Deinonychosauria Colbert and Russell, 1969
Definition- (Deinonychus antirrhopus <- Passer domesticus) (Holtz and Osmolska, 2004; modified from Padian, 1997)
Other definitions- (Troodon formosus + Dromaeosaurus albertensis) (Holtz and Osmolska, 2004; modified from Sereno, 1997)
(Troodon formosus + Velociraptor mongoliensis) (modified from Sereno, 1998)
(Troodon formosus + Velociraptor mongoliensis, - Ornithomimus edmontonicus, Passer domesticus) (Sereno, in press)
= Dromaeosauria Chatterjee and Templin, 2007
Comments- There have been two basic suggested definitions for Deinonychosauria, one stem-based (Deinonychus <- Passer) by Padian (1997) and the other node based (Troodon + dromaeosaurids) by Sereno (1997). Sereno's latest (in press) definition is a modification of the latter, but explicitly excludes birds and ornithomimosaurs. I prefer Padian's because it is based on the eponymous genus, and Colbert and Russell (1969) did not originally specify the inclusion of troodontids. They only include dromaeosaurids in the taxon, and only mention Dromaeosaurus, Deinonychus and Velociraptor as members of that family. Given the trichotomy between dromaeosaurids, troodontids and ornithurines presented on this site, Sereno's (in press) definition of Deinonychosauria cannot be applied anywhere on the tree. The present formulation of Deinonychosauria is synonymous with Dromaeosauridae in the topology presented on this site, but the taxa retain unique entries because I consider it likely eumaniraptoran topology will be labile for the near future.

Dromaeosauridae Matthew and Brown, 1922 sensu Russell, 1969
Definition- (Velociraptor mongoliensis <- Troodon formosus) (modified from Sereno, 1998)
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 <- Ornithomimus edmontonicus, Troodon formosus, Passer domesticus) (Sereno, in press)
= Ornithodesmidae Hooley, 1913
= Itemiridae Kurzanov, 1976
= Dromaeosauridae sensu Hu, Hou, Zhang and Xu, 2009
Definition- (Deinonychus antirrhopus <- Troodon formosus, Passer domesticus)
= 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) topology, and troodontids and birds dromaeosaurids in my corrected version of the TWG matrix. 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, 1993), 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).
Not dromaeosaurids- 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 Eumaniraptora incertae sedis.
References- Hooley, 1913. On the skeleton of Ornithodesmus latidens; an ornithosaur from the Wealden Shales of Athersfield (Isle of Wight). Quarterly Journal of the Geological Society. 69, 372-421.
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).

Archaeornithoides Elzanowski and Wellnhofer, 1992
A. deinosauriscus Elzanowski and Wellnhofer, 1992
Late Campanian, Late Cretaceous
Djadokhta Formation, Mongolia
Holotype- (ZPAL MgD-II/29) (juvenile) (skull ~50 mm) maxillae, maxillary teeth, anterior jugal, possible vomer fragment, palatine, possible ectopterygoid fragment, possible parasphenoid, dentaries
Comments- Discovered in 1965, this specimen was originally mentioned by Elzanowski (1983), then by Paul (1988) as a possible aublysodontine tyrannosaurid. Elzanowski and Wellnhofer (1992, 1993) originally suggested Archaeornithoides was most closely related to spinosaurids, troodontids and Lisboasaurus, and that these were all avialans. Spinosaurids are now known to be basal tetanurines while Lisboasaurus is a crocodilian. However, in their 1993 article, they suggest Archaeornithoides is more closely related to birds than Lisboasaurus, confusing matters slightly. They rejected a troodontid relationship based on the broad maxillary palatal shelf and unserrated teeth, but Currie (2000) noted this is invalid as Troodon has the former, while Clark et al. (2002) noted it's invalid because Byronosaurus has the former and that juvenile Djadokhta troodontids have the latter. Currie further suggested that Archaeornithoides may be a juvenile Saurornithoides mongoliensis, while Averianov and Sues (2007) suggested it could be a juvenile Byronosaurus. An additional possibility is that it may be a juvenile of one of the undescribed basal troodontids from the Djadokhta, such as IGM 100/1005 or 100/1323. Although Averianov and Sues stated that Chiappe et al. (1996) proposed Archaeornithoides was a juvenile dromaeosaurid, the latter authors actually only state they believe the lack of serrations to be a juvenile character of birdlike theropods, as they mistakenly assigned IGM 100/972 and 100/974 to Dromaeosauridae at the time. It is tentatively placed in Dromaeosauridae here based on including in in a revised version of Senter's (2007) matrix, where it emerges as a dromaeosaurid outside the subclade closer to Dromaeosaurus than Microraptor.
References- Elzanowski, 1983. Birds in Cretaceous Ecosystems. Acta Palaeontologia Polonica, 28 (1-2), 75-92.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Elzanowski and Wellnhofer, 1992. A new link between theropods and birds from the Cretaceous of Mongolia. Nature. 359, 821-823.
Elzanowski and Wellnhofer, 1993. Skull of Archaeornithoides from the Upper Cretaceous of Mongolia. American Journal of Science. 293-A, 235-252.
Chiappe, Norell and Clark, 1996. Phylogenetic position of Mononykus (Aves: Alvarezsauridae) from the Late Cretaceous of the Gobi Desert. Memoirs of the
Queensland Museum. 39(3), 557-582.
Currie, 2000. Theropod dinosaurs from the Cretaceous of Mongolia. in Benton, Shishkin, Unwin and Kurochkin (eds). The Age of Dinosaurs in Russia and Mongolia. pp 434-455.
Clark, Norell and Makovicky, 2002. Cladistic approaches to the relationships of birds to other theropod dinosaurs. in Chiappe and Witmer (eds.). Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press. 31-64.
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.

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) four partial dorsal ribs, seven gastralia or sternal ribs, scapulae (65 mm), coracoids, humeri (89 mm), radii (63 mm), ulnae (62, 64 mm), radiale, centrale, ulnare, semilunate carpal, 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), 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). The total combination of characters suggests Yixianosaurus may be a basal dromaeosaurid or troodontid, as it usually emerges in these positions when included in a revised version of Senter's (2007) matrix.
Reference- Xu and Wang, 2003. A new maniraptoran dinosaur from the Early Cretaceous Yixian Formation of Western Liaoning. Vertebrata PalAsiatica. 41(3), 195-202.

Unenlagiinae Bonaparte, 1999 sensu Makovicky et al., 2005
Definition- (Unenlagia comahuensis <- Microraptor zhaoianus, Velociraptor mongoliensis, Dromaeosaurus albertensis, Passer domesticus) (Sereno, in press)
Other definitions- (Unenlagia comahuensis <- Velociraptor mongoliensis) (Makovicky, Apesteguia and Agnolin, 2005)
= Unenlagiidae Bonaparte, 1999
Comments- This clade was erected by Makovicky et al. (2005) to contain Unenlagia, Buitreraptor and Rahonavis at the base of Dromaeosauridae. It has been recovered in every subsequent Theropod Working Group analysis, including the modified version used by Senter (2007).
Sereno's (in press) definition replaces Makovicky et al.'s, which only included Velociraptor as an external specifier. As I detailed in http://dml.cmnh.org/2005Oct/msg00372.html , I don't find support for a clade of Unenlagia, Rahonavis and Buitreraptor exclusive of dromaeosaurids, troodontids and birds very convincing. I also don't find the position of these taxa within Eumaniraptora to be very stable, so I'd recommend adding Troodon and Archaeopteryx as external specifiers.

Unenlagia Novas and Puerta, 1997
Diagnosis- deep lateral fossae at base of posterior dorsal neural spines; marked inflection of posterodorsal ilial edge at level of supratrochanteric process; posteriorly curved distal pubis.
Comments- The posteriorly curved distal pubis cited as an apomorphy by Calvo et al. (2004) is a basal paravian trait also seen in microraptorians, Bambiraptor, Buitreraptor, Rahonavis, Confuciusornis and other taxa. Calvo et al. also cited the marked inflection of posterodorsal ilial edge at level of supratrochanteric process, but this is due to the tall ilium and prominant supratrochanteric process, which are plesiomorphic and apomorphic respectively. Buitreraptor approaches the condition, while Achillobator has it. The dorsal neural spine fossae aren't known in related taxa, but are only present in some vertebrae of Unenlagia comahuensis and may be easily missed. Thus, the referral of U? paynemili to Unenlagia is provisional and may turn out to be incorrect.
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). However, Neuquenraptor's femur is more robust in anterior view than can be accounted for by distal incompleteness (pers. obs.; Novas and Pol, 2005; contra Makovicky et al.), given the position of curvature in lateral view. Neuquenraptor's pedal phalanx II-1 differs from the one tentatively referred to U? paynemili in some minor ways- ligament tubercle more dorsally placed; taller compared to length. Unenlagia? paynemili's preserved ungual appears to be a manual ungual, and is thus not comparable to Neuquenraptor's preserved unguals. Furthermore, the ungual and phalanx were only tentatively referred to Unenlagia, being found 5 and 12.5 meters away from the holotype of U? paynemili. The tibiae of U. comahuensis and Neuquenraptor differ in that Neuquenraptor's has a shorter medially projecting portion just proximal to the tarsus. 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.
References- Novas and Puerta, 1997. New evidence concerning avian origins from the Late Cretaceous of Patagonia. Nature. 387: 390-392.
Calvo, Porfiri and Kellner, 2004. On a new maniraptoran dinosaur (Theropoda) from the Upper Cretaceous of Neuquén, Patagonia, Argentina. Arquivos do Museo Nacional. 62, 549–566.
Makovicky, Apesteguía and Agnolín, 2005. The earliest dromaeosaurid theropod from South America. Nature. 437, 1007-1011.
Novas and Pol, 2005. New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia. Nature 433: 858 - 861.
U. comahuensis Novas and Puerta, 1997
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of the Rio Neuquen Subgroup, Argentina
Holotype- (MCF PVPH 78) eighth dorsal vertebra, tenth dorsal vertebra, thirteenth dorsal vertebra, three dorsal ribs, sacrum, two chevrons, scapula, humerus (270 mm), ilia, pubes, ischium, femur (381 mm), tibia (431 mm)
Diagnosis- (after Calvo et al., 2004) lateral ridge posterior to deltopectoral crest; angle between distal edge of deltopectoral crest and humeral shaft 116 degrees; ridge dividing brevis fossa.
Comments- The phylogenetic placement of Unenlagia is controversial. Originally described as an avialan (Novas and Puerta, 1997), this was coorborated by Novas and Pol (2002) and Novas (2004). Forster et al. (1998) suggested it was an archaeopterygid, while Holtz (2000) found it equally supported as an avialan sister taxon to Rahonavis, an avialan sister to Archaeopteryx + Ornithurae or an ornithurine sister to pygostylians. Norell and Makovicky (1999) noted several traits similar to dromaeosaurids and not as birdlike as originally suggested, which is reflected in the results of the Theropod Working Group (Norell et al., 2001 and modifications) placing it inside Dromaeosaurinae + Velociraptorinae until the publication of Buitreraptor (Makovicky et al., 2005). This latter paper interprets Unenlagia as a basal dromaeosaurid in a clade with Rahonavis and Buitreraptor, termed the Unenlagiinae. Examination of their matrix revealed numerous coding errors, which when corrected and ran with modifications from Mayr et al. (2005) result in a placement inside Avialae, along with Buitreraptor, Rahonavis, Archaeopteryx and Confuciusornis.
Dromaeosaurid characters of Unenlagia include the anteriorly concave preacetabular process (also seen in Shenzhouraptor), and dorsal neural spine tables (also seen in ornithurines, but not basal troodontids, Buitreraptor, Archaeopteryx or Rahonavis). It's placed by Rahonavis and Buitreraptor by Makovicky et al. due to four characters. Achillobator lacks one (supracetabular crest), but has the other three (dorsally concave postacetabular process; elongate obturator process; mesopuby), yet is a dromaeosaurid in Makovicky et al.'s tree (it's miscoded for the first two in their matrix). Unenlagia is sister to Rahonavis in Makovicky et al.'s tree based on five characters. Three of these (all dorsals pleurocoelous; six sacrals; lateral cuppedicus ridge extending posteriorly) are found in some dromaeosaurids too. The other two (elongate preacetabular process; large proximodorsal ischial process) and the supracetabular crest support Makovicky et al.'s hypothesis, but the support for a dromaeosaurid Unenlagia isn't much worse. One might argue placing Achillobator in Unenlagiinae would decrease the reasons for placing Unenlagia in Dromaeosauridae, but then the dromaeosaurid characters of Achillobator (large serrated teeth; short cervicals; highly elongate distal caudal prezygapophyses and chevrons; etc.) would require explanation. In my latest modified versions of the TWG matrix, Unenlagia falls out as a basal dromaeosaurid, but Rahonavis and Buitreraptor are both avialans.
References- Novas and Puerta, 1997. New evidence concerning avian origins from the Late Cretaceous of Patagonia. Nature. 387: 390-392.
Forster, Sampson, Chiappe and Krause, 1998. The Theropod Ancestry of Birds: New Evidence from the Late Cretaceous of Madagascar. Science. 279: 1915-1919.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton II: information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282: 1-45.
Novas, 2000. Avian-like traits in the ilium of Unenlagia comahuensis. The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves Museum of Archaeology and Natural History. 21.
Holtz, 2000. A new phylogeny of the carnivorous dinosaurs. Gaia. 15, 5-61.
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. Yale Peabody Museum. 49-67.
Novas and Pol, 2002. Alvarezsaurid relationships reconsidered: In: Mesozoic Birds, above the heads of Dinosaurs, University of California Press, 121-125.
Novas, 2004. Avian traits in the ilium of Unenlagia comahuensis (Maniraptora: Avialae). In: Feathered Dragons: Studies on the Transition from Dinosaurs to Birds, (eds. by Currie, P.J., Koppelhus, E.B., Shugar, M.A. and Wright, J.L. pp. 150-166.
Makovicky, Apesteguía and Agnolín, 2005. The earliest dromaeosaurid theropod from South America. Nature. 437, 1007-1011.
Mayr, Pohl, and Peters, 2005. A well-preserved Archaeopteryx specimen with theropod features. Science. 310, 1483-1486.
U? paynemili Calvo, Porfiri and Kellner, 2004
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of the Rio Neuquen Subgroup, Argentina
Holotype- (MUCPv-349) humerus (217 mm), pubis (267 mm), distal pubis
Paratypes- ?(MUCPv-343) manual ungual (65 mm on curve)
(MUCPv-409) partial ilium
?(MUCPv-415) pedal phalanx II-1
(MUCPv-416) partial dorsal vertebra
Diagnosis- (after Calvo et al., 2004) angle between distal edge of deltopectoral crest and humeral shaft 116 degrees; postacetabular process posterior to supratrochanteric process taller than U. comahuensis; shallower brevis fossa than U. comahuensis; proximodistally narrow anterior projection of pubic boot.
Comments- Makovicky et al. (2005) are probably correct about Unenlagia? paynemili's preserved phalanx being II-1 instead of I-1 (as identified by Calvo et al., 2004). The preserved ungual appears to be neither pedal I (contra Calvo et al.) nor II (contra Makovicky et al.), but instead a manual ungual. This is due to the enlarged flexor tubercle and dorsal margin which arches high over the proximal surface, characters used by Senter et al. (2004) to identify Achillobator's ungual (which U. paynemili's strongly resembles) as manual.
References- Calvo, Porfiri and Kellner, 2003. A close relative of Unenlagia comahuensis (Theropoda, Maniraptora) from the Upper Cretaceous of Neuquén, Patagonia, Argentina. 18º Congresso Brasileiro de Paleontologia, Brasilia, Resumos. 82-83.
Calvo, Porfiri and Kellner, 2004. On a new maniraptoran dinosaur (Theropoda) from the Upper Cretaceous of Neuquén, Patagonia, Argentina. Arquivos do Museo Nacional. 62, 549–566.
Senter, Barsold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae (Dinosauria, Theropoda). Bulletin of the Gunma Museum of Natural History. 8, 1-20.
Makovicky, Apesteguía and Agnolín, 2005. The earliest dromaeosaurid theropod from South America. Nature. 437, 1007-1011.

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

Itemirus Kurzanov, 1976
I. medullaris Kurzanov, 1976
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (PIN 327/699) (adult) braincase
Referred- several braincases, teeth, postcranial elements (Sues and Averianov, 2004)
Diagnosis- branches of internal carotid artery that enter hypophyseal fossa confluent; small otic recess; posterior foramen for internal carotid artery inside anterior temporal recess.
Comments- The holotype braincase was discovered in 1958, and described by Kurzanov as the new genus Itemirus in 1976. 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 (1993), 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.
Sues and Averianov (2004) referred additional material to this taxon, describing it as a dromaeosaurid and stating it reaches sizes comparable to Utahraptor. These are as yet undescribed, but probably include dromaeosaurid/dromaeosaurine teeth mentioned by Nessov (1995) and Averianov (2007).
Within Dromaeosauridae, Itemirus is more similar to Dromaeosaurus, Tsaagan and Bambiraptor than Velociraptor in having a developed otosphenoid crest; more similar to Bambiraptor in having well defined basipterygoid pockets, which are shallow in Tsaagan and Velociraptor, and absent in Dromaeosaurus; more similar to Velociraptor than Bambiraptor, Tsaagan, Deinonychus and Dromaeosaurus in having subcondylar recesses; intermediate between Dromaeosaurus (no dorsal tympanic recess) and Velociraptor and Tsaagan (deep dorsal tympanic recess) in having a shallow dorsal tympanic recess; more similar to Tsaagan, Velociraptor and Deinonychus than Bambiraptor and Dromaeosaurus in having paracondylar recesses; more similar to Dromaeosaurus and Tsaagan than Deinonychus and Velociraptor in having a foramen magnum taller than wide; more derived than microraptorians in having a smaller foramen magnum.
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.
Currie and Zhao, 1993. 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 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.
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. The Dinosauria Second Edition. University of California Press. 861 pp.
Sues and Averianov, 2004. Dinosaurs from the Upper Cretaceous (Turonian) of Dzharakuduk, Kyzylkum Desert, Uzbekistan. Journal of Vertebrate Paleontology. 24(3).
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.

"Kitadanisaurus" Lambert, 1990
Barremian, 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 (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.

undescribed 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.

Dromaeosauridae indet. (Nelson and Crooks, 1987)
Barremian-Late Albian, Early Cretaceous
Cedar Mountain Formation, Utah, US
Material- teeth
Comments- Currie et al. (1990) said that troodontid teeth reported from the Cedar Mountain Formation by Nelson and Crooks (1987) were more likely velociraptorines because of the serrations are small (12/mm) and elongate.
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, Grand Junction. 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, New York. pp. 107-125.

undescribed Dromaeosauridae (Kirkland et al., 1997)
Early Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- teeth
pes (Kirkland, online)
Comments- Kirkland et al. (1997; 1999) referred teeth to cf. Deinonychus sp., while Kirkland (online) noted a pes is known. This is possible, but cannot be proven until they are described.
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, Cifelli, Britt, Burge, DeCourten, Eaton and Parrish, 1999. Distribution of vertebrate faunas in the Cedar Mountain Formation, east-central Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 201-218.
https://scientists.dmns.org/sites/kencarpenter/Cedar%20Mountain%20storage/Dinosaurs.aspx

undescribed Dromaeosauridae (Chapman, Deck, Varricchio and Jackson, 2004)
Albian, Early Cretaceous
Wayan Formation, Idaho, US
Comments- Stated to be small to medium-sized.
Reference- Chapman, Deck, Varricchio and Jackson, 2004. Creaceous Wayan Formation of Idaho: A preliminary report. 24(3).

unnamed Dromaeosauridae (Kirkland et al., 1997)
Late Albian, Early Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71599) two teeth (Fiorillo, 1999)
teeth (Kirkland et al., 1997)
partial skeleton (Cifelli et al., 1999)
Comments- This 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) note 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.
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.
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.

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 (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.

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.

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

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.

probable Dromaeosauridae indet. (Currie, Rigby and Sloan, 1990)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith 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 Judith 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, Paronychodon, 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.

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)
(UCMP 149171) limb elements (UCMP online)
Comments- The teeth were listed as Velociraptorinae by Parrish (1999).
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.

Dromaeosauridae indet. (Hall, 1991)
Late Campanian, Late Cretaceous
Fruitland Formation, New Mexico, US
Material- 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.
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. Unpublished Masters Thesis. Lawrence, 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.

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.

unnamed Dromaeosauridae (Carpenter, 1982)
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 124983) tooth (2.1 mm) (Carpenter, 1982)
(UCMP 124984) tooth (3 mm) (Carpenter, 1982)
(UCMP 124985) tooth (2.7 mm) (Carpenter, 1982)
(UCMP 124989) tooth (UCMP online)
(UCMP 125238) dentary fragment (Carpenter, 1982)
(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)
material (Triebold and Russell, 1995)
material (Jacobsen and Sroka, 1998)
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 basal dromaeosaurids.
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.
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.

Dromaeosauridae indet. (Kirkland, Lucas and Estep, 1998)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Kirtland Formation, New Mexico, US
Material- Kirkland et al. (1998) list Dromaeosauridae indet. from the Naashoibito Member, but dromaeosaurids are not mentioned by Williamson in his list of dinosaur teet from that level.
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.
Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.

unnamed dromaeosaurid (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- teeth (Baszio, 1997)
Comments- Baszio (1997) reported this was similar to teeth from the Lance Formation, perhaps indicating they are the same species. He referred it to Saurornitholestes, but as with the Lance material, this is considered unlikely.
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.

unnamed Dromaeosauridae (Ostrom, 1969)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- (AMNH 22670) fourteen teeth (AMNH online)
(UA BTB 123, 129-132, 161-164) nine teeth (Baszio, 1997)
(UCM 39502) tooth (1.7 mm) (Carpenter, 1982)
(UCM 45055) tooth (2.6 mm) (Carpenter, 1982)
(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)
(UW 13684) dentary fragment (Carpenter, 1982)
(YPM PU 20589) proximal pedal phalanx II-2 (Ostrom, 1969)
(YPM PU 55007) (YPM online)
(YPM PU 55502) (YPM online)
(YPM PU 55522) (YPM online)
(YPM PU 55525) (YPM online)
?(YPM PU 55535) (YPM online)
(YPM PU 55560) (YPM online)
(YPM PU 55561) (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.
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. As Atrociraptor has teeth nearly identical to Saurornitholestes though, it is likely the Lance material will be placed in a new genus once more material is recovered.
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.
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.

unnamed dromaeosaurid (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
Material- (RTMP 87.16.18) tooth (Ryan and Russell, 2001)
(UA JLE 64:109) tooth (Vaszio, 1997)
teeth (Ryan and Russell, 2001)
Comments- Baszio (1997) reported this was similar to teeth from the Lance Formation, perhaps indicating they are the same species. He referred it to Saurornitholestes, but as with the Lance material, this is considered unlikely.
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.

unnamed possible Dromaeosauridae (Franco-Rosas, 2002)
Turonian-Late Maastrichtian, Late Cretaceous
Adamantina and Marilia 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.

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 dromaeosaurid (van der Lubbe, Richter and Knotschke, 2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry, Germany
Material- six teeth
Reference- van der Lubbe, Richter and Knotschke, 2009. Velociraptorine dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany. Acta Palaeontologica Polonica. in press.

unnamed Dromaeosauridae (Sweetman, 2004)
Barremian, Early Cretaceous
Wessex Formation, England
Material- (BMNH R 16510) lateral tooth (>21.5 mm)
(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
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.

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.

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 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 chaînon de Saint-Chinian. Bull. Soc. Géol. France. 27, 686-709.
Lapparent, 1947. Les dinosauriens du Crétacé supérieur du midi de la France. Mémoire 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 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.

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 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 (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 (Nessov, 1995)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- (CCMGE 461/12457) tooth
(juvenile) maxilla
teeth
Comments- These remains were identified as Deinonychosauria and Dromaeosauridae by Nessov (1995). The maxilla is described as shorter and wider than Deinonychus, while the teeth were described as small. They may belong to the Bissekty dromaeosaurine mentioned by Averianov (2007), which may be Itemirus. A partial braincase referred to as possibly dromaeosaurid (CCMGE 466/12457) was referred to Urbacodon sp. by Averianov and Sues (2007).
Reference- 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.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.
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 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, St. Petersburg State University, St. Petersburg, Russia. 156 pp. + 14 pl. [in Russian with short English, German, and French abstracts].

undescribed Dromaeosauridae (Nessov, Kaznyshkina and Cherepanov, 1987)
Early Santonian, Late Cretaceous
Yalovach Formation, Tajikistan
Material- (PIN 3041/11) anterior dorsal vertebra (Alifanov and Averianov, 2006)
teeth (Nessov et al., 1987)
Comments- The dorsal vertebra is described as relatively short with stout hypapophyses.
References- Nessov, Kaznyshkina and Cherepanov, 1987. [Dinosaurs, crocodiles, and other archosaurs of the late Mesozoic of Middle Asia and their place in ecosystems]. Tyezisy dokladov XXXIII syessii Vsyesoyuznogo palyeontologichyeskogo obshchyestva, Lyeningrad. 46-47.
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.

undescribed dromaeosaurid (Norell et al., 2006)
Campanian, Late Cretaceous
Djadokhta Formation, Mongolia
Holotype- (IGM coll.) material including frontal
Comments- Norell et al. (2006) mention an undescribed dromaeosaurid species from the Zos Wash locality in the Djadokhta Formation that differs from Tsaagan in frontal morphology.
Reference- Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.

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 Dromaeosauridae (Gilmore, 1933)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (AMNH 6572) pedal phalanx II-1 (Ostrom, 1969)
?(AMNH coll.) pedal elements, skeletal fragments (Gilmore, 1933) 6376, 6556, 6744, 6756, 6757, 21552, 21565, 21588, 21597, 21774, 21775, 21776, 21780-21782, 21784
?(IVPP 270790-4) tooth (21 mm) (Currie and Zhao, 1993)
teeth, unguals (Currie and Eberth, 1993)
Comments- Gilmore (1933) noted the presence of small theropod pedal elements and other fragmentary remains from the Iren Dabasu Formation. He believed that they were probably from dromaeosaurids or troodontids but admitted that this is based on temporal evidence. Ostrom (1969) described a pedal phalanx from the Iren Dabasu Formation as being almost exactly identical with that of Deinonychus but about 20% larger. Currie and Zhao (1993) illustrate a dromaeosaurid tooth from the Iren Dabasu, but given the slight constriction below the crown, convex distal edge and seeming lack of serrations, this may be misidentified.
Currie and Eberth (1993) state isolated dromaeosaurid elements are common in the formation, and that most of them can be referred to Velociraptor (except for larger, dromaeosaurine teeth). Yet no rationale was presented, and the only two specifically identified Iren Dabasu dromaeosaurid elements in the literature are clearly not Velociraptor (AMNH 6572 is far too large, while IVPP 270790-4 is different from most dromaeosaurid teeth as noted above). AMNH 6572 may belong to the dromaeosaurine though, based on size.
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.
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. A new troodontid (Dinosauria, Theropoda) braincase from the Dinosaur Park Formation (Campanian) of Alberta. Canadian Journal of Earth Sciences. 30(10-11), 2234-2247.

undescribed Dromaeosauridae (Sidleir, 1998)
Cenomanian, Late Cretaceous
Kem Kem Beds, Morocco
Material- (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)
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- Sidleir, 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.

undescribed Dromaeosauridae (Sidleir, 1998)
Cenomanian, Late Cretaceous
Kem Kem Beds, Morocco
Material- (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)
Comments- Sadleir (1998) reported dromaeosaurids may be represented by the small teeth in his collection of over 100 from the Kem Kem Formation.
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- Sidleir, 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.

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.

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 (Currie, Vickers-Rich and Rich, 1996)
Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P186343) (small) tooth
Comments- Currie et al. (1996) noted this was dromaeosaurid-like. It is possibly the specimen photographed on Pigdon's website, which is highly recurved, with only distal serrations (2.5-3/mm). Besides basal dromaeosaurids, basal coelurosaurs can also have this kind of tooth. They may belong to Timimus, if the latter is indeed an unenlagiine.
References- Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia. Alcheringa. 20(1-2), 73-79.
Rich and Vickers-Rich, 1997. Future directions for dinosaur research in Australia. 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. 275-277.
Long, 1998. Dinosaurs of Australia and New Zealand and other animals of the Mesozoic Era. Harvard University Press. 192 pp.
http://home.alphalink.com.au/~dannj/austdino.htm

undescribed possible Dromaeosauridae (Rich and Vickers-Rich, 1997)
Early Aptian, Early Cretaceous
Wonthoggi Formation of the Strzelecki Group, Victoria, Australia
Material- ten teeth (Kool, 2000)
Comments- Kool (2000) reported ten small theropod teeth found in the 2000 Dinosaur Dreaming field season. All were reported to have mesial serrations and lack distal ones, but this was probably a typo for the opposite condition. Some of the over 100 reported small theropod teeth from the Strzelecki Formation (Rich, 2004) probably belong to the same taxon. Besides basal dromaeosaurids, basal coelurosaurs and troodontids can have teeth with only distal serrations.
References- Rich and Vickers-Rich, 1997. Future directions for dinosaur research in Australia. 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. 275-277.
http://home.alphalink.com.au/~dannj/alozdino.htm
Kool, 2000. Summary of Dinosaur Dreaming 2000.
Rich, 2004. Research Update. Dinosaur Dreaming 2004 Field Report. 7.

unnamed dromaeosaurid (Case, Martin and Caguero, 2007)
Early Maastrichtian, Late Cretaceous
Cape Lamb Member of the Snow Hill Island Formation, Antarctica
Material- two teeth, distal tibiae, distal fibula, astragalus, calcaneum, partial metatarsals II, phalanx II-1, partial pedal ungual II, partial metatarsals III, phalanx III-1, partial metatarsal IV, three pedal phalanx fragments
Reference- Case, Martin and Reguero, 2007. A dromaeosaur from the Maastrichtian of James Ross Island and the Late Cretaceous Antarctic dinosaur fauna. in Cooper and Raymond (eds). Antarctica: A Keystone in a Changing World – Online Proceedings of the 10th ISAES X. USGS Open-File Report 2007-1047, Short Research Paper 083, 4 p.; doi:10.3133/of2007-1047.srp083

undescribed dromaeosaurid (Anonymous, 2009)
Early Cretaceous
Japan
Material- (FPDM coll.) (~1.7 m) premaxilla, maxilla, jugal, braincase, teeth, cervical vertebra, cervical ribs, dorsal vertebra, dorsal ribs, gastralia, sacrum, thirteen caudal vertebrae, five chevrons, proximal humerus, radius, proximal ulna, phalanx I-1, phalanx II-1, phalanx II-2, , femora, tibiae, fibula, metatarsal I, metatarsus, phalanges II-1, phalanges II-2, pedal ungual II, phalanges III-2, phalanges III-3, phalanx IV-1, phalanx IV-2, phalanges IV-3, phalanx IV-4
Comments- This partial skeleton was announced in March, 2009 as a new though unnamed taxon of dromaeosaurid. Preperation should be complete in 2009, and the description will no doubt follow. The braincase has been examined via CT scan, and the translation seems to indicate the teeth are serrationless. It is said to be related to Sinornithosaurus, so may be microraptorian.
Reference- Anonymous, 2009. [3rd new species? Small-sized meat diet dinosaur to restoration Fukui] msn.com 3/18/2009 http://sankei.jp.msn.com/science/science/090318/scn0903182113002-n1.htm

Microraptoria Senter et al., 2004
Definition- (Microraptor zhaoianus <- Velociraptor mongoliensis, Dromaeosaurus albertensis) (modified from Senter et al., 2004)
= "Microraptorinae" Sereno, in press
Definition- (Microraptor zhaoianus <- Velociraptor mongoliensis, Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus) (Sereno, in press)
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 mispelling of their own (Headden, pers. comm. to Makovicky, 2005). Sereno (in press) erected "Microraptorinae", attributing it to Senter et al. (though I don't believe the ICZN allows 'potential' names to be attributed in that way), giving it 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 Sereno'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.

Bambiraptor Burnham, Derstler, Currie, Bakker, Zhou and Ostrom, 2000
B. feinbergi Burnham, Derstler, Currie, Bakker, Zhou and Ostrom, 2000
= "Linsterosaurus" DML, 1997
= Bambiraptor feinbergorum Norell and Makovicky, 2004
Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
Holotype- (AMNH 30554; = 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), radiales, 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 (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. This opinion seems largely due to distaste for the name Bambiraptor combined with stratigraphy however, and has never been defended with shared characters. In fact, Bambiraptor has always claded away from Saurornitholestes in published phylogenetic analyses (Currie and Varricchio, 2004; Senter et al., 2004; Norell et al., 2006; Senter, 2007). This might be explainable by ontogenetic differences, as the Bambiraptor holotype is very young and the adult paratypes have never been described. The description of these paratypes, additional juvenile dromaeosaurids and more complete Saurornitholestes specimens should help resolve matters.
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.
References- Burnham, Derstler and Linster, 1997. A new specimen of Velociraptor (Dinosauria: Theropoda) from the Two Medicine Formation of Montana. Dinofest International Proceedings. 73-75.
http://dml.cmnh.org/1997Jun/msg00518.html
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.
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.
http://dml.cmnh.org/2000Mar/msg00342.html
http://dml.cmnh.org/2000Mar/msg00361.html
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, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History 8: 1-20.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
Senter, 2006. Comparison of forelimb function between Deinonychus and Bambiraptor (Theropoda: Dromaeosauridae). Journal of Vertebrate Paleontology. 26(4), 897-906.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.

unnamed clade

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
Hoovor (=Khovboor, Khoobur, etc.), 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.

undescribed Microraptoria
Early Cretaceous
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
(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.
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 ornithurines 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-http://dml.cmnh.org/2000Jun/msg00268.html
http://dml.cmnh.org/2002Dec/msg00303.html

Graciliraptor Xu and Wang, 2004
G. lujiatunensis Xu and Wang, 2004
Hauterivian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V13474) (~1 m; adult) fragmentary maxilla, teeth, ten caudal vertebrae (33 mm), chevrons, humerus (~110 mm), radius, ulna (128 mm), radiale, ulnare, semilunate carpal, metacarpal I (19 mm), phalanx I-1 (40 mm), manual ungual I (~30 mm), metacarpal II (59 mm), phalanx II-1 (~27 mm), phalanx II-2 (27 mm), proximal manual ungual II, metacarpal III (54 mm), phalanx III-1 (11.5 mm), phalanx III-2 (5.9 mm), phalanx III-3 (18 mm), manual ungual III, partial femur (~130 mm), incomplete tibiae (~200 mm), distal fibula, astragalus, calcaneum, distal metatarsal II, phalanx II-1 (12.7 mm), phalanx II-2 (13.6 mm), proximal pedal ungual II, distal metatarsal III, phalanx III-1 (21 mm), phalanx III-2 (13 mm), distal metatarsal IV
Diagnosis- (after Xu and Wang, 2004) laminal structure connects mid caudal postzygopophyses and extends over an eighth of the succeeding centrum; mid caudals with centrum length/width ratio of 8.6; manual ungual I much smaller than manual ungual II; proximal end of metacarpal III strongly expanded dorsoventrally; tibiotarsus with length / midshaft width ratio of 28; tibial shaft proximally rectangular in cross section; astragalar medial condyle significantly expanded posteriorly; metatarsal II 1.5 times wider than metatarsal III distally; pedal phalanx III-1 long and slender.
elongate ulna (>105% of humeral length).
Reference- Xu and Wang, 2004. A new dromaeosaur (Dinosauria: Theropoda) from the Early Cretaceous Yixian Formation of Western Liaoning. Vertebrata PalAsiatica. 42(2), 111-119.

"Paleopteryx" Jensen, 1981
"P. thomsoni" Jensen, 1981
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US
Holotype- (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. 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.
Relationships- "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.

Sinornithosaurus Xu, Wang and Wu, 1999
= "Sinavisaurus" Skrepnick, DML 2000
Diagnosis- (modified from Xu et al., 1999) antorbital fossa with extensive pitting; elongate posterolateral parietal process. (modified from Xu and Wu, 2001) large promaxillary fenestra.
larger humerus than Microraptor+Cryptovolans or Bambiraptor (humerofemoral ratio >88%).
Comments- Skrepnick (DML, 2000) noted that Sinornithosaurus had the working name of "Sinavisaurus" until right before it was published.
Xu et al. (1999) listed additional synapomorphies of Sinornithosaurus. However, the elongate posterodorsal dentary process, enlarged supracoracoid fenestra, lateral tubercle on the pubic midshaft, and proximodorsal ischial process are also present in Microraptor; 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. These cannot be used to place S. haoiana in Sinornithosaurus, contra Liu et al. (2004).
Ji et al. (2002) referred NGMC 91 to Sinornithosaurus sp., but it is here referred to a new genus more closely related to Microraptor.
References- Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
http://dml.cmnh.org/2000Oct/msg00074.html
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, 6 figs. [phot.]; Beijing.
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.
S. millenii Xu, Wang and Wu, 1999
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Member of the Yixian Formation, Liaoning, China
Holotype- (IVPP V12811) (~1.05 m) skull (143 mm), scleral plates, mandibles, hyoid, cervical vertebrae, dorsal vertebrae (26 mm), dorsal ribs, gastralia, sacrum (65 mm), three distal caudal vertebrae, scapulae (85 mm), coracoids (44 mm), furcula, sternal plates (84 mm), humeri (134 mm), radius, ulna (110 mm), radiale, ulnare, semilunate carpal, metacarpal I, phalanx I-1 (40.7 mm), manual ungual I, metacarpal II (63 mm), phalanx II-1 (32.2 mm), phalanx II-2 (36.2 mm), manual ungual II, metacarpal III, phalanx III-1 (18.1 mm), phalanx III-2 (6.9 mm), phalanx III-3 (23.9 mm), manual ungual III, ilia (85 mm), pubes (116 mm), ischia (52 mm), femora (148 mm), 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), metatarsal III (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- (IG-3) skull, postcranial skeleton (Azuma, 2005)
(IG-5b) skull postcranial skeleton, feathers (Azuma, 2005)
Diagnosis- (modified from Xu and Wu, 2001) groove posterior to the anterior carina on the lingual surface of the premaxillary tooth crowns (unknown in S. haoiana); diastema between the premaxillary and maxillary teeth (unknown in S. haoiana); column-like margin of the pterygoid process of the quadrate (unknown in S. haoiana); large excavation on the posterolateral surface of the parasphenoid process (unknown in S. haoiana).
(modified from Liu et al., 2004) low premaxillary body (~10% of orbit+jugal height); maxillary fenestra large (~15% of orbit+jugal height) and elongate (height/length ratio ~65%); reduced dorsal quadratojugal process (~12% of orbit+jugal height) with no squamosal contact; slender dentary (length/width ratio 13); pubic peduncle of ilium longitudinally longer than acetabulum.
Comments- The species name must be emended to millennii because it's a mispelling of a Latin word (millennium) (ICZN Art. 32.5).
Xu and Wu (2001) listed additional synapomorphies of Sinornithosaurus millenii (S. haoiana was still unpublished). However, the deep excavation on the posteroventral margin of the premaxilla is also present in Bambiraptor.
References- Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
Xu and Wang, 2000. Troodontid-like pes in the dromaeosaurid Sinornithosaurus. Paleont. Soc. 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.
White, 2009. The subarctometatarsus: intermediate metatarsus architecture demonstrating the evolution of the arctometatarsus and advanced agility in theropod dinosaurs. Alcheringa. 33(1), 1-21.
S. haoiana Liu, Ji, Tang and Gao, 2004
Late Valanginian-Early Aptian, Early Cretaceous
Yixian or Jiufotang Formation, Liaoning, China
Holotype- (D2140) (skull ~125 mm) premaxilla, maxilla, nasals, lacrimal, frontals, parietal, partial jugal, postorbital, squamosal, quadratojugal, quadrate, vomer, palatine, pterygoid, parasphenoid-basisphenoid, scleral 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, five proximal caudal vertebrae, several mid caudal vertebrae and chevrons, scapula (84.4 mm), coracoid, furcula, sternal plates (64.1 mm), four sternal ribs (to 42.6 mm), humeri (129 mm), radii, ulnae (105.3 mm), radiale, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II (59 mm), metacarpal III, phalanx III-1, phalanx III-2, ilia (84.5, 86.1 mm), pubis (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, pedal digit III, pedal ungual III, metatarsal IV, pedal digit IV, pedal ungual IV, metatarsal V, feathers
Diagnosis- (modified from Liu et al., 2004) high premaxillary body (~19% of orbit+jugal height); maxillary fenestra small (~11% of orbit+jugal height) and circular (height/length ratio ~92%); tall dorsal quadratojugal process (~41% of orbit+jugal height) with squamosal contact; tall dentary (length/width ratio 7.3); pubic peduncle of ilium longitudinally shorter than acetabulum.
Comments- The species name must be emended to haoianus, to match the gender of the genus name (ICZN Art. 31.2, 34.2).
Liu et al. (2004) listed an additional character to distinguish S. haoiana from S. millenii - the long subnarial process of the premaxilla which excludes the maxilla from the naris. However, this cannot be evaluated in S. millenii.
Reference- 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.
S. sp. (Kaiser, 2007)
Early Cretaceous
China
Material- ?(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 (http://www.geoscience-enterprises.com/geoscience_enterprises/Specimen_pages/D_CH_0001.htm)
(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 dihit 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
?(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)
Comments- 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 lomgate 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.
References- http://dml.cmnh.org/2000Jun/msg00268.html
http://dml.cmnh.org/2002Dec/msg00303.html
Kaiser, 2007. The Inner Bird: Anatomy and Evolution. UBC Press. 386 pp.

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.
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.
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.
R. gilmorei Currie, Rigby and Sloan, 1990
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith 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 81.16.194) premaxillary tooth
(RTMP 83.129.11) tooth
(RTMP 84.89.274) tooth
Referred- (RTMP 65.26.13) tooth (Baszio, 1997)
(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 96.29.2) tooth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
(RTMP 97.39.2) tooth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
(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 toooth (Sankey, 2001)
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). 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.
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 (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. 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 (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 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
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 (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 (Stokosa, 2005)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
(SDSM 64372b) tooth
Reference- Stokosa, 2005. Enamel microstructure variation within the Theropoda. in Carpenter (ed). The Carnivorous Dinosaurs. 163-178.
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, Standhardt and Schiebout, 2005)
(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)
Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
material (Lewis, Heckert and Forys, 2006)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(UA MR-4: 36, 37, 38, 39, 40, 41) six teeth (Bazsio, 1997)
teeth (Russell, 1935)
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 Judith River Group, Alberta, Canada
(RTMP coll.) teeth (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith 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)
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. 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.
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.
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.
R. cf. isosceles (Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997)
Late Albian, Early Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- 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. 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.
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)
Kimmeridgian, Late Jurassic
Guimarota Formation, Portugal
Material- (IPFUB 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 sp. nov. (Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke, 2002)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romanian
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) new comb.
= Asiamericana asiatica Nessov, 1995
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Paratypes- (N 460/12457) tooth
(N 12457 coll.) two teeth
Referred- several dozen teeth (Averianov, 2007)
teeth (Sues, pers. comm. 2007)
Diagnosis- differs from R. isosceles in that serrations are absent (?); differs from R. gilmorei in that no teeth constricted at their base, and the teeth are straight.
Comments- Nessov (1995) considered this taxon to be either a spinosaurid or a saurodontid fish. However, he noted resemblence between the holotype and a tooth described by Estes (1964), which was later referred to Richardoestesia isosceles by Sankey et al. (2002). Sues (pers. comm., 2007) confirms Asiamericana is indistinguishable from Richardoestesia, contra Buffetaut et al. (2008) who stated it was certainly a saurodontid. The holotype teeth are similar to R. isosceles in being straight and lacking a constricted base. They differ from both R. gilmorei and R. isosceles in lacking serrations, though this may be due to wear or positional variation. Ryan (1997) mentions Richardoestesia teeth from this formation, which may belong to the same species. Sues (pers. comm., 2007) notes he has examined "scores of similar teeth", while Averianov (2007) mentions several dozen Richardoestesia teeth are known.
References- Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming. Univ. California Publ. Geol. Sci. 49, 1-180.
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].
Ryan, 1997. Middle Asian Dinosaurs. In Encyclopedia of Dinosaurs (J.Currie and K.Padian, Eds.), pp.442-444 Academic Press, San Diego, California/London, UK.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of Paleontology: Vol. 76, No. 4, pp. 751-763.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.
Buffetaut, Suteethorn, Tong and Amiot, 2008. An Early Cretaceous spinosaurid theropod from southern China. Geological Magazine. 145(5), 745-748.
R? sp. (Prieto-Marquez, Gaete, Galobart and Ardevol, 2000)
Campanian, Late Cretaceous
Aren Sandstone Formation, Spain
Material- tooth
Reference- 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.
R. sp. (Averianov, 2007)
Middle Albian-Early Cenomanian, Early-Late Cretaceous
Khodzhakul Formation, Uzbekistan
Material- dozens of teeth
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.
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. sp. (Kirkland, Lucas and Estep, 1998)
Barremian, Early Cretaceous
Yellow Cat Member of Cedar Mountain Formation, Utah, US
Comments- Kirkland et al. (1998) list cf. Richardoestesia sp. under the Lower Cedar Mountain Formation, whgich corresponds to the Yellow Cat Member.
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.
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. (Girouard, 1997)
Campanian, Late Cretaceous
Trent River Formation, British Columbia, Canada
Material- (The Courtenay and District Museum coll.) tooth
Comments- This was digested, making it resemble Paronychodon teeth.
Reference- Girouard, 1997. First dinosaur remains from the Pacific coast of British Columbia, of the Trent River Formation (Campanian, Late Cretaceous) of Vancouver Island [abstract], in British Columbia Paleontological Symposium, May 9-11, 1997, Vancouver, B.C., Program and Abstracts, p.15.
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. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Foremost Formation of the Judith River Group, Alberta, Canada
(RTMP 88.86.44) tooth
(RTMP coll.) teeth
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, Bloomington, Indiana. pp. 279-297.
R. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Judith 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. (Williamson, 2001)
Late Campanian, Late Cretaceous
Fruitland and/or Lower Kirtland Formation, New Mexico, US
Reference- Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
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. (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. (Wel and Williamson, 2000)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Kirtland Formation, New Mexico, US
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.
R. sp. (Bertini and Franco-Rosas, 2001)
Turonian-Late Maastrichtian, Late Cretaceous
Adamantina and Marilia 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.
R. sp. indet.
Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- two teeth (Mongelli and Varricchio, 1998)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- ?(UCMP 120849) over fifty teeth (UCMP online)
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
Material- (Varricchio, 1995)
Maastrichtian, Late Cretaceous
Fox Hills Formation, South Dakota, US
Material- (SDSM 14516) tooth (Stokosa, 2005)
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)
(Triebold, 1997)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- ?(UCMP 119674) tooth (UCMP online)
Late Cretaceous
US
Material- (YPM 56979) (YPM online)
Comments- These reports could be either cf. gilmorei or cf. isosceles. 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- 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.
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.
Mongelli and Varricchio, 1998. Theropod teeth of the Lower Two Medicine Formation (Campanian) of Northwestern Montana. Journal of Vertebrate Paleontology. 18(3), 64A.
Stokosa, 2005. Enamel microstructure variation within the Theropoda. in Carpenter (ed). The Carnivorous Dinosaurs. 163-178.

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, coracoids, furcula, sternal plate, sternal ribs, humeri (87 mm), ulnae (72 mm), radii, radiale, 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.
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, 6 figs. [phot.]; Beijing.
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. Journal of Systematic Palaeontology.

Microraptor Xu, Zhou and Wang, 2000
= "Archaeoraptor" Sloan, 1999 in part
= Cryptovolans Czerkas, Zhang, Li and Li, 2002
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
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (IVPP V 12330; specimen of "Archaeoraptor liaoningensis" in part) (~420 mm; 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), radiale, ulnare, 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
Referred- (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) incomplete skeleton including femur (61 mm), remiges and leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
(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) 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), radiale, ulnare, semilunate carpal, 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 V13476) (~745 mm) specimen including femur (~94 mm) and leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
(IVPP V13477) (~572 mm) specimen including femur (72 mm), tibiotarsus, metatarsus, pedal digit II, pedal digit III, pedal ungual III, pedal digit IV, pedal ungual IV, metatarsal V and leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
(JI-5) skull postcranial skeleton, feathers (Azuma, 2005)
(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)
(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), radiale, ulnare, semilunate carpal, distal carpal 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)
(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)
Early Cretaceous
Qianyang, Liaoning, China
Referred- (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 II (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
?(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- (modified from 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.
(modified from Xu et al., 2003) manual phalanx III-1 >83% of phalanx III-3 in length.
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).
Comments- 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.
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 an ornithuromorph 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 ornithuromorph 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). There is a prominent biceps tubercle on the proximal radius, which Senter et al. considered individual variation. They 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. 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. 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..
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%).
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%).
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%).
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.
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 CAGS 20-80-001 and NGMC 00-12-A based on their fused sterna, we could equally as well argue the Cryptovolans specimens 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. 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. The more conservative approach taken by Senter et al. 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.
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.
References- Sloan, 1999. Feathers for T. rex?. National Geographic. 196(5), 98-107.
http://dml.cmnh.org/2000Dec/msg00720.html
Olson, 2000. Countdown to Piltdown at National Geographic: the rise and fall of Archaeoraptor. Backbone, newsletter of the Department of Vertebrate Zoology, National Museum of Natural History. 13(2), 1-3.
Stoksad, 2000. Tiny, feathered dino is most birdlike yet. Science. 290(5498), 1871-1872
Xu, Zhou and Wang, 2000. The smallest known non-avian theropod dinosaur. Nature, 408, 705-708.
http://dml.cmnh.org/2001Jan/msg00092.html
Rowe, Ketcham, Deinson, Colbert, Xu and Currie, 2001. The Archaeoraptor forgery. Nature. 410, 539-540.
Czerkas and Ji, 2002. A preliminary report on an omnivorous volant bird from Northeast China. in Czerkas (ed.). Feathered Dinosaurs and the Origin of Flight. 127-135.
Czerkas and Xu, 2002. A new toothed bird from China. Feathered Dinosaurs and the Origin of Flight. 43-61.
Czerkas, Zhang, Li and Li, 2002. Flying dromaeosaurs. in Czerkas (ed.). Feathered Dinosaurs and the Origin of Flight. 97-126.
Hwang, Norell, Qiang and Keqin, 2002. New specimens of Microraptor zhaoianus (Theropoda: Dromaeosauridae) from northeastern China. American Museum Novitates. 3381, 1-44.
Ji, 2002. New data of Early Cretaceous dromaeosaurs from Western Liaoning with comments on the origin of feathers. Unpublished Thesis. 94 pp.
Norell, Ji, Gao, Yuan, Zhao and Wang 2002. 'Modern' feathers on a non-avian dinosaur. Nature. 416, 36-37.
Zhou, Clarke and Zhang, 2002. Archaeoraptor's better half. Nature. 420, 253-344.
http://dml.cmnh.org/2003Jan/msg00416.html
Xu, Zhou, Wang, Kuang, Zhang and Du, 2003. Four-winged dinosaurs from China. Nature. 421, 335-340.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History 8: 1-20.
Xu, Zhou, Zhang, Wang and Kuang, 2004. Functional hind-wings conform to the hip structure in dromaeosaurids. Journal of Vertebrate Paleontology. 24(3).
Azuma, 2005. The Flying Dinosaurs: Fukui Prefectural Dinosaur Museum. 118 pp.
Padian and Dial, 2005. Could 'four-winged' dinosaurs fly? Nature. 438.
Xu, Zhou, Wang, Kuang, Zhang and Du, 2005. Xu et al. reply. Nature. 438.
Zhou and Zhang, 2005. Zhang & Zhou reply. Nature. 438.
Ji, Ji, Lu, You and Yuan, 2006. Embryos of Early Cretaceous Choristodera (Reptilia) from the Jehol Biota in Western Liaoning, China. Journal of the Paleontological Society of Korea. 22(1), 111-118.
Chatterjee and Templin, 2007. Biplane wing planform and flight performance of the feathered dinosaur Microraptor gui. Proceedings of the National Academy of Sciences of the United States of America. 104(5), 1576-1580.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
White, 2009. The subarctometatarsus: intermediate metatarsus architecture demonstrating the evolution of the arctometatarsus and advanced agility in theropod dinosaurs. Alcheringa. 33(1), 1-21.

Hesperonychus Longrich and Currie, 2009
H. elizabethae Longrich and Currie, 2009
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith River Group, Alberta, Canada
Holotype- (UALVP 48778) (1.6 kg) partial ilia, pubes (95 mm)
Paratypes- (RTMP 1988.36.68) pedal ungual II
(RTMP 1989.116.65) pedal phalanx II-1 (14.2 mm)
(RTMP 1991.50.105) incomplete pedal ungual II
(RTMP 2000.12.100) incomplete pedal ungual II
(UALVP 50686; = UALVP 50048 in supplementary info?) pedal ungual II
(UALVP 50687; = UALVP 50049 in supplementary info?) incomplete pedal phalanx II-1
Late Campanian, Late Cretaceous
Oldman Formation of the Judith River Group, Alberta, Canada
Paratype- (RTMP 1995.92.9) pedal ungual II
Late Campanian, Late Cretaceous
Judith River Group, Alberta, Canada
Paratypes- (RTMP 1966.19.22) incomplete pedal phalanx II-1
(RTMP 1979.10.6) pedal ungual II
(RTMP 1980.8.205) pedal ungual II
(RTMP 1980.16.1880) pedal ungual II
(RTMP 1983.67.7) pedal phalanx II-2 (17.1 mm)
(RTMP 1988.50.34) pedal ungual II
(RTMP 1990.107.15) pedal ungual II
(RTMP 1991.50.143) pedal phalanx II-2 (4.8 mm)
(RTMP 1992.36.61; = RTMP 1992.36.31 in supplementary info) pedal phalanx II-2 (11.1 mm)
Diagnosis- (after Longrich and Currie, 2009) pubic peduncle of ilium with medial surface deeply excavated; postacetabular process of ilium with medial shelf split to form anterior and posterior processes; lateral tubercles of pubis wing-like and curving anteriorly; pubis with fossa on lateral surface ventral to acetabulum; pubic apron shifted onto posterior surface of pubis; pubic symphysis teardrop-shaped in lateral view; ischil peduncle of pubis reduced to a narrow lamina.
Comments- Several of the paratype specimens are only listed in the electronic supplementary material, not the hypodigm of the article.
Reference- Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae) from the Late Cretaceous of North America. PNAS Early Edition. www.pnas.org/cgi/doi/10.1073/pnas.0811664106

Austroraptor Novas, Pol, Canale, Porfiri and Calvo, 2008
A. cabazai Novas, Pol, Canale, Porfiri and Calvo, 2008
Campanian-Maastrichtian, Late Cretaceous
Allen Formation, Patagonia, Argentina
Holotype- (MML-195) (~4.9 m; 368 kg) (skull ~800 mm) incomplete maxillae, lacrimals, frontal, postorbital, incomplete dentaries, surangular, prearticular, third cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, second dorsal vertebra, fourth dorsal vertebra, ribs, gastralia, humerus (262 mm), manual ungual III, partial pubis, incomplete femur (~560 mm), tibia (565 mm), astragalus, calcaneum, pedal ungual I, phalanx II-2 (57 mm), metatarsal III (330 mm), pedal ungual III, phalanx IV-2 (48 mm)
Diagnosis- (modified from Novas et al., 2008) 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.
Comments- Note "Austroraptor" was earlier used as an unpublished name for Ozraptor (Pigdon, DML 1997).
Novas et al. (2008) also included the small tooth size and lack of serrations as apomorphies, but these are probably basal for paravians. They assigned the taxon to Unenlagiinae, which was sister to Microraptoria in their cladogram (based on Turner et al.'s 2007 matrix). Including the taxon in a modified version of Senter's (2007) matrix with all of Turner et al.'s data and numerous additional taxa and characters results in a similar placement - Austroraptor, Neuquenraptor and Shanag still fall out as relatives within Microraptoria. However, Unenlagia is more basal within Dromaeosauridae while Buitreraptor and Rahonavis are avialans.
References- 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), 98A.
Novas, Pol, Canale, Porfiri and Calvo, 2008. A bizarre Cretaceous theropod dinosaur from Patagonia and the evolution of Gondwanan dromaeosaurids. Proceedings of the Royal Society B. doi:10.1098/rspb.2008.1554

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, Argentina
Holotype- (MCF PVPH 77) (~2 m) cervical fragments, dorsal ribs, chevrons, proximal radius, incomplete femur, distal tibia, distal fibula, incomplete astragalus, calcaneum, metatarsal I, phalanx I-1, pedal ungual I, incomplete metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, incomplete metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, incomplete metatarsal IV, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
Diagnosis- (modified from Novas and Pol, 2005) metatarsal II with lateral expansion over the caudal surface of metatarsal III; extensor sulcus on proximal half of metatarsus; distal end of metatarsal III is incipiently ginglymoid (to a lesser degree than other dromaeosaurs).
Comments- This was originally reported as a troodontid until its official description identified it as a dromaeosaur. Novas (pers. comm., 2005) confirms that it is the same specimen as "Araucanoraptor". 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. 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.
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 (La Plata). Ameghiniana. 36 (S), p. 17R.
Novas and Pol, 2005. New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia. Nature. 433, 858-861.

Shanag Turner, Hwang and Norell, 2007
S. ashile Turner, Hwang and Norell, 2007
Berriasian-Barremian, Early Cretaceous
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.
Reference- Turner, Hwang and Norell, 2007. A small derived theropod from Oosh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates. 3557, 27 pp.

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)

Adasaurus Barsbold, 1983
= "Adasaurus" Barsbold, 1977
A. mongoliensis Barsbold, 1983
= Dromaeosaurus mongoliensis (Barsbold, 1983) Paul, 1988
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (IGM 100/20) (2.39 m) (old adult) posterior skull, posterior mandible, eight cervical vertebrae, eleven dorsal vertebrae, sacrum, seven caudal vertebrae, chevrons, scapula, coracoid, sternal plate, ilium (202 mm), pubis (241 mm), ischium (118 mm), femora (273 mm), tibiae, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1 (23 mm), phalanx II-2 (14 mm), pedal ungual II (29 mm), metatarsal III, pedal digit III, metatarsal IV, pedal digit IV, metatarsal V
Paratype- (IGM 100/51) posterior postcranial skeleton including hindlimbs
Referred- (IGM 100/22) premaxilla, maxilla, dentary, humeri, ulna (Currie and Varricchio, 2004)
(IGM 100/23) premaxilla, maxilla, dentary (Currie and Varricchio, 2004)
Diagnosis- (after Barsbold, 1983) pedal ungual II reduced in size to be subequal in length to phalanx II-1.
preacetabular processes strongly divergent; preacetabular process strongly notched anteriorly; posterodorsal edge of postacetabular process very thick; distal ischium curved strongly posteriorly; metatarsal II reduced in width.
Comments- This genus was 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. It was later described extremely briefly by Barsbold (1983) and officially named Adasaurus mongoliensis. A photograph of the mounted holotype skeleton is online
The holotype specimen was approximately 2.5 meters long if the ilium is scaled from Deinonychus. It was from an old individual, so this is probably close to the maximum size Adasaurus got. Because the specimen is pathologic (Norell and Makovicky, 1997), some features of its skeleton may not be representative of the species.
The published cranial information comes mostly from the data matrices of the Theropod Working Group (Norell et al., 2001 and variations), Senter et al. (2004) and Senter (2007). Most characters are like other dromaeosaurids. The lacrimal has an elongate posterior process, and there is no lacrimal horn or enlarged lacrimal foramen. The jugal is robust, as in non-microraptorian dromaeosaurids. The lateral temporal fenestra is reduced as in non-oviraptorid maniraptoriforms. The postorbital lacks an orbital process, and has a dorsally projected anterior process. The quadrate is vertical. The braincase lacks a lateral depression, and the paroccipital processes are elongate. In the mandible, the surangular lacks process intruding the mandibular fenestra, while the retroarticular process is short. There is no lateral surangular ridge, unlike Velociraptor, Saurornitholestes, Deinonychus and Dromaesaurus. Currie and Varricchio (2004) report there are four premaxillary teeth, eleven maxillary teeth and thirteen dentary teeth in IGM 100/23 and 100/22. They also state teeth have four serrations per mm in both mesial and distal carinae. The teeth are relatively tall, labiolingually flattened and recurved.
Vertebral information is exclusively from data matrices. Presacral vertebrae are not opisthocoelous. There are ten short cervical vertebrae, with large epipophyses on the axis, tall and axially short neural spines, and no carotid processes. The anterior dorsals have large hypapophyses and all dorsals have hyposphene-hypantra and lack fan-shaped neural spines. Dorsal centra are not short, unlike Deinonychus, Achillobator and Utahraptor. Posterior dorsal postzygapophyses which extend posterior to the centrum are unlike Utahraptor and Achillobator. The short dorsal neural spines are unlike Bambiraptor and more derived dromaeosaurids. There are five sacral vertebrae with a sinuous zygapophyseal ridge and no pleurocoels. The posterior centra are ventrally flattened and grooved, with a flat posterior articulation. Distal caudals have highly elongate prezygapphyses.
Pectoral information is also exclusively from data matrices. There is no marked distal scapular expansion. The glenoid is laterally directed, and the coracoid is subrectangular. Jasinowski et al. (2006) list a scapulocoracoid and sternal plate for the holotype, and humeri and an ulna for IGM 100/22 (which is only labeled 'dromaeosaur').
The pelvis was illustrated by Barsbold (1983) after the initial schematic drawing in Barsbold (1977). Adasaurus has a strongly opisthopubic pelvis as in most dromaeosaurids except Achillobator. The ilium is generally similar to other dromaeosaurids. Both ilia are separated widely from each other, but the preacetabular processes are divergent, unlike Sinornithosaurus and Velociraptor. The preacetabular process is notched anteriorly, as in Achillobator, but to a greater degree. Deinonychus has a very slightly concave anterior edge, while Bambiraptor's is convex. The anterior edge is also sloped strongly posteroventrally with an acute anterodorsal corner, as in Achillobator, Deinonychus and (probably) Sinornithosaurus, and unlike Bambiraptor. There is a shallow brevis fossa. The ventral margin of the pubic peduncle is slightly concave, as in Achillobator and Sinornithosaurus, as opposed to the pronounced notch present in Bambiraptor and Velociraptor. Adasaurus's ilium is also different from both Achillobator in that it was lower and lacks a prominant supratrochanteric process. Thus, Adasaurus resembles Bambiraptor, Deinonychus, Velociraptor and possibly Sinornithosaurus in these features. There is no supracetabular crest or brevis fossa, while the antitrochanter is well developed. The postacetabular process is not a gently curved blade not descending below the ischial peduncle like Bambiraptor, Deinonychus and Velociraptor. Instead, it has a slight concavity on it's posterior edge and lies below the ischial peduncle, as in Achillobator. The postacetabular process also lies below the ischial peduncle in Sinornithosaurus, although it appears to only have a very slight concavity if any.
The straight pubic shaft is unlike Jehol microraptorians. The pubic foot has a slight anterior component, smaller than in Deinonychus and Velociraptor, unlike the enlarged anterior pubic foot in Achillobator and the absent one in Bambiraptor, Sinornithosaurus and Microraptor. The posterior pubic foot is of modest proportions, larger than Microraptor, Sinornithosaurus and Velociraptor, but smaller than other dromaeosaurids.
Adasaurus lacks any posterior ischial processes, unlike all dromaeosaurids except Deinonychus and Velociraptor. Besides the apomorphically posteriorly curved distal end, the ischium is almost identical to Deinonychus, the only other differences being a slight dorsal curvature and slightly smaller obturator process. There is a longitudinal ischial ridge as in dromaeosaurids besides Utahraptor and Achillobator.
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. The anterior trochanter is separated from the greater trochanter by a deep cleft (contra Senter, 2007). There is no accessory trochanter. Adasaurus still has a fourth trochantor (Perle et al. 1999), like "Koreanosaurus", Achillobator, Velociraptor and some specimens of Deinonychus, as opposed to Bambiraptor and most eumaniraptorans, where it is absent. It is positioned more proximally than in Achillobator, as is Velociraptor's. As in other maniraptorans, the anterodistal femoral fossa of Adasaurus is absent. The tibia is longer than the femur, while the metatarsus is longer than 50% of femoral length. The latter is unlike Velociraptor, Deinonychus and Achillobator.
Distal tarsals are not fused to the metatarsus, unlike Microraptor and Velociraptor, but as in Sinornithosaurus, Deinonychus and Achillobator. Metatarsal I is reduced proximally as in non-therizinosauroid theropods. The hallux is neither reversed nor enlarged. The metatarsus is illustrated by Senter et al. (2004). It is unfused and non-arctometatarsalian, the latter unlike Jehol microraptorians. The metatarsus is also more robust than Jehol microraptorians, but less than dromaeosaurines. Metatarsals II and III are both ginglymoid. Barsbold (1983) noted that metatarsal II of Adasaurus was "considerably thinned". Thus, it may have been similar to Tochisaurus in this regard, while in opposition to noasaurids and avisaurids, which reduced their fourth digit. As for the pedal elements, only digit two has been figured or described (by Barsbold, 1983). Like other dromaeosaurids and troodontids, but unlike Rahonavis, phalanx II-2 has a large proximoventral heel participating in the phalangeal articulation. The second digit is often compared to Dromaeosaurus because of the reduced ungual, but the second ungual of Dromaeosaurus is unknown. We can however compare phalanges II-1 and II-2 of Deinonychus and Dromaeosaurus, then see how Adasaurus and other dromaeosaurids compare. Comparison indicates that phalanx II-1 is near identical in the two species. Phalanx II-2 however is much stouter in Dromaeosaurus than Deinonychus. The phalanges of Adasaurus and especially Achillobator are even more stout, while those of Bambiraptor, Pyroraptor, Saurornitholestes, Sinornithosaurus and Velociraptor are elongate. The claw of Adasaurus is reduced compared to most other dromaeosaurs. It also lacks a large flexor tubercle, unlike other dromaeosaurids. Metatarsal V is less than 50% of metatarsal IV's length, unlike microraptorians.
Relationships- Adasaurus was originally identified as a dromaeosaurid (Barsbold, 1977) and later specified to be a dromaeosaurine (Barsbold, 1983; Paul, 1988). The Theropod Working Group analyses (Norell et al., 2001 and variations) have always recovered Adasaurus as a dromaeosaurid, generally in a polytomy with the non-microraptorian taxa. Turner et al. (2007) and several other more recent variations have added many characters relevant to dromaeosaurids and found Adasaurus to be most closely related to Utahraptor, Dromaeosaurus and Achillobator. Senter et al. (2004) also found Adasaurus to be a dromaeosaurine, but a basal one, outside a clade formed of Saurornitholestes, Deinonychus, Achillobator, Utahraptor and Dromaeosaurus. Senter (2007) used a matrix combining the Theropod Working Group characters with his 2004 study and recovered Adasaurus to be one node more basal, outside the Velociraptorinae + Dromaeosaurinae clade. This is tentatively followed here. 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.
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.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian dinosaurs. pp. 49–67 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 Univ. Press.
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.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8, 1-20.
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.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
Turner, Hwang and Norell, 2007. A small derived theropod from Oosh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates. 3557, 27 pp.
Turner, Pol, Clarke, Ericson and Norell, 2007. A basal dromaeosaurid and size evolution preceding avian flight. Science. 317, 1378-1381.

Saurornitholestiinae Longrich and Currie, 2009
Definition- (Saurornitholestes langstoni <- Dromaeosaurus albertensis, Velociraptor mongoliensis, Microraptor zhaoianus) (Longrich and Currie, 2009)

Saurornitholestes Sues, 1978
S. langstoni Sues, 1978
= Laelaps explanatus Cope, 1876
= Laelaps laevifrons Cope, 1876
= Dryptosaurus explanatus (Cope, 1876) Hay, 1902
= Dryptosaurus laevifrons (Cope, 1876) Hay, 1902
= Deinodon explanatus (Cope, 1876) Lambe, 1902
= Deinodon laevifrons (Cope, 1876) Osborn, 1902
= Aublysodon explanatus (Cope, 1876) Hatcher, 1903
= Dromaeosaurus laevifrons (Cope, 1876) Matthew and Brown, 1922
= Dromaeosaurus explanatus (Cope, 1876) Kuhn, 1939
= Velociraptor langstoni (Sues, 1978) Paul, 1988
= Paronychodon explanatus (Cope, 1876) Olshevsky, 1995
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 Judith River Group, Alberta, Canada
Holotype- (RTMP 74.10.5) maxillary tooth (9.2 mm), quadrate, frontals, ectopterygoid, 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)
(RTMP 70.37.1) 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) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 80.16.312) frontal (Currie, 1987)
(RTMP 80.16.996) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.20.259) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.23.7) frontal (Currie, 1987)
(RTMP 82.16.43) tooth (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 82.24.16) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.26.1) incomplete caudal series (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 84.94.6) frontal (Currie, 1987)
(RTMP 85.43.69) frontal (Currie, 1987)
(RTMP 86.36.117) premaxilla (Currie, Rigby and Sloan, 1990)
?(RTMP 86.77.2) ischium (Hutchinson, 2001)
(RTMP 86.77.57) frontal (Currie, 1987)
(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 87.36.11) 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) 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 87.50.38) 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 88.121.39) (128 kg) cranial elements including quadratojugal, mandible (dentary 120 mm), teeth (to 9 mm), three dorsal ribs, gastralia, distal caudal series, scapula, coracoid, humerus, femur (214 mm), tibia (283 mm), fibula, metatarsus, pedal phalanges, several unguals (Currie, Rigby and Sloan, 1990; Currie and Varricchio in prep.)
(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 88.162.1) tooth (Baszio, 1997)
(RTMP 88.215.72) tooth (Baszio, 1997)
(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 92.36.129) sacrum (Rauhut, 2003)
(RTMP 92.36.333) sternal plate (Godfrey and Currie, 2004)
(RTMP 92.83.2) tooth (Currie and Jacobsen, 1995)
(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)
Late Campanian, Late Cretaceous
Oldman or Dinosaur Park Formation of the Judith River Group, Alberta, Canada
(BMNH R4463) sacrum (Howse and Milner, 1993)
?(CMN 12072) metatarsal IV (Colbert and Russell, 1969)
?(CMN 12240) pedal ungual II (Colbert and Russell, 1969)
(UA 12091) dentary (Sues, 1977)
(UA 12339) dentary (Sues, 1977)
tooth (12 mm) (Lambe, 1902)
Middle Campanian, Late Cretaceous
Foremost Formation, Alberta, Canada
(RTMP 88.86.29) tooth (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(AMNH 3958; holotype of Laelaps explanatus) tooth (11 mm), twenty-five teeth (Cope, 1876)
(AMNH 3961; holotype of Laelaps laevifrons) tooth (15 mm) (Cope, 1876)
(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)
(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)
Late Campanian, Late Cretaceous
Oldman Formation of the Judith River Group, Alberta, Canada
(RTMP 94.144.105) tooth (Ryan and Russell, 2001)
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, incomplete forelimb including radius, ulna and manual ungual II, ilium, hindlimb including femur (225 mm), incomplete pes (Britt, 1993)
(MOR 666) tibia (256 mm), fibula (225 mm) (Christiansen, 1999)
Late Cretaceous
North America
(CMN 12410) tooth
(RTMP 67.20.36) last dorsal vertebra, sacrum (Norell and Makovicky, 1997)
(RTMP 81.14.34) (Rauhut, 2003)
(RTMP 85.56.48) (adult) posterior cervical vertebra (Makovicky, 1995)
(RTMP 86.78.1) (Rauhut, 2003)
(RTMP 91.36.112) (Rauhut, 2003)
(RTMP 93.36.98) (Rauhut, 2003)
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), and Makovicky's (1995) as yet unpublished description of vertebrae, 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.
Laelaps explanatus- Cope (1876) 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 Judith 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 (1876) 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.
Early finds- 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 (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.
Russell (1969) listed CMN 12343, 12354 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 BMNH 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.
Recent discoveries- Howse and Milner (1993) illustrated a sacrum (BMNH 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.
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.
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.
References- Cope, 1876. Descriptions of some vertebrate remains from the Fort Union Beds of Montana. Paleontological Bulletin. 22, 1-14.
Cope, 1876. 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, 1876. 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 Catalogue 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.
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, 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, 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, New York. pp. 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.
Christiansen, 1999. Long bone scaling and limb posture in non-avian theropods: Evidence for differential allometry. Journal of Vertebrate Paleontology. 19(4), 666-680.
Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
Tanke and Currie, 2000. Head-biting behavior in theropod dinosaurs: Paleopathological evidence. GAIA. 15, 167-184.
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria, Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences. 38, 1753-1766.
Hutchinson, 2001. The evolution of pelvic osteology and soft tissues on the line to extant birds (Neornithes). Zoological Journal of the Linnean Society. 131, 123-168.
Jacobsen, 2001. Tooth-marked small theropod bone: An extremely rare trace. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana. pp.58-63.
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, 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.
Godfrey and Currie, 2004. A theropod (Dromaeosauridae, Dinosauria) sternal plate from the Dinosaur Park Formation (Campanian, Upper Cretaceous) of Alberta, Canada. In Currie, Koppelhus, Shugar and Wright (eds). Feathered Dragons: Studies on the Transition from Dinosaurs to Birds. 144-149.
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.
Turner, Pol, Clarke, Ericson and Norell, 2007. A basal dromaeosaurid and size evolution preceding avian flight. Science. 317, 1378-1381.
S. robustus Sullivan, 2006
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
Holotype- (SMP VP-1955) frontal (62 mm)
Paratypes- (SMP VP-1270) frontal (42.8 mm) (Sullivan and Lucas, 2000)
?(SMP VP-1741) partial pedal ungual II
?(SMP VP-1901) tooth (15 mm)
Diagnosis- (after Sullivan, 2006) ratio of the length (measured along the midline) to the thickness (posterior part of the frontal) is 6:1.
Comments- The separation of this species from S. langstoni rests entirely on the greater thickness of the frontals, which are both larger than the S. langstoni holotype. It is therefore possible this is an ontogenetic character, though one may expect a separate species due to the geographical distance.
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. pp. 253-256.
S? sp. (Kirkland, Lucas and Estep, 1998)
Late Santonian-Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
Comments- Kirkland et al. (1998) noted fragmentary remains of cf. Saurornitholestes sp. and Lewis et al. (2004) referred remains to Saurornitholestes sp..
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.
Lewis, Heckert, Lucas, Hunt and Hutchison, 2004. The vertebrate fauna and paleoecology of the Upper Cretaceous Menefee Formation (Late Santonian-Early Campanian), Northwestern New Mexico. Journal of Vertebrate Paleontology. 24(3).
S? sp. nov. (Baszio, 1997)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(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.
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.
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. (Williamson, 2001)
Late Campanian, Late Cretaceous
Fruitland and/or Lower Kirtland Formation, New Mexico, US
Material- teeth
Reference- Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
S? sp. (Sankey et al., 2005)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Material- (LSUMG 5659) tooth (2.8 mm) (Sankey, Standhardt and Shiebout, 2005)
(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, Standhardt and Shiebout, 2005)
(LSUMG 5980) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6132) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6139) tooth (~5.2 mm) (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6183) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6184) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6185) tooth (Sankey, Standhardt and Shiebout, 2005)
(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)
S. sp. nov. (Ryan and Russell, 2001)
Maastrichtian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Material- (RTMP 89.55.47) frontal
(RTMP 89.55.1523) tooth
teeth, elements
Comments- Ryan and Russell (2001) list the frontal as "Saurornitholestes undescribed n. sp. (Currie pers. comm.)".
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, Bloomington, Indiana. pp. 279-297.

Velociraptorinae Barsbold, 1978
Definition- (Velociraptor mongoliensis <- Dromaeosaurus albertensis) (modified from Sereno, 1998)
Other definitions- (Velociraptor mongoliensis <- Microraptor zhaoianus, Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus) (Sereno, in press)
= Velociraptorinae sensu Sereno, in press
Definition- (Velociraptor mongoliensis <- Microraptor zhaoianus, Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus)
Comments- Since Sereno (in press) uses a stem-based definition for Dromaeosauridae, the node stem triplet advocated by Padian et al. (1999) no longer functions. Sereno's new definition is slightly superior to his 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.

Velociraptor Osborn, 1924
= "Ovoraptor" Osborn, 1924
?= "Airakoraptor" Perle, Norell and Clark, 1999
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). Currie and Eberth (1993) referred remains from the Iren Dabasu Formation of Inner Mongolia, China to Velociraptor, but there is no published evidence for this, and published specimens differ from Velociraptor in some ways. 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).
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.
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. pp 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.
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. Journal of Systematic Palaeontology.
V. mongoliensis Osborn, 1924
= "Ovoraptor djadochtari" Osborn, 1924
Late Campanian, Late Cretaceous
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 on curve)
Paratype- (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)
Referred- (IGM 94.07.28) scapulocoracoid, sternal plates, humerus, radius, ulna (Jasinowski et al., 2006)
(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 veretebrae 1-10, cervical ribs, dorsal vertebrae 1-13, dorsal ribs, uncinate processes, gastralia, sacrum, caudal vertebrae 1-43, chevrons, scapula, coracoid, sternal plates (95 mm), humeri, radii, ulnae, manus, ilium, 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) (Barsbold, 1974)
(IGM 100/54) twelve dorsal vertebrae, sixteen dorsal ribs, five uncinate processes, gastralia, scapula, partial coracoids, sternal plates (Codd, 2004)
(IGM 100/200) premaxilla, maxilla, teeth (6.17-9.93 mm) (Smith, 2002)
(IGM 100/976) (adult) partial skull including braincase, incomplete mandible, atlas, axis, 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 100/982) (adult) incomplete skull including partial braincase, mandible, scapula, coracoid, partial furcula, partial sternal plates, proximal humeri, radii, ulnae, metacarpal I (19.2 mm), phalanx I-1 (39.9 mm), manual ungual I (39.9 mm curve), metacarpal II (50.8 mm), phalanx II-1 (31.3 mm), phalanx II-2 (45.8 mm), manual ungual II, 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 (32 mm curve), ilia (131.7, 124.6 mm), pubes (132 mm), femur (184 mm), astragalocalcaneum, metatarsal I (22.5 mm), phalanx I-1 (19.8, 19.3 mm), pedal ungual I, metatarsal II (73.7 mm), phalanx II-1 (23.9 mm), phalanx II-2 (26.1 mm), pedal ungual II (66.3 mm curve), metatarsal III, phalanx III-1 (39.6 mm), phalanx III-2 (24.7 mm), metatarsal IV (83.4 mm), phalanx IV-1 (27.1 mm), phalanx IV-2 (21 mm), metatarsal V (Norell and Makovicky, 1999)
(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)
(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, 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, metatarsal V (Norell and Mackovicky, 1999)
(IGM 100/2000) (juvenile) skull, mandible, postcranium (Barsbold and Osmolska, 1999)
(PIN 3143/8) incomplete skull, mandible (Barsbold and Osmolska, 1999)
numerous teeth (Barsbold and Osmolska, 1999)
Late Campanian, Late Cretaceous
Baruungoyot Formation, Mongolia
?(IGM 100/981; "Airakoraptor") partial skeleton including parts of skull, mandible, vertebrae, ulna, manus, ilium, and pes (Norell et al., 1992)
(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)
Comments- The holotype and paratype were discovered in 1923 and originally named "Ovoraptor djadochtari" by Osborn in 1924. However, this publication lacked a proper description, and Osborn 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 1971 (Kielan-Jaworowska and Barsbold, 1972) and described briefly by Barsbold (1974), with additional elements illustrated by him in 1983. This is the "fighting dinosaurs" specimen which is preserved grasping a Protoceratops, and has been analyzed by several authors (Osmolska, 1993; Unwin et al., 1995; Carpenter, 2000). Barsbold (1983) also mentioned another specimen, IGM 100/24, which is represented by a skull. ZPAL MgD-I/97 was found between 1963 and 1971 on the Polish-Mongolian expeditions, and mentioned briefly by Osmolska (1982) in her description of Hulsanpes. The skulls of all of these specimens, as well as PIN 3143/8 (found during a Soviet-Mongolian expediation) and IGM 100/2000 (found in a Mongolian-Japanese expedition), were described in detail by Barsbold and Osmolska (1999), with description of the postcrania in preperation.
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 was described in depth by Norell and Makovicky (1999). 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. IGM 100/982 was discovered in 1995 and described by Norell and Makovicky in 1999. Norell et al. (2004) described the braincases of IGM 100/976 and 100/982.
IGM 100/980 and 100/981 are from the Baruungoyot Formation at Khulsan and may also be Velociraptor. The former has been nicknamed "Ichabodcraniosaurus" (see entry) while the latter is discussed below.
"Airakoraptor"- "Airakoraptor" is only 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.
Three specimens discovered in 1991 are described in the abstract. The first (IGM 100/976) was referred to Velociraptor and was described in detail by Norell et al. (2004). The second (IGM 100/980) was mentioned and illustrated by Norell and Makovicky (1999) as an undescribed dromaeosaurid, and nicknamed "Ichabodcraniosaurus" by Novacek (1996). The third was said to be from Khulsan, very fragmentary and "possess features unknown in described dromaeosaurs." Noted characters are a deep mandible and dual posterior surangular foramina. While either IGM 100/980 or this specimen could be "Airakoraptor", the last choice seems most likely since it was specifically mentioned as being distinct. This specimen is also probably that described by Novacek as discovered by Perle on July 4th - "a fine skeleton of a dromaeosaur, perhaps not Velociraptor but something rare, and new." The deep jaw and dual surangular foramina remind one of Bagaraatan, though that genus is not a dromaeosaurid.
Of the Mongolian dromaeosaurid specimens that have appeared in the literature, only IGM 100/981 seems to be a potential match. This specimen was mentioned as an "unnamed dromaeosaur from Khulsan collected by the Mongolian American Expeditions" (Norell and Makovicky, 1997). It is 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 Dromaeosauridae based on caudal and pedal characters, but that insufficient elements are preserved to refer it to Velociraptor. Finally, Norell et al. (2006) illustrate the surangular as "an undescribed dromaeosaur from Khulson (IGM 100/981)" and it clearly shows dual surangular foramina. This clinches IGM 100/981 as the "Airakoraptor" specimen. However, Turner et al. (2007) described an ulna of a Velociraptor specimen they called IGM 100/981. This specimen was said to be from the Gilvent Wash locality near Ukhaa Tolgod and to "possess several characteristics found in V. mongoliensis." This may mean IGM 100/981 was later determined to just be a Velociraptor specimen.
References- Osborn, 1924. The discovery of an unknown continent. Natural History. 24(2), 133-149.
Osborn, 1924. 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.
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.
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, 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.
"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]
Osmolska, 1993. Were the Mongolian "fighting dinosaurs" really fighting? Rev. Paleobiol. 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.
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, 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.
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.
Carpenter, 2000. Evidence of predatory behavior by carnivorous dinosaurs. GAIA. 15, 135-144.
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. Unpublished dissertation. Rheinischen Friedrich-Wilhelms-Universität Bonn. 108 pp.
Kundrat, 2004. Two morphotypes of the Velociraptor neurocranium. 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. 133-143.
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.
Turner, Makovicky and Norell, 2007. Feather quill knobs in the dinosaur Velociraptor. Science. 317, 1721.
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. doi:10.1371/journal.pone.0007390
V. osmolskae Godefroit, Currie, Li, Shang and Dong, 2008
Late Campanian, Late Cretaceous
Bayan Mandahu Formation, Inner Mongolia, China
Holotype- (IMM 99NM-BYM-3/3) incomplete maxillae, lacrimal
Referred- ?(juvenile) several partial skeletons (Jerzykiewicz et al., 1993)
? teeth (Jerzykiewicz et al., 1993)
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- Jerzykiewicz et al. (1993) refer to several specimens as V. mongoliensis, but these remain undescribed. They may be referrable to the later named species V. osmolskae, though this is only based on provenance.
References- 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, 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.
V? sp. indet. (Bohlin, 1953)
Campanian-Maastrichtian, Late Cretaceous
Minhe Formation, Inner Mongolia, China
Material- two teeth
Comments- 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 mononykine manual ungual I however.
Reference- Bohlin, 1953. Fossil reptiles from Mongolia and Kansu. Sino-Swedish Expedition Publication. 37, 1-105.
V? sp. (Young, 1958)
Late Cretaceous
Tzoyun, Shanxi, China
Material- (IVPP V965) tooth
Comments- Young referred this to cf. Velociraptor mongoliensis, but as it is based on a single tooth, referral to any particular dromaeosaurid species is questionable. Sues (1977) noted it is larger than the V. mongoliensis holotype.
Reference- Young, 1958. The first record of dinosaurian remains from Shansi. Vertebrata PalAsiatica. 2(4), 231-236.
Sues, 1977. The skull of Velociraptor mongoliensis, a small Cretaceous theropod dinosaur from Mongolia. Palontologische Zeitschrift. 51, 173-184.

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- This taxon has yet to be included in a published cladistic analysis, but when run in a modified version of Senter's (2007) 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.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.

Deinonychus Ostrom, 1969
= "Daptosaurus" Brown vide Chure and McIntosh, 1989
D. antirrhopus Ostrom, 1969
= Velociraptor antirrhopus (Ostrom, 1969) Paul, 1988
= "Daptosaurus agilis" Brown vide Chure and McIntosh, 1989
Late Aptian, Early Cretaceous
Cloverly Formation, Montana, Wyoming
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, 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
Referred- (AMNH 3015) (3.06 m, 45 kg) 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), radiale, ulnares, 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 (Ostrom, 1969)
(AMNH 3034) teeth (Ostrom, 1970)
(AMNH 3037) teeth, manual elements, pedal elements (Ostrom, 1969)
(AMNH 3041; holotype of Microvenator celer in part) twenty-five teeth (Ostrom, 1970)
(AMNH 21609) 3 teeth (AMNH online)
(MCZ 4371) (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)
(MCZ 8791) (subadult) vertebrae including posterior dorsal and proximal caudal, coracoids, radius, ulna, manual phalanx II-2, tibia, fibula, pedal elements (Parsons and Parsons, 2004)
(MOR 682) eleven teeth (Maxwell and Ostrom, 1995)
(MOR 747) (several individuals) elements including anterior dentary (Maxwell, 1997)
(MOR 947) frontal (MOR online)
(MOR 1178) (~2.32 m; subadult) 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 (Parsons and Parsons, 2002)
(MOR 1182) caudal series (MOR online)
(MOR coll.) maxilla, nasal, lacrimal, jugal, prefrontal, frontal, parietal, quadrate, laterosphenoid, mesethmoid, braincase, ectopterygoid, palatine, teeth, other skull material, cervical vertebra, several caudal vertebrae, radius, ulna, manus, pes (Maxwell and Witmer, 1996; Maxwell and Witmer, 1996)
(OMNH 50268) (Parsons and Parsons, 2003)
(YPM 4886) (Ostrom, 1970)
(YPM 4887) (Ostrom, 1970)
(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 (Ostrom, 1969)
(YPM 5202) caudal vertebrae, chevrons (Ostrom, 1969)
(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 (Ostrom, 1969)
(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) (Ostrom, 1969)
(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 (Ostrom, 1969)
(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) (Ostrom, 1969)
(YPM 5208) radiale, ulnare (Ostrom, 1969)
(YPM 5209) manual phalanx II-2 (78 mm), manual phalanx III-3 (52 mm), manual ungual III (Ostrom, 1969)
(YPM 5210) (robust) (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) (Ostrom, 1969)
(YPM 5211) radiale (Ostrom, 1969)
(YPM 5212) manual ungual I (Ostrom, 1969)
(YPM 5213) manual phalanx I-1 (72 mm) (Ostrom, 1969)
(YPM 5214) metacarpal III (Ostrom, 1969)
(YPM 5215) manual phalanx III-3 (Ostrom, 1969)
(YPM 5216) manual phalanx II-1 (55.3 mm) (Ostrom, 1969)
(YPM 5217) radiale, 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) (Ostrom, 1969)
(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 (Ostrom, 1969)
(YPM 5219) (Ostrom, 1969)
(YPM 5220) radius (172 mm), ulnae (174.2, 180 mm), manual phalanx I-1 (67.2 mm), manual ungual I (Ostrom, 1969)
(YPM 5221) (Ostrom, 1969)
(YPM 5222) manual ungual II (Ostrom, 1969)
(YPM 5223) distal tarsal IV (Ostrom, 1969)
(YPM 5224) (Ostrom, 1969)
(YPM 5225) calcaneum (Ostrom, 1969)
(YPM 5226) astragalus, calcaneum (Ostrom, 1969)
(YPM 5227) (Ostrom, 1969)
(YPM 5228) ulnare (Ostrom, 1969)
(YPM 5229) distal tarsal (Ostrom, 1969)
(YPM 5230) ulna (Ostrom, 1969)
(YPM 5231) (Ostrom, 1969)
(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 (Ostrom, 1969)
(YPM 5233) ectopterygoid, pterygoid (Ostrom, 1969)
(YPM 5234) surangular (Ostrom, 1969)
(YPM 5235) ischium (161 mm) (Ostrom, 1969)
(YPM 5236) coracoid (Ostrom, 1969)
(YPM 5237) splenial (Ostrom, 1969)
(YPM 5238) splenial (Ostrom, 1969)
(YPM 5239) pterygoid (Ostrom, 1969)
(YPM 5240) proximal metatarsal I (45.5 mm), metatarsal I (40 mm) (Ostrom, 1969)
(YPM 5241) dorsal ribs (Ostrom, 1969)
(YPM 5242) ulnare (Ostrom, 1969)
(YPM 5243) manual phalanx III-3 (48.7 mm), manual ungual III (Ostrom, 1969)
(YPM 5244) chevron (Ostrom, 1969)
(YPM 5245) anterior dorsal rib (Ostrom, 1969)
(YPM 5246) sternal ribs (Ostrom, 1969)
(YPM 5247) gastralia (Ostrom, 1969)
(YPM 5248) (Ostrom, 1969)
(YPM 5249) mid dorsal rib, posterior dorsal rib (Ostrom, 1969)
(YPM 5250) (Ostrom, 1969)
(YPM 5251) (Ostrom, 1969)
(YPM 5252) (Ostrom, 1969)
(YPM 5253) (Ostrom, 1969)
(YPM 5254) (Ostrom, 1969)
(YPM 5255) (Ostrom, 1969)
(YPM 5256) (Ostrom, 1969)
(YPM 5257) (Ostrom, 1969)
(YPM 5258) (Ostrom, 1969)
(YPM 5259) (Ostrom, 1969)
(YPM 5260) (Ostrom, 1969)
(YPM 5261) (Ostrom, 1969)
(YPM 5262) (Ostrom, 1969)
(YPM 5263) (Ostrom, 1969)
(YPM 5264) (Ostrom, 1969)
(YPM 5265) (Ostrom, 1969)
(YPM 5266) (Maxwell and Ostrom, 1995)
(YPM 5267) (Maxwell and Ostrom, 1995)
(YPM 5268) (Maxwell and Ostrom, 1995)
(YPM 5269) (Maxwell and Ostrom, 1995)
(YPM 5270) metacarpal II (~93.5 mm) (Maxwell and Ostrom, 1995)
(YPM 5271) (Maxwell and Ostrom, 1995)
(YPM 5272) (Maxwell and Ostrom, 1995)
(YPM 5273) (Ostrom, 1970)
(YPM 5274) (Ostrom, 1970)
(YPM 5275) (Maxwell and Ostrom, 1995)
(YPM 5278) (Ostrom, 1970)
(YPM 5279) (Maxwell and Ostrom, 1995)
(YPM 5280) (Maxwell and Ostrom, 1995)
(YPM 5281) (Ostrom, 1970)
(YPM 5283) (Maxwell and Ostrom, 1995)
(YPM 5287) (Maxwell and Ostrom, 1995)
(YPM 5288) (Ostrom, 1970)
(YPM 5289) (Maxwell and Ostrom, 1995)
(YPM 5290) (Ostrom, 1970)
(YPM 5291) (Maxwell and Ostrom, 1995)
(YPM 5356) (Ostrom, 1970)
(YPM 5371) (Ostrom, 1970)
(YPM 5376) (Ostrom, 1970)
(YPM 5379) (Ostrom, 1970)
(YPM 5397) (Ostrom, 1970)
(YPM 5420) (Maxwell and Ostrom, 1995)
(YPM 5441) (Ostrom, 1970)
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) 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)
Middle-Late Aptian, Early Cretaceous
Arundel Formation, Maryland, US
?(USNM 497715) tooth (Lipka, 1998)
?(USNM 497719) tooth (Lipka, 1998)
?(USNM 497720) premaxillary tooth (Lipka, 1998)
?(USNM 497727) tooth (Lipka, 1998)
Early Cretaceous
Trinity Group, Texas, US
? seven teeth (Brinkman, Cifelli and Czaplewski, 1998)
Early Cretaceous
Arizona, US
? teeth (Thayer and Ratkevich, 1996)
Comments- 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 subadult Deinonychus (MCZ 8791) which was discovered in 1982. It is smaller than MOR 1178 and has even longer forelimbs.
References- Ostrom, 1969. A new theropod dinosaur from the Lower Cretaceous of Montana. Postilla. 128, 17 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.
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, 1-234.
Ostrom, 1974. The pectoral girdle and forelimb function of Deinonychus (Reptilia: Saurischia): a correction. Postilla. 165, 11 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.
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, Arizona State University, Tempe. 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.
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).
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.
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.

Tsaagan Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006
T. mangas Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006
Campanian, Late Cretaceous
Djadokhta Formation, Mongolia
Holotype- (IGM 100/1015) (adult) skull (201 mm), sclerotic rings, mandibles, ten cervical vertebrae, cervical ribs, proximal scapula, partial coracoid
Diagnosis- (after Norell et al., 2006) paroccipital process pendulous and not twisted distally; basipterygoid process elongate and anteroventrally directed; maxillary fenestra large and located at the anterior edge of the antorbital fossa; jugal meets the squamosal to exclude the postorbital from the margin of the infratemporal fenestra.
Comments- This specimen was originally identified as Velociraptor (Webster, 1996; AMNH website) before being described as a new genus.
References- Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
http://paleo.amnh.org/gobi/gobi.swf
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.

(Dromaeosaurus + Velociraptor) incertae sedis

"Dromaeosaurus" gracilis (Marsh, 1888) Matthew and Brown, 1922
= Coelurus gracilis Marsh, 1888
Late Aptian-Early Albian, Early Cretaceous
Arundel Formation, Maryland, US
Holotype- (USNM 4973) proximal manual ungual III
Referred- ?(USNM 8176; lost) tooth (Lull, 1911)
?(USNM 8444; = Goucher College 3336) tooth (Lull, 1911)
?(USNM 8445; = Goucher College 3338) tooth (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. 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 Judith River Group of Canada in 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 Gilmore's AMNH unguals belonged to Chirostenotes, and referred USNM 4973 to that genus. 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 resenble 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.

"Ichabodcraniosaurus" Novacek, 1996
Late Campanian, Late Cretaceous
Baruungoyot Formation, Mongolia
Material- (IGM 100/980) incomplete postcranial skeleton including atlas, axis, twelfth dorsal vertebra, thirteenth dorsal vertebra (15.1 mm), first sacral vertebra (16.5 mm), second caudal vertebra (19 mm), third caudal vertebra, fourth caudal vertebra (22.8 mm), fifth caudal vertebra (25.2 mm), five chevrons, ilium, pubes, ischium, complete hindlimb (Norell et al., 1992)
Comments- This specimen was discovered by Norell in 1991 and mentioned by Norell et al. (1992). Novacek noted the skeleton was complete except for lacking a skull, so was given the nickname "Ichabodcraniosaurus". 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. It has yet to be described or assigned to a particular genus, but may be Velociraptor and/or "Airakoraptor". The strongly opisthopubic pelvis and lack of dorsal ischial processes suggest it is a derived dromaeosaurid (not a microraptorian or unenlagiine).
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.
http://dml.cmnh.org/2002Feb/msg00648.html

"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.

Ornithodesmus Seeley, 1887
O. cluniculus Seeley, 1887
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (BMNH R187) (sacrum- 96 mm) first sacral vertebra (17 mm), second sacral vertebra (16 mm), third sacral vertebra (18 mm), fourth sacral vertebra (15 mm), fifth sacral vertebra (16 mm), sixth sacral vertebra (13 mm)
Comments- Seeley originally described Ornithodesmus as a bird, but Hulke (in anonymous, 1887) soon suggested it was pterosaurian. Seeley later (1901) referred BMNH R176 to Ornithodesmus as a new species, O. latidens. For over a century, the well known pterosaur Ornithodesmus latidens was used as the standard example of the genus. This ended in 1993 when Howse and Milner reidentified the Ornithodesmus cluniculus holotype as a theropod (the pterosaur was later renamed Istiodactylus latidens by Howse et al., 2001). Specifically they believed it to be a troodontid, based largely on comparison to BMNH R4463 (another supposed troodontid sacrum). Makovicky (1995) and Norell and Makovicky (1997) identified BMNH R4463 as Saurornitholestes, and the latter reference noted Ornithodesmus resembles Dromaeosauridae in possessing a well developed dorsal ridge formed by zygapophyses. Makovicky stated the specimen was "probably neither a troodontid nor a dromaeosaurid" while Naish et al. (2001) stated some characters argue against a dromaeosaurid identity, such as transverse processes which are not dorsoventrally flattened. That character is meaningless outside of a phylogenetic context however. As an alternative, Naish et al. note resemblence to Megapnosaurus and Carnotaurus in the presence of six sacrals, neural spine lamina and neural platform. Yet coelophysoids never have more than five sacrals, and dromaeosaurids like Velociraptor have all three listed characters. In addition, no coelophysoids or ceratosaurs have sacral pleurocoels or flattened ventral sacral surfaces with a median groove, unlike Ornithodesmus and dromaeosaurids. A dromaeosaurid identity seems most likely at this point.
References- Seeley, 1887. On a sacrum, apparently indicating a new type of Bird, Ornithodesmus cluniculus, Seeley, from the Wealden of Brook. Quarterly Journal of the Geological Society of London. 42, 206-211.
Anonymous, 1887. Discussion (on Ornithodesmus and Patricosaurus). Quarterly Journal of the Geological Society of London. 43, 219-220.
Seeley, 1901. Dragons of the Air. Methuen & Co., London, United Kingdom. 239 pp.
Howse and Milner, 1993. Ornithodesmus - a maniraptoran theropod dinosaur from the Lower Cretaceous of the Isle of Wight, England. Palaeontology. 36, 425-437.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Univ. Copenhagen, 311pp.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton: Information from a new specimen. American Museum Novitates. 3215, 1-28.
Howse, Milner and Martill, 2001. Pterosaurs. in Martill and Naish (eds.). Dinosaurs of the Isle of Wight. The Palaeontological Association, London. pp. 324-335.
Naish, Hutt and Martill, 2001. Saurischia dinosaurs: theropods. in Martill and Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association, Field Guides to Fossils. 10, 242-309.

Pyroraptor Allain and Taquet, 2000
P. olympius Allain and Taquet, 2000
Late Campanian-Early Maastrictian, Late Cretaceous
La Boucharde, Bouches-du-Rhone, France
Holotype- (MNHN BO001) pedal ungual II (66 mm)
Paratypes- ....(MNHN BO002) pedal phalanx II-2 (23 mm)
....(MNHN BO003) metatarsal II (118.7 mm)
....(MNHN BO004) pedal ungual II
....(MNHN BO005) ulna (112.8 mm)
....(MNHN BO006-BO010) pedal phalanges
....(MNHN BO011) manual phalanx
....(MNHN BO012) distal metacarpal I
....(MNHN BO013) incomplete radius
....(MNHN BO014) tooth
....(MNHN BO015) tooth
....(MNHN BO016) proximal caudal vertebra (24 mm)
....(MNHN BO017) dorsal vertebra
Diagnosis- deep m. brachialis fossa on lateral surface of proximal ulna; ventrally concave metatarsal II.
Comments- The other characters listed in Allain and Taquet's (2000) diagnosis are common in dromaeosaurids- teeth distally serrated and with restricted mesial serrations; ulna subequal in lenth to metatarsus; asymmetrically ginglymoid metatarsal II; strongly curved pedal ungual II.
Pyroraptor may be a junior synonym of the contemporary dromaeosaurid Variraptor, but this cannot be established until the dorsal vertebra MNHN BO017 or additional specimens are described. Alternatively, the paratypes and referred specimens of Variraptor may belong to Pyroraptor instead. They are not comparable presently.
Senter et al. (2004) included Variraptor in their phylogenetic analysis of coelurosaurs and found it to be a dromaeosaurid excluded from the Jehol microraptorian clade (based on the stout pedal phalanx II-2) and Dromaeosaurus+Utahraptor (based on the high DSDI).
References- 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.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8, 1-20.

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) 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- 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 presence of an ulna from the Variraptor type locality that lacks Pyroraptor's apomorphic deep brachial fossa suggests the two genera may be distinct, however.
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. When added to Senter's later (2007) analysis, which does have these characters, Variraptor emerges as a dromaeosaurine based on the short dorsal centra, and outside the Utahraptor+Achillobator clade based on the posterior dorsal postzygapophyses which extend posterior to the centra.
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). Ph.D. dissertation, University of Bristol, 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. Bull. Soc. géol. Fr.. 180(2), 145-154

Dromaeosaurinae Matthew and Brown, 1922
Definition- (Dromaeosaurus albertensis <- Velociraptor mongoliensis) (modified from Sereno, 1998)
Other definitions- (Dromaeosaurus albertensis <- Microraptor zhaoianus, Velociraptor mongoliensis, Unenlagia comahuensis, Passer domesticus) (Sereno, in press)
= Dromaeosaurinae sensu Sereno, in press
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 Sereno's 2005 definition.

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
Comments- This tooth 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 questinable.
Reference- 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.

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.

Dromaeosaurinae indet. (Fiorillo, 1999)
Late Albian, Early 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)
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. (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.

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 dromaeosaurine (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)
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.
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 possible dromaeosaurine (Averianov, 2007)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- (ZIN PH 33/49) (juvenile) tooth (FABL 3.1 mm) (Averianov, 2007)
Comments- 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.
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.

undescribed dromaeosaurine (Currie and Eberth, 1993)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- teeth (Currie and Eberth, 1993)
Comments- Currie and Eberth (1993) note teeth that indicate an indeterminate dromaeosaurine larger than Velociraptor. 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.

Dromaeosauroides Christiansen and Bonde, 2003
D. bornholmensis Christiansen and Bonde, 2003
Late Berriasian, Early Cretaceous
Jydegaard Formation, Denmark
Holotype- (MGUH DK No. 315) anterior dentary tooth (21.7 mm)
Diagnosis- 25% larger than Dromaeosaurus albertensis; 6 distal serrations per mm compared to 3-3.5/mm in D. albertensis.
Comments- This tooth 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.

Zapsalis Cope, 1876
Z. abradens Cope, 1876
= Dromaeosaurus abradens (Cope, 1876)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Holotype- (AMNH 3953) tooth (12 mm)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith River Group, Alberta, Canada
Material- ?(RTMP 79.15.3) tooth (15.5 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 80.16.833) tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 81.26.37) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.14.45) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.19.7) tooth (15 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.458) tooth (12.5 mm), tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.91.39) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.91.40) tooth (12.5 mm), tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.163.80) tooth (10 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.2) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.131) tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.132) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.133) tooth (~11.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.58.65) tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.9.92) tooth (~14.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.23.104) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.36.425) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.77.112) tooth (~14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.130.219) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.198.46) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.5) tooth (15.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.383) lateral tooth (Baszio, 1997)
(RTMP 89.50.8) tooth (13.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.312) tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.50.202) tooth (9.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.6.15) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.50.208) tooth (~11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.53.21) tooth (~11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 91.50.60) tooth (14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 92.50.23) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.268) tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 97.133.2) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP coll.; lost) lateral tooth (Baszio, 1997)
thirty teeth (Baszio, 1997)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, US
Material- ?(RTMP 1009) lateral tooth (Baszio, 1997)
several teeth (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
Material- (UA 103) lateral tooth (Baszio, 1997)
(UA 121) premaxillary tooth (Baszio, 1997)
several teeth (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- (AMNH 27122; in part) tooth (AMNH online)
(AMNH coll.) teeth (Estes, 1964)
(UA 132) lateral tooth (Baszio, 1997)
Comments- Estes (1964) included all theropod teeth with ridges and a flattened lingual side in Paronychodon, including those with serrations. 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. This makes a pathological explanation unlikely.
The holotype of Zapsalis abradens is a tooth which is flat lingually, with no mesial serrations and 3 distal serrations per mm (Cope, 1876). 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 based on its flat lingual side and ridges, which has been followed in the literature without comment ever since. However, Zapsalis falls outside the current concept of Paronychodon in having serrations, and is more robust than teeth of that genus as well. Yet Zapsalis does match ?Dromaeosaurus morphotype A, which are lingually flat with ridges on both sides, and serrations which can occur on both carinae (3-4/mm). The proportions also match, with an average length of 12.4 mm, FABL of 6.3 mm, and BW of 3.4 mm. Thus Zapsalis is here used as the valid name for these teeth.
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 Zapsalis, it differs in the curvature, 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. The DSDI, serration shape and curvature match Saurornitholestes, but the ridges do not and the serrations are slightly too small.
Another similar tooth (RTMP 79.15.3) was illustrated by Currie et al. as a Saurornitholestes Paronychodon-like tooth. It has four lingual ridges, a straight distal edge with 3 serrations per mm, a curved mesial edge with 6 serrations per mm, and a basal constriction. Posterior serrations are long and appear to be hooked. In this case, the ridges, curvature, and distal serration size match Zapsalis. Yet the mesial serration size and distal serration shape do not. Though the DSDI is more similar to Saurornitholestes and Richardoestesia, the serration size is too large for either. Those two genera also lack the ridges and straight distal edge of this tooth. It was referred to ?Dromaeosaurus morphotype A by Sankey et al., despite the incongruent characters present. It's possible these two teeth belong to another part of the jaw of Zapsalis, or are a distinct species of more basal dromaeosaurid.
References- Cope, 1876. On some extinct reptiles and batrachia from the Judith River and Fox Hills Beds of Montana. Proceedings of the Academy of Natural Sciences, Philadelphia. 28, 340-359.
Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
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.
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, 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.

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), teeth, fragments
Referred- ?(RTMP 97.39.4) tooth (Ryan et al., 2000)
?(RTMP 1005-1009, 1033, 1034) seven teeth (Baszio, 1997)
?(RTMP coll.) seven teeth (Ryan et al., 2000)
Diagnosis- snout shorter and deeper than other dromaeosaurids; subnarial and nasal processes of premaxilla inclined more than 45 degrees dorsally; maxillary fenestra enlarged; maxillary fenestra directly dorsal to promaxillary fenestra.
Comments- 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'. These teeth are all provisionally referred to Atrociraptor here, as no definite Saurornitholestes specimens have been found in this formation.
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.

Dromaeosaurus [UNDER CONSTRUCTION]
Late Campanian, Late Cretaceous
Judith River Formation, Montana
Material- (AMNH 8516) lateral tooth (AMNH online)
(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)
D. sp. (Williamson, 2001)
Late Campanian, Late Cretaceous
Fruitland and/or Lower Kirtland Formation, New Mexico, US
Material- teeth
Reference- Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
D. sp.
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- ?(AMNH 25268) astragalus (AMNH online)
(YPM PU 22312) (YPM online)

Achillobator Perle, Norell and Clark, 1999
A. giganticus Perle, Norell and Clark, 1999
Santonian-Campanian, Late Cretaceous
Bayan Shireh Formation, Mongolia
Holotype- (FR.MNUFR-15) (~5 m; adult) maxilla (290.8 mm), nine teeth (to 38 mm), sixth cervical vertebra (34.6 mm), tenth cervical vertebra (51.3 mm), fourth?dorsal vertebra (53.8 mm), posterior dorsal vertebra (49.5 mm), anterior dorsal rib, posterior dorsal rib, mid caudal vertebra, six distal caudal vertebrae, three chevrons, scapula, coracoid, radius (260 mm), metacarpal III (71 mm), phalanx I-1 (78 mm), two manual unguals (112 mm), ilium (531 mm), pubes (548 mm), ischium (378 mm), femur (505 mm), tibia (490.4 mm), phalanx II-2 (56.4 mm), metatarsal III (234.4 mm), phalanx III-2 (55 mm), metatarsal IV (209.6 mm)
Diagnosis- (modified from Perle et al., 1999) anterior pubic foot slightly longer than posterior foot; very stout pedal phalanx II-2.
very short anterior dorsal centra; two pairs of pleurocoels in posterior dorsal centra; pleurocoel-like foramina on caudal vertebrae; mid caudal chevrons with sinuous ventral margin; manual elements very robust; sinuous ridges present on the lateral surface of the ilium, above the peduncles and acetabulum; metatarsal IV with distal lateral condyle strongly reduced.
Comments- Achillobator was discovered in 1989 and described in an obscure Mongolian journal before its time. Norell and Clark originally intended to publish it in American Museum Novitates with a comparative analysis. Unfortunately, it was released in an extremely preliminary form in Contributions of the Mongolian-American Paleontological Project without their knowledge (pers. comm.). Indeed, the section "Habits and affinities of dromaeosaurian dinosaurs" was neither written nor seen by Norell and Clark prior to publication.
The holotype was found as an associated but disarticulated specimen. No other dinosaurs were found in the area, so it is unlikely this is a chimaera as Burnham et al. (2000) have supposed. They stated that the maxilla, ilium, ischium and caudal vertebrae share no unique characters with dromaeosaurids. This is obviously not true, as the maxilla has fused interdental plates and a dorsally placed maxillary fenestra; the caudal vertebrae have elongate prezygapophyses and chevrons; while the ilium has a concave anterior edge that slants posteroventrally.
Description- The skull is represented by a maxilla and several teeth. The maxilla is deeper than Deinonychus and is missing most of the nasal process. Compared to Deinonychus, the nasal process projects more vertically, the premaxillary suture is more vertical and the jugal process is more ventrally projected. The antorbital fossa appears more recessed and the teeth are more widely spaced. The internal structure of the maxilla is partially known, as there is a medial excavation of the nasal process by a two-chambered sinus that opens laterally to form the maxillary fenestra. There are eleven alveoli and the interdental plates appear to be fused. The teeth are recurved and serrated, with anterior serrations being slightly smaller than posterior serrations (17-20 per 5 mm vs. 15-18 per 5 mm).
The sixth or seventh cervical is preserved. It's centrum is concave in front and taller than wide anteriorly. In addition, there are prominent epipophyses and the ribs are unfused. Compared to Deinonychus, the neural spine is transversely narrower and directed posteriorly, the neural canal is larger, the parapophyses are more pronounced, the diapophyses are smaller, the posterior articular surface is concave dorsally and the anterior articular surface is taller. The interspinal ligament scars extend futher dorsally than in Deinonychus as well.
The tenth cervical vertebra has an amphiplatyan centrum with deep pleurocoels. It is slightly wider than tall anteriorly and slightly taller than wide posteriorly. There is a broad keel ventrally and hyposphene-hypantra articulations are present.
A few dorsal vertebrae are known including a mid-dorsal (possibly fourth) and a posterior dorsal. The centra are amphicoelous with deep pleurocoels, double in posterior dorsals. Anterior pleurocoels are larger when there are two pairs. The articular surfaces are slightly higher than wide in the mid-dorsal. The ventral surface is slightly keeled, but there is no hypapophysis. Hyposphene-hypantrum articulations are present and the vertebral foramen is about 23% of centrum height. Other differences from Deinonychus include shorter centra and almost vertically directed prezygapophyses.
The text states only one rib was recovered, but two are figured. The rib the text mentions is supposed to be either the last cervical or first dorsal, but the two pictured come from the mid-anterior and posterior portion of the vertebral column. They are similar to Deinonychus, but exact comparison is difficult without having ribs from the same vertebrae to compare to.
There are seven caudal vertebrae preserved, one from the middle of the series, the rest from the distal tip. They are amphicoelous and the distal ones lack neural spines and transverse processes. There is a "pleurocoel-like foramen" on the basal surface of the transverse process that probably connects to the neural canal. Another foramen is also present, exiting from the anteroventral base of transverse process next to the pleurocoel-like foramen. Elongate prezygapophyses are present, although the postzygapophyses lack elongate rods. The prezygapophyses bifurcate more distally than Deinonychus, if at all. The neural spine is more prominent than Deinonychus, extending past the postzygapophyseal articular surfaces, and the centra are shorter.
Three mid-caudal and distal chevrons are present in the holotype. They are very short dorsoventrally, but have elongate anterior and posterior processes. The anterior processes are dromaeosaurid-like in their elongation, but must bifurcate more distally than Deinonychus. Other differences from Deinonychus are the sigmoid ventral outline, more widely spaced articular facets and trifurcate anterior process.
A scapulocoracoid is preserved missing only the distal end. It is very similar to Deinonychus, with a very shallow acromial expansion and slender shaft. It narrows distally a bit, unlike Deinonychus. The coracoid is broadly similar to Deinonychus, being elongate with a prominent coracoid tubercle and foramen. The anteroventral surface is not as projecting as Deinonychus and the posterior process is much larger and triangular. The glenoid faces posteroventrally.
A radius is illustrated, but not described. It has a more expanded distal end than Deinonychus and the proximal end is expanded more gradually. It resembles dromaeosaurids in being longer than the metatarsus.
There are four manual elements preserved. The third metacarpal is stouter than in Deinonychus and bowed dorsally, but not laterally. Both manual phalanx I-I and manual ungual I are present, although the latter is not figured. The phalanx differs from Deinonychus in it's stoutness and small details of articulations, but is very similar in shape dorsally. Manual ungual I is reported to be laterally compressed, recurved and have a large flexor tubercle. The supposed second pedal ungual is actually a manual ungual (Senter et al., 2004; Senter, 2007).
The ilium is very unique in structure. It is very tall with a short preacetabular process and longer postacetabular process. The preacetabular process is like a dorsoventrally expanded version of Deinonychus', with a posteroventrally slanted concave anterior edge. The dorsal margin is slightly convex and slopes ventrally over the postacetabular process. The postacetabular process is quite tall, extends ventrally past the ischial peduncle and has a vertical posterior margin, with a posterodorsal and a posteroventral tubercle. The pubic peduncle is nearly vertical and has a concave ventral edge facing slightly posteriorly. The ischial peduncle is reduced, perhaps with a prominent antitrochantor, and the acetabulum is partially closed off medially. Several sinuous ridges are present on the lateral surface above the peduncles and acetabulum. In addition, there are many other ridges and striations for various muscles on different areas of the medial and lateral surfaces.
The pubes are well-preserved and distict from other dromaeosaurids. I estimate they were projected ventrally or slightly posteriorly. They are straight with the proximal end expanded a bit anteriorly and a small obturator notch. There is no pectineal process and the shafts are circular in cross section. The distal foot is slightly longer anteriorly than posteriorly with a pointed anterior foot and a convex ventral edge. The two pubes are joined for 69% of their length and have straight lateral margins the whole way down, unlike the narrow foot of Deinonychus.
The ischium has a more proximally placed obturator process than Deinonychus (18% down shaft) and is longer compared to the pubis (69%). The pubic peduncle is narrower, the obturator process longer and the distal end is blunt. There is a small proximodorsal process. The two ischia were not fused, but may have had a mobile articulation.
The femur is moderately bowed anteriorly and round in cross section. The head is slightly declined and separated from the greater trochantor by a moderately depressed surface, while the lesser trochantor is small and barely separated from the greater trochantor. The posterior trochantor and a distally placed (~40% down shaft) fourth trochantor are also present. The anterodistal fossa is absent and unlike Deinonychus, the tibial condyle is much longer than the fibular condyle.
The tibia is 97% of femoral length and similar to Deinonychus in most respects, although stouter. The cnemial crest is much larger and directed more dorsally and the fibular crest is very proximally placed. What may be the facet for the astragalar ascending process is 29% of tibial length and expanded lateromedially.
Metatarsals III and IV are known. Metatarsal III is 46% of femoral length and clearly not artometatarsalian. The proximal end of metatarsal III is compressed transversely, but not nearly as much as in Deinonychus. Unlike Deinonychus, the distal end of metatarsal IV is ginglymoideal, with a very small lateral condyle.
Several pedal phalanges are mentioned, but only phalanx II-2 and ungual II are figured. Phalanx II-2 is similar to dromaeosaurids in that it has a large proximoventral heel, but it is much stouter than Deinonychus and even Adasaurus.
Relationships- The only comment regarding it's placement within the Dromaeosauridae in the original description is in the abstract, where the authors state it is most closely related to Dromaeosaurus. The Theropod Working Group analyses (Norell et al., 2001 and variations) have always recovered Achillobator as a dromaeosaurid, generally in a polytomy with the non-microraptorian taxa. Turner et al. (2007) and several other more recent variations have added many characters relevant to dromaeosaurids and found Achillobator to be most closely related to Utahraptor, Dromaeosaurus and Adasaurus. Senter et al. (2004) and Senter (2007) both found Achillobator to be the sister taxon to Dromaeosaurus + Utahraptor. Adasaurus was more basal in both. In addition, Britt et al. (2001) indicate new materials of Utahraptor share characters with Achillobator.
References- 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.
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.
Britt, Chure, Stadtman, Madsen, Scheetz and Burge, 2001. New osteological data and the affinities of Utahraptor from the Cedar Mountain Fm. (Early Cretaceous) of Utah. Journa of Vertebrate Paleontology. 21(3), 36A.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian dinosaurs. pp. 49–67 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 Univ. Press.
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. Journal of Systematic Palaeontology.
Turner, Hwang and Norell, 2007. A small derived theropod from Oosh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates. 3557, 27 pp.

Utahraptor [UNDER CONSTRUCTION]
U. ostrommaysi
Referred- (BYUVP 15465) femur (600 mm) (Erickson et al., 2009)
References- 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. doi:10.1371/journal.pone.0007390
U. sp. (Carpenter, online)
Aptian, Early Cretaceous
Poison Strip Member of Cedar Mountain Formation, Utah, US
Material- ? pelvic element
Reference- https://scientists.dmns.org/sites/kencarpenter/Cedar%20Mountain%20storage/Dinosaurs.aspx
U. sp. (Carpenter, online)
Early Albian, Early Cretaceous
Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- ? manual ungual
Reference- https://scientists.dmns.org/sites/kencarpenter/Cedar%20Mountain%20storage/Dinosaurs.aspx