth (Martin and Brett-Surman, 1992 unpublished)
(USNM 529428) tooth (USNM online)
(USNM 604958) tooth (USNM online)
(USNM 604963) tooth (USNM online)
(USNM 604983) tooth (USNM online)
(USNM 604985) partial tooth (USNM online)
(USNM 604987) premaxillary tooth (USNM online)
Late Aptian, Early Cretaceous
Arundel Formation, Prince George's County, Maryland, US
(USNM 508535) tooth (USNM online)
Comments- Numerous teeth have been catalogued as dromaeosaurid from the Arundel Formation, but have not been published or referred to Deinonychus or "Dromaeosaurus" gracilis. Martin and Brett-Surman (1992 unpublished) figured USNM 437630 as a theropod tooth, which the USNM online catalog indicates was discovered in 1989 and identified as dromaeosaurid by Kirkland. Among the other USNM teeth, USNM 508535 was found in 2001 and ID'd by Brett-Surman, 529428 in 2005 and ID'd by Carrano, 604958 in 2012 and also ID'd by Carrano, and 604963, 604983, 604985 and 604987 were all discovered from 2010-2013 and ID'd by Miller (604963) and Carrano (the others). As for the NHRD-AP specimens, NHRD-AP 2011.v.267 was found in 2011, NHRD-AP 2012.v.601 in 2012, and 2014.s.104-105 and 131-141 in 2013.
Reference- Martin and Brett-Surman, 1992 unpublished. A preliminary report on new dinosaur material from the Arundel Clay (Lower Cretaceous, Late Aptian-Early Albian) of Maryland. Smithsonian Institution. 31 pp.

undescribed Dromaeosauridae
?Late Cretaceous
?North America
Material- (AMNH 86344) (AMNH online)
(MOR 140) teeth (MOR online)
(MOR 157) tooth (MOR online)
(YPM 56980) (YPM online)
(YPM 56999) (YPM online)
(YPM PU 55010) (YPM online)
(YPM PU 55011) (YPM online)

undescribed Dromaeosauridae (Rivera-Sylva, Frey, Stinnesbeck, Padilla Gutierrez, Gonzalez Gonzalez and Amezcua Torres, 2015)
Late Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Material- long bone fragments and phalanx
Reference- Rivera-Sylva, Frey, Stinnesbeck, Padilla Gutierrez, Gonzalez Gonzalez and Amezcua Torres, 2015. The Late Cretaceous Las Aguilas dinosaur graveyard, Coahuila, Mexico. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 203.

undescribed dromaeosaurid (Metcalf, Vaughan, Benton, Cole, Simms and Dartnall, 1992)
Early Bathonian, Middle Jurassic
Chipping Norton Formation, England
Material- (GLRCM coll.; F) tooth (3.5 mm)
Comments- This was labeled a dromaeosaur-like tooth and said to be a possible troodontid. It is elongate and moderately recurved, with mesial serrations present apically. Serrations on both carinae are low and rounded, with a DSDI of ~2. The base is not constricted and blood pits are absent. Serration density is 6/mm distally and 13/mm mesially.
This tooth differs from troodontids in lacking a basal constriction and blood pits, as well as having such a high DSDI. The latter is only known in some derived dromaeosaurids and Richardoestesia, but the low serration density excludes it from the latter genus. However, the low rounded morphology of the distal serrations is not seen in other dromaeosaurids.
Reference- Metcalf and Walker, 1994. A new Bathonian microvertebrate locality in the English Midlands. in Fraser and Sues (eds.). In the Shadow of the Dinosaurs- Mesozoic Small Tetrapods, Cambridge (Cambridge University Press). 322-332.

undescribed Dromaeosauridae (Wills, 2015)
Bathonian, Middle Jurassic
Forest Marble Formation, England
Material- (small) sixteen teeth
Reference- Wills, 2015. Isolated dromaeosaurid teeth from the Bathonian (Middle Jurassic) of Dorset, United Kingdom. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 238.

undescribed possible dromaeosaurid (Naish pers. comm. to Sweetman, 2004)
Barremian-Early Aptian, Early Cretaceous
Wealden Group, England
Material- fragmentary remains.
Comments- Sweetman commented on these remains, attributed to Naish pers. comm. 2003.
Reference- Sweetman, 2004. The first record of velociraptorine dinosaurs (Saurischia, Theropoda) from the Wealden (Early Cretaceous, Barremian) of southern England. Cretaceous Research. 25, 353-364.

unnamed Dromaeosauridae (Sweetman, 2004)
Barremian, Early Cretaceous
Wessex Formation, England
Material- (IWCMS 2002.1) lateral tooth (16 mm)
(IWCMS 2002.2) lateral tooth (8.5 mm)
(IWCMS 2002.3) lateral tooth
(IWCMS 2002.4) lateral tooth
(NHMUK R 16510) lateral tooth (>21.5 mm)
Comments- These teeth are moderately to strongly recurved and compressed, with BW/FABL ratios of .4-.6. Mesial serrations are present only apically, and absent in IWCMS 2002.2. There are 3.8-4.8/mm. Distal serrations number 2.7-3.5/mm in most specimens, are rounded and inclined apically. IWCMS 2002.2 has 6.25/mm however, and may be a different taxon. Sweetman (2004) assigned them to Velociraptorinae based on the high DSDI (1.37-1.6)
Reference- Sweetman, 2004. The first record of velociraptorine dinosaurs (Saurischia, Theropoda) from the Wealden (Early Cretaceous, Barremian) of southern England. Cretaceous Research. 25, 353-364.

undescribed Dromaeosauridae (Royo y Gómez, 1927)
Tithonian-Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (CPT-1283) tooth (5.4x4.1x2.3 mm) (Gasco et al., 2012)
(CPT-1327) tooth (14.6x8.4x4 mm) (Gasco et al., 2012)
(CPT-1433) tooth (~10.7x5.1x3.1 mm) (Gasco et al., 2012)
(CPT-1443) tooth (5.5x4.2x2.3 mm) (Gasco et al., 2012)
(CPT-1658) tooth (6.2x3.7x2.4 mm) (Gasco et al., 2012)
(CPT-1659) tooth (6.8x4.3x2.6 mm) (Gasco et al., 2012)
(CPT-2126) tooth (6.3x4x2.1 mm) (Gasco et al., 2012)
(MNCN coll.) tooth (Royo y Gómez, 1927)
Comments- MNCN coll. was originally identified by Royo y Gomez (1927) as Megalosaurus cf. dunkeri, then as a dromaeosaurid by Ruiz-Omeñaca and Pereda-Suberbiola (1999).
References- Royo y Gómez, 1927. Sesión del 6 de Julio de 1927. Boletín de la Real Sociedad Española de Historia Natural. 27.
Ruiz-Omeñaca and Pereda-Suberbiola, 1999. Un documento inédito de Royo y Gómez sobre los dinosaurios del Levante. Temas Geológico-Mineros ITGE. 26, 111-112.
Gascó, Cobos, Royo-Torres, Mampel and Alcalá, 2012. Theropod teeth diversity from the Villar del Arzobispo Formation (Tithonian-Berriasian) at Riodeva (Teruel, Spain). Palaeobiodiversity and Palaeoenvironments. 92(2), 273-285.

unnamed dromaeosaurid (Rauhut and Zinke, 1995)
Barremian, Early Cretaceous
Una Formation, Spain
Material- (IPFUB Una Th 21-36, 38, 40-47, 50-52, 63, 65, 70) sixty-six teeth
(IPFUB Una Th 66) tooth (15 mm)
Comments- These teeth are strongly recurved and highly compressed. Distal serrations are rounded or slightly inclined apically. Mesial serrations are much smaller when present (DSDI 1.11-2). A few have faint longitudnal ridges on both sides. Rauhut (2002) referred them to Velociraptorinae due to the high DSDI, large serrations (~7-7.5/mm) and strong recurvature.
References- Rauhut and Zunke, 1995. A description of the Barremian dinosaur fauna from Una with a comparison of that of Las Hoyas. II. International Symposium of Lithographic Limestone, Extended Abstracts. 123-126.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca, Spain. Cretaceous Research. 23, 255-263.

unnamed dromaeosaurid (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
Late Campanian, Late Cretaceous
Figuerola 2, Tremp Formation, Spain
Material- (DPM-FIG2-T1+T2) tooth (12.1x6.7x4.3 mm)
Comments- This was listed as Dromaeosauridae indet. 2 by Torices (2002), and Dromaeosauridae indet. by Torices et al. (2015).
Reference- Torices, 2002. Los dinosaurios terópodos del Cretácico Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Paleontología. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.

unnamed dromaeosaurid (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
Late Campanian, Late Cretaceous
Laño, Sedano Formation, Spain
Material- (MCNA 14622) tooth (3.4x2.6x1.3 mm)
Reference- Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.

undescribed dromaeosaurid (Company, Torices, Pereda-Suberbiola and Ruiz-Omenaca, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Sierra Perenchiza Formation, Valencia, Spain
Material- two teeth (~10 mm)
Comments- Described as strongly compressed labiolingually, slightly recurved distally, and both mesial and distal carinae have minute serrations (~7 per mm).
Reference- Company, Torices, Pereda-Suberbiola and Ruiz-Omenaca, 2009. Theropod teeth from the Late Cretaceous of Chera (Valencia, Eastern Spain). Journal of Vertebrate Paleontology. 29(3), 81A.

undescribed Dromaeosauridae (Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- teeth, several postcranial elements
Comments- Ortega et al. (2009) state teeth of Velociraptorinae and dromaeosaurid postcrania are present. Torices et al. (2011) mention both cf. Dromaeosauridae indet. and cf. Velociraptorinae indet..
Reference- Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009. The vertebrate diversity of the Upper Campanian-Lower Maastrichtian "Lo Hueco" fossil-site (Cuenca, Spain). Journal of Vertebrate Paleontology. 29(3), 159A-160A.
Torices, Barroso-Barcenilla, Cambra-Moo, Perez and Serrano, 2011. Vertebrate microfossil analysis in the palaeontological site of 'Lo Hueco' (Upper Cretaceous, Cuenca, Spain). Journal of Vertebrate Paleontology. Program and Abstracts 2011, 205.

unnamed Dromaeosauridae (Torices, 2002)
Early Maastrichtian, Late Cretaceous
Fontllonga 6, Tremp Formation, Spain
Material- (DPM-FON6-T1) tooth
(DPM-FON6-T2) tooth (16.3x7.5x5.8 mm)
Comments- DPM-FON6-T1 was noted as Dromaeosauridae indet. 1 by Torices (2002), though not mentioned in Torices et al. (2015). DPM-FON6-T2 was listed by Torices as Dromaeosauridae indet. 2, and by Torices et al. as Dromaeosauridae indet..
Reference- Torices, 2002. Los dinosaurios terópodos del Cretácico Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Paleontología. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.

unnamed dromaeosaurid (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
Late Maastrichtian, Late Cretaceous
Blasi, Aren Formation, Spain
Material- (MPZ2004/6) tooth (17.5x11x4.4 mm)
Reference- Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.

unnamed dromaeosaurid (Garcia, Pincemaille, Vianey-Liaud, Marandat, Lorenz, Cheylan, Capetta, Michaux and Sudre, 1999)
Early Maastrichtian, Late Cretaceous
Vitrolles-Couperigne, Provence, France
Material- few teeth
Comments- Garcia et al. (1999) provisionally assign a few teeth to Dromaeosauridae. The illustrated tooth is highly recurved, not constricted basally, seems to lack mesial serrations, while having 2-3 distal serrations per mm. Distal serrations are hooked apically. They therefore most closely resemble "velociraptorine" dromaeosaurids.
Reference- Garcia, Pincemaille, Vianey-Liaud, Marandat, Lorenz, Cheylan, Capetta, Michaux and Sudre, 1999. Decouverte du premier squelette presque complet de Rhabdodon priscus (Dinosauria, Ornithopoda) du Maastrichtian inferieur de Provence. Les Comptes rendus de l'Académie des sciences. 328: 415-21.

unnamed dromaeosaurid (Deperet, 1899)
Maastrichtian, Late Cretaceous
Gres de Saint-Chinian, Herault, France
Material- maxillary fragment, partial mandible, teeth, manual phalanges, fibula
Comments- Deperet (1899) referred remains from the Masstrichtian of France to Dryptosaurus. These were later referred to Megalosaurus pannoniensis by Lapparent (1946), and then to Dromaeosauridae by Allain and Taquet (2000).
References- Deperet, 1899. Aperçu sur la géologie du chaïnon de Saint-Chinian. Bulletin de la Société Géologique de France. 27, 686-709.
Lapparent, 1947. Les dinosauriens du Crétacé Supérieur du Midi de la France. Mémoires de la Société Géologique de France. 56, 1-54.
Allain and Taquet, 2000. A new genus of Dromaeosauridae (Dinosauria, Theropoda) from the Upper Cretaceous of France. Journal of Vertebrate Paleontology. 20(2), 404-407.

unnamed possible dromaeosaurid (Laurent, Cavin and Bilotte, 1999)
Late Maastrichtian, Late Cretaceous
Lestaillats Mars Formation, France
Material- tooth fragment
Comments- Laurent et al. (1999) report a small serrated, laterally compressed tooth fragment that they believe may come from a dromaeosaurid.
Reference- Laurent, Cavin and Bilotte, 1999. A new Late Maastrichtian vertebrate locality in the French Petites-Pyrenees. Les Comptes rendus de l'Académie des sciences. 328, 781-787.

undescribed Dromaeosauridae (Lubbe, Richter and Knotschke, 2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry, Germany
Material- (DFMMh/FV 383) tooth (9.7x6.9x3.1 mm) (Lubbe, Richter and Knötschke, 2009)
(DFMMh/FV 530) tooth (7.2x5.7x2.7 mm) (Lubbe, Richter and Knötschke, 2009)
(DFMMh/FV 705) tooth (xx mm) (Lubbe, Richter and Knötschke, 2009)
Comments- While seven teeth assigned to Velociraptorinae by Lubbe et al. (2009), Gerke and Wings (2014) found four of these were Neotheropoda indet., megalosaurid and tyrannosauroid. Based on the information in Lubbe et al., FV 658 may be the megalosaurid (larger, elongate crown, more of mesial carina serrated), FV 382 and 790.5 may be tyrannosauroid (less recurved and thicker labiolingually), and FV 707.1 may be indet. (larger and only partially preserved).
References- Lubbe, Richter and Knötschke, 2009. Velociraptorine dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany. Acta Palaeontologica Polonica. 54(3), 401-408.
Gerke and Wings, 2014. Characters versus morphometrics: A case study with isolated theropod teeth from the Late Jurassic of Lower Saxony, Germany, reveals an astonishing diversity of theropod taxa. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 137.

unnamed Dromaeosauridae (Lanser and Heimhofer, 2015)
Late Barremian-Early Aptian, Early Cretaceous
Balve-Beckum quarry, Germany
Material- (LWL MN Ba 5) partial tooth
(LWL MN Ba 6) tooth fragment
(LWL MN Ba 7) tooth fragment
(LWL MN Ba 9) tooth fragment
(LWL MN Ba 10) tooth fragment
(LWL MN Ba 11) tooth fragment
(LWL MN Ba 12) incomplete tooth (?x5x3.8 mm)
(LWL MN Ba 13) tooth (12.8x7.4x4.5 mm)
Reference- Lanser and Heimhofer, 2015. Evidence of theropod dinosaurs from a Lower Cretaceous karst filling in the northern Sauerland (Rhenish Massif, Germany). Paläontologische Zeitschrift. 89(1), 79-94.

unnamed possible Dromaeosauridae (Lindgren, Currie, Rees, Siverson, Lindström and Alwmark, 2008)
Early Berriasian, Early Cretaceous
Skyttegaard Member of the Rabekke Formation, Denmark
Material- (MGUH 28403) incomplete tooth (5x3.5x1.3 mm)
(MGUH 28404) incomplete tooth (4.1x2.5x1.4 mm)
(MGUH 28405) (juvenile?) incomplete tooth (1.5x.1.2x.8 mm)
(MGUH 28406) partial tooth
(MGUH 28407) tooth fragment
(MGUH 28408) tooth fragment
(MGUH 28409) tooth
Comments- Lindgren et al. (2008) referred these to three morphotypes- 28403, 28404 and 28408 are said to be a basal velociraptorine, 28406 and 28407 might be conspecific but differ in serration morphology, and 28405 was a possible dromaeosaurine. Bonde (2012) on the other hand referred 28406 and 28407 to Velociraptorinae, 28403 and 28404 to basal dromaeosaurids, and 28405 to Dromaeosaurinae. The latter also considered 28409 as Dromaeosauridae incertae sedis.
References- Lindgren, Currie, Rees, Siverson, Lindström and Alwmark, 2008. Theropod dinosaur teeth from the lowermost Cretaceous Rabekke Formation on Bornholm, Denmark. Geobios. 41, 253-263.
Bonde, 2012. Danish dinosaurs: A review. In Godefroit (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 434-451.

undescribed Dromaeosauridae (Codrea, Godefroit and Smith, 2012)
Maastrichtian, Late Cretaceous
Rusca Montana Basin, Romania
Material- (UBB NgTh2) tooth
(UBB coll.) two teeth
Reference- Codrea, Godefroit and Smith, 2012. First discovery of Maastrichtian (Latest Cretaceous) terrestrial vertebrates in Rusca Montana Basin (Romania). In Godefroit (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 570-581.

unnamed Dromaeosauridae (Kordikova, Polly, Alifanov, Rocek, Gunnell and Averianov, 2001)
Turonian-Coniacian, Late Cretaceous
Zhirkindek Formation, Kazakhstan
Reference- Kordikova, Polly, Alifanov, Rocek, Gunnell and Averianov, 2001. Small vertebrates from the Late Cretaceous and early Tertiary of the northeastern
Aral Sea Region, Kazakhstan. Journal of Paleontology. 75, 390-400.

unnamed dromaeosaurid (Averianov, 2007)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- (ZIN PH 34/49) posterior tooth (FABL 2.1 mm) (Averianov, 2007)
Comments- This tooth has a BW/FABL of .38, with a centered mesial carina with very small serrations present at least apically. Distal serrations are U-shaped and occur at a density of 7/mm. It was referred to Velociraptorinae by Averianov (2007). The small serration sze may indicate this is microraptorian.
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.

undescribed possible dromaeosaurid (Nessov, 1995)
Santonian, Late Cretaceous
Syuk-Syuk Formation, Kazakhstan
Material- unguals?
Comments- Nessov (1995) notes Prinada (1925, 1927) and/or Riabinin (1938, 1939) identified unguals as Velociraptor. This remains uncertain pending further studies.
References- Prinada, 1925. [Search for remains of large vertebrates of Upper Cretaceous age in Turkestan. Report on the state of activities of the Geological Committee for 1924. Part II, III]. Izvyestiya Gyeologichyeskogo komityeta. 44(2), 257.
Prinada, 1927. [Report on the excavation at the localities where dinosaur bones were discovered. Report on the state of activities of the Geological Committee for 1925. Part II, III]. Izvyestiya Gyeologichyeskogo komityeta. 45(4), 453-454.
Riabinin, 1938. [Some results of the study of the Upper Cretaceous dinosaur fauna from the vicinity of st. Sary-Agachin, Southern Kazakhstan]. Problyemy palyeontologii. 4, 125-135.
Riabinin, 1939. [Vertebrate fauna from the Upper Cretaceous of southern Kazakhstan. I. Reptilia. Part 1. 1. Ornithischia]. Trudy Tsyentral'nogo Nauchno-Isslyedovatyel'skogo Gyeologorazvyedochnogo instituta. 18, 1-40.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology and paleobiogeography. Scientific Research Institute of the Earth's Crust. 156 pp.

undescribed dromaeosaur (Novikov, Lebidev and Alifanov, 1998)
Bathonian, Middle Jurassic
Meshcherskii Gorizont of the Moskvoretskaya Svita, Russia
Material- tooth
Reference- Novikov, Lebedev and Alifanov, 1998. New Mesozoic vertebrate fossil sites of Russia. Third European Workshop on Vertebrate Paleontology, Maastricht, 6-9 May 1998, p. 58.

undescribed Dromaeosauridae (Averianov, Starkov and Skutschas, 2003)
Aptian-Albian, Early Cretaceous
Ilek (=Shestakovo) Formation, Russia
Material- teeth (Averianov et al., 2003)
anterior dorsal vertebra (Averianov et al., 2004)
References- Averianov, Starkov and Skutschas, 2003. Dinosaurs from the Early Cretaceous Murtoi Formation in Buryatia, Eastern Russia. Journal of Vertebrate Paleontology. 23(3):586–594.
Averianov, Leshchinskiy, Skutschas, Fayngertz and Rezvyi, 2004. Dinosaurs from the Early Cretaceous Ilek Formations in West Siberia, Russia. 2nd EAVP Meeting. July 19-24, 2004. Brno, Czech Republic. Abstracts of papers and posters with program, Excursion Guidebook (O. Dostal, R. Gregorova & M. Ivanov, Eds.), 6.

undescribed Dromaeosauridae (Watabe and Suzuki, 2000)
Early Cretaceous
Shine Usny Tolgod, Mongolia
Material- (uncollected?) unguals
Comments- Watabe and Suzuki (2000) report "dromaeosaur claws" collected on June 11 1994.
Reference- Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 30-44.

undescribed dromaeosaurid (Matsukawa and Obata, 1994)
Aptian-Albian, Early Cretaceous
Zouyun Formation, Mongolia
Comments- Matsukawa and Obata (1994) report through personal communication with Mateer (1992) that cf. Velociraptor mongoliensis was discovered in the Zouyun Formation, though this is much too early to actually be that genus.
Reference- Matsukawa and Obata, 1994. Cretaceous, a contribution to dinosaur facies in Asia based on molluscan paleontology and stratigraphy. Cretaceous Research. 15, 101-125.

undescribed Dromaeosauridae (Watabe and Suzuki, 2000)
Cenomanian-Turonian, Late Cretaceous
Burkhant, Bayanshiree Formation, Mongolia
Material- (uncollected?) ungual (Watabe and Suzuki, 2000)
many postcranial elements (Watabe, Tsogtbaatar, Suzuki and Saneyoshi, 2010)
Cenomanian-Turonian, Late Cretaceous
Urlibe Khudak, Bayanshiree Formation, Mongolia
(IGM coll.) material (Kobayashi, Tsogtbaatar, Tsogtbaatar and Barsbold, 2015)
Comments- Watabe and Burkhant (2000a) reported a "dromaeosaur claw" found at Burkhant on August 13 1993. Watabe et al. (2010) stated "many postcranial bones of large-sized dromaeosaurid were also found."
Kobayashi et al. (2015) reported dromaeosaurid remains from Urlibe Khudak found in 2012.
Achillobator is known from Burkhant.
References- Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1993. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 19-29.
Watabe, Tsogtbaatar, Suzuki and Saneyoshi, 2010. Geology of dinosaur-fossil-bearing localities (Jurassic and Cretaceous: Mesozoic) in the Gobi Desert: Results of the HMNS-MPC Joint Paleontological Expedition. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 41-118.
Kobayashi, Tsogtbaatar, Tsogtbaatar and Barsbold, 2015. A new therizinosaur with functionally didactyl hands from the Bayanshiree Formation (Cenomanian-Turonian), Omnogovi Province, southeastern Mongolia. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 157.

undescribed dromaeosaurid (Watabe and Suzuki, 2000)
Late Campanian, Late Cretaceous
Abdrant Nuru, Djadokhta Formation?, Mongolia
Material- (uncollected?) ungual (Watabe and Suzuki, 2000)
Comments- Watabe and Suzuki (2000) reported "a large claw of dromaeosaurid" from the upper area (Toosgod) of Abdrant Nuru found on September 4-5 1996.
Reference- Watabe and Suzuki, 2000c. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1996. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 58-68.

undescribed dromaeosaurid (Watabe and Suzuki, 2000)
Late Campanian-Early Maastrichtian, Late Cretaceous
Shiluut Ula, ?Baruungoyot Formation, Mongolia
Material- (IGM coll.; 970803 SUPJ-5-9TSGT) partial skeleton, many elements including teeth
Comments- Watabe and Suzuki (2000) reported a "dromaeosaurid partial skeleton" discovered on August 1-4 1997, saying "the bed yielded the dromaeosaurid skeleton is a bone bed, including many other isolated bones." The field number is listed as "Doromaeosaurian bone bed", and they later state "from the bone bed, isolated bones and teeth of dromaeosaurid theropod were collected."
Reference- Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1997. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 69-82.

undescribed Dromaeosauridae (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
Material- (uncollected?) cervical vertebra (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Khaichin I, Nemegt Formation, Mongolia
three teeth (Badamkhatan, 2008)
Early Maastrichtian, Late Cretaceous
White Beds of Khermeen Tsav, Nemegt Formation, Mongolia
(IGM coll.) dorsal vertebra (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
?(IGM coll.) dorsal vertebra (Currie, 2001)
elements (Currie et al., 2008)
Comments- Currie (2001) lists two specimens recovered in September 10-24 2000, a "Coelurosaur (dromaeosaur?) dorsal vertebra" from the Central Sayr of the Nemegt locality, and a "Coelurosaur (dromaeosaur) dorsal vertebra" from Khermeen Tsav in the Nemegt Formation. The teeth reported by Badamkhatan (2008) are recurved, with distal serrations and an 8 shape basally. Currie et al.'s (2008) material is from an Avimimus bonebed. Watabe et al. (2010) reported "a cervical vertebra of dromaeosaurid (theropod)" from Bugin Tsav. All may belong to the contemporaneous Adasaurus.
References- Currie, 2001. Nomadic Expeditions, Inc. report on fieldwork in Mongolia, September 2000. Alberta Palaeontological Society, Fifth Annual Symposium. 12-16.
Badamkhatan, 2008. Dinosaurs from the Late Cretaceous Mongolian locality of Khaichin I. Journal of Vertebrate Paleontology. 28(3), 47A.
Currie, Longrich, Ryan, Eberth and Demchig, 2008. A bonebed of Avimimus sp. (Dinosauria: Theropoda) from the Late Cretaceous Nemegt Formation, Gobi Desert: Insights into social behavior and development in a maniraptoran theropod. Journal of Vertebrate Paleontology. 28(3), 67A.
Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010. Report of the HMNS-MPC Joint Paleontological Expedition in 2006. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 11-18.

undescribed dromaeosaurid (Barsbold, 1983)
Late Cretaceous?
Mongolia
Material- (IGM 100/12) pedal phalanx II-1 (25 mm), pedal phalanx II-2 (28 mm), proximal pedal ungual II
Comments- This was illustrated in figure 27 of Barsbold (1983) as an "undescribed Mongolian dromaeosaurid". The specimen number also belongs to a Gallimimus paratype.
Reference- Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.

unnamed possible Dromaeosauridae (Dong, 1997)
Barremian-Albian, Early Cretaceous
Xinminbao Group, Gansu, China
Material- (IVPP V11122) twelve teeth, caudal vertebra, two phalanges
Comments- Dong referred this material to Dromaeosauridae, though he refers two teeth (IVPP V11122-2 and V11122-3) from the same specimen to Troodontidae and Allosauridae. The illustrated supposedly dromaeosaurid teeth are moderately tall, serrated mesially and distally, and with serrations "almost as wide as long".
Reference- Dong, 1997. On small theropods from Mazongshan Area, Gansu Province, China. Pp. 13-18. in Dong (ed). Sino-Japanese Silk Road Dinosaur Expedition. China Ocean Press, Beijing. 114 p.

undescribed Dromaeosauridae (Dong, Currie and Russell, 1989)
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Material- (IVPP coll.; lost?) (juvenile) several partial skeletons (Jerzykiewicz et al., 1993)
(IVPP coll.; lost?) teeth (Dong, Currie and Russell, 1989)
Comments- Dong et al. (1989) reported "isolated bones and teeth of Velociraptor" found in June to July 1988. Jerzykiewicz et al. (1993) clarified that "Velociraptor mongoliensis, is represented by several partial skeletons of juveniles, and by shed teeth." Godefroit et al. (2008) stated "this material is not prepared yet and "these undescribed specimens could not be found in the IVPP collections." Linheraptor, "Velociraptor" osmolskae and mesially serrated teeth with a high DSDI that may be Velociraptor are known from this formation, and it's also possible the supposed juveniles are another variety of paravian as happened with Philovenator and the Almas perinates.
References- Dong, Currie and Russell, 1989. The 1988 field program of the Dinosaur Project. Vertebrata Palasiatica. 27(3), 233-236.
Jerzykiewicz, Currie, Eberth, Johnston, Koster and Zheng, 1993. Djadokhta Formation correlative strata in Chinese Inner Mongolia: An overview of the stratigraphy, sedimentary geology, and paleontology and comparisons with the type locality in the pre-Altai Gobi. Canadian Journal of Earth Sciences. 30(10), 2180-2195.
Godefroit, Currie, Li, Shang and Dong, 2008. A new species of Velociraptor (Dinosauria: Dromaeosauridae) from the Upper Cretaceous of northern China. Journal of Vertebrate Paleontology. 28(2), 432-438.

unnamed Dromaeosauridae (Gilmore, 1933)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (AMNH 6572) pedal phalanx II-1 (Ostrom, 1969)
(AMNH 21781) pedal ungual II (AMNH online)
?(IVPP 270790-4) tooth (~21x~9x? mm) (Currie and Zhao, 1993)
?(IVPP V16334.b) proximal femur (Funston, Currie, Ryan and Dong, 2019)
(IVPP coll.) teeth, elements (Dong, Currie and Russell, 1989)
(IVPP coll.) teeth and/or elements (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)
Comments- Gilmore (1933) noted the presence of "a few foot bones and other fragmentary skeletal parts" of small theropods from the Iren Dabasu Formation that he assigned to "Dromaeosaurinae Genus and species indet.", but admitted that this assignment as opposed to Coeluridae or Compsognathidae is based purely on their Cretaceous age. Thus while it turned out at least two of these elements are apparently dromaeosaurid (see below), Gilmore's rationale is equivalent to Maniraptora indet. today and the fossils referenced plausibly included material now recognized as avimimid, troodontid and ?alvarezsaurid. Ostrom (1969) noted in a discussion of deinonychosaur pedal examples that "E. H. Colbert has also discovered an isolated phalanx (AMNH 6572) in the American Museum collections from the Iren Dabasu Formation of Mongolia which compares almost exactly with the proximal phalanx of digit II of Deinonychus, but is perhaps 20 percent larger", which would make it somewhere around 46-52 mm long. He shows it questionably derived from Velociraptor in his phylogram without explanation, which would not make sense in the most recent interpretation of Iren Dabasu's age being contemporaneous or slightly older than the Djadochta. Paul (1988) states "at the AMNH is a hyper-extendable toe bone from the Late Cretaceous of Mongolia that looks like a Velociraptor somewhat bigger than V. antirrhopus [= Deinonychus]", but it is uncertain whether he saw it independant of Ostrom's text (Paul, pers. comm. 6-2022). The AMNH online catalog lists AMNH 21781 as an Iren Dabasu member of Dromaeosauridae represented by "Ungual of pes (digit II)" and being found by Kaisen. Both AMNH 6572 and 21781 would have been found in Erenhot during the April 22 to May 25 1923 Central Asiatic Expedition.
Dong et al. (1989) state Velociraptor material was discovered in the July 1988 Sino-Canadian expedition to Erenhot, and Dong (1992) specifies "teeth of Velociraptor". Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in the Iren Dabasu" and that "Most of these can be attributed to Velociraptor, although some of the teeth suggest that there was a second, larger species of an indeterminate dromaeosaurine dromaeosaurid." Yet no rationale was presented, and the only two specified Iren Dabasu dromaeosaurid elements in the literature are clearly not Velociraptor (AMNH 6572 is twice the size, while IVPP 270790-4 is different from most dromaeosaurid teeth as noted above). AMNH 6572 may belong to the supposed dromaeosaurine though, based on size. Currie and Zhao (1993) figure "Dromaeosaurid tooth (IVPP 270790-4) from the Iren Dabasu Formation near Erenhot, People's Republic of China, showing replacement pit on medial side of root", but given the slight constriction basal to the crown, convex distal edge and seeming lack of serrations, this may be misidentified. Based on its similar field number to troodontid metatarsal IVPP 230790-16 it was probably also found in the Sino-Canadian expedition of 1990.
Funston et al. (2019) notes a femoral head that supposedly differs from Avimimus in having fused anterior and greater trochanters, "which suggests that it may be oviraptorid or, more likely, dromaeosaur." This is from the Avimimus bonebed (locality K of Currie and Eberth [1993], which was CCDP locality 1 and may correspond to AMNH locality 141) and recovered in July 1988. Interestingly, femur PIN 2549-100 is from the same locality and shares a trochanteric crest, but is here identified as oviraptorid. It's possible they are from the same taxon (or even individual).
Yao et al. (2015) note "small unarticulated bones and teeth, including fossils of ... dromaeosaurids" from "a rare microvertebrate locality within the Iren Dabasu Formation, about 16 km northeast of Erenhot City."
References- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu Formation. Bulletin American Museum of Natural History. 67, 23-78.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin. 30, 1-165.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic of China. Cretaceous Research. 14, 127-144.
Currie and Zhao, 1993 (published 1994). A new troodontid (Dinosauria, Theropoda) braincase from the Dinosaur Park Formation (Campanian) of Alberta. Canadian Journal of Earth Sciences. 30(10-11), 2234-2247.
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. Caenagnathasia sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous: Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.
Funston, Currie, Ryan and Dong, 2019. Birdlike growth and mixed-age flocks in avimimids (Theropoda, Oviraptorosauria). Scientific Reports. 9:18816.

undescribed possible Dromaeosauridae (Prasad, Parmar and Kumar, 2014)
Early Jurassic
Kota Formation, India
Material- teeth
Comments- These were described as "dromaeosaurid-like, and Velociraptorinae-like."
Reference- Prasad, Parmar and Kumar, 2014. Recent vertebrate fossil discoveries from the Jurassic Kota Formation of India. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 208.

unnamed possible Dromaeosauridae (Knoll and Ruiz-Omenaca, 2005)
Beriassian, Early Cretaceous
KM 1983, Ksar Metlili Formation, Morocco
Material- (MNHN SA 2004/3A; Velociraptorinae indet. Morphotype I; lost) lateral tooth (3.20x2.20x.80 mm) (Knoll and Ruiz-Omenaca, 2005)
(MNHN SA mcm 158; Velociraptorinae indet. Morphotype III; lost) tooth (4.32x?x.76 mm) (Knoll and Ruiz-Omenaca, 2005)
(MNHN SA mcm 168; Velociraptorinae indet. Morphotype I; lost) tooth (1.28x1.20x.56 mm) (Knoll and Ruiz-Omenaca, 2005)
Beriassian, Early Cretaceous
KM-A1, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-A1-10) tooth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, Métais, Rage, Vullo and Zouhri, 2020)
Beriassian, Early Cretaceous
KM-A2, Ksar Metlili, Ksar Metlili Formation, Morocco
?(FSAC-KM-A2-9 [1]; Theropoda gen. et sp. indet. morphotype II) lateral tooth (3.34x1.82x1.22 mm) (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-B', Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-B'-26) tooth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, Métais, Rage, Vullo and Zouhri, 2020)
Beriassian, Early Cretaceous
KM-D2, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-D2-11; Theropoda gen. et sp. indet. morphotype II) lateral tooth (9.93x5.49x3.23 mm), tooth (Lasseron, 2020; Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, Métais, Rage, Vullo and Zouhri, 2020)
Comments- The MNHN specimens were collected in 1983, 1986 and 1999, while the FSAC specimens were collected in 2010, 2015 and 2018 (Lasseron, 2020). Lasseron noted the "dinosaur remains (Knoll, 2000; Knoll & Ruiz-Omeñaca, 2009) ... were taken out of the MNHN and lost." The MNHN teeth were referred to Dromaeosauridae by Knoll and Ruiz-Omenaca (2009) based on their high DSDI (1.08-1.45). However, this is also found in many more basal coelurosaurs. While Knoll and Ruiz-Omnaca (2005) originally noted teeth that "could be related to "paronychodontids"", the eventual 2009 publication shows this only refers to GNHN SA mcm 158, which is their Velociraptorinae indet. Morphotype III and has serrated carinae unlike Paronychodon. Knoll and Ruiz-Omenaca considered Morphotype I similar to Deinonychus while Morphotype III is close to specimens now recognized as Saurornitholestes premaxillary teeth (Zapsalis, Sankey et al.'s ?Dromaeosaurus morphotype A). Lasseron et al. (2020) stated "it is possible that those potential dromaeosaurid teeth from the KM fauna belong instead to Noasauridae or a basal tyrannosauroid (C. Hendrickx, pers. comm. to ML, March 2018)." Lasseron's Theropoda gen. et sp. indet. morphotype II is placed here as well due to the high DSDI (1.67-1.75). Lasseron finds quantitative analysis classifies FSAC-KM-A2-9 [1] as the contemporaneous Nuthetes, perhaps indicating a basal tyrannosauroid position if that genus is correctly placed here. Note Lasseron listed FSAC-KM-D2-11 as both a 'Theropoda gen. et sp. indet. morphotype I' and 'Theropoda gen. et sp. indet. morphotype II' tooth, without using a number in brackets to designate multiple teeth falling under one specimen number. As the mesial serration density is listed, the measurements given belong to a morphotype II tooth. Lasseron et al. lists all four teeth under FSAC-KM-B'-26 as Dromaeosauridae, although Lasseron described three as Theropoda gen. et sp. indet. morphotype III, Coelurosauria gen. et sp. indet. morphotype I and Coelurosauria gen. et sp. indet. morphotype III (unnamed Averostra and Coelurosauria here). Similarly, they list FSAC-KM-D1-6 as dromaeosaurid, but it was described as Coelurosauria gen. et sp. indet. morphotype I by Lasseron and is catalogd under unnamed Coelurosauria here. Lasseron et al. list both FSAC-KM-A2-9 teeth as Maniraptora, but they were described as Theropoda gen. et sp. indet. morphotypes I and II by Lasseron.
References- Knoll and Ruiz-Omenaca, 2005. Theropod teeth from the Berriasian of Anoual (Morocco). Journal of Vertebrate Paleontology. 25(3), 78A.
Knoll and Ruiz-Omenaca, 2009. Theropod teeth from the basalmost Cretaceous of Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine. 146(4), 602-616.
Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des vertebres mesozoiques africans et gondwaniens : apport des gisements du Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle. 493 pp.
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, Métais, Rage, Vullo and Zouhri, 2020 (online 2019). New data on the microvertebrate fauna from the Upper Jurassic or lowest Cretaceous of Ksar Metlili (Anoual Syncline, eastern Morocco). Geological Magazine. 157, 367‑392.

undescribed Dromaeosauridae (Sadleir, 1998)
Cenomanian, Late Cretaceous
Kem Kem Beds, Morocco
Material- (GZG.V.19997) tooth (13.8x7.5x3.8 mm) (Richter, Mudroch and Buckley, 2013)
?(GZG.V.19998) tooth (12.5x7.3x3.1 mm) (Richter, Mudroch and Buckley, 2013)
(M-CH-009) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
(M-JQ-012) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
(M-KS-015) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
(M-ZA-014) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
?(NMB-1671-R) tooth (14.5x7.5x3.8 mm) (Richter, Mudroch and Buckley, 2013)
Comments- The teeth reported by Amiot et al. (2004) are 10-16 mm in height, and strongly recurved with BW/FABLs of 0.4-0.6. Mesial serrations are present on all but M-CH-009 and are small (3.4-4/mm). Distal serrations are larger (2.4-3.8/mm) and apically inclined. DSDI ranged between 1.11-1.42. Interdenticle slits are deep. They referred these specimens to Velociraptorinae.
References- Sadleir, 1998. Theropod teeth from the Cretaceous of Morocco. Journal of Vertebrate Paleontology. 18(3), 74A.
Amiot, Buffetaut, Tong, Boudad and Kabiri, 2002. Laurasian theropod dinosaur teeth from the Late Cretaceous of Morocco. Conference abstract, Third Georges Cuvier Symposium, Montbeliard, France.
Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004. Isolated theropod teeth from the Cenomanian of Morocco and their palaeobiogeographical significance. Revue de Paleobiologie, Geneve. 9, 143-149.
Richter, Mudroch and Buckley, 2013. Isolated theropod teeth from the Kem Kem Beds (Early Cenomanian) near Taouz, Morocco. Palaontologische Zeitschrift. 87, 291-309.

unnamed dromaeosaurid (Werner, 1994)
Cenomanian, Late Cretaceous
Wadi Milk Formation, Sudan
Material- (Vb-713) pedal phalanx II-2 (36 mm) (Werner, 1994)
(Vb-714) proximal manual ungual (Werner, 1994)
(Vb-860) distal manual ungual (Rauhut and Werner, 1995)
(Vb-866) distal pedal ungual I (Rauhut and Werner, 1995)
(Vb-867) ungual (Rauhut and Werner, 1995)
(Vb-868) proximal pedal ungual III (Rauhut and Werner, 1995)
(Vb-875) tooth (8.3 mm) (Rauhut and Werner, 1995)
Comments- These unassociated remains were found in 1991 and 1992, and first reported by Werner (1994) before being described in depth by Rauhut and Werner (1995).
Vb-714 was originally identified as a pedal ungual II, but is more likely a manual ungual based on the high dorsal arch and large flexor tubercle. The similar partial ungual Vb-860 (also assigned to pedal digit II by Rauhut and Werner) may also be from the manus then. Vb-867 was also identified as a pedal ungual II, but was not illustrated. The high DSDI (1.33) was used to place this in the Velociraptorinae, but are also known in microraptorians and basal dromaeosaurines. It does exclude the Sudanese taxon from the Dromaeosaurus+Achillobator+Utahraptor subclade, as does the elongate pedal phalanx II-2.
References- Werner, 1994. Die kontinentale Wirbeltierfauna aus der unteren Oberkreide des Sudan (Wadi Milk Formation). Berliner geowiss. Abh.. 13, 221-249.
Rauhut and Werner, 1995. First record of the family Dromaeosauridae (Dinosauria: Theropoda) in the Cretaceous of Gondwana (Wadi Milk Formation, northern Sudan). Palaeontologische Zeitschrift. 69(3/4), 475-489.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.

unnamed possible dromaeosaurid (Fanti and Therrien, 2007)
Middle Maastrichtian, Late Cretaceous
Anembalemba Member of Maevarano Formation, Madagascar
Material- (MSNM V5365) premaxillary tooth
(MSNM V5372) premaxillary tooth
(MSNM V5373) lateral tooth
(MSNM V5394) premaxillary tooth
(MSNM V5590) premaxillary tooth
(MSNM V coll.) 13 teeth (8-15 mm)
Comments- These teeth are high compressed (BW/FABL ~.5), have 3-5.5 mesial serrations and 3-3.5 distal serrations. Mesial serrations are apically inclined, while distal serrations are perpendicular to the distal edge. All serrations are slightly hooked apically and have absent or poorly defined interdenticle slits.
These were referred to Dromaeosauridae based on a morphometric comparision of several measurements to dromaeosaurids, abelisaurids, Masiakasaurus and tyrannosaurids. Thus it is possible they belong to another theropod clade once they are compared to a broader range of taxa.
Reference- Fanti and Therrien, 2007. Theropod tooth assemblages from the Late Cretaceous Maevarano Formation and the possible presence of dromaeosaurids in Madagascar. Acta Palaeontologica Polonica. 52(1), 155-166.

undescribed possible dromaeosaurid (Canale, Carballido, Otero, Canudo and Garrido, 2014)
Albian-Cenomanian, Early Cretaceous-Late Cretaceous
Bayo Overo Member of the Cerro Barcino Formation, Chubut, Argentina
Material- tooth
Reference- Canale, Carballido, Otero, Canudo and Garrido, 2014. Carcharodontosaurid teeth associated with titanosaur carcasses from the Early Cretaceous (Albian) of the Chubut Group, Chubut Province, Patagonia, Argentina. Jornadas Argentinas de Paleontologia de Vertebrados. Ameghiniana. 51(6) suplemento, 6.

undescribed Dromaeosauridae (Casal, Martinez, Candeiro, Lamanna and Ibiricu, 2007)
Mid Cenomanian-Turonian, Late Cretaceous
Lower Bajo Barreal Formation, Chubut, Argentina
Material- three teeth
Reference- Casal, Martinez, Candeiro, Lamanna and Ibiricu, 2007. First record of Dromaeosauridae (Dinosauria: Theropoda) in the Early Late Cretaceous Bajo Barreal Formation of Chubut Province, Argentina. Journal of Vertebrate Paleontology. 27(3), 56A.

unnamed possible Dromaeosauridae (Elias, Bertini and Medeiro, 2007)
Late Albian-Early Cenomanian, Early-Late Cretaceous
Alcantara Formation, Brazil
Material- (UFMA 1.20.194-1) tooth (16.1x7.3x3.4 mm)
(UFMA 1.20.194-2) incomplete tooth (13.1x7.9x3.8 mm)
Comments- These were referred to Velociraptorinae by Elias et al. (2007), but lack e.g. a high DSDI or 8-shaped basal section.
Reference- Elias, Bertini and Medeiro, 2007. Velociraptorinae (Maniraptoriformes) teeth from the Coringa Flagstone outcrop, Middle Cretaceous of the Sao Luis-Grajau basin, Maranhao state, northern-northeastern Brazil. in Carvalho, Cassab, Schwanke, Carvalho, Fernandes, Rodrigues, Carvalho, Arai and Oliveira (eds.). Paleontologia: Cenários de Vida. 1, 315-325.

unnamed possible Dromaeosauridae (Franco-Rosas, 2002)
Turonian-Late Maastrichtian, Late Cretaceous
Adamantina, Marília and/or Serra da Galga Formations of the Bauru Group, Brazil
Material- teeth
Comments- These teeth have long, pointed distal serrations with deep interdenticle slits. They are said to be similar to dromaeosaurids, and the description matches basal forms more than derived dromaeosaurines.
Reference- Franco-Rosas, 2002. Methodological parameters for the identification and taxonomic classification of isolated theropodomorph teeth. Anais da Academia Brasileira de Ciencias. 74(2), 367.

Dromaeosauridae sensu Norell and Makovicky, 2004
Definition- (Microraptor zhaoianus + Sinornithosaurus millenii + Velociraptor mongoliensis)

Microraptoria Senter et al., 2004
Definition- (Microraptor zhaoianus <- Velociraptor mongoliensis, Dromaeosaurus albertensis) (modified from Senter et al., 2004)
= Microraptorinae sensu Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Microraptor zhaoianus <- Dromaeosaurus albertensis)
= "Microraptorinae" Makovicky, Apesteguía and Agnolín, 2005
= Microraptorinae Longrich and Currie, 2009
Definition- (Microraptor zhaoianus <- Velociraptor mongoliensis, Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus) (Turner et al., 2012)
Other definition- (Microraptor zhaoianus <- Dromaeosaurus albertensis) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017)
Comments- Senter et al. (2004) erected Microraptoria, with a non-family level suffix to ensure ICZN rules are followed whether or not the clade was within Dromaeosauridae (depending on both the tree's topology and the latter's definition). They listed Microraptoridae and Microraptorinae as alternatives considered prior to deciding upon Microraptoria, and Microraptorini as a mispelling of Microraptoria. Makovicky et al. (2005) cited Senter et al. for Microraptorinae, but the latter was a misspelling of their own (Headden, pers. comm. to Makovicky, 2005). Turner et al. (2007) followed Makovicky et al. in using Microraptorinae, but neither reference contains a diagnosis or definition so the name is a nomen nudum (ICZN Article 13.1.1). While Makovicky et al. did reference Senter et al.'s paper, which included a definition for Microraptoria, the latter is not an available name under the ICZN so Article 13.1.3 does not apply. In addition, ICZN Article 11.5 states names must be proposed as being valid, so Senter et al. 2004 cannot be cited as the authors for Microraptorinae/idae, contra most authors discussed here. This leaves Longrich and Currie (2009) as the first publication to validy erect Microraptorinae. Turner et al. (2012) gave Microraptorinae a more restrictive definition that supposedly overcomes Senter et al.'s perceived difficulties with family-level names. However, in topologies such as Mayr et al.'s (2005), Microraptorinae does not fall under any named family, which was just the problem Senter et al. anticipated when they rejected the name. On the other hand, Microraptoria includes Confuciusornis and potentially all pygostylians in Mayr et al.'s topology, unlike Microraptorinae. So Turner et al.'s definition is more stable due to its extra specifiers. At present, however, the consensus is that Microraptorinae and Microraptoria are synonymous, and the latter has priority.
References- Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8, 1-20.
Makovicky, Apesteguía and Agnolín, 2005. The earliest dromaeosaurid theropod from South America. Nature. 437, 1007-1011.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399.

Tianyuraptor Zheng, Xu, You, Zhao and Dong, 2010
= "Tianyuraptor" Zheng, Xu, You, Zhao and Dong, 2009 online
= "Zhenyuanlong" Lu and Brusatte, 2015
T. ostromi Zheng, Xu, You, Zhao and Dong, 2010
= "Tianyuraptor ostromi" Zheng, Xu, You, Zhao and Dong, 2009 online
= "Zhenyuanlong suni" Lu and Brusatte, 2015
Early Aptian, Early Cretaceous
Dawangzhangzi Beds of Yixian Formation, Liaoning, China
Holotype- (STM1-3) (subadult) (12.5 kg) skull, mandible, six cervical vertebrae, thirteen dorsal vertebrae, dorsal ribs, uncinate processes, gastralia, sacrum, twenty-five caudal vertebrae (caudal series 960 mm), chevrons, scapulae (121 mm), coracoid, partial furcula, sternal plate, humeri (138 mm), radii (104 mm), ulnae, semilunate carpals fused to metacarpals II, metacarpals I, phalanges I-1, manual unguals I, phalanges II-1, phalanges II-2, manual unguals II, metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3, manual unguals III, ilia (~159 mm), pubes, ischia, femora (212.5 mm), tibiae (283 mm), fibula, astragalus, calcaneum, metatarsal I(?), phalanx I-1(?), metatarsals II, metatarsals III, phalanx III-2, phalanx III-3, pedal ungual III, metatarsals IV, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, pedal phalanges, pedal ungual, metatarsal V
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Referred- (JPM-0008; intended holotype of "Zhenyuanlong suni") (~1.65 m, 11.8 kg, subadult) incomplete skull (166.1 mm), mandible (165.2 mm), hyoid, ten cervical vertebrae (30.3 mm), dorsal series (250 mm; best preserved 23.7 mm), dorsal ribs, gastralia, six sacral vertebrae, first caudal vertebra (23.6 mm), second caudal vertebra (23.6 mm), third caudal vertebra (23.6 mm), fourth caudal vertebra (29.7 mm), fifth caudal vertebra (32.4 mm), sixth caudal vertebra (37.3 mm), seventh caudal vertebra (43.4 mm), eighth caudal vertebra (40.7 mm), ninth caudal vertebra (40.7 mm), tenth caudal vertebra (42.1 mm), eleventh caudal vertebra (46.8 mm), twelfth caudal vertebra (46.8 mm), thirteenth caudal vertebra, fourteenth caudal vertebra, fragmentary fifteenth caudal vertebra, chevrons, scapulae (one partial), incomplete coracoids, sternal plates, sternal ribs, humeri (121.1 mm), radii (96.6 mm), ulnae (102.9 mm), scapholunare, semilunate carpal, distal carpal III, metacarpals I (19.5 mm), phalanges I-1 (40.5 mm), manual unguals I (25.4 mm), metacarpals II (55.7 mm), phalanges II-1 (32.8 mm), phalanges II-2 (39.2 mm), manual unguals II (31.7 mm), metacarpals III (53 mm), phalanges III-1 (14.9 mm), phalanges III-2 (8.1 mm), phalanges III-3 (24.7 mm), manual unguals III (22.8 mm), manual claw sheaths, ilium (128.3 mm), pubes (155.7 mm), ischia (80.1 mm), femora (193.4 mm), tibiae (260.3 mm), fibulae (262.5 mm), astragalocalcanea, distal tarsal III, distal tarsal, metatarsals I (27.7 mm), phalanges I-1, pedal unguals I, metatarsals II (128.3 mm), phalanx II-2, pedal ungual II, metatarsals III (129.8 mm), phalanges III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsals IV (128.2 mm), phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, pedal phalanges, body feathers, remiges, retrices
Early Cretaceous
Jehol Group, China
(PMOL coll.) (2.5 m) complete skeleton (Makovicky, Gorscak and Zhou, 2018)
Diagnosis- (after Zheng et al., 2010) small and extremely slender furcula; elongated hindlimb about three times as long as dorsal series.
(after Lu and Brusatte, 2015) short forelimb (humerofemoral ratio <0.65, ulnofemoral ratio <0.55, manuofemoral ratio <0.90; actual values 0.63, 0.53 and 0.76; values of Tianyuraptor holotype 0.65, 0.49 and 0.86; with the humerofemoral ratio barely excepted) (also in Achillobator - ?, 0.51 and ?).
Other diagnoses- Turner et al. (2012) note Zheng et al.'s character of 'length of mid caudal vertebrae more than twice that of dorsal vertebrae' is present in Microraptor as well.
Lu and Brusatte (2015) listed several characters in their diagnosis. Adasaurus, Velociraptor and Deinonychus also have six sacral vertebrae. Graciliraptor, Microraptor, Velociraptor, Achillobator and some Deinonychus have a radius thinner than manual phalanx I-1. Metacarpal II is shorter than metacarpal I plus manual phalanx I-1 in eudromaeosaurs (Velociraptor, Tsaagan, Deinonychus), Bambiraptor, Tianyuraptor (contra Lu and Brusatte) and some Microraptor (LPM 0159). Lu and Brusatte stated the posteriorly curved pubis differs from the straight element in Tianyuraptor, but the latter is also posteriorly curved. While they said the ilium of Tianyuraptor differs from "Zhenyuanlong" in sharing a lobate brevis shelf with Microraptor, this seems to be absent in Tianyuraptor's type based on available low resolution photos. They also said "Zhenyuanlong" differs from Tianyuraptor in having a ridge along the ventral antorbital fossa edge, but the former has a narrow and convex surface ventral to the fossa, not a ridge compared to the rest of the external surface. Lu and Brusatte also included two characters shared with Tianyuraptor and eudromaeosaurs- mid-pubic tubercle absent (plesiomorphic), and mid dorsal ischial process absent (also in Unenlagia and Buitreraptor).
Comments- The description was released online in August 2009 but not officially published until January 2010.
The holotype of "Zhenyuanlong" was collected by a farmer prior to 2015. Lu and Brusatte (2015) believed the Sihedang locality, which it derives from, to be in the Yixian Formation. It is here assigned to the Jiufotang Formation instead (see Iteravis entry). The taxon was described by Lu and Brusatte on July 16 2015 as a new genus of dromaeosaurid. However, this paper has no mention of ZooBank and as of February 6 2020 "Zhenyuanlong" lacks an entry on the ZooBank website. Thus according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Zhenyuanlong suni" Lu and Brusatte 2015 is a nomen nudum that will only be technically valid pending action on behalf of the authors or ICZN as its journal is not published physically.
Valid differences between Tianyuraptor and "Zhenyuanlong" noted by Lu and Brusatte are limited to- radius thinner than manual phalanx I-1; smaller manus (76% of femoral length vs. 86%). These could easily be explainable as individual variation, as the former varies in Deinonychus. Makovicky et al. (2018) noted a new specimen which "exhibits a combination of traits that were previously used to distinguish Tianyuraptor from Zhenyuanlong."
Zheng et al. (2010) found Tianyuraptor to be more closely related to Dromaeosaurus than Unenlagia, but outside previously described microraptorians, and Saurornitholestes plus Velociraptor plus dromaeosaurines. It was recovered as a microraptorian by Senter et al. (2012), Turner et al. (2012) and Lee et al. (2014), but in a polytomy with microraptorians and more derived dromaeosaurids by Brusatte et al. (2014). Lu and Brusatte added "Zhenyuanlong" to Turner's TWiG analysis and found it to be in a polytomy with the microraptorian genera, Tianyuraptor and Eudromaeosauria.
References- Zheng, Xu, You, Zhao and Dong, 2010 (online 2009). A short-armed dromaeosaurid from the Jehol Group of China with implications for early dromaeosaurid evolution. Proceedings of the Royal Society B. 277, 211-217.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Lu and Brusatte, 2015. A large, short-armed, winged dromaeosaurid (Dinosauria: Theropoda) from the Early Cretaceous of China and its implications for feather evolution. Scientific Reports. 5, 11775.
Makovicky, Gorscak and Zhou, 2018. A new specimen of the large-bodied dromaeosaurid Tiuanyuraptor provides new insights on microraptorine anatomy, taxonomy, and plumage evolution. Journal of Vertebrate Paleontology. Program and Abstracts 2018, 174.

undescribed dromaeosaurid (Li, 2013)
Aptian-Albian, Early Cretaceous
Zhonggou Formation, Gansu, China
Material- (FRDC-GJ(07)11-2) (~2 m) frontal (63.5 mm), mid dorsal vertebra (30 mm), incomplete mid dorsal vertebra (28.5 mm), posterior dorsal vertebra (26.5 mm), posterior dorsal vertebra (23 mm), sacrum (32.5, 21.5, ~20, ~21.5, ~22.5, ~23 mm), ~fifth caudal vertebra (19 mm), distal caudal vertebra (31.5 mm), distal caudal vertebra (26.5 mm), incomplete scapula, humerus (138 mm), manual ungual II (30.5 mm), phalanx III-3 (41.5 mm), manual ungual III (44 mm), ventral fragment of ilium, proximal pubis, proximal ischium, phalanx II-1 (29.5 mm), phalanx II-2 (28 mm), distal metatarsal III(?), phalanx III-1 (53.5 mm), phalanx III-3 (33.5 mm), phalanx IV-4 (27.5 mm)
Comments- Discovered in 2007, this was described as a new taxon in Li's (2013) and Mariko's (2015) theses and recovered as a basal velociraptorine in 2013 based on Senter's TWiG matrix and a eudromaeosaur incertae sedis in 2015 based on Turner's TWiG matrix. When entered into the Hartman et al. maniraptoromorph matrix, FRDC-GJ(07)11-2 falls out as a microraptorian sister to "Zhenyuanlong".
References- Li, 2013. A new dromaeosaurid dinosaur from the Early Cretaceous of Yujingzi basin, Jiuquan area, Gansu Province, China. Masters Thesis, China University of Geosciences. 120 pp.
Mariko, 2015. A study of the new dromaeosaurid dinosaur from the Early Cretaceous of Jiuquan area, Gansu Province, China. Masters Thesis, China University of Geosciences. 112 pp.

"Paleopteryx" Jensen, 1981
"P. thomsoni" Jensen, 1981
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US (Dry Mesa quarry)
Material- (BYU 2022) distal radius
Comments- Jensen (1981) described the holotype (BYU 2022) as a proximal tibiotarsus of a new taxon of bird- "Paleopteryx thomsoni". In addition, he referred a proximal femur (BYU 2023) to Archaeopteryx, and another femur (BYU 2025) and sacrum (BYU 2024) as avian-like. Molnar (1985) showed that "Paleopteryx" was not diagnosed, so is a nomen nudum (ICZN Article 13.1.1). He assigned the BYU material to Theropoda. Jensen and Padian reidentified BYU 2022 as a distal radius, and referred both it and BYU 2023 to Maniraptora indet.. They made BYU 2024 the holotype of a new pterosaur genus Mesadactylus, and referred BYU 2025 to that taxon as well.
"Paleopteryx" actually has an extremely distinctive distal radius, with a medially flared articular surface seen in dromaeosaurids, Sapeornis and Confuciusornis, but absent in therizinosaurs, oviraptorosaurs, Yixianosaurus, Sinornithoides, Archaeopteryx, Rahonavis or Shenzhouraptor. The high angle of flaring relative to the shaft is shared with Graciliraptor and Microraptor among dromaeosaurids, unlike Deinonychus and Bambiraptor. It is more similar to Graciliraptor, in that both lack Microraptor's apomorphic concave surface just distal to the flare. An additional character shared with Microraptor but not other maniraptorans (unknown in Graciliraptor) is a distally projecting lateral process. "Paleopteryx" is thus here assigned to Microraptoria.
References- Jensen, 1981. [A new oldest bird?] Anima (Tokyo). 1981, 33-39. [in Japanese]
Jensen, 1981. Another look at Archaeopteryx as the worlds oldest bird. Encyclia, The Journal of the Utah Academy of Sciences, Arts, and Letters. 58, 109-128.
Molnar, 1985. Alternatives to Archaeopteryx; a survey of proposed early or ancestral birds. in Hecht, Ostrom, Viohl and Wellnhofer (eds). The Beginnings of Birds. Eichstatt, Germany: Freunde des Jura-Museums Eichstatt. 209-217.
Jensen and Padian, 1989. Small Pterosaurs and Dinosaurs from the Uncomphagre fauna (Brushy Basin Member, Morrison Formation: ?Tithonian), Late Jurassic, Western Colorado. Journal of Paleontology. 63(3), 364-373.
Padian, 1998. Pterosaurians and ?avians from the Morrison Formation (Upper Jurassic, Western U.S.). Modern Geology. 23, 57-68.

Shanag Turner, Hwang and Norell, 2007
S. ashile Turner, Hwang and Norell, 2007
= Sinornithosaurus ashile (Turner, Hwang and Norell, 2007) Paul, 2010
Berriasian-Barremian, Early Cretaceous
Oosh, Huhteeg Svita, Mongolia
Holotype- (IGM 100/1119) incomplete maxilla, incomplete dentary, partial splenial
Diagnosis- (after Turner et al., 2007) triangular, anteriorly tapering maxilla; lateral lamina of nasal process of maxilla reduced to small triangular exposure; absence of a promaxillary fenestra; presence of interalveolar pneumatic cavities; incipient dentary groove on posterolateral surface of dentary.
Comments- This specimen was discovered in 1999, first noted by Turner et al. (2006) then described in Turner et al. (2007). The latter analysis found it in an unresolved position among Microraptoria+Eudromaeosauria, though noted greatest similarity to Sinornithosaurus. It has since been recovered as a microraptorian (Longrich and Currie, 2009) or sister to Microraptoria+Eudromaeosauria (Senter et al., 2012).
References- Turner, Pol, Norell and Hwang, 2006. Resolving dromaeosaurid phylogeny: New information and additions to the tree. Journal of Vertebrate Paleontology. 26(3), 133A.
Turner, Hwang and Norell, 2007. A small derived theropod from Oosh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates. 3557, 27 pp.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.

Richardoestesia Currie, Rigby and Sloan, 1990
= Asiamericana Nessov, 1995
Diagnosis- very small serrations on teeth (5-12/mm); apical half of anterior dentary teeth convex distally.
Comments- This genus is known from a large amount of teeth, spanning the Bathonian to Maastrichtian of North and South America, Europe and Asia. The only known nondental remains are the holotype dentaries, though it's probable some of the many unidentified small theropod postcranial remains so far discovered belonged to individuals with Richardoestesia-type teeth. The wide temporal and geographic range suggests Richardoestesia as currently diagnosed is a clade comparable to "families" in scope, and quite possibly polyphyletic. The two named species, R. gilmorei and R. isosceles, show variation between different formations that probably indicate they each contain several species. R. cf. gilmorei is indistinguishable from Saurornitholestes in the Early Campanian Milk River Formation except for serration size, though the possible presence of gilmorei-like teeth from earlier formations may indicate this is not due to R. gilmorei evolving from a dromaeosaurid in the Early Campanian. However, these earlier records are based on unpublished observations from a single figure (Chipping Norton Formation) and a single specimen which seems to have serrations too large for Richardoestesia (Woodbine Formation). R. isosceles-like specimens have a more definitive pre-Campanian record and are the only ones verified from Eurasia, where they extend back to the Kimmeridgian of Portugal. Though the pre-Campanian American records should be examined before any conclusions are made, it seems possible R. isosceles originated much earlier than R. gilmorei and had little to do with that species. It should also be noted that many poorly described records of Richardoestesia may belong to other genera such as basal tyrannosauroids (e.g. Nuthetes, Dilong) or other basal dromaeosaurids (e.g. Sinornithosaurus, Shanag). Indeed, both Shanag and Sinornithosaurus have very small serrations on their teeth, and some teeth whose apical half is convex distally. Hendrickx and Mateus (2014) found Richardoestesia to be a dromaeosaurid in their phylogenetic analysis.
Richardoestesia or Ricardoestesia?- Olshevsky and others have noted that the spelling of this genus was originally supposed to be Ricardoestesia, and was misspelled in all cases except for a figure caption in the original description by an editor. Thus, Olshevsky believes Ricardoestesia should be the correct spelling. However, he acted as first revisor in 1991, listing Richardoestesia as the correct spelling and Ricardoestesia as a misspelling. As Creisler (DML, 2002) noted- "Under ICZN Art. 24.2.3, the first author to have cited two alternate spellings of a name together and to have selected one spelling as correct qualifies as the First Reviser. Under 32.5.1, this spelling could only be changed if it is determined to be "incorrect"--meaning there is evidence of an inadverent error in the original publication itself "without recourse to any external source of information." Crucially, "incorrect transliteration or latinization" is not considered an inadvertent error.
As I pointed out back in Feb. 2001, "Richardus" is a perfectly good latinization for Richard, and one widely used in Medieval and later Latin literature. Since the name was intended to honor Richard Estes, the latinized form Richardoestesia is perfectly good and not in any way readable as "inadvertent error." If one of the authors of the original paper wanted Ricardoestesia instead, his wish qualifies as an "external source of information" which can't be used, since it's not mentioned in the original paper. Moreover, the senior author on the original Richardoestesia paper was Phil Currie, who has used the spelling Richardoestesia in other papers he has authored or coauthored.
Frankly, I can't find any basis in the ICZN for switching the spellings in later editions of Mesozoic Meanderings. George had in fact already been the First Reviser on the name in Oct. 1991. Since generic names differing by one letter are not homonyms, the coexistence of Richardoestesia and Ricardoestesia for the same taxon is becoming more confusing as time goes on, especially with electronic data retrieval and online databases. As the situation is shaping up, nearly all technical papers are using Richardoestesia and a few books have used Ricardoestesia. Maybe the ICZN will have to make a decision, but it seems like a lot of fuss over an issue that was settled just fine by George in 1991."
Not Richardoestesia- Richardoestesia was reported from the Campanian Fort Crittenden Formation of Arizona by Ratkevich and Duffek (1996), but reidentified merely as Dromaeosauridae indet. by Sullivan and Lucas (2006).
References- Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.
Nessov, 1995. Dinozavri severnoi Yevrasii: Novye dannye o sostave kompleksov, ekologii i paleobiogeografii [Dinosaurs of Northern Eurasia: new data about assemblages, ecology and paleobiogeography], Scientific Research Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia: 156 pp. + 14 pl. [in Russian with short English, German, and French abstracts].
Ratkevich and Duffek, 1996. Small macro-and large micro-vertebrate fauna of the Fort Crittenden Formation, Southeast Arizona. Proceedings of Southwest Paleontological Society and Mesa Southwest Museum, Mesa, Arizona. 4, 115-120.
Creisler, DML 2002. https://web.archive.org/web/20200806190728/http://dml.cmnh.org/2002Jul/msg00530.html
Sullivan and Lucas, 2006. The Kirtlandian land-vertebrate "age" - faunal composition, temporal position and biostratigraphic correlation in the nonmarine Upper Cretaceous of western North America. New Mexico Museum of Natural History and Science Bulletin. 35, 7-29.
Hendrickx and Mateus, 2014. Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth. Zootaxa. 3759(1), 1-74.
R. gilmorei Currie, Rigby and Sloan, 1990
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, US
Holotype- (CMN 343) partial dentaries, teeth
Paratypes- (RTMP 83.45.2) sixth dentary tooth
(RTMP 80.8.298) tooth
(RTMP 80.16.1230) tooth
(RTMP 83.129.11) tooth
(RTMP 84.89.274) tooth
Referred- (RTMP 65.26.13) tooth (Baszio, 1997)
(RTMP 81.16.194) premaxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.16.173) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 83.36.233) tooth (12.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 83.36.242) tooth (8.3 mm) (Baszio, 1997)
(RTMP 84.92.268) tooth (~5.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.59.204) tooth (Baszio, 1997)
(RTMP 86.23.90) tooth (10 mm) (Baszio, 1997)
(RTMP 86.23.105) tooth (Baszio, 1997)
(RTMP 86.159.60) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.171.9) tooth (4.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.171.61) tooth (3.4 mm) (Baszio, 1997)
(RTMP 87.116.60) tooth (Baszio, 1997)
(RTMP 87.80.35) tooth (7.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.91.28) tooth (5.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.156) tooth (Baszio, 1997)
(RTMP 89.36.355) tooth (10 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.76.63) tooth (5.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.106.6) tooth (4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.157.29) tooth (2.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.30b) tooth (3.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(ANSP 15824) tooth (Fiorillo and Currie, 1994)
(ANSP 15942) tooth (Fiorillo and Currie, 1994)
(ANSP 15952) tooth (Fiorillo and Currie, 1994)
(ANSP 17647) tooth (Fiorillo and Currie, 1994)
(ANSP 17782) tooth (Fiorillo and Currie, 1994)
(ANSP 17784) tooth (Fiorillo and Currie, 1994)
(ANSP 17983) tooth (Fiorillo and Currie, 1994)
(ANSP 18101) tooth (Fiorillo and Currie, 1994)
(ANSP 18104) tooth (Fiorillo and Currie, 1994)
(ANSP 18114) tooth (Fiorillo and Currie, 1994)
(ANSP 18115) tooth (Fiorillo and Currie, 1994)
(ANSP 18119) tooth (Fiorillo and Currie, 1994)
Middle Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
(RTMP 83.33.17) tooth (7.6x4.4x2.2 mm) (Larson and Currie, 2013)
(RTMP 83.45.11) tooth (6.6x4.4x2.2 mm) (Larson and Currie, 2013)
(RTMP 90.82.15) tooth (11.7x6.2x2.6 mm) (Larson and Currie, 2013)
(RTMP 94.28.1) tooth (10.2x5.3x2.5 mm) (Larson and Currie, 2013)
(RTMP 96.29.2) tooth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
(RTMP 96.39.29a) tooth (3.9x3x1.2 mm) (Larson and Currie, 2013)
(RTMP 97.39.2) tooth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
(RTMP 99.50.113) tooth (Larson et al., 2010)
(RTMP 2000.45.80) tooth (11.9x6.l4x2.7 mm) (Larson et al., 2010)
(RTMP 2001.45.82) tooth (8.9x3.9x2.2 mm) (Larson et al., 2010)
(RTMP 2002.45.49) tooth (4.7x3x1.3 mm) (Larson et al., 2010)
(RTMP 2002.45.53) tooth (4.3x3.4x1.3 mm) (Larson et al., 2010)
(RTMP 2003.15.2) tooth (Larson et al., 2010)
(RTMP 1016) tooth (Baszio, 1997)
(RTMP 1017) tooth (Baszio, 1997)
(RTMP 1018) tooth (Baszio, 1997)
(RTMP 1026) tooth (Baszio, 1997)
(RTMP 1027) tooth (Baszio, 1997)
(RTMP coll.) nine teeth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
(LSUMG V-6237) partial tooth (Sankey, 2001)
Late Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
(RAM 24759) tooth (Farke, Letteau Stallings and Andrews, 2020)
Diagnosis- (from Sankey et al., 2002) differs from R. isosceles in- teeth generally shorter, more recurved and larger; serrations not square shaped, and with small interdenticle spaces; some teeth constricted at base.
differs from Shanag in- shallow Mackelian groove; some teeth constricted at base; all teeth have distal serrations; some teeth have mesial serrations.
differs from Sinornithosaurus in- shallow Mackelian groove; Mackelian groove placed more dorsally; unfused posterior dentary interdental plates; at least some premaxillary teeth serrated distally; some teeth constricted at base; anterior dentary teeth convex distoapically.
Comments- The holotype was discovered in 1917 and described in 1924 by Gilmore as a provisionally referred specimen of Chirostenotes (now known to be an oviraptorosaur whose mandibles were named Caenagnathus). Currie et al. (1990) soon described it as a new genus after Currie and Russell (1988) recognized Chirostenotes as an oviraptorosaur. Currie et al. did leave open the possibility Richardoestesia could be synonymous with Chirostenotes or Elmisaurus however.
Horseshoe Canyon Formation Richardoestesia gilmorei are said to be identical to Judith River specimens by Baszio (1997), but taller by Larson et al. (2010). Though R. gilmorei has been reported from the Cenomanian-Maastrichtian of the US and Canada (e.g. Baszio, 1997), these are slightly different from Judith River specimens when examined and it is likely they are separate species.
References- Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous of Alberta. Canada Department of Mines Geological Survey Bulletin (Geological Series). 38(43), 1-12.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.
Fiorillo and Currie, 1994. Theropod teeth from the Judith River Formation (Upper Cretaceous) of south-central Montana. Journal of Vertebrate Paleontology. 14(1), 74-80.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998. Baby hadrosaurid material associated with an unusually high abundance of Troodon teeth from the Horseshoe Canyon Formation, Upper Cretaceous, Alberta, Canada. Gaia. 15, 123-133.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend, Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology. 76(4), 751-763.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta, with special reference to the Albertosaurus sarcophagus bonebed. Canadian Journal of Earth Sciences. 47(9), 1159-1181.
Larson and Currie, 2013. Multivariate analyses of small theropod dinosaur teeth and implications for paleoecological turnover through time. PloS ONE. 8(1), e54329.
Farke, Letteau Stallings and Andrews, 2020. New vertebrate localities and biostratigraphic interpretations of the Mesaverde Formation (Campanian, Late Cretaceous) in northwestern Wyoming. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 136-137.
R. cf. gilmorei sp. nov. (Baszio, 1997)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(UA MR-4: 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20) seventeen teeth
Comments- Milk River Formation R. gilmorei-like teeth have a greater variety of serration shape than Dinosaur Park examples (rectangular to slightly pointed) and grade into Saurornitholestes sp. of the same formation. They may indicate the R. gilmorei lineage descended from dromaeosaurids in the Santonian-Early Campanian, or they may simply be an example of convergence.
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
(UA KUA-1: 85, 120, 108) three teeth
(UA LSF-140: 105) tooth
(UA KUA-18:114) tooth
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- teeth
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Krumenacker, Simon, Scofield and Varricchio, 2016)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2167/50102; Morph 6) incomplete tooth (?x3.1x1.2 mm)
Reference- Krumenacker, Simon, Scofield and Varricchio, 2016. Theropod dinosaurs from the Albian-Cenomanian Wayan Formation of eastern Idaho. Historical Biology. DOI: 10.1080/08912963.2015.1137913
R. cf. gilmorei (Stokosa, 2005)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Material- (FMNH PR2899) tooth (2.7x1.3x.8 mm) (Gates, Zanno and Makovicky, 2015)
(SDSM 64372b) tooth (Stokosa, 2005)
Comments- Gates et al. (2015) referred FMNH PR2899 to Dromaeosauridae, as a "morphology not previously described in the literature." While it differs from most Richardoestesia teeth in being fluted, the shape and small serrations size (19 per mm) are similar. Although the authors provide a PCA analysis which "found it to be markedly different from Richardoestesia teeth", their figure indicates there are Richardoestesia specimens which are equally distant outliers.
Reference- Stokosa, 2005. Enamel microstructure variation within the Theropoda. in Carpenter (ed). The Carnivorous Dinosaurs. 163-178.
Gates, Zanno and Makovicky, 2015. Theropod teeth from the upper Maastrichtian Hell Creek Formation "Sue" Quarry: New morphotypes and faunal comparisons. Acta Palaeontologica Polonica. 60(1), 131-139.
R. cf. gilmorei (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
(BTB: 135, 154, 155, 158, 159, 160) six teeth
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Diem, 1999)
Campanian, Late Cretaceous
Williams Fork Formation, Colorado, US
Reference- Diem, 1999. Vertebrate faunal analysis of the Upper Cretaceous Williams Fork Formation, northwestern Colorado. Unpublished Masters Thesis. San Diego, San Diego State University. 188 pp.
R. cf. gilmorei (Lewis, Heckert, Lucas and Williamson, 2007)
Late Santonian-Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
Material- teeth
Reference- Lewis, Heckert, Lucas and Williamson, 2007. A diverse new microvertebrate fauna from the Upper Cretaceous (Late Santonian-Early Campanian) Menefee Formation of New Mexico. Journal of Vertebrate Paleontology. 27(3), 105A.
Lewis, Heckert and Lucas, 2008. A mixed marine/non-marine and terrestrial microvertebrate assemblage from the Late Cretaceous (Late Santonian-Early Campanian) Menefee Formation of New Mexico: Fauna, biostratigraphy, and paleoecology. Journal of Vertebrate Paleontology. 28(3), 105A.
R. cf. gilmorei sp. nov. (Lee, 1995)
Cenomanian, Late Cretaceous
Woodbine Formation, Texas, US
Material- (SMU 73779) tooth
Comments- This is straight like R. gilmorei, but with larger serrations (4/mm).
References- Lee, 1995. Mid-Cretaceous archosaur faunal changes in Texas. in Sun and Wang (eds.). Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota, Short Papers. China Ocean Press, Beijing. 143-146.
Lee, 1997. The Archosauria from the Woodbine Formation (Cenomanian) in Texas. Journal of Paleontology. 71(6), 1147-1156.
R. cf. gilmorei (Frederickson, Lipka and Cifelli, 2018)
Late Aptian, Early Cretaceous
Dinosaur Park / Cherokee-Sanford Brick Clay Pit / Muirkirk Clay Pit USNM 41614, Arundel Formation, Prince George's County, Maryland, US
Material- (USNM 497748) incomplete tooth (~10.0x~3.7x? mm)
(USNM 497749) tooth
Comments- Frederickson et al. (2018) state "These teeth range from 5-11 mm in height and maximum widths of 2-4 mm. Denticles are present on both mesial and distal margins but the mesial denticles appear to be smaller. The teeth are mesiodistally thinner than those attributed to Deinonychus sp. and possess much smaller denticles on both margins. Further, the general curvature of the teeth is different, in that teeth from Theropoda indet. A are more evenly curved, while those of Deinonychus sp. have a straight base and curve more sharply higher up the tooth. These teeth are similar to those reported elsewhere as Richardoestesia."
Reference- Frederickson, Lipka and Cifelli, 2018. Faunal composition and paleoenvironment of the Arundel Clay (Potomac Formation; Early Cretaceous), Maryland, USA. Palaeontologia Electronica. 21.2.31A, 1-24.
R. cf. gilmorei (Metcalf and Walker, 1994)
Early Bathonian, Middle Jurassic
Chipping Norton Formation, England
Material- (GLRCM coll.) tooth (2.9 mm; FABL 1.5 mm)
Comments- This tooth was labeled as "dromaeosaur-like" by Metcalf and Walker (1994).
Mesial serrations are absent, and the crown is elongate and recurved. Serrations are fairly flat and not hooked apically, but are taller than wide. Serration density is 19/mm. Blood grooves are not apparent.
The high serration density is characteristic of Richardoestesia, while the serration size compared to FABL and curvature match R. gilmorei more than R. isosceles. No features distinguishing this tooth from R. gilmorei can be determined, though it is smaller than most specimens of that species.
Reference- Metcalf and Walker, 1994. A new Bathonian microvertebrate locality in the English Midlands. in Fraser and Sues (eds.). In the Shadow of the Dinosaurs- Mesozoic Small Tetrapods, Cambridge (Cambridge University Press). 322-332.
R. cf. gilmorei (Hendrickx and Mateus, 2014)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Material- (ML 939) lateral tooth (5.1x2.8x1 mm)
Reference- Hendrickx and Mateus, 2014. Abelisauridae (Dinosauria: Theropoda) from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution for the identification of isolated theropod teeth. Zootaxa. 3759(1), 1-74.
R. cf. gilmorei (Torices, 2002)
Late Campanian, Late Cretaceous
Vicari 4, Tremp Formation, Spain
Material- (DPM-VIR4-T6) tooth (1.5x1.4x.7 mm)
(DPM-VIR4-T7) tooth (2.2x1.6x.6 mm)
Comments- These were referred to Dromaeosauridae indet. 4 by Torices (2002), and cf. Richardoestesia sp. by Torices et al. (2015).
References- Torices, 2002. Los dinosaurios terópodos del Cretácico Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Paleontología. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.
R. cf. gilmorei (Buffetaut, Marandat and Sige, 1986)
Early Campanian, Late Cretaceous
Villeveyrac, Herualt, France
Material- (Universite des Sciences et Techniques de Languedoc VIC 17) tooth (3.5 mm) (Buffetaut, Marandat and Sige, 1986)
?(Costa coll.) several teeth (Buffetaut, Costa, Le Loeuff, Martin, Rage, Valentin and Tong, 1996)
Comments- VIC 17 is short and recurved, with 10 distal serraions per mm, and 15 mesial serrations per mm. The distal serrations seem rounded and perpendicular to the crown, while the mesial serrations are only present in the apical portion. The BW/FABL is about .44. Though assigned to Dromaeosauridae by Buffetaut et al. (1986), the small serration size and high DSDI make it almost identical to Richardoestesia gilmorei. Its curvature, short crown, and perhaps constricted base distinguish it from R. isosceles.
Buffetaut et al. (1996) later noted several small laterally compressed and serrated teeth which they referred to small theropods, possibly dromaeosaurids. These are tentatively retained here untl they are described further.
Reference- Buffetaut, Marandat and Sige, 1986. Decourvert de dents de Deinonychosaures (Saurischia, Theropoda) dans le Creace superieur du Sud de la France. Les Comptes rendus de l'Académie des sciences. 303, Serie II(15), 1393-1396.
Buffetaut, Costa, Le Loeuff, Martin, Rage, Valentin and Tong, 1996. An Early Campanian vertebrate fauna from the Villeveyrac Basin (Herault, southern France). Neues Jahrbuch fur Geologie und Palaontologie, Monatshefte. 1, 1-16.
R. cf. gilmorei (Buffetaut, Marandat and Sige, 1986)
Middle-Late Campanian, Late Cretaceous
La Neuve, Aix, France
Material- (Universite des Sciences et Techniques de Languedoc NEV 12) partial tooth (~3.5 mm)
Comments- This tooth is very similar to VIC 17 where preserved, though the mesial carina is unpreserved apically (making the presence of serrations uncertain), and slightly larger serrations are present distally (8/mm). It was similarly assigned to Dromaeosauridae by Buffetaut et al. (1986), but assigned to Richardoestesia cf. gilmorei here for the same reasons as VIC 17 - small distal serrations, short crown, constricted base.
References- Buffetaut, Marandat and Sige, 1986. Decourvert de dents de Deinonychosaures (Saurischia, Theropoda) dans le Creace superieur du Sud de la France. Les Comptes rendus de l'Académie des sciences. 303, Serie II(15), 1393-1396.
R? isosceles Sankey, 2001
Late Campanian, Late Cretaceous
Lower Aguja Formation, Texas, US
Holotype- (LSUMG 489:6238) tooth
Paratypes- (LSUMG 489:6233) tooth
(LSUMG 489:6234) tooth
(LSUMG 489:6235) tooth
(LSUMG 492:6264) tooth
Referred- (LSUMG 140:6140) tooth (Sankey, 2001)
(LSUMG 140:6051) tooth (Sankey, Standhardt and Schiebout, 2005)
(LSUMG 489:6050) tooth (Sankey, Standhardt and Schiebout, 2005)
(TMM 43057-313) tooth (Rowe, Cifelli, Lehman and Weil, 1992)
Late Santonian-Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
teeth (Lewis, Heckert and Forys, 2006)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(CMN coll.) teeth (Russell, 1935)
(UA MR-4: 36, 37, 38, 39, 40, 41) six teeth (Bazsio, 1997)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
(UW 34821) tooth (Demar and Breithaupt, 2006)
(UW 34822) tooth (Demar and Breithaupt, 2006)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(AMNH 8549) tooth (Sahni, 1972)
(AMNH coll.) eleven teeth (Sahni, 1972)
Late Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
(RTMP 96.62 coll.) <6 teeth (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
(RTMP 89.103.25) tooth (5.4 mm) (Sankey et al., 2002)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
(RTMP 84.1.12) tooth (~11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.36.97) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.23.105) tooth (5.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.33.54) tooth (~4.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.34.43) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.45.46) tooth (~5.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.159.62) tooth (~7.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.195.42) tooth (<5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.16.19) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.99.48) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.65) tooth fragment (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.157.52) tooth fragment (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.36.199) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.86.44) tooth (~8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.136.56) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.79.31) tooth (6.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.177.49a) tooth (~8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.177.49b) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.180.5a) tooth (5.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.180.5b) tooth (4.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.181.10) tooth (7.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.10.35) tooth fragment (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.48.11) tooth (~5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.25) tooth (<4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.30a) tooth (12.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.31a) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.31b) tooth (10.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.142.19) tooth (9.7 mm) (Ryan and Russell, 2001)
teeth (Baszio, 1997)
Early Maastrichtian, Late Cretaceous
Upper Aguja Formation, Texas, US
(LSUMG 113:5939) tooth (Sankey, Standhardt and Schiebout, 2005)
(LSUMG 741:5933) tooth (Sankey, Standhardt and Schiebout, 2005)
(LSUMG 741:5934) tooth (Sankey, Standhardt and Schiebout, 2005)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
teeth (Currie, Rigby and Sloan, 1990)
teeth (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, North Dakota, South Dakota, US
teeth (Currie, Rigby and Sloan, 1990)
teeth (Sankey, 2001)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
(RTMP 1019) tooth (Baszio, 1997)
teeth (Ryan and Russell, 2001)
Maastrichtian, Late Cretaceous
Ocozocoautla Formation, Mexico
(IHNFG-0537) maxillary tooth (11.6x7.3x4.8 mm) (Carbot-Chanona and Rivera-Sylva, 2011)
Late Maasstrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Paratype- (AMNH 8113) tooth (Estes, 1964)
Referred- (AMNH coll.) teeth (Estes, 1964)
(UA BTB: 144, 143) two teeth (Baszio, 1997)
teeth (Derstler, 1994, 1995)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, US
(UA LSF140: 107, 111, 112, 115, 116, 118) six teeth (Baszio, 1997)
Late Cretaceous
North America
Paratype- (RTMP 91.170.9) tooth
Diagnosis- (from Sankey et al., 2002) differs from R. gilmorei in- teeth generally longer, straighter and smaller; serrations square-shaped, and with larger interdenticle spaces; no teeth constricted at base.
differs from Shanag in- teeth straighter and generally longer; all teeth have distal serrations; some teeth have mesial serrations.
differs from Sinornithosaurus in- teeth straighter and generally longer.
Comments- Russell (1935) first reported these kinds of teeth and hypothesized they were tyrannosaurid ("deinodont") premaxillary teeth. Estes (1964) believed them to be from juvenile theropods, while Sahni (1972) tentatively assigned them to Sebecosuchia. They were first connected to Richardoestesia gilmorei remains by Currie et al. (1990), who suspected they belonged to a new species of that genus. It was referred to as Richardoestesia sp. by Baszio (1997) and other sources until being named by Sankey in 2001. Sankey's paratypes are from several formations, and the species has been reported from the Early Campanian to the Late Maastrichtian of North America. These show very little variation (Baszio, 1997) though we may assume more than one species was present in this time interval. Additional similar teeth are known from the Late Jurassic to Late Cretaceous of Eurasia and the Early Cretaceous of the US, but differ slightly and are referred to new unnamed species below. R. isosceles only shares a couple apomorphic features with R. gilmorei (tiny serrations, convex distoapical edge of some teeth) and may not be closely related. The fact its lineage extends back to Kimmeridgian Europe, while R. gilmorei seems to emerge from dromaeosaurid teeth in the Early Campanian of North America may support this view. However, Williamson and Brusatte (2014) note both gilmorei-like and isosceles-like teeth from the Fruitland Formation have a high DSDI, suggesting they derive from the same species. Sankey (2002) has suggested Paronychodon and Richardoestesia teeth are morphotypes of the same taxon, based on morphology and relative abundance. The details of this study have yet to be published, though it does make sense stratigraphically, as both taxa first appear in Late Jurassic Europe and spread to North America in the Albian, with Late Cretaceous examples known from the western North America, central Asia and Europe. It's also logical anatomically, as Richardoestesia? isosceles would be expected to have some unserrated and possibly constricted teeth if it were microraptorian. It should be noted Paronychodon has priority over Richardoestesia, and lacustris and caperatus both have priority over isosceles. Also, Euronychodon has priority over Asiamericana, and portuculensis has priority over both asiatica and asiaticus. So if this synonymy is proven, none of the names associated with straight-toothed Richardoestesia will survive synonymization. Averianov and Sues (2019) believe "R. isosceles cannot be differentiated from R. asiatica", but nonsensically also suggest R. isosceles "should rather be treated as a nomen dubium" because the type tooth "appears to be rather short, with distinctly convex crown sides. This contrasts with the other straight teeth of Richardoestesia, which have a proportionally taller crown with straight or only slightly convex crown margins." The shortness could be simply due to being incomplete basally, it and the degree of convexity are not shown to differ from all other supposed R. isosceles specimens, let alone those from the type Aguja Formation, and if its differences were taxonomically significant it would be a valid taxon instead of a nomen dubium (which is more realistic given the temporal and geographical separation). Further, while a fragmentary type may be problematic in some circumstances, the difference between a complete and incomplete tooth is negligible. In any case, the species are kept separate here based on the previously noted geographical and temporal difference, and this may even be defensible morphologically pending study of the other Aguja Formation R. isosceles specimens.
Ryan and Russell (2001) list "uncatalogd TMP teeth" as "Richardsoestesia n. sp. Baszio 1997b" from the Foremost Formation, which are some of the six Richardoestesia teeth reported by Peng et al. (2001) from the PHR-1 and PHR-2 localities. The latter listed "RTMP96.62.26 and -31" as "Richardoestesia sp. Baszio, 1997" from the Foremost and Oldman Formations, so one of those numbers represents the less than six Foremost Formation R. isosceles teeth.
The tooth LSUMG 140:6140 was first referred to Saurornitholestes cf. langstoni by Sankey (2001) before being identified as R? isosceles by Sankey et al. (2002). Similarly, Oldman tooth RTMP 89.103.25 was identified as Aves indet. by Sankey et al. (2002) before being reidentified as Richardoestesia isolsceles by Larson and Currie (2013) and Dumont et al. (2016).
References- Russell, 1935. Fauna of the Upper Milk River beds, Southern Alberta. Transactions, Royal Society of Canada. 3(29),115-127.
Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin of the AMNH. 147.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press, New York. 107-125.
Rowe, Cifelli, Lehman and Weil, 1992. The Campanian Terlingua Local Fauna, with a summary of other vertebrates from the Aguja Formation, Trans-Pecos Texas. Journal of Vertebrate Paleontology. 12(4), 472-493.
Derstler, 1994. Dinosaurs of the Lance Formation in eastern Wyoming. in Nelson (ed.). Wyoming Geological Association. Forty-Fourth Annual Field Conference Guidebook. 127-146.
Derstler, 1995. The Dragons’ Grave - an Edmontosaurus bonebed containing theropod egg shells and juveniles, Lance Formation, (Uppermost Cretaceous), Niobrara County, Wyoming: Journal of Vertebrate Paleontology. 15(3), 26A.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Sankey, 1997. Late Cretaceous vertebrate paleontology and Paleoecology, Upper Aguja Formation, Big Bend National Park, Texas. Journal of Vertebrate Paleontology. 17(3), 73A.
Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages (exclusive of mammals) from the Foremost and Oldman Formations of the Judith River Group (Campanian) of southeastern Alberta: An illustrated guide. Provincial Museum of Alberta Natural History Occasional Paper. 25, 54 pp.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp. 279-297.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend, Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, 2002. Theropod dinosaur diversity in the latest Cretaceous (Maastrichtian) of North America. Journal of Vertebrate Paleontology. 22(3), 103A.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology. 76(4), 751-763.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous (Campanian-Maastrichtian), Big Bend National Park, Texas. In Carpenter (ed). The Carnivorous Dinosaurs. 127-152.
Demar and Breithaupt, 2006. The nonmammalian vertebrate microfossil assemblages of the Mesaverde Formation (Upper Cretaceous, Campanian) of the Wind River and Bighorn Basin, Wyoming. in Lucas and Sullivan (eds). Late Cretaceous Vertbrates from the Western Interior. New Mexico Museum of Natural History & Science. Bulletin 35, 33-53.
Lewis, Heckert and Forys, 2006. Paleoecology of the aqueous paleoenvironments of the Late Cretaceous (Early Campanian) Allison Member of the Menefee Formation in Northwestern New Mexico. NMGS Annual Spring Meeting, abstracts.
Carbot-Chanona and Rivera-Sylva, 2011. Presence of a maniraptoriform dinosaur in the Late Cretaceous (Maastrichtian) of Chiapas, Southern Mexico. Boletin de la Sociedad Geologica Mexicana. 63(3), 393-398.
Sankey, 2012. Something's fishy: Was one of the most abundant Latest Cretaceous theropods a fish-eater? Journal of Vertebrate Paleontology. Program and Abstracts 2012, 165.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Dumont, Tafforeau, Bertin, Bhullar, Field, Schulp, Strilisky, Thivichon-Prince, Viriot and Louchart, 2016. Synchrotron imaging of dentition provides insights into the biology of Hesperornis and Ichthyornis, the "last" toothed birds. BMC Evolutionary Biology. 16:178.
Averianov and Sues, 2019. Morphometric analysis of the teeth and taxonomy of the enigmatic theropod Richardoestesia from the Upper Cretaceous of Uzbekistan. Journal of Vertebrate Paleontology. 39(3), e1614941.
R? cf. isosceles (Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997)
Cenomanian- Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (NCSM 33274) tooth (4.21x1.65x0.98 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
(NCSM 33288) incomplete tooth (~9x4.05x1.9 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
(OMNH coll.) teeth (Kirkland et al., 1997)
three partial teeth (Garrison et al., 2007)
Comments- Kirkland et al. (1997) and Cifelli et al. (1999) list cf. Richardoestesia sp. teeth. Kirkland (pers. comm. to Demirjian, 9-2019) stated "Most of these reports were based on studies we did on basal Cenomanian age teeth in the Mussentuchit from the Oklahoma Mus. Nat. History in the 1990s." Garrison et al. (2007) described and illustrated three partial teeth which are tall and straight, some having tiny serrations. They referred these to Richardoestesia cf. isosceles.
References- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial Cretaceous vertebrates from the Cedar Mountain Formation, Emery County, Utah: the Mussentuchit Local Fauna. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 219-242.
Garrison, Brinkman, Nichols, Layer, Burge and Thayn, 2007. A multidisciplinary study of the Lower Cretaceous Cedar Mountain Formation, Mussentuchit Wash, Utah: a determination of the paleoenvironment and paleoecology of the Eolambia caroljonesa dinosaur quarry. Cretaceous Research. 28, 461-494.
Avrahami, 2018. Paleobiodiversity of a new microvertebrate locality from the Upper Cretaceous Mussentuchit Member, Cedar Mountain Formation, Utah: Testing morphometric multivariate approaches for quantifying shape variation in microvertebrate specimens. Masters thesis, North Carolina State University. 181 pp.
Avrahami, Gates, Heckert, Makovicky and Zanno, 2018. A new microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain Formation: Insights into the paleobiodiversity and paleobiogeography of early Late Cretaceous ecosystems in western North America. PeerJ. 6:e5883.
R? cf. isosceles sp. nov. (Williamson, 2001)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-52503) tooth (1.5x1x.2 mm) (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
(NMMNH P-32753) tooth (?x3.2x1.5 mm) (Williamson and Brusatte, 2014)
Comments- Williamson and Brusatte (2014) note this differs from other Richardoestesia isosceles in that it has a high DSDI, as do recurved Richardoestesia teeth from the formation.
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
R? cf. isosceles sp. nov. (Lee, 1995)
Cenomanian, Late Cretaceous
Woodbine Formation, Texas, US
Material- (SMU 73778) tooth
Comments- This is straight like R. isosceles, but much shorter and with larger serrations (4/mm).
References- Lee, 1995. Mid-Cretaceous archosaur faunal changes in Texas. in Sun and Wang (eds.). Sixth Symposium on Mesozoic Terrestrial Ecosystems and Biota, Short Papers. China Ocean Press, Beijing. 143-146.
Lee, 1997. The Archosauria from the Woodbine Formation (Cenomanian) in Texas. Journal of Paleontology. 71(6), 1147-1156.
R? cf. isosceles sp. nov. (Zinke, 1998)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Material- (IPFUB Gui D 118-155) forty teeth (~3.42 mm)
Comments- These are very similar to R. isosceles, but are less tall on average and can have more distal serrations (up to 14/mm).
Reference- Zinke, 1998. Small theropod teeth from the Upper Jurassic coal mine of Guimarota (Portugal). Palaontologische Zeitschrift. 72(1/2), 179-189.
R? cf. isosceles sp. nov. (Rauhut and Zinke, 1995)
Barremian, Early Cretaceous
Una Formation, Spain
Material- (IPFUB Una Th 1–20, 64, 68, 81) forty-seven teeth (3-5 mm)
Comments- These are very similar to R. isosceles, but differ in that some teeth lack serrations on both mesial and distal carinae.
References- Rauhut and Zunke, 1995. A description of the Barremian dinosaur fauna from Una with a comparison of that of Las Hoyas. II. International Symposium of Lithographic Limestone, Extended Abstracts. 123-126.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca, Spain. Cretaceous Research. 23, 255-263.
R? cf. isosceles (Torices, 2002)
Late Campanian-Early Maastrichtian, Late Cretaceous
Montrebei, Tremp Formation, Spain
Material- (DPM-MON-T5) tooth (2.4x1.4x.6 mm) (Torices, 2002)
(DPM-MON-T9) tooth (2.8x1.7x.8 mm) (Torices, 2002)
Late Maastrichtian, Late Cretaceous
Blasi 2B, Tremp Formation, Spain
Material- (MPZ98/72) tooth (4.3x2.2x1.1 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MPZ98/73) tooth (3.4x2.2x.9 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MPZ98/74) tooth (3x1.4x.6 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
(MPZ2004/7) tooth (2.1x1x.4 mm) (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
Comments- These are only assigned to cf. Richardoestesia sp. by Torices et al. (2015), but resemble R. isosceles in the convex or straight distal edge. DPM-MON-T5 and T9 were assigned to Dromaeosauridae indet. 4 by Torices (2002).
References- Torices, 2002. Los dinosaurios terópodos del Cretácico Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Paleontología. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.
R? cf. isosceles sp. nov. (Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke, 2002)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Material- teeth
Comments- These are very elongate, straight to slightly recurved, with no mesial serrations and up to 5 distal serrations per mm. They were said to be most similar to the straight type of Richardoestesia teeth, but they differ in all lacking mesial serrations.
Reference- Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke, 2002. Dinosaur egg nests, mammals and other vertebrates from a new Maastrichtian site of the Hateg Basin (Romania). C. R. Palevol. V. 1, p. 173-180.
R? asiatica (Nessov, 1995) Sues and Averianov, 2013
= Asiamericana asiatica Nessov, 1995
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Syntypes- (N 460/12457) tooth
(N 12457 coll.) two teeth
Referred- (ZIN PH 1073/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1074/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1075/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1076/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1077/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1078/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1079/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1080/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1081/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1082/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1083/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1084/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1085/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1086/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1087/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1088/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1089/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1090/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1091/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1092/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1093/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1094/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1095/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1096/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1097/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1098/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1099/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1100/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1101/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1102/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1103/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1104/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1105/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1106/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1107/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1108/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1109/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1110/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1111/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1112/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1113/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1114/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1115/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1116/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1117/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1118/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1119/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1120/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1121/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1122/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1123/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1124/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1125/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1126/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1127/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1128/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1129/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1130/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1131/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1132/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1133/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1134/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1135/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1136/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1137/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1138/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1139/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1140/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1141/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1142/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1143/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1144/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1145/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1231/16) tooth (Sues and Averianov, 2013)
(ZIN PH 2315/16) tooth (Sues and Averianov, 2013)
Diagnosis- indistinguishable from R. isosceles.
Comments- Nessov (1989) first published these teeth as ?Spinosauridae, then (1995) named it Asiamericana asiatica and considered it to be either a spinosaurid or a saurodontid fish. However, he noted resemblence between the holotype and a tooth (AMNH 8113) that has since been referred to Richardoestesia isosceles. Ryan (1997) first mentioned Richardoestesia teeth from this formation. Sues and Averianov (2013) redescribed the teeth as a species of Richardoestesia, finding it's indistinguishable from R. isosceles, contra Buffetaut et al. (2008) who stated it was certainly a saurodontid. The holotype teeth were initially described as lacking serrations, but Sues and Averianov show this is untrue. Notably, asiatica has precedence over isosceles, though the species are unlikely to be synonymous given the geographical and age difference. This contrasts with Averianov and Sues (2019) who while agreeing "R. isosceles cannot be differentiated from R. asiatica", nonsensically also suggest R. isosceles "should rather be treated as a nomen dubium" (see Comments under R. isosceles).
References- Nessov, 1989. [New findings of remains of dinosaurs, crocodiles, and flying reptiles of the Late Mesozoic of USSSR]. [Questions of herpetology. Abstracts of the Seventh Herpetological Conference]. 173-174.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov, ekologii i paleobiogeografii. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Ryan, 1997. Middle Asian dinosaurs. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. Academic Press. 442-444.
Buffetaut, Suteethorn, Tong and Amiot, 2008. An Early Cretaceous spinosaurid theropod from southern China. Geological Magazine. 145(5), 745-748.
Sues and Averianov, 2013. Enigmatic teeth of small theropod dinosaurs from the Upper Cretaceous (Cenomanian–Turonian) of Uzbekistan. Canadian Journal of Earth Sciences. 50, 306-314.
Averianov and Sues, 2019. Morphometric analysis of the teeth and taxonomy of the enigmatic theropod Richardoestesia from the Upper Cretaceous of Uzbekistan. Journal of Vertebrate Paleontology. 39(3), e1614941.
R? cf. asiatica (Averianov, 2007)
Middle Albian-Early Cenomanian, Early-Late Cretaceous
Khodzhakul Formation, Uzbekistan
Material- (ZIN PH 1222/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1223/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1224/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1225/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1226/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1227/16) tooth (Sues and Averianov, 2013)
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research. 28, 532-544.
Sues and Averianov, 2013. Enigmatic teeth of small theropod dinosaurs from the Upper Cretaceous (Cenomanian–Turonian) of Uzbekistan. Canadian Journal of Earth Sciences. 50, 306-314.
R. sp. (Druckenmiller, Erickson, Brinkman and Brown, 2012)
Coniacian-Maastrichtian, Late Cretaceous
Prince Creek Formation, Alaska, US
Reference- Druckenmiller, Erickson, Brinkman and Brown, 2012. Dinosaur diversity in the Arctic: New records of polar dinosaurs based on microvertebrate analysis from the Upper Cretaceous Prince Creek Formation, Northern Alaska. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 88.
R. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
(RTMP 88.86.44) tooth (Ryan and Russell, 2001)
(RTMP 96.62 coll.) <6 teeth (Peng, Russell and Brinkman, 2001)
(?UC coll.) tooth (Frampton, 2006)
Comments- Ryan and Russell (2001) list RTMP 88.86.44 as a reference Richardoestesia sp. specimen from the Foremost Formation with additional teeth being referrable. The latter are some of the Richardoestesia gilmorei teeth described by Peng et al. (2001) from the Foremost Formation, with numbers within the range of RTMP 96.62.23-32 and/or 66. These would represent less than six of the teeth found at sites PHR-1 and PHR-2 (as at least one of these is R. isosceles)
Cullen and Evans (2016) list one Richardoestesia tooth from the PHRN site, which equates with Frampton's (2006) thesis.
References- Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages (exclusive of mammals) from the Foremost and Oldman Formations of the Judith River Group (Campanian) of southeastern Alberta: An illustrated guide. Provincial Museum of Alberta Natural History Occasional Paper. 25, 54 pp.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp. 279-297.
Frampton, 2006. Taphonomy and palaeoecology of mixed invertebrate-vertebrate fossil assemblage in the Foremost Formation (Cretaceous, Campanian), Milk River Valley, Alberta. Masters thesis, University of Calgary. 294+ pp.
Cullen and Evans, 2016. Palaeoenvironmental drivers of vertebrate community composition in the Belly River Group (Campanian) of Alberta, Canada, with implications for dinosaur biogeography. BMC Ecology. 16, 52.
R. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
Material- (RTMP 87.78.3) tooth (Ryan, 2003)
(RTMP 89.79.62) tooth (Ryan and Russell, 2001)
teeth (Ryan and Russell, 2001)
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp. 279-297.
Ryan, 2003. Taxonomy, systematics and evolution of centrosaurine ceratopsids of the Campanian Western Interior basin of North America. Unpublished PhD thesis. Department of Biological Sciences, Calgary, Alberta. 578pp.
R. sp. (Fanti and Miyashita, 2009)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Material- (RTMP 2004.93.3) incomplete tooth
Reference- Fanti and Miyashita, 2009. A high latitude vertebrate fossil assemblage from the Late Cretaceous of west-central Alberta, Canada: Evidence for dinosaur nesting and vertebrate latitudinal gradient. Palaeogeography, Palaeoclimatology, Palaeoecology. 275, 37-53.
R. sp. indet. (Mongelli and Varricchio, 1998)
Early Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- two teeth
Reference- Mongelli and Varricchio, 1998. Theropod teeth of the Lower Two Medicine Formation (Campanian) of northwestern Montana. Journal of Vertebrate Paleontology. 18(3), 64A.
R. sp. indet. (Varricchio, 1995)
Mid-Late Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
Material- teeth? (Varricchio, 1995)
teeth (Redman, Moore and Varricchio, 2015)
References- Varricchio, 1995. Taphonomy of Jack’s Birthday Site, a diverse dinosaur bonebed from the Upper Cretaceous Two Medicine Formation of Montana. Palaeogeography, Palaecolimatology, Palaeoecology. 114, 297-323.
Redman, Moore and Varricchio, 2015. A new vertebrate microfossil locality in the Upper Two Medicine Formation in the vicinity of Egg Mountain. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 201-202.
R. sp. indet. (Triebold, 1997)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, South Dakota, US
Material- ?(UCMP 119718) teeth (UCMP online)
?(UCMP 119733) tooth (UCMP online)
?(UCMP 119920) eight teeth (UCMP online)
?(UCMP 119921) tooth (UCMP online)
?(UCMP 120075) three teeth (UCMP online)
?(UCMP 120132) tooth (UCMP online)
?(UCMP 120153) tooth (UCMP online)
?(UCMP 120191) two teeth (UCMP online)
?(UCMP 120255) three teeth (UCMP online)
?(UCMP 120287) tooth (UCMP online)
?(UCMP 120338) tooth (UCMP online)
?(UCMP 123543) two teeth (UCMP online)
?(UCMP 123565) tooth (UCMP online)
?(UCMP 128913) tooth (UCMP online)
?(UCMP 128941) tooth (UCMP online)
(UCMP 174810) tooth (UCMP online)
teeth? (Triebold, 1997)
teeth (DePalma, 2010)
Comments- Most of the UCMP specimens (except 174810) are identified as Chirostenotes in their collection, so are referred to Richardoestesia here, as the R. gilmorei holotype was originally referred to Chirostenotes (which is toothless).
References- Triebold, 1997. The Sandy Site: Small Dinosaurs from the Hell Creek Formation of South Dakota. in Wolberg, Stump and Rosenberg (eds). Dinofest International, Proceedings of a Symposium sponsered by Arizona State University. A Publication of The Academy of Natural Sciences. 245-248.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis, University of Kansas. 227 pp.
R. sp. indet. (Stokosa, 2005)
Maastrichtian, Late Cretaceous
Fox Hills Formation, South Dakota, US
Material- (SDSM 14516) tooth
Reference- Stokosa, 2005. Enamel microstructure variation within the Theropoda. In Carpenter (ed.). The Carnivorous Dinosaurs. 163-178.
R. sp. indet. (UCMP online)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- ?(UCMP 120849) over fifty teeth
Comments- Most of the UCMP specimens are identified as Chirostenotes in their collection, so are referred to Richardoestesia here, as the R. gilmorei holotype was originally referred to Chirostenotes (which is toothless).
R. sp. (Wroblewski, 1995)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
Material- teeth
References- Wroblewski, 1995. First report of changes in Lower Vertebrate Faunas across the Cretaceous-Tertiary boundary, Western Hanna Basin, Wyoming. Journal of Vertebrate Paleontology. 15(3), 61A.
Wroblewski, 1998. Changing paleoenvironments and paleofaunas across the K-T boundary, Ferris Formation, Southcentral Wyoming. Tate Geological Museum, Casper College, Casper Wyoming. Tate ’98. Life in the Cretaceous. 53-70.
R. sp. indet. (UCMP online)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- ?(UCMP 119674) tooth
Comments- This is identified as Chirostenotes in their collection, so is referred to Richardoestesia here, as the R. gilmorei holotype was originally referred to Chirostenotes (which is toothless).
R. sp. (Kirkland, Lucas and Estep, 1998)
Barremian, Early Cretaceous
Yellow Cat Member of Cedar Mountain Formation, Utah, US
Material- teeth
Comments- Kirkland et al. (1998) list cf. Richardoestesia sp. under the Lower Cedar Mountain Formation, which corresponds to the Yellow Cat Member. Kirkland (pers. comm. to Demirjian, 9-2019) later stated "Richardoestesia-like teeth in the Yellow Cat toward base (fauna 2) maybe primitive ornithominisaur teeth....Nedcolbertia?????", but if these are truly Richardoestesia-like in having serrations it would be more primitive than known ornithomimosaur cranial material and potentially homoplasic if alvarezsaurs are arctometatarsalians.
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
R. sp. (Kirkland, Lucas and Estep, 1998)
Early Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Comments- Kirkland et al. (1998) list cf. Richardoestesia sp. under the Middle Cedar Mountain Formation, which includes the Ruby Ranch and Poison Strip Members.
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
R. sp. (Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997)
Late Cenomanian, Late Cretaceous
Dakota Formation, Utah, US
Material- teeth
Comments- This is listed as cf. Richardoestesia sp. by Kirkland et al. (1997).
Reference- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young University Geology Studies. 42, 69-103.
R. sp. (Kirkland, Lucas and Estep, 1998)
Middle-Late Turonian, Late Cretaceous
Smoky Hollow Member of the Straight Cliffs Formation, Utah, US
Comments- This is listed as cf. Richardoestesia sp. by Kirkland et al. (1998).
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
R. sp. (Kirkland, Lucas and Estep, 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Comments- This is listed as cf. Richardoestesia sp. by Kirkland et al. (1998).
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
R. sp. (Kirkland, Lucas and Estep, 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah
Comments- This is listed as cf. Richardoestesia sp. by Kirkland et al. (1998).
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
R. sp. (Parrish, 1999)
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Material- (UCM 8656) tooth
Comments- This was listed as cf. Richardoestesia by Parrish (1999).
Reference- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.
R. sp. nov. (Williamson, 2001)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-27488) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27489) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27495) tooth (?x~3.9x1.7 mm) (Williamson and Brusatte, 2014)
(NMMNH P-27496) tooth (?x3.7x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-28379) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30004) tooth (4.1x3.3x1.4 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30005) tooth (?x3.4x1.3 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30277) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30331) tooth (?x3.4x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-36552) tooth (Williamson and Brusatte, 2014)
(NMMNH P-38410) tooth (?x1.5x.7 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38428) tooth (Williamson and Brusatte, 2014)
(NMMNH P-52501) tooth (?x?x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-52514) tooth (?x.9x? mm) (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
(NMMNH P-30207) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30208) tooth (5x2.9x1.4 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30209) tooth (?x2.2x.8 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30219) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30221) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30224) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33478) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33479) tooth (?x1.9x1.1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33480) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33481) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33482) tooth (4.5x2.9x1.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33483) tooth (Williamson and Brusatte, 2014)
Comments- Williamson and Brusatte (2014) note these differ from other Richardoestesia in that they have a high DSDI.
References- Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
R. sp. (Wel and Williamson, 2000; described by Williamson and Brusatte, 2014)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (NMMNH P-22567) tooth (?x2.8x1.4 mm) (Williamson and Brusatte, 2014)
(NMMNH P-32742) tooth (?x2x1.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-37793) tooth (?x2.3x1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-46389) tooth (?x2.8x1.2 mm) (Williamson and Brusatte, 2014)
Reference- Weil and Williamson, 2000. Diverse Maastrichtian terrestrial vertebrate fauna of the Naashoibito Member, Kirtland Formation (San Juan Basin, New Mexico) confirms “Lancian� faunal heterogeneity in western North America. Geological Society of America Abstracts with Programs. 32, A-498.
Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
R. sp. indet. (YPM online)
Late Cretaceous
US
Material- (YPM 56979)
R? sp. (Prieto-Marquez, Gaete, Galobart and Ardevol, 2000)
Late Campanian, Late Cretaceous
L'Abeeler, Aren Sandstone Formation, Spain
Material- (IPS 18 372) tooth (14.03x5.85x3.11 mm)
Comments- Prieto-Marquez et al. (2000) concluded "the minute size and the similar morphology of the denticles of R. gilmorei place the Spanish theropod closer to this species than to any other known taxon." Torices et al. (2015) stated "in the discriminant analysis performed, this tooth is clearly isolated by its height from both the North American and Pyrenean samples, which suggests it might be a new taxon."
References- Prieto-Marquez, Gaete, Galobart and Ardevol, 2000. A Richardoestesia-like theropod tooth from the Late Cretaceous foredeep, south-central Pyrenees, Spain. Eclogae Geologicae Helvetiae. 93(3), 497-501.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.
R. spp. (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
Late Campanian, Late Cretaceous
Laño, Sedano Formation, Spain
Material- (MCNA 14566-14621) sixty-four teeth (1.2-6.6 mm) [details in Torices et al., 2015]
(UPVLP 135) tooth (Isasmendi et al., 2020)
Comments- These teeth seem to fall under both gilmorei and isosceles morphotypes.
References- Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica Polonica. 60(3), 611-626.
Isasmendi, Torices, Canudo and Pereda-Suberbiola, 2020. Paleobiodiversity of theropod dinosaurs from the Upper Cretaceous Laño site, northern Iberian peninsula. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 186-187.
R? sp. (Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- teeth
Comments- Ortega et al. (2009) state Richardoestesia teeth are known, calling it a basal tetanurine, and Torices et al. (2011) mention cf. Richardoestesia.
References- Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009. The vertebrate diversity of the Upper Campanian-Lower Maastrichtian "Lo Hueco" fossil-site (Cuenca, Spain). Journal of Vertebrate Paleontology. 29(3), 159A-160A.
Torices, Barroso-Barcenilla, Cambra-Moo, Perez and Serrano, 2011. Vertebrate microfossil analysis in the palaeontological site of 'Lo Hueco' (Upper Cretaceous, Cuenca, Spain). Journal of Vertebrate Paleontology. Program and Abstracts 2011, 205.
R. sp. (Valentin, Godefroit, Tabuce, Vianey-Liaud, Wenhao and Garcia, 2012)
Late Maastrichtian, Late Cretaceous
Vitrolles-La-Plaine, Provence, France
Material- (UP-VLP-08-001) tooth
(UP coll.) several teeth
Reference- Valentin, Godefroit, Tabuce, Vianey-Liaud, Wenhao and Garcia, 2012. First Late Maastrichtian (Latest Cretaceous) vertebrate assemblage from Provence (Vitrolles-la-Plaine, Southern France). In Godefroit (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 582-597.
R? sp. (Debeljak, Kosir and Otonicar, 1999)
Campanian-Maastrichtian, Late Cretaceous
Kozina, Slovenia
Material- (ACKK-D-8/081) tooth
Comments- This tooth was identified as Dromaeosauridae based on the high DSDI, and furthermore seems to be Richardoestesia because of the small serrations (~8 per mm) and elongate shape.
References- Debeljak, Kosir and Otonicar, 1999. A preliminary note on dinosaurs and non-dinosaurian reptiles from the Upper Cretaceous carbonate platform succession at Kozina (SW Slovenia). Razprave IV. razreda SAZU. 40, 3-25.
Debeljak, Kosir, Buffetaut and Otonicar, 2002. The Late Cretaceous dinosaurs and crocodiles of Kozina (SW Slovenia): A preliminary study. Memorie della Societa Geologica Italiana. 57, 193-201.
R. sp. indet. (Nessov, 1995)
Santonian-Early Campanian, Late Cretaceous
Syuk-Syuk Formation, Kazakhstan
Material- jaw, teeth
Comments- These teeth are serrated, unlike R. asiatica.
Reference- Nessov, 1995. Dinozavri severnoi Yevrasii: Novye dannye o sostave kompleksov, ekologii i paleobiogeografii [Dinosaurs of Northern Eurasia: new data about assemblages, ecology and paleobiogeography], Scientific Research Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia: 156 pp. + 14 pl. [in Russian with short English, German, and French abstracts].
R. spp. (Alifanov and Bolotsky, 2002)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- teeth
Comments- Alifanov and Bolotsky (2002) referred teeth to two morphs of Richardoestesia, possibly gilmorei and isosceles.
Reference- Alifanov and Bolotsky, 2002. New data about the assemblages of the Upper Cretaceous carnivorous dinosaurs (Theropoda) from the Amur region. In Kirillova (ed.). Fourth International Symposium of IGCP 434. Cretaceous continental margin of East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
R? sp. (Knoll and Ruiz-Omenaca, 2005)
Beriassian, Early Cretaceous
KM 1983, Ksar Metlili Formation, Morocco
Material- (MNHN SA 2004/5B; Velociraptorinae indet. Morphotype II; lost) incomplete tooth (?x?x1.40 mm)
Comments- This was collected in 1983, 1986 or 1999, and identified by Knoll and Ruiz-Omenaca (2005) as Velociraptorinae indet. Morphotype II. They noted resemblences to European Richardoestesia teeth and the serration density (7.52-8.33 per mm) matches that genus. Lasseron (2020) noted the "dinosaur remains (Knoll, 2000; Knoll & Ruiz-Omeñaca, 2009) ... were taken out of the MNHN and lost" (translated).
References- Knoll and Ruiz-Omenaca, 2009. Theropod teeth from the basalmost Cretaceous of Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine. 146(4), 602-616.
Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des vertebres mesozoiques africans et gondwaniens : apport des gisements du Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle. 493 pp.
R? sp. (Prasad, Parmar and Kumar, 2014)
Early Jurassic
Kota Formation, India
Material- teeth
Comments- These were described as "Richardoestesia-like."
Reference- Prasad, Parmar and Kumar, 2014. Recent vertebrate fossil discoveries from the Jurassic Kota Formation of India. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 208.
R. sp. (Bertini and Franco-Rosas, 2001)
Turonian-Late Maastrichtian, Late Cretaceous
Adamantina, Marília and/or Serra da Galga Formations, Bauru Group, Brazil
Material- teeth
Reference- Bertini and Franco-Rosas, 2001. Scanning electron microscope analysis on Maniraptoriformes teeth from the Upper Cretaceous of southeastern Brazil. JVP 21(3) 33A.

undescribed possible Microraptoria (Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997)
Early Cenomanian, Early Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71599) two teeth (Fiorillo, 1999)
(OMNH coll.; Morphotype 7 of Frederickson et al., 2018) teeth (<5 mm)
Comments- These are identified as cf. Troodon sp. in Kirkland et al. (1997), and Troodontidae gen. et sp. indet. by Cifelli et al. (1999). They are probably "the earliest troodontids" mentioned by Parrish and Eaton (1991) and troodontids of Kirkland and Parrish (1995) from the upper Cedar Mountain Formation. Frederickson et al. (2018) lists these as "?troodontid" but also say they "have relatively smaller denticles than those of most Late Cretaceous troodontids ... ; though more associated material is needed to unequivocally dismiss this identification." They also noted Fiorillo (1999) identified this morphotype as Velociraptorinae. Given the lack of mesial serrations, small size, Middle Cretaceous timing and longitudinal grooves on some specimens, these are here suggested to be microraptorians.
References- Parrish and Eaton, 1991. Diversity and evolution of dinosaurs in the Cretaceous of the Kaipirowits plateau, Utah. Journal of Vertebrate Paleontology. 11(3), 50A.
Kirkland and Parrish, 1995. Theropod teeth from the Lower Cretaceous of Utah. Journal of Vertebrate Paleontology. 15(3), 39A.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the central Colorado plateau: A key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial Cretaceous vertebrates from the Cedar Mountain Formation, Emery County, Utah: The Mussentuchit local fauna. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison eggshell site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 259-268.
Goldberg, 2000. Faunal composition, non-marine vertebrates, of the upper Cedar Mountain Formation (Cretaceous: Albian-Cenomanian), central Utah. PhD thesis, University of Oklahoma. 243 pp.
Frederickson, Engel and Cifelli, 2018. Niche partitioning in pheropod dinosaurs: Diet and habitat preference in predators from the uppermost Cedar Mountain Formation (Utah, U.S.A.). Scientific Reports. 8:17872.

unnamed microraptorian (Averianov, Starkov and Skutschas, 2003)
Late Barremian-Mid Aptian, Early Cretaceous
Mogoito Member of Murtoi Formation, Russia
Material- (ZIN PH 10/13) anterior lateral tooth (FABL 2.65 mm)
(ZIN PH 11/13) posterior lateral tooth (FABL 4.1 mm), posterior lateral tooth (FABL 5.45 mm)
Comments- These teeth were referred to Dromaeosauridae by Averianov et al. (2003), but are here specified as microraptorians based on their small serrations (9-10/mm mesially, 6-9/mm distally). The anterior tooth is elongate and only slightly curved, while the posterior teeth are short and strongly recurved. The BW/FABL of the anterior tooth is .47, and the posterior teeth .52-.56. The mesial carinae follow the midline and are serrated only basally on the anterior tooth. Posterior serrations are rectangular, broad labiolingually, and have short shallow interdenticle slits. They resemble teeth of Sinornithosaurus and Richardoestesia.
Reference- Averianov, Starkov and Skutschas, 2003. Dinosaurs from the Early Cretaceous Murtoi Formation in Buryatia, Eastern Russia. Journal of Vertebrate Paleontology. 23(3):586–594.

undescribed possible microraptorian (Kurochkin, 1988)
Aptian-Albian, Early Cretaceous
Khoboor, Mongolia
Material- teeth
Comments- These have unserrated mesial carinae, and distal carinae with very small serrations. There is a slight basal constriction, and the crown is strongly recurved. They were referred to birds by Kurochkin (1998), but to troodontids by Nessov and Golovneva (1990) and Averianov and Sues (2007). However, they also sound similar to Shanag and perhaps Richardoestesia.
References- Kurochkin, 1988. [Cretaceous Mongolian birds and their significance for the study of bird phylogeny.] Trudy Sovmestnoi Sovetsko-Mongol’skoi Paleontologicheskoi Ekspeditsii. 34, 33–42. [Russian]
Nessov and Golovneva, 1990. [History of the flora, vertebrates and climate in the late Senonian of the north-eastern Koriak Uplands]. in Krasilov (ed). [Continental Cretaceous of the USSR.] Dal’nevostochnoe Otdelenie AN SSSR, Vladivostok. [Russian]. 191-212.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the Cenomanian of Uzbekistan, with a review of troodontid records from the territories of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.

unnamed possible microraptorian (Pittman, Pei and Xu, 2015)
Aptian-Albian, Early Cretaceous
Bayan Gobi Formation, Inner Mongolia, China
Material- (IVPP V22530) partial dorsal rib, manual ungual, manual claw sheath, distal femur, tibia (~75 mm), fibula, metatarsal I, incomplete metatarsal II, fragmentary phalanx II-1, partial phalanx II-2, pedal ungual II, incomplete metatarsal III, incomplete phalanx III-1, incomplete metatarsal IV, phalanx IV-1, phalanx IV-2, two pedal phalanges, partial metatarsal V
Comments- The rib and manual ungual may not belong to the same individual as the hindlimb. Pittman et al. noted the distally appressed metatarsals are similar to non-eudromaeosaurs, but also that the more constricted pedal phalanx II-2 is similar to some eudromaeosaurs.
Reference- Pittman, Pei and Xu, 2015. The first dromaeosaurid (Dinosauria: Theropoda) from the Early Cretaceous Bayan Gobi Formation of Nei Mongol, China. PeerJ PrePrints. https://dx.doi.org/10.7287/peerj.preprints.1340v1

undescribed microraptorian (Lamanna, Li, Harris, Atterholt and You, 2010)
Early Aptian, Early Cretaceous
Xiagou Formation, Gansu, China
Material- (GSGM coll?) pectoral girdle, humerus, radius, ulna
Comments- Lamanna et al. (2010) referred this to Microraptoria based on the supracoracoid fenestra.
References- Lamanna, Li, Harris, Atterholt and You, 2010. First non-avian dinosaur from the Lower Cretaceous (Aptian) Xiagou Formation of the Changma Basin, northwestern China. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 119A.
Zhou, Lamanna, Poust, Li, You and O'Connor, in review. First non-avian theropod (Dromaeosauridae, Microraptorinae) from the bird-bearing Lower Cretaceous Xiagou Formation of the Changma Basin, Gansu Province, northwestern China. Journal of Vertebrate Paleontology.

undescribed Microraptoria
Jehol Group?, Early Cretaceous
Liaoning?, China
Material- (private coll.) fragmentary skull, fragmentary mandibles, cervical vertebrae, dorsal vertebrae, dorsal ribs, sacrum, caudal vertebrae, chevrons, humeri, radius, ulna, metacarpal I, phalanx I-1, metacarpal II, metacarpal III, ilium, pubes, ischia, femora, tibiotarsi, metatarsi, pedal digit II, pedal digit III, pedal digit IV, metatarsal V (http://www.thenaturalcanvas.com/Dinosaurs/pages/4523.html)
(private coll.) skull, cervical series, dorsal series, dorsal ribs, sacrum, caudal series, chevrons, scapulocoracoid, furcula, humeri, radii, ulnae, carpus, metacarpal I, phalanx I-1, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx III-3, manual ungual III, pubis, ischia, femora, tibiotarsi, proximal fibula, metatarsus, pedal digit II, pedal digit III, pedal ungual III, pedal digit IV, pedal ungal IV
(IVPP, D and Tianyu coll.) (over 300 individuals) (Alexander et al., 2010)
(Wenya Museum DNO: 087) (640 mm) incomplete skull, dorsal ribs, caudal series, incomplete humeri, partial radius, incomplete ulnae, metacarpal I, phalanx I-1, manual ungual I, partial manus, pelvic elements, femora, tibiotarsi, metatarsi, pedal ungual II, pedal digit III, pedal ungual III, pedal digit IV, pedal ungual IV, contaur feathers
(Wenya Museum DNO: 090) composite including supposed humerus, radius, ulna, partial manus, femora, tibiae
microraptorian- partial mandible, cervical series, dorsal vertebrae, dorsal ribs, uncinate processes, sacrum, caudal series, radii, ulnae, semilunate carpal, metacarpal I, phalanx I-1, metacarpal II, phalanx II-1, phalanx II-2, metacarpal III
confuciusornithid- humerus, pubes, metatarsi, pedal digit I, pedal ungual I, pedal digit II, pedal ungual II, pedal digit III, pedal ungual III, pedal digit IV, pedal ungual IV
Comments- There are many undescribed microraptorian specimens photographed online, most for sale. These are usually advertised as Microraptor, but may also belong to Sinornithosaurus. They are all probably from the Yixian or perhaps Juifotang Formations of Liaoning. Many are altered or composites. Alexander et al. (2010) reported seeing over three hundred microraptorian specimens at the IVPP, Dalian Natural History Museum and Shandong Tianyu Museum of Natural History.
A specimen on sale at The Natural Canvas is labeled as Microraptor gui. It has a disarticulated skull and its precise affinities cannot be determined without better photographs. Another specimen is preserved spread in dorsal view and seems basically complete besides one distal pes. Its affinities cannot be determined yet either.
Three dromaeosaurid specimens were exhibited at the China Jehol Biota Fossil Exhibition in 2002 (Creisler, DML 2002) from the personal collection of Du Wenya, who runs a museum in Jinzhou. These were available at least in 2000 (Hutchinson, DML 2000). DNO: 087 is fairly complete but very poorly preserved. The specimen is obviously altered, as the tibiotarsi and femora are switched (giving the illusion of long femora), while one pes is far too short. The pelvic elements are mixed up as well, with possible proximal pubes located anteriorly, and two other elements located posteriorly.
DNO: 090 is a composite specimen, as shown by the obviously confuciusornithid pes (Hutchinson, DML 2000). One pes is attached to a broken tibia(?), while the other is placed at the end of what may be another bird tarsmetatarsus. Both are far too short compared to the femora, one of which is much longer and apparently constructed of two limb bones. The pubis appears too slender and pointed for a dromaeosaurid, resembling those of avialans more. One humerus appears to have a proximal foramen like Confuciusornis and Sapeornis. One of the lower arms and manus appears genuinely microraptorian, though the other lower arm is much too short with a far too robust "radius". The other possible manus is too poorly preserved to evaluate. The dentary and tail both appear to be dromaeosaurid.
References- Hutchinson, DML 2000. https://web.archive.org/web/20180115070226/http://dml.cmnh.org/2000Jun/msg00268.html
Creisler, DML 2002. https://web.archive.org/web/20180115070231/http://dml.cmnh.org/2002Dec/msg00303.html
Alexander, Gong, Martin, Burnham and Falk, 2010. Model tests of gliding with different hindwing configurations in the four-winged dromaeosaurid Microraptor gui. Proceedings of the National Academy of Sciences. 107(7), 2972-2976.

Graciliraptor Xu and Wang, 2004
= "Graciliraptor" Xu, 2002
G. lujiatunensis Xu and Wang, 2004
= "Graciliraptor lujiatunensis" Xu, 2002
= Sinornithosaurus lujiatunensis (Xu and Wang, 2004) Paul, 2010
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V13474) (~1 m; 3.3 kg; adult) fragmentary maxilla, eight maxillary teeth, mid caudal vertebra (12.5 mm), nine distal caudal vertebrae (30 mm), chevrons, incomplete humerus (~106 mm), radii (~87 mm; one distal), ulnae (91 mm; one distal), scapholunare, semilunate carpals, distal carpal III, metacarpals I (17, 17 mm), phalanges I-1 (35, 35 mm), manual unguals I (32.5, 33.5 mm), metacarpals II (53.5, 54 mm), phalanges II-1 (25, ~24.5 mm), phalanx II-2 (26.5 mm), proximal manual ungual II, metacarpals III (48.8, 49.4 mm), phalanges III-1 (~13.5, ~12 mm), phalanges III-2 (6.4, 6.2 mm), phalanx III-3 (17.5 mm), manual ungual III, incomplete tibiotarsi (~172 mm), distal fibula (~167 mm), distal metatarsals II, phalanx II-1 (12 mm), phalanges II-2 (13, 12 mm), proximal pedal ungual II, distal metatarsal III, phalanx III-1 (21.8, 22 mm), phalanges III-2 (14, 13 mm), distal metatarsals IV, phalanges IV-1 (17, ~17 mm), phalanx IV-2 (10.8 mm)
Diagnosis- (after Xu and Wang, 2004) astragalar medial condyle significantly expanded posteriorly; metatarsal II 1.5 times wider than metatarsal III distally.
Other diagnoses- Turner et al. (2012) note several characters listed in the original diagnosis are problematic. The elongate caudals, slender tibiotarsus and pedal phalanx III-1 are shared with Microraptor, the postzygapophyseal laminae on distal caudals and rectangular proximal section of the tibia are common in dromaeosaurids, the proximal end of metacarpal III is not very dorsoventrally expanded, and manual ungual I is not necessarily small compared to ungual II due to the latter being crushed.
Comments- Discovered in 2001, this was initially described in Xu's (2002) thesis, then published in 2004. Turner et al. (2012) consider it a possible synonym of Microraptor zhaoianus, though it falls outside the Sinornithosaurus+Microraptor clade in Foth et al. (2014) and Lee et al. (2014).
References- Xu, 2002. Deinonychosaurian fossils from the Jehol Group of Western Liaoning and the coelurosaurian evolution. PhD Thesis. Chinese Academy of Sciences. 325 pp.
Xu and Wang, 2004. A new dromaeosaur (Dinosauria: Theropoda) from the Early Cretaceous Yixian Formation of Western Liaoning. Vertebrata PalAsiatica. 42(2), 111-119.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.

Changyuraptor Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014
C. yangi Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014
= Sinornithosaurus yangi (Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014) Paul, 2016
Barremian-Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Holotype- (HG B016) (~1.32 m; 7.62 kg; five year old adult) fragmentary skull, fragmentary mandibles, cervical vertebrae, dorsal vertebrae, dorsal ribs, uncinate processes, gastralia, twenty caudal vertebrae, chevrons, scapula, fragmentary coracoid, partial furcula, sternum, sternal ribs, humeri (one distal; 148.9 mm), radii, ulnae (126.1 mm), semilunate carpal, metacarpals I (~21.3 mm), phalanges I-1 (~44.6 mm), manual ungual I, metacarpal II (~75.6 mm), phalanx II-1, phalanges II-2, manual unguals II, metacarpal III (73.2 mm), phalanx III-1 (~20.7 mm), phalanx III-2 (~11.5 mm), phalanx III-3 (~20.4 mm), manual unguals III, manual claw sheaths, pubes (117.1 mm), partial ischium, femora (153 mm), tibiae, fibula, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II (32.5 mm), metatarsals III (111.2 mm), pedal phalanges III-1 (21.8 mm), pedal phalanges III-2 (15.3 mm), pedal phalanges III-3 (13.9 mm), pedal unguals III (25.7 mm), metatarsals IV (107.4 mm), metatarsals IV, pedal phalanges IV-1, pedal phalanges IV-2, pedal phalanges IV-3, pedal phalanges IV-4, pedal unguals IV (23.1 mm), pedal claw sheaths, metatarsals V, scales, contour feathers, fragmentary remiges, leg remiges, retrices
Diagnosis- (after Han et al., 2014) furcula more robust than Sinornithosaurus and much larger than Tianyuraptor; forelimb much longer when compared to hindlimb; humerus much longer (>20%) than ulna; metacarpal I shorter than Sinornithosaurus (1/4-1/5 versus 1/3); semilunate carpal covering all of proximal metacarpals I and II; manual ungual II largest; ischium shorter than Microraptor; midshaft of metatarsal IV significantly broader than metatarsal II or III; mid caudals roughly twice length of dorsals unlike Microraptor; fewer (22) caudal vertebrae than Microraptor and Tianyuraptor; rectrices significantly longer than other microraptorines.
References- Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014. A new raptorial dinosaur with exceptionally long feathering provides insights into dromaeosaurid flight performance. Nature Communications. 5, 4382.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press. 360 pp.

Sinornithosaurus Xu, Wang and Wu, 1999
= "Sinavisaurus" Skrepnick, DML 2000
= "Jeholraptor" Paul, 2024
S. millenii Xu, Wang and Wu, 1999
= Sinornithosaurus haoiana Liu, Ji, Tang and Gao, 2004
= "Jeholraptor" haoiana (Liu, Ji, Tang and Gao, 2004) Paul, 2024
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Member of the Yixian Formation, Liaoning, China
Holotype- (IVPP V12811) (~1.05 m, 5.00 kg) skull (152 mm), scleral plates, mandibles (135 mm), hyoid (53 mm), atlantal intercentrum, atlantal neurapophyses, anterior cervical vertebra, three mid cervical vertebrae, three posterior cervical vertebrae, tenth cervical vertebra, two cervical ribs, eleven dorsal vertebrae (13 mm), presacral vertebra, fourteen dorsal ribs, gastralia, sacrum (65 mm), first caudal vertebra (9.5 mm), second caudal vertebra, third caudal vertebra, fourth caudal vertebra, fifth caudal vertebra (15 mm), sixth caudal vertebra (20 mm), seventh caudal vertebra (28 mm), eighth caudal vertebra, ninth caudal vertebra, chevrons, scapulae (85, ~85 mm), coracoids (42 mm), furcula, sternal plates (84 mm), four sternal ribs, humeri (133, 134 mm), radius (~106 mm), ulna (110 mm), semilunate carpal, distal carpal III, metacarpal I (20 mm), phalanx I-1 (41 mm), manual ungual I (27 mm), metacarpal II (63 mm), phalanx II-1 (32 mm), phalanx II-2 (35 mm), manual ungual II (26 mm), metacarpal III (60 mm), phalanx III-1 (18 mm), phalanx III-2 (9 mm), phalanx III-3 (23 mm), manual ungual III (19 mm), ilia (85 mm), pubes (116 mm), ischia (52 mm), incomplete femora (~146 mm), distal tibia, partial fibula, astragalocalcaneum, distal tarsal III, distal tarsal IV, metatarsal I (21 mm), phalanx I-1 (11 mm), pedal ungual I (11 mm), metatarsal II (88 mm), phalanx II-1 (15 mm), phalanx II-2 (17 mm), partial pedal ungual II (35 mm), metatarsals III (93, 93 mm), phalanx III-1 (25 mm), phalanx III-2 (17.5 mm), phalanx III-3 (16.5 mm), pedal ungual III (20 mm), metatarsal IV (91 mm), phalanx IV-1 (20 mm), phalanx IV-2 (14 mm), phalanx IV-3 (10 mm), phalanx IV-4 (12 mm), pedal ungual IV (18.5 mm), metatarsal V (47 mm), feathers, keratin ungual sheaths (m2, p4)
Referred- (D2140; holotype of Sinornithosaurus haoiana) (2 year old subadult) (skull ~125 mm) premaxilla, maxilla, nasals, lacrimal, frontals, parietal, partial jugal, postorbital, squamosal, quadratojugal, quadrate, vomer, palatine, pterygoid, parasphenoid-basisphenoid, sclerotic plates, dentaries, splenial, surangular, angulars, hyoid, teeth, eight or nine cervical vertebrae, three cervical ribs, eleven dorsal vertebrae (13 mm), dorsal ribs, five uncinate processes (to 29 mm), gastralia, sacral vertebrae, five proximal caudal vertebrae, several mid caudal vertebrae and chevrons, scapulacoracoid (scapula 84.4 mm), furcula, sternal plates (64.1 mm), four sternal ribs (to 42.6 mm), humeri (129.5 mm), radii, ulnae (105.2 mm), scapholunare, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II (59 mm), metacarpal III (54.1 mm), phalanx III-1, phalanx III-2, ilia (84.5, 86.1 mm), pubes (116.9 mm), ischium, incomplete femur, incomplete tibia, incomplete fibula, astragalocalcaneum, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II (45.2 mm on curve), metatarsal III (86.8 mm), phalanx III-1 (23.3 mm), phalanx III-2 (14.5 mm), phalanx III-3 (12.8 mm), pedal ungual III (~21 mm), metatarsal IV (84.2 mm), pedal digit IV, pedal ungual IV, metatarsal V, feathers (Liu et al., 2004)
(IG-3) skull, postcranial skeleton (Azuma, 2005)
(IG-5b) skull postcranial skeleton, feathers (Azuma, 2005)
(IVPP V16904) (adult) specimen including skull, mandibles, hyoid, cervical vertebrae, dorsal vertebra, dorsal ribs, scapulae, coracoids, manual phalanx, manual ungual and tibiotarsus (Turner et al., 2012)
(JMP-V-05-8-01; inside holotype of Sinocalliopteryx gigas) tibia (155 mm), partial fibula, phalanx I-1, pedal ungual I, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, feathers (Ji et al., 2007)
(Tianyu Museum of Natural History coll.) multiple specimens including cranial material (Gong et al., 2010)
?(Wenya Museum DNO: 088) skull, dorsal vertebrae, sacrum, caudal series, furcula, humeri, radius, ulnae, manual phalanges, manual unguals, pubes, femora, tibiotarsi, metatarsi, pedal digit III, pedal digit IV (Kaiser, 2007)
?(private coll.) skull, mandibles, hyoid, cervical vertebrae, dorsal vertebrae, gastralia, caudal series, scapulocoracoid, partial furcula, sternum, sternal ribs, humeri, radii, ulnae, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-3, manual ungual III, ilium, pubes, femora, tibiotarsi, metatarsal II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, pedal ungual IV, pedal phalanges, metatarsal V
(private coll.) incomplete skull, mandible, hyoid, cervical series, dorsal vertebrae, dorsal ribs, gastralia, sacrum, caudal series, scapulocoracoid, sternal plate, humeri, radius, ulna, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, ilia, pubes, ischium, femora, tibiotarsi, metatarsi, pedal digit II, pedal digit III, pedal digit IV, unidentified elements
? skull, mandible, cervical vertebrae, dorsal series, dorsal ribs, gastralia, caudal series, coracoids, furcula, humeri, radii, ulnae, manual elements, ilium, pubis, ischium, femora, tibiae, metatarsi, pedal digit II, pedal ungual II, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
specimen including skull, mandible (Zheng, 2009)
Diagnosis- (from Xu et al., 1999) elongate posterolateral parietal process (unknown in other derived microraptorines).
(after Xu and Wu, 2001) groove posterior to the anterior carina on the lingual surface of the premaxillary tooth crowns (unknown in other derived microraptorines); column-like margin of the pterygoid process of the quadrate (unknown in other derived microraptorines); large excavation on the posterolateral surface of the cultriform process (unknown in other derived microraptorines).
(after Turner et al., 2012) thickened posterior rim of promaxillary fenestra (unknown in other derived microraptorines).
(proposed) larger humerus than Microraptor or Bambiraptor (humerofemoral ratio >88%) (also in Wulong).
Other diagnoses- Xu et al. (1999) listed additional synapomorphies of Sinornithosaurus. However, the pitted antorbital fossa, elongate posterodorsal dentary process, enlarged supracoracoid fenestra, lateral tubercle on the pubic midshaft, and proximodorsal ischial process are also present in Microraptor (and the pubic character in Hesperonychus); unserrated premaxillary teeth and a subarctometatarsus are near certainly plesiomorphic for dromaeosaurids and found in other microraptorians; the manual phalanx III-1/III-2 ratio of S. millenii falls between those of Microraptor specimens.
Contra Xu and Wu (2001), the large promaxillary fenestra is now known to be present in Microraptor as well. The deep excavation on the posteroventral margin of the premaxilla is also present in Bambiraptor. Turner et al. (2012) note the supposed diastema between premaxillary and maxillary teeth is actually the ventral edge of the subnarial process.
Comments- Sinornithosaurus' holotype was discovered in 1998. It was named and briefly described by Xu et al. (1999), and described in depth in Xu's (2002) thesis, though only the cranial (Xu and Wu, 2001) and pedal (Wu and Wang, 2000) portions have been published. Skrepnick (DML, 2000) noted that Sinornithosaurus had the working name of "Sinavisaurus" until right before it was published. The species name must be emended to millennii because it's a mispelling of a Latin word (millennium) (ICZN Art. 32.5).
IVPP V16904 was first published as IVPP uncatalogd by Turner et al. (2012), but Xu et al. (2015) reveal its specimen number. Xu et al. do not believe it to be conspecific with S. millenii because none of its teeth are serrated, and its neurocentral sutures are fully closed and a tibiotarsus is developed despite it being "significantly smaller" than the holotype which lacks these fusions. While possibly correct pending further description, it should be noted the referred specimen of Mei long has a tibiotarsus and fused presacral neurocentral sutures but is 80% the size of the holotype that doesn't, the Sinovenator changii paratype has a tibiotarsus but is 89% the size of the holotype that doesn't, and Microraptor zhaoianus specimens can have serrations on both carinae (NGMC 00-12-A), only distal serrations (holotype) or no serrations (IVPP V13320). Thus such variations are known in other basal paravian species, so are expected in S. millenii individuals.
Ji et al. (2002) referred NGMC 91 to Sinornithosaurus sp., and it has been referred to S. millenii by other authors such as Senter (2011) and Turner et al. (2012), but it is here referred to a new genus more closely related to Microraptor. Xu (2002) referred IVPP V13477 to Sinornithosaurus sp., but Xu et al. (2003) later made it a paratype of Microraptor gui. Ji (2002) described NGMC 00-12-A as Sinornithosaurus sp., but it is here referred to Microraptor.
Gong et al. (2010) argued Sinornithosaurus was venomous, but Gianechini et al. (2011) show the paper is highly flawed and its conclusions lack merit.
Several undescribed specimens are photographed online, and appear to be Sinornithosaurus. They are probably all from the Yixian or perhaps Juifotang Formations of Liaoning, China.
One is complete and largely articulated except for the pelvis and hindlimbs. It is being sold by Geosciences Enterprises as Microraptor zhaoianus. The nasal is disarticulated, making the skull appear more slender and troodontid-like, but the proportions all match Sinornithosaurus. The elongate manual phalanx III-3 distinguishes it from Microraptor, while the short ulna and long manual digit II distinguish it from Graciliraptor.
Another is quite disarticulated, but well preserved. The elongate tibiotarsus suggests it is Sinornithosaurus.
A further specimen is almost complete and well preserved, except both manus are fragmentaty and one distal tibia is missing. The pes is positioned directly contacting the broken end, suggesting some alteration. The skull is almost complete and looks particularily elongate.
Three dromaeosaurid specimens were exhibited at the China Jehol Biota Fossil Exhibition in 2002 (Creisler, DML 2002) from the personal collection of Du Wenya, who runs a museum in Jinzhou. These were available at least in 2000 (Hutchinson, DML 2000). DNO: 088 is possibly the specimen illustrated by Kaiser (2007) as an unidentified confuciusornithid. It is obviously dromaeosaurid however, with an elongate tail, slender metatarsus, and slender furcula. The elongate tibia suggests it may be Sinornithosaurus.
Gong et al. (2010) mention specimens from the Tianyu Museum, and later (2011) illustrate a skull from Zheng (2009).
Sinornithosaurus haoiana- Liu et al. (2004) described D2140 as a new species of Sinornithosaurus, S. haoiana. This was based on several proposed differences from S. millenii, but Turner et al. have proposed the two species are synonymous. The long subnarial process of the premaxilla which excludes the maxilla from the naris cannot be evaluated in S. millenii. They further note the maxillary fenestra of the millenii holotype is heavily damaged, so the small and circular morphology of haoiana cannot be compared. Turner et al. find the dorsal quadratojugal process of millenii to be longer than thought by Xu, which can indeed be seen in photos. They state both holotypes have low premaxillae, and yet do not argue against Liu et al.'s correct observation millenii's are much lower. Similarly, they argue all specimens have low dentaries, but do not argue against Liu et al.'s correct observation millenii's are much lower. Turner et al. are wrong when they claim millenii has a pubic peduncle narrower than its acetabulum. I would use these three differences to retain them as separate species if not for IVPP V16904 having both a tall premaxilla and a low dentary (fig. 23C in Turner et al.), which ironically is not used in Turner et al.'s argument. Based on this specimen, I agree the species are synonymous. The species name must be emended to haoianus, to match the gender of the genus name (ICZN Art. 31.2, 34.2). Han et al. (2014) provided more measurements. Poust et al. (2020) examined the histology, finding it to be more mature than the Wulong holotype. Paul (2024) uses Jeholraptor haoiana for the taxon, stating "Usual placement in earlier S. millenii incorrect because of very different quadratojugals and other reasons. L-shaped quadratojugals in this and Wulong suggest they are more closely related to one another than to other microraptorines and probably form their own subgroup" and "Smaller furcula, sternum, and less gracile hand suggest was not as good a flier as Sinornithosaurus." It's a nomen nudum based on ICZN Article 16.1 at least- "Every new name published after 1999, including new replacement names (nomina nova), must be explicitly indicated as intentionally new." I would dispute that haoiana has an L-shaped quadratojugal, it's just that millenii has a shorter dorsal process. And even though Wulong's skull is illustrated that way in its description, the photo suggests that whole corner of the bone is unpreserved. Purported pectoral and forelimb differences deserve more scrutiny, but (for instance) one sternal plate contacts a humerus and could project below it to increase its length past what the text states. It should also be noted that Paul synonymizes Graciliraptor, Microraptor, Cryptovolans and Changyuraptor under Sinornithosaurus, so haoiana being more distinct than these might be especially questionable.
References- Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
Hutchinson, DML 2000. https://web.archive.org/web/20180115070226/http://dml.cmnh.org/2000Jun/msg00268.html
Skrepnick, DML 2000. https://web.archive.org/web/20210605025348/http://dml.cmnh.org/2000Oct/msg00074.html
Xu and Wang, 2000. Troodontid-like pes in the dromaeosaurid Sinornithosaurus. Paleontological Society of Korea Special Publication. 4, 179-188.
Xu and Wu, 2001. Cranial morphology of Sinornithosaurus millenii Xu et al. 1999 (Dinosauria: Theropoda: Dromaeosauridae) from the Yixian Formation of Liaoning, China. Canadian Journal of Earth Sciences. 38, 1739–1752.
Xu, Zhou and Prum, 2001. Branched integumental structures in Sinornithosaurus and the origin of feathers. Nature. 410, 200-204.
Creisler, DML 2002. https://web.archive.org/web/20180115070231/http://dml.cmnh.org/2002Dec/msg00303.html
Ji, 2002. New data of Early Cretaceous dromaeosaurs from western Liaoning with comments on the origin of feathers. Unpublished Thesis. 94 pp.
Ji, Ji, Yuan and Ji, 2002. Restudy on a small dromaeosaurid dinosaur with feathers over its entire body. Earth Science Frontiers. 9(3), 57-63.
Xu, 2002. Deinonychosaurian fossils from the Jehol Group of Western Liaoning and the coelurosaurian evolution. PhD Thesis. Chinese Academy of Sciences. 325 pp.
Xu, Zhou, Wang, Kuang, Zhang and Du, 2003. Four-winged dinosaurs from China. Nature. 421, 335-340.
Liu, Ji, Tang and Gao, 2004. A new species of dromaeosaurids from the Yixian Formation of western Liaoning. Geological Bulletin of China. 23(8), 778-783.
Azuma, 2005. The Flying Dinosaurs: Fukui Prefectural Dinosaur Museum. 118 pp.
Ji, Ji, Lu and Yuan, 2007. A new giant compsognathid dinosaur with long filamentous integuments from Lower Cretaceous of Northeastern China. Acta Geologica Sinica. 81(1), 8-15.
Kaiser, 2007. The Inner Bird: Anatomy and Evolution. UBC Press. 386 pp.
White, 2009. The subarctometatarsus: intermediate metatarsus architecture demonstrating the evolution of the arctometatarsus and advanced agility in theropod dinosaurs. Alcheringa. 33(1), 1-21.
Zheng, 2009. Niaolei Qiyuan. Jinan City, China: Shandong Science and Technology Publishing House Press.
Gong, Martin, Burnham and Falk, 2010. The birdlike raptor Sinornithosaurus was venomous. Proceedings of the National Academy of Sciences. 107, 766-768.
Zhang, Kearns, Orr, Benton, Zhou, Johnson, Xu and Wang, 2010. Fossilized melanosomes and the colour of Cretaceous dinosaurs and birds. Nature. 463, 1075-1078.
Gianechini, Agnolin and Ezcurra, 2011. A reassessment of the purported venom delivery system of the bird-like raptor Sinornithosaurus. Paläontologische Zeitschrift. 85, 103-107.
Gong, Martin, Burnham and Falk, 2011. Evidence for a venomous Sinornithosaurus. Paläontologische Zeitschrift. 85, 109-111.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24, 2197-2216.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Xing, Bell, Persons, Ji, Miyashita, Burns, Ji and Currie, 2012. Abdominal contents from two large Early Cretaceous compsognathids (Dinosauria: Theropoda) demonstrate feeding on confuciusornithids and dromaeosaurids. PLoS ONE. 7(8), e44012.
Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014. A new raptorial dinosaur with exceptionally long feathering provides insights into dromaeosaurid flight performance. Nature Communications. 5, 4382.
Xu, Pittman, Sullivan, Choiniere, Tan, Clark, Norell and Wang, 2015. The taxonomic status of the Late Cretaceous dromaeosaurid Linheraptor exquisitus and its implications for dromaeosaurid systematics. Vertebrata PalAsiatica. 53(1), 29-62.
Poust, Gao, Varricchio, Wu and Zhang, 2020. A new microraptorine theropod from the Jehol biota and growth in early dromaeosaurids. The Anatomical Record. Early View DOI: 10.1002/ar.24343
Paul, 2024. The Princeton Field Guide to Dinosaurs 3rd edition. Princeton University Press. 384 pp.

unnamed microraptorian (Ji, Norell, Gao, Ji and Ren, 2001)
Late Valanginian-Early Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Material- ?(NGMC 91; Dave) (69 cm) skull (~100 mm), sclerotic rings, mandibles, ten cervical vertebrae, cervical ribs, dorsal series, dorsal ribs, uncinate processes, gastralia, caudal series, chevrons, scapulae (~53 mm), coracoids, furcula, sternal plate, sternal ribs, humeri (87 mm), ulnae (72 mm), radii, scapholunare, semilunate carpal, metacarpal I (13 mm), phalanx I-1 (27 mm), manual ungual I (15 mm), metacarpal II (40 mm), phalanx II-1 (22 mm), phalanx II-2 (24 mm), manual ungual II (16 mm), metacarpal III (40 mm), phalanx III-1 (12 mm), phalanx III-2 (7 mm), phalanx III-3 (16 mm), manual ungual III (11 mm), pubis(?), ischium(?), femora (95 mm), tibiae (133 mm), fibulae, astragalus, distal phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1 (10 mm), phalanx II-2 (10.2mm), pedal ungual II, metatarsal III (62 mm), phalanx III-1 (16.2 mm), phalanx III-2 (9.6 mm), phalanx III-3 (10.3 mm), pedal ungual III, metatarsal IV, phalanx IV-1 (7.1 mm), phalanx IV-2 (6 mm), phalanx IV-3 (6.9 mm), phalanx IV-4, pedal ungual IV, metatarsal V, scales, contour feathers, remiges, retrices, ungual sheaths
Comments- NGMC 91 was first reported by Ji et al. (2001) as Dromaeosauridae indet., though they noted strong resemblence to Sinornithosaurus. Ji et al. noted the possibility that some differences, such as the bowed first metacarpal, were due to age. Ji et al. (2002) later assigned NGMC 91 to Sinornithosaurus sp., with the same reservations regarding whether differences from the S. millenii holotype were taxonomic or ontogenetic. Czerkas et al. (2002) tentatively referred NGMC 91 to their new genus Cryptovolans (= Microraptor), though without stated reasons besides "nearly identical wing proportions". In fact, NGMC 91 lacks the proposed apomorphies of Cryptovolans (28-30 caudal vertebrae; manual phalanx III-1 longer than III-3), and several apomorphies of Microraptor- posterior dentary teeth with constricted bases; manual phalanx III-1 >83% of phalanx III-3 in length; manual phalanx I-1 <62% of metacarpal II in length; tibiofemoral ratio <136%; highly elongate leg remiges (>1.5 times femoral length). Senter et al. (2004) found that NGMC 91 claded with Microraptor based on- mesial serrations absent on posterior teeth; dentary with midlength increase in height. Senter has since (2007) stated the dentary character's "apparent presence was an optical illusion caused by differing heights of preserved dentary teeth and the curvature of the dentary." He still found the Microraptor+NGMC 91 clade however, supported by a proximodorsal lip on manual ungual I and strongly curved pedal unguals III and IV. Senter (2011) has since combined NGMC 91 into his Sinornithosaurus OTU without justification, and Turner et al. (2012) also placed the specimen in Sinornithosaurus millenii. This was because they claimed it differed from Microraptor in lacking mid caudal vertebrae 3-4 times length of anterior dorsal vertebrae, but this is untrue. Further, the supposed Sinornithosaurus apomorphy of a posteriorly bifurcated dentary is also present in Microraptor. It is retained as separate here.
References- Ji, Norell, Gao, Ji and Ren, 2001. The distribution of integumentary structures in a feathered dinosaur. Nature. 410 (6832), 1084-1088.
Czerkas, Zhang, Li and Li, 2002. Flying dromaeosaurs. in Czerkas (ed.). Feathered Dinosaurs and the Origin of Flight. 97-126.
Ji, Ji, Yuan and Ji, 2002. Restudy on a small dromaeosaurid dinosaur with feathers over its entire body. Earth Science Frontiers, 9(3), 57-63.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8, 1-20.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24, 2197-2216.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.

Microraptorinae sensu Martyniuk, 2012
Definition- (Microraptor zhaoianus <- Sinornithosaurus millenii, Dromaeosaurus albertensis)

Wulong Poust, Gao, Varricchio, Wu and Zhang, 2020
= "Wulong" Poust, 2014
W. bohaiensis Poust, Gao, Varricchio, Wu and Zhang, 2020
= "Wulong bohaiensis" Poust, 2014
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (D2933) (1 year old juvenile) skull (98.6 mm), sclerotic plates, mandibles, ten cervical vertebrae, thirteen dorsal vertebrae, ~ten-thirteen dorsal ribs, gastralia, three or four sacral centra, sacral neural arches and ribs, thirty or thirty-one caudal vertebrae (distal caudals ~19 mm), chevrons, scapulae (44.8 mm), coracoids, furcula, sternals, four sternal ribs, humeri (75.4, 76.6 mm), radii (61.4 mm), ulnae (63.4, 64.5 mm), semilunate carpals, distal carpal III, metacarpals I (12.3, 12.3 mm), phalanges I-1 (~25 mm), manual unguals I (15.3, 14.8 mm), metacarpals II (39.1, 41.3 mm), phalanges II-1 (20, 19 mm), phalanges II-2 (22.7, 22.7 mm), manual unguals II (15.1, 15.3 mm), metacarpals III (~39, ~36 mm), phalanges III-1 (12.8, 13.1 mm), phalanges III-2 (8.3, 7.5 mm), phalanges III-3 (14.2 mm), manual unguals III (10.1, 9.9 mm), manual claw sheaths, ilium (49.5 mm), pubes (62.4, 68.4 mm), ischia (26.9, 27.5 mm), femora (85.8, 86.5 mm), tibiae (117 mm), fibulae (118.7 mm), astragali, calcanea, metatarsal I, phalanx I-1 (6.5 mm), pedal ungual I (5.2 mm), metatarsals II (53.5, 54 mm), phalanges II-1 (one partial; 9 mm), phalanges II-2 (one partial; 9.2 mm), pedal unguals II (16.4, 16.1 mm), metatarsals III (56.5, 56.9 mm), phalanges III-1 (14.3 mm), phalanges III-2 (10 mm), phalanges III-3 (9.3 mm), pedal unguals III (11.3 mm), metatarsals IV (56.7, 53.5 mm), phalanges IV-1 (10.7, 11.1 mm), phalanges IV-2 (7.6, 8.5 mm), phalanges IV-3 (5.9, 6.5 mm), phalanges IV-4 (6.1, 6.7 mm), pedal unguals IV (10.1 mm), metatarsals V, pedal claw sheaths, body feathers, retrices, remiges, leg remiges
Diagnosis- (after Poust et al., 2020) long anterior process of quadratojugal; anteriorly inclined pleurocoels on anterior half of dorsal centra; transverse processes of proximal caudals significantly longer than width of centrum; 30 caudal vertebrae; distally located and large distodorsal ischial process; large supracoracoid fenestra (>15% of total area).
Comments- This specimen was transferred from a private collector to the Dalian Natural History Museum prior to April 2014 (based on SVP abstract deadlines). Poust (2014) first described and named Wulong in his thesis, with the disclaimer "This thesis is not intended to constitute publication of new names or nomenclatural acts and such names as are given are not here intended for public and permanent scientific record – this is a model of what will be published later." This means the thesis falls under ICZN Article 8.2 "A work that contains a statement to the effect that it is not issued for public and permanent scientific record, or for purposes of zoological nomenclature, is not published within the meaning of the Code", and the name was a nomen nudum until Poust et al. described it officially in 2020.
Poust et al.'s measurements for manual phalanx I-1 (15.9 and 15.8 mm) and metacarpal III (17.1, 18.7 mm) are far too short, and others listed above may end up being off as well once checked. Poust et al. are incorrect in stating "the tiny first phalanges of left and right manual digit three are missing", as shown by their own description, figures and measurement table. Contra Poust et al., only the right ilium seems to be exposed, with the supposed broken dorsal boundary exposing the dorsal left ilium being a crack through the bone. The authors note "restoration efforts by technically proficient but inexpert preparators untrained in anatomy resulted in one noticeable error prior to the specimen’s arrival at the museum: the distal pedal phalanges of the right Digit II (2, 3, and the ungual) have been reconstructed incorrectly."
Poust et al. (2020) entered it into Turner's TWiG analysis and recovered it as a microraptorian sister to Sinornithosaurus, but both characters supporting this are misscored- "the lack of a proximodorsal lip on the manual unguals (153) and a laterally inclined flange along the dorsal edge of the surangular (212)." Both taxa have a proximodorsal lip on at least manual ungual II, while Wulong is too crushed to determine the presence of a surangular flange. Adding it to Hartman et al.'s TWiG analysis results in a most parsimonious position sister to Microraptor plus Zhongjianosaurus, placing all of the Jiufotang microraptorians together.
References- Poust, 2014. Description and ontogenetic assessment of a new Jehol microraptorine. Masters thesis, Montana State University. 105 pp.
Poust, Varricchio and Gao, 2014. A new Jehol microraptorine with implications for secondary feather loss in Sinornithosaurus. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 208.
Poust, Gao, Varricchio, Wu and Zhang, 2020. A new microraptorine theropod from the Jehol biota and growth in early dromaeosaurids. The Anatomical Record. Early View DOI: 10.1002/ar.24343

Zhongjianosaurus Xu and Qin, 2017
Z. yangi Xu and Qin, 2017
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Holotype- (IVPP V22775) (adult, 310 g) seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra (4.7 mm), tenth cervical vertebra (4.4 mm), first dorsal vertebra (4.4 mm), second dorsal vertebra (4.4 mm), third dorsal vertebra (4.7 mm), fourth dorsal vertebra (4.7 mm), fifth dorsal vertebra (4.8 mm), sixth dorsal vertebra (5.0 mm), seventh dorsal vertebra (5.0 mm), thirteen dorsal ribs (d1 ~20, d4 39, d8 25 mm), three free uncinate processes, second caudal vertebra (4.6 mm), third caudal vertebra (5.1 mm), fourth caudal vertebra (5.5 mm), fifth caudal vertebra (6.3 mm), sixth caudal vertebra (9.3 mm), seventh caudal vertebra (13.3 mm), eighth caudal vertebra (14.9 mm), ninth caudal vertebra (~16.5 mm), tenth caudal vertebra, eleventh caudal vertebra, twelfth caudal vertebra, thirteenth caudal vertebra (13.2 mm), fourteenth caudal vertebra (13.1 mm), fifteenth caudal vertebra (13.1 mm), sixteenth caudal vertebra (13.1 mm), seventeenth caudal vertebra (13.2 mm), eighteenth caudal vertebra (13.1 mm), nineteenth caudal vertebra (12.7 mm), twentieth caudal vertebra (12.7 mm), twenty-first caudal vertebra (11.4 mm), twenty-second caudal vertebra (10.8 mm), twenty-third caudal vertebra (9.1 mm), twenty-fourth caudal vertebra (8.7 mm), twenty-fifth caudal vertebra (7.9 mm), twenty-sixth caudal vertebra (6.8 mm), twenty-seventh caudal vertebra (6.3 mm), chevrons, furcula, sternal plate (33 mm), five sternal ribs (r1 6.5, r5 25.5 mm), scapulocoracoids (scapula 33 mm), humeri (43 mm), radius (42 mm), ulna (44 mm), scapholunare, pisiform, carpometacarpus (mc1 8.5, mcII 27 mm), phalanx I-1 (16.3 mm), manual ungual I (12.5 mm), manual claw sheath, femora (59 mm), tibiotarsi (78 mm), fibula, tarsometatarsi (mt II 35.5, mt III 39, mt IV 37.6 mm), distal phalanx III-1, phalanx III-2 (5.5 mm), phalanx III-3 (5.6 mm), pedal ungual III (~8 mm), distal phalanx IV-2, phalanx IV-3 (3.1 mm), phalanx IV-4 (3.2 mm), pedal ungual IV (7.4 mm), pedal ungual sheaths
Diagnosis- (after Xu and Qin, 2017) long ossified uncinate processes fused to dorsal ribs; widely arched furcula with slender and posteriorly curved rami; humeral proximal end strongly offset medially from humeral shaft; humeral internal tuberosity short; large fenestra in humeral deltopectoral crest; humeral ulnar condyle hypertrophied; ulna slightly longer than humerus; ulnar olecranon process with posterior margin mediolaterally pinched; ulnar distal end bending anteriorly and strongly expanded laterally; proximal end of metacarpal I with strong ventrolateral extension; manual ungual I without proximodorsal lip and lacking strong dorsal arching; metacarpal II laterally bowed with longitudinal ventral groove; femoral head stout and lower than trochanteric crest; medial condyle of tibiotarsus distal end with prominent distal extension; arctometatarsalian pes; metatarsal II without ginglymus on distal end.
Comments- While various websites including Wikipedia state the specimen was first reported in 2009, this is not substantiated by any sourced account and is unlikely as specimens from the Sihedang locality were not described until 2012, with no non-bird taxa reported until 2014. Xu and Qin (2017) indicate is that it must have been found prior to March 2017. An advanced online version was published April 10, but the official paginated version was published by April 26 so no separate nomen nudum entry is given here for the two week long gap. While Xu and Qin assigned Zhongjianosaurus to the Yixian Formation, Sihedang is here viewed as belonging to the Jiufotang Formation (see Iteravis entry).
Xu and Qin (2017) assigned Zhongjianosaurus to Microraptoria (their Microraptorinae) without a quantitative analysis, which was confirmed by Hartman et al. (2019) who recovered it sister to Microraptor.
References- Xu and Qin, 2017. A new tiny dromaeosaurid dinosaur from the Lower Cretaceous Jehol Group of western Liaoning and niche differentiation among the Jehol dromaeosaurids. Vertebrata PalAsiatica. 55(2), 129-144.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 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

Microraptor Xu, Zhou and Wang, 2000
= "Archaeoraptor" Sloan, 1999 in part
= Cryptovolans Czerkas, Zhang, Li and Li, 2002
= "Tetrapterornis" Miller, 2004
M. zhaoianus Xu, Zhou and Wang, 2000
= "Archaeoraptor liaoningensis" Sloan, 1999 in part
= Cryptovolans pauli Czerkas, Zhang, Li and Li, 2002
= Microraptor gui Xu, Zhou, Wang, Kuang, Zhang and Du, 2003
= "Tetrapterornis" gui (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003) Miller, 2004
= Sinornithosaurus "zhaoi" Zhang, 2007
= Sinornithosaurus zhaoianus (Xu, Zhou and Wang, 2000) Paul, 2010
= Microraptor hanqingi Gong, Martin, Burnham, Falk and Hou, 2012
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (IVPP V 12330; IVPP V 12444 in part; specimen of "Archaeoraptor liaoningensis" in part) (~420 mm; 200 g; adult) partial skull (45 mm), mandibles (42 mm), four dorsal vertebrae, thirteenth dorsal central fragment, several dorsal ribs, sternal ribs or uncinate processes, gastralia, sacrum (19 mm), (caudal series 232 mm) about twenty-five caudal vertebrae, chevrons, radius (35 mm), ulna (35 mm), scapholunare, pisiform, semilunate carpal, distal carpal III, proximal metacarpal I, proximal metacarpal II, proximal metacarpal III, partial ilia, incomplete pubes, ischia (17.9 mm), femora (one fragmentary) (53 mm), tibiae (one partial) (68 mm), fibula, tarsus, phalanx I-1 (4.4 mm), pedal ungual I, metatarsal II (32.4 mm), phalanx II-1 (5.4 mm), phalanx II-2 (5.2 mm), pedal ungual II, metatarsal III (33 mm), phalanx III-1 (6.6 mm), phalanx III-2, phalanx III-3 (5 mm), pedal ungual III, metatarsal IV (33.2 mm), phalanx IV-1, phalanx IV-2 (4.8 mm), phalanx IV-3 (3.1 mm), phalanx IV-4 (3.8 mm), pedal ungual IV, metatarsal V, pedal ungual sheaths, contour feathers, fragmentary remiges, therian ?cranial elements, ?axial elements, ?limb elements and pes
Referred- (BMNHC PH881) (540 mm) skull (~48.4 mm), mandibles (42.5, 44.2 mm), hyoids (~18.7 mm), ten cervical vertebrae, ~13-14 dorsal vertebrae, ten dorsal ribs, three uncinate processes, gastralia, sacrum, ~25-26 caudal vertebrae, chevrons, incomplete scapula, sternum, sternal ribs, humeri (42.5, 47.4 mm), radii (one incomplete; 41 mm), ulnae (one incomplete; 44.7 mm), metacarpal I (13 mm), phalanx I-1 (12.8 mm), manual unguals I (one fragmentary; 11.9 mm), metacarpal II (31.8 mm), phalanx II-1 (13.4 mm), phalanges II-2 (12.5 mm), manual unguals II (one partial; 12 mm), metacarpal III (~29.8 mm), phalanx III-1 (5.3 mm), phalanx III-2 (7.6 mm), phalanges III-3 (7.2 mm), manual unguals III (5, 5.3 mm), manual claw sheaths, incomplete ilia (~27.8 mm), pubes (46.2 mm), femora (51.8, 52.1 mm), tibiotarsi (70.7, 72.9 mm), fibulae, distal tarsal, metatarsal I, phalanges I-1 (4.6 mm), pedal unguals I (6 mm), metatarsals II (36.2 mm), metatarsals III (39, 38.5 mm), phalanges III-1 (10.4, 8.9 mm), phalanges III-2 (6.4, 5.4 mm), phalanges III-3 (6, ~5.2 mm), pedal unguals III (10.6, 10.5 mm), metatarsals IV (36.1, 36.1 mm), phalanges IV-1 (7.8, 8.2 mm), phalanges IV-2 (4.7, 4.8 mm), phalanges IV-3 I(4, 3.8 mm), phalanges IV-4 (4.8, 4.4 mm), pedal unguals IV (8.3, 8.6 mm), pedal claw sheaths, metatarsals V, body feathers, retrices, remiges, leg remiges (Li et al., 2012; described by Pei et al., 2014)
(CAGS02-IG-gausa-1/DM 609 in part) presacral vertebrae, furcula, humerus (70 mm), radius, ulna (63 mm), carpus, metacarpal I, phalanx I-1 (24 mm), manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, manual digit III, tibiae (106 mm), fibula, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (58 mm), metatarsal IV, pedal phalanges, other elements (Czerkas and Ji, 2002)
(IG-1) skull, postcranial skeleton (Azuma, 2005)
(IVPP V13320; paratype of Microraptor gui) (~485 mm) skull, cervical vertebrae, dorsal vertebrae, dorsal ribs, sacrum, caudal vertebrae, chevrons, humeri, radii, ulnae, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpals II, phalanges II-1, metacarpals III, phalanx III-1, phalanx III-2, ilia, pubis, femora (61 mm), tibiae, fibulae, metatarsi, both pes, body feathers, remiges (150 mm), leg remiges, retrices (Xu, 2002)
(IVPP V13351) (~645 mm) specimen including femur (81 mm), retrices and leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
(IVPP V13352; holotype of Microraptor gui) (770 mm, 1.40 kg) posterior skull, posterior mandible, eighth cervical vertebrae, cervical ribs, thirteen dorsal vertebrae, dorsal ribs, six uncinate processes, gastralia, sacrum, about twenty-six caudal vertebrae, chevrons, scapulocoracoids, furcula, sternum (47.8 mm), two sternal ribs, humeri (77 mm), radii (74.9 mm), ulnae (77.6 mm), scapholunare, pisiform, semilunate carpal, distal carpals III, metacarpal I (13.6 mm), phalanx I-1 (28.7 mm), manual ungual I, metacarpal II (48.6 mm), phalanx II-1 (24.3 mm), phalanx II-2 (25.1 mm), manual ungual II, metacarpal III (47.6 mm), phalanx III-1 (12.9 mm), phalanx III-2 (8.2 mm), phalanx III-3 (14.3 mm), manual ungual sheaths, pubis (74.5 mm), ischia (35.5 mm), femora (97.5 mm), tibiotarsi (126 mm), metatarsal II, phalanx II-1, phalanx II-2 (10.1 mm), pedal ungual II, pedal ungual II sheath, metatarsal III (65 mm), phalanx III-1 (12.8 mm), phalanx III-2 (10.3 mm), phalanx III-3, metatarsal IV, phalanx IV-1 (10.4 mm), phalanx IV-2 (7 mm), phalanx IV-3 (7.3 mm), phalanx IV-4 (7.2 mm), proximal pedal ungual IV, metatarsal V, contour feathers, retrices, remiges, leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
(IVPP V13475) (393 mm; adult) skull (~51 mm), scleral plates, mandibles (41.3 mm), ten cervical vertebrae, two posterior cervical ribs, thirteen dorsal vertebrae, dorsal ribs, uncinate processes, gastralia, sacrum, incomplete caudal series, sternal ribs, scapula (34 mm), coracoids, furcula (23.2 mm wide), humeri (42, ~42 mm), radius (34.5 mm), ulnae (37, 36 mm), metacarpals I (7, 7 mm), phalanx I-1 (14.6 mm), manual ungual I (10.8 mm), metacarpals II (~22.3, 22 mm), phalanges II-1 (12, 11.3 mm), phalanges II-2 (11.2, 11.8 mm), manual unguals II (11.2, ~11 mm), metacarpals III (~21, 21 mm), phalanx III-1 (9.8 mm), phalanx III-3 (10.2 mm), manual ungual III (7.8 mm), ilium, pubes (40 mm), ischium (18 mm), femora (49.7, 49.8 mm), tibiotarsi (66, 66 mm), fibula, distal tarsal III, distal tarsal IV, phalanx I-1 (4.6, 4.7 mm), pedal unguals I (4, 4 mm), metatarsals II (33.2, 33 mm), phalanx II-1 (5.9 mm), phalanx II-2 (5.1 mm), pedal ungual II (11.4 mm), metatarsal III (35.3 mm), phalanges III-1 (8.9, 8.8 mm), phalanges III-2 (5.5, 5.8 mm), phalanges III-3 (5, 5 mm), pedal unguals III (8.7, 9.2 mm), metatarsal IV (35 mm), phalanges IV-1 (6.1, 6.1 mm), phalanges IV-2 (4.1, 4.1 mm), phalanx IV-3 (3.1 mm), phalanx IV-4 (3.1 mm), pedal ungual IV, body feathers, remiges, leg remiges (Xu, 2002)
(IVPP V13476) (~745 mm) specimen including femur (~94 mm) and leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
(IVPP V13477) (~572 mm) skull, cervical vertebrae, dorsal vertebrae, about twenty-six caudal vertebrae, chevrons, ilium, pubis, ischium, femur (72 mm), tibiotarsus, metatarsus, phalanx II-1, phalanx II-2, pedal ungual II, pedal digit III, pedal ungual III, pedal digit IV, pedal ungual IV, metatarsal V, body feathers, leg remiges, retrices (Xu, 2002)
(IVPP V17972) skull (89.9 mm), mandibles, hyoids, several cervical vertebrae, dorsal series, dorsal ribs, uncinate processes, gastralia, partial sacrum, partial caudal series, chevrons, partial scapula, partial coracoid, furcula, sternal plates, humeri (89.4 mm), radii (75.9 mm), ulnae (79.7 mm), semilunate carpal, metacarpal I (14.4 mm), phalanx I-1, manual ungual I, metacarpal II (46.6 mm), phalanx II-1, phalanx II-2, manual ungual II, metacarpal III (~43 mm), phalanx III-1, phalanx III-3, manual ungual III, manual claw sheaths, ilium, pubis (~82.6 mm), ischium, femora (~82.3 mm), tibiotarsi (140.4 mm), metatarsals II, phalanx II-1, phalanx II-2, pedal unguals II, metatarsals III (~73.8 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV, body feathers, enantiornithine humerus, radius, ulna and pes (O'Connor, Zhou and Xu, 2011)
(JI-5) skull postcranial skeleton, feathers (Azuma, 2005)
(JLUM Y-MR160501) skull (57.64 mm), sclerotic plates, mandibles (42.9, 30.91 mm) (Maranga et al., 2020)
(LPM 0159; paratype of Cryptovolans pauli) (~706 mm) dorsal vertebrae, dorsal ribs, uncinate processes, gastralia, sacrum, scapulocoracoid, incomplete furcula, partial sternum, sternal ribs, humeri (74 mm), radii (66 mm), ulnae, carpus, metacarpal I (11.9 mm), phalanx I-1 (30 mm), manual ungual I, metacarpal II (38 mm), phalanx II-1 (16.8 mm), phalanx II-2 (17.6 mm), manual ungual II, metacarpal III, phalanx III-1 (14.2 mm), phalanx III-2 (3.4 mm), phalanx III-3 (10 mm), manual ungual III, manual ungual sheaths, pubis, ischia, femora (89 mm), tibiae (105 mm), metatarsi (~53 mm), phalanx II-2, pedal ungual II, phalanx III-1, phalanx III-2 (7.9 mm), phalanx III-3 (8 mm), pedal ungual III, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, contour feathers, remiges, leg remiges (Czerkas, Zhang, Li and Li, 2002)
(LPM 0200/0201; = BPM 1 3-13; holotype of Cryptovolans pauli) (~950 mm) fragmentary skull, teeth, dorsal vertebrae, dorsal ribs, uncinate processes, twenty-eight to thirty caudal vertebrae, chevrons, furcula, sternum (60 mm), sternal ribs, humeri, radii, ulnae, carpus, manual digit I, metacarpal II, phalanx II-1, phalanx II-2 (26 mm), manual ungual II, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3 (14.2 mm), manual ungual III, manual ungual sheaths, ilium, pubis, ischium, femora (~100 mm), tibiae (132 mm), metatarsus (~66 mm), phalanx II-2, pedal ungual II, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, pedal ungual sheaths, remiges, retrices, leg remiges (Czerkas, Zhang, Li and Li, 2002)
(LVH 0026; holotype of Microraptor hanqingi) (~950 mm) skull, mandibles (92.5 mm), eight cervical vertebrae, cervical ribs, thirteen dorsal vertebrae (~132 mm), dorsal rib fragments, gastralia, sacrum (43.6 mm), twenty-three caudal vertebrae (~535 mm), chevrons, furcula, scapulocoracoids (scapulae 59.9, 54.9 mm), sternal plate (47.3 mm), humeri (92.6, 93.5 mm), radii (79.5, ~77.9 mm), ulnae (82.5, ~81.9 mm), scapholunare, pisiform, semilunate carpal, distal carpal III, metacarpal I (12.98 mm), phalanx I-1 (32.44 mm), manual ungual I (21.98 mm), metacarpal II (54.88 mm), phalanges II-1 (21.96, 23.21 mm), phalanges II-2 (24.2, 24.72 mm), manual unguals II (20.56 mm), metacarpal III (48.17 mm), phalanges III-1 (17.7 mm), phalanges III-2 (4.66 mm), phalanges III-3 (13.1 mm), manual unguals III (9.35 mm), manual claw sheaths, ilium (60 mm), pubes (78 mm), ischia (36 mm), femora (109.8, 111.7 mm), tibiotarsi (137.9, 137.1 mm), metatarsal I, pedal ungual I, metatarsals II, phalanges II-1 (one proximal), phalanges II-1 (one distal), pedal unguals II, metatarsals III (77.7, 75.4 mm), phalanges III-1, phalanges III-2 (one incomplete), phalanges III-3 (one incomplete), pedal unguals III (one incomplete), distal tarsals III+IV fused to metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV (one incomplete, one proximal), pedal claw sheaths, retrices, leg remiges (Alexander et al., 2010)
(NGMC 00-12-A) (~832 mm) incomplete skull, sclerotic plates, incomplete mandible, hyoids, eight cervical vertebrae, second dorsal vertebra (9 mm), third dorsal vertebra (8.1 mm), fourth dorsal vertebra (9.4 mm), fifth dorsal neural arch fragment, sixth dorsal vertebra (8.9 mm), seventh dorsal vertebra (9.3 mm), eighth dorsal vertebra (9.6 mm), ninth dorsal vertebra (9.6 mm), tenth dorsal vertebra (9.6 mm), eleventh dorsal vertebra (9.6 mm), twelfth dorsal vertebra (8.7 mm), partial thirteenth dorsal vertebra, dorsal ribs, uncinate processes, gastralia, first caudal vertebra, third caudal vertebra (11 mm), fourth caudal vertebra (12 mm), fifth caudal vertebra (13.8 mm), sixth caudal vertebra (~18.9 mm), eighth caudal vertebra (26.8 mm), ninth caudal vertebra (27.1 mm), tenth caudal vertebra (26.6 mm), eleventh caudal vertebra (26.1 mm), twelfth caudal vertebra (25.7 mm), thirteenth caudal vertebra (25.2 mm), fourteenth caudal vertebra (26.3 mm), fifteenth caudal vertebra (26 mm), sixteenth caudal vertebra (25.3 mm), seventeenth caudal vertebra (24.8 mm), eighteenth caudal vertebra (23.5 mm), nineteenth caudal vertebra (22.9 mm), twentieth caudal vertebra (21.5 mm), twenty-first caudal vertebra (20.7 mm), twenty-second caudal vertebra (19.2 mm), twenty-third caudal vertebra (17.2 mm), twenty-fourth caudal vertebra (15.5 mm), chevrons, scapulae, incomplete coracoids, incomplete furcula, sternal plates (56 mm), six sternal ribs (15.6, 18.6 mm), humeri (89, ~90.7 mm), radii (78, 75.5 mm), ulnae (79.1, ~77.1 mm), scapholunares, pisiforms, semilunate carpals, distal carpals III, metacarpal I (14.2, 14.2 mm), phalanx I-1 (30, 29.1 mm), manual ungual I, metacarpal II (50.8, 51.9 mm), phalanx II-1 (23.6, 25.6 mm), phalanx II-2 (24.8 mm), manual ungual II, metacarpal III (49.2, 49 mm), phalanx III-1 (19.1, 19.6 mm), phalanx III-2 (4.8, 5.2 mm), phalanx III-3 (14 mm), manual ungual III, manual ungual sheaths, partial ilia, pubes (90 mm), incomplete ischia, femora (~104.8 mm), tibiae (139.9, 137.9 mm), proximal fibula, distal tarsal IV, phalanx I-1, pedal ungual I, incomplete metatarsal II (~71.9 mm), phalanx II-1 (9.7 mm), phalanx II-2 (10.9 mm), pedal ungual II (~23.7 mm), metatarsal III (~77 mm), phalanx III-1 (15.7 mm), phalanx III-2 (11.3, 11.1 mm), phalanx III-3 (11.9, 10.9 mm), pedal ungual III (18.1 mm), metatarsal IV (75.4 mm), phalanx IV-1 (12.8 mm), phalanx IV-2 (8.7, 8.8 mm), phalanx IV-3 (7, 6.7 mm), phalanx IV-4 (7.9, 7.2 mm), pedal ungual IV (17.1, 14.9 mm), metatarsal V (45.7 mm), pedal ungual sheaths, contour feathers, remiges, leg remiges, retrices (Ji, 2002)
(QM V1002; material of Sinornithosaurus "zhaoi") (800 mm adult) skull, mandible, cervical series, dorsal series, dorsal ribs, five uncinate processes, gastralia, sarum, incomplete caudal series, chevrons, scapulae, furcula, sternal plates, humeri (~94 mm), radii, ulnae, metacarpal I, phalanx I-1, manual unguals I, metacarpals II, phalanges II-1, phalanges II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, manual claw sheaths, ilia, pubis, ischia, femora (109 mm), tibiotarsi (one incomplete), fibulae, pedal ungual I, metatarsals II (one incomplete), phalanx II-2, pedal ungual II, metatarsals III (one incomplete), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsals IV (one incomplete), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, pedal claw sheaths, contour feathers, remiges, leg remiges, retrices, actinopterygian neural/haemal arches, ribs and fin rays, teleost cranial elements and two centra (Zhang, 2007)
(TNP00996) (~500 mm) specimen including skull, tail (290 mm), femur (63 mm), contour feathers, retrices and leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
Late Barremian-Early Albian, Early Cretaceous
Yixian or Jiufotang Formation, Liaoning, China
(IVPP 12727) specimen including eight dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, coracoid, proximal humerus, distal radius, ulnar fragment, scapholunare, metacarpal I, pubes, femur and tibia (Turner et al., 2012)
(IVPP V17749) specimen including radius, ulna, scapholunare, semilunate carpal, distal carpal II, proximal metacarpal I, proximal metacarpal II and proximal metacarpal III (Xu et al., 2014)
(IVPP V17750) specimen including radius, scapholunare, semilunate carpal, metacarpal I, phalanx I-1, metacarpal II and metacarpal III (Xu et al., 2014)
(IVPP 200211) specimen including caudal vertebrae and tibia (Turner et al., 2012)
(IVPP coll.) specimen including caudal vertebrae, manual digit I, pubis and tibia (Turner et al., 2012)
(IVPP coll.) specimen including caudal vertebrae, radius, manual digit I, pubis and tibia (Turner et al., 2012)
(IVPP coll.) specimen including teeth, caudal vertebrae, pubis, femur and tibia (Turner et al., 2012)
Early Cretaceous
Qianyang, Liaoning, China
(CAGS 20-7-004) (~590 mm) dentaries (25.1, 33.36 mm), splenials, teeth, tenth cervical vertebra (6.07 mm), tenth cervical rib, first dorsal vertebra (5.97 mm), second dorsal vertebra, third dorsal vertebra (7.4 mm), fourth dorsal vertebra (6.54 mm), fifth dorsal vertebra (7.1 mm), sixth dorsal vertebra (5.39 mm), seventh dorsal vertebra (5.18 mm), eighth dorsal vertebra (6.14 mm), ninth dorsal vertebra (5.76 mm), tenth dorsal vertebra (5.97 mm), eleventh dorsal vertebra (6.08 mm), twelfth dorsal vertebra (6.1 mm), thirteenth dorsal vertebra (6.12 mm), dorsal ribs, two uncinate processes, first caudal vertebra (6.19 mm), second caudal vertebra (7.52 mm), third caudal vertebra (8.58 mm), fourth caudal vertebra (13.12 mm), fifth caudal vertebra (16.54 mm), sixth caudal vertebra (19.42 mm), seventh caudal vertebra (18.49 mm), eighth caudal vertebra (17.90 mm), nineth caudal vertebra (17.16 mm), tenth caudal vertebra (17.24 mm), eleventh caudal vertebra (16.82 mm), twelfth caudal vertebra (17.30 mm), thirteenth caudal vertebra (17.17 mm), fourteenth caudal vertebra (16.38 mm), fifteenth caudal vertebra (15.94 mm), sixteenth caudal vertebra (16.36 mm), seventeenth caudal vertebra (15.48 mm), eighteenth caudal vertebra (14.06 mm), nineteenth caudal vertebra (13.79 mm), twentieth caudal vertebra (12.55 mm), twenty-first caudal vertebra (11.22 mm), twenty-second caudal vertebra (10.64 mm), twenty-third caudal vertebra (9.03 mm), twenty-fourth caudal vertebra (7.44 mm), twenty-fifth caudal vertebra (7.62 mm), twenty-sixth caudal vertebra (5.58 mm), chevrons, partial scapulae, furcula, sternal fragments, four sternal ribs, humeri (61.25, 62.06 mm), radius (~48.01 mm), ulna (53.78 mm), semilunate carpals, metacarpal I (7.84 mm), phalanx I-1 (20.45 mm), proximal metacarpal II, phalanx II-1 (13.26 mm), phalanx II-2 (15.88, 16.39 mm), manual ungual II (9.49, 10.26 mm), proximal metacarpal III, phalanx III-1 (10.12 mm), phalanx III-2 (4.1 mm), phalanx III-3 (10.06, 10.37 mm), fragmentary ilia, incomplete pubes (52.73, ~52.39 mm), incomplete ischia (25.06, 25.22 mm), femora (74.26, 74.75 mm), tibiotarsi (94.22 mm), fibulae (~85.67 mm), metatarsal II (44.51 mm), phalanx II-1 (6.88 mm), phalanx II-2 (7.24, 6.77 mm), pedal ungual II (13.24, 12.88 mm), metatarsal III (47.76 mm), phalanx III-1 (9.95 mm), phalanx III-2 (7.74 mm), phalanx III-3 (6.13 mm), pedal ungual III (9.39 mm), metatarsal IV (46.8 mm), phalanx IV-1 (7.3 mm), phalanx IV-2 (5.4 mm), phalanx IV-3 (5.01 mm), phalanx IV-4 (5.42, 5.01 mm), pedal ungual IV (9.18, 8.95 mm), metatarsal V (22.03 mm) (Hwang, Norell, Qiang and Keqin, 2002)
(CAGS 20-8-001) (~590 mm) ninth cervical vertebra (3.51 mm), tenth cervical vertebrae (4.18 mm), first dorsal vertebra, second dorsal vertebra, third dorsal vertebra, fourth dorsal vertebra, fifth dorsal vertebra (6.38 mm), sixth dorsal vertebra (6.34 mm), seventh dorsal vertebra (6.4 mm), eighth dorsal vertebra (6.42 mm), ninth dorsal vertebra (6.45 mm), tenth dorsal vertebra (6.34 mm), eleventh dorsal vertebra (6.32 mm), twelfth dorsal vertebra (6.24 mm), thirteenth dorsal vertebra (6.12 mm), dorsal ribs, seven uncinate processes, gastralia, sacrum (fourth- 5.28 mm; fifth- 4.91 mm), first caudal vertebra (4.25 mm), second caudal vertebra (5.28 mm), third caudal vertebra (5.63 mm), fourth caudal vertebra (6.16 mm), fifth caudal vertebra (7.42 mm), sixth caudal vertebra (10.32 mm), seventh caudal vertebra (14.7 mm), eighth caudal vertebra (18.47 mm), nineth caudal vertebra (18.91 mm), tenth caudal vertebra (18.45 mm), eleventh caudal vertebra (18.8 mm), twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra (17.64 mm), fifteenth caudal vertebra (18.08 mm), sixteenth caudal vertebra (16.96 mm), seventeenth caudal vertebra (17.08 mm), eighteenth caudal vertebra (16.74 mm), nineteenth caudal vertebra (16.9 mm), twentieth caudal vertebra (~16.06 mm), twenty-first caudal vertebra (15.82 mm), twenty-second caudal vertebra (~15.67 mm), twenty-third caudal vertebra (~14 mm), twenty-fourth caudal vertebra (11.49 mm), twenty-fifth caudal vertebra (~10.66 mm), twenty-sixth caudal vertebra (9.46 mm), chevrons, scapulocoracoids, furcula, sternal plates (~37.99, 38.39 mm), four sternal ribs, humeri (62.88 mm), radii (~48.3 mm), ulnae (~53.5 mm), four manual phalanges, manual ungual I (14.4 mm), manual ungual ?III (10.27 mm), manual ungual sheaths, ilia (40.81, 40.04 mm), pubes (52.77 mm), ischia (26.2, 26.84 mm), femora (74.40, 74.77 mm), tibiotarsi 94.14, ~95.51 mm), fibulae (87.2 mm), distal tarsal III, metatarsal I (8.09 mm), phalanx I-1 (5.26, 5.35 mm), pedal ungual I (3.34 mm), metatarsal II (45.89 mm), phalanx II-1 (7.38, 6.91 mm), phalanx II-2 (7.59 mm), pedal ungual II (~15.2, 16.04 mm), metatarsal III (49.39 mm), phalanx III-1 (10.05, 9.74 mm), phalanx III-2 (~8.24, 8.2 mm), phalanx III-3 (7.18, 8.1 mm), pedal ungual III (9.51, 9.37 mm), metatarsal IV (~48.5 mm), phalanx IV-1 (8.14, 8.39 mm), phalanx IV-2 (5.72, 5.97 mm), phalanx IV-3 (~4.37, 4.16 mm), phalanx IV-4 (5.66, 5.9 mm), pedal ungual IV (9.02 mm), metatarsal V (23.98 mm) (Hwang, Norell, Qiang and Keqin, 2002)
Early Cretaceous
China
(STM 5-11) (subadult) specimen including sternal plates and humeri (O'Connor et al., 2015)
(STM 5-50) (subadult) specimen including scapula, coracoid, furcula and sternal plate (Zhang et al., 2014)
(STM 5-225) specimen including dorsal vertebrae and femur (Gong et al., 2012)
(STM coll.) skeleton including tibiotarsus, metatarsal II, metatarsal III, metatarsal IV, pedal phalanges, body feathers, metatarsal remiges (Alexander et al., 2010)
?(private coll.) skull, mandibles, cervical series, dorsal series, dorsal ribs, sacrum, caudal series, chevrons, furcula, anterior sternum, humeri, radii, ulnae, carpus, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, manual digit III, manual ungual III, ilium, pubis, femora, tibiotarsi, metatarsal II, pedal digit II, metatarsal III, pedal digit III, metatarsal IV, pedal digit IV, remiges (www.paleowonders.com.tw/picture/DN-008.jpg)
(Hong Kong Science Museum coll.) partial skull, cervical series, dorsal series, sacrum, caudal series, scapula, humeri, radii, ulnae, manus, ilia, femora, tibiotarsi, metatarsi, pes, remiges, leg remiges (hk.science.museum/eexhibit/images/sd_mg.jpg)
?(private coll.) skull, mandibles, cervical vertebrae, dorsal series, dorsal ribs, gastralia, sacrum, caudal series, scapula, furcula, humeri, radii, ulnae, carpus, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ischia, femora, tibiotarsi, metatarsi, pedal digit I, pedal digit II, pedal digit III, pedal digit IV (http://www.fossilmall.com/Science/Sites/China/Dromeosauridea/Dromeosauridea.jpg)
Diagnosis- (after Xu et al., 2000) posterior dentary teeth with constricted bases (also in Richardoestesia gilmorei and Paronychodon; may be plesiomorphic for Maniraptoriformes); accessory trochanter on femur.
(after Xu et al., 2003) manual phalanx III-1 >83% of phalanx III-3 in length (also in Wulong).
(after Gong et al., 2012) slender ischial shaft.
(proposed) manual phalanx I-1 <62% of metacarpal II in length; tibiofemoral ratio <136%; metatarsal I <20% of metatarsal III in length; pedal phalanx IV-4 >65% of phalanx IV-1; highly elongate leg remiges (>1.5 times femoral length).
Other diagnoses- Xu et al. (2000) listed "mesial serrations absent from all teeth" as an apomorphy of Microraptor, but large specimen NGMC 00-12-A has mesial serrations on maxillary teeth (Ji, 2002), so this may be size-related. Another possibility is that NGMC 00-12-A is a different species. Xu et al. also listed mid caudal vertebrae 3-4 times length of anterior dorsal vertebrae, but this is also seen in NGMC 91. Another supposed apomorphy listed in the original description is less than 26 caudal vertebrae, but several subsequently discovered specimens have 26 or more caudals. Finally, strongly curved pedal unguals are also seen in NGMC 91 (Senter, 2007).
Contra Xu et al. (2003), the high metacarpal III/II ratio is not diagnostic, being similar to Bambiraptor and matched by NGMC 91. The distal articulation of manual phalanx III-3 does not appear to be smaller or more ventrally skewed than NGMC 91.
Gong et al. (2012) listed the absence of antorbital fossa pitting in their diagnosis, but this is plesiomorphic. Contra their diagnosis, LVH 0026 does appear to have a 'subfenestral fossa', though more anteriorly placed than in Sinornithosaurus. Manual phalanx III-1 is not longer than III-3 in all specimens (e.g. IVPP V13475, CAGS 20-7-004, gui holotype). The ischium is not longer compared to the pubis or femur than in Sinornithosaurus. Not all Microraptor specimens have bowed tibiotarsi (e.g. IVPP coll. in Turner et al., 2012; fig. 20F).
Contra Turner et al. (2012), metatarsal V is not longer than other dromaeosaurids such as Sinornithosaurus and Deinonychus in all specimens (e.g. CAGS 20-7-004 and 20-8-001) and bowing is common in dromaeosaurids.
The "Archaeoraptor" debacle- The holotype was discovered in 1997, though its part and counterpart slabs were quickly separated. One slab contained the tail, which was then fraudulently combined with a euornithine bird skeleton by a Chinese farmer. It was smuggled out of the country then sold at the 1998 Tuscon Gem Show to Czerkas. Currie recognized the legs were part and counterpart slabs of the same bones, while Rowe and Aulenback independently verified the composite nature of the specimen. National Geographic announced the specimen in a press conference in October, and in November, Sloan (1999) published a paper using the name "Archaeoraptor liaoningensis". This was a nomen nudum because it explicitely stated the taxon was to be described formerly in an official publication. That official publication was to have been in Science or Nature, but both journals rejected it. In October, Xu had noticed the tail of "Archaeoraptor" matched the one on the dromaeosaurid skeleton he was describing. Xu located the body which went with this tail by contacting fossil dealers in December. In April, Olson (2000) published an article purporting to officially describe "Archaeoraptor" and attach that name to the dromaeosaurid tail (with the latter as the lectotype). Several months later in December, Xu et al. (2000) officially named Microraptor zhaoianus based on the dromaeosaurid tail and associated anterior part of the skeleton Xu had been studying. At this time, Olshevsky (DML, 2000) noted that Olson's publication predated Xu et al.'s, and he believed that this made Microraptor a junior synonym of "Archaeoraptor". Several days later, Creisler (DML, 2001) pointed out the Olson's attempt to name "Archaeoraptor" was invalid because the ICZN requires a diagnosis in a valid publication, while Olson merely referenced the invalid article by Sloan. Creisler further indicated Olson cannot designate a lectotype without a valid publication defining a holotype first. Thus "Archaeoraptor" is still a nomen nudum, despite Olson's efforts, and Microraptor zhaoianus is the valid name for the IVPP V 12330 dromaeosaurid. The euornithine section (IVPP V 12444) was later described by Czerkas and Xu (2002) as a new taxon- Archaeovolans repatriates, which was in turn synonymized with Yanornis martini by Zhou et al. (2002).
Microraptor gui, Cryptovolans pauli and other specimens- In 2001, IVPP 13476 was discovered, and later mentioned by Xu et al. (2003) as Microraptor sp.. Four additional specimens (IVPP V13320, V13477, V13351, V13352) were purchased by the IVPP in 2001-2002. Xu et al. (2003) mentioned IVPP V13351 as another Microraptor sp. specimen, and IVPP V13477 as Dromaeosauridae gen. et sp. indet., but described V13352 and V13320 as the holotype and paratype of a new species, Microraptor gui. This was based on several characters supposedly distinct from the M. zhaoianus holotype, which have been subsequently questioned by Senter et al. (2004) and Turner et al. (2012). There is a prominent biceps tubercle on the proximal radius of the gui holotype, which Senter et al. and Turner et al. considered individual variation. Both papers correctly noted the bent pubis and low ratio of metacarpal I plus manual phalanx I-1 compared to metacarpal II (~87%) are unknown in the M. zhaoianus holotype, while a curved tibiotarsus is found in that species as well (Turner et al. considered the curvature to be taphonomic). Xu et al. also mention the fused sterna as being different than other dromaeosaurids, but sterna are unknown in the M. zhaoianus holotype. Proportional differences have now been shown to be plausibly allometric, due to intermediate sized Microraptor specimens having intermediate ratios (possibly besides the tibiofemoral ratio; see below). Contrary to popular perception, the famous elongated, asymmetrical feathers attached to M. gui's metatarsi and tibiotarsi were not absent in M. zhaoianus, merely unpreserved in the holotype. Thus there seems to be almost no basis for separating M. gui from M. zhaoianus, let alone as a separate genus "Tetrapterornis" as was suggested by Miller (2004) based on these same characters. The genus is a nomen nudum as Miller always places it in quotation marks, and thus violates ICZN Article 11.5- "a name must be used as valid for a taxon when proposed." IVPP V13477 has the apomorphically long leg remiges of Microraptor, so is here referred to that genus. TNP00996 was purchased in 2002 and briefly described by Xu et al. (2003) as Microraptor sp.. Senter (2011) has since retained gui as a separate OTU than zhaoianus, though there are no differences between the two listed in his table S1. O'Connor et al. (2011) described a new specimen (IVPP V17972) as M. gui based on its large size relative to M. zhaoianus, manual digit proportions, pubic curvature and slight bowing of the tibia. Of these only size is new, which could easily be ontogenetic as sutures are visible in the M. zhaoianus holotype sacrum.
Hwang et al. (2002) described two new Microraptor specimens (CAGS 20-7-004 and 20-8-001) which were collected by farmers from an unknown locality. These were assigned to M. zhaoianus, as M. gui had yet to be described. Although published much later, the M. gui description was accepted for publication several days before Hwang et al.'s paper was published, and Xu et al. (2003) never reference either CAGS specimen. The presence of a biceps tubercle cannot be determined due to preservation, and their tibiotarsi are curved. Though metacarpal II is incomplete distally, 20-7-004 has digit II phalanges placed in a position where the first digit could be as comparatively short as M. gui. The pubic shape cannot be determined as the bones are preserved in anterior view. Size and most proportions (ulnofemoral, ischiofemoral and metatarsofemoral ratios) are intermediate between M. zhaoianus and M. gui, while the tibiofemoral ratio is more similar to zhaoianus (126% compared to 128% in zhaoianus and 133% in gui). The sternal plates of CAGS 20-8-001 are unfused, unlike the M. gui holotype. Proportional differences from M. gui include a smaller ulnohumeral ratio (85-87% vs. 101%), longer phalanx I-1 compared to metacarpal I (261% vs. 211%), longer manual phalanx II-2 compared to II-1 (122% vs. 103%), and longer manual phalanx III-1 compared to III-3 (99% vs. 90%). Senter (2011) viewed the CAGS specimens as distinct from zhaoianus and/or gui based on three characters. He claims the latter two species lack serrated maxillary and dentary teeth, but the zhaoianus type has serrations clearly described and illustrated. IVPP V13320 is indeed unique among reported Microraptor specimens in lacking serrated mid-jaw teeth, but the condition is unpreserved in the gui holotype. Unfused sternal plates do indeed differ from the gui holotype as noted above. Finally, as noted above, manual phalanx I-1 is much longer compared to metacarpal I (261%) than in the gui holotype (211%), but this is also true of Cryptovolans (252%) and hanqingi (250%), though not of NGMC 00-12-A (211%) or IVPP V13475 (209%).
LPM 0200 was first briefly described as an unnamed dromaeosaurid by Norell et al. (2002), then described further and named Cryptovolans pauli by Czerkas et al. (2002) several months later. Most of the bones in the holotype and paratype (LPM 0159) are split between slab and counterslab, leading to very poor preservation and a lack of anatomical detail. Norell et al. correctly described the elongate leg remiges, which Czerkas et al. mistook for forelimb feathers. However, Czerkas et al. were correct in describing the fused sternum and asymmetrical remiges, contra Norell et al.. Czerkas et al. also believed NGMC 91 to possibly be referrable to Cryptovolans, but this specimen is more likely Sinornithosaurus (Ji et al., 2002) or the sister taxon to Microraptor (Senter et al., 2004; Senter, 2007). Most publications have ignored Cryptovolans, perhaps due to the poor description, highly illogical discussion and various personal disputes with Czerkas' publishing practices.
Immediately after Xu et al.'s (2003) publication of M. gui, Holtz (DML, 2003) proposed it is a junior synonym of Cryptovolans pauli (using the new, unpublished combination "Microraptor pauli"). He noted the biceps tubercle cannot be determined as absent in the latter, which also has a bent pubis, curved tibiotarsus and fused sternal plates. The Cryptovolans paratype seems to have a similarly short first manual digit (though poor preservation makes this uncertain), contra Czerkas et al.'s (2002) illustration. Proportional differences between Cryptovolans' paratype and M. gui include a smaller ulnohumeral ratio (~92% vs. 101%) and longer manual phalanx III-1 vs. III-3 (142% vs. 90%). The CAGS specimens are more similar to Cryptovolans in the ulnohumeral ratio, and to M. gui in the manual digit III phalangeal proportions. The Cryptovolans specimens are more similar to the CAGS specimens than M. gui in their ulnohumeral ratios, long manual phalanx I-1 compared to metacarpal I (252%), and long manual phalanx III-1 compared to III-3, but are more similar to M. gui in their shorter manual phalanx II-2 compared to II-1 (105%). Similarly, Miller (2004) stated "Under my own classification, M. gui and C. pauli would both become 'Tetrapterornis' gui", despite the fact both Cryptovolans and pauli have priority over "Tetrapterornis" and gui.
Two of the only papers to mention Cryptovolans (Senter et al., 2004; Senter, 2007) agree it is a junior synonym of Microraptor zhaoianus. Senter et al. state the manual proportions that diagnose Cryptovolans are also seen in other Chinese microraptorians. Czerkas et al. diagnosed Cryptovolans by its high III-1/III-3 ratio (138-142%). If we examine the ratio in other microraptorians, Microraptor's (CAGS 20-7-004) is 98%, Microraptor gui's holotype is 85%, NGMC 91's is 75%, Sinornithosaurus millenii's is 76%, and Graciliraptor's is 64%. Senter et al. are however correct in noting that the Microraptor gui holotype also has fused sternal plates, and that the unfused sternal plates of another Microraptor specimen (CAGS-20-8-001) might be due to ontogeny. Czerkas' final listed diagnostic character (28-30 caudal vertebrae) is similar to Microraptor zhaoianus' holotype (24), Microraptor specimens CAGS 20-7-004 and 20-8-001 (26) and Microraptor gui' holotype (~26). Other maniraptoriform species (Gallimimus bullatus, Shenzhouraptor sinensis) are known to have individual variation in caudal count within 3-4 vertebrae. Czerkas et al. also stated that the tibiofemoral ratio of the paratype scales differently compared to Microraptor, which they viewed as evidence the taxa were not synonymous. Yet the Microraptor gui holotype has almost identical hindlimb size and ratio to the Cryptovolans holotype, while the paratype would seem to have an unsually long femur. It seems unlikely this is due to allometry, as Microraptor shows almost no change in tibiofemoral ratio with age. More likely, individual variation or poor preservation is the cause. Finally, Czerkas et al. stated that Cryptovolans has a longer forelimb than other dromaeosaurids (hum+rad+mcII+pII-1+pII-2 239% of femoral length), but it is subequal or shorter in length to Microraptor (237-258%), Sinornithosaurus millenii (252%), NGMC 91 (256%) and Bambiraptor (244%). Senter (2011) reversed his opinion and retained Cryptovolans as a separate OTU, where it grouped with Sinornithosaurus and Graciliraptor to the exclusion of Microraptor. This was due to several characters. Cryptovolans has a longitudinal labial depression depression on some teeth, but this is polymorphic in many theropods and also found in some teeth of the zhaoianus holotype and M. hanqingi for instance. The second character is the supposed lack of dorsal arching (when the articular surface is held vertically) in manual ungual I, but this is impossible to determine in LPM 0159 due to the proximal portion of ungual being broken (LPM 0200's hands are impossible to evaluate in Czerkas et al.'s photos), and contra Senter is not true of Sinornithosaurus or NGMC 91. Senter also claims Cryptovolans lacks a proximodorsal lip on manual ungual I, but not only is this variable in other taxa (e.g. Archaeopteryx, Sapeornis, Confuciusornis), it is an illusion in Cryptovolans caused by the aforementioned broken proximal portion of that ungual. The final character is tibiotarsal bowing, which Senter incorrectly considered absent in Microraptor.
Czerkas and Ji (2002) illustrate a dromaeosaurid preserved alongside the Omnivoropteryx holotype, referring it tentatively to Cryptovolans. Unfortunately, the illustrations are of x-rays, so little detail can be ascertained. The ulnohumeral and metatarsotibial ratios are more similar to Microraptor specimen NGMC 00-12-A, while the short manual digit I is more similar to Microraptor than Sinornithosaurus, so it is provisionally retained in the former genus until it is described in more detail.
Ji (2002) described a dromaeosaurid (NGMC 00-12-A) found in 2000 as Sinornithosaurus sp.. This was based on several characters, which are all found in other microraptorians. The unserrated premaxillary teeth, posteriorly bifurcated dentary, prominent obturator process and subarctometatarsalian metatarsus are also present in Microraptor, the U shaped furcula is present in all microraptorians, while the extremely short manual phalanx III-2 relative to III-1 is even more similar to Microraptor than Sinornithosaurus. The ischium is more similar to Microraptor than Sinornithosaurus in being slender shafted with a more proximally placed distodorsal process, and the maxillary teeth are more like Microraptor in being mesially unserrated. NGMC 00-12-A seems to lack a pronounced biceps tubercle, but has a short first manual digit, bent pubis and curved tibiotarsus. Unlike Microraptor gui and the holotype and paratype of Cryptovolans, NGMC 00-12-A lacks a fused sternum (though it is the largest well described specimen). In a few proportions, NGMC 00-12-A is more similar to M. gui than Cryptovolans pauli, including the small sternum (53% of femoral length vs. 49% in gui and 60% in pauli), though in others it is more similar to C. pauli. The latter include the (originally supposed to be apomorphic for Cryptovolans) ratio between manual phalanx III-1 and III-3 (139% vs. 142% in pauli and 90% in gui). NGMC 00-12-A is more similar to the CAGS specimens than M. gui in its ulnohumeral ratio (89%) and manual phalanx III-1 compared to III-3, but more similar to M. gui in its short manual phalanx I-1 compared to metacarpal I (208%), and short manual phalanx II-2 compared to II-1 (101%). There are a few differences from smaller Microraptor specimens- more slender dentary, posterior dentary teeth serrated mesially, and taller posterior dorsal neural spines (last dorsal neural spine 72% taller than centrum compared to 63% in CAGS 20-8-001), longer pubofemoral ratio (86% vs. 71-77%), and shorter pedal phalanx III-2 vs. III-1 (59% vs. 78-83%). These are possibly due to age.
Zhang (2007) proposed Sinornithosaurus "zhaoi" for QM V1002 with a short description that claimed to distinguish it from S. millenii based on teeh that are thick and less sharp, and longer arms. However, its humerofemoral ratio (~86%) is actually shorter than the S. millenii holotype (~91%) and Microraptor teeth are typically less elongate than Sinornithosaurus'. The slender ischium, short tibia, elongate pedal phalanx IV-4 and long leg remiges all support assigning QM V1002 to Microraptor instead of Sinornithosaurus, and Xing et al. (2013) described it as a specimen of M. gui. This was based on the biceps tubercle, short manual digit I, bent pubis, and a new character- manual ungual I shorter to be subequal in size to ungual II. The holotype of M. zhaoianus doesn't preserve manual unguals, and this is seemingly based on CAGS 20-8-001 which has been referred to that species without basis. This is the only Microraptor specimen with such a large ungual I compared to II, which is probably due to supposed ungual II belonging to digit III instead. Both unguals are disarticulated and the only ones preserved in that specimen. Notably, the sterna are unfused despite being from an individual 12% larger than the M. gui holotype. Sinornithosaurus "zhaoi" is a nomen nudum as there is no "explicit fixation of a holotype, or syntypes" (ICZN Article 16.4.1), with photos of the skull and pes merely captioned "One skull of Sinornithosaurus zhaoi (sp.nov.)" and "One claw of Sinorni thosaurus [sic] zhaoi (sp.nov.)" respectively and no mention of its catalog number.
Alexander et al. (2010) used LVH 0026 for their biomechanical work, considering it "probably a different species of Microraptor but is morphologically closely similar to M. gui, including the presence of flight-adapted feathers on the tarsometatarsus" (though remember M. zhaoianus' holotype merely doesn't preserve metatarsal remiges, and may have had them in life). Gong et al. (2012) later described it as the new species Microraptor hanqingi. They diagnosed it based on several characters. The larger size (11-14% larger than M. gui or Cryptovolans) could be ontogenetic or individual variation. Unfused sternals are also present in CAGS 20-80-001 and NGMC 00-12-A. The pubis is not more robust than in M. gui, and it cannot be determined if the pubic boot is more squared since most of M. gui's is hidden behind the tibiotarsus. The pubis is indeed bent ~10 degrees less than in the M. gui holotype, but some other specimens lack much bending at all (e.g. IVPP V13475). Contra Gong et al., the ischia of M. hanqingi are posteriorly sinuous and those of M. gui are anteriorly concave, so do not differ in these respects. Additionally, the short manual digit I was said to be like M. gui but unlike M. zhaoianus, and this is true for a referred M. zhaoianus IVPP V13475 and the Cryptovolans paratype, while NGMC 00-12-A is also like M. hanqingi and M. gui. Yet these ratios do not correspond to other skeletal differences as noted in the taxonomy conclusion below. The supposedly longer metatarsal II compared to IV was also supposed to be more similar to M. gui than to M. zhaoianus, but the latter's holotype has a ratio of 98% compared to 91% in M. gui's holotype. In addition, other Microraptor specimens fall between those two ratios, and 7% differences or more are known in other coelurosaurs (e.g. Archaeopteryx lithographica and Dromiceiomimus brevitertius). Finally, its 23 caudal vertebrae were said to be less than M. gui's ~26, but as noted in Cryptovolans' discussion other Microraptor specimens fall between these measures and such variation is known in other species. Pei et al. (2014) agree M. hanqingi is a junior synonym of M. zhaoianus.
In conclusion, the problem with splitting the Microraptor-Cryptovolans clade into species is not the absence of variation, as most of the described specimens have some proportions or characters which differ from all or most other specimens. Instead, the problem is that these differences don't vary in a systematic, congruent way. While we might argue the Cryptovolans specimens and M. gui holotype should be grouped together to the exclusion of M. hanqingi, CAGS 20-80-001, NGMC 00-12-A and QM V1002 based on their fused sterna, we could equally as well argue the Cryptovolans specimens, M. hanqingi and NGMC 00-12-A should be grouped together to the exclusion of the M. gui holotype and CAGS 20-7-004 based on their long manual phalanx III-1. Or that Cryptovolans, M. hanqingi and CAGS 20-7-004 should be grouped together to the exclusion of the M. gui holotype, NGMC 00-12-A and QM V1002 based on their long manual phalanx I-1. There is no obvious answer. One possibility would be to diagnose a separate species for each specimen, and indeed the variation may be due to several different species living in the Jiufotang fauna. This approach has since been taken by Senter (2011), who views M. zhaoianus, M. gui, the CAGS specimens and Cryptovolans pauli as separate species, with the latter closer to Sinornithosaurus and Graciliraptor. The more conservative approach taken by Senter et al. (2004) and Turner et al. (2012) is followed here, where differences are ascribed to individual and ontogenetic variation. Further description of the specimens mentioned by Xu et al. (2003) and others will help confirm or deny this view.
Maranga et al. (2020) preliminarily describe a new skull (JLUM Y-MR160501) from Sihedang they call Microraptor sp., notable for having "denticles on both the mesial and distal carinae of the posterior dentary teeth, a characteristic that has never been observed in Microraptor to date." However as noted above this is also present in NGMC 00-12-A. Oddly they recover the specimen sister to Microraptoria plus eudromaeosaurs using Currie's dromaeosaurid analysis.
A number of undescribed specimens appear to belong to Microraptor as defined here. One on the Paleowonders website is particularily complete, preserving faint remiges. It was advertised as Microraptor gui at a fossil show. The manual phalanx I-1 length suggests it is in fact Microraptor. One photo of the specimen seems more simplified, lacks an ilium, and has contaur feather impressions. It may be a cast. The specimen appears to preserve rather thick, elongate bones in the pelvic and thoracic areas which are near certainly from another animal.
Another specimen is shown at the Hong Kong Science Museum's "Soaring Dinosaurs" exhibit, advertised as Microraptor gui. This specimen is largely complete, but strangely positioned, with the presacral column sharply curved backward and the hindlimbs extending anteriorly. The elongate leg remiges confirm the identification as Microraptor.
Yet another specimen is complete and perfectly articulated, photographed on Fossil Mall's website as Dromaeosauridae cf. Sinornithosaurus. Its manual phalanx III-3 is elongate like Sinornithosaurus, yet its tibiofemoral ratio and manual phalanx I-1 are short as in Microraptor. It is tentaively referred to Microraptor here.
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Xing, Persons, Bell, Xu, Zhang, Miyashita, Wang and Currie, 2013. Piscivory in the feathered dinosaur Microraptor. Evolution. 67(8), 2441-2445.
Evangelista, Cardona, Guenther-Gleason, Huynh, Kwong, Marks, Ray, Tisbe, Tse and Koehl, 2014. Aerodynamic characteristics of a feathered dinosaur measured using physical models. Effects of form on static stability and control effectiveness. PLoS ONE. 9(1), e85203.
Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014. A new raptorial dinosaur with exceptionally long feathering provides insights into dromaeosaurid flight performance. Nature Communications. 5, 4382.
Palmer, 2014. The aerodynamics of gliding flight and its application to the arboreal flight of the Chinese feathered dinosaur Microraptor. Biological Journal of the Linnean Society. 113, 828-835.
Pei, Li, Meng, Gao and Norell, 2014. A new specimen of Microraptor (Theropoda: Dromaeosauridae) from the Lower Cretaceous of Western Liaoning, China. American Museum Novitates. 3821, 28 pp.
Xu, Han and Zhao, 2014. Homologies and homeotic transformation of the theropod 'semilunate' carpal. Nature Scientific Reports. 4, 6042.
Zheng, O'Connor, Wang, Wang, Zhang and Zhou, 2014. On the absence of sternal elements in Anchiornis (Paraves) and Sapeornis (Aves) and the complex early evolution of the avian sternum. Proceedings of the National Academy of Sciences. 111(38), 13900-13905.
O'Connor, Zheng, Sullivan, Chuong, Wang, Li, Wang, Zhang and Zhou, 2015. Evolution and functional significance of derived sternal ossification patterns in ornithothoracine birds. Journal of Evolutionary Biology. 28(8), 1550-1567.
Rowe, Luo, Ketcham, Maisano and Colbert, 2016 online. Archaeoraptor, Digital Morphology. http://digimorph.org/specimens/Archaeoraptor_forgery/
Maranga, Reisz and Evans, 2020. Detailed morphology of the skull and dentition in a new, exceptionally preserved specimen of Microraptor (Theropoda: Dromaeosauridae) from the Early Cretaceous of China. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 234-235.
Wang and Pei, 2024 online. The smallest known specimen of Microraptor (Dinosauria: Dromaeosauridae) from the Jiufotang Formation in northeastern China. Historical Biology. DOI: 10.1080/08912963.2024.2385604

unnamed clade (Bambiraptor feinbergi + Dromaeosaurus albertensis)

unnamed dromaeosaurid (Senter, Kirkland, Deblieux and Madsen, 2010)
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Material- (UMNH VP 21751) incomplete radius
....(UMNH VP 21752) incomplete pubis
Comments- This was initially identified (Senter et al., 2010) as part of what would later be named Yurgovuchia by Senter et al. (2012). The latter referred it to Velociraptorinae based on the large pubic tubercle.
References- Senter, Kirkland, Deblieux and Madsen, 2010. Three new theropods from the Cedar Mountain Formation (Lower Cretaceous) of Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 162A.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.

Bambiraptor Burnham, Derstler, Currie, Bakker, Zhou and Ostrom, 2000
= "Linsterosaurus" Buchholz, DML, 1997
B. feinbergi Burnham, Derstler, Currie, Bakker, Zhou and Ostrom, 2000
= Bambiraptor feinbergorum Norell and Makovicky, 2004
Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
Holotype- (AMNH 30556; = AMNH 001; = FIP 001; Bambi) (subadult) skull (127 mm), sclerotic plates, stapes, incomplete mandibles (~122 mm), hyoid, atlas, axis (13 mm), third cervical vertebra (13.5 mm), fourth cervical vertebra (13.5 mm), fifth cervical vertebra (16 mm), sixth cervical vertebra (14.5 mm), seventh cervical vertebra (~15 mm), eighth cervical vertebra (12 mm), ninth cervical vertebra (13 mm), fragmentary anterior cervical ribs, first dorsal vertebra (11 mm), second dorsal vertebra (12 mm), third dorsal vertebra (12 mm), fourth dorsal vertebra (11.8 mm), fifth dorsal vertebra (11 mm), sixth dorsal vertebra (13 mm), seventh dorsal vertebra (10 mm), eighth dorsal vertebra (10.5 mm), ninth dorsal vertebra (9.5 mm), tenth dorsal vertebra (10 mm), eleventh dorsal vertebra (10.5 mm), twelfth dorsal vertebra (10.8 mm), thirteenth dorsal vertebra (10.5 mm), dorsal ribs, gastralia, first sacral vertebra (11 mm), second sacral vertebra (15 mm), third sacral vertebra (15 mm), fourth sacral vertebra (13 mm), fifth sacral vertebra (11.5 mm), first caudal vertebra (9.1 mm), second caudal vertebra (12.6 mm), third caudal vertebra (13.1 mm), fourth caudal vertebra (14.5 mm), fifth caudal vertebra (15.6 mm), sixth caudal vertebra (16.1 mm), seventh caudal vertebra (17.9 mm), eighth caudal vertebra (20.3 mm), ninth caudal vertebra (21.6 mm), tenth caudal vertebra (24.2 mm), eleventh caudal vertebra (24.6 mm), twelfth caudal vertebra (26 mm), thirteenth caudal vertebra (~27 mm), fourteenth caudal vertebra (27 mm), fifteenth caudal vertebra (~26 mm), sixteenth caudal vertebra (~24 mm), seventeenth caudal vertebra (23 mm), eighteenth caudal vertebra (22 mm), nineteenth caudal vertebra, twentieth caudal vertebra, twenty-first caudal vertebra, twenty-second caudal vertebra, proximal chevron, distal chevrons, scapulae (~83, 85 mm), coracoids, furcula, sternal plates (67, 63 mm), humeri (~105, 100 mm), radii (85, 85 mm), ulnae (95, 93 mm), scapholunares, semilunate carpals, metacarpal I (16.8, 16.8 mm), phalanx I-1 (32.5, 32.3 mm), manual ungual I, metacarpal II (47.8, 46.5 mm), phalanx II-1 (21.1, 21.4 mm), phalanx II-2 (35, 35 mm), manual ungual II (43 mm), metacarpal III (44.9, 43.8 mm), phalanx III-1 (15.5, 16.5 mm), phalanx III-2 (6.3, 5.5 mm), phalanx III-3 (23.5 mm), ilia (86, ~78 mm), pubes (103, 103 mm), ischia (53, 50 mm), femora (118, 118 mm), tibiae (167, 170 mm), fibulae (~170 mm), astragalus, calcaneum, distal tarsal III, distal tarsal IV, metatarsal I (18.3 mm), phalanx I-1 (12.1 mm), pedal ungual I (14 mm), metatarsal II (70, 67.5 mm), phalanx II-1 (14.3, 14 mm), phalanx II-2 (14.6, 14.2 mm), pedal ungual II (46 mm), metatarsal III (77, 81 mm), phalanx III-1 (29.6, 27.8 mm), phalanx III-2 (17.6, 15.4 mm), phalanx III-3 (16.9 mm), pedal ungual III (24 mm), metatarsal IV (70, 74 mm), phalanx IV-1 (23.6, 23.4 mm), phalanx IV-2 (16.6, 17.8 mm), phalanx IV-3 (11.6, 12.3 mm), phalanx IV-4 (12.3, 12.9 mm), pedal ungual IV (22 mm), metatarsal V (33.2 mm)
Paratypes- (FIP 002-136) (at least two adults) thirty-four elements including caudal vertebrae, humerus (145 mm), femur (170 mm), tibia (225 mm), metatarsal III (105 mm)
Referred- (MOR 553S-7-30-91-274) maxilla (~94 mm) (Currie and Varricchio, 2004)
Comments- The holotype was discovered in 1993 and initially believed to be Saurornitholestes, then reported as Velociraptor sp. (Burnham et al., 1997). Buchholz (DML 1997) used the name "Linsterosaurus" for this taxon on the Dinosaur Mailing List between 1997 and 2000, while the Oxford Museum of Natural History labeled their cast of the specimen Velociraptor feinbergi (Taylor, DML 2003). Neither of these names have been published however. It was later described as a new taxon of dromaeosaurid (Burnham et al., 2000), though some believe it should be synonymized with Saurornitholestes. However, Hohman and Varricchio (2021) described juvenile Saurornitholestes maxilla MOR 9753 which is smaller than the Bambiraptor holotype, but shares characters with adult Saurornitholestes maxillae. Similarly, they find larger maxilla MOR 553S-7-30-91-274 groups with Bambiraptor in Currie and Varracchio's dromaeosaurid analysis and differs from Saurornitholestes. Thus the taxa are unlikely to be synonymous. Olshevsky (DML, 2000) noted Bambiraptor feinbergi was named after multiple people, so suggested it be emended to B. feinbergorum, which is followed on several websites and publications. However, the Fourth Edition of the ICZN no longer requires emendations based on this reasoning (Article 31.1.3), as Creisler pointed out later that day (DML, 2000). Norell and Makovicky (2004) first published the name Bambiraptor feinbergorum, which according to ICZN Article 32.2.3 is an available name with its own authorship, though an objective junior synonym of B. feinbergi.
Currie and Evans (2020) identified the first and second premaxillary teeth, the latter of which is catalogd as "Paronychodon?" due to the ridges also found in Saurornitholestes ('Zapsalis') teeth.
Longrich and Currie (2009) found Bambiraptor to be a saurornitholestiine in their analysis, and Brusatte et al. (2014) found the genera to be sister taxa within Velociraptorinae. Foth et al. (2014) and Senter et al. (2012) have both found Bambiraptor to be outside the Velociraptorinae+Dromaeosaurinae clade, though neither included Saurornitholestes.
References- Buchholz, DML 1997. https://web.archive.org/web/20180115091825/http://dml.cmnh.org/1997Jun/msg00518.html
Burnham, Derstler and Linster, 1997. A new specimen of Velociraptor (Dinosauria: Theropoda) from the Two Medicine Formation of Montana. Dinofest International Proceedings. 73-75.
Burnham, Derstler, Currie, Bakker, Zhou and Ostrom, 2000. Remarkable new birdlike dinosaur (Theropoda: Maniraptora) from the Upper Cretaceous of Montana. The University of Kansas Paleontological Contributions. 13, 1-14.
Burnham and Durstler, 2000. Bambiraptor feinbergi (Dinosauria, Theropoda, Dromaeosauridae). The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History. 10.
Cooley, 2000. The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History. 13.
Creisler, DML 2000. https://web.archive.org/web/20180115091851/http://dml.cmnh.org/2000Mar/msg00361.html
Derstler and Burnham, 2000. Phylogenetic Context of Bambiraptor feinbergi. The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History. 15.
Garstka, Marsic, Carroll, Heffelfinger, Lyson and Ng, 2000. Analysis of the structure and articulation of the forelimb bones of the maniraptoran dinosaur, Bambiraptor feinbergi, support predation as the pre-adaptation of flight. The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History. 16.
Olshevsky, DML 2000. https://web.archive.org/web/20180115091846/http://dml.cmnh.org/2000Mar/msg00342.html
Taylor, DML 2003. https://web.archive.org/web/20180115091925/http://dml.cmnh.org/2003Oct/msg00210.html
Burnham, 2004. New information on Bambiraptor feinbergi (Theropoda: Dromaeosauridae) from the Late Cretaceous of Montana. in Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons. Studies on the transition from dinosaurs to birds. Indiana University Press. 67-111.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons. Studies on the transition from dinosaurs to birds. Indiana University Press. 112-132.
Norell and Makovicky, 2004. Dromaeosauridae. in Weishampel, Dodson and Osmólska (eds.). The Dinosauria (second edition). University of California Press, Berkeley. 196-209.
Senter, 2006. Comparison of forelimb function between Deinonychus and Bambiraptor (Theropoda: Dromaeosauridae). Journal of Vertebrate Paleontology. 26(4), 897-906.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Currie and Evans, 2020 (online 2019). Cranial anatomy of new specimens of Saurornitholestes langstoni (Dinosauria, Theropoda, Dromaeosauridae) from the Dinosaur Park Formation (Campanian) of Alberta. The Anatomical Record. 303(4), 691-715.
Hohman and Varricchio, 2021. Ontogenetic implications of two distinct dromaeosaurid maxillae from the Cretaceous (Campanian) Two Medicine Formation of Montana, U.S.A. The Society of Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual Meeting. 141-142.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as 2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters. Journal of Vertebrate Paleontology. 41(5), 2010087.

Variraptor Le Loeuff and Buffetaut, 1998
V. mechinorum Le Loeuff and Buffetaut, 1998
Late Campanian-Early Maastrichtian, Late Cretaceous
Gres a Reptiles Formation, France
Holotype- (MDE-D168) thirteenth dorsal vertebra
....(MDE-D169) (sacrum 160 mm) first sacral vertebra (25 mm), second sacral vertebra (27 mm), third sacral vertebra (32 mm), fourth sacral vertebra (32 mm), fifth sacral vertebra (30 mm)
....(CM-645) ilium (230 mm) (Chanthasit and Buffetaut, 2009)
Paratypes- ?(MDE-D01) anterior dorsal vertebra (29 mm) (Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992)
?(MDE-D158) humerus (195 mm)
?(MDE-D203; cast is MDE-D49 and TMP 1991.185.0001) incomplete femur (~230 mm) (Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992)
(MDE coll.) fifth sacral vertebra (Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992)
Referred- ?(CM-186) tooth (FABL 4.6 mm) (Chanthasit and Buffetaut, 2009)
?(CM-218) tooth (FABL 7.1 mm) (Chanthasit and Buffetaut, 2009)
?(CM-307) ulna (145 mm) (Chanthasit and Buffetaut, 2009)
?(CM-537) tooth (FABL 5.9 mm) (Chanthasit and Buffetaut, 2009)
?(CM-684) tooth (FABL 6.7 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M187) femur (214 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M228) pedal ungual III or IV (46.6 mm on curve) (Chanthasit and Buffetaut, 2009)
?(MC-M324) tooth (FABL 5 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M561) tooth (FABL 4.3 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M938) tooth (Chanthasit and Buffetaut, 2009)
(MC-PSP6) (sacrum 134 mm) first sacral vertebra (20.4 mm), second sacral vertebra (26 mm), third sacral vertebra (28.4 mm), fourth sacral vertebra (28.6 mm), fifth sacral vertebra (22.6 mm) (Chanthasit and Buffetaut, 2009)
?(MC-VC1) tooth (Chanthasit and Buffetaut, 2009)
?(MC-VC2) tooth (FABL 6 mm) (Chanthasit and Buffetaut, 2009)
?(MC coll.) proximal manual ungual (Chanthasit and Buffetaut, 2009)
? several fragmentary dorsal ribs (Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992)
Diagnosis- (proposed) sacral centra three and four transversely compressed (transverse width <60% of first sacral centrum).
Comments- Le Loeuff et al. (1992) described a femur (MDE-D203) from Metisson (Fox-Amphoux, Var), and an anterior dorsal vertebra (MDE-D01), posterior sacral vertebra (MDE coll.) and several fragmentary dorsal ribs from Roques-Hautes (Bouches-du-Rhone). They believed these were congeneric or at least related to Elopteryx, most closely related to dromaeosaurids, though perhaps deserving their own family or subfamily (Elopterygidae or Elopteryginae). The femur was only stated to share general characteristics with Elopteryx (reduced fourth trochanter, posterior trochanter, "shape and size") while differing in having a linear capital ligament fossa and absent fourth trochanter. The other remains are not comparable to Elopteryx. It is here tentatively assigned to Variraptor as it is from the type locality.
Buffetaut et al. (1997) first mention the sacrum (MDE-D169) and humerus (MDE-D158) from Bastide Neuve (Fox-Amphoux, Var) as a new taxon of dromaeosaurid. Le Loeuf and Buffetaut (1998) described and named this taxon Variraptor, with the sacrum and an articulated dorsal vertebra as the holotype. They referred the anterior dorsal and posterior sacral described by Le Loeuff et al. (1992), as well as a femur (MDE-D49) and the aforementioned humerus. The fifth sacral was said to be identical to that of the holotype, though the other remains are not comparable. The femur was not mentioned further, and seems to be a cast of MDE-D203.
Buffetaut et al. (1999) first mentioned teeth (MC-M324, M561 and M938) and a femur (MC-M187) from Massecaps (Cruzy, Herualt) and referred it to Variraptor. Chanthasit and Buffetaut (2009) later described this material, as well as a partial manual ungual (MC coll.) and pedal ungual (MC-M228) from the same locality. Chanthasit and Buffetaut also described two teeth (MC-VC1 and VC2) from Combebelle and a sacrum (MC-PSP6) from Plo Saint Pons, both also in Cruzy. Finally, they describe five teeth (CM-186, 218, 307, 537 and 684), an ulna (CM-307) and an ilium (CM-645) from Bastide Neuve. They noted the ilium matches the holotype sacrum exactly, so probably belongs to the same individual, while the sacrum has Variraptor's apomorphic proportions. Unfortunately, the femur was not compared to MDE-D203, but the ulna does differ from Pyroraptor in lacking a deep brachial fossa. Chanthasit and Buffetaut do not refer most of their material to a particular genus, though it is all provisionally referred to Variraptor here, given its presence in Bastide Neuve, Roques-Hautes and Plo Saint Pons.
Allain and Taquet (2000) correctly noted no diagnostic dorsal or sacral characters were described by Le Loeuff and Buffetaut, so they considered Variraptor a nomen dubium. Yet as noted by Chanthasit and Buffetaut, Variraptor does differ from all other comparable dromaeosaurids. The presence of posterior dorsal pleurocoels distinguishes it from Microraptor and Velociraptor, while it has one less pair of dorsal pleurocoels than Achillobator. The pleurocoels are smaller than in Deinonychus however. The dorsal vertebra is anteroposteriorly compressed, unlike microraptorians. Indeed, the compression is a dromaeosaurine synapomorphy (Senter, 2007). Variraptor has less sacrals than Deinonychus, Ornithodesmus and unenlagiines, and its sacrum is not dorsally arched like those of Saurornitholestes and Ornithodesmus. Variraptor differs from Microraptor, Sinornithosaurus, Saurornitholestes, Ornithodesmus and Velociraptor in having highly compressed mid sacral centra. It may be a synonym of Pyroraptor, but it would be a senior synonym if so. The newly described ilium will probably provide further diagnostic characters.
Rauhut (2000) refers Variraptor to Coelurosauria indet. since some Archaeopteryx specimens and perhaps juvenile individuals of other maniraptoriform clades have five sacral vertebrae, while he finds the rectangular posterior central face of the last sacral (and presumably proximal caudals) to be potentially present in other maniraptorans. Indeed, basal members of Therizinosauria, Oviraptorosauria and Troodontidae all have five sacral vertebrae as well. Also, most non-pygostylian maniraptorans have rectangular proximal caudal centra. Yet troodontids and basal oviraptorosaurs lack posterior dorsal and sacral pleurocoels, and the high degree of fusion in Variraptor suggests it was not a juvenile. The only non-dromaeosaurid taxon with five sacral vertebrae, pleurocoelous posterior dorsals and anterior sacrals, and rectangular proximal caudal vertebrae is Falcarius. Variraptor is more similar to dromaeosaurids in having a hyposphene separated by the posterior ligament groove, and shorter dorsal centra.
Rauhut further suggested the paratype anterior dorsal vertebra is a caenagnathid, based on resemblence to Chirostenotes, especially in having two pairs of pleurocoels. Yet Achillobator and Utahraptor both also have two pairs of pleurocoels in some dorsals, and Variraptor resembles Deinonychus more than Chirostenotes in having a longer infraprezygopophyseal fossa, more poorly developed inradiapophyseal fossa, anteroposteriorly narrower neural spine, and basally restricted posterior interspinous ligament groove. The prominent epipophyses also suggest anterior cervical epipophyses would be well developed, as in dromaeosaurids but not oviraptorosaurs. Rauhut's hypothesis is unsupported, and Variraptor is not demonstrably a chimaera.
Senter et al. (2004) included Variraptor in a phylogenetic analysis and found only that it was a coelurosaur closer to birds than tyrannosauroids, and not a Jehol microraptorian or member of the Achillobator+Utahraptor+Dromaeosaurus clade. However, their analysis lacked many relevent characters (precise sacral number; sacral pleurocoel presence; posterior dorsal pleurocoel presence; proximal caudal centrum shape; etc.), so this result is meaningless.
Hartman et al. (2019) recover the taxon sister to Bambiraptor, while Longrich et al. (2021) suggest several characters are shared with unenlagiines ("enlarged cuppedicus fossa, arched dorsal margin of the ilium and deep anterior blade, short posterior blade, concave posterodorsal margin of the blade, and a brevis fossa that is reduced anteriorly").
References- Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992. The first record of dromaeosaurid dinosaur (Saurichia, Theropoda) in the Maastrichtian of Southern Europe: palaeobiogeographical implications. Bulletin de la Societe Geologique de France. 163(3), 337-343.
Buffetaut, Le Loeuff, Cavin, Duffaud, Gheerbrant, Laurent, Martin, Rage, Tong and Vasse, 1997. Late Cretaceous non-marine vertebrates from southern France: a review of recent finds. Geobios. 20, 101-108.
Le Loeuff and Buffetaut, 1998. A new dromaeosaurid theropod from the Upper Cretaceous of Southern France. Oryctos. 1, 105-112.
Buffetaut, Le Loeuff, Tong, Duffaud, Cavin, Garcia, Ward and L'association Culturelle, Archeologique Et Paleontologique De Cruzy, 1999. A new Late Cretaceous vertebrate locality at Crazy (Harault, southern France). Les Comptes rendus de l'Académie des sciences. 328, 203-208.
Allain and Taquet, 2000. A new genus of Dromaeosauridae (Dinosauria, Theropoda) from the Upper Cretaceous of France. Journal of Vertebrate Paleontology. 20(2), 404-407.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). PhD thesis, University of Bristol. 583 pp.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8, 1-20.
Chanthasit and Buffetaut, 2009. New data on the Dromaeosauridae (Dinosauria: Theropoda) from the Late Cretaceous of southern France. Bulletin de la Societe Geologique de France. 180(2), 145-154.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new paravian dinosaur from the Late Jurassic of North America supports a late acquisition of avian flight. PeerJ. 7:e7247.
Longrich, Martill and Jacobs, 2021. A new dromaeosaurid dinosaur from the Wessex Formation (Lower Cretaceous, Barremian) of the Isle of Wight, and implications for European palaeobiogeography. Cretaceous Research. Journal Pre-proof. DOI: 10.1016/j.cretres.2021.105123

Eudromaeosauria Longrich and Currie, 2009
Definition- (Saurornitholestes langstoni + Velociraptor mongoliensis + Deinonychus antirrhopus + Dromaeosaurus albertensis) (Longrich and Currie, 2009)
= Dromaeosauridae sensu Padian et al., 1999
Definition- (Dromaeosaurus albertensis + Velociraptor mongoliensis) (modified)

"Dromaeosaurus" gracilis (Marsh, 1888) Matthew and Brown, 1922
= Coelurus gracilis Marsh, 1888
= Chirostenotes gracilis (Marsh, 1888) Gilmore, 1924
Late Aptian, Early Cretaceous
near Miurkirk USNM 41615, Arundel Formation, Prince George's County, Maryland, US
Holotype- (USNM 4973) proximal manual ungual III (~12.7 mm tall prox)
Referred- ?(USNM 8176; lost) tooth (Lull, 1911)
?(USNM 8444; = Goucher College 3336) tooth (Lull, 1911)
?(USNM 8445; = Goucher College 3338) tooth (~12.8~6.2x? mm) (Lull, 1911)
? teeth (Lipka, 1998)
Comments- The holotype was discovered in 1887 and described by Marsh (1888) as a new species of Coelurus. In addition to the manual ungual, he referenced metatarsals which were not necessarily based on real specimens. Lull (1911) recognized the ungual as pertaining to digit III and referred three teeth. USNM 8444 and 8445 were found in 1895 (USNM online). One tooth is said to lack mesial serrations while another has extremely small mesial serrations which are restricted apically. Gilmore (1920) noted four unguals from the Belly River Group of Canada at the AMNH (including AMNH 5387) were nearly identical to the holotype. Matthew and Brown (1922) provisionally referred the taxon to their new genus Dromaeosaurus, as (?)Dromaeosaurus gracilis. This was done without justification. Gilmore (1924) believed the AMNH unguals belonged to Chirostenotes, and stated Coelurus gracilis "should now be removed to this genus" but didn't explicitly use the combination Chirostenotes gracilis. Ostrom (1969) notes a strong resemblence of the holotype to the first manual ungual of Deinonychus, but considered it indeterminate. Lipka (1998) reported recovering teeth similar to those described by Lull, which remain undescribed. He noted Coelurus gracilis may relate to the Arundel Deinonychus-like teeth he described.
The extremely large flexor tubercle and proximodorsal lip suggest the ungual is from digit III as suggested by Lull, contra Ostrom. The holotype ungual differs from Chirostenotes in being more curved, having a smaller proximodorsal lip, more proximally placed flexor tubercle and a proximally undivided longitudinal groove. Microvenator's unguals resemble Chirostenotes in being less curved with more distally placed flexor tubercles. Hagryphus is more similar in being more strongly curved with a more proximally placed flexor tubercle than Chirostenotes, but still has a divided groove and larger proximodorsal lip. These oviraptorosaurs also all have lower flexor tubercles than USNM 4973. Among oviraptorids, only Citipati is similar in having strongly curved third manual unguals with a proximodorsal lip, but differs in having a lower flexor tubercle. This also seems to be the case for Yixianosaurus and Tanycolagreus. Archaeopteryx and Confuciusornis unguals are more slender and less curved with much lower flexor tubercles. This leaves dromaeosaurids, among which microraptorians and Saurornitholestes seem less similar than derived taxa like Velociraptor and Deinonychus in having lower flexor tubercles. This suggests USNM 4973 is a derived dromaeosaurid, perhaps a synonym of Deinonychus based on its provenence. Thus Matthew and Brown's generic assignment is more accurate than Marsh's, and is followed here. It should be noted that based on its early age, USNM 4973 is near certainly not Dromaeosaurus however. The unguals mentioned by Gilmore are presumably also dromaeosaurid, perhaps belonging to Saurornitholestes and/or Dromaeosaurus. Lull's and Lipka's referred teeth may also be dromaeosaurid, though basal coelurosaurs may also lack mesial serrations. More information is needed.
References- Marsh, 1888. Notice of a new genus of Sauropoda and other dinosaurs from the Potomac Formation. American Journal of Science, 3rd Series. 35, 89-94.
Lull, 1911. The Reptilia of the Arundel Formation. Maryland Geological Survey, Lower Cretaceous. 174-178.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States National Museum with special reference to the genera Antrodemus (Allosaurus) and Ceratosaurus. United States National Museum Bulletin. 110, l-154.
Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus from Cretaceous of Alberta. Bulletin of the American Museum of Natural History. 66, 367-385.
Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta. Canada Geological Survey, Bulletin 38, geological series 43, 1-13.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin. 30, 1-165.
Lipka, 1998. The affinities of the enigmatic theropods of the Arundel Clay facies (Aptian), Potomac Formation, Atlantic Coastal Plain of Maryland. in Kirkland, Lucas and Estep (eds). Lower to Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Sciences Bulletin. 14, 229-234.

Vectiraptor Longrich, Martill and Jacobs, 2022
= "Vectiraptor" Longrich, Martill and Jacobs, 2021 online
V. greeni Longrich, Martill and Jacobs, 2022
= "Vectiraptor greeni" Longrich, Martill and Jacobs, 2021 online
Barremian, Early Cretaceous
Wessex Formation, England
Material- (IWCMS 2021.31.1) (~7-9 year old adult) incomplete posterior dorsal vertebra (~39 mm)
....(IWCMS 2021.31.2) partial anterior dorsal vertebra (~48 mm)
....(IWCMS 2021.31.3) partial synsacrum
Diagnosis- (after Longrich et al., 2021) small anterior dorsal pleurocoels and large, elliptical posterior dorsal pleurocoels; camellate internal architecture in dorsal vertebrae (also in Unenlagiinae); dorsal vertebrae with large neural canals (also in other paravians); dorsal vertebrae with deep, triangular infradiapophyseal fossae (also in Deinonychus, Utahraptor and ?Itemirus); infradiapophyseal fossae divided by accessory lamina in anterior dorsal vertebrae; deep dorsal infrapostzygapophyseal fossae; massive dorsal neural spines with broad ligament scars (also in Saurornitholestes); sacral centra not pneumatized; ventral median groove not extending the full length of sacrum; sacrum with massive neural canal (also in Saurornitholestes and Rahonavis).
Comments- The holotype was collected in 2004 and described in an online Journal Pre-proof on December 17 2021 by Longrich et al. (2021). As the paper has no mention of ZooBank, according to ICZN Article 8.5.3 (an electronic work must "be registered in the Official Register of Zoological Nomenclature (ZooBank) (see Article 78.2.4) and contain evidence in the work itself that such registration has occurred"), "Vectiraptor greeni" Longrich et al., 2021 was a nomen nudum that became valid in the June 2022 issue.
Longrich et al. proposed "affinities with Eudromaeosauria, perhaps as a primitive member of the group or sister to it" based on broad interspinous ligament scars that differ from Unenlagia, and three characters shared with that genus and Eudromaeosauria (short dorsal centra, posterior dorsal pleurocoels, tall and anteroposteriorly narrow dorsal neural spines). Based on this, they suggested several teeth from the Wessex Formation (IWCMS 2002.1, 3, 4 and NHMUK R 16510) may be referrable as they are the correct size, although the cooccurance of dromaeosaurid species in other formations makes this speculative. Adding it to Hartman et al.'s maniraptoromorph matrix results in it being a velociraptorine sister taxon to IGM 100/22-23.
Reference- Longrich, Martill and Jacobs, 2022 (online 2021). A new dromaeosaurid dinosaur from the Wessex Formation (Lower Cretaceous, Barremian) of the Isle of Wight, and implications for European palaeobiogeography. Cretaceous Research. 134, 105123.

Yurgovuchia Senter, Kirkland, DeBlieux, Madsen and Toth, 2012
Y. doellingi Senter, Kirkland, DeBlieux, Madsen and Toth, 2012
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Holotype- (UMNH VP 20211) (adult) atlas, axis, incomplete third cervical vertebra (45.1 mm), incomplete posterior cervical vertebra (~24.89 mm), incomplete posterior cervical vertebra (~40.95 mm), two partial cervical neural arches, first dorsal vertebra (~26.42 mm), partial dorsal vertebra (32.04 mm), partial dorsal neural arch, incomplete proximal caudal vertebra (40.17 mm), incomplete proximal caudal vertebra (45.95 mm), proximal caudal vertebra (48.03 mm), mid caudal vertebra (52.91 mm), mid caudal vertebra, mid caudal vertebra (55 mm), distal caudal centrum (~48.34 mm), proximal pubis
Diagnosis- (after Senter et al., 2012) centrum of axis with single pneumatopore on each side; cranial end of centrum of third cervical vertebra not beveled; cervical prezygapophyses flexed; epipophyses of cervical vertebrae above postzygapophyseal facets; anterior dorsal vertebrae with hypapophyses and without pneumatopores; anterior faces of centra of proximal caudal vertebrae round; caudal prezygapophyses elongated distal to transition point, but not over the length of a centrum; pubis without pubic tubercle.
Comments- The holotype was discovered in 2005 and initially mentioned by Senter et al. (2010). The latter noted a lacrimal, radius and ilium among the material, which were not in the final description. They state the pubis has a marked pubic tubercle, unlike the holotype. This indicates Senter et al. included UMNH VP 21751 and 21752 in the hypodigm, though they later (2012) described these as a velociraptorine instead. As the supposed lacrimal was stated to lack an anterodorsal process, it's possible this ended up being the holotype proximal pubis. The identity of the supposed ilium is unknown. Senter et al. (2012) found Yurgovuchia to be a dromaeosaurine more derived than Deinonychus in their analysis, using a modified form of Senter's TWiG-based matrix.
References- Senter, Kirkland, Deblieux and Madsen, 2010. Three new theropods from the Cedar Mountain Formation (Lower Cretaceous) of Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 162A.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.

Dromaeosaurinae Matthew and Brown, 1922
Definition- (Dromaeosaurus albertensis <- Velociraptor mongoliensis) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017; modified from Sereno, 1998)
Other definitions- (Dromaeosaurus albertensis <- Microraptor zhaoianus, Velociraptor mongoliensis, Unenlagia comahuensis, Passer domesticus) (Turner et al., 2012)
= Dromaeosaurinae sensu Turner et al., 2012
Definition- (Dromaeosaurus albertensis <- Microraptor zhaoianus, Velociraptor mongoliensis, Unenlagia comahuensis, Passer domesticus)
Comments- The comments for Velociraptorinae apply here as well, though I consider Passer even less necessary in Turner et al.'s definition.
References- Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399.

Dromaeosaurinae indet. (Gangloff, 1998)
Albian-Cenomanian, Early-Late Cretaceous
Chandler Formation, Alaska, US
Material- (AK211-V-001) tooth
Comments- This was originally referred to Alectrosaurus (Gangloff, 1998), and later to Dromaeosaurus albertensis (Fiorillo and Gangloff, 2000) based on strong lateral compression and a lack of blood grooves. However, it seems to early to belong to D. albertensis itself and is probably another genus of dromaeosaurine.
References- Gangloff, 1998. Arctic dinosaurs with emphasis on the Cretaceous record of Alaska and the Eurasian-North American connection. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 211-220.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation (Cretaceous) of northern Alaska, with speculations on Arctic dinosaur paleoecology. Journal of Vertebrate Paleontology. 20(4), 675-682.

Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Late Cenomanian, Late Cretaceous
Dakota Formation, Utah, US
Material- teeth
Comments- These remains were listed as Dromaeosaurinae indet. by Kirkland et al. (1998) and Eaton et al. (1999). They are presumably one of the two Dromaeosauridae indet. gen. et sp. teeth listed by Kirkland et al. (1997).
References- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau: a key to understanding 35 million years of tectonics, sedimentology, evolution, and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Eaton, Cifelli, Hutchison, Kirkland and Parrish, 1999. Cretaceous vertebrate faunas from the Kaiparowits Plateau, south central Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 345-353.

Dromaeosaurinae indet. (Fiorillo, 1999)
Cenomanian-Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71598) teeth and fragments (Fiorillo, 1999)
partial tooth (Garrison et al., 2007)
Comments- Fiorillo (1999) notes laterally compressed teeth which possess labiolingually broad serrations which are not hooked apically. He refers these to Dromaeosaurinae. Garrison et al. (2007) describe and illustrate a partial tooth with flat serrations (6/mm) on the distal carinae with prominent blood pits, which they refer to Dromaeosaurinae indet.. Cifelli et al. (1999) also list Dromaeosaurinae indet. teeth.
References- Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial Cretaceous vertebrates from the Cedar Mountain Formation, Emery County, Utah: the Mussentuchit Local Fauna. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison Eggshell site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 259-268.
Garrison, Brinkman, Nichols, Layer, Burge and Thayn, 2007. A multidisciplinary study of the Lower Cretaceous Cedar Mountain Formation, Mussentuchit Wash, Utah: a determination of the paleoenvironment and paleoecology of the Eolambia caroljonesa dinosaur quarry. Cretaceous Research. 28, 461-494.

Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Middle-Late Turonian, Late Cretaceous
Smoky Hollow Member of the Straight Cliffs Formation, Utah, US
Material- (OMNH 23712) tooth (Parrish, 1999)
(OMNH 25422) tooth (Parrish, 1999)
(OMNH 24437) tooth (Parrish, 1999)
(OMNH 24438) tooth (Parrish, 1999)
(OMNH 25420) tooth (Parrish, 1999)
Comments- These were listed as Dromaeosaurinae by Kirkland et al. (1998) and Parrish (1999).
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.

Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah, US
Material- (MNA NB-9) tooth (Parrish, 1999)
(OMNH 21222, 21318, 21230, 21281, 21335, 21340, 21395, 21868, 21991, 23636, 24307, 24308, 24327) thirteen teeth (Parrish, 1999)
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

Dromaeosaurinae indet. (Parrish, 1999)
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Material- (UCM 83240) tooth (Parrish, 1999)
(UCM 8659; in part) tooth (Parrish, 1999)
(OMNH 21117, 23178, 23527, 23595, 23608, 23854, 23875, 23969-23975, 24157, 24159, 24160, 23851, 23565 (in part), 24381) nineteen teeth (Parrish, 1999)
Reference- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.

unnamed dromaeosaurine (Rauhut and Zinke, 1995)
Barremian, Early Cretaceous
Una Formation, Spain
Material- (IPFUB Una Th 37, 39, 48, 49, 67, 71-80) thirty-seven teeth (<5 mm)
Comments- These teeth are moderately to strongly recurved, and some have a very slight basal constriction. Most specimens have bot mesial and distal serrations, with the mesial serrations restricted apically. Some have a reduced number of distal serrations, and only weakly developed mesial serrations. One lacks serrations entirely. The DSDI varies between .86 and 1.17, with an average of 1.03. Serrations are chisel-shaped. The mesial carina twists lingually in the basal half.
References- Rauhut and Zunke, 1995. A description of the Barremian dinosaur fauna from Una with a comparison of that of Las Hoyas. II. International Symposium of Lithographic Limestone, Extended Abstracts. 123-126.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca, Spain. Cretaceous Research. 23, 255-263.

undescribed Dromaeosaurinae (Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- teeth
Comments- Ortega et al. (2009) state teeth of Dromaeosaurinae are present.
Reference- Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009. The vertebrate diversity of the Upper Campanian-Lower Maastrichtian "Lo Hueco" fossil-site (Cuenca, Spain). Journal of Vertebrate Paleontology. 29(3), 159A-160A.

undescribed Dromaeosaurinae (Averianov, 2007)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- teeth
Comments- Averianov states these are similar to dromaeosaurine teeth from the Bostobe Formation (ZIN PH 27/49 - 31/49). They probably belong to Itemirus.
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.

unnamed Dromaeosaurinae (Nessov and Khisorova, 1988)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- (ZIN PH 27/49) tooth (Averianov, 2007)
(ZIN PH 28/49) tooth (Averianov, 2007)
(ZIN PH 29/49) tooth (Averianov, 2007)
(ZIN PH 30/49) tooth (Averianov, 2007)
(ZIN PH 31/49) tooth (Averianov, 2007)
(ZIN PH 33/49) (juvenile) tooth (FABL 3.1 mm) (Averianov, 2007)
two teeth (Dyke and Malakhov, 2004)
Comments- ZIN PH 27/49 - 31/49 have FABLs between 9.1 and 16.7 mm, BW/FABLs of .54-.86, lingually shifted mesial carinae, 14 mesial serrations per 5 mm and 15-17 distal serrations per 5 mm. Those described by Dyke and Malakhov (2004) as dromaeosaurids were said to be similar. They are probably the same specimens that were mentioned as dromaeosaurids by Nessov and Khisorova (1988) and Nessov (1995). Averianov (2007) noted they had low DSDIs for dromaeosaurids and considered the possibility they are anterior maxillary/dentary teeth from juvenile tyrannosauroids.
ZIN PH 33/49 has a BW/FABL of .48, a lingually displaced mesial carina which may have originally had small serrations, 6 distal serrations per mm, wrinkled enamel and two vertical ridges lingually. Averianov referred it to Dromaeosauridae based on its small serrations, and to Dromaeosaurinae based on its relative thickness and lingually displaced mesial carina. He viewed Paronychodon as juvenile dromaeosaurid teeth, with this specimen as an intemediate age.
References- Nessov and Khisorova, 1988. [New data on vertebrates of the Late Cretaceous in the localities Shakh-Shakh and Baibolat (North-Eastern Aral Region)]. Matyerialy po istorii fauny i flory Kazakhstana. 10, 5-14.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Dyke and Malakhov, 2004. Abundance and taphonomy of dinosaur teeth and other vertebrate remains from the Bostobynskaya Formation, north-east Aral Sea region, Republic of Kazakhstan. Cretaceous Research.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.

unnamed dromaeosaurine (Alifanov and Bolotsky, 2002)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- (1/314) tooth (19x7.5x2.9 mm) (Bolotsky, 2011)
(1/806) tooth (20.2x9.9x3.2 mm) (Bolotsky, 2011)
(1/1079) tooth (23.2x8.8x5.8 mm) (Bolotsky, 2011)
(1/1080) tooth (18x8.8x1.1 mm) (Bolotsky, 2011)
Comments- Alifanov and Bolotsky (2002) referred these remains to cf. Dromaeosaurus sp., while Bolotsky (2011) described them as Dromaeosaurinae.
Reference- Alifanov and Bolotsky, 2002. New data about the assemblages of the Upper Cretaceous carnivorous dinosaurs (Theropoda) from the Amur region. In Kirillova (ed.). Fourth International Symposium of IGCP 434. Cretaceous continental margin of East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
Bolotsky, 2011. On paleoecology of carnivorous dinosaurs (Tyrannosauridae, Dromaeosauridae) from Late Cretaceous fossil deposits of Amur region, Russian far East. Global Geology. 14(1), 1-6.

unnamed possible dromaeosaurine (Maisch and Matzke, 2020)
Oxfordian, Late Jurassic
Qigu Formation, Xinjiang, China
Material- (SGP 2002/6) tooth (10.3x5.15x3.3 mm)
(SGP 2002/7) tooth (9.0x4.8x2.5 mm)
Comments- Maisch and Matzke (2020) refer these to ?Dromaeosauridae indet. based largely on the twisted mesial carina.
Reference- Maisch and Matzke, 2020. Small theropod teeth (Dinosauria) from the Upper Jurassic Qigu Formation of the southern Junggar Basin, NW China. Neues Jahrbuch für Geologie und Paläontologie Abhandlungen. 295(1), 91-100.

unnamed possible dromaeosaurine (Han, Clark, Xu, Sullivan, Choiniere and Hone, 2011)
Late Oxfordian, Late Jurassic
Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Material- (IVPP V15857) tooth (7.1x3.7x2.1 mm)
Comments- Discovered by the Sino-American expeditions between 2001 and 2010, this is called Morphotype 5 by Han et al. (2011), who refer it to a dromaeosaurid. It is placed in that clade here based on the twisted mesial carina.
Reference- Han, Clark, Xu, Sullivan, Choiniere and Hone, 2011. Theropod teeth from the Middle-Upper Jurassic Shishugou Formation of northwest Xinjiang, China. Journal of Vertebrate Paleontology. 31(1), 111-126.

undescribed dromaeosaurine (Watanabe and Sereno, 2010)
Cenomanian, Late Cretaceous
Ulansuhai Formation, Inner Mongolia, China
Material- (juvenile or subadult) premaxilla, maxilla, lacrimal, frontal, postorbital, quadrate, ectopterygoid, atlantal intercentrum, mid dorsal neural arch, ischium
Diagnosis- (after Watanabe and Sereno, 2010) deep sagittal crest extending onto frontals; hourglass-shaped hyposphene-hypantrum articulation in dorsal vertebrae.
Comments- This is described as having a short snout and deep obturator process. It was found to group with Dromaeosaurus, Utahraptor and Achillobator.
Reference- Watanabe and Sereno, 2010. A large short-snouted dromaeosaurid (Theropoda: Maniraptora) from Inner Mongolia. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 184A.

undescribed possible dromaeosaurine (Currie and Eberth, 1993)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- teeth
Comments- Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in the Iren Dabasu" and that "Most of these can be attributed to Velociraptor, although some of the teeth suggest that there was a second, larger species of an indeterminate dromaeosaurine dromaeosaurid." A pedal phalanx II-1 (AMNH 6572) mentioned by Ostrom (1969) as being 20% larger than Deinonychus may belong to the same taxon, based on size.
References- Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin. 30, 1-165.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic of China. Cretaceous Research. 14, 127-144.

unnamed possible dromaeosaurine (Goodwin, Clemens, Hutchinson, Wood, Zavada, Kemp, Duffin and Schaff, 1999)
Tithonian, Late Jurassic
Mugher Mudstone, Ethiopia
Material- (JN-96-2B/UCMP 170803) tooth
several teeth (Hall and Goodwin, 2011)
Comments- JN-96-2B/UCMP 170803 is slightly recurved, with a FABL of 9.3 mm and a BW of 7.3 mm. It has both mesial and distal serrations, but no further details are given or visible in the photograph. The authors state the morphology "compares well with the dentary teeth of Dromaeosaurus albertensis" and describe it as having possible dromaeosaurine affinity. The lack of specific similarities and wide temporal and spatial gulf between this specimen and verified dromaeosaurines makes the assignment questionable.
References- Goodwin, Clemens, Hutchinson, Wood, Zavada, Kemp, Duffin and Schaff, 1999. Mesozoic continental vertebrates with associated palynostratigraphic dates from the northwestern Ethiopian Plateau. Journal of Vertebrate Paleontology. 19(4), 728-741.
Hall and Goodwin, 2011. A diverse dinosaur tooth assemblage from the Upper Jurassic of Ethiopia: Implications for Gondwanan dinosaur biogeography. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 121.

Atrociraptor Currie and Varricchio, 2004
A. marshalli Currie and Varricchio, 2004
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
Holotype- (RTMP 95.166.1) (~1.7 m) (skull ~210 mm) premaxillae, maxilla, dentaries (one partial), ?third premaxillary tooth (10x5x3.5 mm); anterior maxillary tooth (12x5.5x3.5 mm), anterior dentary tooth (?x5.2x2.8 mm), posterior dentary tooth (7x5.2x2.6 mm), fragments
Referred- ?(AMNH coll.) tooth (Bell, 2007)
(RTMP 65.16.150) tooth (7.2x4.7x2.6 mm) (Larson and Currie, 2013)
(RTMP 84.64.3) tooth (5.5x4.3x1.9 mm) (Larson and Currie, 2013)
(RTMP 84.64.12) tooth (7.3x4.5x2.3 mm) (Larson and Currie, 2013)
(RTMP 84.79.4) tooth (8x5.6x2.6 mm) (Larson and Currie, 2013)
(RTMP 85.12.2) tooth (5.1x3.2x1.7 mm) (Larson and Currie, 2013)
(RTMP 85.98.2) tooth (9.5x5.5x2.6 mm) (Larson and Currie, 2013)
(RTMP 86.64.2) tooth (4.1x3.2x1.6 mm) (Larson and Currie, 2013)
(RTMP 86.64.3) tooth (9.6x5.1x2.3 mm) (Larson and Currie, 2013)
(RTMP 90.82.17) tooth (7.6x4.7x2.4 mm) (Larson and Currie, 2013)
(RTMP 90.82.18) tooth (9.1x5.8x2.3 mm) (Larson and Currie, 2013)
(RTMP 90.82.21) tooth (6.6x4.7x2.4 mm) (Larson and Currie, 2013)
(RTMP 90.82.108) tooth (9.4x5.2x2.8 mm) (Larson and Currie, 2013)
(RTMP 93.12.21) tooth (7.7x4.4x2.5 mm) (Larson and Currie, 2013)
?(RTMP 97.39.4) tooth (Ryan et al., 2000)
(RTMP 98.63.4) tooth (Larson et al., 2010)
(RTMP 99.50.117) tooth (6.4x4.1x1.3 mm) (Larson et al., 2010)
?(RTMP 1005-1008, 1033, 1034) six teeth (Baszio, 1997)
?(RTMP 1009) premaxillary tooth (Baszio, 1997)
(RTMP 2000.45.35) tooth (5.8x3.9x2.1 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.40) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.103) tooth (10.2x4.3x2.1 mm) (Larson et al., 2010)
(RTMP 2003.45.49) tooth (6.6x4.2x2.2 mm) (Larson et al., 2010)
(RTMP 2003.45.52) tooth (7.1x4.7x? mm) (Larson et al., 2010)
(RTMP 2003.45.60) tooth (Larson et al., 2010)
(RTMP 2005.7.3) tooth (8.4x5.2x2.4 mm) (Larson and Currie, 2013)
?(RTMP coll.) seven teeth (Ryan et al., 2000)
(RTMP or UALVP coll.) (unassociated) five teeth (one uncollected) (Eberth and Currie, 2010)
?(RTMP or UALVP coll.) (unassociated) manual phalanx, manual ungual (Eberth and Currie, 2010)
?(RTMP coll.; lost) tooth (Torices et al., 2014)
?(UALVP 52075) manual ungual III (Eberth and Currie, 2010)
?(UALVP 55571) tooth (Torices et al., 2014)
? several premaxillary teeth (Baszio, 1997)
Diagnosis- (after Currie and Varricchio, 2004) snout short and deep; premaxilla taller than long (also in Deinonychus); subnarial and nasal processes of premaxilla inclined more than 45 degrees dorsally; large maxillary fenestra; maxillary fenestra directly dorsal to promaxillary fenestra; maxillary teeth isodont.
(after Larson et al., 2010) differs from Saurornitholestes langstoni in- generally larger basal width and serration size; lower crown height compared to FABL.
Other diagnoses- Currie and Varricchio (2004) also listed highly posteriorly inclined maxillary teeth in their diagnosis, supposedly shared with Bambiraptor and Deinonychus, but Senter (2010) showed this was due to taphonomy.
Comments- Discovered in 1995, Atrociraptor was announced in an abstract by Currie and Varricchio (2000), then named and fully described by Currie and Varricchio (2004). Powers et al. (2022) describes the specimen in more detail using CT scanning.
Baszio (1997) noted Horseshoe Canyon teeth he referred to Saurornitholestes sp. differ from Dinosaur Park specimens in being smaller with less variable serration shape. Ryan et al. (1997) listed seven teeth from this formation as Saurornitholestes sp.. Ryan et al. (2000) later described these teeth as cf. Saurornitholestes sp., as they noted the then unnamed Atrociraptor had teeth nearly identical to Saurornitholestes'. Larson and Currie (2013) found that Atrociraptor teeth only partially overlapped Saurornitholestes teeth in morphometric analyses. These teeth are all provisionally referred to Atrociraptor here, as no definite Saurornitholestes specimens have been found in this formation. Eberth and Currie (2010) refer three manual elements to Atrociraptor, but these cannot be compared to the type material and may belong to the dromaeosaurine or Richardoestesia also present in the bonebed, if not the troodontids.
RTMP 1009 was figured by Baszio as a Richardoestesia Paronychodon-like tooth. It has three faint ridges, is moderately recurved, and has a high DSDI (10 mesial serrations per mm vs. 5.7 distal serrations per mm). Distal serrations are taller than apicobasally wide and rounded, while mesial serrations are shorter than wide. Though the ridges and distal serration shape are similar to Saurornitholestes premaxillary teeth, it differs in the curvature, fewer and weaker ridges, small serrations, and high DSDI. The basal constriction, serration size, DSDI and curvature are similar to Richardoestesia gilmorei, but the ridges and serration shape differ. It may be a posterior premaxillary tooth, which have fewer and weaker grooves than anterior teeth in Saurornitholestes. However, Currie and Evans (2020) find in the holotype's second premaxillary tooth "there do not appear to be any apicobasal longitudinal ridges and flutes. This could be a function of wear and/or poor preservation, however." Whether this means Saurornitholestes and Atrociraptor coexisted in the Horseshoe Canyon Formation, Atrociraptor exhibited variation in dental ridges, or taphonomy is responsible is uncertain.
References- Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan, Currie, Gardner and Livigne, 1997. Baby hadrosaurid material associated with an unusually high abundance of troodontid teeth from the Horseshoe Canyon Formation (Early Maastrichtian), Alberta, Canada. Journal of Vertebrate Paleontology. 17(3), 72A.
Currie and Varricchio, 2000. New dromaeosaurids from the Late Cretaceous of western North America. The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History. 14.
Ryan, Currie, Gardner, Vickaryous and Lavigne, 2000. Baby hadrosaurid material associated with an unusually high abundance of Troodon teeth from the Horseshoe Canyon Formation, Upper Cretaceous, Alberta, Canada. Gaia. 15, 123-133.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp. 279-297.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons. Studies on the transition from dinosaurs to birds. Indiana University Press. 112-132.
Bell, 2007. The Danek bonebed: An unusual dinosaur assemblage from the Horseshoe Canyon Formation, Edmonton, Alberta. Journal of Vertebrate Paleontology. 27(3), 46A.
Eberth and Currie, 2010. Stratigraphy, sedimentology, and taphonomy of the Albertosaurus bonebed (upper Horseshoe Canyon Formation; Maastrichtian), southern Alberta, Canada. Canadian Journal of Earth Sciences. 47(9), 1119-1143.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta, with special reference to the Albertosaurus sarcophagus bonebed. Canadian Journal of Earth Sciences. 47(9), 1159-1181.
Senter, 2010. Using creation science to demonstrate evolution: Application of a creationist method for visualizing gaps in the fossil record to a phylogenetic study of coelurosaurian dinosaurs. Journal of Evolutionary Biology. 23(8), 1732-1743.
Larson and Currie, 2013. Multivariate analyses of small theropod dinosaur teeth and implications for paleoecological turnover through time. PloS ONE. 8(1), e54329.
Torices, Funston, Kraichy and Currie, 2014. The first appearance of Troodon in the Upper Cretaceous site of Danek Bonebed, and a reevaluation of troodontid quantitative tooth morphotypes. Canadian Journal of Earth Sciences. 51, 1039-1044.
Currie and Evans, 2020 (online 2019). Cranial anatomy of new specimens of Saurornitholestes langstoni (Dinosauria, Theropoda, Dromaeosauridae) from the Dinosaur Park Formation (Campanian) of Alberta. The Anatomical Record. 303(4), 691-715.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as 2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters. Journal of Vertebrate Paleontology. 41(5), 2010087.

Saurornitholestiinae Longrich and Currie, 2009
Definition- (Saurornitholestes langstoni <- Dromaeosaurus albertensis, Velociraptor mongoliensis, Microraptor zhaoianus) (Longrich and Currie, 2009)
Saurornitholestes Sues, 1978
= Zapsalis Cope, 1876b
S. langstoni Sues, 1978
= Laelaps explanatus Cope, 1876a
= Laelaps laevifrons Cope, 1876b
= Zapsalis abradens Cope, 1876b
= Dryptosaurus explanatus (Cope, 1876a) Hay, 1902
= Dryptosaurus laevifrons (Cope, 1876b) Hay, 1902
= Deinodon explanatus (Cope, 1876a) Lambe, 1902
= Deinodon laevifrons (Cope, 1876b) Osborn, 1902
= Aublysodon explanatus (Cope, 1876a) Hatcher, 1903
= Dromaeosaurus laevifrons (Cope, 1876b) Matthew and Brown, 1922
= Dromaeosaurus explanatus (Cope, 1876a) Kuhn, 1939
= Dromaeosaurus abradens (Cope, 1876b) Matthew and Brown, 1922 vide Kuhn, 1939
= Velociraptor langstoni (Sues, 1978) Paul, 1988
= Paronychodon explanatus (Cope, 1876a) Olshevsky, 1995
= Saurornitholestes explanatus (Cope, 1876a) Martyniuk, 2012
Diagnosis- (after Sullivan, 2006) ratio of the length (measured along the midline) to the thickness (posterior part of the frontal) is 10:1.
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
Holotype- (RTMP 74.10.5) maxillary tooth (9.2 mm), lacrimal, jugal, postorbital, frontals, quadrate, ectopterygoid, angular, anterior dentary tooth (8.9 mm), two vertebrae, dorsal rib fragments, gastralia, caudal prezygopophyses, distal phalanx I-1, manual ungual I, distal phalanx II-2, manual ungual II, distal metacarpal III, phalanx III-1 (17 mm), distal phalanx III-3, manual ungual III
Paratypes- (CMN 12343) frontal
(CMN 12354) frontal
(UA 5283) posterior frontal
Referred- (CMN 2664) second or third premaxillary tooth (Currie, Rigby and Sloan, 1990)
(ROM 57164) incomplete frontal (Currie and Evans, 2020)
(RTMP 67.20.36) last dorsal vertebra, sacrum (Norell and Makovicky, 1997)
(RTMP 70.37.1) first premaxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 78.9.96) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 79.8.643) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 79.15.3) second premaxillary tooth (11.8x6.4x? mm) (Currie, Rigby and Sloan, 1990)
(RTMP 80.16.312) frontal (Currie, 1987)
(RTMP 80.16.833) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 80.16.996) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.14.34) cervical vertebra (Rauhut, 2003)
(RTMP 81.20.259) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.23.7) frontal (Currie, 1987)
(RTMP 81.26.37) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.14.45) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.16.43) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.7) premaxillary tooth (15 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.180) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.336) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.458) premaxillary tooth (12.5 mm), tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.24.16) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.26.1) ribs, incomplete caudal series, distal tarsal, metatarsal II, pedal ungual II (distal caudal vertebra 31.1 mm) (Currie, 1995; Makovicky, 1995)
(RTMP 82.112.10) tooth (8.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.91.39) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.91.40) premaxillary tooth (12.5 mm), tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.94.6) frontal (Currie, 1987)
(RTMP 84.163.80) premaxillary tooth (10 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.2) premaxillary tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.131) premaxillary tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.132) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.133) premaxillary tooth (~11.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.43.69) frontal (Currie, 1987)
(RTMP 85.56.48) (adult) posterior cervical vertebra (Makovicky, 1995)
(RTMP 85.58.65) premaxillary tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.9.92) premaxillary tooth (~14.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.23.104) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.36.117) incomplete premaxilla (Currie, Rigby and Sloan, 1990)
(RTMP 86.36.425) premaxillary tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
?(RTMP 86.77.2) ischium (Hutchinson, 2001)
(RTMP 86.77.57) frontal (Currie, 1987)
(RTMP 86.77.112) premaxillary tooth (~14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.78.1) basioccipital (Rauhut, 2003)
(RTMP 86.130.219) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.198.46) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.4.47) tooth (4.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.19.68) tooth (4.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.31.14) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.31.52) tooth (12.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.31.54) tooth (4.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.33.55) tooth (9.9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.5) premaxillary tooth (15.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.11) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.68) tooth (~11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.70) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.184) tooth (7.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.300) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.383) premaxillary tooth (Baszio, 1997)
(RTMP 87.36.392) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.418) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.39.93) tooth (11.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.43.5) tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.46.53) tooth (9.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.48.77) tooth (6.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.48.86) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.50.8) second premaxillary tooth (13.3x6.8x? mm) (Currie and Evans, 2020)
(RTMP 87.50.38) tooth (14.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.50.100) tooth (8.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.51.23) tooth (8.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.62.87) tooth ( mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.72.4) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.72.23) tooth (~9.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.72.26) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.120) tooth (~11.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.125) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.126) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.79.89) tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.79.90) tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.83.1) tooth (12.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.112.9) tooth (8.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.153.12) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.153.55) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.62) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.63) tooth (8.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.158.78) tooth (4.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.158.80) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.158.81) tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.112.21) tooth (5.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.112.28) tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.64) tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.50.123) tooth (Baszio, 1997)
(RTMP 88.50.126) tooth (Baszio, 1997)
(RTMP 88.87.89) tooth (Baszio, 1997)
(RTMP 88.116.42) tooth (Baszio, 1997)
(RTMP 88.121.39) (128 kg) quadratojugal, quadrate, mandible (dentary 120 mm), teeth (6.2-12.3 mm), three dorsal ribs, gastralia, distal caudal series, scapula (~155 mm), coracoid (~65 mm prox-dist), humerus (~177 mm), femur (214 mm), tibia (283 mm), fibula, astragalus, calcaneum, metatarsal II (101 mm), phalanx II-2 (31 mm), pedal ungual II (69 mm straight), metatarsal III (115 mm), metatarsal IV (107 mm) (Currie, Rigby and Sloan, 1990; Currie and Varricchio in prep.)
(RTMP 88.162.1) tooth (Baszio, 1997)
(RTMP 88.215.72) tooth (Baszio, 1997)
(RTMP 89.36.312) premaxillary tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.353) tooth (Baszio, 1997)
(RTMP 89.36.400) tooth (Baszio, 1997)
(RTMP 89.36.403) tooth (Baszio, 1997)
(RTMP 89.36.410) tooth (Baszio, 1997)
(RTMP 89.50.8) second premaxillary tooth (13.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.50.202) premaxillary tooth (9.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.6.15) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.50.208) premaxillary tooth (~11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.53.21) premaxillary tooth (~11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 91.36.112) dentary, tooth (Rauhut, 2003)
(RTMP 91.50.60) premaxillary tooth (14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 92.36.129) sacrum (Rauhut, 2003)
(RTMP 92.36.333) sternal plate (Godfrey and Currie, 2004)
(RTMP 92.50.23) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 92.83.2) tooth (Currie and Jacobsen, 1995)
(RTMP 93.36.98) tarsal (Rauhut, 2003)
(RTMP 94.12.268) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.844 maxilla (Currie and Varricchio, 2004)
(RTMP 95.2.18) tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.31) tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.33) tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.36) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.38) tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.40) tooth (7.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.42) tooth (7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.43) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.74) tooth (10.2 mm), tooth (6.3 mm), tooth (9.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.109) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.19.4) tooth (8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.21.5) tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.16) tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.25) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.27) tooth (6.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.28) tooth (~5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.54) tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.124.4) tooth (6.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.126.29) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.127.25) tooth (11.8 mm), tooth (~7.8 mm), tooth (6.7 mm), tooth (7.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.129.2) tooth (9.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.131.12) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.137.2) tooth (9.2 mm), tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.147.26) tooth (2.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.151.10) tooth (3.9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.177.48) tooth (6 mm), tooth (4.7 mm), tooth (~3.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.179.3) tooth (10.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.180.4) tooth (~3.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.181.11) tooth (~5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.182.21) tooth (~7.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.184.23) tooth (8.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.406.5) tooth (10.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.34) tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.38) tooth (11.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.102) tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.104) tooth (10.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.112) tooth (16 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.115) tooth (14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.118) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.359) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.360) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.361) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.363) tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.364) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.366) tooth (6.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.422) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 97.80.36) tooth (3.9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 97.133.2) premaxillary tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP coll.; lost) premaxillary tooth (Baszio, 1997)
(UALVP 55425) surangular, articular (Currie and Evans, 2020)
(UALVP 55700) (~2 m, 8 year old subadult) skull (~210 mm), sclerotic plates, mandibles, hyoids, axis, third cervical vertebra, fourth-tenth cervical vertebrae, cervical ribs, dorsal series, dorsal ribs, uncinate processes, gastralia, sacrum, incomplete caudal series, chevrons, scapulae, coracoids, furcula, sternum, sternal ribs, forelimbs missing a few phalanges, pelves, hindlimbs missing a few phalanges (femur ~228 mm), pedal ungual II claw sheath (Currie and Evans, 2015)
thirty premaxillary teeth (Baszio, 1997)
Late Campanian, Late Cretaceous
Oldman or Dinosaur Park Formation of the Belly River Group, Alberta, Canada
(NHMUK R4463) sacrum (Howse and Milner, 1993)
tooth (12 mm) (Lambe, 1902)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
?(CMN 12071; = CMN 12072 in Colbert and Russell?) metatarsal IV (Colbert and Russell, 1969)
?(CMN 12240) pedal ungual II (Colbert and Russell, 1969)
(CMN 12410) tooth (CMN online)
(RTMP 94.144.105) tooth (Ryan and Russell, 2001)
(UA 12091) dentary (Sues, 1977)
(UA 12339) dentary (Sues, 1977)
Middle Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
(RTMP 88.86.29) tooth (Ryan and Russell, 2001)
(RTMP 96.62 coll.) twenty-seven teeth (Peng, Russell and Brinkman, 2001)
(RTMP 2009.37.109) tooth (Cullen, Fanti, Capobianco, Ryan and Evans, 2016)
(RTMP coll.) three teeth (Cullen, Fanti, Capobianco, Ryan and Evans, 2016)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(AMNH 3953; holotype of Zapsalis abradens) second premaxillary tooth (12 mm) (Cope, 1876b)
(AMNH 3958; holotype of Laelaps explanatus) tooth (11 mm), twenty-five teeth (Cope, 1876a)
(AMNH 3961; holotype of Laelaps laevifrons) tooth (15 mm) (Cope, 1876b)
(AMNH 8518) tooth (Sahni, 1972)
(ANSP 15808) tooth (Fiorillo and Currie, 1994)
(ANSP 15810) tooth (Fiorillo and Currie, 1994)
(ANSP 15811) tooth (Fiorillo and Currie, 1994)
(ANSP 15814) tooth (Fiorillo and Currie, 1994)
(ANSP 15815) tooth (Fiorillo and Currie, 1994)
(ANSP 15817) tooth (Fiorillo and Currie, 1994)
(ANSP 15818) tooth (Fiorillo and Currie, 1994)
(ANSP 15819) tooth (Fiorillo and Currie, 1994)
(ANSP 15823) tooth (Fiorillo and Currie, 1994)
(ANSP 15824) tooth (Fiorillo and Currie, 1994)
(ANSP 15825) tooth (Fiorillo and Currie, 1994)
(ANSP 15826) tooth (Fiorillo and Currie, 1994)
(ANSP 15828) tooth (Fiorillo and Currie, 1994)
(ANSP 15829) tooth (Fiorillo and Currie, 1994)
(ANSP 15831) tooth (Fiorillo and Currie, 1994)
(ANSP 15833) tooth (Fiorillo and Currie, 1994)
(ANSP 15838) tooth (Fiorillo and Currie, 1994)
(ANSP 15939) tooth (Fiorillo and Currie, 1994)
(ANSP 15943) tooth (Fiorillo and Currie, 1994)
(ANSP 15945) tooth (Fiorillo and Currie, 1994)
(ANSP 15946) tooth (Fiorillo and Currie, 1994)
(ANSP 15948) tooth (Fiorillo and Currie, 1994)
(ANSP 15949) tooth (Fiorillo and Currie, 1994)
(ANSP 15951) tooth (Fiorillo and Currie, 1994)
(ANSP 15953) tooth (Fiorillo and Currie, 1994)
(ANSP 15954) tooth (Fiorillo and Currie, 1994)
(ANSP 15955) tooth (Fiorillo and Currie, 1994)
(ANSP 15956) tooth (Fiorillo and Currie, 1994)
(ANSP 15957) tooth (Fiorillo and Currie, 1994)
(ANSP 15958) tooth (Fiorillo and Currie, 1994)
(ANSP 15960) tooth (Fiorillo and Currie, 1994)
(ANSP 15975) tooth (Fiorillo and Currie, 1994)
(ANSP 16195) tooth (Fiorillo and Currie, 1994)
(ANSP 16197) tooth (Fiorillo and Currie, 1994)
(ANSP 16198) tooth (Fiorillo and Currie, 1994)
(ANSP 16199) tooth (Fiorillo and Currie, 1994)
(ANSP 16225) tooth (Fiorillo and Currie, 1994)
(ANSP 16226) tooth (Fiorillo and Currie, 1994)
(ANSP 16227) tooth (Fiorillo and Currie, 1994)
(ANSP 16228) tooth (Fiorillo and Currie, 1994)
(ANSP 16229) tooth (Fiorillo and Currie, 1994)
(ANSP 17646) tooth (Fiorillo and Currie, 1994)
(ANSP 17648) tooth (Fiorillo and Currie, 1994)
(ANSP 17649) tooth (Fiorillo and Currie, 1994)
(ANSP 17773) tooth (Fiorillo and Currie, 1994)
(ANSP 17774) tooth (Fiorillo and Currie, 1994)
(ANSP 17775) tooth (Fiorillo and Currie, 1994)
(ANSP 17776) tooth (Fiorillo and Currie, 1994)
(ANSP 17777) tooth (Fiorillo and Currie, 1994)
(ANSP 17778) tooth (Fiorillo and Currie, 1994)
(ANSP 17781) tooth (Fiorillo and Currie, 1994)
(ANSP 17783) tooth (Fiorillo and Currie, 1994)
(ANSP 17785) tooth (Fiorillo and Currie, 1994)
(ANSP 17786) tooth (Fiorillo and Currie, 1994)
(ANSP 17787) tooth (Fiorillo and Currie, 1994)
(ANSP 17788) tooth (Fiorillo and Currie, 1994)
(ANSP 17789) tooth (Fiorillo and Currie, 1994)
(ANSP 17801) tooth (Fiorillo and Currie, 1994)
(ANSP 17802) tooth (Fiorillo and Currie, 1994)
(ANSP 17803) tooth (Fiorillo and Currie, 1994)
(ANSP 17962) tooth (Fiorillo and Currie, 1994)
(ANSP 17963) tooth (Fiorillo and Currie, 1994)
(ANSP 18002) tooth (Fiorillo and Currie, 1994)
(ANSP 18006) tooth (Fiorillo and Currie, 1994)
(ANSP 18100) tooth (Fiorillo and Currie, 1994)
(ANSP 18103) tooth (Fiorillo and Currie, 1994)
(ANSP 18105) tooth (Fiorillo and Currie, 1994)
(ANSP 18109) tooth (Fiorillo and Currie, 1994)
(ANSP 18112) tooth (Fiorillo and Currie, 1994)
(ANSP 18118) tooth (Fiorillo and Currie, 1994)
(ANSP 18122) tooth (Fiorillo and Currie, 1994)
(BDM 005) frontal (Wilson and Fowler, 2020)
(MOR 125) phalanx (MOR online)
(UCMP 137559) tooth (UCMP online)
(UCMP 137560) tooth (UCMP online)
(UCMP 150584) tooth (UCMP online)
(UCMP 150585) teeth (UCMP online)
(UCMP 150586) teeth (UCMP online)
(UCMP 150587) teeth (UCMP online)
(UCMP 150588) ungual fragment (UCMP online)
(YPM PU 24968) (YPM online)
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
(MOR 484) ungual (MOR online)
(MOR 553E-6-26-91-104) metatarsal IV (MOR online)
(MOR 553S-7-10-9-197) pedal phalanx IV-4 (MOR online)
(MOR 660) (19.5 kg) partial skeleton including nine cervical vertebrae, thirteen dorsal vertebrae, gastralia, sacrum, first caudal vertebra, distal caudal vertebrae, chevrons, humerus (178 mm), radius, ulna (135 mm), metacarpal I (30 mm), phalanx I-1 (61 mm), manual ungual I (55 straight), phalanx II-1 (49 mm), phalanx II-2 (43 mm), manual ungual II (35 mm), metacarpal III (67 mm), phalanx III-1 (13 mm), phalanx III-2 (29 mm), phalanx III-3 (40 mm), ilium, femur (225 mm), tibia (257 or 301 mm), metatarsal II (112 mm), phalanx II-1 (32 mm), phalanx II-2 (30 mm), pedal ungual II (74 mm straight), metatarsal III (136 mm) and metatarsal IV (123 mm), (Britt, 1993)
(MOR 666) tibia (256 mm), fibula (225 mm) (Christiansen, 1999)
(MOR 9753) (juvenile) maxilla (55 mm) (Hohman and Varricchio, 2021)
teeth (Redman, Moore and Varricchio, 2015)
Diagnosis- (after Currie and Evans, 2020) relatively short skull (skull/femur length ratio less than 1.0); large maxillary fenestra that is positioned near the dorsal margin of the antorbital fossa; maxillary sinuses that interconnect with those inside the nasal and lacrimal; enlarged second premaxillary tooth has a flat lingual surface and has longitudinal ridges and grooves on both labial and lingual surfaces.
Comments- Unfortunately, besides the original description of the fragmentary holotype (Sues, 1978), Currie et al.'s (1990) description of teeth (see also Sankey et al., 2002), Makovicky's (1995) as yet unpublished description of vertebrae, and very recent cranial description of Currie and Evans (2020), Saurornitholestes has not received a detailed description and much information must be pieced together from sporadic references throughout the literature. Currie and Varricchio (2004) note both RTMP 88.121.39 and MOR 660 are being studied by them, which had been occurring for over a decade as of that date.
Saurornitholestes was historically assigned to Velociraptorinae, but was recently given its own subfamily outside the Velociraptorinae+Dromaeosaurinae clade (Longrich and Currie, 2009; Currie and Evans, 2020). Brusatte et al. (2014) still recovered it in Velociraptorinae however, and here it is placed in Dromaeosaurinae based on Hartman et al. (2019).
Laelaps explanatus- Cope (1876a) described this tooth as a new species of Laelaps (in which he placed all Judithian theropods). Once Marsh provided the replacement name Dryptosaurus for the preoccupied Laelaps, Hay (1902) moved explanatus to that genus. Lambe (1902) meanwhile referred it to Deinodon, while Hatcher referred it to Aublysodon, both contemporary tyrannosaurids. Matthew and Brown (1922) questionably referred the species to their new genus Dromaeosaurus as Dromaeosaurus(?) sp., probably based on size, for it was the smallest of their 'deinodontids'. Kuhn (1939) formalized this as D. explanatus. Ostrom (1969) noted these have a high DSDI, and assigned the species to Dromaeosauridae. Olshevsky's (1995) referral of the species to Paronychodon is almost certainly incorrect, as it is serrated and lacks grooves.
Of Judith River theropods, only Saurornitholestes and Richardoestesia have a comparable DSDI. However, Richardoestesia is smaller (FABL up to 4.7 vs. 6.6 mm) and has more serrations per mm (6+ vs. 4.5). The compression (BW/FABL = 2.8/6.6 mm = .42) is comparable to Saurornitholestes, and while the measured tooth is lower than most sampled by Sankey et al. (2002), it is still within the range of variation (lower 5%). The short mesial carina, which is only serrated apically, is also common in dromaeosaurids and differs from Dromaeosaurus in lacking a lingual twist. As Saurornitholestes langstoni is common in the equivalent Dinosaur Park Formation further north, explanatus is here referred to that species. It should be noted that the name explanatus has over a century of priority over langstoni though, so would technically have precedence as a senior synonym. This is not followed here because from the limited description, explanatus is indistinguishable from not only Saurornitholestes, but Bambiraptor and Velociraptor as well. It is only referred to Saurornitholestes due to provenance.
Lambe (1902) referred another tooth to this species, from the Belly River Group of Alberta. It is also said to have a high DSDI and seems to be Saurornitholestes as well. It has a FABL of 6 mm and a BW of 3 mm.
Laelaps laevifrons- Cope (1876b) described this tooth as a new species of Laelaps (in which he placed all Judithian theropods). Once Marsh provided the replacement name Dryptosaurus for the preoccupied Laelaps, Hay (1902) moved laevifrons to that genus. Osborn (1902) meanwhile referred it to Deinodon. Matthew and Brown (1922) referred the species to their new genus Dromaeosaurus, probably based on size, for it was the smallest of their 'deinodontids'.
Of Judith River theropod taxa, Zapsalis and Paronychodon can be eliminated due to the lack of grooves, while Richardoestesia can be eliminated due to the low serration count and troodontids due to the high serration count. This leaves tyrannosaurids, Dromaeosaurus and Saurornitholestes as potential candidates. Tooth size is within the range of juvenile tyrannosaurids, in the upper 10% of Dromaeosaurus, and the upper 1% of Saurornitholestes. The lack of mesial serrations is known in Saurornitholestes and juvenile tyrannosaurids, but not Dromaeosaurus. Crown compression (BW of 4 mm and FABL of 7 mm) is comparable to Dromaeosaurus and tyrannosaurids when size is taken into account, though hypothetically possible for a Saurornitholestes that size. Crown elongation is similar to all three taxa when compared to FABL. Crown elongation vs. crown compression is however withion the range of Saurornitholestes and Dromaeosaurus, but not tyrannosaurids (which usually have stouter crowns when they are that thick). The serration density (5/mm) compared to crown compression is comparable to tyrannosaurids, and in the outer range of Dromaeosaurus and Saurornitholestes. Serration density compared to crown elongation is comparable to Saurornitholestes, and outside either Dromaeosaurus or tyrannosaurids. When all components are analyzed together, laevifrons falls out within Saurornitholestes. It is here viewed as a particularily large example of that genus. As Saurornitholestes langstoni is common in the equivalent Dinosaur Park Formation further north, laevifrons is here referred to that species. It should be noted that the name laevifrons has over a century of priority over langstoni though, so would technically have precedence as a senior synonym (though explanatus has even more priority). This is not followed here because from the limited description, laevifrons is indistinguishable from not only Saurornitholestes, but Bambiraptor and Velociraptor as well. It is only referred to Saurornitholestes due to provenance.
Zapsalis- The holotype of Zapsalis abradens (AMNH 3953) is a tooth which is flat lingually, with no mesial serrations and 3 distal serrations per mm (Cope, 1876b). There are three lingual ridges and four labial ones. The length is 12 mm, the FABL 6.5 mm, and the BW 3 mm. It was synonymized with Paronychodon lacustris by Estes (1964) based on its flat lingual side and ridges, which was followed in the literature for decades. Currie et al. (1990) later restricted Paronychodon to the unserrated forms, and believed the serrated ones were growth abnormalities of named genera- Dromaeosaurus, Saurornitholestes and Troodon. Baszio (1997) included these in his Paronychodon discussion, referred to them as Paronychodon-like teeth. Sankey et al. (2002) separated the Paronychodon-like teeth with Dromaeosaurus-like and Saurornitholestes-like serrations as ?Dromaeosaurus morphotype A. In addition to their ridges and flat lingual side, they differ from Dromaeosaurus albertensis in being shorter and labiolingually narrower, and having a straight distal edge. They argued complete D. albertensis tooth rows are known, so the morphology of morphotype A teeth is not due to positional variation. Furthermore, they have a different stratigraphic distribution than D. albertensis, being present in the Late Maastrichtian unlike the latter species. In 2007, Mortimer (DML, 2007) noticed that Zapsalis matched ?Dromaeosaurus morphotype A, although which kind of dromaeosaurid it was was still a mystery. The question was solved by the description of a complete Saurornitholestes langstoni skull UALVP 55700 (Currie and Evans, 2020) which featured a tooth nearly identical to Zapsalis' holotype in the second premaxillary position. Its other premaxillary teeth are also ridged, showing Zapsalis consists of premaxillary teeth of Saurornitholestes or closely related taxa. Currie and Evans proposed Zapsalis' holotype differed from Saurornitholestes langstoni (RTMP 79.15.3 and UALVP 55700 at least) in lacking mesial serrations (seemingly not taphonomic) and being concave apicodistally, and thus "recommended that the two genera be kept separate." Given the large amount of dental variation within theropod species and correlation between their formations, this seems flimsy unless more Judith River teeth are discovered that share those characters with AMNH 3953. Indeed, Baszio figures two Dinosaur Park teeth with convex apicodistal margins and no mesial serrations (RTMP 87.36.383, RTMP coll.), blending the morphotypes. It would also lead to such a narrow concept of Saurornitholestes that even the Two Medicine material couldn't justifiably be referred. One motivation for keeping the genera separate is the obvious priority Zapsalis abradens has over Saurornitholestes langstoni, and unlike explanatus or laevifrons, it can be distinguished from all other named taxa (even Atrociraptor and Bambiraptor). Yet it does not seem distinguishable from 'Zapsalis' teeth from the later Scollard and Lance Formations (Baszio, 1997: pl. VI, fig. 87, 88), suggesting Zapsalis' holotype is indeterminate at the species level. While one could argue for a genus Zapsalis containing indeterminate Z. abradens, and diagnostic 'Z.langstoni' and 'Z. sullivani', a petition to the ICZN to suppress Zapsalis abradens would cause far less disruption.
Early finds- Colbert and Russell (1969) tentatively referred a fourth metatarsal (as CMN 12072) and a pedal ungual II (CMN 12240) to Dromaeosaurus, as did Ostrom (1969) for the latter element. Paul (1988) referred the ungual to Saurornitholestes instead, as he hypothesized Dromaeosaurus to have a reduced ungual as in Adasaurus. Both elements were referred to Saurornitholestes by Currie (1995), but as those bones remain undescribed for known Saurornitholestes specimens (and Dromaeosaurus may not be closer to Adasaurus), such referrals are only provisional. The CMN online catalog lists 12072 as a cast of the Dromaeosaurus holotype, but 12071 as a real Dromaeosaurus fossil, so 12072 may have been a typo.
Russell (1969) listed CMN 12343, 12354 (both found in 1968) and UA 5283 as frontals different from Dromaeosaurus, Troodon and a therizinosaur frontal (CMN 12355), but these were later made paratypes of Saurornitholestes when that genus was described.
Sahni (1972) identified AMNH 8518 as Troodon, but the large DSDI matches Saurornitholestes better. Howse and Milner (1993) identified NHMUK R4463 as a troodontid, but Makovicky (1995) recognized it as Saurornitholestes.
Sues (1977) described two dentaries (UA 12091 and 12339) discovered in 1969 and 1974 respectively. He referred these to Dromaeosaurus sp., but they were later realized to be Saurornitholestes (Paul, 1988).
The holotype (RTMP 74.10.5) was discovered in 1974, and described by Sues (1978) as a new genus of dromaeosaurid. Currie and Evans (2020) report "the quarry was reopened in 2002 and additional cranial material was recovered" including a lacrimal, jugal, postorbital and angular described in that paper.
Recent discoveries- Howse and Milner (1993) illustrated a sacrum (NHMUK R4463) and identified it as a troodontid, comparing it favorably to Ornithodesmus. Makovicky (1995) reidentified it as Saurornitholestes, which was first published publically in Norell and Makovicky (1997).
Makovicky (1995) describes the caudal vertebrae of RTMP 82.26.1 in depth. It is also mentioned by Currie (1995, 2005).
RTMP 88.121.39 is a specimen discovered in 1988 and first mentioned by Currie et al. (1990). The dorsal ribs were noted to be apneumatic by Britt (1993). Makovicky (1995) described the elongate distal caudal prezygapophyseal anatomy. Currie (1995) noted it has a T-shaped quadratojugal, fifteen dentary teeth, fused interdental plates, a posterior splenial notch, a collumnar articular process and elongated distal caudal prezygapophyses. Christiansen (1999) lists measurements for the femur and tibia. Xu et al. (1999) noted the glenoid faced laterally and the coracoid was similar to other dromaeosaurids. Jacobsen (2001) describes and illustrates the dentary, noting tooth marks on the bone (these were previously mentioned by Tanke and Currie, 2000). He stated the specimen is under study by Currie and Varricchio. Rauhut (2003) lists the scapular dimensions, and illustrates the splenial, scapulocoracoid and fibula. Codd (2004) notes that there are scars for uncinate processes on three preserved ribs. Jasinowski et al. (2006) illustrate and describe the scapulocoracoid and humerus. Brusatte et al. (2013) list various limb element lengths. Funston and Currie (2018) figure the tibiotarsus in different views. Currie and Evans (2020) describe and figure the cranial material.
MOR 660 is a partial skeleton discovered in 1990. Britt (1993) describes the pneumaticity of MOR 660 and notes it is being studied by Currie and Varricchio. Makovicky (1995) describes the presacral and sacral morphology in depth. Novas (1998) mentions the ulna has a feeble olecranon process and lacks the proximoanterior process for articulation with the radius. Currie and Dong (2001) note that the anterior dorsals have very large hyapophyses. Hutchinson (2001) illustrates the ilium, and notes it has a supratrochanteric process and a reduced but present cuppedicus fossa. Rauhut (2003) notes it has a concave anterior pubic peduncle edge, while he notes the posterior dorsal centra are subcircular, all dorsal centra are pleurocoelous and the presence of five sacral vertebrae in his theropod phylogeny study from that same year. He also illustrates manual ungual II. Claessens (2004) studied the medial gastralia of the specimen. Organ and Adams (2005) studied the histology of elongate prezygapophyses and chevrons from this specimen. Novas and Pol (2005) note the radius has a triangular proximal articular surface and the femur is robust. Martinez and Novas (2006) note the distal ulna has an anteroposteriorly expanded articular surface. Turner et al. (2007) lists the femoral length in their supplementary information table. Norell et al. (2006) note the posterior cervicals lack carotid processes, but have a raised lip instead. Csiki et al. (2010) list the hindlimb measurements. Brusatte et al. (2013) list various limb element lengths.
Norell and Makovicky (1997) note the last dorsal centrum and sacral centra of RTMP 67.20.36 are pleurocoelous.
Christiansen (1999) lists measurements for the tibia and fibula of MOR 666.
Xu et al. (1999) mention a sternal plate found in 1992 (RTMP 92.36.333) that was later described in detail by Godfrey and Currie (2004).
Hutchinson (2001) identified an ischium (RTMP 86.77.2) as Troodon, but it is more likely Saurornitholestes based on comparison to Bambiraptor and Troodon specimen MOR 553L.
Rauhut (2003) mentions there are five sacral vertebrae in RTMP 92.36.129.
Rauhut (2004) lists RTMP 94.12.844 (a maxilla), but it was not described and illustrated until Currie and Varricchio's (2004) paper. Powers et al. (2022) provide CT scans of the element.
Currie and Evans (2015) reported a new incomplete skeleton (UALVP 55700- McFeeters, pers. comm. 2015) found on June 20 2014. It preserves uncinate processes, furcula, sternum and sternal ribs, with a shorter and broader snout then Velociraptor and a larger pedal ungual II. The skull was described in depth by Currie and Evans (2020), who state the mandibular CT scans and postcrania "will be described in a separate paper."
UALVP 55425 is a fused surangular and articular found on May 25 2009 (Currie and Evans, 2020).
Hohman and Varricchio (2021) described juvenile maxilla MOR 9753 found in the 1990 from Bob's Vacation Site, which is smaller than Bambiraptor's holotype but shares characters with Saurornitholestes suggesting the former is not a juvenile of the latter.
References- Cope, 1876a. Descriptions of some vertebrate remains from the Fort Union Beds of Montana. Proceedings of the Academy of Natural Sciences of Philadelphia. 28, 248-261.
Cope, 1876b. On some extinct reptiles and Batrachia from the Judith River and Fox Hills Beds of Montana. Proceedings of the Academy of Natural Sciences of Philadelphia. 28, 340-359.
Hay, 1902. Bibliography and catalog of the Fossil Vertebrata of North America. Bulletin of the United States Geological Survey. 179, 1-868.
Lambe, 1902. New genera and species from the Belly River Series (mid-Cretaceous). Geological Survey of Canada Contributions to Canadian Palaeontology. 3(2), 25-81.
Osborn, 1902. On Vertebrata of the Mid-Cretaceous of the Northwest Territory. I: Distinctive characters of the Mid-Cretaceous fauna. Contrib. Canad. Pal. III. 1-21.
Hatcher, 1903. Osteology of Haplocanthosaurus, with description of a new species, and remarks on the probable habits of the Sauropoda and the age and origin of the Atlantosaurus Beds. Memoirs of the Carnegie Museum. 2(1), 1-72
Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History. 46(6), 367-385.
Kuhn, 1939. Fossillium Catalogus.
Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus. American Museum Novitates. 2380, 49 pp.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History. 30, 165 pp.
Russell, 1969. A new specimen of Stenonychosaurus from the Oldman Foramtion of Alberta. Canadian Journal of Earth Sciences. 6, 595-612.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin of the AMNH. 147.
Sues, 1977. Dentaries of small theropods from the Judith River Formation (Campanian) of Alberta, Canada. Canadian Journal of Earth Sciences. 14(4), 587-592.
Sues, 1978. A new small theropod dinosaur from the Judith River Formation (Campanian) of Alberta, Canada. Zoological Journal of the Linnean Society. 62, 381-400.
Currie, 1987. Theropods of the Judith River Formation. Occasional Paper of the Tyrrell Museum of Palaeontology. 3, 52-60.
Paul. 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus - and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River - Deinonychus, Velociraptor and Saurornitholestes. Hunteria. 2(4), 1-9.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press. 107-125.
Britt, 1993. Pneumatic postcranial bones in dinosaurs and other archosaurs. PhD Thesis, University of Calgary (Canada), Alberta.
Howse and Milner, 1993. Ornithodesmus - a maniraptoran theropod dinosaur from the Lower Cretaceous of the Isle of Wight, England. Palaeontology. 36, 425-437.
Currie, 1995. New information on the anatomy and relationships of Dromaeosaurus albertensis (Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 15(3), 576-591.
Currie and Jacobsen, 1995. An azhdarchid pterosaur eaten by a velociraptorine theropod. Canadian Journal of Earth Sciences. 32, 922-925.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Univ. Copenhagen, 311pp.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku Saizensen [Dino-Frontline]. 9, 92-119 (part 1); 10, 75-99 (part 2) [in Japanese].
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton: Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Novas, 1998. Megaraptor namunhuaiquii, gen. et sp. nov., a large-clawed, Late Cretaceous theropod from Patagonia. Journal of Vertebrate Paleontology. 18(1), 4-9.
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Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology. 76(4), 751-763.
Rauhut, 2003. A tyrannosauroid dinosaur from the Upper Jurassic of Portugal. Palaeontology 46, 903-910.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Palaeontology. 69, 1-213.
Claessens, 2004. Dinosaur gastralia; Origin, morphology and function. Journal of Vertebrate Paleontology. 24(1), 89-106.
Codd, 2004. The uncinate processes in birds and their implications for the breathing mechanics of maniraptoran dinosaurs. Unpublished dissertation. Rheinischen Friedrich-Wilhelms-Universität Bonn. 108 pp.
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Currie, 2005. Theropods, including birds. in Currie and Koppelhus (eds). Dinosaur Provincial Park, a spectacular ecosystem revealed, Part Two, Flora and Fauna from the park. Indiana University Press. 367-397.
Novas and Pol, 2005. New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia. Nature 433: 858 - 861.
Organ and Adams, 2005. The histology of ossified tendon in dinosaur. Journal of Vertebrate Paleontology. 25(3), 602-613.
Jasinowski, Russell and Currie, 2006. An integrative phylogenetic and extrapolatory approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder musculature. Zoological Journal of the Linnean Society. 146, 301-344.
Martínez and Novas, 2006. Aniksosaurus darwini gen. et sp. nov., a new coelurosaurian theropod from the early Late Cretaceous of central Patagonia, Argentina. Revista del Museo Argentino de Ciencias Naturales. 8(2), 243-259.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
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Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Csiki, Vremir, Brusatte and Norell, 2010. An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania. Proceedings of the National Academy of Sciences. Early Edition, 5 pp.
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Redman, Moore and Varricchio, 2015. A new vertebrate microfossil locality in the Upper Two Medicine Formation in the vicinity of Egg Mountain. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 201-202.
Coy and Currie, 2016. Discovery and significance of the University of Alberta’s Saurornitholestes (UALVP 55700). Exhibition catalog, The Dinosaur Expo 2016. Asahi Shimbun. 114-117.
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S? sullivani Jasinski, 2015
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
Holotype- (SMP VP-1270) frontal (42.8 mm) (Sullivan and Lucas, 2000)
Referred- ?(NMMNH P-22578) tooth (8.4x4.4x1.9 mm) (Williamson and Brusatte, 2014)
?(NMMNH P-26201) tooth (Williamson and Brusatte, 2014)
?(NMMNH P-26205) tooth (8.7x4.9x2.2 mm) (Williamson and Brusatte, 2014)
?(NMMNH P-32752) tooth (8.9x4.4x2.4 mm) (Williamson and Brusatte, 2014)
?(NMMNH P-68395) tooth (4.2x3.7x2.1 mm) (Williamson and Brusatte, 2014)
?(SMP VP-1741) partial pedal ungual II (Sullivan, 2006)
?(SMP VP-1901) tooth (15 mm) (Sullivan, 2006)
Diagnosis- (after Jasinski, 2015) differs from S. langstoni in- more constricted anteriorly; less prominent nasal facets; less prominent anterior projection between the nasal and lacrimal facet regions; deeper and less strongly demarcated orbital rim; deeper and more prominent olfactory bulb surface; more pronounced and longer ventrally-directed ridge between the olfactory bulb surface and the cerebral hemisphere surface; more robust frontal median sutural surface; while slightly smaller, still being more robust.
Comments- Sullivan and Lucas (2000) described SMP VP-1270 as an individual of Saurornitholestes langstoni. Later, Sullivan (2006) described another frontal as the new species S. robustus, referring the original frontal and two more elements to this taxon. Evans et al. (2014) later reidentified S. robustus as a troodontid, though the other remains are still dromaeosaurid. Williamson and Brusatte (2014) described more teeth as Dromaeosauridae morphotype A, which they found fall within the range of variation of Dinosaur Park Saurornitholestes. Jasinski (2015) used SMP VP-1270 as the holotype of his new species Saurornitholestes sullivani, and kept other De-na-zin dromaeosaurid remains as indeterminate. However, only the S. langstoni holotype was compared, not other specimens of that species to check for variation which might explain sullivani's differences. Furthermore, most other North American dromaeosaurids from the Late Cretaceous (Acheroraptor, Atrociraptor) don't preserve frontals, making proposed Saurornitholestes frontal synapomorphies difficult to test.
References- Sullivan and Lucas, 2000. First occurrence of Saurornitholestes (Theropoda: Dromaeosauridae) from the Upper Cretaceous of New Mexico. in Lucas and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History and Science, Bulletin 17. pp. 105-108.
Sullivan, 2006. Saurornitholestes robustus, n. sp. (Theropoda: Dromaeosauridae) from the Upper Cretaceous Kirtland Formation (De-na-zin Member), San Juan Basin, New Mexico. In Lucas and Sullivan (eds.). Late Cretaceous vertebrates from the Western Interior. New Mexico Museum of Natural History and Science Bulletin. 35, 253-256.
Evans, Larson, Cullen and Sullivan, 2014. 'Saurornitholestes' robustus is a troodontid (Dinosauria: Theropoda). Canadian Journal of Earth Sciences. 51(7), 730-734.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Jasinski, 2015.
S? sp. (Gangloff, 1998)
Campanian, Late Cretaceous
Prince Creek Formation, Alaska, US
(AK249-V-034) tooth (Fiorillo and Currie, 2000)
(AK249-V-035) tooth (Fiorillo and Currie, 2000)
vertebra (Gangloff, 1998)
References- Gangloff, 1998. Arctic Dinosaurs with Emphasis on the Cretaceous Record of Alaska and the Eurasian-North American Connection. Kirkland, Lucas and Estep (eds). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science, Bulletin. 14, 211-220.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation (Cretaceous) of Northern Alaska, with speculations on Arctic dinosaur paleoecology. Journal of Vertebrate Paleontology. 20(4), 675-682.
S? sp. nov. (Baszio, 1997)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(CMN coll.) teeth (Russell, 1935)
(UA MR-4:1-3, 21-32) fifteen teeth (Baszio, 1997)
Comments- These teeth are highly variable, but have smaller serrations than S. langstoni. Mesial serrations are usually absent, and the teeth are only distinguishable from Richardoestesia based on serration count.
References- Russell, 1935. Fauna of the Upper Milk River beds, Southern Alberta. Transactions of the Royal Society of Canada, series 3, section 4. 29, 115-127.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
S? sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Material- (RTMP 86.55.7) tooth (Bell and Currie, 2015)
(RTMP 86.55.9) tooth (Bell and Currie, 2015)
(RTMP 86.55.27) tooth (Bell and Currie, 2015)
(RTMP 86.55.38) tooth (Bell and Currie, 2015)
(RTMP 88.55.129; paratype of Boreonykus certekorum) incomplete distal caudal vertebra (Bell and Currie, 2015)
(RTMP 87.55.148) tooth (Bell and Currie, 2015)
(RTMP 87.55.149) tooth (Bell and Currie, 2015)
(RTMP 86.55.184; paratype of Boreonykus certekorum) pedal ungual II (Bell and Currie, 2015)
(RTMP 87.55.329) tooth (Bell and Currie, 2015)
(RTMP 87.55.333) tooth (Bell and Currie, 2015)
(RTMP 87.55.1523) tooth (Ryan and Russell, 2001)
(RTMP 89.55.707) tooth (Bell and Currie, 2015)
(RTMP 89.55.1005) tooth (Bell and Currie, 2015)
(RTMP 89.55.1141) tooth (Bell and Currie, 2015)
(RTMP 89.55.1162) tooth (Bell and Currie, 2015)
(RTMP 2004.23.4) tooth (Fanti and Miyashita, 2009)
(UALVP 53597; paratype of Boreonykus certekorum) incomplete manual ungual II (Bell and Currie, 2015)
two teeth (Fanti and Miyashita, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Wapiti Formation, British Columbia, Canada
Material- (PRPRC coll.) tooth (McCrea and Buckley, 2004)
Comments- Ryan and Russell (2001) listed the frontal RTMP 89.55.47 as "Saurornitholestes undescribed n. sp. (Currie pers. comm.)". Bell and Currie (2015) described this and other material (distal caudal RTMP 88.55.129, manual ungual UALVP 53597, and pedal ungual II RTMP 86.55.184) from the Pipestone Creek Pachyrhinosaurus lakustai bonebed as Boreonykus certekorum, noting that preserved elements are not duplicated and are properly proportioned to belong to one individual. The fourteen referred teeth are shed so were admitted to at least belong to another individual. While the authors assigned Boreonykus to Velociraptorinae, Cau (online, 2015) noted the frontal differs from dromaeosaurids and that its association with the postcrania cannot be substantiated. Boreonykus is here placed as ?Paraves incertae sedis, though it may be Paronychodon.
McCrea and Buckley (2004) first reported a theropod tooth excavated with a hadrosaur from British Columbia in 2002-2003. Reid (2016) notes McCrea assigned it to Saurornitholestes online, and he refers it to cf. Boreonykus certekorum based on the formation and lack of distal inclination.
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press. 279-297.
McCrea and Buckley, 2004. Excavating British Columbia's first dinosaurs and other palaeontological projects in the Tumbler Ridge area. Eighth Alberta Palaeontological Society Symposium, Abstracts. 24-33.
Buckley, McCrea and Currie, 2005. Theropod teeth from the Upper Cretaceous Kaskapau (Middle Turonian) and the Wapiti (Upper Campanian - Lower Maastrichtian) formations of north-eastern British Columbia, Canada. Journal of Vertebrate Paleontology. 25(3), 40A-41A.
Fanti and Miyashita, 2009. A high latitude vertebrate fossil assemblage from the Late Cretaceous of west-central Alberta, Canada: Evidence for dinosaur nesting and vertebrate latitudinal gradient. Palaeogeography, Palaeoclimatology, Palaeoecology. 275, 37-53.
Bell and 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. e1034359. DOI: 10.1080/02724634.2015.1034359.
Cau, online 2015. http://theropoda.blogspot.com/2015/12/boreonykus-e-un-dromaeosauridae.html
Reid, 2016. A review of dinosaurian body fossils from British Columbia, Canada. PeerJ Preprints. 4:e1369v3.
S. sp. nov? (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
Material- (RTMP 87.16.18) tooth (Ryan and Russell, 2001)
(UA 103) premaxillary tooth (Baszio, 1997)
(UA 121) premaxillary tooth (Baszio, 1997)
(UA JLE 64:109) lateral tooth (Baszio, 1997)
several premaxillary teeth (Baszio, 1997)
teeth (Ryan and Russell, 2001)
Comments- Baszio (1997) reported UA JLE 64:109 was similar to teeth from the Lance Formation, perhaps indicating they are the same species. He referred it to Saurornitholestes, which may have merit as he also describes several Zapsalis-style premaxillary teeth that are characteristic of that genus. Those teeth have numerous, strong ridges unlike Bambiraptor or Atrociraptor, and may have more ridges than S. langstoni (6-8).
References- Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp. 279-297.
S. sp. (Carpenter, 1982)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (UCMP 124983) tooth (2.1 mm) (Carpenter, 1982)
(UCMP 124984) tooth (3 mm) (Carpenter, 1982)
(UCMP 124985) tooth (2.7 mm) (Carpenter, 1982)
(UCMP 125238) dentary fragment (Carpenter, 1982)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
(FMNH PR2893) tooth (4.4x3.1x1.2 mm) (Gates, Zanno and Makovicky, 2015)
(FMNH PR2895) tooth (4.5x3x1.3 mm) (Gates, Zanno and Makovicky, 2015)
(FMNH PR2897) premaxillary tooth (Gates, Zanno and Makovicky, 2015)
(FMNH PR2898) tooth (5.4x3.7x1.3 mm) (Gates, Zanno and Makovicky, 2015)
teeth and elements (DePalma, 2010)
Comments- Carpenter (1982) considered UCMP 125238 to be probably conspecific with Hell Creek specimen UW 13684. Both have fused interdental plates, subrectangular alveoli and elongate external foramina, which Carpenter compares to Velociraptor (in which he includes Saurornitholestes). These characters are also present in more basal dromaeosaurids such as Sinornithosaurus and Bambiraptor, though the elongate foramina does distinguish the dentary fragments from Dromaeosaurus. The former preserves a short, curved tooth with poorly developed mesial serrations and 5.5 distal serrations per mm. Referred teeth are short, strongly curved and highly compressed, with well developed distal serrations and variably developed mesial ones. Carpenter states they resemble Velociraptor, which may indicate they belong to Saurornitholestes.
FMNH PR2897 is a Zapsalis-style premaxillary tooth, more similar to the Zapsalis holotype than some Saurornitholestes langstoni in lacking mesial serrations and having a concave apicodistal edge. The numerous, strong ridges are unlike Bambiraptor or Atrociraptor.
References- Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek formations and a description of a new species of theropod. Contributions to Geology, University of Wyoming. 20(2), 123-134.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis, University of Kansas. 227 pp.
Gates, Zanno and Makovicky, 2015. Theropod teeth from the upper Maastrichtian Hell Creek Formation "Sue" Quarry: New morphotypes and faunal comparisons. Acta Palaeontologica Polonica. 60(1), 131-139.
S. sp. nov. (Estes, 1964)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- (AMNH 22670) fourteen teeth (AMNH online)
(AMNH 27122; in part) premaxillary tooth (AMNH online)
(AMNH coll.) premaxillary teeth (Estes, 1964)
(UA BTB 132) premaxillary tooth (Baszio, 1997)
(UA BTB 123, 129-131, 161-164) seven lateral teeth (Baszio, 1997)
(UCM 39502) tooth (1.7 mm) (Carpenter, 1982)
(UCM 45055) tooth (2.6 mm) (Carpenter, 1982)
(UW 13684) dentary fragment (Carpenter, 1982)
(YPM PU 55007) (YPM online)
(YPM PU 55502) (YPM online)
(YPM PU 55522) (YPM online)
(YPM PU 55535) (YPM online)
(YPM PU 55560) (YPM online)
(YPM PU 55561) (YPM online)
Comments- Carpenter (1982) considered UW 13684 to be probably conspecific with Hell Creek specimen UCMP 125238. Both have fused interdental plates, subrectangular alveoli and elongate external foramina, which Carpenter compares to Velociraptor (in which he includes Saurornitholestes). These characters are also present in more basal dromaeosaurids such as Sinornithosaurus and Bambiraptor, though the elongate foramina does distinguish the dentary fragments from Dromaeosaurus. Referred teeth are short, strongly curved and highly compressed, with well developed distal serrations and variably developed mesial ones. Carpenter states they resemble Velociraptor, which may indicate they belong to basal dromaeosaurids.
AMNH 22670 is listed as Deinonychus sp. on the AMNH website, but this is unlikely due to the age difference from the Cloverly Formation.
Baszio (1997) refers Lance Formation teeth (UA BTB coll.) to Saurornitholestes sp., but notes they differ from S. langstoni in being smaller, with rounded basal serrations and mesial serrations which are developed as a slight serrated structure if present. The premaxillary tooth UA BTB 132 has numerous, strong ridges unlike Bambiraptor or Atrociraptor.
The YPM specimens are listed as Dromaeosauridae, but photos indicate they are 'velociraptorine'-type teeth with high DSDIs.
References- Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek formations and a description of a new species of theropod. Contributions to Geology, University of Wyoming. 20(2), 123-134.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
S? sp. (Williamson, 1997)
Late Santonian-Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
Material- (NMMNH P-25054) tooth (?x3x1.4 mm)
Comments- Williamson (1997) referred a tooth to cf. Saurornitholestes sp. and Williamson and Brusatte (2014) found it fell within the range of variation of Dinosaur Park Saurornitholestes teeth.
References- Williamson, 1997. A new Late Cretaceous (Early Campanian) vertebrate fauna from the Allison Member, Menefee Formation, San Juan Basin, New Mexico. New Mexico Museum of Natural Hisory and Science Bulletin. 11, 51-59.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
S? sp. (Armstrong-Ziegler, 1980)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-26240) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27486) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27487) tooth (?x2.7x1.3 mm) (Williamson and Brusatte, 2014)
(NMMNH P-27559) tooth (8.6x5.5x2.5 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30001) tooth (7.1x4x2.1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30002) tooth (?x3.4x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30003) tooth (3.8x2.9x1.4 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30330) tooth (3.6x3x1.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38027) tooth (?x4.5x2.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38425) tooth (?x2.5x1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38431) tooth (Williamson and Brusatte, 2014)
(NMMNH P-38432) tooth (Williamson and Brusatte, 2014)
(NMMNH P-49808) tooth (?x5.8x2.5 mm) (Williamson and Brusatte, 2014)
(NMMNH P-66896) tooth (9.1x4.3x2.3 mm) (Williamson and Brusatte, 2014)
teeth (Armstrong-Ziegler, 1980)
three teeth (Hall, 1991)
Comments- Armstrong-Ziegler (1978) reported dromaeosaurid teeth, while three others were referred to Saurornitholestes langstoni by Hall (1991). Sullivan and Lucas (2006) could not confirm the latter identification. Williamson and Brusatte (2014) describe several teeth as falling within the range of variation of Dinosaur Park Saurornitholestes teeth, including some identified by Hall as troodontids.
References- Armstrong-Ziegler, 1980. Amphibia and Reptilia from the Campanian of New Mexico. Fieldiana Geology (new series). 4, 39 pp.
Hall, 1991. Lower vertebrate paleontology of the upper Fruitland Formation, Fossil Forest area, New Mexico, and implications for Late Cretaceous terrestrial biostratigraphy. Masters Thesis. University of Kansas. 126 pp.
Sullivan and Lucas, 2006. The Kirtlandian land-vertebrate "age" - faunal composition, temporal position and biostratigraphic correlation in the nonmarine Upper Cretaceous of western North America. New Mexico Museum of Natural History and Science Bulletin. 35, 7-29.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
S? sp. (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- ?(NMMNH P-07189) tooth (8.3x?x3.1 mm)
(NMMNH P-30220) tooth (?x?x.5 mm)
(NMMNH P-30222) tooth
(NMMNH P-30223) tooth (?x4.1x2.2 mm)
(NMMNH P-30225) tooth (?x4x2 mm)
(NMMNH P-30226) tooth
(NMMNH P-30227) tooth (?x2.6x1.2 mm)
(NMMNH P-30228) tooth (?x3.7x1.7 mm)
(NMMNH P-30229) tooth (?x3.3x1.5 mm)
(NMMNH P-30230) tooth
(NMMNH P-30231) tooth
(NMMNH P-30232) tooth (?x4.7x2.3 mm)
(NMMNH P-32831) tooth (?x4.3x? mm)
(NMMNH P-33144) tooth (?x~4.3x2 mm)
(NMMNH P-33145) tooth (?x3.1x1.5 mm)
(NMMNH P-33146) tooth (?x4.1x~1.7 mm)
(NMMNH P-33149) tooth
(NMMNH P-33473) tooth (6.5x3.2x1.7 mm)
(NMMNH P-33474) tooth
(NMMNH P-33475) tooth (?x4.6x2.1 mm)
(NMMNH P-33476) tooth (3.2x2.5x1.1 mm)
(NMMNH P-33477) tooth (?x?x1.4 mm)
(NMMNH P-33484) tooth (?x3.8x1.8 mm)
(NMMNH P-33485) tooth
(NMMNH P-33486) tooth (6.7x5.1x2.6 mm)
(NMMNH P-33487) tooth
Comments- Williamson and Brusatte (2014) described these as Dromaeosauridae morphotype A, which they found fall within the range of variation of Dinosaur Park Saurornitholestes.
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
S? sp. (Sankey et al., 2005)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Material- (LSUMG 5158) tooth (Sankey, 2001)
(LSUMG 5659) tooth (2.8 mm) (Sankey, 2001)
(LSUMG 5923) tooth (5.3 mm) (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 5924) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 5928) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 5950) tooth (~2.2 mm) (Sankey, 2001)
(LSUMG 5980) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6132) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6139) tooth (~5.2 mm) (Sankey, 2001)
(LSUMG 6183) tooth (Sankey, 2001)
(LSUMG 6184) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6185) tooth (Sankey, 2001)
(LSUMG 6204) tooth (6 mm) (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6229) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6234) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6270) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6280) tooth (~7.5 mm) (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6281) tooth (7.2 mm) (Sankey, Standhardt and Shiebout, 2005)
? metacarpal, phalanx (Jasinski, Sullivan and Dodson, 2015)
Late Campanian, Late Cretaceous
Aguja Formation, Mexico
teeth (Montellano, Monroy, Hernandez-Rivera and Torres, 2009)
Comments- Montellano et al. (2009) reported teeth from S. langstoni, plus two unnamed taxa they called Saurornitholestes n. sp. A? and S. n. sp. C.. Evaluation of these awaits their description.
References- Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend, Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous (Campanian-Maastrichtian), Big Bend National Park, Texas. in Carpenter (ed). The Carnivorous Dinosaurs. 127-152.
Montellano, Monroy, Hernandez-Rivera and Torres, 2009. Late Cretaceous microvertebrate fauna from the Northern state of Coahuila, Mexico. Journal of Vertebrate Paleontology. 29(3), 151A.
Jasinski, Sullivan and Dodson, 2015. Late Cretaceous dromaeosaurid theropod dinosaurs (Dinosauria: Dromaeosauridae) from southern Laramidia and implications for dinosaur faunal provinciality in North America. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 150.
S? sp. (Standhardt, 1986)
Early Maastrichtian, Late Cretaceous
Upper Aguja Formation, Texas, US
Material- (LSUMG 113:coll.) several teeth
Reference- Standhardt, 1986. Vertebrate paleontology of the Cretaceous/Tertiary transition of Big Bend National Park, Texas. Unpublished Ph.D. dissertation, Louisiana State University, Baton Rouge. 298 pp.
S? sp. (Schwimmer, Sanders, Erickson and Weems, 2015)
Middle Campanian, Late Cretaceous
Coachman Formation, South Carolina, US
Material- (ChM PV8674) tooth
(ChM PV8675) tooth
(ChM PV8679) incomplete pedal ungual II
(SCSM 2005.11.1) tooth
Comments- Schwimmer et al. (2015) referred these to Saurornitholestes langstoni.
Reference- Schwimmer, Sanders, Erickson and Weems, 2015. A Late Cretaceous dinosaur and reptile assemblage from South Carolina, USA. Transactions of the American Philosophical Society. 105(2), 157 pp.
S? sp. (Schwimmer, Sanders, Erickson and Weems, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Donaho Creek or Peedee Formation, South Carolina, US
Material- (SCSM 99.55.1) partial pedal ungual II
Comments- Schwimmer et al. (2015) referred this to Saurornitholestes langstoni.
Reference- Schwimmer, Sanders, Erickson and Weems, 2015. A Late Cretaceous dinosaur and reptile assemblage from South Carolina, USA. Transactions of the American Philosophical Society. 105(2), 157 pp.
S? sp. (Alifanov and Bolotsky, 2002)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- (1/1081) tooth (15x10x4 mm) (Bolotsky, 2011)
(1/1082) tooth (15x7x2 mm) (Bolotsky, 2011)
(1/1083) tooth (13x8x1.8 mm) (Bolotsky, 2011)
Reference- Alifanov and Bolotsky, 2002. New data about the assemblages of the Upper Cretaceous carnivorous dinosaurs (Theropoda) from the Amur region. In Kirillova (ed.). Fourth International Symposium of IGCP 434. Cretaceous continental margin of East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
Bolotsky, 2011. On paleoecology of carnivorous dinosaurs (Tyrannosauridae, Dromaeosauridae) from Late Cretaceous fossil deposits of Amur region, Russian far East. Global Geology. 14(1), 1-6.

Tsaagan Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006
= Linheraptor Xu, Choiniere, Pittman, Tan, Xiao, Li, Tan, Clark, Norell, Hone and Sullivan, 2010
T. mangas Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006
= Velociraptor mangas (Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006) Paul, 2010
= Linheraptor exquisitus Xu, Choiniere, Pittman, Tan, Xiao, Li, Tan, Clark, Norell, Hone and Sullivan, 2010
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadokhta Formation, Mongolia
Holotype- (IGM 100/1015) (adult) skull (201 mm), sclerotic rings, mandibles, ten cervical vertebrae, cervical ribs, proximal scapula, partial coracoid
Late Campanian, Late Cretaceous
Zos Wash, Djadokhta Formation, Mongolia
Referred- (IGM 100/3503; "IGM 100/981" of Turner et al., 2007) (~1.5 m, ~15 kg) frontal, jugal fragment, ectopterygoids, pterygoid, incomplete dentary, dorsal vertebra, few proximal dorsal ribs, gastralia, seven distal caudal vertebrae, several chevrons, scapulocoracoid, humeral fragment, radius, ulna (~113 mm), scapholunare, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, distal pubis, ischium, proximal fibula, phalanges I-1, pedal unguals I, propximal metatarsal II, proximal metatarsal III, phalanges III-3, pedal unguals III, metatarsals IV (one proximal), phalanx IV-4, pedal ungual IV, metatarsals V (Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006)
Late Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Referred- (IVPP V16923; holotype of Linheraptor exquisitus) (~1.8 m; adult) skull (225 mm), sclerotic rings, incomplete mandibles, hyoids (one partial), (cervical series 320 mm) atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth cervical vertebra, cervical ribs 2-10, first dorsal vertebra, dorsal ribs 1-8, tenth dorsal rib, uncinate processes, gastralia, seven proximal caudal vertebrae (first caudal 22.2 mm), eleven distal caudal vertebrae (longest 30 mm), seven proximal chevrons, distal chevrons, scapula (~160 mm), sternum (80 mm), humerus (~155 mm), radius (110 mm), metacarpal I (25.3 mm), phalanx I-1 (47.6 mm), metacarpal II (64.4 mm), phalanx II-1 (32.3 mm), phalanx II-2 (53.5 mm), manual ungual II (41 mm), metacarpal III (53.1 mm), phalanx III-1 (15.5 mm), phalanx III-2 (12 mm), phalanx III-3 (33 mm), pubis (240 mm), femur (220 mm), tibiotarsi (252 mm), fibula, distal tarsal IV, phalanx I-1 (15.7 mm), pedal ungual ! (17.5 mm straight), metatarsal II (103.2 mm), phalanx II-2 (39.9 mm), pedal unguals II (55.6 mm), metatarsal III (127.9 mm), phalanx III-1 (54.2 mm), phalanx III-2 (32.9 mm), phalanx III-3 (19.3 mm), pedal ungual III (48.5 mm), metatarsal IV (112.8 mm), phalanx IV-1 (40 mm), phalanx IV-2 (28 mm), phalanx IV-3 (26.1 mm), phalanx IV-4 (26.4 mm), pedal ungual IV (18 mm) (Xu et al., 2010)
Diagnosis- (after Norell et al., 2006) paroccipital process pendulous; maxillary fenestra large; maxillary fenestra located at anterior edge of antorbital fossa (also in Atrociraptor); jugal meets the squamosal to exclude the postorbital from the margin of the infratemporal fenestra.
(after Xu et al., 2010) Xu et al. listed two characters supposedly shared between Linheraptor and Tsaagan, which would be diagnostic of the latter if they are synonymous- lacrimal with relatively broad medial lamina; sharp angle between anterior and dorsal processes of quadratojugal.
(after Turner et al., 2012) oval-shaped foramen magnum; low coracoid tuber; weak subglenoid shelf; dorsoventrally oriented path of supracoracoid nerve through coracoid.
(after Xu et al., 2015) Note Xu et al. viewed Tsaagan and Linheraptor as taxonomically distinct, but listed the following characters as shared between them. Promaxillary fenestra not visible in lateral view; maxilla without ridge dorsal to supralabial foramina (also probably in Achillobator); longitudinal fossa along midline of nasals (also in Velociraptor); jugal without ridge parallel to orbital rim; frontal supraorbital rim rugose (also in Dromaeosaurus); parietal with prominent nuchal crest (also in Adasaurus); parietal forms entire nuchal crest (also in Sinornithosaurus); pterygoid flange small (also in Dromaeosaurus); all teeth without mesial serrations (also in some microraptorians).
Other diagnoses- Norell et al. (2006) originally listed paroccipital process not twisted distally in their diagnosis, but this is absent in the Linheraptor holotype (Xu et al., 2015). Turner et al. (2012) suggested it was taphonomic in the Tsaagan holotype, though Xu et al. view it as a character supporting taxonomic separation of the specimens. Similarly, Norell et al. listed 'basipterygoid process elongate and anteroventrally directed' as being diagnostic, but the Linheraptor holotype has shorter processes which are only ventrally directed. Again, Xu et al. view it as having taxonomic implications, thouigh it could also be individual variation.
Xu et al. (2010) listed two supposed autapomorphies of Linheraptor- greatly enlarged maxillary fenestra subequal in size to external naris; several large foramina on lateral surface of jugal. The former would be individual variation if the genus is a synonym of Tsaagan, while Xu et al. (2015) later noted the jugal foramina are also present in Velociraptor and "might be present in T. mangas but are simply obscured by specimen damage." They also listed 'lacrimal lacking lateral flange over descending process' as shared between Linheraptor and Tsaagan, but this is actually untrue for former's holotype (Xu et al., 2015).
Xu et al. (2015) describe 61 differences between Linheraptor and Tsaagan, interpreting the following as autapomorphies of the former- nasal process of premaxilla transversely compressed along entire length; premaxillary process of nasal terminates at anterior border of external naris; lacrimal with several anteriorly opening foramina on dorsal surface (possibly obscured by poor preservation in Tsaagan); pyramidal projection on anterior border of supratemporal fossa; supraoccipital crest short and sharp; quadratojugal dorsal process distally expanded; basipterygoid process is not inclined anteriorly. If the taxa are synonymous, these are merely examples of individual variation, as seen in other theropods (e.g. Allosaurus, Tyrannosaurus, Microraptor, Archaeopteryx).
Comments- The holotype was discovered in 1993 and originally identified as Velociraptor (e.g. Webster, 1996) before being identified as a new genus (Turner et al., 2006) then described by Norell et al. (2006). Tsaagan (under its specimen number) was used as an OTU in the Theropod Working Group analyses since 2001 however. Powers et al. (2022) provide CT scan data on the maxilla. Linheraptor's holotype was discovered in 2008 and described by Xu et al. (2010) as a new taxon of dromaeosaurid. Both Senter (2011) and Turner et al. (2012) synonymized Linheraptor with Tsaagan, though only the latter justified it. This is provisionally accepted here despite a recent paper by Xu et al. (2015) defending Linheraptor's validity based on 61 differences from Tsaagan. While these differences are real, Xu et al. do not establish they are outside the range of individual variation expected for a dromaeosaurid species. To the contrary, they dismiss the idea of utilizing individual variation present in Velociraptor mongoliensis until authors "determine how observed morphological variations relate to inter- or possibly intraspecific factors" ... "by including all Velociraptor specimens in a specimen-level phylogenetic analysis, by using morphometric methods to quantify the variation present and by deepening our understanding of the biological significance of the variations observed." Yet while ideal, such studies have not been completed for any Mesozoic theropod. Thus insisting "noticeable variations between L. exquisitus and T. mangas are grounds for taxonomic separation" until those studies are done on Velociraptor would logically result in the numerous differences present between almost all described individuals of e.g. Allosaurus and Tyrannosaurus being viewed as taxonomically significant until the studies are done on them as well. Such a conclusion is highly unrealistic, and given the large amount of individual variation present in all theropod species including V. mongoliensis, Tsaagan and Linheraptor are accepted as synonymous here until either further specimens show their differences covary or phylogenetic analyses find them to not be sister taxa even when their shared characters are included.
Tsaagan has been recovered as a velociraptorine (Senter et al., 2012; Brusatte et al., 2014) and a saurornitholestiine (Longrich and Currie, 2009).
IGM 100/3503- Discovered in 1998, Norell et al. (2006) mention "A second, as yet undescribed species of dromaeosaurid, which differs from Tsaagan mangas in frontal morphology [that] is known from the nearby locality of Zos Wash." Turner et al. (2007) describe an ulna as IGM 100/981, referring it to Velociraptor mongoliensis based on sharing several unlisted characteristics. This specimen was said to be from "the Gilvent Wash locality near Ukhaa Tolgod", but IGM 100/981 is a specimen from Khulsan later described as Kuru. Napoli et al. (2021) clarified the matter, stating "Turner et al. (2007a) erroneously identified the Zos Wash specimen as IGM 100/981. The correct specimen number for the Zos Wash specimen is IGM 100/3503." Ironically Ruebenstahl et al.'s (2021) abstract also switches 100/981 and 100/3503, but they do say the Zos Wash specimen "appears referrable to Velociraptor, although it differs from other Velociraptor mongoliensis specimens in some respects such as the absence of a dorsal ectopterygoid recess." Napoli et al. state "This individual is currently referred to Velociraptor mongoliensis and is under renewed study." Based on data presented in Ruebenstahl et al.'s SVP presentation and preliminary inclusion in the Hartman et al. maniraptoromorph analysis, I believe it is closest to Tsaagan and falls within the range of variation of the latter and Linheraptor.
References- Webster, 1996. Dinosaurs of the Gobi: Unearthing a fossil trove. National Geographic. 190(1), 70-89.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
Turner, Pol, Norell and Hwang, 2006. Resolving dromaeosaurid phylogeny: New information and additions to the tree. Journal of Vertebrate Paleontology. 26(3), 133A.
Turner, Makovicky and Norell, 2007. Feather quill knobs in the dinosaur Velociraptor. Science. 317, 1721.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Xu, Choiniere, Pittman, Tan, Xiao, Li, Tan, Clark, Norell, Hone and Sullivan, 2010. A new dromaeosaurid (Dinosauria: Theropoda) from the Upper Cretaceous Wulansuhai Formation of Inner Mongolia, China. Zootaxa. 2403, 1-9.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological continuity within Dinosauria. Journal of Evolutionary Biology. 24, 2197-2216.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2012 online. Tsaagan mangas, Digital Morphology. http://digimorph.org/specimens/Tsaagan_mangas/
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Brusatte, Vremir, Csiki-Sava, Turner, Watanabe, Erickson and Norell, 2013. The osteology of Balaur bondoc, an island-dwelling dromaeosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Romania. Bulletin of the American Museum of Natural History. 374, 1-100.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Xu, Pittman, Sullivan, Choiniere, Tan, Clark, Norell and Wang, 2015. The taxonomic status of the Late Cretaceous dromaeosaurid Linheraptor exquisitus and its implications for dromaeosaurid systematics. Vertebrata PalAsiatica. 53(1), 29-62.
Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021. A new dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central Mongolia. American Museum Novitates. 3982, 47 pp.
Ruebenstahl, Napoli, Bhullar, Turner and Norell, 2021. Two new eudromaeosaurs from Khulsan (central Mongolia) reveal modern-like faunal predatory structure amoung non-avian dinosaurs. The Society of Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual Meeting. 222-223.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as 2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters. Journal of Vertebrate Paleontology. 41(5), 2010087.

undescribed dromaeosaurine (Powers, Sullivan and Currie, 2020)
Late Campanian, Late Cretaceous
Djadokhta Formation, Mongolia
Material- (private coll.; cast UALVP 49389) skull (~193 mm), sclerotic plates, mandibles, pes
Comments- Powers et al. (2020) included in their analysis "a cast of a complete skull belonging to an undescribed eudromaeosaurian", which "is of a specimen in private collections and has no available locality data. The likeness of this specimen is akin to that of an Asian dromaeosaurid. and has been frequently labeled as Velociraptor, however, its morphology is quite different from other specimens referred to this genus." They report that "In the cluster analysis UALVP 49389 was found to be most like [Tsaagan and Linheraptor] and often plotted close to them but toward the extreme of variation along each PC." The maxilla is illustrated in their figure 1M and differs from most other dromaeosaurids in lacking an externally obvious promaxillary fenestra and having a slender maxillary fenestra along the ventral edge of the ascending process, and is similar to Tsaagan and Velociraptor in having an elongated anterior ramus. Powers' (2020) thesis this paper is based on listed the specimen as Velociraptor sp. and noted it consists of "Cast of articulated skull and foot", while the UA online catalog lists it being from the Djadokhta Formation of Mongolia. Powers et al.'s figure allows one to identify the cast as one labeled Velociraptor mongoliensis in the WitmerLab Collections (Witmer, online 2012) and ultimately sold by Gaston Design Inc. (2022 online; but available since at least 2004- https://web.archive.org/web/20040609002120/http://gastondesign.com/dinoskulls.htm), although no pes is associated with either cast.
References- Witmer, online 2012. https://people.ohio.edu/witmerl/collections/Theropods/Velociraptor_gaston.htm
Powers, 2020. The evolution of snout shape in eudromaeosaurians and its ecological significance. Masters thesis, University of Alberta. 437 pp.
Powers, Sullivan and Currie, 2020. Re-examining ratio based premaxillary and maxillary characters in Eudromaeosauria (Dinosauria : Theropoda): Divergent trends in snout morphology between Asian and North American taxa. Palaeogeography, Palaeoclimatology, Palaeoecology. 547, 109704.
Gaston Design Inc., 2022 online. https://gastondesign.com/product/velociraptor-skull/

undescribed dromaeosaurine (Currie and Varricchio, 2004)
Cenomanian-Turonian, Late Cretaceous
Teel Ulaan Jalzai, Bayanshiree Formation, Mongolia
Material- (IGM 100/22) premaxillae, partial maxillae, four dorsal vertebrae (25, 25 mm), five proximal dorsal ribs, first caudal vertebra, second caudal vertebra (29 mm), third caudal vertebra (29 mm), fourth caudal vertebra (31 mm), fifth caudal vertebra (32 mm), sixth caudal vertebra (32 mm), seventh caudal vertebra, eighth caudal vertebra (35 mm), ninth caudal vertebra (37 mm), tenth caudal vertebra (37 mm), eleventh caudal vertebra (37 mm), twelfth caudal vertebra (36 mm), thirteenth caudal vertebra (34 mm), fourteenth caudal vertebra (35 mm), fifteenth caudal vertebra (34 mm), sixteenth caudal vertebra (33 mm), seventeenth caudal vertebra, eighteenth caudal vertebra (30 mm), nineteenth caudal vertebra (29 mm), twentieth caudal vertebra (26 mm), twenty-first caudal vertebra, four chevrons, proximal scapulae, proximal coracoids, incomplete humeri (~186 mm), radii (one partial; 134 mm), ulnae (one incomplete; 146 mm), scapholunares, semilunate carpals, metacarpals I (34, 34 mm), phalanges I-1 (59, 61 mm), incomplete manual unguals I, metacarpals II (one incomplete; 72 mm), phalanges II-1 (one incomplete; 45 mm), phalanges II-2 (one incomplete; 59 mm), incomplete manual unguals II, metacarpals III (one incomplete; 59 mm), phalanges III-1 (one incomplete; 24 mm), phalanges III-2 (17, 17 mm), phalanges III-3 (one incomplete; 43 mm), incomplete manual unguals III, ischium (130 mm), distal femur, tibia (279 mm), incomplete fibula (249 mm), incomplete astragalus, metatarsals I (one incomplete; 39 mm), phalanx I-1 (26 mm), pedal ungual I (~28 mm), distal tarsal III fused to metatarsal II and III (mtII 111, mtIII 134 mm), incomplete pedal ungual II, phalanges III-1 (one partial; 52 mm), partial phalanges III-2, proximal phalanx III-3, pedal ungual III (~44 mm), metatarsal IV (121 mm), partial phalanges IV-1, phalanges IV-2 (31, 30 mm), phalanges IV-3 (24 mm), phalanges IV-4 (22 mm), pedal unguals IV, pedal phalanx x-? (33 mm)
Cenomanian-Turonian, Late Cretaceous
Shine Us Khuduk, Bayanshiree Formation, Mongolia
(IGM 100/23; intended holotype) partial skull, incomplete mandibles, posterior synsacrum, first caudal vertebra, second caudal vertebra (28 mm), third caudal vertebra (33 mm), fourth caudal vertebra (35 mm), two proximal chevrons, posterior ilium, incomplete ischium, proximal metatarsal II, proximal metatarsal III, proximal metatarsal IV
Diagnosis (after Kubota and Barsbold, 2007) convex posterodorsal edge of ilium with posteriorly curved distal end; transversely wide distal end of pedal phalanx II-1.
Other diagnoses- Turner et al. (2012) noted other dromaeosaurids like Velociraptor and Dromaeosaurus also possess two shallow subalveolar grooves on the labial surface of the dentary, which Kubota and Barsbold considered diagnostic.
Comments- These specimens were discovered in the 1970s and must have been referred to Adasaurus early on, as the ischium of IGM 100/22 was used in Barsbold's (1983) pelvic illustration. Similarly, scorings indicate Norell et al. (2001) and Senter (2007) used the specimens when scoring Adasaurus into their TWiG analyses. However, the fact these were new specimens besides the Adasaurus holotype and paratype was not published until Currie and Varricchio (2004), who provided some dental information and specimen numbers. Shortly after in an SVP abstract, Kubota and Barsbold (2007) determined they are from a different taxon, with Adasaurus being a velociraptorine which lived later. They find it to group with Dromaeosaurus and Achillobator based on the low DSDI, and differ from Dromaeosaurus in having less anterior maxillary teeth with lingually twisted mesial carinae, and from Achillobator in lacking double dorsal pleurocoels, lacking caudal pleurocoels, having a low ilium with tapered postacetabular process and a distally placed obturator process. Evans et al. (2013) separated IGM 100/23 (as "Bayanshiree sp. nov.") and Adasaurus in a version of Longrich and Currie's dromaeosaurid analysis, recovering the former as the basalmost velociraptorine and Adasaurus more nested in the subfamily. Kubota (2015) describes and names the taxon in his thesis.
Currie and Varricchio (2004) report there are four premaxillary teeth, eleven maxillary teeth and thirteen dentary teeth in IGM 100/23 and 100/22, but 100/22 doesn't preserve a dentary and only has fragmentary maxillae (Kubota, 2015). They also state teeth have four serrations per mm in both mesial and distal carinae, but the actual values are 3.2-5 and 3.5-5.5 respectively (average 4.25 and 4.19). Jasinowski et al. (2006) list humeri and an ulna for IGM 100/22 (which is only labeled 'dromaeosaur').
References- Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian dinosaurs. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. Yale University Press. 49-67.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons. Studies on the transition from dinosaurs to birds. Indiana University Press. 112-132.
Jasinowski, Russell and Currie, 2006. An integrative phylogenetic and extrapolatory approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder musculature. Zoological Journal of the Linnean Society. 146, 301-344.
Kubota and Barsbold, 2007. New dromaeosaurid (Dinosauria Theropoda) from the Upper Cretaceous Bayanshiree Formation of Mongolia. Journal of Vertebrate Paleontology. 27(3), 102A.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Evans, Larson and Currie, 2013. A new dromaeosaurid (Dinosauria: Theropoda) with Asian affinities from the latest Cretaceous of North America. Naturwissenschaften. 100(11), 1041-1049.
Kubota, 2015. Descriptions of Mongolian dromaeosaurids (Dinosauria: Theropoda) and phylogeny of Dromaeosauridae. PhD thesis, University of Tsubaka. 397 pp.

Dromaeosaurus Matthew and Brown, 1922
Not Dromaeosaurus- The teeth referred to Dromaeosaurus by Russell (1935) from the Milk River Formation of Alberta seem to have a large DSDI, so are more probably Saurornitholestes. Colbert and Russell (1969) tentatively referred a fourth metatarsal (CMN 12072) and a pedal ungual II (CMN 12240) to Dromaeosaurus, as did Ostrom (1969) for the latter element. Paul (1988b) referred the ungual to Saurornitholestes instead, as he hypothesized Dromaeosaurus to have a reduced ungual as in Adasaurus. Both elements were referred to Saurornitholestes by Currie (1995).
Sues (1977) described two dentaries (UA 12091 and 12339) discovered in 1969 and 1974 respectively. He referred these to Dromaeosaurus sp., but they were later realized to be Saurornitholestes (Paul, 1988b).
Rowe et al. (1992) identified cf. Dromaeosaurus teeth (including TMM 43057-314) from the Aguja Formation of Texas. Sankey (1998) later identified Dromaeosaurus teeth from another area of that formation, but these and Rowe et al.'s specimens were referred to Theropoda "family and genus undetermined." They consisted of two tooth fragments (LSUMG 5483 and 6239) which were similar to Dromaeosaurus except in lacking a lingually twisted mesial carina, and were reidentified as tyrannosaurid teeth by Sankey et al. (2005).
Matthew and Brown (1922) referred Lalaeps laevifrons to Dromaeosaurus as D. laevifrons, while Sahni (1972) synonymized it with D. albertensis. It is properly referred to Saurornitholestes langstoni. Matthew and Brown also tentatively referred Laelaps cristatus to the genus as D? cristatus, but this is a Troodon specimen as recognized by Olshevsky (1995). They referred Laelaps explanatus to Dromaeosaurus? sp., which Kuhn (1939) formalized as Dromaeosaurus explanatus. Sahni synonymized it with D. albertensis, but is is a Saurornitholestes specimen. Matthew and Brown also referred Laelaps falculus to Dromaeosaurus? sp., which Olshevsky (1978) formalized as Dromaeosaurus falculus. Sahni also synonymized this species with D. albertensis, but is it probably a juvenile Gorgosaurus or Daspletosaurus. Zapsalis abradens was considered comparable to Dromaeosaurus by Matthew and Brown, and Tracy Ford's taxonomic lists include the combination Dromaeosaurus abradens. More recently, Zapsalis teeth were called ?Dromaeosaurus morphotype A by Sankey et al. (2002), but Zapsalis is now known to be Saurornitholestes premaxillary teeth (Currie and Evans, 2020). Matthew and Brown referred "Coelurus" gracilis to Dromaeosaurus as D? gracilis, but it cannot be compared to Dromaeosaurus though it does seem to be dromaeosaurid. Tracy Ford's lists also include the combination Dromaeosaurus minutus for "Ornithomimus" minutus, an alvarezsaurid. Finally, Paul (1988a) referred Adasaurus to Dromaeosaurus as D. mongoliensis, but now that the genus is better understood it appears to be a more basal dromaeosaurid.
References- Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History. 46(6), 367-385.
Russell, 1935. Fauna of the Upper Milk River beds, Southern Alberta. Transactions of the Royal Society of Canada, series 3, section 4. 29, 115-127.
Kuhn, 1939. Fossillium Catalogus.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus. American Museum Novitates. 2380, 49 pp.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History. 30, 165 pp.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin of the AMNH. 147.
Sues, 1977. Dentaries of small theropods from the Judith River Formation (Campanian) of Alberta, Canada. Canadian Journal of Earth Sciences. 14, 587-592.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus - and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River - Deinonychus, Velociraptor and Saurornitholestes. Hunteria. 2(4), 1-9.
Rowe, Ciffelli, Lehman and Weil, 1992. The Campanian Terlingua local fauna, with a summary of other vertebrates from the Aguja Formation, Trans-Pecos, Texas. Journal of Vertebrate Paleontology. 12, 472-493.
Currie, 1995. New information on the anatomy and relationships of Dromaeosaurus albertensis (Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 15(3), 576-591.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku Saizensen [Dino-Frontline]. 9, 92-119 (part 1); 10, 75-99 (part 2) [in Japanese].
Sankey, 1998. Vertebrate paleontology and magnetostratigraphy of the upper Aguja Formation (Late Campanian), Talley Mountain area, Big Bend National Park, Texas. Unpublished Ph.D. dissertation, Louisiana State University, Baton Rouge. 263 pp.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend, Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology. 76(4), 751-763.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous (Campanian-Maastrichtian), Big Bend National Park, Texas. in Carpenter (ed). The Carnivorous Dinosaurs. 127-152.
Currie and Evans, 2020 (online 2019). Cranial anatomy of new specimens of Saurornitholestes langstoni (Dinosauria, Theropoda, Dromaeosauridae) from the Dinosaur Park Formation (Campanian) of Alberta. The Anatomical Record. 303(4), 691-715.
D. albertensis Matthew and Brown, 1922
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
Holotype- (AMNH 5356) (adult) incomplete skull (240 mm) partial sclerotic ring, mandibles (202.5 mm), hyoids, metacarpal I, distal metatarsal I, phalanx I-1 (27 mm), distal metatarsal II, phalanx II-1 (37 mm), phalanx II-2 (39 mm), distal metatarsal III, distal phalanx III-1, phalanx III-2 (32 mm), partial pedal ungual III, distal phalanx IV-2, phalanx IV-3 (24 mm)
Referred- (ANSP 15827) tooth (Fiorillo and Currie, 1994)
(ANSP 15959) premaxillary tooth (Fiorillo and Currie, 1994)
(ANSP 15963) anterior maxillary tooth (Fiorillo and Currie, 1994)
(ANSP 17639) tooth (Fiorillo and Currie, 1994)
(ANSP 17805) tooth (Fiorillo and Currie, 1994)
(ANSP 17960) tooth (Fiorillo and Currie, 1994)
(ANSP 17798) tooth (Fiorillo and Currie, 1994)
(ANSP 18108) tooth (Fiorillo and Currie, 1994)
(ANSP 18110) tooth (Fiorillo and Currie, 1994)
(CMN 12349) frontal (Sues, 1978)
(RTMP 66.25.16) lateral tooth (14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 66.31.117) tooth (Baszio, 1997)
(RTMP 74.10.91) lateral tooth (Baszio, 1997)
(RTMP 78.9.41) tooth (Baszio, 1997)
(RTMP 79.8.732) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 79.11.165) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 79.14.100) premaxillary tooth (8.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 79.14.1007) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 79.15.2) premaxillary tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 80.8.298) lateral tooth (9.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 80.8.308) lateral tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 80.13.35) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 80.16.2094) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 81.14.60) lateral tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 81.16.161) premaxillary tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 81.16.281) lateral tooth (10 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 81.16.461) premaxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.22.93) premaxillary tooth (7.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 81.26.48) anterior dentary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.26.175) lateral tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 81.27.66) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.11.2) lateral tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.16.361) lateral tooth (Baszio, 1997)
(RTMP 82.18.137) lateral tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.18.250) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.19.185) partial dentary (Currie, 1987a)
(RTMP 83.36.8) maxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 83.67.38) lateral tooth (13.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.67.115) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.89.47) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.89.48) lateral tooth (14.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.92.267) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.135) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.36.332) lateral tooth (6.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.36.336) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.36.337) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.36.338) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.43.4) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.52.10) premaxillary tooth (6.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.56.200) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.59.11) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.59.81) premaxillary tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.66.56) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.68.32) premaxillary tooth (14.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.68.47) premaxillary tooth (9.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.5.39) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.11.20) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.18.99) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.33.52) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.54.67) premaxillary tooth (8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.76.11) lateral tooth (17.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.76.13) premaxillary tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.130.211) lateral tooth (9.5 mm) (Baszio, 1997)
(RTMP 86.130.218) lateral tooth (10.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.184.44) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.136) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.153.56) premaxillary tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.157.30) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.50.45) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.50.66) lateral tooth (Baszio, 1997)
(RTMP 88.50.127) premaxillary tooth (10 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.77.46) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.86.30) premaxillary tooth (7.3 mm) (Baszio, 1997)
(RTMP 88.87.87) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.215.73) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.354) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.404) lateral tooth (Baszio, 1997)
(RTMP 89.77.6) lateral tooth (11 mm), lateral tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.103.11) lateral tooth (15.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.155.2) lateral tooth (14.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.145.1) premaxillary tooth (10.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 92.77.2) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 93.36.460) lateral tooth (16.2), lateral tooth (16.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 93.36.462) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 93.36.472) lateral tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.241) lateral tooth (~15.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.243) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.247) lateral tooth (15.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.266) premaxillary tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.652) lateral tooth (12.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.92.2) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.99.2) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.99.3) premaxillary tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.142.4) lateral tooth (18 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.172.39) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.127.26a) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.127.26b) lateral tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.134.5) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.137.1) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.143.45) lateral tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.151.8) premaxillary tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.151.10) jaw bone with lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.171.40) jaw bone with two lateral teeth (14, 12.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.181.9) premaxillary tooth (7.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.406.4) lateral tooth (11.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.665.59) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.362) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 98.93.172) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 99.55.328) lateral tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP coll.; Bob's site) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP coll.) three teeth (Ryan et al., 2001)
(RTMP coll.) almost a dozen postcranial elements (Currie, 2005)
thirteen specimens (Brinkman et al., 1998)
Late Campanian, Late Cretaceous
Judith River Formation, Montana
Material- (AMNH 8516) lateral tooth (Sahni, 1972)
(AMNH 8517) sixteen teeth (AMNH online)
?(AMNH 8519) tooth (AMNH online)
?(AMNH 8520) posterior dentary tooth (AMNH online)
(AMNH 8521) thirty-four teeth (AMNH online)
(UCMP 137558) tooth (UCMP online)
?(UCMP 154575) manual ungual III (UCMP online)
(YPM PU 21847) (YPM online)
(YPM PU 22245) (YPM online)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- (AMNH 12884) ten teeth (AMNH online)
(AMNH 12885) seven teeth (AMNH online)
(AMNH 12886) fourteen teeth (AMNH online)
(AMNH 12887) six teeth (AMNH online)
(AMNH 12888) tooth (AMNH online)
(UCMP 120850) two teeth (UCMP online)
material (Demar and Breithaupt, 2006)
Campanian-Maastrichtian, Late Cretaceous
Prince Creek Formation, Alaska, US
Material- (AK232-V-073) tooth (Fiorillo and Gangloff, 2000)
(AK238-V-009) tooth (Fiorillo and Gangloff, 2000)
(AK292-V-03) tooth (Fiorillo and Gangloff, 2000)
(AK381-V-052) tooth (Fiorillo and Gangloff, 2000)
(AK306-V-035) tooth (Fiorillo and Gangloff, 2000)
(AK335-V-013) tooth (Fiorillo and Gangloff, 2000)
(AK383-V-180) tooth (Fiorillo and Gangloff, 2000)
(AK385-V-002) tooth (Fiorillo and Gangloff, 2000)
(AK390-V-041) tooth (Fiorillo and Gangloff, 2000)
(AK456-V-024) tooth (Fiorillo and Gangloff, 2000)
(AK456-V-025) tooth (Fiorillo and Gangloff, 2000)
(AK490-V-170) tooth (Fiorillo and Gangloff, 2000)
(AK497-V-001FT) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK83.V-090) tooth (Fiorillo and Gangloff, 2000)
Diagnosis- (after Turner et al., 2012) premaxilla deeper and thicker than other dromaeosaurids; presence of either an enlarged promaxillary fenestra or an extremely anteroventrally placed maxillary fenestra with no promaxillary fenestra depending on interpretation of identity of opening; anteroposterior short lateral lamina of maxilla anterior to antorbital fossa; nine maxillary teeth; nasals with V-shaped suture posteriorly between frontals; deep notches anteriorly on frontal for articulation with lacrimal; tip of frontal flatter and margin of supratemporal fossa less pronounced; postorbital process of frontal sharply demarcated from the dorsomedial orbital margin; quadratojugal stout; dorsal tympanic recess very weakly expressed; moderately developed preotic pendant; expression of anterior tympanic recess and/or basipterygoid recess absent on the basisphenoid or basipterygoid processes; posteromedial process of palatine slender; anterior and posterior tooth denticles subequal in size; anterior carina of maxillary or dentary tooth close to midline of tooth near tip, twists toward lingual surface.
Comments- The holotype was discovered in 1914 but not described until 1922 (Matthew and Brown, 1922) as a new subfamily of tyrannosaurs. Currie (1987a) stated that Dinosaur Park Dromaeosaurus teeth suggest more than one species was present, citing the in prep. at the time Currie et al. (1990). He noted CMN 12349 is more robust than the holotype and that the supratemporal fossa ridge is more strongly curved, implying these could be reason for referring it to the other Dromaeosaurus species. However, Currie et al. never proposed more than one morphotype of Dromaeosaurus once it was published, though they did refer to teeth with Dromaeosaurus-like serrations which lack a twisted mesial carina that "may represent a distinct species of dromaeosaurid, or a gracile, small form of tyrannosaurid." Similar teeth from the Aguja Formation have been referred to juvenile tyrannosaurids.
Is CMN 12349 a therizinosaur frontal?- Sues (1978) identified frontal CMN 12349 as Dromaeosaurus and frontal CMN 12355 as Theropoda indet., although note his plates 7 and 8 are switched so that they are given each others' captions. Currie (1987b) accepted CMN 12349 as Dromaeosaurus, but with regard to CMN 12355 stated that "comparison with Mongolian specimens suggests that it may represent Erlicosaurus (Currie, in preparation)." Currie (1992) again stated CMN 12355 "may represent the segnosaurid Erlicosaurus" (although note that as in Sues' paper it is mislabeled 12349 in his Figure 2) and also referred TMP 1981.016.0231 to Segnosauridae. Currie (2005) listed CMN 12349 as a tentative "therizinosauroid similar to Erlikosaurus", but figured CMN 12355 so probably meant that specimen. Indeed, it seems CMN 12349 was correctly identified as Dromaeosaurus in the first place as it is similar to the holotype and has only been referred to Therizinosauroidea accidentally due to Sues' original plate caption mistake.
References- Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum of Natural History. 46(6), 367-385.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus. American Museum Novitates. 2380, 49 pp.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin of the AMNH. 147.
Sues, 1978. A new small theropod dinosaur from the Judith River Formation (Campanian) of Alberta, Canada. Journal of the Linnean Society: Zoology. 62, 381-400.
Currie, 1987a. Bird-like characteristics of the jaws and teeth of troodontid theropods (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. 7, 72-81.
Currie, 1987b. Theropods of the Judith River Formation of Dinosaur Frovincial Park, Alberta. In Currie and Koster (eds). 4th Symposium of Mesozoic Terrestrial Ecosystems Short Papers. Tyrell Museum of Palaeontology, Drumheller, Alberta. 52-60.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press. 107-125.
Currie, 1992. Saurischian dinosaurs of the Late Cretaceous of Asia and North America. In Mateer and Chen (eds.). Aspects of Nonmarine Cretaceous Geology. China Ocean Press. 237-249.
Fiorillo and Currie, 1994. Theropod teeth from the Judith River Formation (Upper Cretaceous) of south-central Montana. Journal of Vertebrate Paleontology. 14(1), 74-80.
Currie, 1995. New information on the anatomy and relationships of Dromaeosaurus albertensis (Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 15(3), 576-591.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Brinkman, Ryan, and Eberth, 1998. The paleogeographic and stratigraphic distribution of ceratopsids (Ornithischia) in the Upper Judith River Group of western Canada. Palaios. 13, 160-169.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation (Cretaceous) of northern Alaska, with speculations on Arctic dinosaur paleoecology. Journal of Vertebrate Paleontology. 20(4), 675-682.
Ryan, Russell, Eberth and Currie, 2001. The taphonomy of a Centrosaurus (Ornithischia: Ceratopsidae) bone bed from th Dinosaur Park Formation (Upper Campanian), Alberta, Canada, with comments on cranial ontogeny. Palaios. 16, 482-506.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology. 76(4), 751-763.
Currie, 2005. Theropods, including birds. In Currie and Koppelhus (eds.). Dinosaur Provincial Park, a spectacular ecosystem revealed. Part Two, Flora and Fauna from the park. Indiana University Press. 367-397.
Demar and Breithaupt, 2006. The nonmammalian vertebrate microfossil assemblages of the Mesaverde Formation (Upper Cretaceous, Campanian) of the Wind River and Bighorn Basin, Wyoming. in Lucas and Sullivan (eds). Late Cretaceous Vertbrates from the Western Interior. New Mexico Museum of Natural History & Science. Bulletin 35, 33-53.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
D. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
Material- (RTMP 97.99.4) phalanx
Reference- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press. 279-297.
D. sp. nov. (Peng, Russell and Brinkman, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
Material- (RTMP 96.62.1) tooth (Peng et al., 2001)
(RTMP 96.62.2) tooth (Peng et al., 2001)
(RTMP 96.62.3) tooth (Peng et al., 2001)
(RTMP 96.62.69) tooth (Peng et al., 2001)
(RTMP 96.62.70) tooth (Peng et al., 2001)
(RTMP 96.103.1) tooth (Ryan and Russell, 2001)
four specimens (Brinkman et al., 1998)
Comments- These were referred to D. albertensis by Peng et al. (2001), but the teeth have less distinctively displaced mesial carinae than that species.
References- Brinkman, Ryan, and Eberth, 1998. The paleogeographic and stratigraphic distribution of ceratopsids (Ornithischia) in the Upper Judith River Group of western Canada. Palaios. 13, 160-169.
Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages (exclusive of mammals) from the Foremost and Oldman Formations of the Judith River Group (Campanian) of southeastern Alberta: An illustrated guide. Provincial Museum of Alberta, Natural History Occasional Paper. 25, 1-546.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp. 279-297.
D. spp. nov. (Russell, 1933)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
Material- (RTMP 1010) tooth (Baszio, 1997)
(RTMP 1011) lateral tooth (Baszio, 1997)
(RTMP 1012) tooth (Baszio, 1997)
(RTMP 1013) lateral tooth (Baszio, 1997)
(RTMP 1029) lateral tooth (Baszio, 1997)
(RTMP 1035) tooth (Baszio, 1997)
(RTMP 83.45.3) tooth (11.2x4.8x4 mm) (Larson and Currie, 2013)
(RTMP 85.98.5) tooth (9.3x4.7x3 mm) (Larson and Currie, 2013)
(RTMP 98.63.32) tooth (11.8x7.4x4.1 mm) (Larson et al., 2010)
(RTMP 98.63.71) tooth (6.6x4.6x2 mm) (Larson et al., 2010)
(RTMP 98.64.17) tooth (9.6x6x3.2 mm) (Larson et al., 2010)
(RTMP 98.84.3) tooth (8.9x7.4x3.4 mm) (Larson and Currie, 2013)
(RTMP 99.50.116) tooth (11.7x7.7x4 mm) (Larson et al., 2010)
(RTMP 2000.45.40) tooth (10x6.6x2.8 mm) (Larson et al., 2010)
(RTMP 2000.45.82) tooth (11x6.1x3.4 mm) (Larson et al., 2010)
(RTMP 2000.45.102) tooth (9.6x6.5x2.9 mm) (Larson et al., 2010)
(RTMP 2001.45.83) tooth (16.6x7.9x4.7 mm) (Larson et al., 2010)
(RTMP 2002.45.50) tooth (11.2x6.7x3.3 mm) (Larson et al., 2010)
(RTMP 2003.45.60) tooth (9.6x5.4x2.4 mm) (Larson et al., 2010)
(RTMP 2005.7.5) tooth (8.2x3.6x2.4 mm) (Larson and Currie, 2013)
(RTMP 2009.122.2) tooth (8.8x5.4x3 mm) (Larson and Currie, 2013)
(RTMP 2009.122.4) tooth (9.9x6.6x3.2 mm) (Larson and Currie, 2013)
(RTMP 2009.136.8) tooth (Larson et al., 2010)
(RTMP coll.) two teeth (Baszio, 1997)
tooth (Russell, 1933)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- tooth (Baszio, 1997)
three specimens (Tokaryk and Bryant, 2004)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta
Material- (RTMP 81.31.99) tooth (Ryan and Russell, 2001)
(UA Alberta 1:98) tooth (Baszio, 1997)
(UA Alberta 1:99) tooth (Baszio, 1997)
(UA Alberta 3:103) tooth (Baszio, 1997)
(UA Alberta 3:104) lateral tooth (Baszio, 1997)
(UA JLE 62:113) tooth (Baszio, 1997)
(UA KUA-1:106) lateral tooth (Baszio, 1997)
(UA KUA-1:119) tooth (Baszio, 1997)
(UA KUA-2:110) tooth (Baszio, 1997)
Comments- Baszio (1997) notes these teeth are smaller than Dinosaur Park specimens and differ in that they are slightly more convex labially than lingually. Larson and Currie (2013) found that upper Horseshoe Canyon teeth are quantitatively different from Dinosaur Park teeth, and that lower Horseshoe Canyon teeth are not but lack the twisted mesial carina present in D. albertensis.
References- Russell, 1933. The Cretaceous-Tertiary transition of Alberta. Transactions of the Royal Society of Canada, series 3. 26(4), 121-156.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp. 279-297.
Tokaryk and Bryant, 2004. The fauna from the Tyrannosaurus rex excavation, Frenchman Formation (Late Maastrichtian), Saskatchewan. Summary of Investigations 2004, Volume 1. Saskatchewan Geological Survey, Saskatchewan Industry Resources, Miscellaneous Report 2004-4, 1-12.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta, with special reference to the Albertosaurus sarcophagus bonebed. Canadian Journal of Earth Sciences. 47(9), 1159-1181.
Larson and Currie, 2013. Multivariate analyses of small theropod dinosaur teeth and implications for paleoecological turnover through time. PloS ONE. 8(1), e54329.
D. sp. (Mongelli and Varricchio, 1998)
Early Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- teeth
Reference- Mongelli and Varricchio, 1998. Theropod teeth of the Lower Two Medicine Formation (Campanian) of northwestern Montana. Journal of Vertebrate Paleontology. 18(3), 64A.
D. sp. (Wilson, 2008)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, South Dakota, US
Material- two teeth (Wilson, 2008)
teeth, elements (DePalma, 2010)
References- Wilson, 2008. Comparative taphonomy and paleoecological reconstruction of two microvertebrate accumulations from the Late Cretaceous Hell Creek Formation (Maastrichtian), eastern Montana. Palaios. 23, 289-297.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis, University of Kansas. 227 pp.
D. sp. (Stokosa, 2005)
Maastrichtian, Late Cretaceous
Fox Hills Formation, South Dakota, US
Material- (SDSM 14516) tooth fragment (Stokosa, 2005)
Reference- Stokosa, 2005. Enamel microstructure variation within the Theropoda. in Carpenter (ed). The Carnivorous Dinosaurs. 163-178.
D. sp. (Lillegraven and Eberle, 1999)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
Material- (UW 26254)
(UW 26580)
(UW 27203)
Reference- Lillegraven and Eberle, 1999. Vertebrate faunal changes through Lancian and Puercan time in southern Wyoming. Journal of Paleontology. 73(4), 691-710.
D. sp. nov. (Breithaupt, 1982)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- ?(AMNH 25268) astragalus (AMNH online)
(UA 125) lateral tooth (Baszio, 1997)
(UA 126) lateral tooth (Baszio, 1997)
(UA 128) lateral tooth (Baszio, 1997)
(YPM PU 22312) (YPM online)
two specimens (Breithaupt, 1982)
?specimen (Derstler, 1995)
Comments- Baszio (1997) notes these teeth are much smaller than D. albertensis with more (6) serrations per mm.
References- Breithaupt, 1982. Paleontology and paleoecology of the Lance Formation (Maastrichtian), east flank of Rock Springs Uplift, Sweetwater County, Wyoming. Contributions to Geology, University of Wyoming. 21(2), 123-151.
Derstler, 1995. The Dragons' Grave: An Edmontosaurus bonebed containing theropod egg shells and juveniles, Lance Formation (uppermost Cretaceous), Niobrara County, Wyoming. Journal of Vertebrate Paleontology. 15(3), 26A.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
D. sp. (Williamson, 2001; described by Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-33148) anterior tooth (6.5x3.8x2.5 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33489) tooth (Williamson and Brusatte, 2014)
References- Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.

Dromaeosauroides Christiansen and Bonde, 2003
D. bornholmensis Christiansen and Bonde, 2003
Late Berriasian, Early Cretaceous
Jydegaard Formation, Denmark
Holotype- (MGUH DK 315) anterior dentary tooth (21.7x9.7x6.6 mm)
Referred- (MGUH DK 559) tooth (15 mm) (Bonde, 2012)
Diagnosis- 25% larger than Dromaeosaurus albertensis; 6 distal serrations per mm compared to 3-3.5/mm in D. albertensis.
Comments- The holotype was discovered in 2000 and mentioned as Dromaeosauridae indet. by Bonde (2001). The BW/FABL is .68, while the crown is fairly tall and recurved. Both carinae are lingually displaced, the mesial carina only extending down 1/3 of the crown. Mesial serrations decrease in size toard the base, with 6.1/mm present apically. Distal serrations are square, labiolingually broad, lack blood grooves and have a density of 6/mm.
References- Bonde, 2001. A Berriasian "Wealden fauna" from Bornholm, Denmark. Palaeontological Association 45th annual meeting. 4.
Christiansen and Bonde, 2003. The first dinosaur from Denmark. Neues Jahrbuch der Geologie und Palaontologie Abhandlungen. 227(2), 287-299.
Bonde and Christiansen, 2003. New dinosaurs from Denmark. Comptes Rendus Palevol. 2, 13-26.
Bonde, 2012. Danish dinosaurs: A review. In Godefroit (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 434-451.

Velociraptorinae Barsbold, 1983
Definition- (Velociraptor mongoliensis <- Dromaeosaurus albertensis) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017; modified from Sereno, 1998)
Other definitions- (Velociraptor mongoliensis <- Microraptor zhaoianus, Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus) (Turner et al., 2012)
= Itemirinae Kurzanov, 1976 vide Martyniuk, 2012
= Velociraptorinae sensu Turner et al., 2012
Definition- (Velociraptor mongoliensis <- Microraptor zhaoianus, Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus)
Comments- Since Turner et al. (2012) uses a stem-based definition for Dromaeosauridae, the node stem triplet advocated by Padian et al. (1999) no longer functions. Turner et al.'s definition is slightly superior to Sereno's 1998 one by adding Microraptor, Unenlagia and Passer, but I think the chance of a (Dromaeosaurus (Velociraptor, Passer)) topology is very low, and wouldn't have a problem with microraptorians being velociraptorines. But since Unenlagiinae is active, I suppose it's best to keep subfamilies out of subfamilies.
Kurzanov (1976) erected Itemiridae for Itemirus only, sister to tyrannosaurids than to dromaeosaurids. Martynuik (2012) both defined Itemiridae as excluding Dromaeosaurus, and created an undefined Itemirinae within Dromaeosauridae with no explicit content (besides Itemirus). Here (as in Longrich and Currie, 2009), Itemirus is recovered as closer to Velociraptor than to Dromaeosaurus. This would technically make Itemirinae a senior synonym of Velociraptorinae, as the latter family-level taxon was named 7 years later. Yet the Itemirus holotype has only been found to have this position in one published analysis, while it was recovered closer to Dromaeosaurus by Sues and Averianov (2014). Thus until there is a consensus on the position of Itemirus, the subfamily Velociraptorinae is used here.
References- Kurzanov, 1976. Braincase structure in the carnosaur Itemirus n. gen., and some aspects of the cranial anatomy of dinosaurs. Paleontological Journal. 1976, 361-369.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Sues and Averianov, 2014. Dromaeosauridae (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan and the phylogenetic position of Itemirus medullaris Kurzanov, 1976. Cretaceous Research. 51, 225-240.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold, Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature. 552, 395-399.

Shri Turner, Montanari and Norell, 2021
= "Ichabodcraniosaurus" Novacek, 1996
S. devi Turner, Montanari and Norell, 2021
Late Campanian, Late Cretaceous
Khulsan, Baruungoyot Formation, Mongolia
Material- (IGM 100/980) third cervical vertebra (28 mm), fourth cervical vertebra (28.3 mm), fifth cervical vertebra (30.6 mm), sixth cervical vertebra (24.2 mm), seventh cervical vertebra (19.55 mm), eighth cervical vertebra (18.5 mm), ninth cervical vertebra (19.6 mm), tenth cervical vertebra (19.1 mm), few cervical ribs, first dorsal vertebra (21.5 mm), second dorsal vertebra (19.5 mm), third dorsal vertebra (17.5 mm), fourth dorsal vertebra (16.3 mm), fifth dorsal vertebra (17.25 mm), sixth dorsal vertebra (17.5 mm), seventh dorsal vertebra (17 mm), eighth dorsal vertebra (16.9 mm), ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra, twelfth dorsal vertebra, thirteenth dorsal vertebra (15.1 mm), five proximal dorsal ribs, fragmentary sacrum, first caudal vertebra (18.8 mm), second caudal vertebra (18.8 mm), third caudal vertebra (20 mm), fourth caudal vertebra (21.05 mm), fifth caudal vertebra (23.4 mm), sixth caudal vertebra (~21 mm), seventh caudal vertebra, eighth caudal vertebra, five chevrons, ilia (183.8 mm), pubes (~245 mm; one partial), ischia (~118 mm), femur (205 mm), incomplete tibiae (246 mm), incomplete fibula, partial astragali, partial calcaneum, metatarsal I (30.9 mm), phalanx I-1 (22.5 mm), pedal ungual I, incomplete metatarsal II (~90 mm), phalanx II-1 (25.8 mm), phalanx II-2, pedal ungual II (91.3 mm on curve), incomplete metatarsal III (~117 mm), distal phalanx III-1 (~47 mm), phalanx III-2 (30 mm), phalanx III-3 (28.4 mm), pedal ungual III (49.4 mm on curve), incomplete metatarsal IV (~124 mm), fragmentary phalanx IV-1 (~41.5 mm), phalanx IV-2 (27.1 mm), phalanx IV-3 (23.4 mm), phalanx IV-4 (22.4 mm), pedal ungual IV (36.5 mm on curve), partial metatarsal V
Diagnosis- (after Turner et al., 2021) first dorsal epipophysis large and overhangs posterior margin of postzygapophysis; two dorsoventrally aligned pleurocoels on first and second dorsals; posteriorly inclined scar on lateral surface of neural arch.
Comments- This specimen was discovered by Norell in July 1991 and mentioned by Norell et al. (1992) as "a very well preserved, articulated postcranial skeleton". Novacek (1996) wrote in a popular book ""It lacked a precious head. Although Mark [Norell] was understandably upset, it was still an important find. Informally called "Ichabodcraniosaurus," the skeleton remains under investigation in New York to this day." Norell and Makovicky (1999) mention IGM 100/980 several times, stating it can be referred to Dromaeosauridae based on caudal and pedal characters, but that insufficient elements are preserved to refer it to Velociraptor. A photo of the pelvis and adjacent vertebrae is included as figure 24. Norell (DML, 2002) confirmed that "Ichabodcraniosaurus" and IGM 100/980 are the same specimen. Turner (2008) described its anatomy as a new taxon of dromaeosaurid, though he did not name it. While IGM 100/980 is partially scored in his matrix, Turner didn't include the specimen in his analysis. It was officially described and named Shri devi by Turner et al. (2021), who included it in a version of the TWiG analysis which recovered it as a velociraptorine sister to Velociraptor.
References- Norell, Clark and Perle, 1992. New dromaeosaur material from the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 12(3), 45A.
Novacek, 1996. Dinosaurs of the Flaming Cliffs. Anchor Books. 367 pp.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 45 pp.
Norell, DML 2002. https://web.archive.org/web/20210120042909/http://dml.cmnh.org/2002Feb/msg00648.html
Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis. Columbia University. 666 pp.
Turner, Montanari and Norell, 2021. A new dromaeosaurid from the Late Cretaceous Khulsan locality of Mongolia. American Museum Novitates. 3965, 46 pp.

Velociraptor Osborn, 1924b
= "Ovoraptor" Osborn, 1924a
Diagnosis- (after Turner et al., 2012; for Velociraptor mongoliensis, at the time including IGM 100/982) premaxilla with long maxillary process reaching well beyond caudal margin of external nares; first and second premaxillary teeth larger than third and fourth; anterior border of internal antorbital fenestra broadly rounded; maxillary fenestra not located in a posteriorly open depression; frontal long, almost four times longer than wide across orbital portion, and almost four times as long as parietal; dentary very shallow, its depth constituting 1/8 to 1/7 of its length, ventral margin convex; flange-like m. ambiens tubercle located proximally on anterior face of pubis.
Not Velociraptor- Besides the remains listed below, V. osmolskae, and Bohlin's (1953) and Young's (1958) questionably referred specimens, many remains have been referred to Velociraptor that probably do not belong there. In 1982, Carpenter stated that he and Paul (in prep.) would show Velociraptor occurred in North America. Though the collaborative publication never surfaced, Paul (1984, 1988, 1988) did synonymize Deinonychus and Saurornitholestes with Velociraptor. Recent analyses indicate these do not form a monophyletic clade with respect to Dromaeosaurus however (Currie and Varricchio, 2004; Senter, 2007), and thus almost all non-microraptorian, non-unenlagiine dromaeosaurids would be a single genus under this scheme. Still, this has led to the occassional identification of Velociraptor in American sediments (eg. AMNH website). Dong et al. (1989) state Velociraptor material was discovered in the Iren Dabasu Formation of Inner Mongolia, China, and Dong (1992) specifies "teeth of Velociraptor". Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in the Iren Dabasu" and that "Most of these can be attributed to Velociraptor..." Yet no rationale was presented, and the only two specified Iren Dabasu dromaeosaurid elements in the literature are clearly not Velociraptor (AMNH 6572 is twice the size, while IVPP 270790-4 is different from most dromaeosaurid teeth). Matsukawa and Obata (1994) report through personal communication with Mateer (1992) that cf. Velociraptor mongoliensis was discovered in the Zouyun Formation, though this is much too early to actually be that genus. Norell et al. (1994) identified two juvenile skulls as Velociraptor, but these seem more likely to be Byronosaurus (Norell and Makovicky, 1999). Nessov (1995) notes that unguals attributed to Velociraptor are known in the Santonian Syuk-Syuk Formation of Kazakhstan, but these reports have not been verified. The Tsaagan holotype was originally referred to Velociraptor (e.g. Webster, 1996) before it was identified as a new genus by Norell et al. (2006). By 2004 a skull cast was in circulation from Gaston Design Inc., generally referred to Velociraptor (e.g. Witmer, online 2012), but Powers et al. (2020) used maxillary proportions to find it is closer to Tsaagan. Turner et al. (2007) describe an ulna as IGM 100/981, referring it to Velociraptor mongoliensis based on sharing several unlisted characteristics, but this specimen is actually IGM 100/3503 (Napoli et al., 2021) and is here placed in Tsaagan.
References- Osborn, 1924a. The discovery of an unknown continent. Natural History. 24(2), 133-149.
Osborn, 1924b. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
Bohlin, 1953. Fossil reptiles from Mongolia and Kansu. Sino-Swedish Expedition Publication. 37, 1-105.
Young, 1958. The first record of dinosaurian remains from Shansi. Vertebrata PalAsiatica. 2(4), 231-236.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek formations and a description of a new species of theropod. Contributions to Geology, University of Wyoming. 20(2), 123-134.
Paul, 1984. The archosaurs: A phylogenetic study. In Reif and Westphal (eds.). Third Symposium on Mesozoic Terrestrial Ecosystems. Tubingen. 175-180.
Paul, 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon and Schuster, New York. 464 pp.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus - and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River - Deinonychus, Velociraptor and Saurornitholestes. Hunteria. 2(4), 1-9.
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic of China. Cretaceous Research. 14, 127-144.
Matsukawa and Obata, 1994. Cretaceous, a contribution to dinosaur facies in Asia based on molluscan paleontology and stratigraphy. Cretaceous Research. 15, 101-125.
Norell, Clark, Dashzeveg, Barsbold, Chiappe, Davidson, McKenna and Novacek, 1994. A theropod dinosaur embryo, and the affinities of the Flaming Cliffs dinosaur eggs. Science 266, 779-782.
Nessov, 1995. Dinosaurs of Northern Eurasia: new data about assemblages, ecology and paleobiogeography. Scientific Research Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia. 156 pp. + 14 pl. [in Russian with short English, German, and French abstracts].
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 45 pp.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons. Studies on the transition from dinosaurs to birds. Indiana University Press. 112-132.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda). Journal of Systematic Palaeontology. 5(4), 429-463.
Turner, Makovicky and Norell, 2007. Feather quill knobs in the dinosaur Velociraptor. Science. 317, 1721.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Witmer, online 2012. https://people.ohio.edu/witmerl/collections/Theropods/Velociraptor_gaston.htm
Powers, Sullivan and Currie, 2020. Re-examining ratio based premaxillary and maxillary characters in Eudromaeosauria (Dinosauria : Theropoda): Divergent trends in snout morphology between Asian and North American taxa. Palaeogeography, Palaeoclimatology, Palaeoecology. 547, 109704.
Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021. A new dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central Mongolia. American Museum Novitates. 3982, 47 pp.
V. mongoliensis Osborn, 1924b
= "Ovoraptor djadochtari" Osborn, 1924a
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Holotype- (AMNH 6515) skull (176 mm), mandibles (175 mm), teeth (4.15-7.91 mm), manual phalanx III-3 (39 mm), manual ungual III (45 mm on curve)
Referred- (IGM coll.; 980826 BDz Velocira ENKH) specimen including dentary, radius, ulna, semilunate carpal, metacarpal I, distal phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, femur, tibia, fibula and fragments (Suzuki and Watabe, 2000)
Late Campanian, Late Cretaceous
Chimney Buttes, Djadokhta Formation, Mongolia
(IGM 100/986) (~25 kg) cranial fragments, cervical vertebrae, three anterior dorsal vertebrae, dorsal vertebrae 8-13, dorsal ribs, sacrum, caudal vertebrae 1-26, chevrons, proximal scapulacoracoid, sternal plate, distal humerus, proximal radius, proximal ulnae, metacarpal I, phalanx I-1 (47.1 mm), manual ungual I, phalanx II-1 (33.2 mm), manual ungual II (48.8 mm curve), phalanx III-1 (19 mm), phalanx III-2 (11.1 mm), phalanx III-3 (33.2 mm), manual ungual III, ilia (145.8 mm), pubes (213, 208 mm), ischia (115.1, 118.8 mm), femur (238 mm), tibia (255 mm), fibula, astragalocalcaneum, distal tarsal III, distal tarsal IV, metatarsal I (27.9 mm), metatarsal II (84.8 mm), phalanx II-1 (26.6 mm), phalanx II-2 (28 mm), pedal ungual II, metatarsal III (99.1 mm), phalanx III-1 (44 mm), phalanx III-2 (27.4 mm), phalanx III-3 (12.2 mm), metatarsal IV (91.6 mm), phalanx IV-1 (30.1 mm), phalanx IV-2 (23 mm), phalanx IV-3 (18.6 mm), phalanx IV-4 (19.6, 18.6 mm), pedal ungual IV (25.6 mm straight), metatarsal V (40.1 mm) (Norell and Mackovicky, 1999)
Late Campanian, Late Cretaceous
Tugrikin Shire, Djadokhta Formation, Mongolia
(IGM 100/24) incomplete skull, mandibles, teeth (7.35-9.44 mm), few fragmentary postcranial elements (Barsbold, 1983)
(IGM 100/25; fighting dinosaurs specimen) (2.07 m, 15 kg) skull (230 mm), mandibles, cervical vertebrae 1-10, cervical ribs, dorsal vertebrae 1-13, dorsal ribs, five uncinate processes, gastralia, sacrum, caudal vertebrae 1-43, chevrons, scapulae, coracoids, sternal plates (95 mm), humeri (~147 mm), radii, ulnae, semilunate carpal, metacarpal I (25 mm), phalanx I-1 (44 mm), manual ungual I (47 mm), metacarpal II (58 mm), phalanx II-1 (35 mm), phalanx II-2 (48 mm), manual ungual II (~51 mm), metacarpal III (50 mm), phalanx III-1 (20 mm), phalanx III-2 (12 mm), phalanx III-3 (35 mm), manual ungual III (33 mm), ilium (~188 mm), pubis, ischium, femur (208 mm), tibia (243 mm), fibula, metatarsus (101 mm), pes including phalanx II-1 (21 mm), phalanx II-2 (24 mm), pedal ungual II (52 mm) (Kielan-Jaworowska and Barsbold, 1972; described by Barsbold, 1974)
(IGM 100/54; 940728 TS-V NAR) (subadult) skull (~230 mm), mandibles, first-ninth cervical vertebrae, twelve dorsal vertebrae, sixteen dorsal ribs, five uncinate processes, gastralia, sacrum, first-third caudal vertebrae, scapula, partial coracoids, sternal plates, humerus, radius, ulna, semilunate carpal, metacarpal I (26 mm), phalanx I-1 (49 mm), manual ungual I (46 mm), metacarpal II (56 mm), phalanx II-1 (37 mm), phalanx II-2 (52 mm), manual ungual II (~37 mm), metacarpal III (50 mm), phalanx III-1 (20 mm), phalanx III-2 (14 mm), phalanx III-3 (36 mm), manual ungual III (30 mm), ilia, proximal pubis, femora (192, 194 mm), tibiae, fibulae, astragalocalcanea, metatarsal I, proximal pedal ungual I, distal tarsal III fused with metatarsals II and metatarsals III (one incomplete), phalanx II-1, phalanges II-2, pedal ungual II, phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV, metatarsal V (Watabe and Suzuki, 2000)
(IGM 100/976) (adult) partial skull including premaxillae, incomplete maxillae, partial nasals, frontals, partial parietals, braincase, incomplete mandible, atlas, axis, third cervical vertebra, fourth cervical vertebra, seventh cervical vertebra (23.6 mm), eighth cervical vertebra, ninth cervical vertebra (26.3 mm), cervical ribs, third dorsal vertebra (15.2 mm), scapulae, coracoids, furcula, sternal plates, sternal ribs, proximal humeri (Norell et al., 1992)
(IGM 100/985) (subadult) skull fragments, vertebral fragments, dorsal rib fragments, twelve rows of gastralia, sacrum (89.2 mm), first caudal vertebra (17 mm), first chevron (33.6 mm), second caudal vertebra (19.1 mm), third caudal vertebra (20.3 mm), fourth caudal vertebra (23.4 mm), fifth caudal vertebra (24.5 mm), sixth caudal vertebra (24.1 mm), sixth chevron (26.4 mm), seventh caudal vertebra (23.9 mm), eighth caudal vertebra (25.9 mm), sternal plates (71 mm), proximal humeri, manual elements including unguals, ilia (126.8, 132.8 mm), pubis (167 mm), ischia (97.7 mm), femur, partial tibia, partial fibula, partial astragalus, distal tarsal III, metatarsi I (22, 21.4 mm), phalanges I-1 (16.9 mm), pedal ungual I (20.3 mm), metatarsi II (71, 71.2 mm), phalanges II-1 (23.6 mm), phalanges II-2 (24.2, 24 mm), pedal ungual II (64.8 mm on curve), metatarsi III (85.6, 86.2 mm), phalanges III-1 (37.6 mm), phalanges III-2 (24.7, 24.9 mm), pedal ungual III (32.3 mm), metatarsi IV (79, 78.2 mm), phalanges IV-1 (26.8 mm), phalanges IV-2 (21, 21.3 mm), phalanges IV-3 (17, 16.8 mm), phalanges IV-4 (16, 16.1 mm), pedal unguals IV (32.3, 31 mm on curve), metatarsal V (Norell and Mackovicky, 1997)
(PIN 3143/8; Moscow specimen) incomplete skull, mandible (Barsbold and Osmolska, 1999)
Diagnosis- (after Kundrat, 2004) differs from V. sp. nov. in- triangle-shaped auricular fossa; the utriculo-saccular and anterior semicircular canal prominence does not project too far medially; hollowed area behind posterior internal acoustic fossa absent; minor expansion of medulla oblongata either anteriorly between the roots of III-VI and VII, and ventrolaterally below the metotic fissure; endoneurocranial base not clearly corrugated longitudinally; transverse convexity of endoneurocranial base at anteriormost point of metotic fissure absent; dorsal tympanic recess less expanded anteriorly; absence of crista intertuberalis; shallow parabasisphenoid recess is separated by low median ridge; paracondylar recesses projected ventrally; foramen magnum considerably compressed dorsoventrally.
(after Turner et al., 2012) supratemporal fossa and fenestra subcircular, bound by laterally convex supratemporal arcade (unknown in IGM 100/982); lateral wall of braincase possesses a deep prootic recess (unknown in IGM 100/982); V-shaped furcula with reduced and asymmetrically developed hypocledium (unknown in IGM 100/982); well-developed obturator tuberosity (unknown in IGM 100/982); rounded longitudinal ischial ridge (also in Deinonychus; unknown in IGM 100/982).
Comments- The holotype was discovered on August 11 1923 and originally named Ovoraptor djadochtari by Osborn (1924a) in a magazine article. Osborn merely describes it as "a wonderfully alert little egg-snatcher, hence Ovoraptor of the Djadochta formation", a "swift-moving, carnivorous dinosaur." However, this publication lacked a proper description, and Osborn (1924b) decided to name the taxon Velociraptor mongoliensis in his official description later that year. The skull was redescribed by Sues in 1977.
IGM 100/25 was discovered in early August 1971 (Kielan-Jaworowska and Barsbold, 1972), initially mentioned as "the skeleton of a Velociraptor mongoliensis preserved together with the skeleton of Protoceratops andrewsi" and photographed as plate II figure 2. It was described briefly by Barsbold (1974), with the skull, sterna, pedal digit II illustrated by him in 1983. Now known as the "fighting dinosaurs" specimen, it has been analyzed by several authors (Osmolska, 1993; Unwin et al., 1995; Carpenter, 2000). Barsbold and Osmolska (1999) described the skull in detail, while Currie et al. (2016) provide manual measurements for it. Paul (1984, 1988) first noticed unicinate processes, which were described in detail by Codd (2004).
Barsbold (1983) also mentioned another specimen, IGM 100/24, a "partially destroyed skull and remains of postcranial skeleton". He illustrates the medial mandible. Barsbold and Osmolska (1999) note it was discovered by a Mongolian expedition, and illustrate the skull and braincase.
PIN 3143/8 was discovered at Tugrikin Shire by the Soviet-Mongolian expedition, who also found material at Ikh Shunkht (Ivakhnenko and Kurzanov, 1988 misstyped as 1982 in Watabe et al., 2010). Barsbold and Osmolska (1999) state the postcrania of IGM 100/24 and 100/25 "will be published at a later date (Barsbold & Osmolska in preparation)."
IGM 100/976 was discovered in 1991 and mentioned by Norell et al. (1992). Norell et al. (1997) briefly described the furcula, while the other postcrania were described in depth by Norell and Makovicky (1999), the braincase was described by Norell et al. (2004) and the endocast by King et al. (2020). IGM 100/985 and 100/986 were discovered in 1993, and described by Norell and Makovicky in 1997 and 1999 respectively. Though IGM 100/985 was originally only referred to Dromaeosauridae indet., it was referred to Velociraptor specifically by 1999.
Three Velociraptor specimens were listed by Watabe and Suzuki (2000) as being from Tugrikin Shire found between July 21 and August 1 1994. The best described is IGM 100/54, with the field number 940728 TS-V NAR from Tugrikin Shire V, photographed by them as "complete skeleton (fore limb shown) of Velociraptor in Tugrikin Shire". This is complete except the distal tail and left forelimb, with the skull and mandible photographed as plate 1 figure 3 in Tsogtbaatar (2004), the skeleton as figure 2 in that publication and in higher resolution in Saneyoshi et al. (2011). Hone et al. (2012) and Codd (2004) provide good photographs of the torso, while Hattori (2016) describes and illustrates a metatarsus. Hone et al. described a pterosaur (?azhdarchid) long bone fragment eaten by the specimen, while Saneyoshi et al. described taphonomy. Jasinoski et al. (2006) used this specimen (listed as IGM 94.07.28) for pectoral and forelimb information. Currie et al. (2016) lists manual measurements for MPC-D unnumbered specimen, which are here guessed to be IGM 100/54 given the <10% length difference using the un-ideal scale in Senoyoshi et al.'s photo (scale bar inserted digitally over an oblique photograph).
A specimen collected in August 26 1998 and reported by Suzuki and Watabe (2000), "Velociraptor forelimbs" with field number 980826 BDz Velocira ENKH discovered at Bayn Dzak, was listed as "Velociraptor lower jaw and partial postcranial bones." The in situ specimen 980826 BDz Velocira ENKH is figured as "Velociraptor skeleton in Bayn Dzak (central Gobi region)."
References- Osborn, 1924a. The discovery of an unknown continent. Natural History. 24(2), 133-149.
Osborn, 1924b. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
Kielan-Jaworowska and Barsbold, 1972. Narrative of the Polish-Mongolian Palaeontological Expeditions, 1967-1972. Palaeontologia Polonica. 27, 1-12.
Barsbold, 1974. Dueling dinosaurs. Priroda. 2, 81-83. [in Russian]
Sues, 1977. The skull of Velociraptor mongoliensis, a small Cretaceous theropod dinosaur from Mongolia. Palontologische Zeitschrift. 51, 173-184.
Barsbold, 1983. Avian features in the morphology of predatory dinosaurs. Transactions of the Joint Soviet Mongolian Paleontological Expedition. 24, 96-103.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions of the Joint Soviet-Mongolian Paleontological Expedition. 19, 5-119.
Paul, 1984. The archosaurs: A phylogenetic study. In Reif and Westphal (eds.). Third Symposium on Mesozoic Terrestrial Ecosystems. Tubingen. 175-180.
Ivakhnenko and Kurzanov, 1988. Geological structure and age of Udan-Sair and Shara-Tsav localities. Trudy Sovmestnoi Sovetsko-Mongolʼkoi Paleontologicheskoi Ekspeditsii. 34, 100105.
Paul, 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide. Simon and Schuster, New York. 464 pp.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus - and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River - Deinonychus, Velociraptor and Saurornitholestes. Hunteria. 2(4), 1-9.
Norell, Clark and Perle, 1992. New dromaeosaur material from the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 12(3), 45A.
Osmolska, 1993. Were the Mongolian "fighting dinosaurs" really fighting? Revue de Paleobiologie. 7, 161-162.
Unwin, Perle and Trueman, 1995. Protoceratops and Velociraptor preserved in association: Evidence from predatory behavior in predatory dinosaurs? Journal of Vertebrate Paleontology. 15(3), 57A.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton: Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Norell, Makovicky and Clark, 1997. A Velociraptor wishbone. Nature. 389, 447.
Feduccia and Martin, 1998. Theropod-bird link reconsidered. Nature. 391, 754.
Norell and Makovicky, 1998. A revised look at the osteology of dromaeosaurs: evidence from new specimens of Velociraptor. Journal of Vertebrate Paleontology. 18(3), 66A.
Barsbold and Osmólska, 1999. The skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica. 44(2), 189-219.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 45 pp.
Carpenter, 2000. Evidence of predatory behavior by carnivorous dinosaurs. GAIA. 15, 135-144.
Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1998. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 83-98.
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 30-44.
Smith, 2002. An examination of dental morphology and variation in theropod dinosaurs: Implications for the identification of shed teeth. PhD dissertation. University of Pennsylvania.
Codd, 2004. The uncinate processes in birds and their implications for the breathing mechanics of maniraptoran dinosaurs. PhD thesis, Rheinischen Friedrich-Wilhelms-Universität. 108 pp.
Kundrat, 2004. Two morphotypes of the Velociraptor neurocranium. ICVM-7 Abstracts. Journal of Morphology. 260(3), 305.
Norell, Makovicky and Clark, 2004. The braincase of Velociraptor. In Currie, Koppelhus, Shugar and Wright (eds.). Feathered Dragons: Studies on the Transition from Dinosaurs to Birds. Indiana University Press. 133-143.
Tsogtbaatar, 2004. Fossil specimens prepared in Mongolian Paleontological Center 1993-2001. Hayashibara Museum of Natural Sciences Research Bulletin. 2, 123-128.
Parsons and Parsons, 2005. A comparison of postcranial features found within the ontogenies of the maniraptoran theropod dinosaurs Deinonychus antirrhopus (Saurischia, Theropoda) and Velociraptor mongoliensis (Saurischia, Theropoda). Journal of Vertebrate Paleontology. 25(3), 99A.
Jasinowski, Russell and Currie, 2006. An integrative phylogenetic and extrapolatory approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder musculature. Zoological Journal of the Linnean Society. 146, 301-344.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
Garcia, Erickson, Curry Rogers and Norell, 2007. Longevity and growth patterns in the dromaeosaurid Velociraptor mongoliensis inferred from long bone histology. Journal of Vertebrate Paleontology. 27(3), 79A.
Gombert, 2008. Craniofacial ontogeny in Velociraptor mongoliensis. Journal of Vertebrate Paleontology. 28(3), 85A.
Watabe, Tsogtbaatar, Suzuki and Saneyoshi, 2010. Geology of dinosaur-fossil-bearing localities (Jurassic and Cretaceous: Mesozoic) in the Gobi Desert: Results of the HMNS-MPC Joint Paleontological Expedition. Hayashibara Museum of Natural Sciences Research Bulletin. 3, 41-118.
Saneyoshi, Watabe, Suzuki and Tsogtbaatar, 2011. Trace fossils on dinosaur bones from Upper Cretaceous eolian deposits in Mongolia: Taphonomic interpretation of paleoecosystems in ancient desert environments. Palaeogeography, Palaeoclimatology, Palaeoecology. 311, 38-47.
Hone, Tsuihiji, Watabe and Tsogtbaatar, 2012. Pterosaurs as a food source for small dromaeosaurs. Palaeogeography, Palaeoclimatology, Palaeoecology. 331-332, 27-30.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Barsbold, 2016. "The Fighting Dinosaurs": The position of their bodies before and after death. Paleontological Journal. 50(12), 1412-1417.
Currie, Funston and Osmólska, 2016. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
King, Sipla, Georgi, Balanoff and Neenan, 2020. The endocranium and trophic ecology of Velociraptor mongoliensis. Journal of Anatomy. 237(5), 861-869.
V. sp. nov. (Norell and Makovicky, 1997)
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Material- (IGM 100/982) (adult) incomplete skull (200 mm), mandible (190 mm), five or six incomplete cervical vertebrae, cervical ribs, four incomplete anterior dorsal vertebrae, two partial posterior dorsal vertebrae, partial dorsal ribs, gastralia, synsacrum, incomplete first-sixth caudal vertebrae, proximal chevrons, incomplete scapulae, incomplete coracoids, partial furcula, partial sternal plates, incomplete humeri (119 mm), incomplete radii, incomplete ulnae, metacarpal I (19.2 mm), phalanx I-1 (39.9 mm), manual ungual I (30.2 mm straight), metacarpal II (50.8 mm), phalanx II-1 (31.3 mm), phalanx II-2 (45.8 mm), manual ungual II (33 mm), metacarpal III (45 mm), phalanx III-1 (17.6 mm), phalanx III-2 (10.1 mm), phalanx III-3 (32.2 mm), manual ungual III (25 mm), incomplete ilia (131.7, 124.6 mm), incomplete pubes (132 mm), ischial fragment, incomplete femora (184 mm), incomplete tibiae (182 mm), incomplete fibulae, astragalocalcaneum, distal tarsal IV, metatarsal I (22.5 mm), phalanges I-1 (19.8, 19.3 mm), pedal ungual I (16 mm straight), metatarsals II (73.7 mm), phalanges II-1 (23.9 mm), phalanges II-2 (26.1 mm), pedal unguals II (66.3 mm curve), metatarsals III (88.1 mm), phalanges III-1 (39.6 mm), phalanx III-2 (24.7 mm), phalanx III-1 fragment, metatarsals IV (83.4 mm), phalanx IV-1 (27.1 mm), phalanx IV-2 (21 mm), proximal phalanx IV-3, metatarsal V
Diagnosis- (after Norell and Makovicky, 1999) vertical ridge dorsal to acetabulum on ilium.
(after Powers et al., 2020) elongate antorbital fossa.
(after Powers et al., 2021a) anteriorly abbreviated cerebellar fossa with elongate olfactory canal; lacrimal notch of the frontal that is more vertically oriented as opposed to transversely in V. mongoliensis.
(after Powers et al., 2022) maxillary fenestra within a broad, shallow maxillary fossa that extends anterior to the maxillary fenestra.
Comments- IGM 100/982 was discovered in 1995 and first mentioned by Norell and Makovicky (1997) as an undescribed dromaeosaur specimen collected by the MAS-AMNH expeditions possessing a wide flat brevis shelf on its ilium. The postcranium was described by Norell and Makovicky in 1999, while the braincase was described by Norell et al. (2004). The skull was first figured in figure 6C of Norell et al. (2006). Norell and Makovicky referred it "to Velociraptor mongoliensis based on comparisons of cranial material with the holotype (AMNH 6515)", although even this early on they did note "A poorly defined vertical crest above the acetabulum that divides the anterior and posterior muscle fossae on the lateral surface of the pelvis is present on IGM 100/982 but is absent on other specimens." In an abstract, Kundrat (2004) noted several braincase characters which differed between it and IGM 100/976, concluding "IGM 100/982 represents a more advanced velociraptorine morphotype than does IGM 100/976" although he wrongly thought it was 100/982 that was confirmed to be V. mongoliensis and 100/976 that "might be referable to a new velociraptorine taxon." Powers (2020) names and describes the new species in his thesis, which also has sections which have been published since. Powers et al. (2020) discovered that using maxillary morphometrics "MPC-D 100/982 plotted far away from all other specimens referred to the genus Velociraptor, the separation being greater than that seen among many specimens representing different species" and states "A description of this specimen and review of variation within the genus Velociraptor is currently underway." Similarly, Powers et al. (2022) provides CT data on the maxilla and mentions it as "a possible third [species of Velociraptor] based on a currently undescribed specimen (MPC-D 100/982, description in prep.)", although as noted above many aspects of the specimen have been described in the literature. Two abstracts devoted to the new species were published in 2021 (Powers et al., 2021a, b).
References- Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton: Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 45 pp.
Kundrat, 2004. Two morphotypes of the Velociraptor neurocranium. ICVM-7 Abstracts. Journal of Morphology. 260(3), 305.
Norell, Makovicky and Clark, 2004. The braincase of Velociraptor. In Currie, Koppelhus, Shugar and Wright (eds.). Feathered Dragons: Studies on the Transition from Dinosaurs to Birds. Indiana University Press. 133-143.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS ONE. 4(10), e7390.
Powers, 2020. The evolution of snout shape in eudromaeosaurians and its ecological significance. Masters thesis, University of Alberta. 437 pp.
Powers, Sullivan and Currie, 2020. Re-examining ratio based premaxillary and maxillary characters in Eudromaeosauria (Dinosauria : Theropoda): Divergent trends in snout morphology between Asian and North American taxa. Palaeogeography, Palaeoclimatology, Palaeoecology. 547, 109704.
Powers, Norell and Currie, 2021a. New shallow snouted species of Velociraptor sheds light on intraspecific variation in Velociraptor mongoliensis and possible niche partitioning between species. 9th Annual Meeting Canadian Society of Vertebrate Palaeontology Online Abstracts. 31.
Powers, Norell and Currie, 2021b. Examination of morphological variation across Velociraptor mongoliensis specimens reveals a new species with possible ecomorphological variation in snout dimensions. The Society of Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual Meeting. 210-211.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as 2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters. Journal of Vertebrate Paleontology. 41(5), 2010087.
V. sp. indet. (Bohlin, 1953)
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Material- ?(AMNH 6518) partial maxilla, distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1 (16.2 mm), pedal ungual I, incomplete metatarsal II, phalanx II-1 (28 mm), phalanx II-2 (36 mm), pedal ungual II (56 mm), incomplete metatarsal III (~250 mm), phalanx III-?, metatarsal IV, phalanx IV-?, metatarsal V (40.1 mm) (Ostrom, 1969)
numerous teeth (9-14 mm) (Barsbold and Osmolska, 1999)
Late Campanian, Late Cretaceous
Tugrikin Shire, Djadokhta Formation, Mongolia
?(IGM 100/2000; 940725 TS-V OTGN or 940801 TS-I WTB) (juvenile) skull, mandible, postcranium (Barsbold and Osmolska, 1999)
?(IGM coll.; 940725 TS-V OTGN or 940801 TS-I WTB) skeleton (Watabe and Suzuki, 2000)
?(IGM coll.; 980725 TS-I Velocira NAR) dorsal vertebrae, ribs, caudal series, partial pelvis, hindlimb (Suzuki and Watabe, 2000)
Late Campanian, Late Cretaceous
Udyn Sayr, Djadokhta Formation, Mongolia
?(uncollected?) forelimbs (Suzuki and Watabe, 2000)
?(uncollected?) teeth (Watabe and Tsogtbaatar, 2004)
Late Campanian, Late Cretaceous
Ikh Shunkht, Baruungoyot Formation, Mongolia
?(IGM or PIN coll.) (Ivakhnenko and Kurzanov, 1988)
Late Campanian, Late Cretaceous
Khulsan, Baruungoyot Formation, Mongolia
?(ZPAL MgD-I/97) partial skull, partial mandibles, distal tibia, astragalus, metatarsal I, phalanx I-1, pedal ungual I, metatarsus (~80 mm), phalanx II-1, phalanx II-2, pedal ungual II, phalanx III-1, phalanx III-2, phalanx III-3, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV (Osmolska, 1982)
? several incomplete skeletons (Osmolska, 1982)
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
?(IVPP V16138) premaxillary tooth, incomplete lateral tooth (Hone, Choiniere, Sullivan, Xu, Pittman and Tan, 2010)
Campanian-Maastrichtian, Late Cretaceous
Minhe Formation, Inner Mongolia, China
? two teeth (Bohlin, 1953)
Late Cretaceous
Tzoyun, Shanxi, China
?(IVPP V965) tooth (Young, 1958)
Late Cretaceous?
Mongolia?
?(IGM 100/200) third premaxillary tooth (6.17x4.03x1.63 mm), partial maxilla, maxillary teeth (7.35-9.93 mm) (Smith, 2002)
?(University of Manchester LL.12392) manual ungual (Johnson et al., 2009)
Comments- These are specimens referred to Velociraptor which have not been described, figured or preserved well enough to refer to either V. mongoliensis or V. sp. nov.. Indeed, basically none of them are complete enough or have been described well enough to even refer to Velociraptor, and could easily be Tsaagan, Shri, Kuru or another eudromaeosaur.
Bohlin (1953) referred two teeth, and more questionably a penultimate phalanx and ungual, to Velociraptor mongoliensis (mispelled V. mongoliense). The teeth are indeed probably dromaeosaurid, and roughly similar to Velociraptor. While they lack mesial serrations like the geographically close V. osmolskae, V. mongoliensis teeth often lack mesial serrations as well (perhaps due to wear in some cases). They are thus indeterminate at least within Velociraptor, and probably within basal Dromaeosauridae. The phalanx is said to be broader than Velociraptor's III-3 and the illustrated ungual is clearly a parvicursorine manual ungual I however.
Young (1958) referred IVPP V965 to cf. Velociraptor mongoliensis, although stated it is "somewhat larger than the type". As it is based on a single tooth, referral to any particular dromaeosaurid species is questionable.
AMNH 6518 was discovered on August 11 1923, but not published until Ostrom (1969) who illustrated several phalanges. Images of the entire specimen are available at the AMNH online catalog.
ZPAL MgD-I/97 was found between 1963 and 1971 on the Polish-Mongolian expeditions, and is mentioned briefly by Osmolska (1982) in her description of Hulsanpes. The crania are figured by Barsbold and Osmolska (1999), who say the postcrania "will be published at a later date (Barsbold & Osmolska in preparation)."
IGM 100/2000 was listed as "the complete skeleton of a young individual. It includes skull with mandible and postcranium" by Barsbold and Osmolska (1999), who also wrote it was found "by the Mongolian-Japanaese Palaeontological Expeditions" at Tugrikin Shire. It is possibly the "Velociraptor skull and small skeleton" reported by Watabe and SuzukI (2000). Those authors list 940725 TS-V OTGN and 940801 TS-I WTB as "Velociraptor skeleton" from Tugrikin Shire V and "Velociraptor skeleton" from Tugrikin Shire I respectively, but which specimen became IGM 100/2000 is unknown and no description or illustration of either has been published.
Suzuki and Watabe (2000) reported two new Velociraptor specimens collected in 1998. "The postcranial skeleton of Velociraptor" with field number 980725 TS-I Velocira NAR discovered on July 25 at Tugrikin Shire I, described as consisting of "dorsal vertebrae, ribs, pelvic part, hind limb, and complete tail"; and "Velociraptor forelimbs" discovered between August 16-23 at Udyn Sayr.
Smith (2002) uses IGM 100/200 as one of his Velociraptor dental specimens used to measure tooth shape in his gigantic database of theropods, which includes a premaxillary tooth and a maxillary fragment that includes at least six tooth positions.
Watabe and Tsogtbaatar (2004) report "teeth of Velociraptor" discovered on July 23 2000 at Udyn Sayr.
Hone et al. (2010) describe two teeth found in 2008-2009 from the Wulansuhai Formation of Inner Mongolia as Velociraptor, associated with a cf. Protoceratops skeleton exhibiting tooth-marked bones. These teeth have mesial serrations, so are unlikely to be Tsaagan or "Velociraptor" osmolskae which are both known from this formation. Yet no characters are described that would refer these to Velociraptor over another dromaeosaurid with mesial serrations and a high DSDI, so they are only tentatively referred here.
References- Bohlin, 1953. Fossil reptiles from Mongolia and Kansu. Sino-Swedish Expedition Publication. 37, 1-105.
Young, 1958. The first record of dinosaurian remains from Shansi. Vertebrata PalAsiatica. 2(4), 231-236.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History. 30, 165 pp.
Osmolska, 1982. Hulsanpes perlei n. g. n. sp. (Deinonychosauria, Saurischia, Dinosauria) from the Upper Cretaceous Barun Goyot Formation of Mongolia. Neues Jahrbuch fur Geologie und Palaeontologie, Monatshefte. 1982(7), 440-448.
Ivakhnenko and Kurzanov, 1988. Geological structure and age of Udan-Sair and Shara-Tsav localities. Trudy Sovmestnoi Sovetsko-Mongolʼkoi Paleontologicheskoi Ekspeditsii. 34, 100105.
Barsbold and Osmólska, 1999. The skull of Velociraptor (Theropoda) from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica. 44(2), 189-219.
Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1998. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 83-98.
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum of Natural Sciences Research Bulletin. 1, 30-44.
Smith, 2002. An examination of dental morphology and variation in theropod dinosaurs: Implications for the identification of shed teeth. PhD dissertation. University of Pennsylvania.
Watabe and Tsogtbaatar, 2004. Report on the Japan - Mongolia Joint Paleontological Expedition to the Gobi desert, 2000. Hayashibara Museum of Natural Sciences Research Bulletin. 2, 45-67.
Johnson, Mustansar, Manning, Margetts and Mummery, 2009. Virtual repair of fossil CT scan data. Journal of Vertebrate Paleontology. 29(3), 123A.
Hone, Choiniere, Sullivan, Xu, Pittman and Tan, 2010. New evidence for a trophic relationship between the dinosaurs Velociraptor and Protoceratops. Palaeogeography, Palaeoclimatology, Palaeoecology. 291, 488-492.

Luanchuanraptor Lu, Xu, Zhang, Ji, Jia, Hu, Zhang and Wu, 2007
L. henanensis Lu, Xu, Zhang, Ji, Jia, Hu, Zhang and Wu, 2007
Late Cretaceous
Qiupa Formation, Henan, China
Holotype- (41HIII-0100) incomplete frontal, premaxillary tooth, three lateral teeth, posterior cervical vertebra, anterior cervical rib, posterior cervical rib, first dorsal vertebra, anterior dorsal rib, two proximal caudal vertebrae, two middle caudal vertebrae, thirteen distal caudal vertebrae, three proximal chevrons, middle chevron, two distal chevrons, scapulocoracoid, humerus, manual phalanx I-1, incomplete manual ungual, ilium, pubis, femoral shaft, fragments
Diagnosis- (after Lu et al., 2007) distal caudal prezygapophyses equal to centra in length; distal caudal vertebrae with neural spine or dorsal groove; elongated proximodorsal processes on distal chevrons; elongate median posterior process on distal chevrons (also in some Deinonychus chevrons); deep concavity present on medial coracoid; deltopectoral crest extends over half of humerus.
Comments- High resolution photos are available in Xu's (2007) thesis, as compared to the low quality of Lu et al.'s (2007) officially published pdf. This taxon has yet to be included in a published cladistic analysis, but when run in a version of the TWiG matrix, it emerges as the sister group to Velociraptor. The plesiomorphic characters may suggest a more basal placement, however.
References- Lu, Xu, Zhang, Ji, Jia, Hu, Zhang and Wu, 2007. New dromaeosaurid dinosaur from the Late Cretaceous Qiupa Formation of Luanchuan area, western Henan, China. Geological Bulletin of China. 26(7), 777-786.
Xu, 2007. The scientific characters and protection of dinosaurs from Henan Province. China University of Geosciences. 120 pp.

Daurlong Wang, Cau, Guo, Ma, Qing and Liu, 2022
D. wangi Wang, Cau, Guo, Ma, Qing and Liu, 2022
Early Aptian, Early Cretaceous
Pigeon Hill, Longjiang Formation, Inner Mongolia, China
Holotype- (IMMNH-PV00731) (1.5 m) skull (170 mm), sclerotic ring, mandible, ten cervical vertebrae, twelve dorsal vertebrae, dorsal ribs, gastralia, synsacrum, caudal series (800 mm), chevrons, scapulocoracoids (one partial; scapula 112 mm), sternum (75 mm), four sternal ribs, incomplete humerus (120 mm), radius (90 mm), ulna (96 mm), scapholunare, semilunate carpal, metacarpal I (20 mm), phalanges I-1 (one incomplete; 36 mm), manual unguals I (30 mm), metacarpal II (50 mm), phalanges II-1 (one incomplete; 30 mm), phalanges II-2 (43 mm), manual unguals II (37 mm), metacarpal III (48 mm), phalanges III-1 (one incomplete; 17 mm), phalanges III-2 (10 mm), phalanges III-3 (20 mm), manual unguals III (17 mm), partial ilium (~115 mm), incomplete pubis (~147 mm), ischium (~92 mm), femora (one incomplete; 180 mm), tibiae (198 mm), incomplete fibula, incomplete metatarsal III (110 mm), phalanx III-1 (35 mm), phalanges III-2 (30 mm), phalanges III-3 (23 mm), pedal unguals III (30 mm), partial metatarsal IV (95 mm), phalanx IV-1 (30 mm), phalanx IV-2 (20 mm), phalanx IV-3 (20 mm), phalanges IV-4 (13 mm), pedal unguals IV (25 mm), pedal claw sheaths, body feathers, intestine
Diagnosis- (after Wang et al., 2022) large, trapezoid promaxillary fenestra placed at anteroventral corner of antorbital fossa; maxillary fenestra large, shallow and posterodorsally located, so that the pila promaxillaris is wider than the pila interfenestralis; fan-shaped distal end of first sternal rib.
Comments- Discovered between 2018 and 2019 (Cau, pers. comm. 4-23), Wang et al. (2023) included this in a version of Cau's megamatrix and recovered it sister to "Zhenyuanlong", then Tianyuraptor in a clade sister to Eudromaeosauria. Added to Hartman et al.'s maniraptoromorph matrix it resolves as a velociraptorine closest to Kuru and Adasaurus.
Reference- Wang, Cau, Guo, Ma, Qing and Liu, 2022. Intestinal preservation in a birdlike dinosaur supports conservatism in digestive canal evolution among theropods. Scientific Reports. 12:19965.

Kuru Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021
= "Airakoraptor" Perle, Norell and Clark, 1999
K. kulla Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021
Late Campanian, Late Cretaceous
Khulsan, Baruungoyot Formation, Mongolia
Holotype- (IGM 100/981) premaxilla, partial lacrimal, incomplete dentary, incomplete surangular, three lateral teeth, incomplete ~third/fourth cervical vertebra, partial ~fourth/fifth cervical vertebra, fragmentary ~fifth/sixth cervical vertebra, tenth cervical vertebra, incomplete first dorsal vertebra, second dorsal vertebra, third dorsal vertebra, incomplete fourth dorsal vertebra, partial fifth dorsal vertebra, five incomplete posterior dorsal vertebrae, ~sixth caudal vertebra, two distal caudal vertebrae (one lost), proximal humerus, distal humeri, distal radius, proximal ulna, semilunate carpal, metacarpal I, phalanx ?I-1, incomplete manual ungual I, proximal metacarpal II, phalanx II-2, manual ungual II, proximal metacarpal III, phalanx ?III-1, phalanx ?III-2, posterior ilium, partial pubes, femora (one incomplete; ~220 mm), proximal tibia, partial astragalus, metatarsal I, pedal ungual I, partial metatarsals II, phalanx II-1, phalanges II-2, partial pedal unguals II, partial metatarsal III, phalanges III-1 (one incomplete), phalanx III-2, partial pedal ungual III, incomplete metatarsal IV (~113 mm), phalanx IV-1, five pedal phalanges
Diagnosis- (after Napoli et al., 2021) deeply incised groove framing anterior and ventral margins of premaxillary narial fossa; lacrimal with posterolateral hornlet arising from posterodorsal process; lacrimal with poorly developed boss at intersection of ventral, posterodorsal, and anterior processes; dentary lacking ventral secondary row of nutrient foramina; dentary nutrient foramina set in shallow groove posteriorly; two posterior surangular foramina; proportionally deep surangular; dentary teeth serrated on both mesial and distal carinae; anteriorly displaced dorsal pleurocoels; dorsal hyposphenes widely separated and joined by web of bone; distal tarsals unfused to metatarsus; mediolaterally narrow metatarsal II; reduced pedal digit II.
Comments- While the description merely states that in the manus "most phalanges are not confidently assignable", the reconstruction in Ruebenstahl et al.'s (2021) SVP presentation shows phalanges I-1, III-3 and III-2 outlined.
"Airakoraptor" was originally listed in a bibliographic entry in Perle et al.'s (1999) description of Achillobator. This supposed paper is listed as authored by Norell, Clark and Perle in a 1992 issue of the Journal of Vertebrate Paleontology. No pages numbers or issue are listed, though it does say "Vol. 11 Toronto. Canada", which indicates it refers to an abstract from the Toronto Society of Vertebrate paleontology meeting in 1992 (volume 11 covered the 1991 meeting in San Diego, so must be a mistake). There is an abstract in that issue by those authors, but it has a different title which does not mention "Airakoraptor" by name. Instead of "Morphology dromaeosaurian dinosaur- Airakoraptor from the Upper Cretaceous of Mongolia", it is simply "New dromaeosaur material from the Late Cretaceous of Mongolia". So Perle et al.'s listing must refer to an earlier version of the abstract. Napoli et al. (2021) confirm that of the specimens noted in the abstract, IGM 100/981 was "familiar (erroneously) under the provisional name "Airakoraptor," and that "the name is a nomen nudum."
IGM 100/981 was initially mentioned by Norell et al. (1992) as "a partial skeleton including portions of the skull, mandible, vertebral column, manus and pes" and said to "possess features unknown in described dromaeosaurs" such as a deep mandible and dual posterior surangular foramina. This specimen is also probably that described by Novacek (1996) as discovered by Perle on July 4th - "a fine skeleton of a dromaeosaur, perhaps not Velociraptor but something rare, and new", since Shri was found by Norell. It was later mentioned as an "unnamed dromaeosaur from Khulsan collected by the Mongolian American Expeditions" by Norell and Makovicky (1997), stated to have a wide and flat brevis shelf and share an anterior tubercle on metatarsal III with Velociraptor and Deinonychus. In Norell and Makovicky (1999) it is noted to lack a fourth trochanter, unlike Velociraptor. They state it "can be referred to the Dromaeosauridae on the basis of pedal and tail morphology, but insufficient elements are present to refer these specimens to Velociraptor mongoliensis." Norell et al. (2006) illustrate the surangular as "an undescribed dromaeosaur from Khulson (IGM 100/981)" and it clearly shows dual surangular foramina, matching figure 6A of Napoli et al. (2021) who finally described the specimen as Kuru kulla. Confusingly, Turner et al. (2007) described an ulna of a supposed Velociraptor specimen they called IGM 100/981. This specimen was said to be from the Gilvent Wash locality near Ukhaa Tolgod, and Napoli et al. state the specimen number was erroneous and is actually IGM 100/3503 which is here referred to Tsaagan.
References- Norell, Clark and Perle, 1992. New dromaeosaur material from the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 12(3), 45A.
"Norell, Clark and Perle, 1992. Morphology dromaeosaurian dinosaur- Airakoraptor from the Upper Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 11. Toronto, Canada." [not a real paper]
Novacek, 1996. Dinosaurs of the Flaming Cliffs. Anchor Books. 367 pp.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton: Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: Information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 45 pp.
Perle, Norell and Clark, 1999. A new maniraptoran Theropod - Achillobator giganticus (Dromaeosauridae) - from the Upper Cretaceous of Burkhant, Mongolia. Contribution no. 101 of the Mongolian-American Paleontological Project. 1-105.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
Turner, Makovicky and Norell, 2007. Feather quill knobs in the dinosaur Velociraptor. Science. 317, 1721.
Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021. A new dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central Mongolia. American Museum Novitates. 3982, 47 pp.
Ruebenstahl, Napoli, Bhullar, Turner and Norell, 2021. Two new eudromaeosaurs from Khulsan (central Mongolia) reveal modern-like faunal predatory structure amoung non-avian dinosaurs. The Society of Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual Meeting. 222-223.
Turner, Montanari and Norell, 2021. A new dromaeosaurid from the Late Cretaceous Khulsan locality of Mongolia. American Museum Novitates. 3965, 46 pp.

Adasaurus Barsbold, 1983
= "Adasaurus" Barsbold, 1977
A. mongoliensis Barsbold, 1983
= Dromaeosaurus mongoliensis (Barsbold, 1983) Paul, 1988
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
Holotype- (IGM 100/20) (~2.7 m) (adult) posterior skull, posterior mandibles, atlantal neurapophyses, atlantal intercentrum, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra (38 mm), tenth cervical vertebra (35 mm), first dorsal vertebra (31 mm), second dorsal vertebra (31 mm), third dorsal vertebra (28 mm), fourth dorsal vertebra (28 mm), fifth dorsal vertebra (27 mm), sixth dorsal vertebra (27 mm), seventh dorsal vertebra (27 mm), eighth dorsal vertebra (27 mm), ninth dorsal vertebra (27 mm), tenth dorsal vertebra (26 mm), eleventh dorsal vertebra (27 mm), synsacrum (217 mm; 29, 31, ?, ?, 31, 29 mm), proximal caudal vertebra (36 mm), sixth-eleventh caudal vertebrae (39, 42, 42, 43, 43, 43 mm), six chevrons, incomplete scapulacoracoid, fragmentary sternal plate, incomplete ilia, incomplete pubes, proximal ischium, femora (267, 263 mm), tibiae (one partial, one incomplete; 296 mm), fibulae, incomplete astragalocalcaneum, fused distal tarsals III and IV, incomplete metatarsus (III 146, IV 130 mm), phalanx III-1, phalanx IV-1 (40 mm), phalanx IV-2 (31 mm), phalanx IV-3 (26 mm), phalanx IV-4 (23 mm), pedal ungual IV, metatarsal V
Paratype- (IGM 100/21) (~2.7 m) two mid caudal vertebrae, metatarsal I (34 mm), phalanx I-1 (27 mm), pedal ungual I, tarsometatarsus (II 122, III 145, IV 130 mm), phalanx II-1 (34 mm), phalanx II-2 (32 mm), phalanx III-1 (55 mm), phalanx III-2 (36 mm), phalanx III-3, pedal ungual III, phalanx IV-1 (39 mm), phalanx IV-2 (30 mm), phalanx IV-3 (25 mm), phalanx IV-4 (32 mm), pedal ungual IV
Diagnosis- (after Kubota and Barsbold, 2006) dorsoventrally expanded anterior process of jugal; low dorsal ridge on median frontals (continuous with sagittal crest on parietals); untwisted paroccipital process; large surangular foramen; pleurocoels only on anterior sacrals; preacetabular process strongly notched anteriorly.
(after Turner et al., 2012) ventral process of lacrimal strongly curved anteriorly; dorsally displaced triangular process along lateral edge of quadrate shaft.
(proposed) metatarsal II reduced in width.
Other diagnoses- Contra Barsbold (1983) and most later references, the supposed small pedal ungual II does not belong to digit II (Senter, 2010).
Comments- This genus was discovered between 1964 and 1966, and first mentioned by Barsbold (1977) as a "latest (Maastrichtian) Mongolian Dromaeosauridae" with a reduced pedal ungual II, with the pelvis illustrated and labeled as Adasaurus. This does not count as a proper description however, due to the lack of a type species or holotype. Osmolska (1980) lists one dromaeosaurid taxon and individual from the Nemegt Formation (as ?Velociraptor sp.), which may be based on Adasaurus and may be the source of Glut's (1982) reference to "a possible dromaeosaurid, mentioned by Osmolska (1978), from the Nemegt Formation of Mongolia." Note no Osmolska reference from 1978 exists and Adasaurus was already listed by Glut on page 40 however. The taxon was later described extremely briefly by Barsbold (1983) and officially named Adasaurus mongoliensis, though the paratype was mistyped as IGM 100/51. While remaining poorly described and virtually unillustrated in the literature for decades due to an embargo, Adasaurus has finally been partially illustrated and described by Turner et al. (2012) with a full description in Kubota's (2015) thesis. Although Currie and Varricchio (2004) referred snout and forelimb remains to Adasaurus based on IGM 100/22 and 100/23, Kubota and Barsbold (2007) have determined these belong to a new taxon from the earlier Bayanshiree Formation.
The holotype specimen was approximately 2.7 meters long if the femur is scaled from Velociraptor. It was from an old individual, so this is probably close to the maximum size Adasaurus got. Contra Norell and Makovicky (1997), the specimen is not pathological (Turner et al., 2012).
Jasinowski et al. (2006) list a scapulocoracoid and sternal plate for the holotype. The pelvis was illustrated by Barsbold (1983) after the initial schematic drawing in Barsbold (1977), but this is quite different from the actual material as it is a composite with the holotype's ilium and pubis and IGM 100/22's ischium.. The femoral morphology was briefly noted by Perle et al. (1999) and Kim et al. (2005). The latter also illustrate the proximal femur in two views. While Adasaurus is perhaps most famous due to its supposedly reduced second pedal ungual, Senter (2010 as a 2006 pers. comm. from Kubota) notes that ungual does not actually pertain to the holotype's digit II.
Relationships- Adasaurus was originally identified as a dromaeosaurid (Barsbold, 1977) and later specified to be a dromaeosaurine (Barsbold, 1983; Paul, 1988). More recently, Longrich and Currie (2009) found it to be a velociraptorine, which was also found by Kubota and Barsbold (2006, 2007), Senter et al. (2012), Turner et al. (2012), Brusatte et al. (2014) and Foth et al. (2014). There is certainly no reason to synonymize Adasaurus with Dromaeosaurus as Paul (1988) did, as many other dromaeosaurid genera seem to be more closely related to Dromaeosaurus.
References- Barsbold, 1977. [On the evolution of the carnivorous dinosaurs]. Transactions of the Joint Soviet Mongolian Paleontological Expedition. 4, 48-56.
Osmolska, 1980. The Late Cretaceous vertebrate assemblages of the Gobi Desert, Mongolia. Memoires de la Societe Geologique de France. 139, 145-150.
Glut, 1982. The New Dinosaur Dictionary. Citadel Press. 288 pp.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton: Information from a new specimen. American Museum Novitates. 3215, 1-28.
Perle, Norell and Clark, 1999. A new maniraptoran Theropod - Achillobator giganticus (Dromaeosauridae) - from the Upper Cretaceous of Burkhant, Mongolia. Contribution no. 101 of the Mongolian-American Paleontological Project. 1-105.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons. Studies on the transition from dinosaurs to birds. Indiana University Press. 112-132.
Kim, Gishlick and Tsuihiji, 2005. The first non-avian maniraptoran skeletal remains from the Lower Cretaceous of Korea. Cretaceous Research. 26, 299-306.
Jasinowski, Russell and Currie, 2006. An integrative phylogenetic and extrapolatory approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder musculature. Zoological Journal of the Linnean Society. 146, 301-344.
Kubota and Barsbold, 2006. Reexamination of Adasaurus mongoliensis (Dinosauria: Theropoda) from the Upper Cretaceous Nemegt Formation of Mongolia. Journal of Vertebrate Paleontology. 26(3), 88A.
Kubota and Barsbold, 2007. New dromaeosaurid (Dinosauria Theropoda) from the Upper Cretaceous Bayanshiree Formation of Mongolia. Journal of Vertebrate Paleontology. 27(3), 102A.
Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis. Columbia University. 666 pp.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Senter, 2010. Using creation science to demonstrate evolution: Application of a creationist method for visualizing gaps in the fossil record to a phylogenetic study of coelurosaurian dinosaurs. Journal of Evolutionary Biology. 23(8), 1732-1743.
Nesbitt, Clarke, Turner and Norell, 2011. A small alvarezsaurid from the Eastern Gobi Desert offers insight into evolutionary patterns in the Alvarezsauroidea. Journal of Vertebrate Paleontology. 31(1), 144-153.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Brusatte, Vremir, Csiki-Sava, Turner, Watanabe, Erickson and Norell, 2013. The osteology of Balaur bondoc, an island-dwelling dromaeosaurid (Dinosauria: Theropoda) from the Late Cretaceous of Romania. Bulletin of the American Museum of Natural History. 374, 1-100.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Kubota, 2015. Descriptions of Mongolian dromaeosaurids (Dinosauria: Theropoda) and phylogeny of Dromaeosauridae. PhD thesis, University of Tsubaka. 397 pp.

"Koreanosaurus" Kim, 1979
"K. koreanensis" (Kim, 1993) new comb.
= Deinonychus "koreanensis" Kim, 1993
Aptian-Early Albian, Early Cretaceous
Lower Gugyedong Formation, South Korea
Material- (DGBU-78) femur (~400 mm)
Comments- Originally referred to the Deinodontidae (Kim, 1979), and after that Hypsilophodontidae or Hadrosauridae (Kim, 1983). By 1986, Kim had decided it was perhaps synonymous with Deinonychus (pers. comm. to Olshevsky). In 1993, the name Deinonychus "koreanensis" was used in a photo caption and faunal list. Lee et al. (2001) remove the specimen from Deinonychus, based on the presence of a fourth trochanter. However, Deinonychus varies in this character. Kim et al. (2005) refer the specimen to Eumaniraptora based on a proximolateral ridge, shelf-like posterior trochanter, and absence of an accessory trochanter and mediodistal crest. The presence of a large fourth trochanter was noted to be similar to Adasaurus and Velociraptor.
References- Kim, 1979. [Dinosaur and volcano discovered from Tabri, Euiseong, Korea].
Kim, 1983. Cretaceous dinosaurs from Korea. Journal of the Geological Society of Korea. 19(3), 115-126.
Kim, 1993. Journal of Natural History and Environments Vol 1 #1, June 1993. Published by the World Society of Natural History and Environments, Pusan University, Pusan, Korea. ISSN 1225-6404.
Lee, Yu and Wood. 2001. A review of vertebrate faunas from the Gyeongsang Supergroup (Cretaceous) in South Korea. Palaeogeography, Palaeoclimatology, Palaeoecology. 165, 357-373.
Kim, Gishlick and Tsuihiji, 2005. The first non-avian maniraptoran skeletal remains from the Lower Cretaceous of Korea. Cretaceous Research. 26, 299-306.

unnamed clade (Deinonychus antirrhopus + Utahraptor ostrommaysi)
= Megaraptora Miller, 2004 (non Benson, Carrano and Brusatte, 2009)
Comments- This clade is based on Hartman et al.'s maniraptoromorph analysis, notable for excluding Dromaeosaurus unlike e.g. Senter's coelurosaur analyses. Miller (2004) had previously suggested a pairing of Deinonychus and Utahraptor, which he called Megaraptora despite it not including Megaraptor which was named six years earlier and which he had to be aware of. Benson et al.'s Megaraptora defined to include Megaraptor wasn't proposed until five years later, but is used here based on being universally accepted and including the eponymous genus.
References- Miller, 2004. A new phylogeny of the Dromaeosauridae. 2004 Student Showcase Journal. 20, 123-158.
Benson, Carrano and Brusatte, 2010 (2009 online). A new clade of archaic large-bodied predatory dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften. 97(1), 71-78.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 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

Pneumatoraptor Osi, Apesteguia and Kowalewski, 2010
P. fodori Osi, Apesteguia and Kowalewski, 2010
Santonian, Late Cretaceous
Csehbanya Formation, Hungary
Holotype- (MTM V.2008.38.1) (adult) scapulocoracoid
Diagnosis- (after Osi et al., 2010) large, circular pneumatic foramen ventral to the coracoid foramen (2 mm in diameter) that opens towards the coracoid tubercle and is also in connection with the hollow scapular blade.
Comments- While only assigned to Paraves by Osi et al. (2010), unpublished analyses using TWiG characters suggest Pneumatoraptor may be a dromaeosaurid.
Reference- Osi, Apesteguia and Kowalewski, 2010. Non-avian theropod dinosaurs from the early Late Cretaceous of Central Europe. Cretaceous Research. 31(3), 304-320.

Yixianosaurus Xu and Wang, 2003
Y. longimanus Xu and Wang, 2003
Late Valanginian-Early Aptian, Early Cretaceous
Lujiatun or Jianshangou Beds of the Yixian Formation, Liaoning, China
Holotype- (IVPP V12638) (subadult or adult) (~1.3-2 kg) four partial dorsal ribs, seven gastralia or sternal ribs, furcula, scapulae (65 mm), coracoids, humeri (89 mm), radii (63 mm), ulnae (62, 64 mm), scapholunare, semilunate carpal, distal carpal III, metacarpal I (14.5 mm), phalanx I-1 (33, 34 mm), manual ungual I (26, 25 mm), metacarpal II (36, 35 mm), phalanx II-1 (26, 25 mm), phalanx II-2 (38, 36 mm), manual ungual II (28 mm), metacarpal III (34 mm), phalanx III-1 (~9 mm), phalanx III-2 (8 mm), phalanx III-3 (22 mm), manual ungual III (22.5, 23 mm), manual claw sheaths, fragments, feathers
Comments- This specimen was discovered in 2001 and described in 2003 as a maniraptoran perhaps most closely related to scansoriopterygids. This was based on the elongate manus (compared to humerus) and phalanx II-2 (compared to metacarpal II). These are correlated with each other however, as an elongate phalanx II-2 will lead to a long manus. In addition to scansoriopterygids, tyrannosauroids, ornithomimosaurs, troodontids, and some dromaeosaurids, basal avialans and basal coelurosaurs have similarly elongate II-2 (compared to II-1). Dececchi et al. (2010) restudied the specimen for an SVP abstract and while they were secretive regarding its phylogenetic position, do indicate it is not a paravian. Dececchi et al. (2012) later published a full paper, noting the presence of a furcula, and finding it to be a maniraptoran in a polytomy with alvarezsaurids, therizinosaurs and pennaraptorans. Xu et al. (2013) replied, describing misinterpretations of anatomy by Dececchi et al., including the supposed intermedium being a broken part of the semilunate carpal. They noted Dececchi et al. coded Yixianosaurus for many characters that could not be determined from the specimen, and their recoded matrix found it to be a deinonychosaur outside of Unenlagiinae, Eudromaeosauria and Troodontidae. Foth et al. (2014) recovered the genus as a basal paravian.
References- Xu and Wang, 2003. A new maniraptoran dinosaur from the Early Cretaceous Yixian Formation of Western Liaoning. Vertebrata PalAsiatica. 41(3), 195-202.
Dececchi, Hone, Sullivan and Xu, 2010. A re-analysis of the "coeluriasaurian pit-bull" Yixianosaurus longimanus with implications for the theropod dinosaur diversity of the Jehol Biota. Journal of Vertebrate Paleontology. Program and Abstracts 2010, 81A.
Dececchi, Larsson and Hone, 2012. Yixianosaurus longimanus (Theropoda: Dinosauria) and its bearing on the evolution of Maniraptora and ecology of the Jehol fauna. Vertebrata PalAsiatica. 59(2), 111-139.
Xu, Sullivan and Wang, 2013. The systematic position of the enigmatic theropod dinosaur Yixianosaurus longimanus. Vertebrata PalAsiatica. 51(3), 169-183.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature. 511, 79-82.

Deinonychus Ostrom, 1969a
= "Daptosaurus" Brown vide Chure and McIntosh, 1989
D. antirrhopus Ostrom, 1969a
= Velociraptor antirrhopus (Ostrom, 1969a) Paul, 1988
= "Daptosaurus agilis" Brown vide Chure and McIntosh, 1989
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Montana, US
Holotype- (YPM 5205) distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (146.5 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, pedal ungual IV
?... (YPM 5203) seventh caudal vertebra (45.5 mm), eighth caudal vertebra (47.6 mm), ninth caudal vertebra (50.2 mm), eleventh caudal vertebra (53 mm), twelfth caudal vertebra (53.1 mm), thirteenth caudal vertebra (53.2 mm), fourteenth caudal vertebra (53 mm), fifteenth caudal vertebra (~53.5 mm), sixteenth caudal vertebra (52.6 mm), seventeenth caudal vertebra (51.7 mm), eighteenth caudal vertebra (50 mm), nineteenth caudal vertebra (50.2 mm), twentieth caudal vertebra (48.3 mm), twenty-first caudal vertebra (47.5 mm), twenty-second caudal vertebra (46.9 mm), twenty-third caudal vertebra (45.2 mm), twenty-fourth caudal vertebra (42.8 mm), twenty-fifth caudal vertebra (42 mm), twenty-sixth caudal vertebra (41.5 mm), twenty-eighth caudal vertebra (37.8 mm), twenty-ninth caudal vertebra (37.5 mm), thirtieth caudal vertebra (33.6 mm), thirty-first caudal vertebra (32.1 mm), thirty-second caudal vertebra (32.3 mm), thirty-third caudal vertebra (28.7 mm), thirty-fourth caudal vertebra (24.8 mm), thirty-fifth caudal vertebra (23.3 mm), thirty-sixth caudal vertebra (20.1 mm), chevrons
?... (YPM 5206) radii (one partial) (176.5 mm), ulna, metacarpal I (45.8, 43.2 mm), phalanx I-1 (74.1 mm), manual ungual I, metacarpal II (93.7, ~88.3 mm), phalanx II-1 (54 mm), phalanx II-2 (76.5 mm), manual ungual II, metacarpal III (82 mm), phalanx III-1 (29.9, 30.6 mm), phalanx III-2 (20.5 mm), manual ungual III
?... (YPM 5210) (partial skull 364 mm) jugal, postorbital, quadratojugals, squamosals, ectopterygoid, pterygoids, vomer, dentary, splenial, surangular, angular, prearticulars, articular, 39 teeth, atlas intercentrum, partial atlantic neural arch, axis, seventh cervical vertebra (41.8 mm), tenth cervical vertebra (29.5 mm), cervical ribs, ninth dorsal vertebra (30 mm), fourth caudal vertebra (39.6 mm)
Paratypes- (YPM 5201) thirteenth caudal vertebra (48.3 mm), fifteenth caudal vertebra (46.6 mm), sixteenth caudal vertebra (46.4 mm), seventeenth caudal vertebra (47.5 mm), eighteenth caudal vertebra (46 mm), nineteenth caudal vertebra (45 mm), twentieth caudal vertebra (41.6 mm), twenty-first caudal vertebra (42.6 mm), twenty-second caudal vertebra (43.2 mm), twenty-third caudal vertebra (~41.6 mm), twenty-fourth caudal vertebra (~40 mm), twenty-fifth caudal vertebra (39.4 mm), twenty-sixth caudal vertebra (~35 mm), twenty-eighth caudal vertebra (35 mm), twenty-ninth caudal vertebra (34.3 mm), chevrons
(YPM 5202) ~twenty caudal vertebrae, chevrons
(YPM 5204) partial atlas neural arch and odontoid, axis (31.5 mm), fourth cervical vertebra (33 mm), fifth cervical vertebra (32 mm), third dorsal vertebra (28.4 mm), fifth dorsal vertebra (28.6 mm), sixth dorsal vertebra (28.5 mm), eighth dorsal vertebra (28.6 mm)
(YPM 5207) distal tarsal III, distal tarsal IV, phalanx I-1 (33.1 mm), phalanx III-2 (39 mm), phalanx III-3 (37.4 mm)
(YPM 5208) scapholunare, semilunate carpal
(YPM 5209) manual phalanx II-2 (78 mm), manual phalanx III-3 (52 mm), manual ungual III
(YPM 5211) semilunate carpal
(YPM 5212) manual ungual I
(YPM 5213) manual phalanx I-1 (72 mm)
(YPM 5214) metacarpal III
(YPM 5215) manual phalanx III-3
(YPM 5216) manual phalanx II-1 (55.3 mm)
(YPM 5217) semilunate carpal, distal tarsal III, metatarsal I (39.1 mm), phalanx I-1, pedal ungual I, phalanx II-1 (43.3, 42.8 mm), metatarsal III (150.5 mm), phalanx III-1, phalanx III-3 (35.4 mm), pedal ungual III, phalanx IV-1 (46.5 mm), phalanx IV-2 (33.6 mm), phalanx IV-3, phalanx IV-4 (28 mm), pedal ungual IV, metatarsal V (69.1 mm)
(YPM 5218) pedal ungual II, phalanx IV-1 (46.6 mm), phalanx IV-2 (36.9 mm), phalanx IV-3 (27.8 mm), pedal ungual IV
(YPM 5219) pedal ungual I or II
(YPM 5220) radius (172 mm), ulnae (174.2, 180 mm), manual phalanx I-1 (67.2 mm), manual ungual I
(YPM 5221) manual ungual II
(YPM 5222) manual ungual II
(YPM 5223) distal tarsal IV
(YPM 5224) manual ungual I
(YPM 5225) calcaneum
(YPM 5226) astragalus, calcaneum
(YPM 5227) pedal ungual IV
(YPM 5228) scapholunare
(YPM 5229) distal tarsal
(YPM 5230) ulna
(YPM 5231) twelve teeth
(YPM 5232) (3.06 m, 45 kg) partial skull (332 mm) including premaxillae, maxilla, nasals, lacrimal, jugals, postorbitals, squamosal, quadratojugal, palatine, vomer, dentaries, angular, articulars, 33 teeth
(YPM 5233) ectopterygoid, pterygoid
(YPM 5234) surangular
(YPM 5235) ischium (161 mm)
(YPM 5236) coracoid
(YPM 5237) splenial
(YPM 5238) splenial
(YPM 5239) pterygoid
(YPM 5240) proximal metatarsal I (45.5 mm), metatarsal I (40 mm)
(YPM 5241) dorsal ribs
(YPM 5242) scapholunare
(YPM 5243) manual phalanx III-3 (48.7 mm), manual ungual III
(YPM 5244) chevron
(YPM 5245) anterior dorsal rib
(YPM 5246) sternal ribs
(YPM 5247) gastralia
(YPM 5248) gastralia
(YPM 5249) mid dorsal rib, posterior dorsal rib
(YPM 5250) dorsal ribs, sternal ribs
(YPM 5251) gastralia
(YPM 5252) gastralia?
(YPM 5253) fragmentary caudal vertebra
(YPM 5254) cervical ribs?
(YPM 5255) rib
(YPM 5256) ribs
(YPM 5257) anterior dorsal rib, posterior dorsal rib
(YPM 5258) dorsal rib
(YPM 5259) two anterior dorsal ribs
(YPM 5260) dorsal rib
(YPM 5261) epipterygoid, gastralia
(YPM 5262) rib
(YPM 5263) metatarsal V
(YPM 5264) ribs, gastralial fragments
(YPM 5265) ribs, gastralial fragments
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Montana, US
(AMNH 3015; intended holotype of "Daptosaurus agilis") (3.06 m, 45 kg, >6 year old adult) incomplete atlas intercentrum, axial fragment, fourth cervical vertebra, fifth cervical vertebra (36.7 mm), sixth cervical vertebra (38.5 mm), seventh cervical vertebra (~39 mm), eighth cervical vertebra (~29 mm), ninth cervical vertebra (~31 mm), tenth cervical vertebra (~30 mm), (dorsal series ~374.5 mm) first dorsal vertebra (~31 mm), second dorsal vertebra (~30 mm), third dorsal vertebra (~30 mm), fourth dorsal vertebra (~30 mm), fifth dorsal vertebra (~27 mm), sixth dorsal vertebra (~28.5 mm), seventh dorsal vertebra (~27.5 mm), eighth dorsal vertebra (~31.5 mm), ninth dorsal vertebra (~30 mm), tenth dorsal vertebra (~28 mm), eleventh dorsal vertebra (~27 mm), twelfth dorsal vertebra, thirteenth dorsal vertebra, gastralia, second caudal vertebra (33.1 mm), third caudal vertebra (37.4 mm), fourth caudal vertebra (38.3 mm), fifth caudal vertebra (39 mm), sixth caudal vertebra (41.4 mm), seventh caudal vertebra (43.6 mm), eighth caudal vertebra (44.5 mm), ninth caudal vertebra (47.6 mm), tenth caudal vertebra (~49.4 mm), eleventh caudal vertebra (~50.6 mm), twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra, fifteenth caudal vertebra (~47.2 mm), sixteenth caudal vertebra (48 mm), seventeenth caudal vertebra (47.8 mm), eighteenth caudal vertebra (46.2 mm), nineteenth caudal vertebra (47.1 mm), twentieth caudal vertebra (46.6 mm), twenty-first caudal vertebra, twenty-second caudal vertebra, twenty-third caudal vertebra (43.5 mm), twenty-fourth caudal vertebra (43.2 mm), twenty-fifth caudal vertebra (43 mm), twenty-sixth caudal vertebra (41.8 mm), chevrons, scapulae (~190 mm), coracoid fragment, humeri (~237, 227 mm), radii (~172 mm), ulnae (186), scapholunares, semilunate carpal, metacarpal I (35.5 mm), manual ungual I (90 mm), phalanx II-1 (62.2 mm), phalanx II-2 (70.7 mm), metacarpal III (73.4 mm), phalanx III-1 (21.7 mm), phalanx III-2 (~15.5 mm), phalanx III-3 (47.3 mm), ilium (~245 mm), ischia (158, ~161 mm), femur (284 mm), tibia with astragalus (324 mm, 312 without), fibula (~297 mm), astragalus (62 mm wide, 77 mm high), calcaneum, distal tarsal III, distal tarsal IV, metatarsal II (129 mm), phalanx II-1 (37.7 mm), phalanx II-2 (42.2 mm), pedal ungual II (92 mm), metatarsal III (151 mm), phalanx III-1 (52.5 mm), phalanx III-2 (33 mm), phalanx III-3 (28 mm), pedal ungual III (62 mm), metatarsal IV (134 mm), phalanx IV-1 (44.7 mm), phalanx IV-2 (35.9 mm), phalanx IV-3 (32.2 mm), phalanx IV-4 (26.3 mm), pedal ungual IV, metatarsal V, eggshells
(AMNH 3037) teeth, manual elements, pedal elements
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Montana, US
Referred- (AMNH 3041 in part) twenty-five teeth (Ostrom, 1970)
(AMNH 30832) teeth (AMNH online)
(MCZ 8791) (one year old juvenile; ~7 kg) partial maxilla, articular, partial ?seventh cervical vertebra, five dorsal centra (~eleventh 17.4 mm), three proximal caudal centra, three mid caudal centra, incomplete coracoids, proximal radius, partial ulna, phalanx II-2 (56.2 mm), ilial fragment, distal femur, partial tibia, proximal fibula, proximal phalanx II-1, partial pedal ungual II, phalanx IV-4 (14.5 mm), gut contents (Parsons and Parsons, 2004; described by Parson and Parsons, 2015)
(MOR 682) eleven teeth (Maxwell and Ostrom, 1995)
(MOR 747) (at least four individuals) ~320 elements including maxilla, nasal, lacrimal, jugal, prefrontal, frontal, parietal, quadrate, laterosphenoid, mesethmoid, braincase, ectopterygoid, palatine, anterior dentary, teeth, other skull material, cervical vertebra, several caudal vertebrae, radius, ulna, manus, pes (Maxwell and Witmer, 1996)
(MOR 1178) (~2.32 m) maxillary tooth fragment, squamosal fragment, (?)ectopterygoid, dentary fragment, axis, sixth cervical vertebra, anterior dorsal vertebra, mid dorsal vertebra, sacral neural arch fragment, proximal caudal vertebra, two mid caudal vertebrae, distal caudal vertebra, scapula, coracoid, sternal(?) fragment, partial humerus, semilunate carpal, manual ungual I, ilium fragment, partial femora, proximal fibula, tarsal, pedal phalanx I-1, proximal metatarsal II, pedal ungual II, peal ungual III, phalanx III-1, metatarsal (?)IV, phalanx IV-4, pedal ungual IV, gut contents (Parsons and Parsons, 2002)
(MOR coll.) two teeth (Maxwell, 1993)
(YPM 5266) gastralia (Maxwell and Ostrom, 1995)
(YPM 5267) two ?anterior dorsal ribs (Maxwell and Ostrom, 1995)
(YPM 5268) dorsal rib fragments (Maxwell and Ostrom, 1995)
(YPM 5269) dorsal rib fragments (Maxwell and Ostrom, 1995)
(YPM 5270) metacarpal II (~93.5 mm) (Maxwell and Ostrom, 1995)
(YPM 5272) five teeth (Ostrom, 1970)
(YPM 5273) tooth (Ostrom, 1970)
(YPM 5278) tooth (Ostrom, 1970)
(YPM 5279) tooth (Ostrom, 1970)
(YPM 5280) tooth, partial tooth (Ostrom, 1970)
(YPM 5281) caudal vertebra, pedal ungual (Ostrom, 1970)
(YPM 5283) two partial teeth (Ostrom, 1970)
(YPM 5287) caudal vertebra (Ostrom, 1970)
(YPM 5288) four partial dorsal and caudal vertebrae, fragments (Ostrom, 1970)
?(YPM 5289) tooth (Ostrom, 1970)
?(YPM 5291) two tooth fragments (Ostrom, 1970)
(YPM 5397) tooth (Ostrom, 1970)
(YPM 5420) eleven teeth (Ostrom, 1970)
(YPM 5538 in part) proximal fibula, fibular fragment? (YPM online)
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Wyoming, US
(YPM 5271) partial tooth (Ostrom, 1970)
(YPM 5274) tooth (Ostrom, 1970)
?(YPM 5290) cervical centrum (Ostrom, 1970)
(YPM 5356) tooth fragment, fragment (Ostrom, 1970)
(YPM 5371) incomplete tooth (Ostrom, 1970)
(YPM 5376) tooth (Ostrom, 1970)
(YPM 5379) tooth (Ostrom, 1970)
(YPM 5441) two teeth (Ostrom, 1970)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Montana, US
(MCZ 4371; field number 74M #7) (3.43 m, 73 kg) snout fragments, mandibular fragments, teeth, nearly complete cervical series, nearly complete dorsal series, rib fragments, gastralia fragments, sacrum, complete caudal series, humerus (254 mm), radius (192 mm), ulna (208 mm), phalanx I-1 (77.3 mm), manual ungual I, metacarpal II, phalanx II-1 (64.2 mm), phalanx II-2 (83.4 mm), manual ungual II, metacarpal III (~90 mm), phalanx III-1 (35 mm), phalanx III-2 (23.2 mm), ilia (325 mm), pubes (380+ mm), ischium (175 mm), femora (336 mm), tibia with astragalus (382 mm, 368 without), metatarsal I (45.6 mm), phalanx I-1 (34.8 mm), pedal ungual I, metatarsal II (144.3 mm), phalanx II-1 (47 mm), phalanx II-2 (49.9 mm), pedal phalanx II, metatarsal III (164.4 mm), phalanx III-1 (64.4 mm), phalanx III-2 (44 mm), phalanx III-3 (41.3 mm), pedal ungual III, metatarsal IV (150.4 mm), phalanx IV-1 (55.5 mm), phalanx IV-2 (41.8 mm), phalanx IV-3 (35.2 mm), phalanx IV-4 (32.6 mm), pedal ungual IV, metatarsal V (78 mm) (Ostrom, 1976)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Wyoming, US
(FMNH 2262) (juvenile) premaxillary tooth (?x4.4x? mm; three lateral teeth (~12x5.5x? mm; 8.1x4.3x? mm; ?x4x? mm) (Gignac, Makovicky, Erickson and Walsh, 2010)
(YPM 4886) distal metatarsal IV (Ostrom, 1970)
(YPM 4887) distal femur (Ostrom, 1970)
(YPM 5275) tooth (Ostrom, 1970)
Aptian-Albian, Early Cretaceous
Cloverly Formation, Montana, US
(AMNH 21609; = AMNH uncatalogd in Ostrom, 1970 p 209?) 3 teeth (AMNH online)
(MOR 947) frontal (MOR online)
(MOR 1182) (adult) specimen including caudal series, radius and manual phalanx (Parsons and Parsons, 2006)
(YPM 64996) bone fragments (YPM online)
Early Cretaceous
Arizona, US
? teeth (Thayer and Ratkevich, 1996)
Aptian-Middle Albian, Early Cretaceous
Trinity Group, Texas
(FMNH 2-51#1) premaxillary tooth (9 mm), seven lateral teeth (4, 5, 12.2, 16.2, 21.3, 21.8, 22.5 mm), four tooth fragments (one juvenile) (Gallup, 1975)
(FMNH 3-51#1) eight teeth (4.6, 4.7, 5, 5.6, 6.9, 6.9, 7, 11.9 mm), eight tooth fragments (Gallup, 1975)
(FMNH 8s-52#1) three teeth (6.1, 12.3, 13.5 mm) (Gallup, 1975)
(FMNH 9s-51#1) two teeth (7.3, 10 mm) (Gallup, 1975)
(FMNH 11s-52#1) three teeth (6.8, 9, 9 mm), tooth fragment (Gallup, 1975)
(FMNH 121-50) tooth (2 mm) (Gallup, 1975)
(FMNH 123-50/124-50) juvenile tooth (8.4 mm), two teeth (13.6, 19.5 mm) (Gallup, 1975)
(FMNH 202-50) tooth (17.9 mm), tooth fragment (Gallup, 1975)
(FMNH 219-50) tooth (Gallup, 1975)
(FMNH 222-50) tooth fragment (Gallup, 1975)
(FMNH Turtle Gully) tooth (8.5 mm), tooth fragment (Gallup, 1975)
(FMNH UvZ) tooth (17 mm), incomplete pedal phalanx III-3 (36 mm) (Gallup, 1975)
(FMNH coll.) dorsal zygapophyseal fragment (Gallup, 1975)
? seven teeth (Brinkman, Cifelli and Czaplewski, 1998)
Cenomanian, Early Cretaceous
Woodbine Formation, Texas, US
Material- ? teeth, limb fragments, manual ungual and/or pedal ungual (Main, Noto and Scotese, 2011)
? teeth (Bennett et al., 2012)
Aptian-Albian, Early Cretaceous
Antlers Formation, Oklahoma, US
(OMNH 16564) tooth (FABL 6.65 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 16565) tooth (FABL 7.09 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 16566) tooth (FABL 3.64 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 17709) tooth (FABL 9.06 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 34203) tooth (FABL 6.19 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49410) tooth (FABL 9.56 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49411) tooth (FABL 9.24 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49412) tooth (FABL 7.32 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49415) tooth (FABL 7.26 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 50268) (subadult) two teeth, partial prootic, occiput, two partial mid dorsal ribs, fifth sacral vertebra (22.5 mm), incomplete proximal chevron, coracoid (73.27 mm), humeral fragments, partial metacarpals I (31.17, 31.04 mm), partial phalanges I-1 (~54.04 mm), incomplete manual ungual I, partial metacarpals II, partial phalanx II-1, partial phalanx II-2, phalanx III-2 (14.56 mm), phalanx III-3 (39.34 mm), femoral fragments, partial metatarsal I, incomplete phalanx II-2 (33.7 mm), partial pedal unguals II, partial phalanx III-3, partial phalanx IV-3 (27.26 mm), phalanx IV-4 (23.94 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 52711) tooth (FABL 6.51 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 53492) tooth (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 54004) three teeth (FABL 8.15, 6.47, 4.75 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 63061) material including femur and tibia (Pascucci, D'Emic and Turner, 2020)
Late Aptian, Early Cretaceous
Dinosaur Park / Cherokee-Sanford Brick Clay Pit / Muirkirk Clay Pit USNM 41614, Arundel Formation, Prince George's County, Maryland, US
?(USNM 497715) tooth (~6.6x~5.2x? mm) (Lipka, 1998)
?(USNM 497716) incomplete tooth (~8.5x~3.5x? mm) (USNM online)
?(USNM 497717) tooth (~10.3x~8.2x? mm) (Brownstein, 2018)
?(USNM 497719) tooth (~8.1x~5.3x? mm) (Lipka, 1998)
?(USNM 497720) premaxillary tooth (Lipka, 1998)
?(USNM 497721) incomplete tooth (?x~4.6x? mm) (Brownstein, 2018)
?(USNM 497727) partial tooth (?x~5.7x? mm) (Lipka, 1998)
?(USNM 497742) tooth (Frederickson, Lipka and Cifelli, 2018)
?(USNM 497743) tooth (Frederickson, Lipka and Cifelli, 2018)
?(USNM 497744) tooth (Frederickson, Lipka and Cifelli, 2018)
?(USNM 497745) tooth (Frederickson, Lipka and Cifelli, 2018)
?(USNM 497746) tooth (Frederickson, Lipka and Cifelli, 2018)
?(USNM 497747) tooth (Frederickson, Lipka and Cifelli, 2018)
Early Cretaceous?
US?
(YPM 64979) vertebra (YPM online)
Diagnosis- (after Turner et al., 2012) four asymmetrical, subincisiform premaxillary teeth; maxillary fenestra located dorsal to level of promaxillary fenestra; 15 maxillary teeth; lobate anteriormost process of jugal beneath antorbital fenestra; lacrimal boss prominent; prefrontal greatly reduced; 16 nearly isodont dentary teeth; all teeth with anterior and posterior serrations; denticles of posterior serrations nearly twice as large as denticles of anterior serrations on all teeth; large, wide calcaneum.
Montana and Wyoming specimens- Brown discovered AMNH 3015 in 1931, and AMNH 3037 in 1932. As Chure and McIntosh (1989) stated, Brown proposed "Daptosaurus agilis" with the intended holotype AMNH 3015, but "never got around to describing these animals, and it was left to John Ostrom to describe and name them many years later in 1970, based in many cases on much better and more diagnostic material which he had collected." "None of Brown's names were ever published (until Glut 1972 [for "Peltosaurus"]) and it does no harm to mention them here, as Ostrom's names are well established taxa having full priority. Brown's names, however, were used in public lectures, and for a time on some exhibited specimens. In addition, they were written on photographs and preliminary skeletal reconstructions made under Brown's direction." In his 1931 field report, Brown first mentioned Deinonychus in saying "closely associated with one Tentontosaurus skeleton is a small carnivorous dinosaur but encased in lime difficult to prepare" (Grellet-Tinner and Makovicky, 2006).
The YPM paratypes were found in 1964-1966 (YPM online), with the holotype being discovered in late August 1964. The YPM quarry was reopened in 1993 by the MOR, resulting in abundant new material (MOR 747; Maxwell and Witmer, 1996)- "one cervical vertebra, several caudal vertebrae, a radius and ulna, two manus, two pes, and portions of the skull, including ... the prefrontal, frontal, parietal, quadrate, laterosphenoid, and portions of the braincase. Other cranial elements in the new sample, such as the nasal, lacrimal, and ectopterygoid are the best samples collected to date." Powers et al. (2022) describe the maxilla of YPM 5232 in detail based on CT scans.
MCZ 4371 was found in July 1974 and described by Ostrom (1976). Ostrom (1969b) believed that "YPM 5203 (a caudal sequence), 5205 (left and right pes), 5206 (left and right manus) and 52010 (several vertebrae and an incomplete skull and jaws) belong to a single individual because of distinct preservation common to these alone and the fact that they are all slightly larger than other comparable lements obtained at the Yale site" but could not prove it. The three articulated caudal series (YPM 5201-5203) show at least three individuals were present among YPM 5201-5265, and Maxwell and Witmer report four right splenials showing at least four were present.
Ostrom (1969a, b) misidentified the scapholunare as an ulnare, the semilunate carpal as the radiale (= scapholunare in maniraptoriforms), and the coracoid as the pubis.
Parsons and Parsons (2002) initially identified MOR 1178 as a new troodontid-like taxon based on an apparent parietal sagittal crest, dorsal groove on distal caudal vertebrae, robust metatarsal IV and pedal ungual ratios similar to Troodon. However, in the period between abstract submission and conference poster, they had reidentified it as a subadult Deinonychus. This was published in their 2003 paper, with no mention of the ectopterygoid or metatarsal IV that were noted in 2002. They may have been reidentified as the squamosal fragment and metatarsal II. The supposed parietal fragment was reidentified as a sacral fragment. Some features such as highly elongated forelimbs and curved manual ungual III are seen as ontogenetic differences from adult specimens.
Parsons and Parsons (2004) describe another young Deinonychus (MCZ 8791) which was discovered in 1982. It is smaller than MOR 1178 and has even longer forelimbs. Parsons and Parsons (2013) describe further details of this specimen.
Parsons and Parsons (2006) report on "newly discovered specimen" MOR 1182
Makovicky and Grellet-Tinner (2000) and Grellet-Tinner's (2005) thesis described Deinonychus eggshells preserved with AMNH 3015 but left unidentified for decades, which were officially published as Grellet-Tinner and Makovicky (2006).
Texas specimens- Main et al. (2011) reported material of dromaeosaurids and basal tetanurines from the Woodbine Formation of Texas, the former of which are provisionally listed as Deinonychus here.
Maryland specimens- USNM 497715-497717, 497719-497721 and 497727 were discovered in 1998. Lipka (1998) described UNSM 497715, 497719-497720 and 497727 as 'deinonychids' that likely "represent an East Coast occurrence of D. antirrhopus", with a figure of USNM 497727 labeled "Probable deinonychid lateral tooth." Brownstein's (2018) Figure 2 includes "Teeth of Deinonychus sp. (1-4)", which are USNM 497719, 497717, 497721 and 497715 respectively based on the USNM online catalog.
Relationships- Deinonychus was traditionally identified as a velociraptorine, but has most recently been recovered as a dromaeosaurine (Senter et al., 2012; Foth et al., 2014; Lee et al., 2014), sister taxon to Velociraptorinae+Dromaeosaurinae (Longrich and Currie, 2009), or a velociraptorine after all (Brusatte et al., 2014).
References- Ostrom, 1969a. A new theropod dinosaur from the Lower Cretaceous of Montana. Postilla. 128, 17 pp.
Ostrom, 1969b. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural History. 30, 165 pp.
Ostrom, 1970. Stratigraphy and paleontology of the Cloverly Formation (lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Bulletin of the Peabody Museum of Natural History. 35, 234 pp.
Ostrom, 1974. The pectoral girdle and forelimb function of Deinonychus (Reptilia: Saurischia): a correction. Postilla. 165, 11 pp.
Gallup, 1975. Lower Cretaceous dinosaurs and associated vertebrates from north-central Texas in the Field Museum of Natural History. MS thesis, University of Texas at Austin. 159 pp.
Ostrom, 1976. On a new specimen of the Lower Cretaceous theropod dinosaur Deinonychus antirrhopus. Breviora. 439, 21 pp.
Paul. 1988. The small predatory dinosaurs of the mid-Mesozoic: The horned theropods of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus - and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River - Deinonychus, Velociraptor and Saurornitholestes. Hunteria. 2(4), 1-9.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves) 1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Maxwell, 1993. Neonate dinosaur remains and dinosaur eggshell from the Lower Cretaceous Cloverly Formation of Montana. Journal of Vertebrate Paleontology. 13(3), 48A.
Maxwell and Ostrom, 1995. Taphonomy and paleobiological implications of Tenontosaurus-Deinonychus associations. Journal of Vertebrate Paleontology. 15, 606-712.
Maxwell and Hallas, 1996. First skull roof and braincase of Deinonychus. Journal of the American Osteopathic Association. 96(9), 563.
Maxwell and Witmer, 1996. New material of Deinonychus (Dinosauria; Theropoda). Journal of Vertebrate Paleontology. 16(3), 51A.
Thayer and Ratkevich, 1996. Dinosaur remains in southern Arizona. In Wolberg and Stump (eds.). Dinofest International Symposium Program and Abstracts, 108.
Witmer and Maxwell, 1996. The skull of Deinonychus (Dinosauria; Theropoda): new evidence and insights. Journal of Vertebrate Paleontology. 16(3), 73A.
Maxwell, 1997. New insights into Deinonychus, Tenontosaurus, and other dinosaurs from the Cloverly Formation (Lower Cretaceous) of south-central Montana. The Dinosaur Report. Spring, 16-17.
Brinkman, Cifelli and Czaplewski, 1998. First occurrence of Deinonychus antirrhopus (Dinosauria: Theropoda) from the Antlers Formation (Lower Cretaceous: Aptian-Albian) of Oklahoma. Bulletin of the Oklahoma Geological Survey. 146, 1-27.
Lipka, 1998. The affinities of the enigmatic theropods of the Arundel Clay facies (Aptian), Potomac Formation, Atlantic Coastal Plain of Maryland. in Lucas, Kirkland and Estep (eds). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 229-234.
Chiappe and Grellet-Tinner, 2000. Dinosaur eggshells and the origin of birds. The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History 12.
Makovicky and Grellet-Tinner, 2000. Association between theropod eggshell and a specimen of Deinonychus antirrhopus. First International Symposium on Dinosaur eggs and babies. Extended abstracts, 123-128.
Gishlick, 2001. The function of the manus and forelimb of Deinonychus antirrhopus and its importance for the origin of avian flight. in Gauthier and Gall (eds). New Perspectives on the Origin and Early Evolution of Birds. 301-318.
Parsons and Parsons, 2002. Data from new dinosaur material discovered in the Cloverly Formation of Central Montana. Journal of Vertebrate Paleontology. 22(3), 95A.
Parsons and Parsons, 2003. Description of a new immature specimen of Deinonychus antirrhopus (Saurischia, Theropoda). Journal of Vertebrate Paleontology. 23(3), 86A.
Carney and Gisklick, 2004. Utilizing digital techniques within an extant phylogenetic bracketing paradigm to reconstruct and analyze the role of articular cartilaginous structures in dromaeosaur forelimb function. Journal of Vertebrate Paleontology. 24(3).
Parsons and Parsons, 2004. Postcranial ontogeny of Deinonychus antirrhopus (Saurischia, Theropoda). Journal of Vertebrate Paleontology. 24(3), 178A-179A.
Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case study of saurischian dinosaur relationships and avian evolution. PhD thesis, University of Southern California. 221 pp.
Parsons and Parsons, 2005. A comparison of some of the important postcranial features found within the ontogenies of Deinonychus antirrhopus (Saurischia, Theropoda) and Velociraptor mongoliensis (Saurischia, Theropoda). Journal of Vertebrate Paleontology. 25(3), 99A.
Grellet-Tinner and Makovicky, 2006. A possible egg of the dromaeosaur Deinonychus antirrhopus: Phylogenetic and biological implications. Canadian Journal of Earth Sciences. 43, 705-719.
Parsons and Parsons, 2006. Morphology and size of an adult specimen of Deinonychus antirrhopus, (Saurischia, Theropoda). Journal of Vertebrate Paleontology. 26(3), 109A.
Gignac, Makovicky, Erickson and Walsh, 2007. The effect of tooth morphology on indentation force in non-avian theropod dinosaurs. Journal of Vertebrate Paleontology. 27(3), 81A.
Parsons and Parsons, 2007. Avian-like manual phalanx found within gut contents of Lower Cretaceous dromaeosaurid: New data on the feeding behavior of Deinonychus antirrhopus (Saurischia: Theropoda). Journal of Vertebrate Paleontology. 27(3), 128A.
Roach and Brinkman, 2007. A reevaluation of cooperative pack hunting and gregariousness in Deinonychus antirrhopus and other nonavian theropod dinosaurs. Bulletin of the Peabody Museum of Natural History. 48(1), 103-138.
Gignac, Makovicky, Erickson and Walsh, 2008. Bite force estimates for Deinonychus antirrhopus using tooth indentation simulations. Journal of Vertebrate Paleontology. 28(3), 84A.
Fowler, 2009. The grasping foot of Deinonychus: Implications for predator ecology, evolution of the perching foot, and a new hypothesis for the origin of flight in birds. Journal of Vertebrate Paleontology. 29(3), 98A.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Parsons and Parsons, 2009. Further descriptions of the osteology of Deinonychus antirrhopus (Saurischia, Theropoda). Bulletin of the Buffalo Society of Natural Sciences. 38, 43-54.
Gignac, Makovicky, Erickson and Walsh, 2010. A description of Deinonychus antirrhopus bite marks and estimates of bite force using tooth indentation simulations. Journal of Vertebrate Paleontology. 30(4), 1169-1177.
Main, Noto and Scotese, 2011. New theropod material from the Cretaceous (Cenomanian) Woodbine Formation of North Central Texas: Paleobiogeographic and paleoecological implications. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 150.
Bennett, Main, Noto, Anderson and Vranken, 2012. Microvertebrate paleoecology, wildfires and biodiversity of coastal Appalachia in the Cretaceous (Cenomanian) Woodbine Formation at the Arlington archosaur site, North Texas. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 63.
Parsons and Parsons, 2012. The first intact scapular glenoid region of Deinonychus antirrhopus and the consequent re-interpretation of dromaeosaurid features that enhanced the evolution of avian flight. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 155.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria: Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 1-206.
Parsons and Parsons, 2013. Ontogeny or phylogeny? Cladistic placement of a juvenile dromaeosaurid from the Lower Cretaceous of Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 188.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan culminated in rapid rates of evolution across the dinosaur-bird transition. Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Parsons and Parsons, 2015. Morphological variations within the ontogeny of Deinonychus antirrhopus (Theropoda, Dromaeosauridae). PLoS ONE. 10(4), e0121476.
Brownstein, 2018. The biogeography and ecology of the Cretaceous non-avian dinosaurs of Appalachia. Palaeontologia Electronica. 21.1.5A, 1-56.
Frederickson, Lipka and Cifelli, 2018. Faunal composition and paleoenvironment of the Arundel Clay (Potomac Formation; Early Cretaceous), Maryland, USA. Palaeontologia Electronica. 21.2.31A, 1-24.
Pascucci, 2019. Bone Histology of the theropod dinosaur Deinonychus antirrhopus. Masters thesis, Adelphi University. 38 pp.
Pascucci, D'Emic and Turner, 2020. Osteohistology of the Early Cretaceous theropod Deinonychus anitirrhopus. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 265.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as 2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist in testing and constructing morphological characters. Journal of Vertebrate Paleontology. 41(5), 2010087.

Itemirus Kurzanov, 1976a
I. medullaris Kurzanov, 1976a
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (PIN 327/699) (adult) braincase
Referred- ?(CCMGE 456/12457) anterior dorsal centrum (Nessov, 1995)
?(CCMGE 461/12457) tooth (Nessov, 1995)
?(CCMGE? ?/12457; lost?) (juvenile) maxilla (Nessov, 1995)
?(CCMGE? ?/12457) teeth (Nessov, 1995)
?(ZIN PH 11/16) pedal phalanx II-2 (96.4 mm) (Sues and Averianov, 2014)
?(ZIN PH 12/16) pedal ungual III (Sues and Averianov, 2014)
?(ZIN PH 13/16) incomplete first dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 73/16) incomplete posterior dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 74/16) anterior dorsal centrum (Sues and Averianov, 2014)
?(ZIN PH 75/16) posterior dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 89/16) anterior cervical vertebra (Sues and Averianov, 2014)
?(ZIN PH 90/16) incomplete posterior cervical vertebra (Sues and Averianov, 2014)
?(ZIN PH 92/16) incomplete anterior cervical vertebra (Sues and Averianov, 2014)
?(ZIN PH 96/16) first dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 182/16) metacarpal I (39 mm) (Sues and Averianov, 2014)
?(ZIN PH 183/16) partial metacarpal I (~76 mm) (Sues and Averianov, 2014)
?(ZIN PH 967/16) frontal (Sues and Averianov, 2014)
?(ZIN PH 2263/16) posterior cervical vertebra (Sues and Averianov, 2014)
?(ZIN PH 2269/16) incomplete posterior dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 2273/16) proximal caudal vertebra (Sues and Averianov, 2014)
?(ZIN PH 2338/16) dentary fragment (Sues and Averianov, 2014)
?(ZIN PH 2344/16) premaxillary tooth (Sues and Averianov, 2014)
?(ZIN PH 2346/16) incomplete distal caudal vertebra (Sues and Averianov, 2014)
?(ZIN PH 2351/16) lateral tooth (Sues and Averianov, 2014)
?(ZIN PH 2352/16) lateral tooth (Sues and Averianov, 2014)
?(ZIN PH 2353/16) lateral tooth (Sues and Averianov, 2014)
?(ZIN PH? ?/16) teeth (Sues and Averianov, 2014)
?(ZIN PH? ?/16) pedal phalanx II-2 fragment (Sues and Averianov, 2014)
Diagnosis- (after Sues and Averianov, 2014) differs from Dromaeosaurus in- basipterygoid recesses on dorsolateral surfaces of basipterygoid processes present (also in Velociraptor, Tsaagan and Bambiraptor); mesial serrations of teeth significantly smaller than distal serrations (also in most dromaeosaurids except Utahraptor and Achillobator; teeth tentatively referred). Differs from Utahraptor in- pleurocoels present in anterior dorsals (also in other dromaeosaurids except Yurgovuchia; vertebrae tentatively referred). Differs from both Utahraptor and Achillobator in lateral surfaces of dorsal centra have deep, emarginated fossae (also in Deinonychus; vertebrae tentatively referred).
(suggested) occipital condyle wider than foramen magnum.
Comments- The holotype braincase was discovered in 1958, and described by Kurzanov as the new genus Itemirus in 1976(a,b). Kurzanov assigned Itemirus to its own family, Itemiridae, which he placed sister to Tyrannosauridae, with these both sister to Dromaeosauridae (including Stenonychosaurus but not Saurornithoides) within the Carnosauria. Theropod braincases were poorly known at the time so there was little to compare Itemirus to and much of the description is outdated. Some aspects were described in a more modern perspective by Currie and Zhao (1994), in comparison with Troodon. Chure and Madsen (1998) found the deep basipterygoid recesses to be similar to their cf. Stokesosaurus braincase, but noted several characters were less derived than Stokesosaurus and tyrannosaurids. This feature led Holtz (2004) to refer to Itemirus as a possible tyrannosauroid, though it has been since noted in most dromaeosaurids and basal paravians. Longrich and Currie (2009) found Itemirus to clade in Velociraptorinae, based on a paravian matrix. Miyashita (2011) used a coelurosaur matrix which grouped Itemirus with tyrannosauroids, but noted (DML, 2011) that few steps are needed to constrain it to Dromaeosauridae.
Sues and Averianov (2004) referred additional material to Itemirus, describing it as a dromaeosaurid and stating it reaches sizes comparable to Utahraptor. Nessov (1995) referred the maxilla CCMGE 600/12457 to Alectrosaurus, but it was reidentified as dromaeosaurid by Averianov and Sues (2012) and described as cf. Itemirus by Sues and Averianov (2014). Tanaka et al. (2021) referred it to their new carcharodontosaur Ulughbegsaurus along with maxillary fragment ZIN PH 357/16 that was described as cf. Itemirus by Sues and Averianov. CCMGE 456/12457 was figured by Nessov as a segnosaur, but later reidentified as cf. Itemirus by Sues and Averianov (2014). The unnumbered maxilla and teeth listed above were identified as Deinonychosauria and Dromaeosauridae by Nessov. The maxilla is described as shorter and wider than Deinonychus, though not mentioned by Sues and Averianov (2014) as no obviously corresponding specimen was found, while the teeth (including that in plate 1 figure 4 of Nessov, assigned to cf. Itemirus by Sues and Averianov, 2014) were described as small. While Sues and Averianov (2004) refer "several braincases" to Itemirus in their abstract, only the holotype was noted in the resulting publication. Sues (pers. comm. 2014) stated these were fragments that showed few diagnostic features once prepared, so their identification remains uncertain. ZIN PH 73/16 was mislabeled ZIN PH 2273/16 in Sues and Averianov's (2014) figure. The final publication of the Bissekty dromaeosaurid was Sues and Averianov's (2014), which tentatively referred all material to Itemirus because "there are no morphological differences among the few comparable remains." Based on the limited materal, this must remain provisional. Several of the characters in this publication are miscoded (dorsal tympanic recess is present; supratemporal fossa border is simply curved; frontal's postorbital process not sharply demarcated; dentary has a labial groove; posterior dorsal parapophyses are not on prominent pedicles; hyposphenes are not separated), most favoring a dromaeosaurid identity. Given these miscodings, Sues and Averianov's recovery of Itemirus as a dromaeosaurine more derived than Tsaagan may be questioned.
References- Kurzanov, 1976a. Stroyeniye mozgovoy korobki karnozavra Itemirus gen. nov. i nekotoryye voprosy kranial-noy anatomii dinozavrov. Paleontologicheskii Zhurnal. 1976(3), 127-137.
Kurzanov, 1976b. Brain-case structure in the carnosaur Itemirus n. gen. and some aspects of the cranial anatomy of dinosaurs. Paleontological Journal. 1976(3), 361-369.
Martinson, Nessov and Starobogatov, 1986. Unique find of gill apparatus of bivalve molluscs Trigonioidoidea from the Cretaceous deposits of Kyzylkum. Byuleten' Moskovskogo Obschestva Ispytatelei Prirody, Otdel Geologicheskii. 61, 94-98.
Currie and Zhao, 1994. A new troodontid (Dinosauria, Theropoda) braincase from the Dinosaur Park Formation (Campanian) of Alberta. Canadian Journal of Earth Sciences. 30(10-11), 2234-2247.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology, and paleobiogeography. Institute for Scientific Research on the Earth's Crust. 1-156.
Chure and Madsen, 1998. An unusual braincase (?Stokesosaurus clevelandi) from the Cleveland-Lloyd Dinosaur Quarry, Utah (Morrison Formation; Late Jurassic). Journal of Vertebrate Paleontology. 18(1), 115-125.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 111-136.
Sues and Averianov, 2004. Dinosaurs from the Upper Cretaceous (Turonian) of Dzharakuduk, Kyzylkum Desert, Uzbekistan. Journal of Vertebrate Paleontology. 24(3), 51A-52A.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research. 28(3), 532-544.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. Proceedings of the National Academy of Sciences. 106(13), 5002-5007.
Miyashita, 2011. Cranial morphology of the basal tyrannosauroid Itemirus medullaris and evolution of the braincase pneumaticity in non-avian coelurosaurs. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 159.
Miyashita, DML 2011. https://web.archive.org/web/20180115091932/http://dml.cmnh.org/2011Nov/msg00368.html
Averianov and Sues, 2012. Skeletal remains of Tyrannosauroidea (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan. 34, 284-297.
Sues and Averianov, 2014. Dromaeosauridae (Dinosauria: Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan and the phylogenetic position of Itemirus medullaris Kurzanov, 1976. Cretaceous Research. 51, 225-240.
Tanaka, Anvarov, Zelenitsky, Ahmedshaev and Kobayashi, 2021. A new carcharodontosaurian theropod dinosaur occupies apex predator niche in the early Late Cretaceous of Uzbekistan. Royal Society Open Science. 8: 210923.