Ningyuansaurus Ji, Lu, Wei
and Wang, 2012
N. wangi Ji, Lu, Wei and Wang, 2012
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (Xingcheng Confuciusornis Museum coll.) skull, mandibles,
nine cervical vertebrae, cervical ribs, ten dorsal vertebrae, dorsal ribs, gastralia,
~22 caudal vertebrae, chevrons, scapula (73 mm), coracoid, (?)sternal plates,
humerus (75 mm), radii (55 mm), ulnae (58 mm), scapholunare, semilunate carpal,
distal carpal III, metacarpals I (12 mm), phalanx I-1 (25 mm), manual ungual I, metacarpals II
(28 mm), phalanx II-1, phalanx II-2, metacarpal III (29 mm), two manual phalanges,
three manual unguals, ilia (70 mm), pubes (109 mm), ischia (41 mm), femora (135
mm), tibiae (172 mm), fibula (172 mm), astragali (17 mm wide), metatarsals II
(88 mm), metatarsals III (105 mm), phalanx III-1 (24 mm), metatarsals IV (102
mm), pedal phalanges, pedal unguals, metatarsal V, body feathers, seeds
Diagnosis- (after Ji et al., 2012) at least 10 upper and 14 lower jaw
teeth on each side; femur much longer than ilium; distal caudal vertebrae much
elongated.
Comments-
Ji et al. state the holotype, discovered prior to April 2012, was
offered for study by a private individual. The locality was listed as
"Lamadong, Jianchang County, Liaoning Province; Yixian Formation", but
Li et al. (2014) state "recent fieldwork confirmed the outcrops of
fossil beds exposed in Lamadong Village mainly belong to the Jiufotang
Formation (X. L. Wang, personal commun.)." The description is
brief and the illustration and photo quality poor, so that not much
information is determinable. While the authors believe it is an
oviraptorosaur more basal than Caudipteryx or Incisivosaurus,
the low iliofemoral ratio, low ischiopubic ratio and enlarged pedal ungual ?II
suggest possible paravian relations. Funston and Currie (2016) added Ningyuansaurus to Osmolska et al.'s oviraptorosaur matrix lineage and recovered it in a polytomy with Caudipteryx, Similicaudipteryx and Luoyanggia, but other than oviraptorosaurs only include Protarchaeopteryx, Archaeopteryx, Velociraptor and Herrerasaurus, not even recovering a paravian Velociraptor.
Hartman et al. (2019) used a full taxon sample and recovered the taxon
as a halszkaraptorine paravian, requiring 4 steps to move to
Oviraptorosauria.
References- Ji, Lu, Wei and Wang, 2012. A new oviraptorosaur from the
Yixian Formation of Jianchang, western Liaoning Province, China. Geological
Bulletin of China. 31(12), 2102-2107.
Li, Zhou, Wang and Clarke, 2014. A new specimen of large-bodied basal enantiornithine
Bohaiornis from the Early Cretaceous of China and the inference of feeding
ecology in Mesozoic birds. Journal of Paleontology. 88(1), 99-108.
Funston and Currie, 2016. A new caenagnathid (Dinosauria:
Oviraptorosauria) from the Horseshoe Canyon Formation of Alberta,
Canada, and a reevaluation of the relationships of Caenagnathidae.
Journal of Vertebrate Paleontology. 36(4), e1160910. 36(4), e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Hulsanpes Osmolska, 1982
H. perlei Osmolska, 1982
Late Campanian, Late Cretaceous
Khulsan, Baron Goyot Formation, Mongolia
Holotype- (ZPAL MgD-I/173) (~560 mm; juvenile) prootic-opisthotic fragment,
incomplete metatarsal II (34 mm), phalanx II-1 (6.5 mm), proximal phalanx II-2, incomplete metatarsal
III (~39 mm), proximal phalanx III-1, incomplete metatarsal IV (~36 mm)
Diagnosis- (after Cau and Madzia, 2018) distal third of
metatarsal II shaft extensively overlapped by metatarsal III in
extensor view; proximal shaft of metatarsal III unconstricted and wider
than adjacent metatarsals; metatarsal III-IV contact straight in
extensor view; distal end of metatarsal IV diverges laterodistally.
Other diagnoses- Osmolska (1982) originally diagnosed Hulsanpes
based on symplesiomorphies for Neotheropoda (functionally tridactyl pes;
metatarsals II-IV subequally thick), Tyrannoraptora (metatarsus slender
[width: length ratio 0.16]), Deinonychosauria (second pedal digit
specialized), and Halszkarptorinae (metatarsal III not wedged in
anterior view).
Comments- This specimen was discovered in Summer 1970 (Kielan-Jaworowska and Barsbold, 1972) and described twelve years
later. Using Halszkaraptor as a guide, we can estimate Hulsanpes was about 560 mm long, but it was juvenile based on the rough bone texture and the badly
abraded articular joints.
Osmolska allied this species with dromaeosaurids based
on the ginglymoid second and third metatarsals and subequally developed metatarsals
II and IV, but noted it resembled troodontids in the narrow metatarsus and weakly
developed second pedal digit. She dismissed avian origins based on the lack
of fusion. Chiappe and Norell (Norell pers. comm. to
Currie, 2001) think Hulsanpes is not a dromaeosaurid, but from "another
more speciose branch of the Maniraptora". Senter et al. (2004) was the first published analysis to include Hulsanpes,
which emerged as a dromaeosaurid because they misscored the ginglymoid metatarsal
II as absent in Sinovenator. Cau et al. (2017) described the more complete taxon Halszkaraptor and created Halszkaraptorinae for these and Mahakala finding them sister to unenlagiids plus dromaeosaurids using his Megamatrix.
References- Kielan-Jaworowska and Barsbold, 1972.
Narrative of the Polish-Mongolian Palaeontological Expeditions
1967-1971. Palaeontologica Polonica. 27, 5-13.
Osmolska, 1982. Hulsanpes perlei n. g. n. sp. (Deinonychosauria,
Saurischia, Dinosauria) from the Upper Cretaceous Barun Goyot Formation of Mongolia.
Neues Jahrbuch fur Geologie und Palaeontologie, Monatshefte. 1982(7), 440-448.
Currie, 2001. Theropod dinosaurs from the Cretaceous of Mongolia. in Benton,
Shishkin, Unwin and Kurochkin, eds. The Age of Dinosaurs in Russia and Mongolia.
434-455.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae.
Bulletin of Gunma Natural History Museum. 8, 1-20.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Cau and Madzia, 2018. Redescription and affinities of Hulsanpes perlei (Dinosauria, Theropoda) from the Upper Cretaceous of Mongolia. PeerJ. 6:e4868.
Mahakala Turner, Pol, Clarke,
Ericson and Norell, 2007
M. omnogovae Turner, Pol, Clarke, Ericson and Norell, 2007
Late Campanian, Late Cretaceous
Tugriken Shire, Djadokhta Formation, Mongolia
Holotype- (IGM 100/1033) (~700 mm; adult) maxillary fragment, frontals
(25.2 mm), partial quadrate, ectopterygoid, pterygoid fragment, posterior braincase,
mandibular fragment, articular? fragment, dentary tooth, atlas, partial axis,
third cervical prezygapophysis, incomplete fourth cervical vertebra (~16.6 mm),
partial fifth cervical vertebra, two fragmentary posterior cervical vertebrae,
parts of twelve dorsal vertebrae including first dorsal neural arch, second
dorsal neural arch, third dorsal vertebra, anterior dorsal centrum (~9.5 mm),
six posterior dorsal vertebrae (~9.2, ~9.8, ~10 mm), incomplete sacrum, first
caudal vertebra, second caudal vertebra, third caudal vertebra, fourth caudal
vertebra, fifth caudal vertebra, sixth caudal vertebra, seventh caudal vertebra,
eighth caudal vertebra, ninth caudal vertebra, tenth caudal vertebra, eleventh
caudal vertebra, twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth
caudal vertebra, fifteenth caudal vertebra, sixteenth caudal vertebra, seventeenth
caudal vertebra, eighteenth caudal vertebra, nineteenth caudal vertebra, twentieth
caudal vertebra, six chevrons, partial scapulae, incomplete humeri (~35-40 mm),
radius (36 mm), partial ulnae (~40 mm), semilunate carpal, distal carpal III, metacarpal
I (~6.8 mm), phalanx I-1 (~13.1 mm), metacarpals II (18 mm), phalanx II-1 (~9.5
mm), phalanx II-2 (~13.2 mm), manual ungual II (~12.5 mm), metacarpals III (16
mm), phalanx III-1, manual ungual III (~11.4 mm), manual ungual, incomplete
ilium (52.5 mm), femur (76.2 mm), tibiae (110 mm; one fragmentary), proximal
fibula, astragalocalcaneum, metatarsals I (one incomplete), distal phalanx I-1,
pedal ungual I, metatarsal II (~74 mm), phalanx II-2 (9 mm), pedal ungual II
(16 mm straight), metatarsals III (82 mm; one fragmentary), phalanges III-1
(19.1 mm), phalanx III-2 (16 mm), phalanges III-3 (13 mm), incomplete pedal
ungual III (10.5 mm), partial metatarsals IV (~77 mm), phalanx IV-1, phalanx
IV-2, phalanx IV-3, phalanx IV-4, three phalanges IV-?, pedal unguals IV
Diagnosis- (after Turner et al., 2007) proximal caudal vertebrae with
subhorizontal, laterally directed prezygapophyses (unknown in Hulsanpes); strongly compressed and anteroposteriorly
broad ulna tapering posteriorly to a narrow edge; elongate lateral crest on
the posterodistal part of the femur.
(after Turner et al., 2011) ledgelike depression at the confluence of metotic
strut and posterior tympanic recess on the anterior face of the paroccipital
process; posteriorly tapering scapula; short forelimb (humerus 50% femur length);
prominent supratrochanteric process; absence of a cuppedicus fossa (unknown in Hulsanpes).
(after Cau et al., 2017) pendant paroccipital processes (unknown in Hulsanpes); short forelimb (humerus not longer than 50% of femur; (unknown in Hulsanpes); distal ends of metatarsals I and II
diverge medially (unknown in Hulsanpes); metatarsal III longer than femur (unknown in Hulsanpes); distal end of metatarsal III with symmetrical constriction proximal to trochlea.
Comments- Discovered in 1992 and first noted by Turner et al. (2006).
This is generally recovered as sister to unenlagiids plus
dromaeosaurids using the TwiG matrix (as in Turner et al. 2007 and 2011), although Senter et
al. (2012) recovered it as a microraptorian. Cau et al. (2017) described the more complete Halszkaraptor and created Halszkaraptorinae for these and Hulsanpes, finding them sister to unenlagiids plus dromaeosaurids using his Megamatrix and Brusatte's TWiG analysis.
References- Turner, Pol, Norell and Hwang, 2006. Resolving dromaeosaurid
phylogeny: New information and additions to the tree. Journal of Vertebrate
Paleontology. 26(3), 133A.
Turner, Pol, Clarke, Ericson and Norell, 2007. A basal dromaeosaurid and size
evolution preceding avian flight. Science. 317, 1378-1381.
Turner, Pol, Conicet, Clarke and Norell, 2007. The basal-most dromaeosaurid:
A new species from Tugrugyin Shireh, Mongolia. Journal of Vertebrate Paleontology.
27(3), 161A.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian
physiology inherited by birds? Reconciling slow growth in Archaeopteryx.
PLoS ONE. 4(10), e7390.
Turner, Pol and Norell, 2011. Anatomy of Mahakala omnogovae (Theropoda:
Dromaeosauridae), T�gr�giin Shiree, Mongolia. American Museum Novitates.
3722, 66 pp.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Unenlagiinae Makovicky et al., 2005
Definition- (Unenlagia comahuensis <- Halszkaraptor escuilliei, Microraptor zhaoianus, Velociraptor mongoliensis, Dromaeosaurus albertensis, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019)
Other definitions- (Unenlagia comahuensis <- Velociraptor
mongoliensis) (Makovicky, Apesteguia and Agnolin, 2005)
(Unenlagia comahuensis <- Microraptor zhaoianus, Velociraptor
mongoliensis, Dromaeosaurus albertensis, Passer domesticus) (
Turner, Makovicky and Norell, 2012)
(Unenlagia comahuensis <- Microraptor zhaoianus, Dromaeosaurus albertensis, Vultur gryphus) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
Comments- Unenlagiinae was
erected by Makovicky et al. (2005) to contain Unenlagia, Buitreraptor
and Rahonavis at the base of Dromaeosauridae. It has been recovered in
this position in most subsequent Theropod Working Group analyses, though sometimes
in Avialae instead (Agnolin and Novas, 2011, 2013).
Turner et al.'s (2012) definition replaced Makovicky et al.'s, which only included
Velociraptor as an external specifier, while Hartman et al. (2019) added Halszkaraptor so that halszkaraptorines could be a subfamiliy of unenlagiines.
References- Bonaparte, 1999. Tetrapod faunas from South America and India:
A paleobiogeographic interpretation. PINSA. 65A(3), 427-437.
Makovicky, Apestegu�a and Agnol�n, 2005. The earliest dromaeosaurid
theropod from South America. Nature. 437, 1007-1011.
Novas and Agnolin, 2010. Phylogenetic relationships of Unenlagiidae theropods:
Are they members of Dromaeosauridae?
Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the Dromaeosauridae
(Theropoda, Maniraptora)? Anais da Academia Brasileira de Ci�ncias. 83(1),
117-162.
Gianechini and Apesteguiia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods
from South America. Anais da Academia Brasileira de Ci�ncias. 83(1), 163-195.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships
of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae.
Springer. 96 pp.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
unnamed possible unenlagiine (Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton,
Zhang, Wolfe, Yi, Tseng, Ran and Currie, 2016)
Early Cenomanian, Late Cretaceous
Angbamo, Myanmar
Material- (DIP-V-15103) (juvenile?) about nine mid to distal caudal vertebrae (2.97, 3.41, 3.41, 3.76, 4.01 mm), skin, feathers
Reference- Xing, McKellar, Xu, Li, Bai, Persons IV, Miyashita, Benton,
Zhang, Wolfe, Yi, Tseng, Ran and Currie, 2016. A feathered dinosaur
tail with primitive plumage trapped in Mid-Cretaceous amber. Current
Biology. 26(24), 3352-3360.
unnamed unenlagiine (Langston, 1953)
Maastrichtian, Late Cretaceous
Ortega, Columbia
Material- (UCMP 39649b) incomplete lateral tooth
Comments- Initially identified as a carnosaur (Langston, 1953), this
was described as an unenlagiine similar to Austroraptor by Ezcurra (2009).
References- Langston, 1953. Cretaceous terrestrial vertebrates from Colombia,
South America. Bulletin of the Geological Society of America. 64, 1519.
Ezcurra, 2009. Theropod remains from the latest Cretaceous of Colombia and their
implications on the palaeozoogeography of western Gondwana. Cretaceous Research.
30, 1339-1344.
unnamed unenlagiine (Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Briss�n-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, �lvarez-Herrera, Gentil, Bogan, Chimento,
Garc�a-Mars�, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019)
Late Campanian-Early Maastrichtian, Late Cretaceous
Chorrillo Formation, Santa Cruz, Argentina
Material- (MPM 21548) partial pedal ungual II
?...(MPM 21549) pedal phalanx III-2 (45 mm)
Comments- Discovered between
January and March 2019, Novas et al. assign MPM 21545 and 21546 to
Unenlagiidae.
Reference- Novas, Agnolin,
Rozadilla, Aranciaga-Rolando, Briss�n-Eli, Motta, Cerroni, Ezcurra,
Martinelli, D'Angelo, �lvarez-Herrera, Gentil, Bogan, Chimento,
Garc�a-Mars�, Lo Coco, Miquel, Brito, Vera, Perez Loinaze, Fernandez
and Salgado, 2019. Paleontological discoveries in the Chorrillo
Formation (upper Campanian-lower Maastrichtian, Upper Cretaceous),
Santa Cruz Province, Patagonia, Argentina. Revista del Museo Argentino
de Ciencias Naturales. 21(2), 217-293.
unnamed unenlagiine (Candeiro, Cau, Fanti, Nava and Novas, 2012)
Turonian-Santonian, Late Cretaceous
Adamantina Formation of the Bauru Group, Brazil
Material- (MPM 011) (~1 m; adult) anterior-mid dorsal vertebra
Reference- Candeiro, Cau, Fanti, Nava and Novas, 2012. First evidence
of an unenlagiid (Dinosauria, Theropoda, Maniraptora) from the Bauru Group,
Brazil. Cretaceous Research. 37, 223-226.
unnamed unenlagiine (Benson, Rich, Vickers-Rich and Hall, 2012)
Late Aptian-Early Albian, Early Cretaceous
Eumeralla Formation of the Otway Group, Victoria, Australia
Material- (NMV P180889) proximal femur
Reference- Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna
from Southern Australia indicates high polar diversity and climate-driven dinosaur
provinciality. PLoS ONE. 7(5), e37122.
"Lopasaurus" Price vide Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021 online
Maastrichtian, Late Cretaceous
Ponto 1 do Price, Serra da Galga Formation, Brazil
Holotype- (DNPM coll.; lost) metatarsal II (~162 mm), distal metatarsal III, distal metatarsal IV
Comments- In 2021 the Serra da
Galga and Ponte Alta Members of the Marilia Formation were recognized
as the Serra da Galga Formation. Collected
in the 1950s and only surviving as an unfinished plate drawn by Price
labeled 'Lopasaurus', this is presumably the "metatarsals (all in the
DNPM, Rio de Jineiro) which Price believed included at least two new
genera and families." Brum et al. (2021 online) stated it
disappeared after Price's death and first published the figure and
name, identifying it as an unenlagiine based on "the anterior surface
of metatarsal III transversally expanded; the expansion of the
posterolateral region of metatarsal II; and the expansion of the
posteromedial region of metatarsal IV. In posterior view, the
expansions observed in metatarsals II and IV overlap metatarsal
III." While this could belong to the contemporaneous Ypupiara,
Brum et al. note it was not associated with the type skull fragments,
and multiple unenlagiines may have coexisted as in the Portezuelo
Formation. "Lopasaurus" is a nomen nudum, as it was not used as
valid when
proposed (ICZN Article 11.5), "accompanied by a description or
definition that states in words characters that are purported to
differentiate the taxon" (Article 13.1.1), or "accompanied by the
fixation of a type species in the original publication" (Article 13.3).
References- Bertini, Marshall,
Gayet and Brito, 1993. Vertebrate faunas from the Adamantina and
Mar�lia formations (Upper Bauru Group, Late Cretaceous, Brazil) in
their stratigraphic and paleobiogeographic context. Neues Jahrbuch f�r
Geologie und Pal�ontologie Abhandlungen. 188(1), 71-101.
Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021 online. A new
unenlagiine (Theropoda, Dromaeosauridae) from the Upper Cretaceous of
Brazil. Papers in Palaeontology. DOI: 10.1002/spp2.1375
Austroraptor Novas, Pol, Canale,
Porfiri and Calvo, 2009
= "Austroraptor" Novas, Pol, Canale, Porfiri and Calvo, 2008 online
A. cabazai Novas, Pol, Canale, Porfiri and Calvo, 2009
= "Austroraptor cabazai" Novas, Pol, Canale, Porfiri and Calvo, 2008
online
Campanian-Maastrichtian, Late Cretaceous
Allen Formation, Rio Negro, Argentina
Holotype- (MML-195) (~4.9 m; 368 kg) (skull ~800 mm) incomplete maxillae,
lacrimals, frontal, postorbital, incomplete dentaries, surangular, prearticular,
third cervical vertebra (~111 mm), fifth cervical vertebra, sixth cervical vertebra,
seventh cervical vertebra, eighth cervical vertebra (~79 mm), second dorsal
vertebra, fourth dorsal vertebra (~68 mm), ribs, gastralia, humerus (262 mm),
manual ungual III (53 mm), partial pubis, incomplete femur (~560 mm), tibia
(565 mm), astragalus, calcaneum, pedal ungual I, metatarsal II (325 mm), phalanx
II-2 (57 mm), metatarsal III (330 mm), pedal ungual III (67 mm), phalanx IV-2
(48 mm)
Referred- (MML-220) (adult) fragmentary maxilla, two teeth, incomplete
mid-posterior dorsal vertebra (96 mm), twelve distal caudal centra (32-82 mm),
incomplete humerus, radii (161 mm), ulnae (one incomplete, one partial; 188
mm), metacarpals I (39 mm), phalanx I-1 (90.5 mm), incomplete manual ungual
I, manual unguals II (one partial; 67 mm), incomplete manual ungual III, tibia
(~560 mm), phalanx I-1 (41 mm), proximal metatarsal II, distal metatarsal II,
phalanx II-1 (61.2 mm), phalanx II-2 (58.1 mm), distal metatarsals III, phalanx
III-1 (98.2 mm), phalanx III-2 (56.2 mm), phalanx III-3 (48.5 mm), phalanx IV-1
(49.3 mm), phalanx IV-2 (33.7 mm), pedal ungual IV (Carabajal et al., 2009;
described in Currie and Paulina Carabajal, 2012)
Diagnosis- (modified from Novas et al., 2009) lacrimal highly pneumatized;
ventral process strongly curved anteriorly; posterior process flaring out horizontally
above orbit; postorbital lacking dorsomedial process for articulation with the
frontal; squamosal process of postorbital extremely reduced; maxillary and dentary
teeth fluted; humerus short, representing slightly less than 50 per cent of
femur length; pedal phalanx II-2 transversely narrow, contrasting with the extremely
robust phalanx IV-2.
Other diagnoses- Novas et al. (2009) also included the small tooth size
and lack of serrations as apomorphies, but these are probably basal for paravians.
Comments- Note "Austroraptor" was earlier used as an unpublished
name for Ozraptor (Pigdon, DML 1997). Austroraptor's name and
description appeared online in December 2008 before being officially published
in March 2009.
Novas et al. (2009) assigned the taxon to Unenlagiinae based on a TWiG analysis,
which has been the result of all subsequent published analyses as well.
References- Pigdon, DML 1997. https://web.archive.org/web/20191030133231/http://dml.cmnh.org/1997Sep/msg00942.html
Novas, Canale and Isasi, 2004. Giant deinonychosaurian theropod from the Late
Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 24(3), 26A-27A.
Carabajal, Currie, Garcia, Salgado, Cerda, Fernandez, Reichel, Sissons, Koppelhus
and Cabaza, 2009. Un nuevo especimen de Austroraptor (Dinosauria: Theropoda:
Dromaeosauridae) del Cret�cico Tard�o (Maastrichtiano) de R�o
Negro, Argentina. Ameghiniana. 46(S), 41R.
Novas, Pol, Canale, Porfiri and Calvo, 2009. A bizarre Cretaceous theropod dinosaur
from Patagonia and the evolution of Gondwanan dromaeosaurids. Proceedings of
the Royal Society B. 276(1659), 1101-1107.
Gianechini and Apesteguiia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods
from South America. Anais da Academia Brasileira de Ci�ncias. 83(1), 163-195.
Currie and Paulina Carabajal, 2012. A new specimen of Austroraptor cabazai
Novas, Pol, Canale, Porfiri and Calvo, 2008 (Dinosauria, Theropoda, Unenlagiidae)
from the Latest Cretaceous (Maastrichtian) of Rio Negro, Argentina. Ameghiniana.
49(4), 662-667.
Ypupiara Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021
Y. lopai Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021
Maastrichtian, Late Cretaceous
Ponto 1 do Price, Serra da Galga Formation, Brazil
Holotype- (DGM 921-R; destroyed) anterior maxillary fragment, dentary fragment
Diagnosis- (after Brum et al.,
2021) preantorbital body of maxilla with only one neurovascular foramen
(unknown in other unenlagiines); maxillary interdental plate
rectangular and anteroposteriorly expanded (unknown in other
unenlagiines); maxillary teeth with a CBR of between 0.6 and 1 (also in
Austroraptor); teeth of maxilla widely spaced (also in Buitreraptor).
Comments- In 2021 the Serra da
Galga and Ponte Alta Members of the Marilia Formation were recognized
as the Serra da Galga Formation. Originally collected
in the 1950s, Holgado et al. (2018) announced this in an abstract as a
new taxon of unenlagiine. Brum et al. state "After the great fire
that destroyed the main building of the MN-UFRJ in September 2018 ...
the specimen was not recovered and is here considered as lost."
They added Ypupiara to Cau's matrix and recovered it as an unenlagiine sister to Austroraptor, which is also the result when added to the Hartman et al. matrix.
References- Holgado, Brum,
Pegas, Bandeira, Souza, Kellner and Campos, 2018. A new Unenlagiinae
(Theropoda: Dromaeosauridae) from the Maastrichtian of Brazil. Journal
of Vertebrate Paleontology, Program and Abstracts, 2018. 148.
Brum, Pegas, Bandeira, Souza, Campos and Kellner, 2021 online. A new
unenlagiine (Theropoda, Dromaeosauridae) from the Upper Cretaceous of
Brazil. Papers in Palaeontology. DOI: 10.1002/spp2.1375
Alcmonavis Rauhut, Tischlinger and Foth, 2019
A. poeschli Rauhut, Tischlinger and Foth, 2019
Early Tithonian, Late Jurassic
M�rnsheim Formation, Germany
Holotype- (SNSB-BSPG 2017 I 133; Muhlheim specimen; 13th specimen)
humerus (~90 mm), radius (~80-82 mm), ulna (82 mm), scapholunare, pisiform,
semilunate carpal, distal carpal III, incomplete metacarpal I (~7.1
mm), phalanx I-1 (28.5 mm), manual ungual I (16 mm), metacarpal II
(40.9 mm), phalanx II-1 (22.8 mm), phalanx II-2 (28 mm), manual ungual
II (17.5 mm), metacarpal III (36.8 mm), phalanx III-1 (8.8 mm), phalanx
III-2 (5.9 mm), phalanx III-3 (17.9 mm), manual ungual III (11 mm),
manual claw sheaths
Diagnosis- (after Rauhut et al., 2019) humerus with large
deltopectoral crest, with a maximal expansion that exceeds the width of
the humeral shaft; proximal part of humerus strongly angled at
approximately 38 degrees with respect to distal shaft; ulna with
well-defined, single, oval, concave proximal cotyle and small lateral
tubercle; distal end of ulna slightly asymmetrically expanded; large,
crest-like biceps tubercle on proximal radius; longitudinal groove
along medial side of radial shaft; metacarpal II considerably more
robust than metacarpals I and III; manual phalanx I-1 with longitudinal
groove; manual phalanx II-1 very robust, but with rounded, rather than
flattened cross-section; manual phalanx II-1 slightly twisted; manual
unguals with strongly developed and palmarly transversely expanded
flexor tubercles.
Comments- This was discovered in 2017 and initially thought to be a new Archaeopteryx specimen.
Rauhut et al. (2019) added it to Foth et al.'s TWiG analysis and
recovered it as an avialan more derived than archaeopterygids but
outside Shemzhouraptor+Sapeornis+Pygostylia.
When added to Hartman et al.'s TWiG analysis, it emerges as an
unenlagiine, which can be avialans with the addition of a single step
(albeit further from Aves than archaeopterygids).
References- Rauhut, Tischlinger and Foth, 2019. A non-archaeopterygid avialan
theropod from the Late Jurassic of southern Germany. eLife. 8:e43789.
Buitreraptor Makovicky,
Apesteguia and Agnolin, 2005
B. gonzalezorum Makovicky, Apesteguia and Agnolin, 2005
Cenomanian-Turonian, Late Cretaceous
Candeleros Formation of the Rio Neuquen Subgroup, Rio Negro, Argentina
Holotype- (MPCA 245) (>4 year old subadult) incomplete skull (~190
mm), incomplete mandibles (~166 mm), five teeth (2.5-4.6 mm), incomplete axis,
partial third cervical vertebra, partial fourth cervical vertebra, fifth cervical
vertebra, partial sixth cervical vertebra, eighth cervical vertebra, ninth cervical
vertebra, tenth cervical vertebra, partial first dorsal vertebra, second dorsal
vertebra, third dorsal vertebra, partial fourth dorsal vertebra, fifth dorsal
vertebra, incomplete sixth dorsal vertebra, seventh dorsal vertebra, eighth
dorsal vertebra, ninth dorsal vertebra, incomplete tenth dorsal vertebra, incomplete
eleventh dorsal vertebra, partial twelfth dorsal vertebra, partial thirteenth
dorsal vertebra, fifteen dorsal ribs, incomplete sacrum (57 mm), first caudal
vertebra, second caudal vertebra, third caudal vertebra, fourth caudal vertebra,
fifth caudal vertebra, sixth caudal vertebra, seventh caudal vertebra, eighth
caudal vertebra, ninth caudal vertebra, tenth caudal vertebra, eleventh caudal
vertebra, twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal
vertebra, fifteenth caudal vertebra, nine chevrons, incomplete scapulae (~96
mm), coracoids (45 mm), furcula (~73 mm), humeri (136 mm), incomplete radius
(95 mm), incomplete ulna (112.2 mm), semilunate carpal?, partial metacarpal
I?, metacarpal II (20.1 mm), partial metacarpal III?, manual phalanges, partial
ilia, ischium (54 mm), femora (149.5 mm), tibiae (one incomplete; >158 mm),
fibulae (one incomplete; >127 mm), incomplete metatarsal II, phalanx II-1
(22.8 mm), phalanx II-2 (19.3 mm), phalanx IV-1 (23.9 mm), fragments
Paratype- (MPCA 238) (>4 year old subadult) incomplete sacrum, ilium,
pubis, femur, tibiotarsus, metatarsal I (20 mm), phalanx I-1 (16.2 mm), incomplete
metatarsal II (90.6 mm), phalanx II-1 (27.9 mm), phalanx II-2 (18.3 mm), pedal
ungual II, incomplete metatarsal III (99.4 mm), incomplete metatarsal IV (90.6
mm)
Referred- (MPCA 471-C) material including partial phalanx I-1, partial
metacarpal II, partial phalanx III-3, four manual fragments (Anonymous, 2005)
(MPCA 471-D) material including appendicular element (Cerda and Gianechini,
2015)
(MPCA? coll.) (Anonymous, 2005)
(MPCN-PV-598) (8+ year old subadult) three incomplete anterior cervical vertebrae
with fused cervical ribs, sixth dorsal vertebra, partial seventh dorsal vertebra,
eighth dorsal centrum, ninth dorsal centrum, tenth dorsal centrum, eleventh
dorsal centrum, partial twelfth dorsal centrum, partial thirteenth dorsal vertebra,
dorsal rib, fragmentary gastralia, partial synsacrum, fragmentary first caudal
vertebra, fragmentary second caudal vertebra, partial third caudal vertebra,
incomplete fourth caudal vertebra, incomplete fifth caudal vertebra, sixth caudal
vertebra, seventh caudal vertebra, eighth caudal vertebra, ninth caudal vertebra,
tenth caudal vertebra, incomplete eleventh caudal vertebra, twelfth caudal vertebra,
thirteenth caudal vertebra, fourteenth caudal centrum, anterior fifteenth caudal
centrum, two distal caudal vertebrae, chevrons, incomplete scapula, incomplete
coracoid, incomplete furcula, incomplete humerus, incomplete radius, incomplete
ulna, semilunate carpal, incomplete metacarpal I, phalanx I-1, manual ungual
I, partial metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, partial
metacarpal III, incomplete phalanges III-1, phalanx III-2, phalanx III-3, manual
elements, incomplete ilia, incomplete pubes, fragmentary ischia, femur (148
mm), incomplete tibia (~185 mm), incomplete fibula, partial astragalus, metatarsal
I, phalanx I-1, pedal ungual I, metatarsals II (one distal), phalanx II-1, phalanx
II-2, pedal ungual II, phalanges II-?, metatarsals III (one distal; ~104 mm),
phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals
IV (one distal), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal
ungual IV, phalanges IV-? (Novas et al., 2017)
Diagnosis- (after Turner et al., 2012) skull long, exceeding femoral
length by 25%; large maxillary fenestra; continuous transition from frontal
margin to postorbital process; quadrate with large lateral flange and pneumatic
foramen; dentary bears a deep subalveolar groove; teeth small, unserrated, without
root-crown constriction; posterior cervical centra with ventrolateral ridge;
furcula pneumatic; flexor process present on humerus; brevis shelf expanded
and lobate, projects laterally from posterior end of ilium.
(after Novas et al., 2017) dorsal vertebrae lacking pleurocoels; mid and distal
caudals with lateral complex of ridges (transverse process with ridge that extends
as feebly developed rims that reachs the prezygapophyses and postzygapophyses;
anterior half of centrum with small ridge running towards the transverse process
and a similar ridge is located at the posterior end of the centrum; sub-triangular
fossa on anterior and posterior lateral margins, delimited by tiny ridges);
scapular blade axially expanded at mid-length; extremely slender manual elements;
manus longer than femur (117% of femoral length); penultimate pedal and manual
phalanges with dorsally displaced collateral ligament pits that are medially
displaced and nearly contact each other.
Comments- Briefly described by Makovicky et al. (2005) and Gianechini
and Apesteguia (2011), Gianechini is studying this taxon for his thesis. While
Novas et al. (2017) identified the most complete dorsaal vertebra as the seventh
"based on comparisons with [the] holotype specimen", but given the
two posteriormost dorsals (12 and 13) articulated with the sacrum and the five
articulated posterior dorsals, it must be at least as anterior as the sixth.
Makovicky et al. found Buitreraptor to be an unenlagiine using the TWiG
matrix, which has been recovered in all subsequent analyses as well.
References- Anonymous, 2005. Newly discovered birdlike dinosaur is oldest
raptor ever found in South America.
Makovicky, Apestegu�a and Agnol�n, 2005. The earliest dromaeosaurid
theropod from South America. Nature. 437, 1007-1011.
Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the Dromaeosauridae
(Theropoda, Maniraptora)? Anais da Academia Brasileira de Ci�ncias. 83(1),
117-162.
Gianechini and Apesteguiia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods
from South America. Anais da Academia Brasileira de Ci�ncias. 83(1), 163-195.
Gianechini, Makovicky and Apesteguia, 2011. The teeth of the unenlagiine theropod
Buitreraptor from the Cretaceous of Patagonia, Argentina, and the unusual
dentition of the Gondwanan dromaeosaurids. Acta Palaeontologica Polonica. 56(2),
279-290.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships
of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae.
Springer. 96 pp.
Cerda and Gianechini, 2015. Determinacion de estadios ontogeneticos en Buitreraptor
gonzalezorum (Dinosauria: Theropoda) a partir de la microestructura osea.
XXIX Jornadas Argentinas de Paleontolog�a de Vertebrados, resumenes.
Ameghiniana. 52(4) suplemento, 11-12.
Novas, Briss�n Egli, Agnolin, Gianechini and Cerda, 2017. Postcranial
osteology of a new specimen of Buitreraptor gonzalezorum (Theropoda,
Coelurosauria). Cretaceous Research. doi: 10.1016/j.cretres.2017.06.003.
Pamparaptor Porfiri, Calvo and
dos Santos, 2011
P. micros Porfiri, Calvo and dos Santos, 2011
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of the Rio Neuquen Subgroup, Neuquen, Argentina
Holotype- (MUCPv-1163) (~500-700 mm) metatarsal II (82.1 mm), phalanx
II-1 (16.6 mm), phalanx II-2 (18.1 mm), proximal pedal ungual II, metatarsal
III (93 mm), phalanx III-1 (26.5 mm), phalanx III-2 (17.4 mm), metatarsal IV
(92.5 mm), distal phalanx IV-1, proximal phalanx IV-2
Diagnosis- (after Porfiri et al., 2011) slender metatarsus; metatarsal
II approximately twice the proximal width of III or IV; metatarsal II distally
overlapping metatarsal III; distal end of metatarsal II with small medially
directed sulcus anteriorly; pedal phalanx II-2 longer than II-1; proximal half
of metatarsal III narrow and with subparallel margins; metatarsal III not ginglymoid;
metatarsals IV and III subequal in length; distal half of metatarsal IV strongly
compressed transversely, acquiring a blade-like shape in posterior view.
Comments- This was discovered in 2005 and initially referred to Neuquenraptor
by Porfiri et al. (2007), with the differences considered ontogenetic. Porfiri
et al. (2011) later described it as a new taxon of dromaeosaurid, though they
noted troodontid similarities and did not run a phylogenetic analysis. Novas
et al. (2013) questioned the unenlagiine identification, stating the elongate
metatarsus, "distally extended metatarsal IV, subequal in distal extension
to metatarsal III", non-ginglymoid metatarsals II and IV, and more elongate
and acute pedal ungual were more similar to birds. They regarded it as Paraves
incertae sedis.
References- Porfiri, Calvo, dos Santos and Valieri, 2007. New record
of Neuquenraptor (Theropoda, Dromaeosauridae) from the Late Cretaceous
of Patagonia. Ameghiniana. 44(S), 34R.
Porfiri, Calvo and dos Santos, 2011. A new small deinonychosaur (Dinosauria:
Theropoda) from the Late Cretaceous of Patagonia, Argentina. Anais da Academia
Brasileira de Ci�ncias. 83(1), 109-116.
Novas, Agnolin, Ezcurra, Porfiri and Canale, 2013. Evolution of the carnivorous
dinosaurs during the Cretaceous: The evidence from Patagonia. Cretaceous Research.
45, 174-215.
Overoraptor Motta, Agnolin, Briss�n Egli and Novas, 2020
O. chimentoi Motta, Agnolin, Briss�n Egli and Novas, 2020
Middle Cenomanian-Early Turonian, Late Cretaceous
Huincul Formation of Rio Limay Subgroup, Rio Negro, Argentina
Holotype- (MPCA-Pv 805) (~1.3 m) proximal caudal vertebra, two mid
caudal vertebrae (20.5 mm), distal caudal centrum (16.6 mm), two
chevrons, incomplete scapula, ulna (~116 mm), incomplete metacarpal I
(~14.9 mm), partial phalanx I-1, manual ungual I (~25.1 mm), manual
ungual II, ilial fragment, pubic shaft, phalanx I-1 (13.6 mm),
incomplete pedal ungual I (~14.5 mm), metatarsals II (100.6 mm),
phalanx II-2 (20.0 mm), incomplete pedal ungual II, metatarsals III
(106.5 mm)
Paratype- (MPCA-Pv 818) (~1 m)
manual phalanx I-1, phalanx III-2, ilial fragment, incomplete pubis,
metatarsal II (~79 mm), phalanx II-1, phalanx III-1
Diagnosis- (after Motta et al., 2020) distal caudal centra with complex system of lateral longitudinal ridges and concavities (also in Buitreraptor and Rahonavis);
scapula with medially deflected distal end; acromion reduced and
ridge-like; robust ulna; radial cotyle of ulna saddle-shaped and
prominent; metacarpal I with extensive medioventral crest; metatarsal
II with longitudinal lateroventral crest on distal half, ending
distally in a posterior tubercle; strongly dorsally displaced
collateral pits on pedal phalanx II-2; metatarsal III distal end
non-ginglymoid (also in Pamparaptor); metatarsal III distal end dorsoventrally deeper than transversely wide.
Comments- The mid and distal
caudal vertebrae, metacarpal I and pedal ungual II of the holotype were
originally described as ?Unenlagiidae gen. et sp. indet. by Motta et
al. (2016). Note the measurements in Motta et al.'s (2020)
Supplementary Table 1 are in cm, not mm as listed.
Motta et al. (2020) described the taxon and added it to Agnolin and Novas' TWiG analysis where it emerged sister to Rahonavis,
one node closer to Ornithes than unenlagiids. Adding it to
Hartman et al.'s maniraptoromorph analysis results in it being an
unenlagiine close to Rahonavis, but making it sister to that henus takes 4 more steps.
References- Motta, Aranciaga Rolando, Rozadilla, Agnolin, Chimento, Brisson
Egli and Novas, 2016. New theropod fauna from the Upper Cretaceous (Huincul
Formtation) of northwestern Patagonia, Argentina. In Khosla and Lucas (eds.).
Cretaceous period: Biotic diversity and biogeography. New Mexico Museum of Natural
History and Science Bulletin. 71, 231-253.
Motta, Agnolin, Briss�n Egli and Novas, 2020. New theropod dinosaur
from the Upper Cretaceous of Patagonia sheds light on the paravian
radiation in Gondwana. The Science of Nature. 107(3), 24.
Rahonaviformes Livezey and Zusi, 2007
"Rahonavidae" Miller, 2004
Comments- Miller (2004) proposed Rahonavidae for Rahonavis and Jeholornis,
but did not indicate the family was new (e.g. on page 142 it lacks the
asterisk he used to indicate which taxa were new), so is a nomen nudum
(ICZN Article 16.1). Livezey and Zusi (2007) erected
Rahonaviformes and Rahonavidae containing only Rahonavis itself. As these were only used in a table
that lacked a diagnosis or definition, "Rahonavidae" was still a nomen nudum
(ICZN Article 13.1.1).
References- Miller, 2004. A new phylogeny of the Dromaeosauridae. 2004 Student Showcase Journal. 20, 123-158.
Livezey and Zusi, 2007. Higher-order phylogeny of modern birds
(Theropoda, Aves: Neornithes) based on comparative anatomy. II. Analysis and
discussion. Zoological Journal of the Linnean Society. 149 (1), 1-95.
Rahonavis Forster, Sampson, Chiappe
and Krause, 1998b
= Rahona Forster, Sampson, Chiappe and Krause, 1998a (preoccupied Griveaud,
1975)
R. ostromi (Forster, Sampson, Chiappe and Krause, 1998) Forster,
Sampson, Chiappe and Krause, 1998b
= Rahona ostromi Forster, Sampson, Chiappe and Krause, 1998a
Maastrichtian, Late Cretaceous
Anembalemba Member of Maevarano Formation, Madagascar
Holotype-
(UA 8656) (adult) incomplete ~first dorsal vertebra (12.0 mm), seventh
dorsal vertebra (10.8 mm), eighth dorsal vertebra (10.7 mm), partial
ninth dorsal neural arch, partial tenth dorsal vertebra, incomplete
eleventh dorsal vertebra (9.8 mm), incomplete twelfth dorsal vertebra
(9.1 mm), incomplete synsacrum (42.3 mm), incomplete first caudal
vertebra, second caudal vertebra (6.3 mm), third caudal vertebra (6.4
mm), fourth caudal vertebra (6.9 mm), fifth caudal vertebra (8.1 mm),
sixth caudal vertebra (9.4 mm), seventh caudal vertebra (9.5 mm),
eighth caudal vertebra (12.2 mm), ninth caudal vertebra (14.9 mm),
tenth caudal vertebra (18.5 mm), eleventh caudal vertebra (22.6 mm),
twelfth caudal vertebra (24.7 mm), thirteenth caudal vertebra (25.8
mm), twelve chevrons (2.8-11.6 mm dorsoventrally), scapula (82.2 mm),
radius (126.5 mm), ulna (132.3 mm), ilia (66.5, 67.1 mm), pubes (one
incomplete; 69.1 mm), incomplete ischia (27.4 mm), femora (87.6, 86.9
mm), tibiae (one incomplete; 118.1mm), incomplete fibulae, incomplete
astragalocalcaneum (10.7 mm trans), distal tarsal IV, metatarsal I (8.6
mm), phalanx I-1 (10.2 mm), pedal ungual I (11.1 mm), metatarsals II
(44.8, 43.9 mm), phalanx II-1 (12.8 mm), phalanx II-2 (11.3 mm), pedal
ungual II (22.8 mm), metatarsals III (47.8, 47.6 mm), phalanx III-1
(19.2 mm), phalanx III-2 (12.9 mm), phalanx III-3 (10.5 mm),
metatarsals IV (44.8, 43.1 mm), phalanx IV-1 (14.7 mm), phalanx IV-2
(9.6 mm), phalanx IV-3 (7.3 mm), pedal claw II sheath,
Referred- (FMNH PA 740) incomplete dentary (Forster et al., 2020)
(FMNH PA 746) distal humerus (Forster and O'Conner, 2000;
described by O'Connor and Forster, 2010)
(FMNH PR 2821) incomplete ulna (Forster et al., 2020)
(UA 9604) distal humerus (Forster and O'Conner, 2000; described by O'Connor
and Forster, 2010)
Diagnosis- (after Turner et al., 2012) neural canal at least 40% of the
height of the dorsal vertebral centra; six sacral vertebrae; deeply concave
glenoid fossa on scapula; elongate acromion process (length greater than length
of glenoid); long muscle scar above glenoid on lateral face of scapula; proximally
kinked scapular shaft; elongate ulna with ulnar papillae; ulnar distal condyle
subtriangular in distal view and twisted more than 54 degrees with respect to
the proximal end; undivided trochanteric crest, that is distally shifted on
the femoral shaft; prominent and proximally placed fingerlike lateral ridge
on femur; mediolaterally broad cnemial crest with short lateral process; proximal
end of tibia of equal width and length; greatly reduced fibula lacking contact
with the calcaneum; lack of a medial fossa on the fibula.
Comments- The genus Rahona is preoccupied by limantriid moths
(Griveaud, 1975), so Forster et al. (1998b) renamed their genus Rahonavis.
All elements of Rahonavis
were found in 1995. The distal humeri were both found near to the
holotype and articulate perfectly with the type ulna, so may belong to
the same individual (O'Connor and Forster, 2010). O'Connor and Forster
call them Humeral Taxon B. However, a second right ulna was also
found in the same layer, so another individual of the same size was
present who the humeri could also belong to.
Originally recovered as an avialan by Forster et al (1998a) and the TWiG matrix
(Norell et al., 2001), Makovicky et al. (2005) and several other iterations
of the TWiG matrix instead place it as a basal dromaeosaurid in Unenlagiinae.
This was also recovered by Foth et al. (2014), and the most recent versions
of the TWiG matrix using Turner's paravian updates (Brusatte et al., 2014). Agnolin
and Novas (2011) recover a compromise phylogeny where unenlagiines include Rahonavis
and are placed as basal avialans, while Xu et al. (2008) and Lee et al. (2014)
again found it to be an avialan, apart from deinonychosaurian unenlagiines.
References- Griveaud, 1975. Descriptions pr�liminaires de nouveaux
genres et esp�ces de Lymantriidae malgaches (Lep.). Bulletin de la Soci�t�
entomologique de France. 80, 225-232.
Forster, Sampson, Chiappe and Krause, 1998a. The theropod ancestry of birds:
New evidence from the Late Cretaceous of Madagascar. Science. 279, 1915-1919.
Forster, Sampson, Chiappe and Krause, 1998b. Genus correction. Science. 280(5361),
179.
Schweitzer, Watt, Avci, Forster, Krause, Knapp, Rogers, Beech and Marshall,
1999. Keratin immunoreactivity in the Late Cretaceous bird Rahonavis ostromi.
Journal of Vertebrate Paleontology. 19(4), 712-722.
Forster and O'Conner, 2000. The avifauna of the Upper Cretaceous Maevarano Formation,
Madagascar. Journal of Vertebrate Paleontology. 20(3), 41A-42A.
Chinsamy and Elzanowski, 2001. Evolution of growth pattern in birds. Nature. 412, 402-403.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian
theropods. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early
Evolution of Birds: Proceedings of the International Symposium in Honor of John
H. Ostrom. 49-67.
Makovicky, Apestegu�a and Agnol�n, 2005. The earliest dromaeosaurid
theropod from South America. Nature. 437, 1007-1011.
Xu, Zhao, Norell, Sullivan, Hone, Erickson, Wang, Han and Guo, 2008. A new feathered
maniraptoran dinosaur fossil that fills a morphological gap in avian origin.
Chinese Science Bulletin. 54(3), 430-435.
O'Connor and Forster, 2010. A Late Cretaceous (Maastrichtian) avifauna from
the Maevarano Formation, Madagascar. Journal of Vertebrate Paleontology. 30(4),
1178-1201.
Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the Dromaeosauridae
(Theropoda, Maniraptora)? Anais da Academia Brasileira de Ci�ncias. 83(1),
117-162.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Forster, O'Connor, Chiappe and Turner, 2020. The osteology of the Late Cretaceous paravian Rahonavis ostromi from Madagascar. Palaeontologia Electronica. 23(2):a31.
Dakotaraptor DePalma, Burnham,
Martin, Larson and Bakker, 2015
D. steini DePalma, Burnham, Martin, Larson and Bakker, 2015
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Holotype- (PBMNH.P.10.113.T) (robust adult) fragmentary dorsal centrum (~70
mm), mid caudal centrum (~75 mm), distal caudal vertebra (70 mm), four incomplete
distal caudal vertebra, two distal caudal centra, partial distal caudal centrum,
incomplete humeri (~320 mm), incomplete radii (~320 mm), ulnae (360 mm), metacarpal
I (75 mm), distal phalanx I-1, metacarpal II (130 mm), distal phalanx II-2,
distal phalanx III-2, femur (558 mm), tibiae (673 mm), astragalocalcaneum (~120
mm trans), metatarsal II (247 mm), pedal ungual II (160 mm straight), fragmentary
metatarsal III (~320 mm), pedal ungual III (70 mm), metatarsals IV (one fragmentary;
290 mm)
Paratypes- ?(KUVP 156045) tooth
(PBMNH.P.10.115.T) (gracile adult) tibia (485 mm)
(PBMNH.P.10.118.T) (gracile) astragalocalcaneum (60 mm trans)
?(PBMNH.P.10.119.T) tooth
?(PBMNH.P.10.121.T) tooth
?(PBMNH.P.10.122.T) tooth
?(PBMNH.P.10.124.T) tooth
Diagnosis- (after DePalma et al., 2015) large size (~5.5 m long); teeth
have 15-20 denticles/5 mm on distal carina and 20-27 denticles/5 mm on mesial
carina; lateral and medial condyles of metacarpal II subequal in size; proximodorsal
margin of metacarpal II straight in dorsal view; shallow lateral ligament fossa
on metacarpal II; pedal ungual II (straight measurement) ~29% of femur length;
fibular crest of tibia long and gracile (height does not exceed 9% of total
length); proximal margin of fibular crest hooked; sharp ventral keel on pedal
unguals II and III; reduced flexor tubercle on pedal ungual IV; lateral groove
of pedal ungual IV fully enclosed in bone for about its distal half.
Comments- The holotype was discovered in 2005. Arbour et al. (2015) reidentified
the three supposed furcula (one in the holotype, KUVP 152429 and NCSM 13170)
as entoplastra of the trionychid turtle cf. Axestemys splendida.
DePalma et al. (2015) found it to be a dromaeosaurine sister to Dromaeosaurus
using a version of the TWiG analysis. However, Cau (online, 2015) recovered it
as sister to Velociraptorinae+Dromaeosaurinae in his larger unpublished analysis,
and my own unpublished analysis agrees it is outside that clade.
References- Arbour, Zanno, Larson, Evans and Sues, 2015. The furculae
of the dromaeosaurid dinosaur Dakotaraptor steini are trionychid turtle
entoplastra. PeerJ PrePrints. https://doi.org/10.7287/peerj.preprints.1570v1
Cau, online 2015. http://theropoda.blogspot.com/2015/10/dakotaraptor-un-acheroraptor-gigante.html
DePalma, Burnham, Martin, Larson and Bakker, 2015. The first giant raptor (Theropoda:
Dromaeosauridae) from the Hell Creek Formation. Paleontological Contributions.
14, 16 pp.
Dineobellator Jasinski, Sullivan and Dodson, 2020
= "Dineobellator" Jasinski, 2018
D. notohesperis Jasinski, Sullivan and Dodson, 2020
= "Dineobellator notohesperis" Jasinski, 2018
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Holotype-
(SMP VP-2430) premaxillary fragment, maxillary fragment, ?maxillary
tooth (11.3x7.78x4.15 mm), ?nasal fragment, lacrimal fragment, jugal
fragment, partial basisphenoid, occipital condyle, several cranial
fragments, partial dorsal rib, several rib fragments, first caudal
vertebra (26.4 mm), ~tenth caudal vertebra (32.56 mm), several
vertebral fragments, incomplete humerus (~215 mm), incomplete ulna
(~140 mm), incomplete manual ungual ?II (~50 mm), proximal metacarpal
III, incomplete manual ungual III (~55 mm), incomplete femur (~275 mm),
partial ?astragalus, incomplete metatarsal I, incomplete metatarsal II,
proximal metatarsal III, fragments
Diagnosis- (after Jasinski et al., 2020) short and robust neural spine
on proximal caudal vertebrae; sharp angle of distal deltopectoral crest of humerus (also in Dakotaraptor);
vertical offset of side grooves on manual ungual II; dorsodistally
oriented groove just distal to articular surface on medial side of
manual ungual II; tall and distally wide flexor tubercle on manual
ungual II.
(suggested) accessory ventral side grooves on manual ungual III.
Other diagnoses- Jasinski et al. describe the first caudal as opisthocoelous but state
"the anterior surface is flat, the posterior is concave", making it
platycoelous as is common in dromaeosaurids (e.g. Deinonychus, Adasaurus). They list "proximal caudal vertebrae
with curved ventral surface", but this is equally true of Bambiraptor, Velociraptor, Deinonychus, Adasaurus, IGM 100/22 and Yurgovuchia.
Similarly, 'proximal caudal vertebrae
with oval to subrectangular anterior and
posterior centrum surfaces' describes all dromaeosaurids. The proximal
caudal transverse processes are "gracile and subrectangular" in Bambiraptor, Deinonychus, Adasaurus and IGM 100/22 as well, but are actually broken distally in Dineobellator
(Jasinski et al., 2011- fig. 10C-D) so that their shape is
indeterminate. 'Distinct round concavities on anterior
and posterior centrum surfaces in mid caudal vertebrae' are on rarely
exposed surfaces in dromaeosaurid tails, but a similar structure is
present at least posteriorly in Deinonychus, and anteriorly in Utahraptor.
The supposed enlarged flexor tubercle on pedal ungual III is actually
manual ungual III, and not larger than in other dromaeosaurids. This
also affects their character of a secondary side groove on pedal ungual
III, which is instead found in manual ungual III.
Comments- Discovered in 2008, Jasinski et al.
(2011) briefly describe SMP VP-2430 as Dromaeosauridae indet., noting
it "probably represents a new taxon and will be described in detail
elsewhere." The dorsal vertebra they note and figure seems to be the
centrum of the first caudal vertebra. Jasinoski et al. (2015) mention
it as "a specimen with both cranial and postcranial material [that] is
known from the Maastrichtian of New Mexico, probably representing a
distinct taxon, and is by far the most complete dromaeosaurid material
recovered from Late Cretaceous strata in southern Laramidia."Jasinski
(2018) described the specimen in his thesis as Dineobellator notohesperis,
which is a nomen nudum as theses are generally considered not "issued
for the purpose of providing a public and permanent scientific record"
(ICZN Areticle 8.1.1). He used Longrich and Currie's dromaeosaurid
analysis to recover it as a eudromaeosaur closer to dromaeosaurines and
velociraptorines than to saurornitholestiines. The taxon was
officially named and described by Jasinski et al. (2020) who used the
same analysis to recover it as a velociraptorine sister to Tsaagan sensu lato. Adding it to the Hartman et al. maniraptoromorph analysis results in it being an unenlagiid close to Dakotaraptor with 2 steps and 3 steps needed respectively to move it to the positions of Jasinski's thesis and eventual description.
The left lacrimal fragment is supp. fig. 2A would seem to be in medial
view based on comparison with Saurornitholestes, making the labeled
lacrimal recess either autapomorphic or taphonomic. Jasinoski et al.
describe an articulated string of four vertebrae as fused distal
caudals, but they are about half as long as tall (43-54%) unlike
theropod distal caudals which are always much longer than tall. For a
comparison, their centra are 75% as tall as the anterior face of the
first caudal vertebra but 16-19% of its length, whereas in Deinonychus
the caudal 75% as tall as the first (~caudal 15) is 143% its length.
Instead, these small (4.12, 5.10 mm) and tall centra resemble
actinopterygians, in both size and the presence of foramina and
anteroposterior ridges which the authors attributed to prezygapophyseal
rods. Multiple fish are present in the Alamo Wash local fauna
including lepisosteids, although amiid and esocid vertebrae are also
common in similar formations. The supposed pedal ungual III is not
only far more curved than non-avialan pedal unguals (except
deinonychosaur pedal unguals II), its dorsal margin arches far dorsal
to the articular surface, which Senter described as distinguishing
dromaeosaurid manual unguals from pedal unguals. Its articular surface
is much smaller (59% as tall, 49% as wide) than the other ungual,
suggesting it is from manual digit III.
References-
Jasinski, Sullivan and Lucas, 2011. Taxonomic composition of the Alamo Wash
local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member),
San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann (eds.). Fossil
Record 3. New Mexico Museum of Natural History and Science Bulletin. 53, 216-271.
Jasinski, Sullivan and Dodson, 2015. Late Cretaceous dromaeosaurid theropod
dinosaurs (Dinosauria: Dromaeosauridae) from southern Laramidia and implications
for dinosaur faunal provinciality in North America. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 150.
Jasinski, 2018. The integration of morphology, variation, and
phylogenetics to better understand fossil taxa and their modern
relatives. PhD thesis, University of Pennsylvania. 564 pp.
Jasinski, Sullivan and Dodson, 2020. New dromaeosaurid dinosaur
(Theropoda, Dromaeosauridae) from New Mexico and biodiversity of
dromaeosaurids at the end of the Cretaceous. Scientific Reports.
10:5105.
Neuquenraptor Novas and
Pol, 2005
= "Araucanoraptor" Novas vide anonymous, 1997
N. argentinus Novas and Pol, 2005
= "Araucanoraptor argentinus" Novas vide anonymous, 1997
Late Turonian-Early Coniacian, Late Cretaceous
Portezuelo Formation of the Rio Neuquen Subgroup, Neuquen, Argentina
Holotype- (MCF PVPH 77) (~2 m) fragmentary cervical vertebra, dorsal
ribs, chevrons, proximal radius, incomplete femur (~250 mm), distal tibia, distal
fibula, incomplete astragalus, calcaneum, metatarsal I (30.4 mm), phalanx I-1
(27 mm), pedal ungual I (26.9 mm straight), incomplete metatarsal II (~147.9
mm), phalanx II-1 (39.1 mm), phalanx II-2 (36.2 mm), pedal ungual II (39.7 mm),
incomplete metatarsal III (~172.7 mm), phalanx III-1 (57.4 mm), phalanx III-2
(36.6 mm), phalanx III-3 (34.4 mm), incomplete metatarsal IV (~149.6 mm), phalanx
IV-2 (32.3 mm), phalanx IV-3 (25.4 mm), phalanx IV-4 (25.5 mm), pedal ungual
IV (21.7 mm)
Other diagnoses- All of the characters listed in Novas and Pol's (2005)
diagnosis are actually more widely distributed. A subarctometatarsus is present
in most basal paravians. Metatarsal II laterally expands over metatarsal III
posteriorly in Buitreraptor and Pamparaptor. An extensor sulcus
on metatarsal III is also present in Buitreraptor, troodontids and microraptorians.
Metatarsal III is also incipiently ginglymoid in Buitreraptor, Microraptor
and several other paravians. Subequal pedal phalanges II-1 and II-2 are also
present in several paravians including Adasaurus, Archaeopteryx,
Microraptor, NGMC 91, Bambiraptor and Velociraptor. A strongly
curved pedal ungual II is found in most dromaeosaurids and basal avialans.
Comments- This was discovered in 1996 and originally reported as a troodontid
until its official description identified it as a dromaeosaurid. Novas (pers.
comm., 2005) confirms that it is the same specimen as "Araucanoraptor".
Gianechini and Apesteguia (2011) illustrate the proximal ungual originally placed
as ungual I by Novas and Pol (2005) as ungual IV. The distal ungual originally
placed as IV may be I or III, but is not illustrated by Gianechini and Apesteguia.
Makovicky et al. (2005) suggested that Neuquenraptor and Unenlagia
may be synonymous based on supposedly identical femur (U. comahuensis),
pedal phalanx II-1 and pedal ungual II (U. paynemili). See the comments
under Unenlagia for details, but Neuquenraptor seems distinct
from U. comahuensis, but cannot be compared to U. paynemili, except
for the possibly referrable phalanx II-1, which differ between taxa. Though
usually referred to Unenlagiinae, without the addition of information from Unenlagia,
Neuquenraptor can be placed in any of several basal paravian positions
outside derived dromaeosaurids, derived troodontids and derived avialans.
Porfiri et al. (2007) reported another specimen of Neuquenraptor (MUCPv-1163),
but this has since been made the holotype of Pamparaptor by Porfiri et
al. (2011).
References- Novas, Cladera and Puerta, 1996. New theropods from the Late
Cretaceous of Patagonia. Journal of Vertebrate Paleontology. 16(3), 56A.
Anonymous, 1997. Clarin (Argentine newspaper). 7-23-97.
Novas, Apesteguia, Pol and Cambiaso, 1999. Un probable Troodontido (Theropoda
- Coelurosauria) del Cretacico Tardio de Patagonia. XV Jornadas Argentinas de
Paleontologia de Vertebrados. Ameghiniana. 36(S), 17R.
Novas and Pol, 2005. New evidence on deinonychosaurian dinosaurs from the Late
Cretaceous of Patagonia. Nature. 433, 858-861.
Porfiri and Calvo, 2007. La validez taxon�mica de Neuquenraptor argentinus
(Theropoda, Dromaeosauridae) y la monofilia del nodo Unenlagiinae. Ameghiniana.
44(S), 34R.
Porfiri, Calvo, dos Santos and Valieri, 2007. New record of Neuquenraptor
(Theropoda, Dromaeosauridae) from the Late Cretaceous of Patagonia. Ameghiniana.
44(S), 34R.
Gianechini and Apesteguiia, 2011. Unenlagiinae revisited: Dromaeosaurid theropods
from South America. Anais da Academia Brasileira de Ci�ncias. 83(1), 163-195.
Porfiri, Calvo and dos Santos, 2011. A new small deinonychosaur (Dinosauria:
Theropoda) from the Late Cretaceous of Patagonia, Argentina. Anais da Academia
Brasileira de Ci�ncias. 83(1), 109-116.
Dromaeosauridae Matthew and Brown, 1922
sensu Russell, 1969
Definition- (Dromaeosaurus albertensis <- Unenlagia comahuensis, Troodon formosus, Archaeopteryx lithographica, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019)
Other definitions- (Dromaeosaurus albertensis + Velociraptor
mongoliensis) (modified from Padian et al., 1999)
(Microraptor zhaoianus + Sinornithosaurus millenii + Velociraptor
mongoliensis) (Norell and Makovicky, 2004)
(Deinonychus antirrhopus <- Troodon formosus, Passer domesticus)
(modified from Hu, Hou, Zhang and Xu, 2009)
(Dromaeosaurus albertensis <- Troodon formosus,
Passer domesticus) (Turner et al., 2012)
(Dromaeosaurus albertensis <- Saurornithoides mongoliensis, Vultur gryphus) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
(Dromaeosaurus albertensis <- Ornithomimus edmontonicus, Troodon
formosus, Passer domesticus) (Sereno, in press)
= Itemiridae Kurzanov, 1976
Definition- (Itemirus medullaris <- Dromaeosaurus albertensis,
Stenonychosaurus inequalis, Tyrannosaurus rex) (Martyniuk, 2012)
= Dromaeosauridae sensu Sereno, 1998
Definition- (Velociraptor mongoliensis <- Troodon formosus)
(modified from Sereno, 1998)
= Avenychia Miller, 2004
= Enantiavenychia Miller, 2004
= "Tetrapteryxidae" Miller, 2004
= Dromaeosauroidea Matthew and Brown, 1922 sensu Livezey and Zusi, 2007
= Dromaeosauridae sensu Hu, Hou, Zhang and Xu, 2009
Definition- (Deinonychus antirrhopus <- Troodon formosus, Passer
domesticus)
= Dromaeosauridae sensu
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Dromaeosaurus albertensis <- Saurornithoides mongoliensis, Vultur gryphus)
= Dromaeosauridae sensu Sereno, in press
Definition- (Dromaeosaurus albertensis <- Ornithomimus edmontonicus,
Troodon formosus, Passer domesticus)
Comments- Sereno's new (in press) definition modifies his earlier (1998)
one by using the eponymous genus as an internal specifier, and adding Ornithomimus
and Passer as external specifiers. This contrasts with the two published
node-based definitions- (Dromaeosaurus albertensis + Velociraptor
mongoliensis) by Padian et al. (1999) and (Microraptor zhaoianus
+ Sinornithosaurus millenii + Velociraptor mongoliensis)
by Norell et al. (2004). Norell et al.'s is problematic for not including Dromaeosaurus,
and for explicitly including Microraptor. The latter would make pygostylians
dromaeosaurids in Mayr et al.'s (2005) and Agnolin and Novas' (2013) topology.
Padian et al.'s is equivalent to the use of Dromaeosauridae prior to 1999, but
microraptorians weren't known and 'unenlagiines' were considered avialans. Once
microraptorians were discovered, they were assigned to Dromaeosauridae by most
authors except Senter et al. (2004). So using a definition which won't probably
exclude them does have some precedence, but Sereno's should have more external
specifiers if it is adopted, namely Archaeopteryx (Paul, 1988; 2002),
Tyrannosaurus (Matthew and Brown, 1922; Russell and Dong, 1994), Ornitholestes
(Makovicky, 1995) and Oviraptor (Barsbold et al., 1990). I actually think
this stem-based definition is better than Padian et al.'s node-based one for
the additional reason that exactly which taxa fall into the (Dromaeosaurus
+ Velociraptor) crown is uncertain. With the recognition that velociraptorine
sensu lato characters are symplesiomorphic for dromaeosaurs (e.g. found in Bambiraptor
and microraptorians), taxa like Deinonychus or Saurornitholestes
could easily fall just outside the defined node, not to mention more fragmentary
taxa such as Variraptor or Pyroraptor. So I advocate the following
redefinition of Sereno's- (Dromaeosaurus albertensis <- Tyrannosaurus
rex, Ornithomimus velox, Oviraptor philoceratops, Troodon formosus, Archaeopteryx
lithographica, Passer domesticus).
Kurzanov (1976) erected Itemiridae for Itemirus only, sister to tyrannosaurids
than to dromaeosaurids. Martynuik (2012) defined Itemiridae as excluding Dromaeosaurus,
but since the ICZN states families cannot contain other families, Itemiridae
is listed as a synonym of Dromaeosauridae here.
Miller (2004) stated "I would ascribe Microraptor gui, and possibly Cryptovolans, to its own family, the Tetrapteryxidae", but Tetrapteryx
is a gruid genus whose name cannot be used for a family of
dromaeosaurs, so Miller's taxon is invalid. Miller also proposed
Avenychia for "The most inclusive dromaeosaurid clade, including the
Tetrapteryxidae* plus all later dromaeosaurid genera", which would make
it a junior synonym of Dromaeosauridae here. Similarly, Miller
proposes Enantiavenychia for the clade including Microraptor, Sinornithosaurus, Bambiraptor and eudromaeosaurs but not "tetrapteryxids", which is not recognized here as Cryptovolans and gui are synonymized with Microraptor zhaoianus.
Not dromaeosaurids- Teeth from the Albian Eumeralla Formation and Aptian
Wonthoggi Formation of Victoria, Australia have been commonly referred to Dromaeosauridae
(Rich and Vickers-Rich, 1994), but are retained as Coelurosauria incertae sedis
here as the lack of mesial serrations is known in many other coelurosaurs (e.g.
Orkoraptor, compsognathids, most troodontids).
A partial braincase referred to as possibly dromaeosaurid (CCMGE 466/12457)
by Nessov (1995) was referred to Urbacodon sp. by Averianov and Sues
(2007).
A partial pedal phalanx II-2 (IGM-7715) was tentatively referred to Dromaeosauridae
by Rodriguez de la Rosa and Cevallos-Ferriz (1998) based on the dorsally placed
collateral ligament pit. However, this character is also present in basal troodontids
(e.g. IGM 100/44, Sinornithoides, Borogovia), Neuquenraptor
and Rahonavis. It is here referred to Paraves incertae sedis.
Tavares et al. (2014) described two teeth from the Adamantina Formation of Brazil
as dromaeosaurid, but these are here considered more likely to be abelisaurid.
References- Kurzanov, 1976. Braincase structure in the carnosaur Itemirus n. gen.,
and some aspects of the cranial anatomy of dinosaurs. Paleontological Journal.
1976, 361-369.
Rich and Vickers-Rich, 1994. Digs at Dinosaur Cove and Flat Rocks 1994. Excavation
Report. Dinosaur Cove 1993 - 1994 & Inverloch 1994. 10-13.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
Rodriguez de la Rosa and Cevallos-Ferriz, 1998. Vertebrates of the El Pelillal
locality (Campanian, Cerro del Pueblo Formation), Southeastern Coahuila, Mexico.
Journal of Vertebrate Paleontology. 18, 751-764.
Norell, Makovicky and Clark, 2000. A Review of the Dromaeosauridae. The Florida
Symposium on Dinosaur Bird Evolution. Publications in Paleontology No.2, Graves
Museum of Archaeology and Natural History 20.
Burnham, Senter, Barsbold and Britt, 2004. Phylogeny of the Dromaeosauridae.
Journal of Vertebrate Paleontology. 24(3), 204A-205A.
Miller, 2004. A new phylogeny of the Dromaeosauridae. 2004 Student Showcase Journal. 20, 123-158.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
Livezey and Zusi, 2007. Higher-order phylogeny of modern birds (Theropoda, Aves:
Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological
Journal of the Linnean Society. 149 (1), 1-95.
Manning, Ali, McDonald, Mummery and Sellers, 2007. Biomechanics of dromaeosaurid
claws: Application of x-ray microtomography, nanoidentification and finite element
analysis. Journal of Vertebrate Paleontology. 27(3), 111A-112A.
Hoffman and Hwang, 2009. Identifying diagnostic characters in the tooth enamel
microstructure of dromaeosaurid dinosaurs. Journal of Vertebrate Paleontology.
29(3), 115A.
Longrich and Currie, 2009. A microraptorine (Dinosauria - Dromaeosauridae) from
the Late Cretaceous of North America. Proceedings of the National Academy of
Sciences. 106(13), 5002-5007.
Manning, Margetts, Johnson, Mustansar and Mummery, 2009. A finite approach to
the biomechanics of dromaeosaurid dinosaur claws. Journal of Vertebrate Paleontology.
29(3), 141A.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Pan Aves. 189 pp.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic relationships
of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae.
Springer. 96 pp.
Tavares, Branco and Santucci, 2014. Theropod teeth from the Adamantina Formation
(Bauru Group, Upper Cretaceous), Monte Alto, S�o Paulo, Brazil. Cretaceous
Research. 50, 59-71.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
"Velociraptor" osmolskae Godefroit, Currie,
Li, Shang and Dong, 2008
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype- (IMM 99NM-BYM-3/3) incomplete maxillae, lacrimal
Diagnosis- (after Godefroit et al., 2008) long anterior ramus of maxilla,
with elongation index (L/H ratio) 1.38; promaxillary fenestra subequal in size
to maxillary fenestra and teardrop-shaped; long axis of promaxillary fenestra
perpendicular to dorsal border of the maxilla; long axis of maxillary fenestra
parallel to this border; ten maxillary teeth with short unserrated carina on
the apical end of the mesial edge and with incipient serrations on the distal
carina.
Comments- Note this species' inclusion in Velociraptor has never been tested in
a phylogenetic analysis, and that some characters such as the lack of mesial
serrations are more similar to basal dromaeosaurids.
References- Godefroit, Currie, Li, Shang and Dong, 2008. A new species of Velociraptor
(Dinosauria: Dromaeosauridae) from the Upper Cretaceous of northern China. Journal
of Vertebrate Paleontology. 28(2), 432-438.
Acheroraptor Evans, Larson
and Currie, 2013
A. temertyorum Evans, Larson and Currie, 2013
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Holotype- (ROM 63777) incomplete maxilla, maxillary tooth
Referred- ?...(ROM 63778) incomplete dentary (Evans, Larson and Currie,
2013)
Diagnosis- (after Evans et al., 2013) relatively small anterior region
of antorbital fossa (defined as the region between the anterior borders of the
antorbital fenestra and the antorbital fossa); maxillary fenestra that almost
reaches ventralmargin of antorbital fossa and is positioned directly posterior
to promaxillary fenestra; extensive, posteriorly projected postantral wall that
is visible laterally through antorbital fenestra (also in Austroraptor);
maxillary dentition with unique configuration of prominent apicobasal ridges
on both labial and lingual surfaces of tooth.
Comments- Powers et al. (2022) described maxillary characters in detail using CT scans.
Evans et al. (2013) entered Acheroraptor into a modified version
of Longrich and Currie's dromaeosaurid analysis and found it to be a basal velociraptorine.
References- Evans, Larson and Currie, 2013. A new dromaeosaurid (Dinosauria:
Theropoda) with Asian affinities from the latest Cretaceous of North America.
Naturwissenschaften. 100(11), 1041-1049.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as
2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist
in testing and constructing morphological characters. Journal of
Vertebrate Paleontology. 41(5), 2010087.
Boreonykus Bell and Currie,
2015
B. certekorum Bell and Currie, 2015
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Holotype- (RTMP 89.55.47) incomplete frontal (52.4 mm)
Diagnosis- (after Bell and Currie, 2015) supratemporal ridge on frontals
forming acute angle (55 degrees) anteriorly.
Combination of- frontal elongate and slender; low supratemporal ridge forming
arc that is concave relative to interfrontal suture in dorsal view; area immediately
posterior to supratemporal ridge smooth and posteroventrally sloping.
Comments- Ryan and Russell (2001) listed the holotype frontal as "Saurornitholestes
undescribed n. sp. (Currie pers. comm.)". Bell and Currie (2015) described
this and other material (distal caudal RTMP 88.55.129, manual ungual UALVP 53597,
and pedal ungual II RTMP 86.55.184) from the Pipestone Creek Pachyrhinosaurus
lakustai bonebed as Boreonykus certekorum, noting that preserved
elements are not duplicated and are properly proportioned to belong to one individual.
The fourteen referred teeth are shed so were admitted to at least belong to
another individual. When entered in Longrich and Currie's dromaeosaurid analysis,
it emerged as a velociraptorine if the teeth were included, and a member of
Velociraptorinae+Dromaeosaurinae if they were excluded. However, Cau (online,
2015) noted that given the paucity of remains and number of small Judithian
theropod taxa, it should not be assumed that the frontal belongs to the same
individual as the dromaeosaurid postcrania. Indeed, he reported that when the
frontal is entered into his theropod supermatrix, it emerges in several position
within Maniraptoriformes but never as a dromaeosaurid. My own analysis finds
it to be a coelurosaur excluded from ornithomimosaurs, alvarezsauroids, therizinosaurs,
oviraptorosaurs, unenlagiines, microraptorians, dromaeosaurines, derived troodontids
and birds. As it is clearly not a member of contemporaneous clades known from
frontals, Boreonykus may belong to one of the problematic taxa such as
Richardoestesia or Paronychodon which are both placed in Paraves
here. If I'm correct and Richardoestesia is a microraptorian, Boreonykus
may be Paronychodon.
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press.
279-297.
Bell and Currie, 2015. A high-latitude dromaeosaurid, Boreonykus certekorum,
gen. et sp. nov. (Theropoda), from the upper Campanian Wapiti Formation, west-central
Alberta. Journal of Vertebrate Paleontology. e1034359. DOI: 10.1080/02724634.2015.1034359.
Cau, online 2015. http://theropoda.blogspot.com/2015/12/boreonykus-e-un-dromaeosauridae.html
"Julieraptor" Thompson vide Robbins, 2009
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Material-
(ROM coll.; Sid Vicious) non-jaw cranial elements, cervical series,
dorsal series, dorsal ribs, incomplete caudal series, distal chevrons,
pectoral girdles, forelimbs (missing four manual elements), ilia,
pubes, hindlimbs (missing two pedal elements)
Comments- As detailed by Bakker
(online 2012) and Robbins (online 2009), material exposed on the
surface (cranial elements, manual phalanges and unguals) were collected
in 2002 and nicknamed "Julieraptor" by Thompson, while the rest of the
specimen was excavated by 2006 and nicknamed Sid Vicious by Murphy of
the Judith River Dinosaur Institute. Once reuninted and after various
legal issues "the skeleton was returned to the land owner, who arranged
to sell the specimen to the Royal Ontario Museum" (Bakker, online
2012).
Described in BHI's website as having "many differences from another significant small raptor, Bambiraptor;
primarily in the ilia, scapula, arms, hands, feet and size (about 35%
larger than Bambiraptor)."
Whether these differences prove convincing and whether "Julieraptor"
can be distinguished from other contemporaneous dromaeosaurids such as Saurornitholestes and Dromaeosaurus remains to be seen.
References- Robbins, 2009. Dinosaur fossil faces charges of theft. The New York Times. January 21.
Bakker, online 2012. http://blog.hmns.org/2012/06/bakker-blogs-the-kleptomania-continues-with-a-sid-vicious-julieraptor-dino-rustlers-part-ii/
BHI online, 2012. http://www.bhigr.com/store/product.php?productid=679
Kansaignathus Averianov and Lopatin, 2021a
K. sogdianus Averianov and Lopatin, 2021a
Early Santonian, Late Cretaceous
Kansai, Yalovach Formation, Tajikistan
Holotype- (PIN 2398/15) (~2 m) dentary (140 mm)
Referred- ?(PIN 2398/4) manual ungual II (Averianov and Lopatin, 2021)
?(PIN 3041/11) anterior dorsal vertebra (Alifanov and Averianov,
2006)
?(PIN 3041/45) anterior tooth (Averianov and Lopatin, 2021)
Early Santonian, Late Cretaceous
Kyzylpilyal, Yalovach Formation, Tajikistan
? teeth (Nessov, Kaznyshkina and Cherepanov, 1987)
Diagnosis- (after Averianov and
Lopatin, 2021) twelve dentary teeth; differs from ?Itemirus ZIN PH
2338/16 in being straight in dorsal view and lacking vertical grooves
in its interdental plates; differs from Dromaeosaurus and Deinonychus
in being more ventrally convex and having chin; differs from
Saurornitholestes and Linheraptor in lacking ventral curvature at the
symphysis; differs from Atrociraptor in being more concave dorsally and
lacking grooves between interdental plates; differs from Velociraptor
in being taller, lacking ventral curvature at the symphysis, and
lacking a paradental groove medially; differs from Acheroraptor in
lesser development of lateral foramina.
(suggested) differs from Bambiraptor in lacking ventral curvature at symphysis.
Other diagnoses- Averianov and Lopatin (2021b) claim Kansaignathus differs from Bambiraptor
"in the presence of the ventral row of vascular foramina in the middle
of the dentary and a smaller difference in the depth between the
anterior and posterior parts of the dentary", but Bambiraptor also has the ventral
row of foramina in the middle and is not notably different in anterior
taper (pers. obs.). They also claim it differs from Velociraptor
in having the dorsal row of lateral foramina be closer to the alveolar
edge, but this doesn't appear to be true. The authors also say it
differs from Tsaagan "in the absence of a common groove for the dorsal row of vascular foramina on the labial side of the dentary", but the grooves in both taxa are not
continuous.
Comments- The holotype dentary was discovered in 1963-1964
and described by Averianov and Lopatin (2021a) as a new taxon of
dromaeosaurid. When they entered it into Currie and Longrich's
dromaeosaurid matrix it emerged as a velociraptorine closer to Velociraptor than Deinonychus but further than Acheroraptor.
Nessov (1995) noted (translated) "dromaeosaurids (D. A. Russell,
personal
communication, 1988)" from Kansai. Alifanov and Averianov (2006)
stated that at Kansai "The family Dromaeosauridae is probably
represented by a corpus of a relatively short anterior thoracic
vertebra with a stout hypapophysis (PIN, no. 3041/11)." Averianov and Loptain (2021b) stated the vertebra may belong to Kansaignathus,
as well as "a tooth with considerably lingually displaced carina
(specimen PIN, no. 3041/45), ... and an ungual phalanx of manual finger
II (specimen PIN, no. 2398/4)." Nessov et
al.
(1987) had earlier reported deinonychosaurian teeth from a different
locality. While these materials could easily belong to other
dromaeosaurid taxa given the presence of multiple genera in well
sampled formations, they are listed here for convenience.
References- Nessov, Kaznyshkina and Cherepanov, 1987. Dinosaurs, crocodiles,
and other archosaurs of the late Mesozoic of Middle Asia and their place in
ecosystems. Abstracts of the XXXII Session of the All-Union Paleontological Society. 46-47.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology
and paleobiogeography. Scientific Research Institute of the Earth's Crust. 156 pp.
Alifanov and Averianov, 2006. On the finding of ornithomimid dinosaurs (Saurischia,
Ornithomimosauria) in the Upper Cretaceous beds of Tajikistan. Paleontological
Journal. 40(1),103-108.
Averianov and Lopatin, 2021a. A new carnivorous dinosaur (Theropoda,
Dromaeosauridae) from the Late Cretaceous of Tajikistan. Reports of the
Russian Academy of Sciences. Earth Sciences. 499(1), 49-53.
Averianov and Lopatin, 2021b. A new theropod dinosaur (Theropoda,
Dromaeosauridae) from the Late Cretaceous of Tajikistan. Doklady Earth
Sciences. 499(1), 570-574.
"Kitadanisaurus" Lambert, 1990
Middle-Late Aptian, Early Cretaceous
Kitadani Formation of the Akaiwa Subgroup of the Tetori Group, Japan
Material- (Kitadani-ryu) tooth (Dong et al., 1990)
?(FPDM-V96082619) dorsal vertebra (Azuma and Currie, 2000)
?(FPDM-V9812638, 96072901, 97082906, 980815181, 98092604) five teeth (Azuma
and Currie, 2000)
?(FPDM-V98081115) humerus (144 mm) (Currie and Azuma, 2006)
?(FPDM-V980801141) manual ungual (Currie and Azuma, 2006)
? several unguals (Currie and Azuma, 2006)
Comments- The original tooth was informally called "Kitadani-ryu",
as found in Azuma (1991). Lambert (1990) inappropriately made it into a genus
name, listing it as "Kitadanisaurus" in a childrens' book. Dong et
al. (1990) published photos of the tooth, labeling it Dromaeosauridae indet..
In 1991, jaw fragments were found in the quarry and identified as dromaeosaurid
based on their fused interdental plates. This was followed by the discovery
of a manual ungual I, astragalus and metatarsal III in 1993. Azuma and Currie
(1995) described these remains in an abstract as those of a giant dromaeosaurid,
which was associated in the paleontological community with "Kitadanisaurus"
through the late 1990's. Azuma and Currie (2000) later described the skeletal
remains as a new taxon of carnosaur, Fukuiraptor. Yet they also noted
smaller teeth are known, similar in size and shape to dromaeosaurids and sharing
Dromaeosaurus' mesial carina twist, as well as a dorsal vertebra "suggestive
of dromaeosaurid affinities." Currie and Azuma (2006) list the teeth which
are not referrable to Fukuiraptor, and describe and illustrate a humerus
they refer to Dromaeosauridae indet.. In addition, they state several unguals
seem to be dromaeosaurid, though these remain undescribed. One manual ungual
is listed as having a proximodorsal lip, which is present in most dromaeosaurids
as well as some other coelurosaurs. It is probably one of the several, though
the only other listed ungual is a distal tip (FPDM-V9912141). Even if these
remains are all dromaeosaurid, they do not necessarily belong to the same taxon.
"Kitadanisaurus" remains a nomen nudum, as the original tooth has
never been described.
References- Dong, Hasegawa and Azuma, 1990. The Age of Dinosaurs in Japan
and China. Fukui, Japan: Fukui Prefectural Museum. 65 pp.
Lambert, 1990. The Dinosaur Data Book. New York: Avon Books, 66. ISBN 0-380-75896-3.
Azuma, 1991. Early Cretaceous Dinosaur Fauna from the Tetori Group, central
Japan. Research on Dinosaurs from the Tetori Group (1). Professor S. Miura Memorial
Volume, 55-69.
Azuma and Currie, 1995. A new giant dromaeosaurid from Japan. Journal of Vertebrate
Paleontology. 15(3), 17A.
Azuma and Currie, 2000. A new carnosaur (Dinosauria: Theropoda) from the Lower
Cretaceous of Japan. Canadian Journal of Earth Sciences. 37(12), 1735-1753.
Currie and Azuma, 2006. New specimens, including a growth series of Fukuiraptor
(Dinosauria, Theropoda) from the Lower Cretaceous Kitadani Quarry of Japan.
J. Paleont. Soc. Korea. 22(1), 173-193.
undescribed Dromaeosauridae (Chiba, Ryan, Braman, Eberth, Scott, Brown,
Kobayashi and Evans, 2015)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
Material- (RTMP coll.) two teeth
Reference- Chiba, Ryan, Braman, Eberth, Scott, Brown, Kobayashi and Evans,
2015. Taphonomy of a monodominant Centrosaurus apertus (Dinosauria: Ceratopsia)
bonebed from the upper Oldman Formation of southeastern Alberta. Palaios. 30,
655-667.
probable Dromaeosauridae indet. (Currie, Rigby and Sloan, 1990)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
Material- (AMNH 5387) manual ungual (Gilmore, 1920)
?(AMNH coll.) three manual unguals (Gilmore, 1920)
(CMN 12413) presacral vertebra (Russell, 1969)
(CMN 12415) presacral vertebra (Russell, 1969)
?(CMN 12438) sacrum (Russell, 1969)
?(CMN 12439) sacrum (Russell, 1969)
(RTMP 80.13.34) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.180) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.20.255) tooth (Currie, Rigby and Sloan, 1990)
(UA 5286) presacral vertebra
Comments- Gilmore (1920) noted four unguals from the Belly River Group
of Canada in the AMNH (including AMNH 5387) were nearly identical to the holotype
of Coelurus gracilis. Gilmore (1924) believed they belonged to Chirostenotes,
but at least AMNH 5387 more closely resembles derived dromaeosaurids in being
more curved, having no proximodorsal lip, a more proximally placed flexor tubercle
and a proximally undivided longitudinal groove. It is likely Saurornitholestes
or Dromaeosaurus, and the other unguals may be as well.
Russell (1969) listed the presacral vertebrae (CMN 12413 and 12414, UA 5286)
as being comparable to Oviraptor in their short centra and pleurocoels.
The sacra (CMN 12438 and 12439) were tentatively referred to the same taxon,
and described as being strongly concave ventrally and containing five vertebrae.
The latter resembles dromaeosaurids more closely (oviraptorids have at least
six sacrals), many of which also share the presacral characters listed above.
They probably belong to Richardoestesia, Dromaeosaurus or Saurornitholestes
based on provenance. Another possibility is that they are from juvenile tyrannosaurids.
The three RTMP teeth have a chalky appearence and lack serrations due to being
swallowed and digested. They are thus difficult to assign to particular genera,
but are near certainly coelurosaurs, and probably dromaeosaurids based on provenence
and elongate shape.
References- Gilmore, 1920. Osteology of the carnivorous Dinosauria in
the United States National Museum with special reference to the genera Antrodemus
(Allosaurus) and Ceratosaurus. United States National Museum Bulletin.
110, l-154.
Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta.
Canada Geological Survey, Bulletin 38, geological series 43, 1-13.
Russell, 1969. A new specimen of Stenonychosaurus from the Oldman Formation
(Cretaceous) of Alberta. Canadian Journal of Earth Science. 6, 595-612.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.
undescribed Dromaeosauridae (Chapman, Deck, Varricchio and Jackson,
2004)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2240/45084; Morph 4) tooth (?x4x2.6 mm) (Krumenacker et
al., 2016)
(IMNH 2251/49826; Morph 4) tooth (8.4x4.2x2.5 mm) (Krumenacker et al., 2016)
(IMNH 2251/50034; Morph 4) tooth (~9.4x~10.6x~5.2 mm) (Krumenacker et al., 2016)
(IMNH 2251/50832; Morph 4) tooth (8.9x5.3x~2.5 mm) (Krumenacker et al., 2016)
(IMNH 2251/50849; Morph 4) tooth (~3x~1.9x? mm) (Krumenacker et al., 2016)
(IMNH 2167/50104; Morph 4) tooth (4.1x2.1x1 mm) (Krumenacker et al., 2016)
caudal vertebrae (Krumenacker, 2005)
material (Chapman, Deck, Varricchio and Jackson, 2004)
Comments- Stated to have a low HCR and high amount of labiolingual compression.
References- Chapman, Deck, Varricchio and Jackson, 2004. Cretaceous Wayan
Formation of Idaho: A preliminary report. 24(3), 151-152.
Krumenacker, 2005. Preliminary report on new vertebrate fossils from the Draney
Limestone (Aptian) and Wayan Formation (Albian) of east Idaho. Journal of Vertebrate
Paleontology. 25(3), 80A.
Krumenacker and Scofield, 2015. A diverse theropod tooth assemblage from the
Mid-Cretaceous (Albian-Cenomanian) Wayan Formation of Idaho. Journal of Vertebrate
Paleontology. Program and Abstracts 2015, 158.
Krumenacker, Simon, Scofield and Varricchio, 2016. Theropod dinosaurs from the
Albian-Cenomanian Wayan Formation of eastern Idaho. Historical Biology. DOI:
10.1080/08912963.2015.1137913
undescribed possible dromaeosaurid (Turner and Peterson, 1999)
Late Oxfordian-Tithonian, Late Jurassic
Mother's Day Quarry, Morrison Formation, Montana, US
Material- tooth
Comments- Turner and Patterson reference a dromaeosaur tooth from this
undescribed quarry.
Reference- Turner and Peterson, 1999. Biostratigraphy of dinosaurs in
the Upper Jurassic Morrison Formation of the Western Interior U.S.A. in Gillette
(ed). Vertebrate Paleontology in Utah. Utah Geological Survey Miscellaneous
Publication. 99-1, 77-114.
undescribed dromaeosaurid (Ullmann, Varricchio, Knell and Lacovara,
2010)
Albian, Early Cretaceous
Vaughn Member of the Blackleaf Formation, Montana, US
Reference- Ullmann, Varricchio, Knell and Lacovara, 2010. Taphonomy and
taxonomy of a vertebrate microsite in the Cretaceous Blacklaef Formation in
Southwest Montana. Journal of Vertebrate Paleontology. Program and Abstracts
2010, 179A.
undescribed Dromaeosauridae (MOR online)
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
Material- (MOR 721) partial skeleton
Comments- This probably belongs to Saurornitholestes and/or Bambiraptor.
undescribed Dromaeosauridae (MOR online)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Material- (MOR 029) teeth (MOR online)
(MOR 040) ungual fragment (MOR online)
(MOR 119) digit, two phalanges (MOR online)
(MOR 126) manual ungual (MOR online)
(MOR 156) tooth (MOR online)
(MOR 176) tooth (MOR online)
(UCMP 129723) maxillary fragment (UCMP online)
(UCMP 140582) tooth (UCMP online)
(UCMP 154576) ungual (UCMP online)
(UCMP 154577) phalanx (UCMP online)
(UCMP 154578) phalanx (UCMP online)
(YPM PU 22301) (YPM online)
(YPM PU 22302) (YPM online)
Comments- These remains likely belong to Richardoestesia, Paronychodon,
Saurornitholestes and/or Dromaeosaurus.
unnamed Dromaeosauridae (Triebold and Russell, 1995)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (MOR 059) tooth (MOR online)
(MOR 061) ungual (MOR online)
(UCMP 64139) phalanx (UCMP online)
(UCMP 64140) phalanx (UCMP online)
(UCMP 119675) tooth (UCMP online)
(UCMP 119679) pedal ungual (UCMP online)
(UCMP 119751) tooth (UCMP online)
(UCMP 119924) three teeth (UCMP online)
(UCMP 119925) four teeth (UCMP online)
(UCMP 119926) tooth (UCMP online)
(UCMP 120077) two teeth (UCMP online)
(UCMP 120256) two teeth (UCMP online)
(UCMP 120304) ungual (UCMP online)
(UCMP 123338) over twenty teeth (UCMP online)
(UCMP 123339) twenty-four teeth (UCMP online)
(UCMP 123505) tooth (UCMP online)
(UCMP 123506) tooth (UCMP online)
(UCMP 123507) three teeth (UCMP online)
(UCMP 123525) two teeth (UCMP online)
(UCMP 123544) tooth (UCMP online)
(UCMP 123566) tooth (UCMP online)
(UCMP 124989) tooth (UCMP online)
(UCMP 129085) two teeth (UCMP online)
(UCMP 134801) tooth (UCMP online)
(UCMP 145737) tooth (UCMP online)
(UCMP 145877) two teeth (UCMP online)
(UCMP 145968) tooth (UCMP online)
(UCMP 174752) tooth (UCMP online)
(UCMP 174796) tooth (UCMP online)
(UCMP 186838) tooth (UCMP online)
(UCMP 186870) teeth (UCMP online)
(UCMP 186874) teeth (UCMP online)
(UCMP 186877) teeth (UCMP online)
(UCMP 186882) teeth (UCMP online)
(UCMP 186893) teeth (UCMP online)
(UCMP 186899) tooth (UCMP online)
(UCMP 186903) teeth (UCMP online)
(UCMP 186909) teeth (UCMP online)
(UCMP 186912) teeth (UCMP online)
(UCMP 186929) teeth (UCMP online)
(UCMP 186944) teeth (UCMP online)
(UCMP 186954) teeth (UCMP online)
(UCMP 186956-186962) seven teeth (UCMP online)
(UCMP 186963) teeth (UCMP online)
(UCMP 186964) tooth (UCMP online)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Material- (FMNH PR2896) tooth (6.5x3.1x1.3 mm) (Gates, Zanno and Makovicky, 2015)
material (Triebold and Russell, 1995)
material (Jacobsen and Sroka, 1998)
Comments- The above material has merely been identified as Dromaeosauridae.
References- Triebold and Russell, 1995. A new small dinosaur locality in the Hell Creek
Formation: Journal of Vertebrate Paleontology. 15(3), 57A.
Jacobson and Sroka, 1998. Preliminary assement of a Hell Creek Dinosaurian Fauna
from sites in Corson County, South Dakota. Journal of Vertebrate Paleontology.
18(3), 53A.
Gates, Zanno and Makovicky, 2015. Theropod teeth from the upper Maastrichtian
Hell Creek Formation "Sue" Quarry: New morphotypes and faunal comparisons.
Acta Palaeontologica Polonica. 60(1), 131-139.
undescribed Dromaeosauridae (AMNH online)
Cretaceous
Montana, US
Material- (AMNH 2363) six teeth
Comments- These are listed on the AMNH wbsite as cf. Velociraptor
sp., but are more likely another basal dromaeosaurid taxon based on provenence.
undescribed Dromaeosauridae (UCMP online)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- (UCMP 120851) four teeth
(UCMP 120852) over thirty-five teeth
unnamed Dromaeosauridae (Ostrom, 1969)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- (UCMP 73079) phalanx (UCMP online)
(UCMP 186861) teeth (UCMP online)
(UCMP 186862) teeth (UCMP online)
(UCMP 186888) teeth (UCMP online)
(UCMP 186891) tooth (UCMP online)
(UCMP 186901) teeth (UCMP online)
(UCMP 186917) teeth (UCMP online)
(UCMP 186921) teeth (UCMP online)
(YPM PU 20589) proximal pedal phalanx II-2 (Ostrom, 1969)
(YPM PU 55525) (YPM online)
Comments- Ostrom (1969) notes PU 20589 is almost identical to the element
in Deinonychus except that the proximoventral heel is a third larger.
He considered it an indeterminate new dromaeosaurid taxon.
The UCMP specimens and YPM PU 55525 are merely listed as Dromaeosauridae.
References- Ostrom, 1969. Osteology of Deinonychus antirrhopus,
an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of
Natural History Bulletin. 30, 1-165.
undescribed possible Dromaeosauridae (Chure, Madsen and Britt, 1993)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of Morrison Formation, Utah, US
Material- teeth
Comments- Chure et al. (1993) reported small teeth with large hooked
distal serrations and small or absent mesial serrations.
References- Chure, Madsen and Britt, 1993. New data on theropod dinosaurs
from the Late Jurassic Morrison FM. (MF). Journal of Vertebrate Paleontology.
13(3), 30A.
Chure, 1995. The teeth of small theropods from the Morrison Formation (Upper
Jurassic: Kimmeridgian), UT. Journal of Vertebrate Paleontology. 15(3), 23A.
unnamed dromaeosaurid (Senter, Kirkland, Deblieux and Madsen, 2010)
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Material- (UMNH VP 20209) proximal caudal vertebra (45 mm), six distal
caudal vertebrae, distal caudal vertebra (41 mm), distal caudal vertebra (~40
mm), distal caudal vertebra (~43 mm), distal caudal vertebra (~46 mm), distal
chevrons
Comments- This was initially identified (Senter et al., 2010) as part
of what would later be named Yurgovuchia by Senter et al. (2012).
References- Senter, Kirkland, Deblieux and Madsen, 2010. Three new theropods
from the Cedar Mountain Formation (Lower Cretaceous) of Utah. Journal of Vertebrate
Paleontology. Program and Abstracts 2010, 162A.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
undescribed Dromaeosauridae (Kirkland, 1996)
Early Albian, Early Cretaceous
Dinosaur National Monument, Ruby Ranch Member of Cedar Mountain Formation, Utah, US
Material- partial skeleton including mid dorsal vertebra, femur (~290 mm), astragalus
and pes (Chure, Britt and Scheetz, 2007)
multiple elements (Chure, Britt and Scheetz, 2007)
Early Albian, Early Cretaceous
Hotel Mesa Site, Ruby Ranch Member of Cedar Mountain Formation, Utah, US
(OMNH coll.) teeth (Kirkland, 1996)
Early Albian, Early Cretaceous
Lorrie's Site, Dinosaur National Monument, Ruby Ranch Member of Cedar Mountain Formation, Utah, US
(DMNH coll.) manual ungual (Kirkland, online 2007)
Comments-
Kirkland (1996) initially reported dromaeosaurid teeth from the "middle
fauna" of the Cedar Mountain Formation, seemingly referring to "teeth
possibly referable to the dromaeosaurid Deinonychus" from Hotel Mesa mentioned by Kirkland et al. (1997). Chure et al. (2007) reported a non-Utahraptor
taxon from Dinosaur National Monument "represented by a partial
skeleton plus isolated elements of a second individual. Femoral length
is ~290 mm. The single mid-dorsal bear a pneumatic foramen on its
centrum. The Monument specimen differs from Utahraptor
in the extreme height of the ascending process of the astragalus plus
its symmetrical shape, with the apex at mid-width." Carpenter
(online 2007) stated "A single hand claw of Utahraptor is known from near the base of the Ruby Ranch at Lorrie's Site", and "The foot of Deinonychus is known from high in the Ruby Ranch at
References-
Kirkland, 1996. Biogeography of western North America's Mid-Cretaceous
dinosaur faunas: Losing European ties and the first great Asian-North
American interchange. Journal of Vertebrate Paleontology. 16(3), 45.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis
and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the
central Colorado plateau: A key to understanding 35 million years of
tectonics, sedimentology, evolution, and biogeography. Brigham Young
University Geology Studies. 42, 69-103.
Chure, Britt and Scheetz, 2007. Sickle-claw theropod dinosaurs of the Lower
Cretaceous Cedar Mountain Formation from the Dalton Wells Quarry and Dinosaur
National Monument, Utah. Journal of Vertebrate Paleontology. 27(3), 59A.
Carpenter, online 2007. https://web.archive.org/web/20090517064058/http://scientists.dmns.org/kenCarpenter/cedar-mountain-project/dinosaurs-of-the-cedar-mountain-formation/
Dromaeosauridae indet. (Kirkland, Lucas and Estep, 1998)
Late Cenomanian, Late Cretaceous
Dakota Formation, Utah, US
Material- teeth
Comments- These remains were listed as Velociraptorinae indet. by Kirkland
et al. (1998) and Eaton et al. (1999). They are presumably one of the two Dromaeosauridae
indet. gen. et sp. teeth listed by Kirkland et al. (1997).
References- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton,
Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the
Central Colorado Plateau: a key to understanding 35 million years of tectonics,
sedimentology, evolution, and biogeography. Brigham Young University Geology
Studies. 42, 69-103.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau.
in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial
Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Eaton, Cifelli, Hutchison, Kirkland and Parrish, 1999. Cretaceous vertebrate
faunas from the Kaiparowits Plateau, south central Utah. in Gillette (ed.).
Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication.
99-1, 345-353.
unnamed Dromaeosauridae (Nelson and Crooks, 1987)
Cenomanian-Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71599) two teeth (Fiorillo, 1999)
(FHSU coll.) two teeth (~1.1x~1.2x? mm, ~2.3x~2.1x? mm) (Nelson and Crooks, 1987)
(NCSM 33267) lateral tooth (10.15x5.03x3.32 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
(NCSM 33275) lateral tooth (5.83x3.23x1.36 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
teeth (Kirkland et al., 1997)
partial skeleton (Cifelli et al., 1999)
Comments- Nelson and Crooks (1987) figure two teeth as "Theropoda, Troodontidae
(= Saurornithoididae)" from the Rough Road Quarry. Currie et al.
(1990) said these "are more likely from a velociraptorine because the
denticles are too small (there are 12 posterior denticles per
millimeter in their figures) and elongate."
Material was called cf. Deinonychus sp. nov. by Kirkland
et al. (1997), and noted to be large. Fiorillo (1999) described two teeth which
were strongly compressed, lack mesial serrations, and have distal serrations
which are taller than wide and are straight with a slight apically oriented
hook. He referred these to Velociraptorinae. Cifelli et al. (1999) state a partial
skeleton is known, though do not say if it is of the "velociraptorine"
or "dromaeosaurine" taxon. Cifelli et al. also list Velociraptorinae
indet. teeth.
Avrahami et al. (2016) reported "four complete and several fragmentary
dromaeosaur teeth from an as-yet indeterminate species" from the Cliffs
of Insanity microsite, which by the time of the material's description
(Avrahamai, 2018; Avrahami et al., 2018) included NCSM 33267 and 33275,
and perhaps also Richardoestesia teeth NCSM 33274 and 33288.
References- Nelson and Crooks, 1987. Stratigraphy and paleontology of
the Cedar Mountain Formation (Lower Cretaceous), eastern Emery County, Utah.
In Averett (ed.). Paleontology and Geology of the Dinosaur Triangle: Guidebook
for 1987 Field Trip. Museum of Western Colorado. 55-63.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press. 107-125.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton,
Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the
Central Colorado Plateau: a key to understanding 35 million years of tectonics,
sedimentology, evolution, and biogeography. Brigham Young University Geology
Studies. 42, 69-103.
Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial Cretaceous
vertebrates from the Cedar Mountain Formation, Emery County, Utah: the Mussentuchit
Local Fauna. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison Eggshell
site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. in Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 259-268.
Avrahami, Humphrey, Heckert, Gates, Makovicky and Zanno, 2016. The
early Late Cretaceous paleofaunal vertebrate assemblage of the Cliffs
of Insanity microsite in the Mussentuchit Member of the Cedar Mountain
Formation, Utah. Journal of Vertebrate Paleontology. Program and
Abstracts, 90.
Avrahami, 2018. Paleobiodiversity of a new microvertebrate locality
from the Upper Cretaceous Mussentuchit Member, Cedar Mountain
Formation, Utah: Testing morphometric multivariate approaches for
quantifying shape variation in microvertebrate specimens. Masters
thesis, North Carolina State University. 181 pp.
Avrahami, Gates, Heckert, Makovicky and Zanno, 2018. A new
microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain
Formation: Insights into the paleobiodiversity and paleobiogeography of
early Late Cretaceous ecosystems in western North America. PeerJ.
6:e5883.
Dromaeosauridae indet. (Kirkland, Lucas and Estep, 1998)
Middle-Late Turonian, Late Cretaceous
Smoky Hollow Member of the Straight Cliffs Formation, Utah, US
Material- (MNA 994) tooth (Parrish, 1999)
(OMNH 24439) tooth (Parrish, 1999)
(OMNH 24441) tooth (Parrish, 1999)
(OMNH 24442) tooth (Parrish, 1999)
Comments- These were listed as Velociraptorinae by Kirkland et al. (1998)
and Parrish (1999).
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian)
of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
Dromaeosauridae indet. (Kirkland, Lucas and Estep 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Material- (OMNH 21238) tooth (Parrish, 1999)
(OMNH 21673) tooth (Parrish, 1999)
? material (Eaton et al., 1999)
Comments- These were listed as Velociraptorinae by Kirkland et al. (1998)
and Parrish (1999). In addition, Eaton et al. (1999) listed ?Dromaeosauridae
indet. from a Santonian possible Straight Cliffs (or Wahweap?) locality.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Eaton, Diem, Archibald, Schierup and Munk, 1999. Vertebrate paleontology of
the Upper Cretaceous rocks of the Markagunt Plateau, southwestern Utah. in Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 323-333.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian)
of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
possible undescribed dromaeosaurid (Eaton, 1999)
Santonian-Campanian, Late Cretaceous
Iron Springs Formation, Utah, US
Material- tooth
Reference- Eaton, 1999. Vertebrate paleontology of the Iron Springs Formation,
Upper Cretaceous, southwestern Utah. in Gillette (ed.). Vertebrate Paleontology
in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 339-343.
Dromaeosauridae indet. (Kirkland, Lucas and Estep 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah, US
Material- (UCM 8613) tooth (Parrish, 1999)
? material (Eaton et al., 1999)
Comments- UCM 8613 was listed as Velociraptorinae by Kirkland et al.
(1998) and Parrish (1999). In addition, Eaton et al. (1999) listed ?Dromaeosauridae
indet. from an Early Campanian possible Wahweap locality.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Eaton, Diem, Archibald, Schierup and Munk, 1999. Vertebrate paleontology of
the Upper Cretaceous rocks of the Markagunt Plateau, southwestern Utah. in Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 323-333.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian)
of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
Dromaeosauridae indet. (Parrish, 1999)
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Material- (MNA HM-6) tooth (Parrish, 1999)
(OMNH 21240) tooth (Parrish, 1999)
(OMNH 24158) tooth (Parrish, 1999)
(UCM 8312, 83240 (in part), 8642 (in part), 8655, 8659 (in part), 8663) six
teeth (Parrish, 1999)
Comments- The teeth were listed as Velociraptorinae by Parrish (1999).
UCMP 149171 was listed in the UCMP online database as dromaeosaurid, but is
a troodontid (Zanno et al., 2011).
References- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-.
Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah.
Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.
Zanno, Varricchio, O’Connor, Titus and Knell, 2011. A new troodontid theropod,
Talos sampsoni gen. et sp. nov., from the Upper Cretaceous western interior
basin of North America. PLoS ONE. 6(9), e24487.
unnamed Dromaeosauridae (Bonde, 2008)
Albian, Early Cretaceous
Willow Tank Formation, Nevada, US
Material- teeth (?x7x3.5, ?x6x3.5 mm), incomplete femur, partial tibia
Reference- Bonde, 2008. Paleoecology and taphonomy of the Willow Tank
Formation (Albian), southern Nevada. Masters thesis, Montana State University.
96 pp.
Dromaeosauridae indet. (Ratkevich and Duffek, 1996)
Campanian, Late Cretaceous
Fort Crittenden Formation, Arizona, US
Material- two teeth
Comments- These were identified as cf. Richardoestesia and cf.
Saurornitholestes by Ratkevich and Duffek (1996), but reidentified merely
as Dromaeosauridae indet. by Sullivan and Lucas (2006).
References- Ratkevich and Duffek, 1996. Small macro-and large micro-vertebrate
fauna of the Fort Crittenden Formation, Southeast Arizona. Proceedings of Southwest
Paleontological Society and Mesa Southwest Museum, Mesa, Arizona. 4, 115-120.
Sullivan and Lucas, 2006. The Kirtlandian land-vertebrate "age" -
faunal composition, temporal position and biostratigraphic correlation in the
nonmarine Upper Cretaceous of western North America. New Mexico Museum of Natural
History and Science Bulletin. 35, 7-29.
undescribed dromaeosaurid (Lucas et al., 1988)
Mid Santonian, Late Cretaceous
Point Lookout Sandstone, New Mexico, US
Material- (NMMNH P-27481) tooth (8.9x5.2x2.7 mm) (Williamson and Brusatte,
2014)
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the
Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications
for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Dromaeosauridae indet. (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-33147) tooth
References- Williamson and Brusatte, 2014. Small theropod teeth from
the Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their
implications for understanding Latest Cretaceous dinosaur evolution. PLoS ONE.
9(4), e93190.
Dromaeosauridae indet. (Lehman, 1981)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- ?(Lafon coll.) metatarsal I (Lehman, 1981)
(NMMNH P-32814) tooth (5.2x3.3x1.6 mm) (Williamson and Brusatte, 2014)
(SMP VP-2505) tooth (34 mm) (Jasinski, Sullivan and Lucas, 2011)
(SMP VP-2595) tooth (14 mm) (Jasinski, Sullivan and Lucas, 2011)
Comments- Williamson and Brusatte (2014) state NMMNH P-32814 is more
similar to Acheroraptor than Saurornitholestes due to rounded
distal serrations and lingual ridges.
References- Lehman, 1981. The Alamo Wash local fauna: A new look at the old Ojo Alamo fauna.
In Lucas, Rigby and Kues (eds.). Advances in San Juan Basin paleontology. University
of New Mexico Press. 189-221.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Jasinski, Sullivan and Lucas, 2011. Taxonomic composition of the Alamo Wash
local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member),
San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann (eds.). Fossil
Record 3. New Mexico Museum of Natural History and Science Bulletin. 53, 216-271.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
undescribed Dromaeosauridae (Fix, Darrough, Parris and Grandstaff, 2012)
Campanian, Late Cretaceous
Chronister site, Missouri, US
Reference- Fix, Darrough, Parris and Grandstaff, 2012. Western Appalachia
Dinosauria and associated vertebrates of the Late Cretaceous of Southeast Missouri.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 94.
undescribed Dromaeosauridae
?Late Cretaceous
?North America
Material- (AMNH 86344) (AMNH online)
(MOR 140) teeth (MOR online)
(MOR 157) tooth (MOR online)
(YPM 56980) (YPM online)
(YPM 56999) (YPM online)
(YPM PU 55010) (YPM online)
(YPM PU 55011) (YPM online)
undescribed Dromaeosauridae (Rivera-Sylva, Frey, Stinnesbeck, Padilla
Gutierrez, Gonzalez Gonzalez and Amezcua Torres, 2015)
Late Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Material- long bone fragments and phalanx
Reference- Rivera-Sylva, Frey, Stinnesbeck, Padilla Gutierrez, Gonzalez
Gonzalez and Amezcua Torres, 2015. The Late Cretaceous Las Aguilas dinosaur
graveyard, Coahuila, Mexico. Journal of Vertebrate Paleontology. Program and
Abstracts 2015, 203.
undescribed dromaeosaurid (Metcalf, Vaughan, Benton, Cole, Simms and
Dartnall, 1992)
Early Bathonian, Middle Jurassic
Chipping Norton Formation, England
Material- (GLRCM coll.; F) tooth (3.5 mm)
Comments- This was labeled a dromaeosaur-like tooth and said to be a
possible troodontid. It is elongate and moderately recurved, with mesial serrations
present apically. Serrations on both carinae are low and rounded, with a DSDI
of ~2. The base is not constricted and blood pits are absent. Serration density
is 6/mm distally and 13/mm mesially.
This tooth differs from troodontids in lacking a basal constriction and blood
pits, as well as having such a high DSDI. The latter is only known in some derived
dromaeosaurids and Richardoestesia, but the low serration density excludes
it from the latter genus. However, the low rounded morphology of the distal
serrations is not seen in other dromaeosaurids.
Reference- Metcalf and Walker, 1994. A new Bathonian microvertebrate
locality in the English Midlands. in Fraser and Sues (eds.). In the Shadow of
the Dinosaurs- Mesozoic Small Tetrapods, Cambridge (Cambridge University Press).
322-332.
undescribed Dromaeosauridae (Wills, 2015)
Bathonian, Middle Jurassic
Forest Marble Formation, England
Material- (small) sixteen teeth
Reference- Wills, 2015. Isolated dromaeosaurid teeth from the Bathonian
(Middle Jurassic) of Dorset, United Kingdom. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 238.
undescribed possible dromaeosaurid (Naish pers. comm. to Sweetman, 2004)
Barremian-Early Aptian, Early Cretaceous
Wealden Group, England
Material- fragmentary remains.
Comments- Sweetman commented on these remains, attributed to Naish pers.
comm. 2003.
Reference- Sweetman, 2004. The first record of velociraptorine dinosaurs
(Saurischia, Theropoda) from the Wealden (Early Cretaceous, Barremian) of southern
England. Cretaceous Research. 25, 353-364.
unnamed Dromaeosauridae (Sweetman, 2004)
Barremian, Early Cretaceous
Wessex Formation, England
Material- (IWCMS 2002.1) lateral tooth (16 mm)
(IWCMS 2002.2) lateral tooth (8.5 mm)
(IWCMS 2002.3) lateral tooth
(IWCMS 2002.4) lateral tooth
(NHMUK R 16510) lateral tooth (>21.5 mm)
Comments- These teeth are moderately to strongly recurved and compressed,
with BW/FABL ratios of .4-.6. Mesial serrations are present only apically, and
absent in IWCMS 2002.2. There are 3.8-4.8/mm. Distal serrations number 2.7-3.5/mm
in most specimens, are rounded and inclined apically. IWCMS 2002.2 has 6.25/mm
however, and may be a different taxon. Sweetman (2004) assigned them to Velociraptorinae
based on the high DSDI (1.37-1.6)
Reference- Sweetman, 2004. The first record of velociraptorine dinosaurs
(Saurischia, Theropoda) from the Wealden (Early Cretaceous, Barremian) of southern
England. Cretaceous Research. 25, 353-364.
undescribed Dromaeosauridae (Royo y G�mez, 1927)
Tithonian-Berriasian, Late Jurassic-Early Cretaceous
Villar del Arzobispo Formation, Spain
Material- (CPT-1283) tooth (5.4x4.1x2.3 mm) (Gasco et al., 2012)
(CPT-1327) tooth (14.6x8.4x4 mm) (Gasco et al., 2012)
(CPT-1433) tooth (~10.7x5.1x3.1 mm) (Gasco et al., 2012)
(CPT-1443) tooth (5.5x4.2x2.3 mm) (Gasco et al., 2012)
(CPT-1658) tooth (6.2x3.7x2.4 mm) (Gasco et al., 2012)
(CPT-1659) tooth (6.8x4.3x2.6 mm) (Gasco et al., 2012)
(CPT-2126) tooth (6.3x4x2.1 mm) (Gasco et al., 2012)
(MNCN coll.) tooth (Royo y G�mez, 1927)
Comments- MNCN coll. was originally identified by Royo y Gomez (1927)
as Megalosaurus cf. dunkeri, then as a dromaeosaurid by Ruiz-Ome�aca
and Pereda-Suberbiola (1999).
References- Royo y G�mez, 1927. Sesi�n del 6 de Julio de
1927. Bolet�n de la Real Sociedad Espa�ola de Historia Natural.
27.
Ruiz-Ome�aca and Pereda-Suberbiola, 1999. Un documento in�dito
de Royo y G�mez sobre los dinosaurios del Levante. Temas Geol�gico-Mineros
ITGE. 26, 111-112.
Gasc�, Cobos, Royo-Torres, Mampel and Alcal�, 2012. Theropod teeth
diversity from the Villar del Arzobispo Formation (Tithonian-Berriasian) at
Riodeva (Teruel, Spain). Palaeobiodiversity and Palaeoenvironments. 92(2), 273-285.
unnamed dromaeosaurid (Rauhut and Zinke, 1995)
Barremian, Early Cretaceous
Una Formation, Spain
Material- (IPFUB Una Th 21-36, 38, 40-47, 50-52, 63, 65, 70) sixty-six
teeth
(IPFUB Una Th 66) tooth (15 mm)
Comments- These teeth are strongly recurved and highly compressed. Distal
serrations are rounded or slightly inclined apically. Mesial serrations are
much smaller when present (DSDI 1.11-2). A few have faint longitudnal ridges
on both sides. Rauhut (2002) referred them to Velociraptorinae due to the high
DSDI, large serrations (~7-7.5/mm) and strong recurvature.
References- Rauhut and Zunke, 1995. A description of the Barremian dinosaur
fauna from Una with a comparison of that of Las Hoyas. II. International Symposium
of Lithographic Limestone, Extended Abstracts. 123-126.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca,
Spain. Cretaceous Research. 23, 255-263.
unnamed dromaeosaurid (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Late Campanian, Late Cretaceous
Figuerola 2, Tremp Formation, Spain
Material- (DPM-FIG2-T1+T2) tooth (12.1x6.7x4.3 mm)
Comments- This was listed as Dromaeosauridae indet. 2 by Torices (2002),
and Dromaeosauridae indet. by Torices et al. (2015).
Reference- Torices, 2002. Los dinosaurios ter�podos del Cret�cico
Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de
Paleontolog�a. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
unnamed dromaeosaurid (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Late Campanian, Late Cretaceous
La�o, Sedano Formation, Spain
Material- (MCNA 14622) tooth (3.4x2.6x1.3 mm)
Reference- Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod
dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta
Palaeontologica Polonica. 60(3), 611-626.
undescribed dromaeosaurid (Company, Torices, Pereda-Suberbiola and Ruiz-Omenaca,
2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Sierra Perenchiza Formation, Valencia, Spain
Material- two teeth (~10 mm)
Comments- Described as strongly compressed labiolingually, slightly recurved
distally, and both mesial and distal carinae have minute serrations (~7 per
mm).
Reference- Company, Torices, Pereda-Suberbiola and Ruiz-Omenaca, 2009.
Theropod teeth from the Late Cretaceous of Chera (Valencia, Eastern Spain).
Journal of Vertebrate Paleontology. 29(3), 81A.
undescribed Dromaeosauridae (Ortega, Escaso, Perez Garcia, Torices and
Sanz, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- teeth, several postcranial elements
Comments- Ortega et al. (2009) state teeth of Velociraptorinae and dromaeosaurid
postcrania are present. Torices et al. (2011) mention both cf. Dromaeosauridae
indet. and cf. Velociraptorinae indet..
Reference- Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009. The
vertebrate diversity of the Upper Campanian-Lower Maastrichtian "Lo Hueco"
fossil-site (Cuenca, Spain). Journal of Vertebrate Paleontology. 29(3), 159A-160A.
Torices, Barroso-Barcenilla, Cambra-Moo, Perez and Serrano, 2011. Vertebrate
microfossil analysis in the palaeontological site of 'Lo Hueco' (Upper Cretaceous,
Cuenca, Spain). Journal of Vertebrate Paleontology. Program and Abstracts 2011,
205.
unnamed Dromaeosauridae (Torices, 2002)
Early Maastrichtian, Late Cretaceous
Fontllonga 6, Tremp Formation, Spain
Material- (DPM-FON6-T1) tooth
(DPM-FON6-T2) tooth (16.3x7.5x5.8 mm)
Comments- DPM-FON6-T1 was noted as Dromaeosauridae indet. 1 by Torices
(2002), though not mentioned in Torices et al. (2015). DPM-FON6-T2 was listed
by Torices as Dromaeosauridae indet. 2, and by Torices et al. as Dromaeosauridae
indet..
Reference- Torices, 2002. Los dinosaurios ter�podos del Cret�cico
Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de
Paleontolog�a. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
unnamed dromaeosaurid (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Late Maastrichtian, Late Cretaceous
Blasi, Aren Formation, Spain
Material- (MPZ2004/6) tooth (17.5x11x4.4 mm)
Reference- Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod
dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta
Palaeontologica Polonica. 60(3), 611-626.
unnamed dromaeosaurid (Garcia, Pincemaille, Vianey-Liaud, Marandat,
Lorenz, Cheylan, Capetta, Michaux and Sudre, 1999)
Early Maastrichtian, Late Cretaceous
Vitrolles-Couperigne, Provence, France
Material- few teeth
Comments- Garcia et al. (1999) provisionally assign a few teeth to Dromaeosauridae.
The illustrated tooth is highly recurved, not constricted basally, seems to
lack mesial serrations, while having 2-3 distal serrations per mm. Distal serrations
are hooked apically. They therefore most closely resemble "velociraptorine"
dromaeosaurids.
Reference- Garcia, Pincemaille, Vianey-Liaud, Marandat, Lorenz, Cheylan,
Capetta, Michaux and Sudre, 1999. Decouverte du premier squelette presque complet
de Rhabdodon priscus (Dinosauria, Ornithopoda) du Maastrichtian inferieur
de Provence. Les Comptes rendus de l'Acad�mie 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. Aper�u sur la g�ologie du cha�non
de Saint-Chinian. Bull. Soc. G�ol. France. 27, 686-709.
Lapparent, 1947. Les dinosauriens du Cr�tac� sup�rieur
du midi de la France. M�moire de la Soci�t� g�ologique
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'Acad�mie
des sciences. 328, 781-787.
undescribed Dromaeosauridae (Lubbe, Richter and Knotschke, 2009)
Kimmeridgian, Late Jurassic
Langenberg Quarry, Germany
Material- (DFMMh/FV 383) tooth (9.7x6.9x3.1 mm) (Lubbe, Richter and Kn�tschke,
2009)
(DFMMh/FV 530) tooth (7.2x5.7x2.7 mm) (Lubbe, Richter and Kn�tschke, 2009)
(DFMMh/FV 705) tooth (xx mm) (Lubbe, Richter and Kn�tschke, 2009)
Comments- While seven teeth assigned to Velociraptorinae by Lubbe et
al. (2009), Gerke and Wings (2014) found four of these were Neotheropoda indet.,
megalosaurid and tyrannosauroid. Based on the information in Lubbe et al., FV
658 may be the megalosaurid (larger, elongate crown, more of mesial carina serrated),
FV 382 and 790.5 may be tyrannosauroid (less recurved and thicker labiolingually),
and FV 707.1 may be indet. (larger and only partially preserved).
References- Lubbe, Richter and Kn�tschke, 2009. Velociraptorine
dromaeosaurid teeth from the Kimmeridgian (Late Jurassic) of Germany. Acta Palaeontologica
Polonica. 54(3), 401-408.
Gerke and Wings, 2014. Characters versus morphometrics: A case study with isolated
theropod teeth from the Late Jurassic of Lower Saxony, Germany, reveals an astonishing
diversity of theropod taxa. Journal of Vertebrate Paleontology. Program and
Abstracts 2014, 137.
unnamed Dromaeosauridae (Lanser and Heimhofer, 2015)
Late Barremian-Early Aptian, Early Cretaceous
Balve-Beckum quarry, Germany
Material- (LWL MN Ba 5) partial tooth
(LWL MN Ba 6) tooth fragment
(LWL MN Ba 7) tooth fragment
(LWL MN Ba 9) tooth fragment
(LWL MN Ba 10) tooth fragment
(LWL MN Ba 11) tooth fragment
(LWL MN Ba 12) incomplete tooth (?x5x3.8 mm)
(LWL MN Ba 13) tooth (12.8x7.4x4.5 mm)
Reference- Lanser and Heimhofer, 2015. Evidence of theropod dinosaurs
from a Lower Cretaceous karst filling in the northern Sauerland (Rhenish Massif,
Germany). Pal�ontologische Zeitschrift. 89(1), 79-94.
unnamed possible Dromaeosauridae (Lindgren, Currie, Rees, Siverson,
Lindstr�m and Alwmark, 2008)
Early Berriasian, Early Cretaceous
Skyttegaard Member of the Rabekke Formation, Denmark
Material- (MGUH 28403) incomplete tooth (5x3.5x1.3 mm)
(MGUH 28404) incomplete tooth (4.1x2.5x1.4 mm)
(MGUH 28405) (juvenile?) incomplete tooth (1.5x.1.2x.8 mm)
(MGUH 28406) partial tooth
(MGUH 28407) tooth fragment
(MGUH 28408) tooth fragment
(MGUH 28409) tooth
Comments- Lindgren et al. (2008) referred these to three morphotypes-
28403, 28404 and 28408 are said to be a basal velociraptorine, 28406 and 28407
might be conspecific but differ in serration morphology, and 28405 was a possible
dromaeosaurine. Bonde (2012) on the other hand referred 28406 and 28407 to Velociraptorinae,
28403 and 28404 to basal dromaeosaurids, and 28405 to Dromaeosaurinae. The latter
also considered 28409 as Dromaeosauridae incertae sedis.
References- Lindgren, Currie, Rees, Siverson, Lindstr�m and Alwmark,
2008. Theropod dinosaur teeth from the lowermost Cretaceous Rabekke Formation
on Bornholm, Denmark. Geobios. 41, 253-263.
Bonde, 2012. Danish dinosaurs: A review. In Godefroit (ed.). Bernissart Dinosaurs
and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 434-451.
undescribed Dromaeosauridae (Codrea, Godefroit and Smith, 2012)
Maastrichtian, Late Cretaceous
Rusca Montana Basin, Romania
Material- (UBB NgTh2) tooth
(UBB coll.) two teeth
Reference- Codrea, Godefroit and Smith, 2012. First discovery of Maastrichtian
(Latest Cretaceous) terrestrial vertebrates in Rusca Montana Basin (Romania).
In Godefroit (ed.). Bernissart Dinosaurs and Early Cretaceous Terrestrial Ecosystems.
Indiana University Press. 570-581.
unnamed Dromaeosauridae (Kordikova, Polly, Alifanov, Rocek, Gunnell
and Averianov, 2001)
Turonian-Coniacian, Late Cretaceous
Zhirkindek Formation, Kazakhstan
Reference- Kordikova, Polly, Alifanov, Rocek, Gunnell and Averianov,
2001. Small vertebrates from the Late Cretaceous and early Tertiary of the northeastern
Aral Sea Region, Kazakhstan. Journal of Paleontology. 75, 390-400.
unnamed dromaeosaurid (Averianov, 2007)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- (ZIN PH 34/49) posterior tooth (FABL 2.1 mm) (Averianov, 2007)
Comments- This tooth has a BW/FABL of .38, with a centered mesial carina
with very small serrations present at least apically. Distal serrations are
U-shaped and occur at a density of 7/mm. It was referred to Velociraptorinae
by Averianov (2007). The small serration sze may indicate this is microraptorian.
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits
in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.
undescribed possible dromaeosaurid (Nessov, 1995)
Santonian, Late Cretaceous
Syuk-Syuk Formation, Kazakhstan
Material- unguals?
Comments- Nessov (1995) notes Prinada (1925, 1927) and/or Riabinin (1938,
1939) identified unguals as Velociraptor. This remains uncertain pending
further studies.
References- Prinada, 1925. [Search for remains of large vertebrates of
Upper Cretaceous age in Turkestan. Report on the state of activities of the
Geological Committee for 1924. Part II, III]. Izvyestiya Gyeologichyeskogo komityeta.
44(2), 257.
Prinada, 1927. [Report on the excavation at the localities where dinosaur bones
were discovered. Report on the state of activities of the Geological Committee
for 1925. Part II, III]. Izvyestiya Gyeologichyeskogo komityeta. 45(4), 453-454.
Riabinin, 1938. [Some results of the study of the Upper Cretaceous dinosaur
fauna from the vicinity of st. Sary-Agachin, Southern Kazakhstan]. Problyemy
palyeontologii. 4, 125-135.
Riabinin, 1939. [Vertebrate fauna from the Upper Cretaceous of southern Kazakhstan.
I. Reptilia. Part 1. 1. Ornithischia]. Trudy Tsyentral'nogo Nauchno-Isslyedovatyel'skogo
Gyeologorazvyedochnogo instituta. 18, 1-40.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology
and paleobiogeography. Scientific Research Institute of the Earth's Crust. 156 pp.
undescribed dromaeosaur (Novikov, Lebidev and Alifanov, 1998)
Bathonian, Middle Jurassic
Meshcherskii Gorizont of the Moskvoretskaya Svita, Russia
Material- tooth
Reference- Novikov, Lebedev and Alifanov, 1998. New Mesozoic vertebrate
fossil sites of Russia. Third European Workshop on Vertebrate Paleontology,
Maastricht, 6-9 May 1998, p. 58.
undescribed Dromaeosauridae (Averianov, Starkov and Skutschas, 2003)
Aptian-Albian, Early Cretaceous
Ilek (=Shestakovo) Formation, Russia
Material- teeth (Averianov et al., 2003)
anterior dorsal vertebra (Averianov et al., 2004)
References- Averianov, Starkov and Skutschas, 2003. Dinosaurs from the
Early Cretaceous Murtoi Formation in Buryatia, Eastern Russia. Journal of Vertebrate
Paleontology. 23(3):586–594.
Averianov, Leshchinskiy, Skutschas, Fayngertz and Rezvyi, 2004. Dinosaurs from
the Early Cretaceous Ilek Formations in West Siberia, Russia. 2nd EAVP Meeting.
July 19-24, 2004. Brno, Czech Republic. Abstracts of papers and posters with
program, Excursion Guidebook (O. Dostal, R. Gregorova & M. Ivanov, Eds.),
6.
undescribed Dromaeosauridae (Watabe and Suzuki, 2000)
Early Cretaceous
Shine Usny Tolgod, Mongolia
Material- (uncollected?) unguals
Comments- Watabe and Suzuki (2000) report "dromaeosaur claws" collected on June 11 1994.
Reference-
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 30-44.
undescribed dromaeosaurid (Matsukawa and Obata, 1994)
Aptian-Albian, Early Cretaceous
Zouyun Formation, Mongolia
Comments- Matsukawa and Obata (1994) report through personal communication
with Mateer (1992) that cf. Velociraptor mongoliensis was discovered
in the Zouyun Formation, though this is much too early to actually be that genus.
Reference- Matsukawa and Obata, 1994. Cretaceous, a contribution to dinosaur
facies in Asia based on molluscan paleontology and stratigraphy. Cretaceous
Research. 15, 101-125.
undescribed Dromaeosauridae (Watabe and Suzuki, 2000)
Cenomanian-Turonian, Late Cretaceous
Burkhant, Bayanshiree Formation, Mongolia
Material- (uncollected?) ungual (Watabe and Suzuki, 2000)
many postcranial elements (Watabe,
Tsogtbaatar, Suzuki and Saneyoshi, 2010)
Cenomanian-Turonian, Late Cretaceous
Urlibe Khudak,
Bayanshiree Formation, Mongolia
(IGM coll.) material (Kobayashi, Tsogtbaatar, Tsogtbaatar and Barsbold,
2015)
Comments-
Watabe and Burkhant (2000a) reported a "dromaeosaur claw" found at
Burkhant on August 13 1993. Watabe et al. (2010) stated "many
postcranial bones of large-sized dromaeosaurid were also found."
Kobayashi et al. (2015) reported dromaeosaurid remains from Urlibe Khudak found in 2012.
Achillobator is known from Burkhant.
References-
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1993. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 19-29.
Watabe,
Tsogtbaatar, Suzuki and Saneyoshi, 2010. Geology of
dinosaur-fossil-bearing localities (Jurassic and Cretaceous: Mesozoic)
in the Gobi Desert: Results of the HMNS-MPC Joint Paleontological
Expedition. Hayashibara Museum of Natural Sciences Research Bulletin.
3, 41-118.
Kobayashi, Tsogtbaatar, Tsogtbaatar and Barsbold, 2015. A
new therizinosaur with functionally didactyl hands from the Bayanshiree Formation
(Cenomanian-Turonian), Omnogovi Province, southeastern Mongolia. Journal of
Vertebrate Paleontology. Program and Abstracts 2015, 157.
undescribed dromaeosaurid (Watabe and Suzuki, 2000)
Late Campanian, Late Cretaceous
Abdrant Nuru, Djadokhta Formation?, Mongolia
Material- (uncollected?) ungual (Watabe and Suzuki, 2000)
Comments-
Watabe and Suzuki (2000) reported "a large claw of dromaeosaurid" from
the upper area (Toosgod) of Abdrant Nuru found on September 4-5 1996.
Reference-
Watabe and Suzuki, 2000c. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1996. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 58-68.
undescribed dromaeosaurid (Watabe and Suzuki, 2000)
Late Campanian-Early Maastrichtian, Late Cretaceous
Shiluut Ula, ?Barun Goyot Formation, Mongolia
Material- (IGM coll.; 970803 SUPJ-5-9TSGT) partial skeleton, many elements including teeth
Comments- Watabe and Suzuki
(2000) reported a "dromaeosaurid partial skeleton" discovered on August
1-4 1997, saying "the bed yielded the dromaeosaurid skeleton is a bone
bed, including many other isolated bones." The field number is
listed as "Doromaeosaurian bone bed", and they later state "from the
bone bed, isolated bones and teeth of dromaeosaurid theropod were
collected."
Reference-
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1997. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 69-82.
undescribed Dromaeosauridae (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
Material- (uncollected?) cervical vertebra (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Khaichin I, Nemegt Formation, Mongolia
three teeth (Badamkhatan, 2008)
Early Maastrichtian, Late Cretaceous
White Beds of Khermeen Tsav, Nemegt Formation, Mongolia
(IGM coll.) dorsal vertebra (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Nemegt,
Nemegt Formation, Mongolia
?(IGM coll.) dorsal vertebra (Currie, 2001)
elements (Currie et al., 2008)
Comments- Currie
(2001) lists two specimens recovered in September 10-24 2000, a
"Coelurosaur (dromaeosaur?) dorsal vertebra" from the Central Sayr of
the Nemegt locality, and a "Coelurosaur (dromaeosaur) dorsal vertebra"
from Khermeen Tsav in the Nemegt Formation. The teeth reported by
Badamkhatan (2008) are recurved, with distal serrations and an 8 shape
basally. Currie et al.'s (2008) material is from an Avimimus
bonebed. Watabe et al. (2010) reported "a cervical vertebra of
dromaeosaurid (theropod)" from Bugin Tsav. All may belong to the
contemporaneous Adasaurus.
References-
Currie, 2001. Nomadic Expeditions, Inc. report on fieldwork in Mongolia, September
2000. Alberta Palaeontological Society, Fifth Annual Symposium. 12-16.
Badamkhatan, 2008. Dinosaurs from the Late Cretaceous Mongolian
locality of Khaichin I. Journal of Vertebrate Paleontology. 28(3), 47A.
Currie, Longrich, Ryan, Eberth and Demchig, 2008. A bonebed of Avimimus
sp. (Dinosauria: Theropoda) from the Late Cretaceous Nemegt Formation, Gobi
Desert: Insights into social behavior and development in a maniraptoran theropod.
Journal of Vertebrate Paleontology. 28(3), 67A.
Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010. Report of
the HMNS-MPC Joint Paleontological Expedition in 2006. Hayashibara
Museum of Natural Sciences Research Bulletin. 3, 11-18.
undescribed dromaeosaurid (Barsbold, 1983)
Late Cretaceous?
Mongolia
Material- (IGM 100/12) pedal phalanx II-1 (25 mm), pedal phalanx II-2 (28
mm), proximal pedal ungual II
Comments- This was illustrated in figure 27 of Barsbold (1983) as an
"undescribed Mongolian dromaeosaurid". The specimen number also belongs
to a Gallimimus paratype.
Reference- Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous
of Mongolia. Transactions of the Joint Soviet-Mongolian Palaeontological Expedition.
19, 117 pp.
unnamed possible Dromaeosauridae (Dong, 1997)
Barremian-Albian, Early Cretaceous
Xinminbao Group, Gansu, China
Material- (IVPP V11122) twelve teeth, caudal vertebra, two phalanges
Comments- Dong referred this material to Dromaeosauridae, though he refers
two teeth (IVPP V11122-2 and V11122-3) from the same specimen to Troodontidae
and Allosauridae. The illustrated supposedly dromaeosaurid teeth are moderately
tall, serrated mesially and distally, and with serrations "almost as wide
as long".
Reference- Dong, 1997. On small theropods from Mazongshan Area, Gansu
Province, China. Pp. 13-18. in Dong (ed). Sino-Japanese Silk Road Dinosaur Expedition.
China Ocean Press, Beijing. 114 p.
undescribed Dromaeosauridae (Dong, Currie and Russell, 1989)
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Material- (IVPP coll.; lost?) (juvenile) several partial skeletons (Jerzykiewicz et al.,
1993)
(IVPP coll.; lost?) teeth (Dong, Currie and Russell, 1989)
Comments- Dong et al. (1989) reported "isolated bones and teeth of Velociraptor" found in June to July 1988. Jerzykiewicz et al. (1993) clarified that "Velociraptor mongoliensis,
is represented by several partial skeletons of juveniles, and by shed
teeth." Godefroit et al. (2008) stated "this material is not prepared
yet and "these undescribed specimens could not be found in the IVPP
collections." Linheraptor, "Velociraptor" osmolskae and mesially serrated teeth with a high DSDI that may be Velociraptor are known from this formation, and it's also possible the supposed juveniles are another variety of paravian as happened with Philovenator and the Almas perinates.
References- Dong, Currie and Russell, 1989. The 1988 field program of the Dinosaur Project. Vertebrata Palasiatica. 27(3), 233-236.
Jerzykiewicz, Currie, Eberth, Johnston, Koster and Zheng,
1993. Djadokhta Formation correlative strata in Chinese Inner Mongolia: An overview
of the stratigraphy, sedimentary geology, and paleontology and comparisons with
the type locality in the pre-Altai Gobi. Canadian Journal of Earth Sciences.
30(10), 2180-2195.
Godefroit, Currie, Li, Shang and Dong, 2008. A new species of Velociraptor
(Dinosauria: Dromaeosauridae) from the Upper Cretaceous of northern China. Journal
of Vertebrate Paleontology. 28(2), 432-438.
unnamed Dromaeosauridae (Gilmore, 1933)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (AMNH 6572) pedal phalanx II-1 (Ostrom, 1969)
(AMNH 21781) pedal ungual II (AMNH online)
?(IVPP 270790-4) tooth (~21x~9x? mm) (Currie and Zhao, 1993)
?(IVPP V16334.b) proximal femur (Funston, Currie, Ryan and Dong, 2019)
(IVPP coll.) teeth, elements (Dong, Currie and Russell, 1989)
(IVPP coll.) teeth and/or elements (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)
Comments-
Gilmore (1933) noted the presence of "a few foot bones and other
fragmentary skeletal parts" of small theropods from the Iren Dabasu
Formation that he assigned to "Dromaeosaurinae Genus and species
indet.", but admitted that this assignment as opposed to Coeluridae or
Compsognathidae is based purely on their Cretaceous age. Thus
while it turned out at least two of these elements are apparently
dromaeosaurid (see below), Gilmore's rationale is equivalent to
Maniraptora indet. today and the fossils referenced plausibly included
material now recognized as avimimid, troodontid and
?alvarezsaurid. Ostrom (1969) noted in a discussion of
deinonychosaur pedal examples that "E. H. Colbert has also discovered
an isolated phalanx (AMNH 6572) in the American Museum collections from
the Iren Dabasu Formation of Mongolia which compares almost exactly
with the proximal phalanx of digit II of Deinonychus,
but is perhaps 20 percent larger", which would make it somewhere around
46-52 mm long. He shows it questionably derived from Velociraptor
in his phylogram without explanation, which would not make sense in the
most recent interpretation of Iren Dabasu's age being contemporaneous
or slightly older than the Djadochta. Paul (1988) states "at the
AMNH is a hyper-extendable toe bone from the Late Cretaceous of
Mongolia that looks like a Velociraptor somewhat bigger than V. antirrhopus [= Deinonychus]",
but it is uncertain whether he saw it independant of Ostrom's text
(Paul, pers. comm. 6-2022). The AMNH online catalogue lists AMNH
21781 as an Iren Dabasu member of Dromaeosauridae represented by
"Ungual of pes (digit II)" and being found by Kaisen. Both AMNH
6572 and 21781 would have been found in Erenhot during the April 22 to
May 25 1923 Central Asiatic Expedition.
Dong et al. (1989) state Velociraptor material was discovered in the July 1988 Sino-Canadian expedition to Erenhot, and Dong (1992) specifies "teeth of Velociraptor". Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in
the Iren Dabasu" and that "Most of these can be attributed to Velociraptor,
although some of the teeth suggest that there was a second, larger
species of an indeterminate dromaeosaurine dromaeosaurid." Yet no
rationale was presented, and the only two specified Iren Dabasu
dromaeosaurid elements in the literature are clearly not Velociraptor (AMNH 6572 is twice
the size, while IVPP 270790-4 is different from most dromaeosaurid
teeth as noted above). AMNH 6572 may belong to the supposed
dromaeosaurine though, based on size. Currie and Zhao (1993)
figure "Dromaeosaurid tooth (IVPP 270790-4) from
the Iren Dabasu Formation near Erenhot, People's Republic of China,
showing replacement pit on medial side of root", but given the slight
constriction basal to the
crown, convex distal edge and seeming lack of serrations, this may be
misidentified. Based on its similar field number to troodontid
metatarsal
IVPP 230790-16 it was probably also found in the Sino-Canadian
expedition of 1990.
Funston et al. (2019) notes a femoral head that
supposedly differs from Avimimus
in having fused anterior and greater trochanters, "which suggests that
it may be oviraptorid or, more likely, dromaeosaur." This is from the Avimimus
bonebed (locality K of Currie and Eberth [1993], which was CCDP
locality 1 and may correspond to AMNH locality 141) and recovered in
July 1988. Interestingly, femur PIN 2549-100 is from the same
locality and shares a trochanteric crest, but is here identified as
oviraptorid. It's possible they are from the same taxon (or even
individual).
Yao et al. (2015) note "small unarticulated bones and teeth, including
fossils of ... dromaeosaurids" from "a rare microvertebrate locality
within the Iren Dabasu Formation, about 16 km northeast of Erenhot
City."
References- Gilmore, 1933. On the dinosaurian fauna of the Iren Dabasu
Formation. Bulletin American Museum of Natural History. 67, 23-78.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod
from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin.
30, 1-165.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster. 464 pp.
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic
of China. Cretaceous Research. 14, 127-144.
Currie and Zhao, 1993 (published 1994). A new troodontid (Dinosauria, Theropoda) braincase from
the Dinosaur Park Formation (Campanian) of Alberta. Canadian Journal of Earth
Sciences. 30(10-11), 2234-2247.
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. Caenagnathasia
sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous:
Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.
Funston, Currie, Ryan and Dong, 2019. Birdlike growth and mixed-age
flocks in avimimids (Theropoda, Oviraptorosauria). Scientific Reports.
9:18816.
undescribed possible Dromaeosauridae (Prasad, Parmar and Kumar, 2014)
Early Jurassic
Kota Formation, India
Material- teeth
Comments- These were described as "dromaeosaurid-like, and Velociraptorinae-like."
Reference- Prasad, Parmar and Kumar, 2014. Recent vertebrate fossil discoveries
from the Jurassic Kota Formation of India. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 208.
unnamed possible Dromaeosauridae (Knoll and Ruiz-Omenaca, 2005)
Beriassian, Early Cretaceous
KM 1983, Ksar Metlili Formation, Morocco
Material- (MNHN SA 2004/3A; Velociraptorinae
indet. Morphotype I; lost) lateral tooth (3.20x2.20x.80 mm) (Knoll and Ruiz-Omenaca, 2005)
(MNHN SA mcm 158; Velociraptorinae
indet. Morphotype III; lost) tooth (4.32x?x.76 mm) (Knoll and Ruiz-Omenaca, 2005)
(MNHN SA mcm 168; Velociraptorinae
indet. Morphotype I; lost) tooth (1.28x1.20x.56 mm) (Knoll and Ruiz-Omenaca, 2005)
Beriassian, Early Cretaceous
KM-A1, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-A1-10) tooth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
Beriassian, Early Cretaceous
KM-A2, Ksar Metlili, Ksar Metlili Formation, Morocco
?(FSAC-KM-A2-9 [1]; Theropoda gen. et sp. indet. morphotype II) lateral tooth (3.34x1.82x1.22 mm) (Lasseron, 2020)
Beriassian, Early Cretaceous
KM-B', Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-B'-26) tooth (Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
Beriassian, Early Cretaceous
KM-D2, Ksar Metlili, Ksar Metlili Formation, Morocco
(FSAC-KM-D2-11; Theropoda gen. et sp. indet. morphotype II) lateral tooth (9.93x5.49x3.23 mm), tooth (Lasseron, 2020;
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020)
Comments- The MNHN specimens were collected in 1983, 1986 and 1999, while the
FSAC specimens were collected in 2010, 2015 and 2018 (Lasseron, 2020). Lasseron noted the "dinosaur remains (Knoll, 2000; Knoll & Ruiz-Ome�aca, 2009) ... were taken out of the MNHN and lost." The MNHN teeth were referred to Dromaeosauridae by Knoll and Ruiz-Omenaca
(2009) based on their high DSDI (1.08-1.45). However, this is also found in many more basal
coelurosaurs. While Knoll and Ruiz-Omnaca (2005) originally noted teeth that
"could be related to "paronychodontids"", the eventual 2009
publication shows this only refers to GNHN SA mcm 158, which is their Velociraptorinae
indet. Morphotype III and has serrated carinae unlike Paronychodon.
Knoll and Ruiz-Omenaca considered Morphotype I similar to Deinonychus while Morphotype III is close to specimens now recognized as Saurornitholestes premaxillary teeth (Zapsalis, Sankey et al.'s ?Dromaeosaurus
morphotype A). Lasseron et al. (2020) stated "it is possible that those potential
dromaeosaurid teeth from the KM fauna belong instead to Noasauridae or
a basal tyrannosauroid (C. Hendrickx, pers. comm. to ML, March 2018)." Lasseron's
Theropoda gen. et sp. indet. morphotype II is placed here as well due
to the high DSDI (1.67-1.75). Lasseron finds quantitative
analysis classifies FSAC-KM-A2-9 [1] as the contemporaneous Nuthetes,
perhaps indicating a basal tyrannosauroid position if that genus is
correctly placed here. Note Lasseron listed FSAC-KM-D2-11 as both
a 'Theropoda gen. et sp.
indet. morphotype I' and 'Theropoda gen. et sp. indet. morphotype II'
tooth, without using a number in brackets to designate multiple teeth
falling under one specimen number. As the mesial serration
density is
listed, the measurements given belong to a morphotype II tooth.
Lasseron et al. lists all four teeth under FSAC-KM-B'-26 as
Dromaeosauridae, although Lasseron described three as Theropoda gen. et
sp. indet. morphotype III, Coelurosauria gen. et sp. indet. morphotype
I and Coelurosauria gen. et sp. indet. morphotype III (unnamed
Averostra and Coelurosauria here). Similarly, they list
FSAC-KM-D1-6 as dromaeosaurid, but it was described as Coelurosauria
gen. et sp. indet. morphotype I by Lasseron and is catalogued under
unnamed Coelurosauria here. Lasseron et al. list both
FSAC-KM-A2-9 teeth as Maniraptora, but they were described as Theropoda
gen. et sp. indet. morphotypes I and II by Lasseron.
References- Knoll and Ruiz-Omenaca, 2005. Theropod teeth from the Berriasian
of Anoual (Morocco). Journal of Vertebrate Paleontology. 25(3), 78A.
Knoll and Ruiz-Omenaca, 2009. Theropod teeth from the basalmost Cretaceous of
Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine.
146(4), 602-616.
Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.
Lasseron, Allain, Gheerbrant, Haddoumi, Jalil, M�tais, Rage, Vullo and
Zouhri, 2020 (online 2019). New data on the microvertebrate fauna from
the Upper Jurassic or lowest Cretaceous of Ksar Metlili (Anoual
Syncline, eastern Morocco). Geological Magazine. 157, 367‑392.
undescribed Dromaeosauridae (Sadleir, 1998)
Cenomanian, Late Cretaceous
Kem Kem Beds, Morocco
Material- (GZG.V.19997) tooth (13.8x7.5x3.8 mm) (Richter, Mudroch and Buckley,
2013)
?(GZG.V.19998) tooth (12.5x7.3x3.1 mm) (Richter, Mudroch and Buckley, 2013)
(M-CH-009) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
(M-JQ-012) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
(M-KS-015) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
(M-ZA-014) tooth (Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004)
?(NMB-1671-R) tooth (14.5x7.5x3.8 mm) (Richter, Mudroch and Buckley, 2013)
Comments- The teeth reported by Amiot et al. (2004) are 10-16 mm in height,
and strongly recurved with BW/FABLs of 0.4-0.6. Mesial serrations are present
on all but M-CH-009 and are small (3.4-4/mm). Distal serrations are larger (2.4-3.8/mm)
and apically inclined. DSDI ranged between 1.11-1.42. Interdenticle slits are
deep. They referred these specimens to Velociraptorinae.
References- Sadleir, 1998. Theropod teeth from the Cretaceous of Morocco.
Journal of Vertebrate Paleontology. 18(3), 74A.
Amiot, Buffetaut, Tong, Boudad and Kabiri, 2002. Laurasian theropod dinosaur
teeth from the Late Cretaceous of Morocco. Conference abstract, Third Georges
Cuvier Symposium, Montbeliard, France.
Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004. Isolated theropod teeth from
the Cenomanian of Morocco and their palaeobiogeographical significance. Revue
de Paleobiologie, Geneve. 9, 143-149.
Richter, Mudroch and Buckley, 2013. Isolated theropod teeth from the Kem Kem
Beds (Early Cenomanian) near Taouz, Morocco. Palaontologische Zeitschrift. 87,
291-309.
unnamed dromaeosaurid (Werner, 1994)
Cenomanian, Late Cretaceous
Wadi Milk Formation, Sudan
Material- (Vb-713) pedal phalanx II-2 (36 mm) (Werner, 1994)
(Vb-714) proximal manual ungual (Werner, 1994)
(Vb-860) distal manual ungual (Rauhut and Werner, 1995)
(Vb-866) distal pedal ungual I (Rauhut and Werner, 1995)
(Vb-867) ungual (Rauhut and Werner, 1995)
(Vb-868) proximal pedal ungual III (Rauhut and Werner, 1995)
(Vb-875) tooth (8.3 mm) (Rauhut and Werner, 1995)
Comments- These unassociated remains were found in 1991 and 1992, and
first reported by Werner (1994) before being described in depth by Rauhut and
Werner (1995).
Vb-714 was originally identified as a pedal ungual II, but is more likely a
manual ungual based on the high dorsal arch and large flexor tubercle. The similar
partial ungual Vb-860 (also assigned to pedal digit II by Rauhut and Werner)
may also be from the manus then. Vb-867 was also identified as a pedal ungual
II, but was not illustrated. The high DSDI (1.33) was used to place this in
the Velociraptorinae, but are also known in microraptorians and basal dromaeosaurines.
It does exclude the Sudanese taxon from the Dromaeosaurus+Achillobator+Utahraptor
subclade, as does the elongate pedal phalanx II-2.
References- Werner, 1994. Die kontinentale Wirbeltierfauna aus der unteren
Oberkreide des Sudan (Wadi Milk Formation). Berliner geowiss. Abh.. 13, 221-249.
Rauhut and Werner, 1995. First record of the family Dromaeosauridae (Dinosauria:
Theropoda) in the Cretaceous of Gondwana (Wadi Milk Formation, northern Sudan).
Palaeontologische Zeitschrift. 69(3/4), 475-489.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
unnamed possible dromaeosaurid (Fanti and Therrien, 2007)
Middle Maastrichtian, Late Cretaceous
Anembalemba Member of Maevarano Formation, Madagascar
Material- (MSNM V5365) premaxillary tooth
(MSNM V5372) premaxillary tooth
(MSNM V5373) lateral tooth
(MSNM V5394) premaxillary tooth
(MSNM V5590) premaxillary tooth
(MSNM V coll.) 13 teeth (8-15 mm)
Comments- These teeth are high compressed (BW/FABL ~.5), have 3-5.5 mesial
serrations and 3-3.5 distal serrations. Mesial serrations are apically inclined,
while distal serrations are perpendicular to the distal edge. All serrations
are slightly hooked apically and have absent or poorly defined interdenticle
slits.
These were referred to Dromaeosauridae based on a morphometric comparision of
several measurements to dromaeosaurids, abelisaurids, Masiakasaurus and
tyrannosaurids. Thus it is possible they belong to another theropod clade once
they are compared to a broader range of taxa.
Reference- Fanti and Therrien, 2007. Theropod tooth assemblages from
the Late Cretaceous Maevarano Formation and the possible presence of dromaeosaurids
in Madagascar. Acta Palaeontologica Polonica. 52(1), 155-166.
undescribed possible dromaeosaurid (Canale, Carballido, Otero, Canudo
and Garrido, 2014)
Albian-Cenomanian, Early Cretaceous-Late Cretaceous
Bayo Overo Member of the Cerro Barcino Formation, Chubut, Argentina
Material- tooth
Reference- Canale, Carballido, Otero, Canudo and Garrido, 2014. Carcharodontosaurid
teeth associated with titanosaur carcasses from the Early Cretaceous (Albian)
of the Chubut Group, Chubut Province, Patagonia, Argentina. Jornadas Argentinas
de Paleontologia de Vertebrados. Ameghiniana. 51(6) suplemento, 6.
undescribed Dromaeosauridae (Casal, Martinez, Candeiro, Lamanna and
Ibiricu, 2007)
Mid Cenomanian-Turonian, Late Cretaceous
Lower Bajo Barreal Formation, Chubut, Argentina
Material- three teeth
Reference- Casal, Martinez, Candeiro, Lamanna and Ibiricu, 2007. First
record of Dromaeosauridae (Dinosauria: Theropoda) in the Early Late Cretaceous
Bajo Barreal Formation of Chubut Province, Argentina. Journal of Vertebrate
Paleontology. 27(3), 56A.
unnamed possible Dromaeosauridae (Elias, Bertini and Medeiro, 2007)
Late Albian-Early Cenomanian, Early-Late Cretaceous
Alcantara Formation, Brazil
Material- (UFMA 1.20.194-1) tooth (16.1x7.3x3.4 mm)
(UFMA 1.20.194-2) incomplete tooth (13.1x7.9x3.8 mm)
Comments- These were referred to Velociraptorinae by Elias et al. (2007),
but lack e.g. a high DSDI or 8-shaped basal section.
Reference- Elias, Bertini and Medeiro, 2007. Velociraptorinae (Maniraptoriformes)
teeth from the Coringa Flagstone outcrop, Middle Cretaceous of the Sao Luis-Grajau
basin, Maranhao state, northern-northeastern Brazil. in Carvalho, Cassab, Schwanke,
Carvalho, Fernandes, Rodrigues, Carvalho, Arai and Oliveira (eds.). Paleontologia:
Cen�rios de Vida. 1, 315-325.
unnamed possible Dromaeosauridae (Franco-Rosas, 2002)
Turonian-Late Maastrichtian, Late Cretaceous
Adamantina, Marilia and/or Serra da Galga Formations of the Bauru Group, Brazil
Material- teeth
Comments- These teeth have long, pointed distal serrations with deep
interdenticle slits. They are said to be similar to dromaeosaurids, and the
description matches basal forms more than derived dromaeosaurines.
Reference- Franco-Rosas, 2002. Methodological parameters for the identification
and taxonomic classification of isolated theropodomorph teeth. Anais da Academia
Brasileira de Ciencias. 74(2), 367.
Dromaeosauridae sensu Norell and Makovicky, 2004
Definition- (Microraptor zhaoianus + Sinornithosaurus millenii
+ Velociraptor mongoliensis)
Microraptoria Senter et al., 2004
Definition- (Microraptor zhaoianus <- Velociraptor mongoliensis,
Dromaeosaurus albertensis) (modified from Senter et al., 2004)
= Microraptorinae sensu Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Microraptor zhaoianus <- Dromaeosaurus albertensis)
= "Microraptorinae" Makovicky, Apestegu�a and Agnol�n,
2005
= Microraptorinae Longrich and Currie, 2009
Definition- (Microraptor zhaoianus <- Velociraptor mongoliensis,
Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus) (Turner
et al., 2012)
Other definition- (Microraptor zhaoianus <- Dromaeosaurus albertensis) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
Comments- Senter et al. (2004) erected Microraptoria, with a non-family
level suffix to ensure ICZN rules are followed whether or not the clade was
within Dromaeosauridae (depending on both the tree's topology and the latter's
definition). They listed Microraptoridae and Microraptorinae as alternatives
considered prior to deciding upon Microraptoria, and Microraptorini as a mispelling
of Microraptoria. Makovicky et al. (2005) cited Senter et al. for Microraptorinae,
but the latter was a misspelling of their own (Headden, pers. comm. to Makovicky,
2005). Turner et al. (2007) followed Makovicky et al. in using Microraptorinae,
but neither reference contains a diagnosis or definition so the name is a nomen
nudum (ICZN Article 13.1.1). While Makovicky et al. did reference Senter et
al.'s paper, which included a definition for Microraptoria, the latter is not
an available name under the ICZN so Article 13.1.3 does not apply. In addition,
ICZN Article 11.5 states names must be proposed as being valid, so Senter et
al. 2004 cannot be cited as the authors for Microraptorinae/idae, contra most
authors discussed here. This leaves Longrich and Currie (2009) as the first
publication to validy erect Microraptorinae. Turner et al. (2012) gave Microraptorinae
a more restrictive definition that supposedly overcomes Senter et al.'s perceived
difficulties with family-level names. However, in topologies such as Mayr et
al.'s (2005), Microraptorinae does not fall under any named family, which was
just the problem Senter et al. anticipated when they rejected the name. On the
other hand, Microraptoria includes Confuciusornis and potentially all
pygostylians in Mayr et al.'s topology, unlike Microraptorinae. So Turner et
al.'s definition is more stable due to its extra specifiers. At present, however,
the consensus is that Microraptorinae and Microraptoria are synonymous, and
the latter has priority.
References- Senter, Barsbold, Britt and Burnham, 2004. Systematics and
evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8,
1-20.
Makovicky, Apestegu�a and Agnol�n, 2005. The earliest dromaeosaurid
theropod from South America. Nature. 437, 1007-1011.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
from the Late Cretaceous of North America. Proceedings of the National Academy
of Sciences. 106(13), 5002-5007.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Tianyuraptor Zheng, Xu, You,
Zhao and Dong, 2010
= "Tianyuraptor" Zheng, Xu, You, Zhao and Dong, 2009 online
= "Zhenyuanlong" Lu and Brusatte,
2015
T. ostromi Zheng, Xu, You, Zhao and Dong, 2010
= "Tianyuraptor ostromi" Zheng, Xu, You, Zhao and Dong, 2009 online
= "Zhenyuanlong suni" Lu and Brusatte, 2015
Early Aptian, Early Cretaceous
Dawangzhangzi Beds of Yixian Formation, Liaoning, China
Holotype- (STM1-3) (subadult) (12.5 kg) skull, mandible, six cervical vertebrae, thirteen
dorsal vertebrae, dorsal ribs, uncinate processes, gastralia, sacrum, twenty-five
caudal vertebrae (caudal series 960 mm), chevrons, scapulae (121 mm), coracoid,
partial furcula, sternal plate, humeri (138 mm), radii (104 mm), ulnae, semilunate
carpals fused to metacarpals II, metacarpals I, phalanges I-1, manual unguals
I, phalanges II-1, phalanges II-2, manual unguals II, metacarpals III, phalanges
III-1, phalanges III-2, phalanges III-3, manual unguals III, ilia (~159 mm),
pubes, ischia, femora (212.5 mm), tibiae (283 mm), fibula, astragalus, calcaneum,
metatarsal I(?), phalanx I-1(?), metatarsals II, metatarsals III, phalanx III-2,
phalanx III-3, pedal ungual III, metatarsals IV, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal ungual IV, pedal phalanges, pedal ungual, metatarsal V
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Referred-
(JPM-0008; intended holotype of "Zhenyuanlong suni") (~1.65 m, 11.8 kg,
subadult) incomplete skull (166.1 mm), mandible (165.2 mm), hyoid, ten
cervical vertebrae (30.3 mm), dorsal series (250 mm; best preserved
23.7 mm), dorsal ribs, gastralia, six sacral vertebrae, first caudal
vertebra (23.6 mm), second caudal vertebra (23.6 mm), third caudal
vertebra (23.6 mm), fourth caudal vertebra (29.7 mm), fifth caudal
vertebra (32.4 mm), sixth caudal vertebra (37.3 mm), seventh caudal
vertebra (43.4 mm), eighth caudal vertebra (40.7 mm), ninth caudal
vertebra (40.7 mm), tenth caudal vertebra (42.1 mm), eleventh caudal
vertebra (46.8 mm), twelfth caudal vertebra (46.8 mm), thirteenth
caudal vertebra, fourteenth caudal vertebra, fragmentary fifteenth
caudal vertebra, chevrons, scapulae (one partial), incomplete
coracoids, sternal plates, sternal ribs, humeri (121.1 mm), radii (96.6
mm), ulnae (102.9 mm), scapholunare, semilunate carpal, distal carpal
III, metacarpals I (19.5 mm), phalanges I-1 (40.5 mm), manual unguals I
(25.4 mm), metacarpals II (55.7 mm), phalanges II-1 (32.8 mm),
phalanges II-2 (39.2 mm), manual unguals II (31.7 mm), metacarpals III
(53 mm), phalanges III-1 (14.9 mm), phalanges III-2 (8.1 mm), phalanges
III-3 (24.7 mm), manual unguals III (22.8 mm), manual claw sheaths,
ilium (128.3 mm), pubes (155.7 mm), ischia (80.1 mm), femora (193.4
mm), tibiae (260.3 mm), fibulae (262.5 mm), astragalocalcanea, distal
tarsal III, distal tarsal, metatarsals I (27.7 mm), phalanges I-1,
pedal unguals I, metatarsals II (128.3 mm), phalanx II-2, pedal ungual
II, metatarsals III (129.8 mm), phalanges III-1, phalanx III-2, phalanx
III-3, pedal ungual III, metatarsals IV (128.2 mm), phalanx IV-2,
phalanx IV-3, phalanx IV-4, pedal ungual IV, pedal phalanges, body
feathers, remiges, retrices
Early Cretaceous
Jehol Group, China
(PMOL coll.) (2.5 m) complete skeleton (Makovicky, Gorscak and Zhou, 2018)
Diagnosis- (after Zheng et al., 2010) small and extremely slender furcula;
elongated hindlimb about three times as long as dorsal series.
(after Lu and Brusatte, 2015) short forelimb (humerofemoral ratio <0.65, ulnofemoral ratio <0.55, manuofemoral
ratio <0.90; actual values 0.63, 0.53 and 0.76; values of Tianyuraptor
holotype 0.65, 0.49 and 0.86; with the humerofemoral ratio barely excepted) (also in Achillobator - ?, 0.51 and ?).
Other diagnoses- Turner et al. (2012) note Zheng et al.'s character of
'length of mid caudal vertebrae more than twice that of dorsal vertebrae' is
present in Microraptor as well.
Lu and Brusatte (2015) listed several characters in
their diagnosis. Adasaurus,
Velociraptor and Deinonychus also have six sacral vertebrae. Graciliraptor, Microraptor, Velociraptor,
Achillobator and some Deinonychus have a radius thinner than manual
phalanx I-1. Metacarpal II is shorter than metacarpal I plus manual phalanx
I-1 in eudromaeosaurs (Velociraptor, Tsaagan, Deinonychus),
Bambiraptor, Tianyuraptor (contra Lu and Brusatte) and some Microraptor
(LPM 0159). Lu and Brusatte stated the posteriorly
curved pubis differs from the straight element in Tianyuraptor, but the
latter is also posteriorly curved. While they said the ilium of Tianyuraptor
differs from "Zhenyuanlong" in sharing a lobate brevis shelf with Microraptor,
this seems to be absent in Tianyuraptor's type based on available low resolution
photos. They also said "Zhenyuanlong" differs from Tianyuraptor
in having a ridge along the ventral antorbital fossa edge, but the former has
a narrow and convex surface ventral to the fossa, not a ridge compared to the
rest of the external surface. Lu and Brusatte also included two characters shared
with Tianyuraptor and eudromaeosaurs- mid-pubic tubercle absent (plesiomorphic),
and mid dorsal ischial process absent (also in Unenlagia and Buitreraptor).
Comments- The description was released online in August 2009 but not
officially published until January 2010.
The holotype of "Zhenyuanlong" was collected by a farmer prior to 2015. Lu and
Brusatte (2015) believed the Sihedang locality, which it derives
from, to be in the Yixian Formation. It is here assigned to the Jiufotang Formation instead (see Iteravis entry). The taxon was described by Lu and Brusatte on
July 16 2015 as a new genus
of dromaeosaurid. However, this paper has no mention of ZooBank
and as of February 6 2020
"Zhenyuanlong" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Zhenyuanlong suni" Lu and
Brusatte 2015 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
Valid differences between Tianyuraptor and "Zhenyuanlong" noted by Lu and
Brusatte are limited to- radius thinner than manual phalanx I-1; smaller manus
(76% of femoral length vs. 86%). These could easily be explainable as individual
variation, as the former varies in Deinonychus.
Makovicky et al. (2018) noted a new specimen which "exhibits a
combination of traits that were previously used to distinguish Tianyuraptor from Zhenyuanlong."
Zheng et al. (2010) found Tianyuraptor to be more closely related to
Dromaeosaurus than Unenlagia, but outside previously described
microraptorians, and Saurornitholestes plus Velociraptor plus
dromaeosaurines. It was recovered as a microraptorian by Senter et al. (2012),
Turner et al. (2012) and Lee et al. (2014), but in a polytomy with microraptorians
and more derived dromaeosaurids by Brusatte et al. (2014).
Lu and Brusatte added "Zhenyuanlong" to Turner's TWiG analysis and found
it to be in a polytomy with the microraptorian genera, Tianyuraptor and
Eudromaeosauria.
References- Zheng, Xu, You, Zhao and Dong, 2010 (online 2009). A short-armed dromaeosaurid
from the Jehol Group of China with implications for early dromaeosaurid evolution.
Proceedings of the Royal Society B. 277, 211-217.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Lu and Brusatte, 2015. A large, short-armed, winged dromaeosaurid
(Dinosauria: Theropoda) from the Early Cretaceous of China and its implications
for feather evolution. Scientific Reports. 5, 11775.
Makovicky, Gorscak and Zhou, 2018. A new specimen of the large-bodied dromaeosaurid Tiuanyuraptor
provides new insights on microraptorine anatomy, taxonomy, and plumage
evolution. Journal of Vertebrate Paleontology. Program and Abstracts
2018, 174.
undescribed dromaeosaurid (Li, 2013)
Aptian-Albian, Early Cretaceous
Zhonggou Formation, Gansu, China
Material- (FRDC-GJ(07)11-2) (~2
m) frontal (63.5 mm), mid dorsal vertebra (30 mm), incomplete mid
dorsal vertebra (28.5 mm), posterior dorsal vertebra (26.5 mm),
posterior dorsal vertebra (23 mm), sacrum (32.5, 21.5, ~20, ~21.5,
~22.5, ~23 mm), ~fifth caudal vertebra (19 mm), distal caudal vertebra
(31.5 mm), distal caudal vertebra (26.5 mm), incomplete scapula,
humerus (138 mm), manual ungual II (30.5 mm), phalanx III-3 (41.5 mm),
manual ungual III (44 mm), ventral fragment of ilium, proximal pubis,
proximal ischium, phalanx II-1 (29.5 mm), phalanx II-2 (28 mm), distal
metatarsal III(?), phalanx III-1 (53.5 mm), phalanx III-3 (33.5 mm),
phalanx IV-4 (27.5 mm)
Comments- Discovered in 2007,
this was described as a new taxon in Li's (2013) and Mariko's (2015)
theses
and recovered as a basal velociraptorine in 2013 based on Senter's TWiG
matrix and a eudromaeosaur incertae sedis in 2015 based on Turner's
TWiG matrix. When entered into the Hartman et al. maniraptoromorph
matrix, FRDC-GJ(07)11-2 falls out as a microraptorian sister to "Zhenyuanlong".
References- Li, 2013. A new
dromaeosaurid dinosaur from the Early Cretaceous of Yujingzi basin,
Jiuquan area, Gansu Province, China. Masters Thesis, China University of Geosciences. 120 pp.
Mariko, 2015. A study of the new dromaeosaurid dinosaur from the
Early Cretaceous of Jiuquan area, Gansu Province, China. Masters
Thesis, China University of Geosciences. 112 pp.
"Paleopteryx" Jensen, 1981
"P. thomsoni" Jensen, 1981
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US (Dry Mesa quarry)
Material- (BYU 2022) distal radius
Comments- Jensen (1981) described the holotype (BYU 2022) as a proximal
tibiotarsus of a new taxon of bird- "Paleopteryx thomsoni". In addition,
he referred a proximal femur (BYU 2023) to Archaeopteryx, and another
femur (BYU 2025) and sacrum (BYU 2024) as avian-like. Molnar (1985) showed that
"Paleopteryx" was not diagnosed, so is a nomen nudum (ICZN Article
13.1.1). He assigned the BYU material to Theropoda. Jensen and Padian reidentified
BYU 2022 as a distal radius, and referred both it and BYU 2023 to Maniraptora
indet.. They made BYU 2024 the holotype of a new pterosaur genus Mesadactylus,
and referred BYU 2025 to that taxon as well.
"Paleopteryx" actually has an extremely distinctive distal radius,
with a medially flared articular surface seen in dromaeosaurids, Sapeornis
and Confuciusornis, but absent in therizinosaurs, oviraptorosaurs, Yixianosaurus,
Sinornithoides, Archaeopteryx, Rahonavis or Shenzhouraptor.
The high angle of flaring relative to the shaft is shared with Graciliraptor
and Microraptor among dromaeosaurids, unlike Deinonychus and Bambiraptor.
It is more similar to Graciliraptor, in that both lack Microraptor's
apomorphic concave surface just distal to the flare. An additional character
shared with Microraptor but not other maniraptorans (unknown in Graciliraptor)
is a distally projecting lateral process. "Paleopteryx" is thus here
assigned to Microraptoria.
References- Jensen, 1981. [A new oldest bird?] Anima (Tokyo). 1981, 33-39.
[in Japanese]
Jensen, 1981. Another look at Archaeopteryx as the worlds oldest bird.
Encyclia, The Journal of the Utah Academy of Sciences, Arts, and Letters. 58,
109-128.
Molnar, 1985. Alternatives to Archaeopteryx; a survey of proposed early
or ancestral birds. in Hecht, Ostrom, Viohl and Wellnhofer (eds). The Beginnings
of Birds. Eichstatt, Germany: Freunde des Jura-Museums Eichstatt. 209-217.
Jensen and Padian, 1989. Small Pterosaurs and Dinosaurs from the Uncomphagre
fauna (Brushy Basin Member, Morrison Formation: ?Tithonian), Late Jurassic,
Western Colorado. Journal of Paleontology. 63(3), 364-373.
Padian, 1998. Pterosaurians and ?avians from the Morrison Formation (Upper Jurassic,
Western U.S.). Modern Geology. 23, 57-68.
Shanag Turner, Hwang and Norell, 2007
S. ashile Turner, Hwang and Norell, 2007
= Sinornithosaurus ashile (Turner, Hwang and Norell, 2007) Paul, 2010
Berriasian-Barremian, Early Cretaceous
Oosh, Huhteeg Svita, Mongolia
Holotype- (IGM 100/1119) incomplete maxilla, incomplete dentary, partial
splenial
Diagnosis- (after Turner et al., 2007) triangular, anteriorly tapering
maxilla; lateral lamina of nasal process of maxilla reduced to small triangular
exposure; absence of a promaxillary fenestra; presence of interalveolar pneumatic
cavities; incipient dentary groove on posterolateral surface of dentary.
Comments- This specimen was discovered in 1999, first noted by Turner
et al. (2006) then described in Turner et al. (2007). The latter analysis found
it in an unresolved position among Microraptoria+Eudromaeosauria, though noted
greatest similarity to Sinornithosaurus. It has since been recovered
as a microraptorian (Longrich and Currie, 2009) or sister to Microraptoria+Eudromaeosauria
(Senter et al., 2012).
References- Turner, Pol, Norell and Hwang, 2006. Resolving dromaeosaurid
phylogeny: New information and additions to the tree. Journal of Vertebrate
Paleontology. 26(3), 133A.
Turner, Hwang and Norell, 2007. A small derived theropod from Oosh, Early Cretaceous,
Baykhangor Mongolia. American Museum Novitates. 3557, 27 pp.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
from the Late Cretaceous of North America. Proceedings of the National Academy
of Sciences. 106(13), 5002-5007.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Richardoestesia Currie, Rigby and Sloan, 1990
= Asiamericana Nessov, 1995
Diagnosis- very small serrations on teeth (5-12/mm); apical half of anterior
dentary teeth convex distally.
Comments- This genus is known from a large amount of teeth, spanning
the Bathonian to Maastrichtian of North and South America, Europe and Asia.
The only known nondental remains are the holotype dentaries, though it's probable
some of the many unidentified small theropod postcranial remains so far discovered
belonged to individuals with Richardoestesia-type teeth. The wide temporal
and geographic range suggests Richardoestesia as currently diagnosed
is a clade comparable to "families" in scope, and quite possibly polyphyletic.
The two named species, R. gilmorei and R. isosceles, show variation
between different formations that probably indicate they each contain several
species. R. cf. gilmorei is indistinguishable from Saurornitholestes
in the Early Campanian Milk River Formation except for serration size, though
the possible presence of gilmorei-like teeth from earlier formations
may indicate this is not due to R. gilmorei evolving from a dromaeosaurid
in the Early Campanian. However, these earlier records are based on unpublished
observations from a single figure (Chipping Norton Formation) and a single specimen
which seems to have serrations too large for Richardoestesia (Woodbine
Formation). R. isosceles-like specimens have a more definitive pre-Campanian
record and are the only ones verified from Eurasia, where they extend back to
the Kimmeridgian of Portugal. Though the pre-Campanian American records should
be examined before any conclusions are made, it seems possible R. isosceles
originated much earlier than R. gilmorei and had little to do with that
species. It should also be noted that many poorly described records of Richardoestesia
may belong to other genera such as basal tyrannosauroids (e.g. Nuthetes,
Dilong) or other basal dromaeosaurids (e.g. Sinornithosaurus,
Shanag). Indeed, both Shanag and Sinornithosaurus have
very small serrations on their teeth, and some teeth whose apical half is convex
distally. Hendrickx and Mateus (2014) found Richardoestesia to be a dromaeosaurid
in their phylogenetic analysis.
Richardoestesia or Ricardoestesia?- Olshevsky and others
have noted that the spelling of this genus was originally supposed to be Ricardoestesia,
and was misspelled in all cases except for a figure caption in the original
description by an editor. Thus, Olshevsky believes Ricardoestesia should
be the correct spelling. However, he acted as first revisor in 1991, listing
Richardoestesia as the correct spelling and Ricardoestesia as
a misspelling. As Creisler (DML, 2002) noted- "Under ICZN Art. 24.2.3,
the first author to have cited two alternate spellings of a name together and
to have selected one spelling as correct qualifies as the First Reviser. Under
32.5.1, this spelling could only be changed if it is determined to be "incorrect"--meaning
there is evidence of an inadverent error in the original publication itself
"without recourse to any external source of information." Crucially,
"incorrect transliteration or latinization" is not considered an inadvertent
error.
As I pointed out back in Feb. 2001, "Richardus" is a perfectly good
latinization for Richard, and one widely used in Medieval and later Latin literature.
Since the name was intended to honor Richard Estes, the latinized form Richardoestesia
is perfectly good and not in any way readable as "inadvertent error."
If one of the authors of the original paper wanted Ricardoestesia instead,
his wish qualifies as an "external source of information" which can't
be used, since it's not mentioned in the original paper. Moreover, the senior
author on the original Richardoestesia paper was Phil Currie, who has
used the spelling Richardoestesia in other papers he has authored or
coauthored.
Frankly, I can't find any basis in the ICZN for switching the spellings in later
editions of Mesozoic Meanderings. George had in fact already been the First
Reviser on the name in Oct. 1991. Since generic names differing by one letter
are not homonyms, the coexistence of Richardoestesia and Ricardoestesia
for the same taxon is becoming more confusing as time goes on, especially with
electronic data retrieval and online databases. As the situation is shaping
up, nearly all technical papers are using Richardoestesia and a few books
have used Ricardoestesia. Maybe the ICZN will have to make a decision,
but it seems like a lot of fuss over an issue that was settled just fine by
George in 1991."
Not Richardoestesia- Richardoestesia was reported from
the Campanian Fort Crittenden Formation of Arizona by Ratkevich and Duffek (1996),
but reidentified merely as Dromaeosauridae indet. by Sullivan and Lucas
(2006).
References- Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith
River Formation of southern Alberta, Canada. in Carpenter and Currie (eds.).
Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press,
New York. pp. 107-125.
Nessov, 1995. Dinozavri severnoi Yevrasii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii [Dinosaurs of Northern Eurasia: new data about
assemblages, ecology and paleobiogeography], Scientific Research Institute of
the Earth's Crust, St. Petersburg State University, St. Petersburg, Russia:
156 pp. + 14 pl. [in Russian with short English, German, and French abstracts].
Ratkevich and Duffek, 1996. Small macro-and large micro-vertebrate fauna of
the Fort Crittenden Formation, Southeast Arizona. Proceedings of Southwest Paleontological
Society and Mesa Southwest Museum, Mesa, Arizona. 4, 115-120.
Creisler, DML 2002. https://web.archive.org/web/20200806190728/http://dml.cmnh.org/2002Jul/msg00530.html
Sullivan and Lucas, 2006. The Kirtlandian land-vertebrate "age" -
faunal composition, temporal position and biostratigraphic correlation in the
nonmarine Upper Cretaceous of western North America. New Mexico Museum of Natural
History and Science Bulletin. 35, 7-29.
Hendrickx and Mateus, 2014. Abelisauridae (Dinosauria: Theropoda) from the Late
Jurassic of Portugal and dentition-based phylogeny as a contribution for the
identification of isolated theropod teeth. Zootaxa. 3759(1), 1-74.
R. gilmorei Currie, Rigby
and Sloan, 1990
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, US
Holotype- (CMN 343) partial dentaries, teeth
Paratypes- (RTMP 83.45.2) sixth dentary tooth
(RTMP 80.8.298) tooth
(RTMP 80.16.1230) tooth
(RTMP 83.129.11) tooth
(RTMP 84.89.274) tooth
Referred- (RTMP 65.26.13) tooth (Baszio, 1997)
(RTMP 81.16.194) premaxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.16.173) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 83.36.233) tooth (12.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 83.36.242) tooth (8.3 mm) (Baszio, 1997)
(RTMP 84.92.268) tooth (~5.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.59.204) tooth (Baszio, 1997)
(RTMP 86.23.90) tooth (10 mm) (Baszio, 1997)
(RTMP 86.23.105) tooth (Baszio, 1997)
(RTMP 86.159.60) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.171.9) tooth (4.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.171.61) tooth (3.4 mm) (Baszio, 1997)
(RTMP 87.116.60) tooth (Baszio, 1997)
(RTMP 87.80.35) tooth (7.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.91.28) tooth (5.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.156) tooth (Baszio, 1997)
(RTMP 89.36.355) tooth (10 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.76.63) tooth (5.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.106.6) tooth (4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.157.29) tooth (2.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.30b) tooth (3.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(ANSP 15824) tooth (Fiorillo and Currie, 1994)
(ANSP 15942) tooth (Fiorillo and Currie, 1994)
(ANSP 15952) tooth (Fiorillo and Currie, 1994)
(ANSP 17647) tooth (Fiorillo and Currie, 1994)
(ANSP 17782) tooth (Fiorillo and Currie, 1994)
(ANSP 17784) tooth (Fiorillo and Currie, 1994)
(ANSP 17983) tooth (Fiorillo and Currie, 1994)
(ANSP 18101) tooth (Fiorillo and Currie, 1994)
(ANSP 18104) tooth (Fiorillo and Currie, 1994)
(ANSP 18114) tooth (Fiorillo and Currie, 1994)
(ANSP 18115) tooth (Fiorillo and Currie, 1994)
(ANSP 18119) tooth (Fiorillo and Currie, 1994)
Middle Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
(RTMP 83.33.17) tooth (7.6x4.4x2.2 mm) (Larson and Currie, 2013)
(RTMP 83.45.11) tooth (6.6x4.4x2.2 mm) (Larson and Currie, 2013)
(RTMP 90.82.15) tooth (11.7x6.2x2.6 mm) (Larson and Currie, 2013)
(RTMP 94.28.1) tooth (10.2x5.3x2.5 mm) (Larson and Currie, 2013)
(RTMP 96.29.2) tooth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
(RTMP 96.39.29a) tooth (3.9x3x1.2 mm) (Larson and Currie, 2013)
(RTMP 97.39.2) tooth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
(RTMP 99.50.113) tooth (Larson et al., 2010)
(RTMP 2000.45.80) tooth (11.9x6.l4x2.7 mm) (Larson et al., 2010)
(RTMP 2001.45.82) tooth (8.9x3.9x2.2 mm) (Larson et al., 2010)
(RTMP 2002.45.49) tooth (4.7x3x1.3 mm) (Larson et al., 2010)
(RTMP 2002.45.53) tooth (4.3x3.4x1.3 mm) (Larson et al., 2010)
(RTMP 2003.15.2) tooth (Larson et al., 2010)
(RTMP 1016) tooth (Baszio, 1997)
(RTMP 1017) tooth (Baszio, 1997)
(RTMP 1018) tooth (Baszio, 1997)
(RTMP 1026) tooth (Baszio, 1997)
(RTMP 1027) tooth (Baszio, 1997)
(RTMP coll.) nine teeth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
(LSUMG V-6237) partial tooth (Sankey, 2001)
Late Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
(RAM 24759) tooth (Farke, Letteau Stallings and Andrews, 2020)
Diagnosis- (from Sankey et al., 2002) differs from R. isosceles
in- teeth generally shorter, more recurved and larger; serrations not square
shaped, and with small interdenticle spaces; some teeth constricted at base.
differs from Shanag in- shallow Mackelian groove; some teeth constricted
at base; all teeth have distal serrations; some teeth have mesial serrations.
differs from Sinornithosaurus in- shallow Mackelian groove; Mackelian
groove placed more dorsally; unfused posterior dentary interdental plates; at
least some premaxillary teeth serrated distally; some teeth constricted at base;
anterior dentary teeth convex distoapically.
Comments- The holotype was discovered in 1917 and described in 1924 by
Gilmore as a provisionally referred specimen of Chirostenotes (now known
to be an oviraptorosaur whose mandibles were named Caenagnathus). Currie
et al. (1990) soon described it as a new genus after Currie and Russell (1988)
recognized Chirostenotes as an oviraptorosaur. Currie et al. did leave
open the possibility Richardoestesia could be synonymous with Chirostenotes
or Elmisaurus however.
Horseshoe Canyon Formation Richardoestesia gilmorei are said to be identical
to Judith River specimens by Baszio (1997), but taller by Larson et al. (2010).
Though R. gilmorei has been reported from the Cenomanian-Maastrichtian
of the US and Canada (e.g. Baszio, 1997), these are slightly different from
Judith River specimens when examined and it is likely they are separate species.
References- Gilmore, 1924. A new coelurid dinosaur from the Belly River
Cretaceous of Alberta. Canada Department of Mines Geological Survey Bulletin
(Geological Series). 38(43), 1-12.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.
Fiorillo and Currie, 1994. Theropod teeth from the Judith River Formation (Upper
Cretaceous) of south-central Montana. Journal of Vertebrate Paleontology. 14(1),
74-80.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998. Baby hadrosaurid material
associated with an unusually high abundance of Troodon teeth from the
Horseshoe Canyon Formation, Upper Cretaceous, Alberta, Canada. Gaia. 15, 123-133.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend,
Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe
Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta,
with special reference to the Albertosaurus sarcophagus bonebed. Canadian
Journal of Earth Sciences. 47(9), 1159-1181.
Larson and Currie, 2013. Multivariate analyses of small theropod dinosaur teeth
and implications for paleoecological turnover through time. PloS ONE. 8(1),
e54329.
Farke, Letteau Stallings and Andrews, 2020. New vertebrate localities
and biostratigraphic interpretations of the Mesaverde Formation
(Campanian, Late Cretaceous) in northwestern Wyoming. The Society of
Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 136-137.
R. cf. gilmorei sp. nov. (Baszio, 1997)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(UA MR-4: 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20) seventeen
teeth
Comments- Milk River Formation R. gilmorei-like teeth have a greater
variety of serration shape than Dinosaur Park examples (rectangular to slightly
pointed) and grade into Saurornitholestes sp. of the same formation.
They may indicate the R. gilmorei lineage descended from dromaeosaurids
in the Santonian-Early Campanian, or they may simply be an example of convergence.
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
(UA KUA-1: 85, 120, 108) three teeth
(UA LSF-140: 105) tooth
(UA KUA-18:114) tooth
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- teeth
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Krumenacker, Simon, Scofield and Varricchio,
2016)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2167/50102; Morph 6) incomplete tooth (?x3.1x1.2 mm)
Reference- Krumenacker, Simon, Scofield and Varricchio, 2016. Theropod
dinosaurs from the Albian-Cenomanian Wayan Formation of eastern Idaho. Historical
Biology. DOI: 10.1080/08912963.2015.1137913
R. cf. gilmorei (Stokosa, 2005)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Material- (FMNH PR2899) tooth (2.7x1.3x.8 mm) (Gates, Zanno and Makovicky,
2015)
(SDSM 64372b) tooth (Stokosa, 2005)
Comments- Gates et al. (2015) referred FMNH PR2899 to Dromaeosauridae,
as a "morphology not previously described in the literature." While
it differs from most Richardoestesia teeth in being fluted, the shape
and small serrations size (19 per mm) are similar. Although the authors provide
a PCA analysis which "found it to be markedly different from Richardoestesia
teeth", their figure indicates there are Richardoestesia specimens
which are equally distant outliers.
Reference- Stokosa, 2005. Enamel microstructure variation within the
Theropoda. in Carpenter (ed). The Carnivorous Dinosaurs. 163-178.
Gates, Zanno and Makovicky, 2015. Theropod teeth from the upper Maastrichtian
Hell Creek Formation "Sue" Quarry: New morphotypes and faunal comparisons.
Acta Palaeontologica Polonica. 60(1), 131-139.
R. cf. gilmorei (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
(BTB: 135, 154, 155, 158, 159, 160) six teeth
Reference- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
R. cf. gilmorei (Diem, 1999)
Campanian, Late Cretaceous
Williams Fork Formation, Colorado, US
Reference- Diem, 1999. Vertebrate faunal analysis of the Upper Cretaceous
Williams Fork Formation, northwestern Colorado. Unpublished Masters Thesis.
San Diego, San Diego State University. 188 pp.
R. cf. gilmorei (Lewis, Heckert, Lucas and Williamson, 2007)
Late Santonian-Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
Material- teeth
Reference- Lewis, Heckert, Lucas and Williamson, 2007. A diverse new
microvertebrate fauna from the Upper Cretaceous (Late Santonian-Early Campanian)
Menefee Formation of New Mexico. Journal of Vertebrate Paleontology. 27(3),
105A.
Lewis, Heckert and Lucas, 2008. A mixed marine/non-marine and terrestrial microvertebrate
assemblage from the Late Cretaceous (Late Santonian-Early Campanian) Menefee
Formation of New Mexico: Fauna, biostratigraphy, and paleoecology. Journal of
Vertebrate Paleontology. 28(3), 105A.
R. cf. gilmorei sp. nov. (Lee, 1995)
Cenomanian, Late Cretaceous
Woodbine Formation, Texas, US
Material- (SMU 73779) tooth
Comments- This is straight like R. gilmorei, but with larger serrations
(4/mm).
References- Lee, 1995. Mid-Cretaceous archosaur faunal changes in Texas.
in Sun and Wang (eds.). Sixth Symposium on Mesozoic Terrestrial Ecosystems and
Biota, Short Papers. China Ocean Press, Beijing. 143-146.
Lee, 1997. The Archosauria from the Woodbine Formation (Cenomanian) in Texas.
Journal of Paleontology. 71(6), 1147-1156.
R. cf. gilmorei (Metcalf and Walker, 1994)
Early Bathonian, Middle Jurassic
Chipping Norton Formation, England
Material- (GLRCM coll.) tooth (2.9 mm; FABL 1.5 mm)
Comments- This tooth was labeled as "dromaeosaur-like" by Metcalf
and Walker (1994).
Mesial serrations are absent, and the crown is elongate and recurved. Serrations
are fairly flat and not hooked apically, but are taller than wide. Serration
density is 19/mm. Blood grooves are not apparent.
The high serration density is characteristic of Richardoestesia, while
the serration size compared to FABL and curvature match R. gilmorei more
than R. isosceles. No features distinguishing this tooth from R. gilmorei
can be determined, though it is smaller than most specimens of that species.
Reference- Metcalf and Walker, 1994. A new Bathonian microvertebrate
locality in the English Midlands. in Fraser and Sues (eds.). In the Shadow of
the Dinosaurs- Mesozoic Small Tetrapods, Cambridge (Cambridge University Press).
322-332.
R. cf. gilmorei (Hendrickx and Mateus, 2014)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Material- (ML 939) lateral tooth (5.1x2.8x1 mm)
Reference- Hendrickx and Mateus, 2014. Abelisauridae (Dinosauria: Theropoda)
from the Late Jurassic of Portugal and dentition-based phylogeny as a contribution
for the identification of isolated theropod teeth. Zootaxa. 3759(1), 1-74.
R. cf. gilmorei (Torices, 2002)
Late Campanian, Late Cretaceous
Vicari 4, Tremp Formation, Spain
Material- (DPM-VIR4-T6) tooth (1.5x1.4x.7 mm)
(DPM-VIR4-T7) tooth (2.2x1.6x.6 mm)
Comments- These were referred to Dromaeosauridae indet. 4 by Torices
(2002), and cf. Richardoestesia sp. by Torices et al. (2015).
References- Torices, 2002. Los dinosaurios ter�podos del Cret�cico
Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de
Paleontolog�a. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
R. cf. gilmorei (Buffetaut, Marandat and Sige, 1986)
Early Campanian, Late Cretaceous
Villeveyrac, Herualt, France
Material- (Universite des Sciences et Techniques de Languedoc VIC 17) tooth
(3.5 mm) (Buffetaut, Marandat and Sige, 1986)
?(Costa coll.) several teeth (Buffetaut, Costa, Le Loeuff, Martin, Rage, Valentin
and Tong, 1996)
Comments- VIC 17 is short and recurved, with 10 distal serraions per
mm, and 15 mesial serrations per mm. The distal serrations seem rounded and
perpendicular to the crown, while the mesial serrations are only present in
the apical portion. The BW/FABL is about .44. Though assigned to Dromaeosauridae
by Buffetaut et al. (1986), the small serration size and high DSDI make it almost
identical to Richardoestesia gilmorei. Its curvature, short crown, and
perhaps constricted base distinguish it from R. isosceles.
Buffetaut et al. (1996) later noted several small laterally compressed and serrated
teeth which they referred to small theropods, possibly dromaeosaurids. These
are tentatively retained here untl they are described further.
Reference- Buffetaut, Marandat and Sige, 1986. Decourvert de dents de
Deinonychosaures (Saurischia, Theropoda) dans le Creace superieur du Sud de
la France. Les Comptes rendus de l'Acad�mie 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'Acad�mie des sciences. 303, Serie
II(15), 1393-1396.
R? isosceles Sankey, 2001
Late Campanian, Late Cretaceous
Lower Aguja Formation, Texas, US
Holotype- (LSUMG 489:6238) tooth
Paratypes- (LSUMG 489:6233) tooth
(LSUMG 489:6234) tooth
(LSUMG 489:6235) tooth
(LSUMG 492:6264) tooth
Referred- (LSUMG 140:6140) tooth (Sankey, 2001)
(LSUMG 140:6051) tooth (Sankey, Standhardt and Schiebout, 2005)
(LSUMG 489:6050) tooth (Sankey, Standhardt and Schiebout, 2005)
(TMM 43057-313) tooth (Rowe, Cifelli, Lehman and Weil, 1992)
Late Santonian-Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
teeth (Lewis, Heckert and Forys, 2006)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(CMN coll.) teeth (Russell, 1935)
(UA MR-4: 36, 37, 38, 39, 40, 41) six teeth (Bazsio, 1997)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
(UW 34821) tooth (Demar and Breithaupt, 2006)
(UW 34822) tooth (Demar and Breithaupt, 2006)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(AMNH 8549) tooth (Sahni, 1972)
(AMNH coll.) eleven teeth (Sahni, 1972)
Late Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
(RTMP 96.62 coll.) <6 teeth (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
(RTMP 89.103.25) tooth (5.4 mm) (Sankey et al., 2002)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
(RTMP 84.1.12) tooth (~11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.36.97) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.23.105) tooth (5.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.33.54) tooth (~4.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.34.43) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.45.46) tooth (~5.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.159.62) tooth (~7.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.195.42) tooth (<5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.16.19) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.99.48) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.65) tooth fragment (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.157.52) tooth fragment (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.36.199) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.86.44) tooth (~8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.136.56) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.79.31) tooth (6.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.177.49a) tooth (~8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.177.49b) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.180.5a) tooth (5.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.180.5b) tooth (4.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.181.10) tooth (7.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.10.35) tooth fragment (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.48.11) tooth (~5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.25) tooth (<4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.30a) tooth (12.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.31a) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.62.31b) tooth (10.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.142.19) tooth (9.7 mm) (Ryan and Russell, 2001)
teeth (Baszio, 1997)
Early Maastrichtian, Late Cretaceous
Upper Aguja Formation, Texas, US
(LSUMG 113:5939) tooth (Sankey, Standhardt and Schiebout, 2005)
(LSUMG 741:5933) tooth (Sankey, Standhardt and Schiebout, 2005)
(LSUMG 741:5934) tooth (Sankey, Standhardt and Schiebout, 2005)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
teeth (Currie, Rigby and Sloan, 1990)
teeth (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, North Dakota, South Dakota, US
teeth (Currie, Rigby and Sloan, 1990)
teeth (Sankey, 2001)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
(RTMP 1019) tooth (Baszio, 1997)
teeth (Ryan and Russell, 2001)
Maastrichtian, Late Cretaceous
Ocozocoautla Formation, Mexico
(IHNFG-0537) maxillary tooth (11.6x7.3x4.8 mm) (Carbot-Chanona and Rivera-Sylva,
2011)
Late Maasstrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Paratype- (AMNH 8113) tooth (Estes, 1964)
Referred- (AMNH coll.) teeth (Estes, 1964)
(UA BTB: 144, 143) two teeth (Baszio, 1997)
teeth (Derstler, 1994, 1995)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, US
(UA LSF140: 107, 111, 112, 115, 116, 118) six teeth (Baszio, 1997)
Late Cretaceous
North America
Paratype- (RTMP 91.170.9) tooth
Diagnosis- (from Sankey et al., 2002) differs from R. gilmorei
in- teeth generally longer, straighter and smaller; serrations square-shaped,
and with larger interdenticle spaces; no teeth constricted at base.
differs from Shanag in- teeth straighter and generally longer; all teeth
have distal serrations; some teeth have mesial serrations.
differs from Sinornithosaurus in- teeth straighter and generally longer.
Comments- Russell (1935) first reported these kinds of teeth and hypothesized
they were tyrannosaurid ("deinodont") premaxillary teeth. Estes (1964)
believed them to be from juvenile theropods, while Sahni (1972) tentatively
assigned them to Sebecosuchia. They were first connected to Richardoestesia
gilmorei remains by Currie et al. (1990), who suspected they belonged
to a new species of that genus. It was referred to as Richardoestesia sp.
by Baszio (1997) and other sources until being named by Sankey in 2001. Sankey's
paratypes are from several formations, and the species has been reported from
the Early Campanian to the Late Maastrichtian of North America. These show very
little variation (Baszio, 1997) though we may assume more than one species was
present in this time interval. Additional similar teeth are known from the Late
Jurassic to Late Cretaceous of Eurasia and the Early Cretaceous of the US, but
differ slightly and are referred to new unnamed species below. R. isosceles
only shares a couple apomorphic features with R. gilmorei (tiny serrations,
convex distoapical edge of some teeth) and may not be closely related. The fact
its lineage extends back to Kimmeridgian Europe, while R. gilmorei seems
to emerge from dromaeosaurid teeth in the Early Campanian of North America may
support this view. However, Williamson and Brusatte (2014) note both gilmorei-like
and isosceles-like teeth from the Fruitland Formation have a high DSDI,
suggesting they derive from the same species. Sankey (2002) has suggested Paronychodon
and Richardoestesia teeth are morphotypes of the same taxon, based on
morphology and relative abundance. The details of this study have yet to be
published, though it does make sense stratigraphically, as both taxa first appear
in Late Jurassic Europe and spread to North America in the Albian, with Late
Cretaceous examples known from the western North America, central Asia and Europe.
It's also logical anatomically, as Richardoestesia? isosceles would be
expected to have some unserrated and possibly constricted teeth if it were microraptorian.
It should be noted Paronychodon has priority over Richardoestesia,
and lacustris and caperatus both have priority over isosceles.
Also, Euronychodon has priority over Asiamericana, and portuculensis
has priority over both asiatica and asiaticus. So if this synonymy
is proven, none of the names associated with straight-toothed Richardoestesia
will survive synonymization.
Ryan and Russell (2001) list "uncatalogued TMP teeth" as "Richardsoestesia n. sp. Baszio 1997b" from the Foremost Formation, which are some of the six Richardoestesia teeth reported by Peng et al. (2001) from the PHR-1 and PHR-2 localities. The latter listed "RTMP96.62.26 and -31" as "Richardoestesia
sp. Baszio, 1997" from the Foremost and Oldman Formations, so one of
those numbers represents the less than six Foremost Formation R. isosceles teeth.
The tooth LSUMG 140:6140 was first referred to Saurornitholestes cf. langstoni
by Sankey (2001) before being identified as R? isosceles
by Sankey et al. (2002). Similarly, Oldman tooth RTMP 89.103.25 was
identified as Aves indet. by Sankey et al. (2002) before being
reidentified as Richardoestesia isolsceles by Larson and Currie (2013) and Dumont et al. (2016).
References- Russell, 1935. Fauna of the Upper Milk River beds, Southern
Alberta. Transactions, Royal Society of Canada. 3(29),115-127.
Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern
Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin
of the AMNH. 147.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press, New York. 107-125.
Rowe, Cifelli, Lehman and Weil, 1992. The Campanian Terlingua Local Fauna, with
a summary of other vertebrates from the Aguja Formation, Trans-Pecos Texas.
Journal of Vertebrate Paleontology. 12(4), 472-493.
Derstler, 1994. Dinosaurs of the Lance Formation in eastern Wyoming. in Nelson
(ed.). Wyoming Geological Association. Forty-Fourth Annual Field Conference
Guidebook. 127-146.
Derstler, 1995. The Dragons’ Grave - an Edmontosaurus bonebed containing
theropod egg shells and juveniles, Lance Formation, (Uppermost Cretaceous),
Niobrara County, Wyoming: Journal of Vertebrate Paleontology. 15(3), 26A.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Sankey, 1997. Late Cretaceous vertebrate paleontology and Paleoecology, Upper
Aguja Formation, Big Bend National Park, Texas. Journal of Vertebrate Paleontology.
17(3), 73A.
Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages
(exclusive of mammals) from the Foremost and Oldman Formations of the
Judith River Group (Campanian) of southeastern Alberta: An illustrated
guide. Provincial Museum of Alberta Natural History Occasional Paper. 25, 54 pp.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend,
Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, 2002. Theropod dinosaur diversity in the latest Cretaceous (Maastrichtian)
of North America. Journal of Vertebrate Paleontology. 22(3), 103A.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous
(Campanian-Maastrichtian), Big Bend National Park, Texas. In Carpenter (ed).
The Carnivorous Dinosaurs. 127-152.
Demar and Breithaupt, 2006. The nonmammalian vertebrate microfossil assemblages
of the Mesaverde Formation (Upper Cretaceous, Campanian) of the Wind River and
Bighorn Basin, Wyoming. in Lucas and Sullivan (eds). Late Cretaceous Vertbrates
from the Western Interior. New Mexico Museum of Natural History & Science.
Bulletin 35, 33-53.
Lewis, Heckert and Forys, 2006. Paleoecology of the aqueous paleoenvironments
of the Late Cretaceous (Early Campanian) Allison Member of the Menefee Formation
in Northwestern New Mexico. NMGS Annual Spring Meeting, abstracts.
Carbot-Chanona and Rivera-Sylva, 2011. Presence of a maniraptoriform dinosaur
in the Late Cretaceous (Maastrichtian) of Chiapas, Southern Mexico. Boletin
de la Sociedad Geologica Mexicana. 63(3), 393-398.
Sankey, 2012. Something's fishy: Was one of the most abundant Latest Cretaceous
theropods a fish-eater? Journal of Vertebrate Paleontology. Program and Abstracts
2012, 165.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Dumont, Tafforeau, Bertin, Bhullar, Field, Schulp, Strilisky,
Thivichon-Prince, Viriot and Louchart, 2016. Synchrotron imaging of
dentition provides insights into the biology of Hesperornis and Ichthyornis, the "last" toothed birds. BMC Evolutionary Biology. 16:178.
R? cf. isosceles (Kirkland, Britt, Burge, Carpenter, Cifelli,
DeCourten, Eaton, Hasiotis and Lawton, 1997)
Cenomanian- Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (NCSM 33274) tooth (4.21x1.65x0.98 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
(NCSM 33288) incomplete tooth (~9x4.05x1.9 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
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.
Avrahami, 2018. Paleobiodiversity of a new microvertebrate locality
from the Upper Cretaceous Mussentuchit Member, Cedar Mountain
Formation, Utah: Testing morphometric multivariate approaches for
quantifying shape variation in microvertebrate specimens. Masters
thesis, North Carolina State University. 181 pp.
Avrahami, Gates, Heckert, Makovicky and Zanno, 2018. A new
microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain
Formation: Insights into the paleobiodiversity and paleobiogeography of
early Late Cretaceous ecosystems in western North America. PeerJ.
6:e5883.
R? cf. isosceles sp. nov. (Williamson, 2001)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-52503) tooth (1.5x1x.2 mm) (Williamson and Brusatte,
2014)
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
(NMMNH P-32753) tooth (?x3.2x1.5 mm) (Williamson and Brusatte, 2014)
Comments- Williamson and Brusatte (2014) note this differs from other
Richardoestesia isosceles in that it has a high DSDI, as do recurved
Richardoestesia teeth from the formation.
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the
Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications
for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
R? cf. isosceles sp. nov. (Lee, 1995)
Cenomanian, Late Cretaceous
Woodbine Formation, Texas, US
Material- (SMU 73778) tooth
Comments- This is straight like R. isosceles, but much shorter
and with larger serrations (4/mm).
References- Lee, 1995. Mid-Cretaceous archosaur faunal changes in Texas.
in Sun and Wang (eds.). Sixth Symposium on Mesozoic Terrestrial Ecosystems and
Biota, Short Papers. China Ocean Press, Beijing. 143-146.
Lee, 1997. The Archosauria from the Woodbine Formation (Cenomanian) in Texas.
Journal of Paleontology. 71(6), 1147-1156.
R? cf. isosceles sp. nov. (Zinke, 1998)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Material- (IPFUB Gui D 118-155) forty teeth (~3.42 mm)
Comments- These are very similar to R. isosceles, but are less
tall on average and can have more distal serrations (up to 14/mm).
Reference- Zinke, 1998. Small theropod teeth from the Upper Jurassic
coal mine of Guimarota (Portugal). Palaontologische Zeitschrift. 72(1/2), 179-189.
R? cf. isosceles sp. nov. (Rauhut and Zinke, 1995)
Barremian, Early Cretaceous
Una Formation, Spain
Material- (IPFUB Una Th 1–20, 64, 68, 81) forty-seven teeth (3-5
mm)
Comments- These are very similar to R. isosceles, but differ in
that some teeth lack serrations on both mesial and distal carinae.
References- Rauhut and Zunke, 1995. A description of the Barremian dinosaur
fauna from Una with a comparison of that of Las Hoyas. II. International Symposium
of Lithographic Limestone, Extended Abstracts. 123-126.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca,
Spain. Cretaceous Research. 23, 255-263.
R? cf. isosceles (Torices, 2002)
Late Campanian-Early Maastrichtian, Late Cretaceous
Montrebei, Tremp Formation, Spain
Material- (DPM-MON-T5) tooth (2.4x1.4x.6 mm) (Torices, 2002)
(DPM-MON-T9) tooth (2.8x1.7x.8 mm) (Torices, 2002)
Late Maastrichtian, Late Cretaceous
Blasi 2B, Tremp Formation, Spain
Material- (MPZ98/72) tooth (4.3x2.2x1.1 mm) (Torices, Currie, Canudo
and Pereda-Suberbiola, 2015)
(MPZ98/73) tooth (3.4x2.2x.9 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
(MPZ98/74) tooth (3x1.4x.6 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
(MPZ2004/7) tooth (2.1x1x.4 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Comments- These are only assigned to cf. Richardoestesia sp. by
Torices et al. (2015), but resemble R. isosceles in the convex or straight
distal edge. DPM-MON-T5 and T9 were assigned to Dromaeosauridae indet. 4 by
Torices (2002).
References- Torices, 2002. Los dinosaurios ter�podos del Cret�cico
Superior de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de
Paleontolog�a. 53, 139-146.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
R? cf. isosceles sp. nov. (Codrea, Smith, Dica, Folie, Garcia,
Godefroit and Van Itterbeecke, 2002)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Material- teeth
Comments- These are very elongate, straight to slightly recurved, with
no mesial serrations and up to 5 distal serrations per mm. They were said to
be most similar to the straight type of Richardoestesia teeth, but they
differ in all lacking mesial serrations.
Reference- Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke,
2002. Dinosaur egg nests, mammals and other vertebrates from a new Maastrichtian
site of the Hateg Basin (Romania). C. R. Palevol. V. 1, p. 173-180.
R? asiatica (Nessov, 1995)
Sues and Averianov, 2013
= Asiamericana asiatica Nessov, 1995
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Syntypes- (N 460/12457) tooth
(N 12457 coll.) two teeth
Referred- (ZIN PH 1073/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1074/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1075/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1076/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1077/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1078/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1079/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1080/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1081/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1082/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1083/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1084/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1085/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1086/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1087/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1088/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1089/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1090/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1091/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1092/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1093/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1094/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1095/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1096/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1097/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1098/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1099/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1100/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1101/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1102/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1103/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1104/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1105/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1106/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1107/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1108/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1109/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1110/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1111/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1112/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1113/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1114/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1115/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1116/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1117/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1118/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1119/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1120/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1121/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1122/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1123/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1124/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1125/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1126/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1127/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1128/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1129/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1130/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1131/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1132/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1133/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1134/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1135/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1136/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1137/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1138/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1139/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1140/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1141/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1142/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1143/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1144/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1145/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1231/16) tooth (Sues and Averianov, 2013)
(ZIN PH 2315/16) tooth (Sues and Averianov, 2013)
Diagnosis- indistinguishable from R. isosceles.
Comments- Nessov (1989) first published these teeth as ?Spinosauridae,
then (1995) named it Asiamericana asiatica and considered it to be either
a spinosaurid or a saurodontid fish. However, he noted resemblence between the
holotype and a tooth (AMNH 8113) that has since been referred to Richardoestesia
isosceles. Ryan (1997) first mentioned Richardoestesia teeth from
this formation. Sues and Averianov (2013) redescribed the teeth as a species
of Richardoestesia, finding it's indistinguishable from R. isosceles,
contra Buffetaut et al. (2008) who stated it was certainly a saurodontid. The
holotype teeth were initially described as lacking serrations, but Sues and
Averianov show this is untrue. Notably, asiatica has precedence over
isosceles, though the species are unlikely to be synonymous given the
geographical and age difference.
References- Nessov, 1989. [New findings of remains of dinosaurs, crocodiles,
and flying reptiles of the Late Mesozoic of USSSR]. [Questions of herpetology.
Abstracts of the Seventh Herpetological Conference]. 173-174.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii. Institute for Scientific Research on the Earth's
Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Ryan, 1997. Middle Asian dinosaurs. In Currie and Padian (eds.). Encyclopedia
of Dinosaurs. Academic Press. 442-444.
Buffetaut, Suteethorn, Tong and Amiot, 2008. An Early Cretaceous spinosaurid
theropod from southern China. Geological Magazine. 145(5), 745-748.
Sues and Averianov, 2013. Enigmatic teeth of small theropod dinosaurs from the
Upper Cretaceous (Cenomanian–Turonian) of Uzbekistan. Canadian Journal
of Earth Sciences. 50, 306-314.
R? cf. asiatica (Averianov, 2007)
Middle Albian-Early Cenomanian, Early-Late Cretaceous
Khodzhakul Formation, Uzbekistan
Material- (ZIN PH 1222/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1223/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1224/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1225/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1226/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1227/16) tooth (Sues and Averianov, 2013)
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits
in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research. 28, 532-544.
Sues and Averianov, 2013. Enigmatic teeth of small theropod dinosaurs from the
Upper Cretaceous (Cenomanian–Turonian) of Uzbekistan. Canadian Journal
of Earth Sciences. 50, 306-314.
R. sp. (Druckenmiller, Erickson, Brinkman and Brown, 2012)
Coniacian-Maastrichtian, Late Cretaceous
Prince Creek Formation, Alaska, US
Reference- Druckenmiller, Erickson, Brinkman and Brown, 2012. Dinosaur
diversity in the Arctic: New records of polar dinosaurs based on microvertebrate
analysis from the Upper Cretaceous Prince Creek Formation, Northern Alaska.
Journal of Vertebrate Paleontology. Program and Abstracts 2012, 88.
R. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
(RTMP 88.86.44) tooth (Ryan and Russell, 2001)
(RTMP 96.62 coll.) <6 teeth (Peng, Russell and Brinkman, 2001)
(?UC coll.) tooth (Frampton, 2006)
Comments- Ryan and Russell (2001) list RTMP 88.86.44 as a reference Richardoestesia sp. specimen from the Foremost Formation with additional teeth being referrable. The latter are some of the Richardoestesia gilmorei
teeth described by Peng et al. (2001) from the Foremost Formation, with
numbers within the range of RTMP 96.62.23-32 and/or 66. These
would represent less than six of the teeth found at sites PHR-1 and
PHR-2 (as at least one of these is R. isosceles)
Cullen and Evans (2016) list one Richardoestesia tooth from the PHRN site, which equates with Frampton's (2006) thesis.
References- Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages
(exclusive of mammals) from the Foremost and Oldman Formations of the
Judith River Group (Campanian) of southeastern Alberta: An illustrated
guide. Provincial Museum of Alberta Natural History Occasional Paper. 25, 54 pp.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press,
Bloomington, Indiana. pp. 279-297.
Frampton, 2006. Taphonomy and palaeoecology of mixed
invertebrate-vertebrate fossil assemblage in the Foremost Formation
(Cretaceous, Campanian), Milk River Valley, Alberta. Masters thesis, University
of Calgary. 294+ pp.
Cullen and Evans, 2016. Palaeoenvironmental drivers of vertebrate
community composition in the Belly River Group (Campanian) of Alberta,
Canada, with implications for dinosaur biogeography. BMC Ecology. 16,
52.
R. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
Material- (RTMP 87.78.3) tooth (Ryan, 2003)
(RTMP 89.79.62) tooth (Ryan and Russell, 2001)
teeth (Ryan and Russell, 2001)
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press,
Bloomington, Indiana. pp. 279-297.
Ryan, 2003. Taxonomy, systematics and evolution of centrosaurine ceratopsids
of the Campanian Western Interior basin of North America. Unpublished PhD thesis.
Department of Biological Sciences, Calgary, Alberta. 578pp.
R. sp. (Fanti and Miyashita, 2009)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Material- (RTMP 2004.93.3) incomplete tooth
Reference- Fanti and Miyashita, 2009. A high latitude vertebrate fossil
assemblage from the Late Cretaceous of west-central Alberta, Canada: Evidence
for dinosaur nesting and vertebrate latitudinal gradient. Palaeogeography, Palaeoclimatology,
Palaeoecology. 275, 37-53.
R. sp. indet. (Mongelli and Varricchio, 1998)
Early Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- two teeth
Reference-
Mongelli and Varricchio, 1998. Theropod teeth of the Lower Two Medicine Formation
(Campanian) of northwestern Montana. Journal of Vertebrate Paleontology. 18(3),
64A.
R. sp. indet. (Varricchio, 1995)
Mid-Late Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
Material- teeth? (Varricchio, 1995)
teeth (Redman, Moore and Varricchio, 2015)
References- Varricchio, 1995. Taphonomy of Jack’s Birthday Site,
a diverse dinosaur bonebed from the Upper Cretaceous Two Medicine Formation
of Montana. Palaeogeography, Palaecolimatology, Palaeoecology. 114, 297-323.
Redman, Moore and Varricchio, 2015. A new vertebrate microfossil locality in
the Upper Two Medicine Formation in the vicinity of Egg Mountain. Journal of
Vertebrate Paleontology. Program and Abstracts 2015, 201-202.
R. sp. indet. (Triebold, 1997)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, South Dakota, US
Material- ?(UCMP 119718) teeth (UCMP online)
?(UCMP 119733) tooth (UCMP online)
?(UCMP 119920) eight teeth (UCMP online)
?(UCMP 119921) tooth (UCMP online)
?(UCMP 120075) three teeth (UCMP online)
?(UCMP 120132) tooth (UCMP online)
?(UCMP 120153) tooth (UCMP online)
?(UCMP 120191) two teeth (UCMP online)
?(UCMP 120255) three teeth (UCMP online)
?(UCMP 120287) tooth (UCMP online)
?(UCMP 120338) tooth (UCMP online)
?(UCMP 123543) two teeth (UCMP online)
?(UCMP 123565) tooth (UCMP online)
?(UCMP 128913) tooth (UCMP online)
?(UCMP 128941) tooth (UCMP online)
(UCMP 174810) tooth (UCMP online)
teeth? (Triebold, 1997)
teeth (DePalma, 2010)
Comments- Most of the UCMP specimens (except 174810) are identified as
Chirostenotes in their collection, so are referred to Richardoestesia
here, as the R. gilmorei holotype was originally referred to Chirostenotes
(which is toothless).
References-
Triebold, 1997. The Sandy Site: Small Dinosaurs from the Hell Creek Formation
of South Dakota. in Wolberg, Stump and Rosenberg (eds). Dinofest International,
Proceedings of a Symposium sponsered by Arizona State University. A Publication
of The Academy of Natural Sciences. 245-248.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous
bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis,
University of Kansas. 227 pp.
R. sp. indet. (Stokosa, 2005)
Maastrichtian, Late Cretaceous
Fox Hills Formation, South Dakota, US
Material- (SDSM 14516) tooth
Reference-
Stokosa, 2005. Enamel microstructure variation within the Theropoda. In Carpenter
(ed.). The Carnivorous Dinosaurs. 163-178.
R. sp. indet. (UCMP online)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- ?(UCMP 120849) over fifty teeth
Comments- Most of the UCMP specimens are identified as
Chirostenotes in their collection, so are referred to Richardoestesia
here, as the R. gilmorei holotype was originally referred to Chirostenotes
(which is toothless).
R. sp. (Wroblewski, 1995)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
Material- teeth
References- Wroblewski, 1995. First report of changes in Lower Vertebrate
Faunas across the Cretaceous-Tertiary boundary, Western Hanna Basin, Wyoming.
Journal of Vertebrate Paleontology. 15(3), 61A.
Wroblewski, 1998. Changing paleoenvironments and paleofaunas across the K-T
boundary, Ferris Formation, Southcentral Wyoming. Tate Geological Museum, Casper
College, Casper Wyoming. Tate ’98. Life in the Cretaceous. 53-70.
R. sp. indet. (UCMP online)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- ?(UCMP 119674) tooth
Comments- This is identified as
Chirostenotes in their collection, so is referred to Richardoestesia
here, as the R. gilmorei holotype was originally referred to Chirostenotes
(which is toothless).
R. sp. (Kirkland, Lucas and Estep, 1998)
Barremian, Early Cretaceous
Yellow Cat Member of Cedar Mountain Formation, Utah, US
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. (Kirkland, Lucas and Estep, 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah
Comments- This is listed as cf. Richardoestesia sp. by Kirkland
et al. (1998).
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the
Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous
Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin.
14, 79-89.
R. sp. (Parrish, 1999)
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Material- (UCM 8656) tooth
Comments- This was listed as cf. Richardoestesia by Parrish (1999).
Reference- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-.
Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah.
Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.
R. sp. nov. (Williamson, 2001)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-27488) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27489) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27495) tooth (?x~3.9x1.7 mm) (Williamson and Brusatte, 2014)
(NMMNH P-27496) tooth (?x3.7x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-28379) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30004) tooth (4.1x3.3x1.4 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30005) tooth (?x3.4x1.3 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30277) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30331) tooth (?x3.4x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-36552) tooth (Williamson and Brusatte, 2014)
(NMMNH P-38410) tooth (?x1.5x.7 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38428) tooth (Williamson and Brusatte, 2014)
(NMMNH P-52501) tooth (?x?x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-52514) tooth (?x.9x? mm) (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
(NMMNH P-30207) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30208) tooth (5x2.9x1.4 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30209) tooth (?x2.2x.8 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30219) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30221) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30224) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33478) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33479) tooth (?x1.9x1.1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33480) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33481) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33482) tooth (4.5x2.9x1.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33483) tooth (Williamson and Brusatte, 2014)
Comments- Williamson and Brusatte (2014) note these differ from other
Richardoestesia in that they have a high DSDI.
References- Williamson, 2001. Dinosaurs from microvertebrate sites in
the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New
Mexico. 2001 GSA abstracts.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
R. sp. (Wel and Williamson, 2000; described by Williamson and
Brusatte, 2014)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (NMMNH P-22567) tooth (?x2.8x1.4 mm) (Williamson and Brusatte,
2014)
(NMMNH P-32742) tooth (?x2x1.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-37793) tooth (?x2.3x1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-46389) tooth (?x2.8x1.2 mm) (Williamson and Brusatte, 2014)
Reference- Weil and Williamson, 2000. Diverse Maastrichtian terrestrial
vertebrate fauna of the Naashoibito Member, Kirtland Formation (San Juan Basin,
New Mexico) confirms “Lancian” faunal heterogeneity in western North
America. Geological Society of America Abstracts with Programs. 32, A-498.
Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous
Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
R. sp. indet. (YPM online)
Late Cretaceous
US
Material- (YPM 56979)
R? sp. (Prieto-Marquez, Gaete, Galobart and Ardevol, 2000)
Late Campanian, Late Cretaceous
L'Abeeler, Aren Sandstone Formation, Spain
Material- (IPS 18 372) tooth (14.03x5.85x3.11 mm)
Comments- Prieto-Marquez et al. (2000) concluded "the minute size and the similar morphology of the denticles of R. gilmorei
place the Spanish theropod closer to this species than to any other
known taxon." Torices et al. (2015) stated "in the discriminant
analysis performed, this tooth is clearly isolated by its height from
both the North American and Pyrenean samples, which suggests it might
be a new taxon."
References- Prieto-Marquez, Gaete, Galobart and Ardevol, 2000. A Richardoestesia-like
theropod tooth from the Late Cretaceous foredeep, south-central Pyrenees, Spain.
Eclogae Geologicae Helvetiae. 93(3), 497-501.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod
dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta
Palaeontologica Polonica. 60(3), 611-626.
R. spp. (Torices, Currie, Canudo and Pereda-Suberbiola, 2015)
Late Campanian, Late Cretaceous
La�o, Sedano Formation, Spain
Material- (MCNA 14566-14621) sixty-four teeth (1.2-6.6 mm) [details
in Torices et al., 2015]
(UPVLP 135) tooth (Isasmendi et al., 2020)
Comments- These teeth seem to fall under both gilmorei and isosceles
morphotypes.
References- Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod
dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta
Palaeontologica Polonica. 60(3), 611-626.
Isasmendi, Torices, Canudo and Pereda-Suberbiola, 2020.
Paleobiodiversity of theropod dinosaurs from the Upper Cretaceous La�o
site, northern Iberian peninsula. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 186-187.
R? sp. (Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- teeth
Comments- Ortega et al. (2009) state Richardoestesia teeth are
known, calling it a basal tetanurine, and Torices et al. (2011) mention cf.
Richardoestesia.
References- Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009. The
vertebrate diversity of the Upper Campanian-Lower Maastrichtian "Lo Hueco"
fossil-site (Cuenca, Spain). Journal of Vertebrate Paleontology. 29(3), 159A-160A.
Torices, Barroso-Barcenilla, Cambra-Moo, Perez and Serrano, 2011. Vertebrate
microfossil analysis in the palaeontological site of 'Lo Hueco' (Upper Cretaceous,
Cuenca, Spain). Journal of Vertebrate Paleontology. Program and Abstracts 2011,
205.
R. sp. (Valentin, Godefroit, Tabuce, Vianey-Liaud, Wenhao and
Garcia, 2012)
Late Maastrichtian, Late Cretaceous
Vitrolles-La-Plaine, Provence, France
Material- (UP-VLP-08-001) tooth
(UP coll.) several teeth
Reference- Valentin, Godefroit, Tabuce, Vianey-Liaud, Wenhao and Garcia,
2012. First Late Maastrichtian (Latest Cretaceous) vertebrate assemblage from
Provence (Vitrolles-la-Plaine, Southern France). In Godefroit (ed.). Bernissart
Dinosaurs and Early Cretaceous Terrestrial Ecosystems. Indiana University Press.
582-597.
R? sp. (Debeljak, Kosir and Otonicar, 1999)
Campanian-Maastrichtian, Late Cretaceous
Kozina, Slovenia
Material- (ACKK-D-8/081) tooth
Comments- This tooth was identified as Dromaeosauridae based on the high
DSDI, and furthermore seems to be Richardoestesia because of the small
serrations (~8 per mm) and elongate shape.
References- Debeljak, Kosir and Otonicar, 1999. A preliminary note on
dinosaurs and non-dinosaurian reptiles from the Upper Cretaceous carbonate platform
succession at Kozina (SW Slovenia). Razprave IV. razreda SAZU. 40, 3-25.
Debeljak, Kosir, Buffetaut and Otonicar, 2002. The Late Cretaceous dinosaurs
and crocodiles of Kozina (SW Slovenia): A preliminary study. Memorie della Societa
Geologica Italiana. 57, 193-201.
R. sp. indet. (Nessov, 1995)
Santonian-Early Campanian, Late Cretaceous
Syuk-Syuk Formation, Kazakhstan
Material- jaw, teeth
Comments- These teeth are serrated, unlike R. asiatica.
Reference- Nessov, 1995. Dinozavri severnoi Yevrasii: Novye dannye o
sostave kompleksov, ekologii i paleobiogeografii [Dinosaurs of Northern Eurasia:
new data about assemblages, ecology and paleobiogeography], Scientific Research
Institute of the Earth's Crust, St. Petersburg State University, St. Petersburg,
Russia: 156 pp. + 14 pl. [in Russian with short English, German, and French
abstracts].
R. spp. (Alifanov and Bolotsky, 2002)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- teeth
Comments- Alifanov and Bolotsky (2002) referred teeth to two morphs of
Richardoestesia, possibly gilmorei and isosceles.
Reference- Alifanov and Bolotsky, 2002. New data about the assemblages
of the Upper Cretaceous carnivorous dinosaurs (Theropoda) from the Amur region.
In Kirillova (ed.). Fourth International Symposium of IGCP 434. Cretaceous continental
margin of East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
R? sp. (Knoll and Ruiz-Omenaca, 2005)
Beriassian, Early Cretaceous
KM 1983, Ksar Metlili Formation, Morocco
Material- (MNHN SA 2004/5B; Velociraptorinae
indet. Morphotype II; lost) incomplete tooth (?x?x1.40 mm)
Comments- This was collected in 1983, 1986 or 1999, and identified by Knoll and Ruiz-Omenaca (2005) as Velociraptorinae
indet. Morphotype II. They noted resemblences to European Richardoestesia teeth and the serration density (7.52-8.33 per mm) matches that genus. Lasseron (2020) noted the "dinosaur remains (Knoll, 2000; Knoll & Ruiz-Ome�aca, 2009) ... were taken out of the MNHN and lost" (translated).
References-
Knoll and Ruiz-Omenaca, 2009. Theropod teeth from the basalmost Cretaceous of
Anoual (Morocco) and their palaeobiogeographical significance. Geological Magazine.
146(4), 602-616.
Lasseron, 2020. Paleobiodiversite, evolution et paleobiogeographie des
vertebres mesozoiques africans et gondwaniens : apport des gisements du
Maroc oriental. Doctoral thesis, Museum National D'Histoire Naturelle.
493 pp.
R? sp. (Prasad, Parmar and Kumar, 2014)
Early Jurassic
Kota Formation, India
Material- teeth
Comments- These were described as "Richardoestesia-like."
Reference- Prasad, Parmar and Kumar, 2014. Recent vertebrate fossil discoveries
from the Jurassic Kota Formation of India. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 208.
R. sp. (Bertini and Franco-Rosas, 2001)
Turonian-Late Maastrichtian, Late Cretaceous
Adamantina, Marilia and/or Serra da Galga Formations, Bauru Group, Brazil
Material- teeth
Reference- Bertini and Franco-Rosas, 2001. Scanning electron microscope
analysis on Maniraptoriformes teeth from the Upper Cretaceous of southeastern
Brazil. JVP 21(3) 33A.
undescribed possible Microraptoria (Kirkland, Britt, Burge, Carpenter,
Cifelli, DeCourten, Eaton, Hasiotis and Lawton, 1997)
Early Cenomanian, Early Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71599) two teeth (Fiorillo, 1999)
(OMNH coll.; Morphotype 7 of Frederickson et al., 2018) teeth (<5 mm)
Comments- These are identified as cf. Troodon sp.
in Kirkland et al. (1997), and Troodontidae gen. et sp. indet. by
Cifelli et al. (1999). They are probably "the earliest troodontids"
mentioned by Parrish and Eaton (1991) and troodontids of Kirkland and
Parrish (1995) from the upper Cedar Mountain Formation. Frederickson
et al. (2018) lists these as "?troodontid" but also say they "have
relatively smaller denticles than those of most Late Cretaceous
troodontids ... ; though more associated material is needed to
unequivocally dismiss this identification." They also noted Fiorillo
(1999) identified this morphotype as Velociraptorinae. Given the lack
of mesial serrations, small size, Middle Cretaceous timing and
longitudinal grooves on some specimens, these are here suggested to be
microraptorians.
References- Parrish and Eaton, 1991. Diversity and evolution of dinosaurs
in the Cretaceous of the Kaipirowits plateau, Utah. Journal of Vertebrate Paleontology.
11(3), 50A.
Kirkland and Parrish, 1995. Theropod teeth from the Lower Cretaceous of Utah.
Journal of Vertebrate Paleontology. 15(3), 39A.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis
and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the
central Colorado plateau: A key to understanding 35 million years of
tectonics, sedimentology, evolution, and biogeography. Brigham Young
University Geology Studies. 42, 69-103.
Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial
Cretaceous vertebrates from the Cedar Mountain Formation, Emery County,
Utah: The Mussentuchit local fauna. In Gillette (ed.). Vertebrate
Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison
eggshell site, Cedar Mountain Formation (Lower Cretaceous), Emery
County, Utah. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah
Geological Survey, Miscellaneous Publication. 99-1, 259-268.
Goldberg, 2000. Faunal composition, non-marine vertebrates, of the
upper Cedar Mountain Formation (Cretaceous: Albian-Cenomanian), central
Utah. PhD thesis, University of Oklahoma. 243 pp.
Frederickson, Engel and Cifelli, 2018. Niche partitioning in pheropod
dinosaurs: Diet and habitat preference in predators from the uppermost
Cedar Mountain Formation (Utah, U.S.A.). Scientific Reports. 8:17872.
unnamed microraptorian (Averianov, Starkov and Skutschas, 2003)
Late Barremian-Mid Aptian, Early Cretaceous
Mogoito Member of Murtoi Formation, Russia
Material- (ZIN PH 10/13) anterior lateral tooth (FABL 2.65 mm)
(ZIN PH 11/13) posterior lateral tooth (FABL 4.1 mm), posterior lateral tooth
(FABL 5.45 mm)
Comments- These teeth were referred to Dromaeosauridae by Averianov et
al. (2003), but are here specified as microraptorians based on their small serrations
(9-10/mm mesially, 6-9/mm distally). The anterior tooth is elongate and only
slightly curved, while the posterior teeth are short and strongly recurved.
The BW/FABL of the anterior tooth is .47, and the posterior teeth .52-.56. The
mesial carinae follow the midline and are serrated only basally on the anterior
tooth. Posterior serrations are rectangular, broad labiolingually, and have
short shallow interdenticle slits. They resemble teeth of Sinornithosaurus
and Richardoestesia.
Reference- Averianov, Starkov and Skutschas, 2003. Dinosaurs from the
Early Cretaceous Murtoi Formation in Buryatia, Eastern Russia. Journal of Vertebrate
Paleontology. 23(3):586–594.
undescribed possible microraptorian (Kurochkin, 1988)
Aptian-Albian, Early Cretaceous
Khoboor, Mongolia
Material- teeth
Comments- These have unserrated mesial carinae, and distal carinae with
very small serrations. There is a slight basal constriction, and the crown is
strongly recurved. They were referred to birds by Kurochkin (1998), but to troodontids
by Nessov and Golovneva (1990) and Averianov and Sues (2007). However, they
also sound similar to Shanag and perhaps Richardoestesia.
References- Kurochkin, 1988. [Cretaceous Mongolian birds and their significance
for the study of bird phylogeny.] Trudy Sovmestnoi Sovetsko-Mongol’skoi
Paleontologicheskoi Ekspeditsii. 34, 33–42. [Russian]
Nessov and Golovneva, 1990. [History of the flora, vertebrates and climate in
the late Senonian of the north-eastern Koriak Uplands]. in Krasilov (ed). [Continental
Cretaceous of the USSR.] Dal’nevostochnoe Otdelenie AN SSSR, Vladivostok.
[Russian]. 191-212.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
unnamed possible microraptorian (Pittman, Pei and Xu, 2015)
Aptian-Albian, Early Cretaceous
Bayan Gobi Formation, Inner Mongolia, China
Material- (IVPP V22530) partial dorsal rib, manual ungual, manual claw
sheath, distal femur, tibia (~75 mm), fibula, metatarsal I, incomplete metatarsal
II, fragmentary phalanx II-1, partial phalanx II-2, pedal ungual II, incomplete
metatarsal III, incomplete phalanx III-1, incomplete metatarsal IV, phalanx
IV-1, phalanx IV-2, two pedal phalanges, partial metatarsal V
Comments- The rib and manual ungual may not belong to the same individual
as the hindlimb. Pittman et al. noted the distally appressed metatarsals are
similar to non-eudromaeosaurs, but also that the more constricted pedal phalanx
II-2 is similar to some eudromaeosaurs.
Reference- Pittman, Pei and Xu, 2015. The first dromaeosaurid (Dinosauria:
Theropoda) from the Early Cretaceous Bayan Gobi Formation of Nei Mongol, China.
PeerJ PrePrints. https://dx.doi.org/10.7287/peerj.preprints.1340v1
undescribed microraptorian (Lamanna, Li, Harris, Atterholt and You,
2010)
Early Aptian, Early Cretaceous
Xiagou Formation, Gansu, China
Material- (GSGM coll?) pectoral girdle, humerus, radius, ulna
Comments- Lamanna et al. (2010) referred this to Microraptoria based
on the supracoracoid fenestra.
References- Lamanna, Li, Harris, Atterholt and You, 2010. First non-avian
dinosaur from the Lower Cretaceous (Aptian) Xiagou Formation of the Changma
Basin, northwestern China. Journal of Vertebrate Paleontology. Program and Abstracts
2010, 119A.
Zhou, Lamanna, Poust, Li, You and O'Connor, in review. First non-avian
theropod (Dromaeosauridae, Microraptorinae) from the bird-bearing Lower
Cretaceous Xiagou Formation of the Changma Basin, Gansu Province,
northwestern China. Journal of Vertebrate Paleontology.
undescribed Microraptoria
Jehol Group?, Early Cretaceous
Liaoning?, China
Material- (private coll.) fragmentary skull, fragmentary mandibles, cervical
vertebrae, dorsal vertebrae, dorsal ribs, sacrum, caudal vertebrae, chevrons,
humeri, radius, ulna, metacarpal I, phalanx I-1, metacarpal II, metacarpal III,
ilium, pubes, ischia, femora, tibiotarsi, metatarsi, pedal digit II, pedal digit
III, pedal digit IV, metatarsal V (http://www.thenaturalcanvas.com/Dinosaurs/pages/4523.html)
(private coll.) skull, cervical series, dorsal series, dorsal ribs, sacrum,
caudal series, chevrons, scapulocoracoid, furcula, humeri, radii, ulnae, carpus,
metacarpal I, phalanx I-1, metacarpal II, phalanx II-1, phalanx II-2, manual
ungual II, metacarpal III, phalanx III-1, phalanx III-3, manual ungual III,
pubis, ischia, femora, tibiotarsi, proximal fibula, metatarsus, pedal digit
II, pedal digit III, pedal ungual III, pedal digit IV, pedal ungal IV
(IVPP, D and Tianyu coll.) (over 300 individuals) (Alexander et al., 2010)
(Wenya Museum DNO: 087) (640 mm) incomplete skull, dorsal ribs, caudal series,
incomplete humeri, partial radius, incomplete ulnae, metacarpal I, phalanx I-1,
manual ungual I, partial manus, pelvic elements, femora, tibiotarsi, metatarsi,
pedal ungual II, pedal digit III, pedal ungual III, pedal digit IV, pedal ungual
IV, contaur feathers
(Wenya Museum DNO: 090) composite including supposed humerus, radius, ulna,
partial manus, femora, tibiae
microraptorian- partial mandible, cervical series, dorsal vertebrae, dorsal
ribs, uncinate processes, sacrum, caudal series, radii, ulnae, semilunate carpal,
metacarpal I, phalanx I-1, metacarpal II, phalanx II-1, phalanx II-2, metacarpal
III
confuciusornithid- humerus, pubes, metatarsi, pedal digit I, pedal ungual I,
pedal digit II, pedal ungual II, pedal digit III, pedal ungual III, pedal digit
IV, pedal ungual IV
Comments- There are many undescribed microraptorian specimens photographed
online, most for sale. These are usually advertised as Microraptor, but
may also belong to Sinornithosaurus. They are all probably from the Yixian
or perhaps Juifotang Formations of Liaoning. Many are altered or composites.
Alexander et al. (2010) reported seeing over three hundred microraptorian specimens
at the IVPP, Dalian Natural History Museum and Shandong Tianyu Museum of Natural
History.
A specimen on sale at The Natural Canvas is labeled as Microraptor gui.
It has a disarticulated skull and its precise affinities cannot be determined
without better photographs. Another specimen is preserved spread in dorsal view
and seems basically complete besides one distal pes. Its affinities cannot be
determined yet either.
Three dromaeosaurid specimens were exhibited at the China Jehol Biota Fossil
Exhibition in 2002 (Creisler, DML 2002) from the personal collection of Du Wenya,
who runs a museum in Jinzhou. These were available at least in 2000 (Hutchinson,
DML 2000). DNO: 087 is fairly complete but very poorly preserved. The specimen
is obviously altered, as the tibiotarsi and femora are switched (giving the
illusion of long femora), while one pes is far too short. The pelvic elements
are mixed up as well, with possible proximal pubes located anteriorly, and two
other elements located posteriorly.
DNO: 090 is a composite specimen, as shown by the obviously confuciusornithid
pes (Hutchinson, DML 2000). One pes is attached to a broken tibia(?), while
the other is placed at the end of what may be another bird tarsmetatarsus. Both
are far too short compared to the femora, one of which is much longer and apparently
constructed of two limb bones. The pubis appears too slender and pointed for
a dromaeosaurid, resembling those of avialans more. One humerus appears
to have a proximal foramen like Confuciusornis and Sapeornis.
One of the lower arms and manus appears genuinely microraptorian, though the
other lower arm is much too short with a far too robust "radius".
The other possible manus is too poorly preserved to evaluate. The dentary and
tail both appear to be dromaeosaurid.
References- Hutchinson, DML 2000. https://web.archive.org/web/20180115070226/http://dml.cmnh.org/2000Jun/msg00268.html
Creisler, DML 2002. https://web.archive.org/web/20180115070231/http://dml.cmnh.org/2002Dec/msg00303.html
Alexander, Gong, Martin, Burnham and Falk, 2010. Model tests of gliding with
different hindwing configurations in the four-winged dromaeosaurid Microraptor
gui. Proceedings of the National Academy of Sciences. 107(7), 2972-2976.
Graciliraptor Xu and Wang,
2004
= "Graciliraptor" Xu, 2002
G. lujiatunensis Xu and Wang, 2004
= "Graciliraptor lujiatunensis" Xu, 2002
= Sinornithosaurus lujiatunensis (Xu and Wang, 2004) Paul, 2010
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V13474) (~1 m; 3.3 kg; adult) fragmentary maxilla, eight maxillary
teeth, mid caudal vertebra (12.5 mm), nine distal caudal vertebrae (30 mm),
chevrons, incomplete humerus (~106 mm), radii (~87 mm; one distal), ulnae (91
mm; one distal), scapholunare, semilunate carpals, distal carpal III, metacarpals I (17, 17
mm), phalanges I-1 (35, 35 mm), manual unguals I (32.5, 33.5 mm), metacarpals
II (53.5, 54 mm), phalanges II-1 (25, ~24.5 mm), phalanx II-2 (26.5 mm), proximal
manual ungual II, metacarpals III (48.8, 49.4 mm), phalanges III-1 (~13.5, ~12
mm), phalanges III-2 (6.4, 6.2 mm), phalanx III-3 (17.5 mm), manual ungual III,
incomplete tibiotarsi (~172 mm), distal fibula (~167 mm), distal metatarsals
II, phalanx II-1 (12 mm), phalanges II-2 (13, 12 mm), proximal pedal ungual
II, distal metatarsal III, phalanx III-1 (21.8, 22 mm), phalanges III-2 (14,
13 mm), distal metatarsals IV, phalanges IV-1 (17, ~17 mm), phalanx IV-2 (10.8
mm)
Diagnosis- (after Xu and Wang, 2004) astragalar medial condyle significantly
expanded posteriorly; metatarsal II 1.5 times wider than metatarsal III distally.
Other diagnoses- Turner et al. (2012) note several characters listed
in the original diagnosis are problematic. The elongate caudals, slender tibiotarsus
and pedal phalanx III-1 are shared with Microraptor, the postzygapophyseal
laminae on distal caudals and rectangular proximal section of the tibia are
common in dromaeosaurids, the proximal end of metacarpal III is not very dorsoventrally
expanded, and manual ungual I is not necessarily small compared to ungual II
due to the latter being crushed.
Comments- Discovered in 2001, this was initially described in Xu's (2002)
thesis, then published in 2004. Turner et al. (2012) consider it a possible
synonym of Microraptor zhaoianus, though it falls outside the Sinornithosaurus+Microraptor
clade in Foth et al. (2014) and Lee et al. (2014).
References- Xu, 2002. Deinonychosaurian fossils from the Jehol Group
of Western Liaoning and the coelurosaurian evolution. PhD Thesis. Chinese Academy
of Sciences. 325 pp.
Xu and Wang, 2004. A new dromaeosaur (Dinosauria: Theropoda) from the Early
Cretaceous Yixian Formation of Western Liaoning. Vertebrata PalAsiatica. 42(2),
111-119.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Changyuraptor Han, Chiappe,
Ji, Habib, Turner, Chinsamy, Liu and Han, 2014
C. yangi Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han,
2014
= Sinornithosaurus yangi (Han, Chiappe,
Ji, Habib, Turner, Chinsamy, Liu and Han, 2014) Paul, 2016
Barremian-Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Holotype- (HG B016) (~1.32 m; 7.62 kg; five year old adult) fragmentary skull, fragmentary
mandibles, cervical vertebrae, dorsal vertebrae, dorsal ribs, uncinate processes,
gastralia, twenty caudal vertebrae, chevrons, scapula, fragmentary coracoid,
partial furcula, sternum, sternal ribs, humeri (one distal; 148.9 mm), radii,
ulnae (126.1 mm), semilunate carpal, metacarpals I (~21.3 mm), phalanges I-1
(~44.6 mm), manual ungual I, metacarpal II (~75.6 mm), phalanx II-1, phalanges
II-2, manual unguals II, metacarpal III (73.2 mm), phalanx III-1 (~20.7 mm),
phalanx III-2 (~11.5 mm), phalanx III-3 (~20.4 mm), manual unguals III, manual
claw sheaths, pubes (117.1 mm), partial ischium, femora (153 mm), tibiae, fibula,
metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II (32.5 mm),
metatarsals III (111.2 mm), pedal phalanges III-1 (21.8 mm), pedal phalanges
III-2 (15.3 mm), pedal phalanges III-3 (13.9 mm), pedal unguals III (25.7 mm),
metatarsals IV (107.4 mm), metatarsals IV, pedal phalanges IV-1, pedal phalanges
IV-2, pedal phalanges IV-3, pedal phalanges IV-4, pedal unguals IV (23.1 mm),
pedal claw sheaths, metatarsals V, scales, contour feathers, fragmentary remiges,
leg remiges, retrices
Diagnosis- (after Han et al., 2014) furcula more robust than Sinornithosaurus
and much larger than Tianyuraptor; forelimb much longer when compared
to hindlimb; humerus much longer (>20%) than ulna; metacarpal I shorter than
Sinornithosaurus (1/4-1/5 versus 1/3); semilunate carpal covering all
of proximal metacarpals I and II; manual ungual II largest; ischium shorter
than Microraptor; midshaft of metatarsal IV significantly broader than
metatarsal II or III; mid caudals roughly twice length of dorsals unlike Microraptor;
fewer (22) caudal vertebrae than Microraptor and Tianyuraptor;
rectrices significantly longer than other microraptorines.
References- Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014.
A new raptorial dinosaur with exceptionally long feathering provides insights
into dromaeosaurid flight performance. Nature Communications. 5, 4382.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press.
360 pp.
Sinornithosaurus Xu, Wang and Wu, 1999
= "Sinavisaurus" Skrepnick, DML 2000
S. millenii Xu, Wang and
Wu, 1999
= Sinornithosaurus haoiana Liu, Ji, Tang and Gao, 2004
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou Member of the Yixian Formation, Liaoning, China
Holotype-
(IVPP V12811) (~1.05 m, 5.00 kg) skull (152 mm), scleral plates,
mandibles (135 mm), hyoid (53 mm), atlantal intercentrum, atlantal
neurapophyses, anterior cervical vertebra, three mid cervical
vertebrae, three posterior cervical vertebrae, tenth cervical vertebra,
two cervical ribs, eleven dorsal vertebrae (13 mm), presacral vertebra,
fourteen dorsal ribs, gastralia, sacrum (65 mm), first caudal vertebra
(9.5 mm), second caudal vertebra, third caudal vertebra, fourth caudal
vertebra, fifth caudal vertebra (15 mm), sixth caudal vertebra (20 mm),
seventh caudal vertebra (28 mm), eighth caudal vertebra, ninth caudal
vertebra, chevrons, scapulae (85, ~85 mm), coracoids (42 mm), furcula,
sternal plates (84 mm), four sternal ribs, humeri (133, 134 mm), radius
(~106 mm), ulna (110 mm), semilunate carpal, distal carpal III,
metacarpal I (20 mm), phalanx I-1 (41 mm), manual ungual I (27 mm),
metacarpal II (63 mm), phalanx II-1 (32 mm), phalanx II-2 (35 mm),
manual ungual II (26 mm), metacarpal III (60 mm), phalanx III-1 (18
mm), phalanx III-2 (9 mm), phalanx III-3 (23 mm), manual ungual III (19
mm), ilia (85 mm), pubes (116 mm), ischia (52 mm), incomplete femora
(~146 mm), distal tibia, partial fibula, astragalocalcaneum, distal
tarsal III, distal tarsal IV, metatarsal I (21 mm), phalanx I-1 (11
mm), pedal ungual I (11 mm), metatarsal II (88 mm), phalanx II-1 (15
mm), phalanx II-2 (17 mm), partial pedal ungual II (35 mm), metatarsals
III (93, 93 mm), phalanx III-1 (25 mm), phalanx III-2 (17.5 mm),
phalanx III-3 (16.5 mm), pedal ungual III (20 mm), metatarsal IV (91
mm), phalanx IV-1 (20 mm), phalanx IV-2 (14 mm), phalanx IV-3 (10 mm),
phalanx IV-4 (12 mm), pedal ungual IV (18.5 mm), metatarsal V (47 mm),
feathers, keratin ungual sheaths (m2, p4)
Referred- (D2140; holotype of Sinornithosaurus haoiana)
(2 year old subadult) (skull ~125 mm) premaxilla, maxilla, nasals,
lacrimal, frontals, parietal, partial jugal, postorbital, squamosal,
quadratojugal, quadrate, vomer, palatine, pterygoid,
parasphenoid-basisphenoid, sclerotic plates, dentaries, splenial,
surangular, angulars, hyoid, teeth, eight or nine cervical vertebrae,
three cervical ribs, eleven dorsal vertebrae (13 mm), dorsal ribs, five
uncinate processes (to 29 mm), gastralia, sacral vertebrae, five
proximal caudal vertebrae, several mid caudal vertebrae and chevrons,
scapulacoracoid (scapula 84.4 mm), furcula, sternal plates (64.1 mm),
four sternal ribs (to 42.6 mm), humeri (129.5 mm), radii, ulnae (105.2
mm), scapholunare, semilunate carpal, metacarpal I, phalanx I-1, manual
ungual I, metacarpal II (59 mm), metacarpal III (54.1 mm), phalanx
III-1, phalanx III-2, ilia (84.5, 86.1 mm), pubes (116.9 mm), ischium,
incomplete femur, incomplete tibia, incomplete fibula,
astragalocalcaneum, metatarsal II, phalanx II-1, phalanx II-2, pedal
ungual II (45.2 mm on curve), metatarsal III (86.8 mm), phalanx III-1
(23.3 mm), phalanx III-2 (14.5 mm), phalanx III-3 (12.8 mm), pedal
ungual III (~21 mm), metatarsal IV (84.2 mm), pedal digit IV, pedal
ungual IV, metatarsal V, feathers (Liu et al., 2004)
(IG-3) skull, postcranial skeleton (Azuma, 2005)
(IG-5b) skull postcranial skeleton, feathers (Azuma, 2005)
(IVPP V16904) (adult) specimen including skull, mandibles, hyoid, cervical vertebrae,
dorsal vertebra, dorsal ribs, scapulae, coracoids, manual phalanx, manual ungual
and tibiotarsus (Turner et al., 2012)
(JMP-V-05-8-01; inside holotype of Sinocalliopteryx gigas) tibia (155
mm), partial fibula, phalanx I-1, pedal ungual I, phalanx II-1, phalanx II-2,
pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, metatarsal IV,
phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, feathers (Ji et al.,
2007)
(Tianyu Museum of Natural History coll.) multiple specimens including cranial
material (Gong et al., 2010)
?(Wenya Museum DNO: 088) skull, dorsal vertebrae, sacrum, caudal series, furcula,
humeri, radius, ulnae, manual phalanges, manual unguals, pubes, femora, tibiotarsi,
metatarsi, pedal digit III, pedal digit IV (Kaiser, 2007)
?(private coll.) skull, mandibles, hyoid, cervical vertebrae, dorsal vertebrae,
gastralia, caudal series, scapulocoracoid, partial furcula, sternum, sternal
ribs, humeri, radii, ulnae, semilunate carpal, metacarpal I, phalanx I-1, manual
ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal
III, phalanx III-3, manual ungual III, ilium, pubes, femora, tibiotarsi, metatarsal
II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual
III, metatarsal IV, pedal ungual IV, pedal phalanges, metatarsal V
(private coll.) incomplete skull, mandible, hyoid, cervical series, dorsal vertebrae,
dorsal ribs, gastralia, sacrum, caudal series, scapulocoracoid, sternal plate,
humeri, radius, ulna, metacarpal I, phalanx I-1, manual ungual I, metacarpal
II, phalanx II-1, phalanx II-2, manual ungual II, ilia, pubes, ischium, femora,
tibiotarsi, metatarsi, pedal digit II, pedal digit III, pedal digit IV, unidentified
elements
? skull, mandible, cervical vertebrae, dorsal series, dorsal ribs, gastralia,
caudal series, coracoids, furcula, humeri, radii, ulnae, manual elements, ilium,
pubis, ischium, femora, tibiae, metatarsi, pedal digit II, pedal ungual II,
phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-1,
phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
specimen including skull, mandible (Zheng, 2009)
Diagnosis- (from Xu et al., 1999) elongate posterolateral parietal process
(unknown in other derived microraptorines).
(after Xu and Wu, 2001) groove posterior to the anterior carina on the lingual
surface of the premaxillary tooth crowns (unknown in other derived microraptorines);
column-like margin of the pterygoid process of the quadrate (unknown in other
derived microraptorines); large excavation on the posterolateral surface of
the cultriform process (unknown in other derived microraptorines).
(after Turner et al., 2012) thickened posterior rim of promaxillary fenestra
(unknown in other derived microraptorines).
(proposed) larger humerus than Microraptor or Bambiraptor (humerofemoral
ratio >88%) (also in Wulong).
Other diagnoses- Xu et al. (1999) listed additional synapomorphies of
Sinornithosaurus. However, the pitted antorbital fossa, elongate posterodorsal
dentary process, enlarged supracoracoid fenestra, lateral tubercle on the pubic
midshaft, and proximodorsal ischial process are also present in Microraptor
(and the pubic character in Hesperonychus); unserrated premaxillary teeth
and a subarctometatarsus are near certainly plesiomorphic for dromaeosaurids
and found in other microraptorians; the manual phalanx III-1/III-2 ratio of
S. millenii falls between those of Microraptor specimens.
Contra Xu and Wu (2001), the large promaxillary fenestra is now known to be
present in Microraptor as well. The deep excavation on the posteroventral
margin of the premaxilla is also present in Bambiraptor. Turner et al.
(2012) note the supposed diastema between premaxillary and maxillary teeth is
actually the ventral edge of the subnarial process.
Comments- Sinornithosaurus' holotype was discovered in 1998. It
was named and briefly described by Xu et al. (1999), and described in depth
in Xu's (2002) thesis, though only the cranial (Xu and Wu, 2001) and pedal (Wu
and Wang, 2000) portions have been published. Skrepnick (DML, 2000) noted that
Sinornithosaurus had the working name of "Sinavisaurus" until
right before it was published. The species name must be emended to millennii
because it's a mispelling of a Latin word (millennium) (ICZN Art. 32.5).
IVPP V16904 was first published as IVPP uncatalogued by Turner et al. (2012),
but Xu et al. (2015) reveal its specimen number. Xu et al. do not believe it
to be conspecific with S. millenii because none of its teeth are serrated,
and its neurocentral sutures are fully closed and a tibiotarsus is developed
despite it being "significantly smaller" than the holotype which lacks
these fusions. While possibly correct pending further description, it should
be noted the referred specimen of Mei long has a tibiotarsus and fused
presacral neurocentral sutures but is 80% the size of the holotype that doesn't,
the Sinovenator changii paratype has a tibiotarsus but is 89% the size
of the holotype that doesn't, and Microraptor zhaoianus specimens can
have serrations on both carinae (NGMC 00-12-A), only distal serrations (holotype)
or no serrations (IVPP V13320). Thus such variations are known in other basal
paravian species, so are expected in S. millenii individuals.
Ji et al. (2002) referred NGMC 91 to Sinornithosaurus sp., and it has
been referred to S. millenii by other authors such as Senter (2011) and
Turner et al. (2012), but it is here referred to a new genus more closely related
to Microraptor. Xu (2002) referred IVPP V13477 to Sinornithosaurus
sp., but Xu et al. (2003) later made it a paratype of Microraptor gui.
Ji (2002) described NGMC 00-12-A as Sinornithosaurus sp., but it is here
referred to Microraptor.
Gong et al. (2010) argued Sinornithosaurus was venomous, but Gianechini
et al. (2011) show the paper is highly flawed and its conclusions lack merit.
Several undescribed specimens are photographed online, and appear to be Sinornithosaurus.
They are probably all from the Yixian or perhaps Juifotang Formations of Liaoning,
China.
One is complete and largely articulated except for the pelvis and hindlimbs.
It is being sold by Geosciences Enterprises as Microraptor zhaoianus.
The nasal is disarticulated, making the skull appear more slender and troodontid-like,
but the proportions all match Sinornithosaurus. The elongate manual phalanx
III-3 distinguishes it from Microraptor, while the short ulna and long
manual digit II distinguish it from Graciliraptor.
Another is quite disarticulated, but well preserved. The elongate tibiotarsus
suggests it is Sinornithosaurus.
A further specimen is almost complete and well preserved, except both manus
are fragmentaty and one distal tibia is missing. The pes is positioned directly
contacting the broken end, suggesting some alteration. The skull is almost complete
and looks particularily elongate.
Three dromaeosaurid specimens were exhibited at the China Jehol Biota Fossil
Exhibition in 2002 (Creisler, DML 2002) from the personal collection of Du Wenya,
who runs a museum in Jinzhou. These were available at least in 2000 (Hutchinson,
DML 2000). DNO: 088 is possibly the specimen illustrated by Kaiser (2007) as
an unidentified confuciusornithid. It is obviously dromaeosaurid however, with
an elongate tail, slender metatarsus, and slender furcula. The elongate tibia
suggests it may be Sinornithosaurus.
Gong et al. (2010) mention specimens from the Tianyu Museum, and later (2011)
illustrate a skull from Zheng (2009).
Sinornithosaurus haoiana- Liu et al. (2004) described D2140 as
a new species of Sinornithosaurus, S. haoiana. This was based
on several proposed differences from S. millenii, but Turner et al. have
proposed the two species are synonymous. The long subnarial process of the premaxilla
which excludes the maxilla from the naris cannot be evaluated in S. millenii.
They further note the maxillary fenestra of the millenii holotype is
heavily damaged, so the small and circular morphology of haoiana cannot
be compared. Turner et al. find the dorsal quadratojugal process of millenii
to be longer than thought by Xu, which can indeed be seen in photos. They state
both holotypes have low premaxillae, and yet do not argue against Liu et al.'s
correct observation millenii's are much lower. Similarly, they argue
all specimens have low dentaries, but do not argue against Liu et al.'s correct
observation millenii's are much lower. Turner et al. are wrong when they
claim millenii has a pubic peduncle narrower than its acetabulum. I would
use these three differences to retain them as separate species if not for IVPP
V16904 having both a tall premaxilla and a low dentary (fig. 23C in Turner et
al.), which ironically is not used in Turner et al.'s argument. Based on this
specimen, I agree the species are synonymous. The species name must be emended
to haoianus,
to match the gender of the genus name (ICZN Art. 31.2, 34.2). Han et
al. (2014) provided more measurements. Poust et al. (2020
examined the histology, finding it to be more mature than the Wulong holotype.
References- Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous
integument from the Yixian Formation of China. Nature. 401, 262-266.
Hutchinson, DML 2000. https://web.archive.org/web/20180115070226/http://dml.cmnh.org/2000Jun/msg00268.html
Skrepnick, DML 2000. https://web.archive.org/web/20210605025348/http://dml.cmnh.org/2000Oct/msg00074.html
Xu and Wang, 2000. Troodontid-like pes in the dromaeosaurid Sinornithosaurus.
Paleontological Society of Korea Special Publication. 4, 179-188.
Xu and Wu, 2001. Cranial morphology of Sinornithosaurus millenii Xu et
al. 1999 (Dinosauria: Theropoda: Dromaeosauridae) from the Yixian Formation
of Liaoning, China. Canadian Journal of Earth Sciences. 38, 1739–1752.
Xu, Zhou and Prum, 2001. Branched integumental structures in Sinornithosaurus
and the origin of feathers. Nature. 410, 200-204.
Creisler, DML 2002. https://web.archive.org/web/20180115070231/http://dml.cmnh.org/2002Dec/msg00303.html
Ji, 2002. New data of Early Cretaceous dromaeosaurs from western Liaoning with
comments on the origin of feathers. Unpublished Thesis. 94 pp.
Ji, Ji, Yuan and Ji, 2002. Restudy on a small dromaeosaurid dinosaur with feathers
over its entire body. Earth Science Frontiers. 9(3), 57-63.
Xu, 2002. Deinonychosaurian fossils from the Jehol Group of Western Liaoning
and the coelurosaurian evolution. PhD Thesis. Chinese Academy of Sciences. 325
pp.
Xu, Zhou, Wang, Kuang, Zhang and Du, 2003. Four-winged dinosaurs from China.
Nature. 421, 335-340.
Liu, Ji, Tang and Gao, 2004. A new species of dromaeosaurids from the Yixian
Formation of western Liaoning. Geological Bulletin of China. 23(8), 778-783.
Azuma, 2005. The Flying Dinosaurs: Fukui Prefectural Dinosaur Museum. 118 pp.
Ji, Ji, Lu and Yuan, 2007. A new giant compsognathid dinosaur with long filamentous
integuments from Lower Cretaceous of Northeastern China. Acta Geologica Sinica.
81(1), 8-15.
Kaiser, 2007. The Inner Bird: Anatomy and Evolution. UBC Press. 386 pp.
White, 2009. The subarctometatarsus: intermediate metatarsus architecture demonstrating
the evolution of the arctometatarsus and advanced agility in theropod dinosaurs.
Alcheringa. 33(1), 1-21.
Zheng, 2009. Niaolei Qiyuan. Jinan City, China: Shandong Science and Technology
Publishing House Press.
Gong, Martin, Burnham and Falk, 2010. The birdlike raptor Sinornithosaurus
was venomous. Proceedings of the National Academy of Sciences. 107, 766-768.
Zhang, Kearns, Orr, Benton, Zhou, Johnson, Xu and Wang, 2010. Fossilized melanosomes
and the colour of Cretaceous dinosaurs and birds. Nature. 463, 1075-1078.
Gianechini, Agnolin and Ezcurra, 2011. A reassessment of the purported venom
delivery system of the bird-like raptor Sinornithosaurus. Pal�ontologische
Zeitschrift. 85, 103-107.
Gong, Martin, Burnham and Falk, 2011. Evidence for a venomous Sinornithosaurus.
Pal�ontologische Zeitschrift. 85, 109-111.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological
continuity within Dinosauria. Journal of Evolutionary Biology. 24, 2197-2216.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Xing, Bell, Persons, Ji, Miyashita, Burns, Ji and Currie, 2012. Abdominal contents
from two large Early Cretaceous compsognathids (Dinosauria: Theropoda) demonstrate
feeding on confuciusornithids and dromaeosaurids. PLoS ONE. 7(8), e44012.
Han, Chiappe, Ji, Habib, Turner, Chinsamy, Liu and Han, 2014. A new raptorial
dinosaur with exceptionally long feathering provides insights into dromaeosaurid
flight performance. Nature Communications. 5, 4382.
Xu, Pittman, Sullivan, Choiniere, Tan, Clark, Norell and Wang, 2015. The taxonomic
status of the Late Cretaceous dromaeosaurid Linheraptor exquisitus and
its implications for dromaeosaurid systematics. Vertebrata PalAsiatica. 53(1),
29-62.
Poust, Gao, Varricchio, Wu and Zhang, 2020. A new microraptorine
theropod from the Jehol biota and growth in early dromaeosaurids. The
Anatomical Record. Early View DOI: 10.1002/ar.24343
unnamed microraptorian (Ji, Norell, Gao, Ji and Ren, 2001)
Late Valanginian-Early Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Material-
?(NGMC 91; Dave) (69 cm) skull (~100 mm), sclerotic rings, mandibles,
ten cervical vertebrae, cervical ribs, dorsal series, dorsal ribs,
uncinate processes, gastralia, caudal series, chevrons, scapulae (~53
mm), coracoids, furcula, sternal plate, sternal ribs, humeri (87 mm),
ulnae (72 mm), radii, scapholunare, semilunate carpal, metacarpal I (13
mm), phalanx I-1 (27 mm), manual ungual I (15 mm), metacarpal II (40
mm), phalanx II-1 (22 mm), phalanx II-2 (24 mm), manual ungual II (16
mm), metacarpal III (40 mm), phalanx III-1 (12 mm), phalanx III-2 (7
mm), phalanx III-3 (16 mm), manual ungual III (11 mm), pubis(?),
ischium(?), femora (95 mm), tibiae (133 mm), fibulae, astragalus,
distal phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1 (10
mm), phalanx II-2 (10.2mm), pedal ungual II, metatarsal III (62 mm),
phalanx III-1 (16.2 mm), phalanx III-2 (9.6 mm), phalanx III-3 (10.3
mm), pedal ungual III, metatarsal IV, phalanx IV-1 (7.1 mm), phalanx
IV-2 (6 mm), phalanx IV-3 (6.9 mm), phalanx IV-4, pedal ungual IV,
metatarsal V, scales, contour feathers, remiges, retrices, ungual
sheaths
Comments- NGMC 91 was first reported by Ji et al. (2001) as Dromaeosauridae
indet., though they noted strong resemblence to Sinornithosaurus. Ji
et al. noted the possibility that some differences, such as the bowed first
metacarpal, were due to age. Ji et al. (2002) later assigned NGMC 91 to Sinornithosaurus
sp., with the same reservations regarding whether differences from the S.
millenii holotype were taxonomic or ontogenetic. Czerkas et al. (2002) tentatively
referred NGMC 91 to their new genus Cryptovolans (= Microraptor),
though without stated reasons besides "nearly identical wing proportions".
In fact, NGMC 91 lacks the proposed apomorphies of Cryptovolans (28-30
caudal vertebrae; manual phalanx III-1 longer than III-3), and several apomorphies
of Microraptor- posterior dentary teeth with constricted bases; manual
phalanx III-1 >83% of phalanx III-3 in length; manual phalanx I-1 <62%
of metacarpal II in length; tibiofemoral ratio <136%; highly elongate leg
remiges (>1.5 times femoral length). Senter et al. (2004) found that NGMC
91 claded with Microraptor based on- mesial serrations absent on posterior
teeth; dentary with midlength increase in height. Senter has since (2007) stated
the dentary character's "apparent presence was an optical illusion caused
by differing heights of preserved dentary teeth and the curvature of the dentary."
He still found the Microraptor+NGMC 91 clade however, supported by a
proximodorsal lip on manual ungual I and strongly curved pedal unguals III and
IV. Senter (2011) has since combined NGMC 91 into his Sinornithosaurus
OTU without justification, and Turner et al. (2012) also placed the specimen
in Sinornithosaurus millenii. This was because they claimed it differed
from Microraptor in lacking mid caudal vertebrae 3-4 times length of
anterior dorsal vertebrae, but this is untrue. Further, the supposed Sinornithosaurus
apomorphy of a posteriorly bifurcated dentary is also present in Microraptor.
It is retained as separate here.
References- Ji, Norell, Gao, Ji and Ren, 2001. The distribution of integumentary
structures in a feathered dinosaur. Nature. 410 (6832), 1084-1088.
Czerkas, Zhang, Li and Li, 2002. Flying dromaeosaurs. in Czerkas (ed.). Feathered
Dinosaurs and the Origin of Flight. 97-126.
Ji, Ji, Yuan and Ji, 2002. Restudy on a small dromaeosaurid dinosaur with feathers
over its entire body. Earth Science Frontiers, 9(3), 57-63.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae.
Bulletin of Gunma Museum of Natural History. 8, 1-20.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 5(4), 429-463.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological
continuity within Dinosauria. Journal of Evolutionary Biology. 24, 2197-2216.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Microraptorinae sensu Martyniuk, 2012
Definition- (Microraptor zhaoianus <- Sinornithosaurus millenii,
Dromaeosaurus albertensis)
Wulong Poust, Gao, Varricchio, Wu and Zhang, 2020
= "Wulong" Poust, 2014
W. bohaiensis Poust, Gao, Varricchio, Wu and Zhang, 2020
= "Wulong bohaiensis" Poust, 2014
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype-
(D2933) (1 year old juvenile) skull (98.6 mm), sclerotic plates,
mandibles, ten cervical vertebrae, thirteen dorsal vertebrae,
~ten-thirteen dorsal ribs, gastralia, three or four sacral centra,
sacral neural arches and ribs, thirty or thirty-one caudal vertebrae
(distal caudals ~19 mm), chevrons, scapulae (44.8 mm), coracoids,
furcula, sternals, four sternal ribs, humeri (75.4, 76.6 mm), radii
(61.4 mm), ulnae (63.4, 64.5 mm), semilunate carpals, distal carpal
III, metacarpals I (12.3, 12.3 mm), phalanges I-1 (~25 mm), manual
unguals I (15.3, 14.8 mm), metacarpals II (39.1, 41.3 mm), phalanges
II-1 (20, 19 mm), phalanges II-2 (22.7, 22.7 mm), manual unguals II
(15.1, 15.3 mm), metacarpals III (~39, ~36 mm), phalanges III-1 (12.8,
13.1 mm), phalanges III-2 (8.3, 7.5 mm), phalanges III-3 (14.2 mm),
manual unguals III (10.1, 9.9 mm), manual claw sheaths, ilium (49.5
mm), pubes (62.4, 68.4 mm), ischia (26.9, 27.5 mm), femora (85.8, 86.5
mm), tibiae (117 mm), fibulae (118.7 mm), astragali, calcanea,
metatarsal I, phalanx I-1 (6.5 mm), pedal ungual I (5.2 mm),
metatarsals II (53.5, 54 mm), phalanges II-1 (one partial; 9 mm),
phalanges II-2 (one partial; 9.2 mm), pedal unguals II (16.4, 16.1 mm),
metatarsals III (56.5, 56.9 mm), phalanges III-1 (14.3 mm), phalanges
III-2 (10 mm), phalanges III-3 (9.3 mm), pedal unguals III (11.3 mm),
metatarsals IV (56.7, 53.5 mm), phalanges IV-1 (10.7, 11.1 mm),
phalanges IV-2 (7.6, 8.5 mm), phalanges IV-3 (5.9, 6.5 mm), phalanges
IV-4 (6.1, 6.7 mm), pedal unguals IV (10.1 mm), metatarsals V, pedal
claw sheaths, body feathers, retrices, remiges, leg remiges
Diagnosis-
(after Poust et al., 2020) long anterior process of quadratojugal;
anteriorly inclined pleurocoels on anterior half of dorsal centra;
transverse processes of proximal caudals significantly longer than
width of centrum; 30 caudal vertebrae; distally located and large
distodorsal ischial process; large supracoracoid fenestra (>15% of
total area).
Comments-
This specimen was transferred from a private collector to the Dalian
Natural History Museum prior to April 2014 (based on SVP abstract
deadlines). Poust (2014) first described and
named Wulong in his thesis,
with the disclaimer "This thesis is not
intended to constitute publication of new names or nomenclatural acts
and such names as are given are not here intended for public and
permanent scientific record – this is a model of what will be published
later." This means the thesis falls under ICZN Article 8.2 "A
work that contains a statement to the effect that it is not issued for
public and permanent scientific record, or for purposes of zoological
nomenclature, is not published within the meaning of the Code", and the
name was a nomen nudum until Poust et al. described it officially in
2020.
Poust et al.'s measurements for manual phalanx I-1 (15.9 and 15.8 mm)
and metacarpal III (17.1, 18.7 mm) are far too short, and others listed
above may end up being off as well once checked. Poust et al. are
incorrect in stating "the tiny first phalanges of left and right manual
digit three are missing", as shown by their own description, figures
and measurement table. Contra Poust et al., only the right ilium
seems
to be exposed, with the supposed broken dorsal boundary exposing the
dorsal left ilium being a crack through the bone. The authors
note
"restoration efforts by technically proficient but inexpert preparators
untrained in anatomy resulted in one noticeable error prior to the
specimen’s arrival at the museum: the distal pedal phalanges of the
right Digit II (2, 3, and the ungual) have been reconstructed
incorrectly."
Poust et al. (2020) entered it into Turner's TWiG analysis and recovered it as a microraptorian sister to Sinornithosaurus,
but both characters supporting this are misscored- "the lack of a
proximodorsal lip on the manual unguals (153) and a laterally inclined
flange along the dorsal edge of the surangular (212)." Both taxa have
a proximodorsal lip on at least manual ungual II, while Wulong
is too crushed to determine the presence of a surangular flange.
Adding it to Hartman et al.'s TWiG analysis results in a most
parsimonious position sister to Microraptor plus Zhongjianosaurus, placing all of the Jiufotang microraptorians together.
References-
Poust, 2014. Description and ontogenetic assessment of a new Jehol
microraptorine. Masters thesis, Montana State University. 105 pp.
Poust, Varricchio and Gao, 2014. A new Jehol microraptorine
with implications for secondary feather loss in Sinornithosaurus. Journal
of Vertebrate Paleontology. Program and Abstracts 2014, 208.
Poust, Gao, Varricchio, Wu and Zhang, 2020. A new microraptorine
theropod from the Jehol biota and growth in early dromaeosaurids. The
Anatomical Record. Early View DOI: 10.1002/ar.24343
Zhongjianosaurus Xu and Qin, 2017
Z. yangi Xu and Qin, 2017
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Holotype- (IVPP V22775) (adult,
310 g) seventh cervical vertebra, eighth cervical vertebra, ninth
cervical vertebra (4.7 mm), tenth cervical vertebra (4.4 mm), first
dorsal vertebra (4.4 mm), second dorsal vertebra (4.4 mm), third dorsal
vertebra (4.7 mm), fourth dorsal vertebra (4.7 mm), fifth dorsal
vertebra (4.8 mm), sixth dorsal vertebra (5.0 mm), seventh dorsal
vertebra (5.0 mm), thirteen dorsal ribs (d1 ~20, d4 39, d8 25 mm),
three free uncinate processes, second caudal vertebra (4.6 mm), third
caudal vertebra (5.1 mm), fourth caudal vertebra (5.5 mm), fifth caudal
vertebra (6.3 mm), sixth caudal vertebra (9.3 mm), seventh caudal
vertebra (13.3 mm), eighth caudal vertebra (14.9 mm), ninth caudal
vertebra (~16.5 mm), tenth caudal vertebra, eleventh caudal vertebra,
twelfth caudal vertebra, thirteenth caudal vertebra (13.2 mm),
fourteenth caudal vertebra (13.1 mm), fifteenth caudal vertebra (13.1
mm), sixteenth caudal vertebra (13.1 mm), seventeenth caudal vertebra
(13.2 mm), eighteenth caudal vertebra (13.1 mm), nineteenth caudal
vertebra (12.7 mm), twentieth caudal vertebra (12.7 mm), twenty-first
caudal vertebra (11.4 mm), twenty-second caudal vertebra (10.8 mm),
twenty-third caudal vertebra (9.1 mm), twenty-fourth caudal vertebra
(8.7 mm), twenty-fifth caudal vertebra (7.9 mm), twenty-sixth caudal
vertebra (6.8 mm), twenty-seventh caudal vertebra (6.3 mm), chevrons,
furcula, sternal plate (33 mm), five sternal ribs (r1 6.5, r5 25.5 mm),
scapulocoracoids (scapula 33 mm), humeri (43 mm), radius (42 mm), ulna
(44 mm), scapholunare, pisiform, carpometacarpus (mc1 8.5, mcII 27 mm),
phalanx I-1 (16.3 mm), manual ungual I (12.5 mm), manual claw sheath,
femora (59 mm), tibiotarsi (78 mm), fibula, tarsometatarsi (mt II 35.5,
mt III 39, mt IV 37.6 mm), distal phalanx III-1, phalanx III-2 (5.5
mm), phalanx III-3 (5.6 mm), pedal ungual III (~8 mm), distal phalanx
IV-2, phalanx IV-3 (3.1 mm), phalanx IV-4 (3.2 mm), pedal ungual IV
(7.4 mm), pedal ungual sheaths
Diagnosis- (after Xu and Qin,
2017) long ossified uncinate processes fused to dorsal ribs; widely
arched furcula with slender and posteriorly curved rami; humeral
proximal end strongly offset medially from humeral shaft; humeral
internal tuberosity short; large fenestra in humeral deltopectoral
crest; humeral ulnar condyle hypertrophied; ulna slightly longer than
humerus; ulnar olecranon process with posterior margin mediolaterally
pinched; ulnar distal end bending anteriorly and strongly expanded
laterally; proximal end of metacarpal I with strong ventrolateral
extension; manual ungual I without proximodorsal lip and lacking strong
dorsal arching; metacarpal II laterally bowed with longitudinal ventral
groove; femoral head stout and lower than trochanteric crest; medial
condyle of tibiotarsus distal end with prominent distal extension;
arctometatarsalian pes; metatarsal II without ginglymus on distal end.
Comments- While various
websites including Wikipedia state the specimen was first reported in
2009, this is not substantiated by any sourced account and is unlikely
as specimens from the Sihedang locality were not described until 2012,
with no non-bird taxa reported until 2014. Xu and Qin (2017)
indicate is that it must have been found prior to March 2017. An
advanced online version was published April 10, but the official
paginated version was published by April 26 so no separate nomen nudum
entry is given here for the two week long gap. While Xu and Qin
assigned Zhongjianosaurus
to the Yixian Formation, Sihedang is here viewed as belonging
to the Jiufotang Formation (see Iteravis entry).
Xu and Qin (2017) assigned Zhongjianosaurus
to Microraptoria (their Microraptorinae) without a quantitative
analysis, which was confirmed by Hartman et al. (2019) who recovered it
sister to Microraptor.
References- Xu and Qin, 2017. A new tiny dromaeosaurid dinosaur from the
Lower Cretaceous Jehol Group of western Liaoning and niche
differentiation among the Jehol dromaeosaurids. Vertebrata PalAsiatica. 55(2), 129-144.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Microraptor Xu, Zhou and Wang, 2000
= "Archaeoraptor" Sloan, 1999 in part
= Cryptovolans Czerkas, Zhang, Li and Li, 2002
= "Tetrapterornis" Miller, 2004
M. zhaoianus Xu, Zhou and Wang,
2000
= "Archaeoraptor liaoningensis" Sloan, 1999 in part
= Cryptovolans pauli Czerkas, Zhang, Li and Li, 2002
= Microraptor gui Xu, Zhou, Wang, Kuang, Zhang and Du, 2003
= "Tetrapterornis" gui (Xu, Zhou, Wang, Kuang, Zhang and Du, 2003) Miller, 2004
= Sinornithosaurus "zhaoi" Zhang, 2007
= Sinornithosaurus zhaoianus (Xu, Zhou and Wang, 2000) Paul, 2010
= Microraptor hanqingi Gong, Martin, Burnham, Falk and Hou, 2012
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype-
(IVPP V 12330; IVPP V 12444 in part; specimen of "Archaeoraptor liaoningensis" in part) (~420
mm; 200 g; adult) partial skull (45 mm), mandibles (42 mm), four dorsal
vertebrae, thirteenth dorsal central fragment, several dorsal ribs,
sternal ribs or uncinate processes, gastralia, sacrum (19 mm), (caudal
series 232 mm) about twenty-five caudal vertebrae, chevrons, radius (35
mm), ulna (35 mm), scapholunare, pisiform, semilunate carpal, distal
carpal III, proximal metacarpal I, proximal metacarpal II, proximal
metacarpal III, partial ilia, incomplete pubes, ischia (17.9 mm),
femora (one fragmentary) (53 mm), tibiae (one partial) (68 mm), fibula,
tarsus, phalanx I-1 (4.4 mm), pedal ungual I, metatarsal II (32.4 mm),
phalanx II-1 (5.4 mm), phalanx II-2 (5.2 mm), pedal ungual II,
metatarsal III (33 mm), phalanx III-1 (6.6 mm), phalanx III-2, phalanx
III-3 (5 mm), pedal ungual III, metatarsal IV (33.2 mm), phalanx IV-1,
phalanx IV-2 (4.8 mm), phalanx IV-3 (3.1 mm), phalanx IV-4 (3.8 mm),
pedal ungual IV, metatarsal V, pedal ungual sheaths, contour feathers,
fragmentary remiges, therian ?cranial elements, ?axial elements, ?limb
elements and pes
Referred- (BMNHC PH881) (540 mm) skull (~48.4 mm), mandibles (42.5, 44.2
mm), hyoids (~18.7 mm), ten cervical vertebrae, ~13-14 dorsal vertebrae, ten
dorsal ribs, three uncinate processes, gastralia, sacrum, ~25-26 caudal vertebrae,
chevrons, incomplete scapula, sternum, sternal ribs, humeri (42.5, 47.4 mm),
radii (one incomplete; 41 mm), ulnae (one incomplete; 44.7 mm), metacarpal I
(13 mm), phalanx I-1 (12.8 mm), manual unguals I (one fragmentary; 11.9 mm),
metacarpal II (31.8 mm), phalanx II-1 (13.4 mm), phalanges II-2 (12.5 mm), manual
unguals II (one partial; 12 mm), metacarpal III (~29.8 mm), phalanx III-1 (5.3
mm), phalanx III-2 (7.6 mm), phalanges III-3 (7.2 mm), manual unguals III (5,
5.3 mm), manual claw sheaths, incomplete ilia (~27.8 mm), pubes (46.2 mm), femora
(51.8, 52.1 mm), tibiotarsi (70.7, 72.9 mm), fibulae, distal tarsal, metatarsal
I, phalanges I-1 (4.6 mm), pedal unguals I (6 mm), metatarsals II (36.2 mm),
metatarsals III (39, 38.5 mm), phalanges III-1 (10.4, 8.9 mm), phalanges III-2
(6.4, 5.4 mm), phalanges III-3 (6, ~5.2 mm), pedal unguals III (10.6, 10.5 mm),
metatarsals IV (36.1, 36.1 mm), phalanges IV-1 (7.8, 8.2 mm), phalanges IV-2
(4.7, 4.8 mm), phalanges IV-3 I(4, 3.8 mm), phalanges IV-4 (4.8, 4.4 mm), pedal
unguals IV (8.3, 8.6 mm), pedal claw sheaths, metatarsals V, body feathers,
retrices, remiges, leg remiges (Li et al., 2012; described by Pei et al., 2014)
(CAGS02-IG-gausa-1/DM 609 in part) presacral vertebrae, furcula, humerus (70
mm), radius, ulna (63 mm), carpus, metacarpal I, phalanx I-1 (24 mm), manual
ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal
III, manual digit III, tibiae (106 mm), fibula, phalanx I-1, pedal ungual I,
metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (58
mm), metatarsal IV, pedal phalanges, other elements (Czerkas and Ji, 2002)
(IG-1) skull, postcranial skeleton (Azuma, 2005)
(IVPP V13320; paratype of Microraptor gui) (~485 mm) skull, cervical
vertebrae, dorsal vertebrae, dorsal ribs, sacrum, caudal vertebrae, chevrons,
humeri, radii, ulnae, semilunate carpal, metacarpal I, phalanx I-1, manual ungual
I, metacarpals II, phalanges II-1, metacarpals III, phalanx III-1, phalanx III-2,
ilia, pubis, femora (61 mm), tibiae, fibulae, metatarsi, both pes, body feathers,
remiges (150 mm), leg remiges, retrices (Xu, 2002)
(IVPP V13351) (~645 mm) specimen including femur (81 mm), retrices and leg remiges
(Xu, Zhou, Wang, Kuang, Zhang and Du, 2003)
(IVPP V13352; holotype of Microraptor gui) (770 mm, 1.40 kg) posterior skull,
posterior mandible, eighth cervical vertebrae, cervical ribs, thirteen dorsal
vertebrae, dorsal ribs, six uncinate processes, gastralia, sacrum, about twenty-six
caudal vertebrae, chevrons, scapulocoracoids, furcula, sternum (47.8 mm), two
sternal ribs, humeri (77 mm), radii (74.9 mm), ulnae (77.6 mm), scapholunare, pisiform,
semilunate carpal, distal carpals III, metacarpal I (13.6 mm), phalanx I-1 (28.7 mm), manual ungual
I, metacarpal II (48.6 mm), phalanx II-1 (24.3 mm), phalanx II-2 (25.1 mm),
manual ungual II, metacarpal III (47.6 mm), phalanx III-1 (12.9 mm), phalanx
III-2 (8.2 mm), phalanx III-3 (14.3 mm), manual ungual sheaths, pubis (74.5
mm), ischia (35.5 mm), femora (97.5 mm), tibiotarsi (126 mm), metatarsal II,
phalanx II-1, phalanx II-2 (10.1 mm), pedal ungual II, pedal ungual II sheath,
metatarsal III (65 mm), phalanx III-1 (12.8 mm), phalanx III-2 (10.3 mm), phalanx
III-3, metatarsal IV, phalanx IV-1 (10.4 mm), phalanx IV-2 (7 mm), phalanx IV-3
(7.3 mm), phalanx IV-4 (7.2 mm), proximal pedal ungual IV, metatarsal V, contour
feathers, retrices, remiges, leg remiges (Xu, Zhou, Wang, Kuang, Zhang and Du,
2003)
(IVPP V13475) (393 mm; adult) skull (~51 mm), scleral plates, mandibles (41.3
mm), ten cervical vertebrae, two posterior cervical ribs, thirteen dorsal vertebrae,
dorsal ribs, uncinate processes, gastralia, sacrum, incomplete caudal series,
sternal ribs, scapula (34 mm), coracoids, furcula (23.2 mm wide), humeri (42,
~42 mm), radius (34.5 mm), ulnae (37, 36 mm), metacarpals I (7, 7 mm), phalanx
I-1 (14.6 mm), manual ungual I (10.8 mm), metacarpals II (~22.3, 22 mm), phalanges
II-1 (12, 11.3 mm), phalanges II-2 (11.2, 11.8 mm), manual unguals II (11.2,
~11 mm), metacarpals III (~21, 21 mm), phalanx III-1 (9.8 mm), phalanx III-3
(10.2 mm), manual ungual III (7.8 mm), ilium, pubes (40 mm), ischium (18 mm),
femora (49.7, 49.8 mm), tibiotarsi (66, 66 mm), fibula, distal tarsal III, distal
tarsal IV, phalanx I-1 (4.6, 4.7 mm), pedal unguals I (4, 4 mm), metatarsals
II (33.2, 33 mm), phalanx II-1 (5.9 mm), phalanx II-2 (5.1 mm), pedal ungual
II (11.4 mm), metatarsal III (35.3 mm), phalanges III-1 (8.9, 8.8 mm), phalanges
III-2 (5.5, 5.8 mm), phalanges III-3 (5, 5 mm), pedal unguals III (8.7, 9.2
mm), metatarsal IV (35 mm), phalanges IV-1 (6.1, 6.1 mm), phalanges IV-2 (4.1,
4.1 mm), phalanx IV-3 (3.1 mm), phalanx IV-4 (3.1 mm), pedal ungual IV, body
feathers, remiges, leg remiges (Xu, 2002)
(IVPP V13476) (~745 mm) specimen including femur (~94 mm) and leg remiges (Xu,
Zhou, Wang, Kuang, Zhang and Du, 2003)
(IVPP V13477) (~572 mm) skull, cervical vertebrae, dorsal vertebrae, about twenty-six
caudal vertebrae, chevrons, ilium, pubis, ischium, femur (72 mm), tibiotarsus,
metatarsus, phalanx II-1, phalanx II-2, pedal ungual II, pedal digit III, pedal
ungual III, pedal digit IV, pedal ungual IV, metatarsal V, body feathers, leg
remiges, retrices (Xu, 2002)
(IVPP V17972) skull (89.9 mm), mandibles, hyoids, several cervical vertebrae,
dorsal series, dorsal ribs, uncinate processes, gastralia, partial sacrum, partial
caudal series, chevrons, partial scapula, partial coracoid, furcula, sternal
plates, humeri (89.4 mm), radii (75.9 mm), ulnae (79.7 mm), semilunate carpal,
metacarpal I (14.4 mm), phalanx I-1, manual ungual I, metacarpal II (46.6 mm),
phalanx II-1, phalanx II-2, manual ungual II, metacarpal III (~43 mm), phalanx
III-1, phalanx III-3, manual ungual III, manual claw sheaths, ilium, pubis (~82.6
mm), ischium, femora (~82.3 mm), tibiotarsi (140.4 mm), metatarsals II, phalanx
II-1, phalanx II-2, pedal unguals II, metatarsals III (~73.8 mm), phalanges
III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV,
phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals
IV, body feathers, enantiornithine humerus, radius, ulna and pes (O'Connor,
Zhou and Xu, 2011)
(JI-5) skull postcranial skeleton, feathers (Azuma, 2005)
(JLUM Y-MR160501) skull (57.64 mm), sclerotic plates, mandibles (42.9, 30.91 mm) (Maranga et al., 2020)
(LPM 0159; paratype of Cryptovolans pauli) (~706 mm) dorsal vertebrae,
dorsal ribs, uncinate processes, gastralia, sacrum, scapulocoracoid, incomplete
furcula, partial sternum, sternal ribs, humeri (74 mm), radii (66 mm), ulnae,
carpus, metacarpal I (11.9 mm), phalanx I-1 (30 mm), manual ungual I, metacarpal
II (38 mm), phalanx II-1 (16.8 mm), phalanx II-2 (17.6 mm), manual ungual II,
metacarpal III, phalanx III-1 (14.2 mm), phalanx III-2 (3.4 mm), phalanx III-3
(10 mm), manual ungual III, manual ungual sheaths, pubis, ischia, femora (89
mm), tibiae (105 mm), metatarsi (~53 mm), phalanx II-2, pedal ungual II, phalanx
III-1, phalanx III-2 (7.9 mm), phalanx III-3 (8 mm), pedal ungual III, phalanx
IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, contour feathers,
remiges, leg remiges (Czerkas, Zhang, Li and Li, 2002)
(LPM 0200/0201; = BPM 1 3-13; holotype of Cryptovolans pauli) (~950 mm)
fragmentary skull, teeth, dorsal vertebrae, dorsal ribs, uncinate processes,
twenty-eight to thirty caudal vertebrae, chevrons, furcula, sternum (60 mm),
sternal ribs, humeri, radii, ulnae, carpus, manual digit I, metacarpal II, phalanx
II-1, phalanx II-2 (26 mm), manual ungual II, metacarpal III, phalanx III-1,
phalanx III-2, phalanx III-3 (14.2 mm), manual ungual III, manual ungual sheaths,
ilium, pubis, ischium, femora (~100 mm), tibiae (132 mm), metatarsus (~66 mm),
phalanx II-2, pedal ungual II, phalanx III-2, phalanx III-3, pedal ungual III,
phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal
V, pedal ungual sheaths, remiges, retrices, leg remiges (Czerkas, Zhang, Li
and Li, 2002)
(LVH 0026; holotype of Microraptor hanqingi) (~950 mm) skull, mandibles
(92.5 mm), eight cervical vertebrae, cervical ribs, thirteen dorsal vertebrae
(~132 mm), dorsal rib fragments, gastralia, sacrum (43.6 mm), twenty-three caudal
vertebrae (~535 mm), chevrons, furcula, scapulocoracoids (scapulae 59.9, 54.9
mm), sternal plate (47.3 mm), humeri (92.6, 93.5 mm), radii (79.5, ~77.9 mm),
ulnae (82.5, ~81.9 mm), scapholunare, pisiform, semilunate carpal, distal carpal III,
metacarpal I (12.98 mm), phalanx I-1 (32.44 mm), manual ungual I (21.98 mm),
metacarpal II (54.88 mm), phalanges II-1 (21.96, 23.21 mm), phalanges II-2 (24.2,
24.72 mm), manual unguals II (20.56 mm), metacarpal III (48.17 mm), phalanges
III-1 (17.7 mm), phalanges III-2 (4.66 mm), phalanges III-3 (13.1 mm), manual
unguals III (9.35 mm), manual claw sheaths, ilium (60 mm), pubes (78 mm), ischia
(36 mm), femora (109.8, 111.7 mm), tibiotarsi (137.9, 137.1 mm), metatarsal
I, pedal ungual I, metatarsals II, phalanges II-1 (one proximal), phalanges
II-1 (one distal), pedal unguals II, metatarsals III (77.7, 75.4 mm), phalanges
III-1, phalanges III-2 (one incomplete), phalanges III-3 (one incomplete), pedal
unguals III (one incomplete), distal tarsals III+IV fused to metatarsals IV,
phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals
IV (one incomplete, one proximal), pedal claw sheaths, retrices, leg remiges
(Alexander et al., 2010)
(NGMC 00-12-A) (~832 mm) incomplete skull, sclerotic plates, incomplete mandible,
hyoids, eight cervical vertebrae, second dorsal vertebra (9 mm), third dorsal
vertebra (8.1 mm), fourth dorsal vertebra (9.4 mm), fifth dorsal neural arch
fragment, sixth dorsal vertebra (8.9 mm), seventh dorsal vertebra (9.3 mm),
eighth dorsal vertebra (9.6 mm), ninth dorsal vertebra (9.6 mm), tenth dorsal
vertebra (9.6 mm), eleventh dorsal vertebra (9.6 mm), twelfth dorsal vertebra
(8.7 mm), partial thirteenth dorsal vertebra, dorsal ribs, uncinate processes,
gastralia, first caudal vertebra, third caudal vertebra (11 mm), fourth caudal
vertebra (12 mm), fifth caudal vertebra (13.8 mm), sixth caudal vertebra (~18.9
mm), eighth caudal vertebra (26.8 mm), ninth caudal vertebra (27.1 mm), tenth
caudal vertebra (26.6 mm), eleventh caudal vertebra (26.1 mm), twelfth caudal
vertebra (25.7 mm), thirteenth caudal vertebra (25.2 mm), fourteenth caudal
vertebra (26.3 mm), fifteenth caudal vertebra (26 mm), sixteenth caudal vertebra
(25.3 mm), seventeenth caudal vertebra (24.8 mm), eighteenth caudal vertebra
(23.5 mm), nineteenth caudal vertebra (22.9 mm), twentieth caudal vertebra (21.5
mm), twenty-first caudal vertebra (20.7 mm), twenty-second caudal vertebra (19.2
mm), twenty-third caudal vertebra (17.2 mm), twenty-fourth caudal vertebra (15.5
mm), chevrons, scapulae, incomplete coracoids, incomplete furcula, sternal plates
(56 mm), six sternal ribs (15.6, 18.6 mm), humeri (89, ~90.7 mm), radii (78,
75.5 mm), ulnae (79.1, ~77.1 mm), scapholunares, pisiforms, semilunate carpals, distal
carpals III, metacarpal I (14.2, 14.2 mm), phalanx I-1 (30, 29.1 mm), manual
ungual I, metacarpal II (50.8, 51.9 mm), phalanx II-1 (23.6, 25.6 mm), phalanx
II-2 (24.8 mm), manual ungual II, metacarpal III (49.2, 49 mm), phalanx III-1
(19.1, 19.6 mm), phalanx III-2 (4.8, 5.2 mm), phalanx III-3 (14 mm), manual
ungual III, manual ungual sheaths, partial ilia, pubes (90 mm), incomplete ischia,
femora (~104.8 mm), tibiae (139.9, 137.9 mm), proximal fibula, distal tarsal
IV, phalanx I-1, pedal ungual I, incomplete metatarsal II (~71.9 mm), phalanx
II-1 (9.7 mm), phalanx II-2 (10.9 mm), pedal ungual II (~23.7 mm), metatarsal
III (~77 mm), phalanx III-1 (15.7 mm), phalanx III-2 (11.3, 11.1 mm), phalanx
III-3 (11.9, 10.9 mm), pedal ungual III (18.1 mm), metatarsal IV (75.4 mm),
phalanx IV-1 (12.8 mm), phalanx IV-2 (8.7, 8.8 mm), phalanx IV-3 (7, 6.7 mm),
phalanx IV-4 (7.9, 7.2 mm), pedal ungual IV (17.1, 14.9 mm), metatarsal V (45.7
mm), pedal ungual sheaths, contour feathers, remiges, leg remiges, retrices
(Ji, 2002)
(QM V1002; material of Sinornithosaurus "zhaoi") (800 mm adult) skull, mandible, cervical series, dorsal series, dorsal
ribs, five uncinate processes, gastralia, sarum, incomplete caudal series, chevrons,
scapulae, furcula, sternal plates, humeri (~94 mm), radii, ulnae, metacarpal I, phalanx
I-1, manual unguals I, metacarpals II, phalanges II-1, phalanges II-2, manual
ungual II, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual
ungual III, manual claw sheaths, ilia, pubis, ischia, femora (109 mm), tibiotarsi
(one incomplete), fibulae, pedal ungual I, metatarsals II (one incomplete),
phalanx II-2, pedal ungual II, metatarsals III (one incomplete), phalanx III-1,
phalanx III-2, phalanx III-3, pedal ungual III, metatarsals IV (one incomplete),
phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, pedal
claw sheaths, contour feathers, remiges, leg remiges, retrices, actinopterygian
neural/haemal arches, ribs and fin rays, teleost cranial elements and two centra
(Zhang, 2007)
(TNP00996) (~500 mm) specimen including skull, tail (290 mm), femur (63 mm),
contour feathers, retrices and leg remiges (Xu, Zhou, Wang, Kuang, Zhang and
Du, 2003)
Late Barremian-Early Albian, Early Cretaceous
Yixian or Jiufotang Formation, Liaoning, China
(IVPP 12727) specimen including eight dorsal vertebrae, dorsal ribs,
gastralia, caudal vertebrae, coracoid, proximal humerus, distal radius,
ulnar fragment, scapholunare, metacarpal I, pubes, femur and tibia
(Turner et al., 2012)
(IVPP V17749) specimen including radius, ulna, scapholunare, semilunate carpal, distal
carpal II, proximal metacarpal I, proximal metacarpal II and proximal metacarpal
III (Xu et al., 2014)
(IVPP V17750) specimen including radius, scapholunare, semilunate carpal, metacarpal
I, phalanx I-1, metacarpal II and metacarpal III (Xu et al., 2014)
(IVPP 200211) specimen including caudal vertebrae and tibia (Turner et al.,
2012)
(IVPP coll.) specimen including caudal vertebrae, manual digit I, pubis and
tibia (Turner et al., 2012)
(IVPP coll.) specimen including caudal vertebrae, radius, manual digit I, pubis
and tibia (Turner et al., 2012)
(IVPP coll.) specimen including teeth, caudal vertebrae, pubis, femur and tibia
(Turner et al., 2012)
Early Cretaceous
Qianyang, Liaoning, China
(CAGS 20-7-004) (~590 mm) dentaries (25.1, 33.36 mm), splenials, teeth, tenth
cervical vertebra (6.07 mm), tenth cervical rib, first dorsal vertebra (5.97
mm), second dorsal vertebra, third dorsal vertebra (7.4 mm), fourth dorsal vertebra
(6.54 mm), fifth dorsal vertebra (7.1 mm), sixth dorsal vertebra (5.39 mm),
seventh dorsal vertebra (5.18 mm), eighth dorsal vertebra (6.14 mm), ninth dorsal
vertebra (5.76 mm), tenth dorsal vertebra (5.97 mm), eleventh dorsal vertebra
(6.08 mm), twelfth dorsal vertebra (6.1 mm), thirteenth dorsal vertebra (6.12
mm), dorsal ribs, two uncinate processes, first caudal vertebra (6.19 mm), second
caudal vertebra (7.52 mm), third caudal vertebra (8.58 mm), fourth caudal vertebra
(13.12 mm), fifth caudal vertebra (16.54 mm), sixth caudal vertebra (19.42 mm),
seventh caudal vertebra (18.49 mm), eighth caudal vertebra (17.90 mm), nineth
caudal vertebra (17.16 mm), tenth caudal vertebra (17.24 mm), eleventh caudal
vertebra (16.82 mm), twelfth caudal vertebra (17.30 mm), thirteenth caudal vertebra
(17.17 mm), fourteenth caudal vertebra (16.38 mm), fifteenth caudal vertebra
(15.94 mm), sixteenth caudal vertebra (16.36 mm), seventeenth caudal vertebra
(15.48 mm), eighteenth caudal vertebra (14.06 mm), nineteenth caudal vertebra
(13.79 mm), twentieth caudal vertebra (12.55 mm), twenty-first caudal vertebra
(11.22 mm), twenty-second caudal vertebra (10.64 mm), twenty-third caudal vertebra
(9.03 mm), twenty-fourth caudal vertebra (7.44 mm), twenty-fifth caudal vertebra
(7.62 mm), twenty-sixth caudal vertebra (5.58 mm), chevrons, partial scapulae,
furcula, sternal fragments, four sternal ribs, humeri (61.25, 62.06 mm), radius
(~48.01 mm), ulna (53.78 mm), semilunate carpals, metacarpal I (7.84 mm), phalanx
I-1 (20.45 mm), proximal metacarpal II, phalanx II-1 (13.26 mm), phalanx II-2
(15.88, 16.39 mm), manual ungual II (9.49, 10.26 mm), proximal metacarpal III,
phalanx III-1 (10.12 mm), phalanx III-2 (4.1 mm), phalanx III-3 (10.06, 10.37
mm), fragmentary ilia, incomplete pubes (52.73, ~52.39 mm), incomplete ischia
(25.06, 25.22 mm), femora (74.26, 74.75 mm), tibiotarsi (94.22 mm), fibulae
(~85.67 mm), metatarsal II (44.51 mm), phalanx II-1 (6.88 mm), phalanx II-2
(7.24, 6.77 mm), pedal ungual II (13.24, 12.88 mm), metatarsal III (47.76 mm),
phalanx III-1 (9.95 mm), phalanx III-2 (7.74 mm), phalanx III-3 (6.13 mm), pedal
ungual III (9.39 mm), metatarsal IV (46.8 mm), phalanx IV-1 (7.3 mm), phalanx
IV-2 (5.4 mm), phalanx IV-3 (5.01 mm), phalanx IV-4 (5.42, 5.01 mm), pedal ungual
IV (9.18, 8.95 mm), metatarsal V (22.03 mm) (Hwang, Norell, Qiang and Keqin,
2002)
(CAGS 20-8-001) (~590 mm) ninth cervical vertebra (3.51 mm), tenth cervical
vertebrae (4.18 mm), first dorsal vertebra, second dorsal vertebra, third dorsal
vertebra, fourth dorsal vertebra, fifth dorsal vertebra (6.38 mm), sixth dorsal
vertebra (6.34 mm), seventh dorsal vertebra (6.4 mm), eighth dorsal vertebra
(6.42 mm), ninth dorsal vertebra (6.45 mm), tenth dorsal vertebra (6.34 mm),
eleventh dorsal vertebra (6.32 mm), twelfth dorsal vertebra (6.24 mm), thirteenth
dorsal vertebra (6.12 mm), dorsal ribs, seven uncinate processes, gastralia,
sacrum (fourth- 5.28 mm; fifth- 4.91 mm), first caudal vertebra (4.25 mm), second
caudal vertebra (5.28 mm), third caudal vertebra (5.63 mm), fourth caudal vertebra
(6.16 mm), fifth caudal vertebra (7.42 mm), sixth caudal vertebra (10.32 mm),
seventh caudal vertebra (14.7 mm), eighth caudal vertebra (18.47 mm), nineth
caudal vertebra (18.91 mm), tenth caudal vertebra (18.45 mm), eleventh caudal
vertebra (18.8 mm), twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth
caudal vertebra (17.64 mm), fifteenth caudal vertebra (18.08 mm), sixteenth
caudal vertebra (16.96 mm), seventeenth caudal vertebra (17.08 mm), eighteenth
caudal vertebra (16.74 mm), nineteenth caudal vertebra (16.9 mm), twentieth
caudal vertebra (~16.06 mm), twenty-first caudal vertebra (15.82 mm), twenty-second
caudal vertebra (~15.67 mm), twenty-third caudal vertebra (~14 mm), twenty-fourth
caudal vertebra (11.49 mm), twenty-fifth caudal vertebra (~10.66 mm), twenty-sixth
caudal vertebra (9.46 mm), chevrons, scapulocoracoids, furcula, sternal plates
(~37.99, 38.39 mm), four sternal ribs, humeri (62.88 mm), radii (~48.3 mm),
ulnae (~53.5 mm), four manual phalanges, manual ungual I (14.4 mm), manual ungual
?III (10.27 mm), manual ungual sheaths, ilia (40.81, 40.04 mm), pubes (52.77
mm), ischia (26.2, 26.84 mm), femora (74.40, 74.77 mm), tibiotarsi 94.14, ~95.51
mm), fibulae (87.2 mm), distal tarsal III, metatarsal I (8.09 mm), phalanx I-1
(5.26, 5.35 mm), pedal ungual I (3.34 mm), metatarsal II (45.89 mm), phalanx
II-1 (7.38, 6.91 mm), phalanx II-2 (7.59 mm), pedal ungual II (~15.2, 16.04
mm), metatarsal III (49.39 mm), phalanx III-1 (10.05, 9.74 mm), phalanx III-2
(~8.24, 8.2 mm), phalanx III-3 (7.18, 8.1 mm), pedal ungual III (9.51, 9.37
mm), metatarsal IV (~48.5 mm), phalanx IV-1 (8.14, 8.39 mm), phalanx IV-2 (5.72,
5.97 mm), phalanx IV-3 (~4.37, 4.16 mm), phalanx IV-4 (5.66, 5.9 mm), pedal
ungual IV (9.02 mm), metatarsal V (23.98 mm) (Hwang, Norell, Qiang and Keqin,
2002)
Early Cretaceous
China
(STM 5-11) (subadult) specimen including sternal plates and humeri (O'Connor
et al., 2015)
(STM 5-50) (subadult) specimen including scapula, coracoid, furcula and sternal plate (Zhang et al., 2014)
(STM 5-225) specimen including dorsal vertebrae and femur (Gong et al., 2012)
(STM coll.) skeleton including tibiotarsus, metatarsal II, metatarsal III, metatarsal
IV, pedal phalanges, body feathers, metatarsal remiges (Alexander et al., 2010)
?(private coll.) skull, mandibles, cervical series, dorsal series, dorsal ribs,
sacrum, caudal series, chevrons, furcula, anterior sternum, humeri, radii, ulnae,
carpus, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1,
phalanx II-2, manual ungual II, metacarpal III, manual digit III, manual ungual
III, ilium, pubis, femora, tibiotarsi, metatarsal II, pedal digit II, metatarsal
III, pedal digit III, metatarsal IV, pedal digit IV, remiges (www.paleowonders.com.tw/picture/DN-008.jpg)
(Hong Kong Science Museum coll.) partial skull, cervical series, dorsal series,
sacrum, caudal series, scapula, humeri, radii, ulnae, manus, ilia, femora, tibiotarsi,
metatarsi, pes, remiges, leg remiges (hk.science.museum/eexhibit/images/sd_mg.jpg)
?(private coll.) skull, mandibles, cervical vertebrae, dorsal series,
dorsal ribs, gastralia, sacrum, caudal series, scapula, furcula,
humeri, radii, ulnae, carpus, metacarpal I, phalanx I-1, manual ungual
I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II,
metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual
ungual III, ischia, femora, tibiotarsi, metatarsi, pedal digit I, pedal
digit II, pedal digit III, pedal digit IV
(http://www.fossilmall.com/Science/Sites/China/Dromeosauridea/Dromeosauridea.jpg)
Diagnosis- (after Xu et al., 2000) posterior dentary teeth with constricted
bases (also in Richardoestesia gilmorei and Paronychodon; may
be plesiomorphic for Maniraptoriformes); accessory trochanter on femur.
(after Xu et al., 2003) manual phalanx III-1 >83% of phalanx III-3 in length (also in Wulong).
(after Gong et al., 2012) slender ischial shaft.
(proposed) manual phalanx I-1 <62% of metacarpal II in length; tibiofemoral
ratio <136%; metatarsal I <20% of metatarsal III in length; pedal phalanx
IV-4 >65% of phalanx IV-1; highly elongate leg remiges (>1.5 times femoral
length).
Other diagnoses- Xu et al. (2000) listed "mesial serrations absent
from all teeth" as an apomorphy of Microraptor, but large specimen
NGMC 00-12-A has mesial serrations on maxillary teeth (Ji, 2002), so this may
be size-related. Another possibility is that NGMC 00-12-A is a different species.
Xu et al. also listed mid caudal vertebrae 3-4 times length of anterior dorsal
vertebrae, but this is also seen in NGMC 91. Another supposed apomorphy listed
in the original description is less than 26 caudal vertebrae, but several subsequently
discovered specimens have 26 or more caudals. Finally, strongly curved pedal
unguals are also seen in NGMC 91 (Senter, 2007).
Contra Xu et al. (2003), the high metacarpal III/II ratio is not diagnostic,
being similar to Bambiraptor and matched by NGMC 91. The distal articulation
of manual phalanx III-3 does not appear to be smaller or more ventrally skewed
than NGMC 91.
Gong et al. (2012) listed the absence of antorbital fossa pitting in their diagnosis,
but this is plesiomorphic. Contra their diagnosis, LVH 0026 does appear to have
a 'subfenestral fossa', though more anteriorly placed than in Sinornithosaurus.
Manual phalanx III-1 is not longer than III-3 in all specimens (e.g. IVPP V13475,
CAGS 20-7-004, gui holotype). The ischium is not longer compared to the
pubis or femur than in Sinornithosaurus. Not all Microraptor specimens
have bowed tibiotarsi (e.g. IVPP coll. in Turner et al., 2012; fig. 20F).
Contra Turner et al. (2012), metatarsal V is not longer than other dromaeosaurids
such as Sinornithosaurus and Deinonychus in all specimens (e.g.
CAGS 20-7-004 and 20-8-001) and bowing is common in dromaeosaurids.
The "Archaeoraptor" debacle- The holotype was discovered in
1997, though its part and counterpart slabs were quickly separated. One slab
contained the tail, which was then fraudulently combined with a euornithine
bird skeleton by a Chinese farmer. It was smuggled out of the country then sold
at the 1998 Tuscon Gem Show to Czerkas. Currie recognized the legs were part
and counterpart slabs of the same bones, while Rowe and Aulenback independently
verified the composite nature of the specimen. National Geographic announced
the specimen in a press conference in October, and in November, Sloan (1999)
published a paper using the name "Archaeoraptor liaoningensis". This
was a nomen nudum because it explicitely stated the taxon was to be described
formerly in an official publication. That official publication was to have been
in Science or Nature, but both journals rejected it. In October, Xu had noticed
the tail of "Archaeoraptor" matched the one on the dromaeosaurid skeleton
he was describing. Xu located the body which went with this tail by contacting
fossil dealers in December. In April, Olson (2000) published an article purporting
to officially describe "Archaeoraptor" and attach that name to the
dromaeosaurid tail (with the latter as the lectotype). Several months later
in December, Xu et al. (2000) officially named Microraptor zhaoianus
based on the dromaeosaurid tail and associated anterior part of the skeleton
Xu had been studying. At this time, Olshevsky (DML, 2000) noted that Olson's
publication predated Xu et al.'s, and he believed that this made Microraptor
a junior synonym of "Archaeoraptor". Several days later, Creisler
(DML, 2001) pointed out the Olson's attempt to name "Archaeoraptor"
was invalid because the ICZN requires a diagnosis in a valid publication, while
Olson merely referenced the invalid article by Sloan. Creisler further indicated
Olson cannot designate a lectotype without a valid publication defining a holotype
first. Thus "Archaeoraptor" is still a nomen nudum, despite
Olson's efforts, and Microraptor zhaoianus is the valid name for the
IVPP V 12330 dromaeosaurid. The euornithine section (IVPP V 12444) was later
described by Czerkas and Xu (2002) as a new taxon- Archaeovolans repatriates,
which was in turn synonymized with Yanornis martini by Zhou et al. (2002).
Microraptor gui, Cryptovolans pauli and other specimens-
In 2001, IVPP 13476 was discovered, and later mentioned by Xu et al. (2003)
as Microraptor sp.. Four additional specimens (IVPP V13320, V13477, V13351,
V13352) were purchased by the IVPP in 2001-2002. Xu et al. (2003) mentioned
IVPP V13351 as another Microraptor sp. specimen, and IVPP V13477 as Dromaeosauridae
gen. et sp. indet., but described V13352 and V13320 as the holotype and paratype
of a new species, Microraptor gui. This was based on several characters
supposedly distinct from the M. zhaoianus holotype, which have been subsequently
questioned by Senter et al. (2004) and Turner et al. (2012). There is a prominent
biceps tubercle on the proximal radius of the gui holotype, which Senter
et al. and Turner et al. considered individual variation. Both papers correctly
noted the bent pubis and low ratio of metacarpal I plus manual phalanx I-1 compared
to metacarpal II (~87%) are unknown in the M. zhaoianus holotype, while
a curved tibiotarsus is found in that species as well (Turner et al. considered
the curvature to be taphonomic). Xu et al. also mention the fused sterna as
being different than other dromaeosaurids, but sterna are unknown in the M.
zhaoianus holotype. Proportional differences have now been shown to be plausibly
allometric, due to intermediate sized Microraptor specimens having intermediate
ratios (possibly besides the tibiofemoral ratio; see below). Contrary to popular
perception, the famous elongated, asymmetrical feathers attached to M. gui's
metatarsi and tibiotarsi were not absent in M. zhaoianus, merely unpreserved
in the holotype. Thus there seems to be almost no basis for separating M.
gui from M. zhaoianus,
let alone as a separate genus "Tetrapterornis" as was suggested by
Miller (2004) based on these same characters. The genus is a
nomen nudum as Miller always places it in quotation marks, and thus
violates ICZN Article 11.5- "a name must be used as valid for a taxon
when proposed." IVPP V13477 has the apomorphically long leg remiges of Microraptor, so is here referred to that genus. TNP00996 was
purchased in 2002 and briefly described by Xu et al. (2003) as Microraptor
sp.. Senter (2011) has since retained gui as a separate OTU than
zhaoianus, though there are no differences between the two listed in
his table S1. O'Connor et al. (2011) described a new specimen (IVPP V17972)
as M. gui based on its large size relative to M. zhaoianus, manual
digit proportions, pubic curvature and slight bowing of the tibia. Of these
only size is new, which could easily be ontogenetic as sutures are visible in
the M. zhaoianus holotype sacrum.
Hwang et al. (2002) described two new Microraptor specimens (CAGS 20-7-004
and 20-8-001) which were collected by farmers from an unknown locality. These
were assigned to M. zhaoianus, as M. gui had yet to be described.
Although published much later, the M. gui description was accepted for
publication several days before Hwang et al.'s paper was published, and Xu et
al. (2003) never reference either CAGS specimen. The presence of a biceps tubercle
cannot be determined due to preservation, and their tibiotarsi are curved. Though
metacarpal II is incomplete distally, 20-7-004 has digit II phalanges placed
in a position where the first digit could be as comparatively short as M.
gui. The pubic shape cannot be determined as the bones are preserved in
anterior view. Size and most proportions (ulnofemoral, ischiofemoral and metatarsofemoral
ratios) are intermediate between M. zhaoianus and M. gui, while
the tibiofemoral ratio is more similar to zhaoianus (126% compared to
128% in zhaoianus and 133% in gui). The sternal plates of CAGS
20-8-001 are unfused, unlike the M. gui holotype. Proportional differences
from M. gui include a smaller ulnohumeral ratio (85-87% vs. 101%), longer
phalanx I-1 compared to metacarpal I (261% vs. 211%), longer manual phalanx
II-2 compared to II-1 (122% vs. 103%), and longer manual phalanx III-1 compared
to III-3 (99% vs. 90%). Senter (2011) viewed the CAGS specimens as distinct
from zhaoianus and/or gui based on three characters. He claims
the latter two species lack serrated maxillary and dentary teeth, but the zhaoianus
type has serrations clearly described and illustrated. IVPP V13320 is indeed
unique among reported Microraptor specimens in lacking serrated mid-jaw
teeth, but the condition is unpreserved in the gui holotype. Unfused
sternal plates do indeed differ from the gui holotype as noted above.
Finally, as noted above, manual phalanx I-1 is much longer compared to metacarpal
I (261%) than in the gui holotype (211%), but this is also true of Cryptovolans
(252%) and hanqingi (250%), though not of NGMC 00-12-A (211%) or IVPP
V13475 (209%).
LPM 0200 was first briefly described as an unnamed dromaeosaurid by Norell et
al. (2002), then described further and named Cryptovolans pauli by Czerkas
et al. (2002) several months later. Most of the bones in the holotype and paratype
(LPM 0159) are split between slab and counterslab, leading to very poor preservation
and a lack of anatomical detail. Norell et al. correctly described the elongate
leg remiges, which Czerkas et al. mistook for forelimb feathers. However, Czerkas
et al. were correct in describing the fused sternum and asymmetrical remiges,
contra Norell et al.. Czerkas et al. also believed NGMC 91 to possibly be referrable
to Cryptovolans, but this specimen is more likely Sinornithosaurus
(Ji et al., 2002) or the sister taxon to Microraptor (Senter et al.,
2004; Senter, 2007). Most publications have ignored Cryptovolans, perhaps
due to the poor description, highly illogical discussion and various personal
disputes with Czerkas' publishing practices.
Immediately after Xu et al.'s (2003) publication of M. gui, Holtz (DML,
2003) proposed it is a junior synonym of Cryptovolans pauli (using the
new, unpublished combination "Microraptor pauli"). He noted
the biceps tubercle cannot be determined as absent in the latter, which also
has a bent pubis, curved tibiotarsus and fused sternal plates. The Cryptovolans
paratype seems to have a similarly short first manual digit (though poor preservation
makes this uncertain), contra Czerkas et al.'s (2002) illustration. Proportional
differences between Cryptovolans' paratype and M. gui include
a smaller ulnohumeral ratio (~92% vs. 101%) and longer manual phalanx III-1
vs. III-3 (142% vs. 90%). The CAGS specimens are more similar to Cryptovolans
in the ulnohumeral ratio, and to M. gui in the manual digit III phalangeal
proportions. The Cryptovolans specimens are more similar to the CAGS
specimens than M. gui in their ulnohumeral ratios, long manual phalanx
I-1 compared to metacarpal I (252%), and long manual phalanx III-1 compared
to III-3, but are more similar to M. gui in their shorter manual phalanx
II-2 compared to II-1 (105%). Similarly, Miller (2004) stated "Under my own classification, M. gui and C. pauli would both become 'Tetrapterornis' gui", despite the fact both Cryptovolans and pauli have priority over "Tetrapterornis" and gui.
Two of the only papers to mention Cryptovolans (Senter et al., 2004;
Senter, 2007) agree it is a junior synonym of Microraptor zhaoianus.
Senter et al. state the manual proportions that diagnose Cryptovolans
are also seen in other Chinese microraptorians. Czerkas et al. diagnosed Cryptovolans
by its high III-1/III-3 ratio (138-142%). If we examine the ratio in other microraptorians,
Microraptor's (CAGS 20-7-004) is 98%, Microraptor gui's holotype
is 85%, NGMC 91's is 75%, Sinornithosaurus millenii's is 76%, and Graciliraptor's
is 64%. Senter et al. are however correct in noting that the Microraptor
gui holotype also has fused sternal plates, and that the unfused sternal
plates of another Microraptor specimen (CAGS-20-8-001) might be due to
ontogeny. Czerkas' final listed diagnostic character (28-30 caudal vertebrae)
is similar to Microraptor zhaoianus' holotype (24), Microraptor
specimens CAGS 20-7-004 and 20-8-001 (26) and Microraptor gui' holotype
(~26). Other maniraptoriform species (Gallimimus bullatus, Shenzhouraptor
sinensis) are known to have individual variation in caudal count within
3-4 vertebrae. Czerkas et al. also stated that the tibiofemoral ratio of the
paratype scales differently compared to Microraptor, which they viewed
as evidence the taxa were not synonymous. Yet the Microraptor gui holotype
has almost identical hindlimb size and ratio to the Cryptovolans holotype,
while the paratype would seem to have an unsually long femur. It seems unlikely
this is due to allometry, as Microraptor shows almost no change in tibiofemoral
ratio with age. More likely, individual variation or poor preservation is the
cause. Finally, Czerkas et al. stated that Cryptovolans has a longer
forelimb than other dromaeosaurids (hum+rad+mcII+pII-1+pII-2 239% of femoral
length), but it is subequal or shorter in length to Microraptor (237-258%),
Sinornithosaurus millenii (252%), NGMC 91 (256%) and Bambiraptor
(244%). Senter (2011) reversed his opinion and retained Cryptovolans
as a separate OTU, where it grouped with Sinornithosaurus and Graciliraptor
to the exclusion of Microraptor. This was due to several characters.
Cryptovolans has a longitudinal labial depression depression on some
teeth, but this is polymorphic in many theropods and also found in some teeth
of the zhaoianus holotype and M. hanqingi for instance. The second
character is the supposed lack of dorsal arching (when the articular surface
is held vertically) in manual ungual I, but this is impossible to determine
in LPM 0159 due to the proximal portion of ungual being broken (LPM 0200's hands
are impossible to evaluate in Czerkas et al.'s photos), and contra Senter is
not true of Sinornithosaurus or NGMC 91. Senter also claims Cryptovolans
lacks a proximodorsal lip on manual ungual I, but not only is this variable
in other taxa (e.g. Archaeopteryx, Sapeornis, Confuciusornis),
it is an illusion in Cryptovolans caused by the aforementioned broken
proximal portion of that ungual. The final character is tibiotarsal bowing,
which Senter incorrectly considered absent in Microraptor.
Czerkas and Ji (2002) illustrate a dromaeosaurid preserved alongside the Omnivoropteryx
holotype, referring it tentatively to Cryptovolans. Unfortunately, the
illustrations are of x-rays, so little detail can be ascertained. The ulnohumeral
and metatarsotibial ratios are more similar to Microraptor specimen NGMC
00-12-A, while the short manual digit I is more similar to Microraptor
than Sinornithosaurus, so it is provisionally retained in the former
genus until it is described in more detail.
Ji (2002) described a dromaeosaurid (NGMC 00-12-A) found in 2000 as Sinornithosaurus
sp.. This was based on several characters, which are all found in other
microraptorians. The unserrated premaxillary teeth, posteriorly bifurcated dentary,
prominent obturator process and subarctometatarsalian metatarsus are also present
in Microraptor, the U shaped furcula is present in all microraptorians,
while the extremely short manual phalanx III-2 relative to III-1 is even more
similar to Microraptor than Sinornithosaurus. The ischium is more
similar to Microraptor than Sinornithosaurus in being slender
shafted with a more proximally placed distodorsal process, and the maxillary
teeth are more like Microraptor in being mesially unserrated. NGMC 00-12-A
seems to lack a pronounced biceps tubercle, but has a short first manual digit,
bent pubis and curved tibiotarsus. Unlike Microraptor gui and the holotype
and paratype of Cryptovolans, NGMC 00-12-A lacks a fused sternum (though
it is the largest well described specimen). In a few proportions, NGMC 00-12-A
is more similar to M. gui than Cryptovolans pauli, including the
small sternum (53% of femoral length vs. 49% in gui and 60% in pauli),
though in others it is more similar to C. pauli. The latter include the
(originally supposed to be apomorphic for Cryptovolans) ratio between
manual phalanx III-1 and III-3 (139% vs. 142% in pauli and 90% in gui).
NGMC 00-12-A is more similar to the CAGS specimens than M. gui in its
ulnohumeral ratio (89%) and manual phalanx III-1 compared to III-3, but more
similar to M. gui in its short manual phalanx I-1 compared to metacarpal
I (208%), and short manual phalanx II-2 compared to II-1 (101%). There are a
few differences from smaller Microraptor specimens- more slender dentary,
posterior dentary teeth serrated mesially, and taller posterior dorsal neural
spines (last dorsal neural spine 72% taller than centrum compared to 63% in
CAGS 20-8-001), longer pubofemoral ratio (86% vs. 71-77%), and shorter pedal
phalanx III-2 vs. III-1 (59% vs. 78-83%). These are possibly due to age.
Zhang (2007) proposed Sinornithosaurus "zhaoi" for QM V1002 with a short description that claimed to distinguish it from S. millenii
based on teeh that are thick and less sharp, and longer
arms. However, its humerofemoral ratio (~86%) is actually
shorter than the S. millenii holotype (~91%) and Microraptor teeth are typically less elongate than Sinornithosaurus'. The slender ischium, short tibia, elongate pedal phalanx IV-4 and long leg remiges all support assigning QM V1002 to Microraptor instead of Sinornithosaurus, and Xing et al. (2013) described it as a specimen of M. gui. This was based on the biceps tubercle, short manual
digit I, bent pubis, and a new character- manual ungual I shorter to be subequal
in size to ungual II. The holotype of M. zhaoianus doesn't preserve manual
unguals, and this is seemingly based on CAGS 20-8-001 which has been referred
to that species without basis. This is the only Microraptor specimen
with such a large ungual I compared to II, which is probably due to supposed
ungual II belonging to digit III instead. Both unguals are disarticulated and
the only ones preserved in that specimen. Notably, the sterna are unfused despite
being from an individual 12% larger than the M. gui holotype. Sinornithosaurus
"zhaoi" is a nomen nudum as there is no "explicit fixation of a
holotype, or syntypes" (ICZN Article 16.4.1), with photos of the skull
and pes merely captioned "One skull of Sinornithosaurus zhaoi (sp.nov.)" and "One claw of Sinorni thosaurus [sic] zhaoi (sp.nov.)" respectively and no mention of its catalogue number.
Alexander et al. (2010) used LVH 0026 for their biomechanical work, considering
it "probably a different species of Microraptor but is morphologically
closely similar to M. gui, including the presence of flight-adapted feathers
on the tarsometatarsus" (though remember M. zhaoianus' holotype
merely doesn't preserve metatarsal remiges, and may have had them in life).
Gong et al. (2012) later described it as the new species Microraptor hanqingi.
They diagnosed it based on several characters. The larger size (11-14% larger
than M. gui or Cryptovolans) could be ontogenetic or individual
variation. Unfused sternals are also present in CAGS 20-80-001 and NGMC 00-12-A.
The pubis is not more robust than in M. gui, and it cannot be determined
if the pubic boot is more squared since most of M. gui's is hidden behind
the tibiotarsus. The pubis is indeed bent ~10 degrees less than in the M.
gui holotype, but some other specimens lack much bending at all (e.g. IVPP
V13475). Contra Gong et al., the ischia of M. hanqingi are posteriorly
sinuous and those of M. gui are anteriorly concave, so do not differ
in these respects. Additionally, the short manual digit I was said to be like
M. gui but unlike M. zhaoianus, and this is true for a referred
M. zhaoianus IVPP V13475 and the Cryptovolans paratype, while
NGMC 00-12-A is also like M. hanqingi and M. gui. Yet these ratios
do not correspond to other skeletal differences as noted in the taxonomy conclusion
below. The supposedly longer metatarsal II compared to IV was also supposed
to be more similar to M. gui than to M. zhaoianus, but the latter's
holotype has a ratio of 98% compared to 91% in M. gui's holotype. In
addition, other Microraptor specimens fall between those two ratios,
and 7% differences or more are known in other coelurosaurs (e.g. Archaeopteryx
lithographica and Dromiceiomimus brevitertius). Finally, its 23 caudal
vertebrae were said to be less than M. gui's ~26, but as noted in Cryptovolans'
discussion other Microraptor specimens fall between these measures and
such variation is known in other species. Pei et al. (2014) agree M. hanqingi
is a junior synonym of M. zhaoianus.
In conclusion, the problem with splitting the Microraptor-Cryptovolans
clade into species is not the absence of variation, as most of the described
specimens have some proportions or characters which differ from all or most
other specimens. Instead, the problem is that these differences don't vary in
a systematic, congruent way. While we might argue the Cryptovolans specimens
and M. gui holotype should be grouped together to the exclusion of M.
hanqingi, CAGS 20-80-001, NGMC 00-12-A and QM V1002 based on their fused
sterna, we could equally as well argue the Cryptovolans specimens, M.
hanqingi and NGMC 00-12-A should be grouped together to the exclusion of
the M. gui holotype and CAGS 20-7-004 based on their long manual phalanx
III-1. Or that Cryptovolans, M. hanqingi and CAGS 20-7-004 should
be grouped together to the exclusion of the M. gui holotype, NGMC 00-12-A
and QM V1002 based on their long manual phalanx I-1. There is no obvious answer.
One possibility would be to diagnose a separate species for each specimen, and
indeed the variation may be due to several different species living in the Jiufotang
fauna. This approach has since been taken by Senter (2011), who views M.
zhaoianus, M. gui, the CAGS specimens and Cryptovolans pauli
as separate species, with the latter closer to Sinornithosaurus and Graciliraptor.
The more conservative approach taken by Senter et al. (2004) and Turner et al.
(2012) is followed here, where differences are ascribed to individual and ontogenetic
variation. Further description of the specimens mentioned by Xu et al. (2003)
and others will help confirm or deny this view.
Maranga et al. (2020) preliminarily describe a new skull (JLUM Y-MR160501) from Sihedang they call Microraptor
sp., notable for having "denticles on both the mesial and distal
carinae of the posterior dentary teeth, a characteristic that has never
been observed in Microraptor
to date." However as noted above this is also present in NGMC
00-12-A. Oddly they recover the specimen sister to Microraptoria
plus eudromaeosaurs using Currie's dromaeosaurid analysis.
A number of undescribed specimens appear to belong to Microraptor as
defined here. One on the Paleowonders website is particularily complete, preserving
faint remiges. It was advertised as Microraptor gui at a fossil show.
The manual phalanx I-1 length suggests it is in fact Microraptor. One
photo of the specimen seems more simplified, lacks an ilium, and has contaur
feather impressions. It may be a cast. The specimen appears to preserve rather
thick, elongate bones in the pelvic and thoracic areas which are near certainly
from another animal.
Another specimen is shown at the Hong Kong Science Museum's "Soaring Dinosaurs"
exhibit, advertised as Microraptor gui. This specimen is largely complete,
but strangely positioned, with the presacral column sharply curved backward
and the hindlimbs extending anteriorly. The elongate leg remiges confirm the
identification as Microraptor.
Yet another specimen is complete and perfectly articulated, photographed on
Fossil Mall's website as Dromaeosauridae cf. Sinornithosaurus. Its manual
phalanx III-3 is elongate like Sinornithosaurus, yet its tibiofemoral
ratio and manual phalanx I-1 are short as in Microraptor. It is tentaively
referred to Microraptor here.
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unnamed clade (Bambiraptor feinbergi + Dromaeosaurus albertensis)
unnamed dromaeosaurid (Senter, Kirkland, Deblieux and Madsen, 2010)
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Material- (UMNH VP 21751) incomplete radius
....(UMNH VP 21752) incomplete pubis
Comments- This was initially identified (Senter et al., 2010) as part
of what would later be named Yurgovuchia by Senter et al. (2012). The
latter referred it to Velociraptorinae based on the large pubic tubercle.
References- Senter, Kirkland, Deblieux and Madsen, 2010. Three new theropods
from the Cedar Mountain Formation (Lower Cretaceous) of Utah. Journal of Vertebrate
Paleontology. Program and Abstracts 2010, 162A.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Bambiraptor Burnham, Derstler,
Currie, Bakker, Zhou and Ostrom, 2000
= "Linsterosaurus" Buchholz, DML, 1997
B. feinbergi Burnham, Derstler, Currie, Bakker, Zhou and Ostrom,
2000
= Bambiraptor feinbergorum Norell and Makovicky, 2004
Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
Holotype- (AMNH 30556; = AMNH 001; = FIP 001; Bambi) (subadult) skull
(127 mm), sclerotic plates, stapes, incomplete mandibles (~122 mm), hyoid, atlas,
axis (13 mm), third cervical vertebra (13.5 mm), fourth cervical vertebra (13.5
mm), fifth cervical vertebra (16 mm), sixth cervical vertebra (14.5 mm), seventh
cervical vertebra (~15 mm), eighth cervical vertebra (12 mm), ninth cervical
vertebra (13 mm), fragmentary anterior cervical ribs, first dorsal vertebra
(11 mm), second dorsal vertebra (12 mm), third dorsal vertebra (12 mm), fourth
dorsal vertebra (11.8 mm), fifth dorsal vertebra (11 mm), sixth dorsal vertebra
(13 mm), seventh dorsal vertebra (10 mm), eighth dorsal vertebra (10.5 mm),
ninth dorsal vertebra (9.5 mm), tenth dorsal vertebra (10 mm), eleventh dorsal
vertebra (10.5 mm), twelfth dorsal vertebra (10.8 mm), thirteenth dorsal vertebra
(10.5 mm), dorsal ribs, gastralia, first sacral vertebra (11 mm), second sacral
vertebra (15 mm), third sacral vertebra (15 mm), fourth sacral vertebra (13
mm), fifth sacral vertebra (11.5 mm), first caudal vertebra (9.1 mm), second
caudal vertebra (12.6 mm), third caudal vertebra (13.1 mm), fourth caudal vertebra
(14.5 mm), fifth caudal vertebra (15.6 mm), sixth caudal vertebra (16.1 mm),
seventh caudal vertebra (17.9 mm), eighth caudal vertebra (20.3 mm), ninth caudal
vertebra (21.6 mm), tenth caudal vertebra (24.2 mm), eleventh caudal vertebra
(24.6 mm), twelfth caudal vertebra (26 mm), thirteenth caudal vertebra (~27
mm), fourteenth caudal vertebra (27 mm), fifteenth caudal vertebra (~26 mm),
sixteenth caudal vertebra (~24 mm), seventeenth caudal vertebra (23 mm), eighteenth
caudal vertebra (22 mm), nineteenth caudal vertebra, twentieth caudal vertebra,
twenty-first caudal vertebra, twenty-second caudal vertebra, proximal chevron,
distal chevrons, scapulae (~83, 85 mm), coracoids, furcula, sternal plates (67,
63 mm), humeri (~105, 100 mm), radii (85, 85 mm), ulnae (95, 93 mm), scapholunares,
semilunate carpals, metacarpal I (16.8, 16.8 mm), phalanx I-1 (32.5, 32.3 mm),
manual ungual I, metacarpal II (47.8, 46.5 mm), phalanx II-1 (21.1, 21.4 mm),
phalanx II-2 (35, 35 mm), manual ungual II (43 mm), metacarpal III (44.9, 43.8
mm), phalanx III-1 (15.5, 16.5 mm), phalanx III-2 (6.3, 5.5 mm), phalanx III-3
(23.5 mm), ilia (86, ~78 mm), pubes (103, 103 mm), ischia (53, 50 mm), femora
(118, 118 mm), tibiae (167, 170 mm), fibulae (~170 mm), astragalus, calcaneum,
distal tarsal III, distal tarsal IV, metatarsal I (18.3 mm), phalanx I-1 (12.1
mm), pedal ungual I (14 mm), metatarsal II (70, 67.5 mm), phalanx II-1 (14.3,
14 mm), phalanx II-2 (14.6, 14.2 mm), pedal ungual II (46 mm), metatarsal III
(77, 81 mm), phalanx III-1 (29.6, 27.8 mm), phalanx III-2 (17.6, 15.4 mm), phalanx
III-3 (16.9 mm), pedal ungual III (24 mm), metatarsal IV (70, 74 mm), phalanx
IV-1 (23.6, 23.4 mm), phalanx IV-2 (16.6, 17.8 mm), phalanx IV-3 (11.6, 12.3
mm), phalanx IV-4 (12.3, 12.9 mm), pedal ungual IV (22 mm), metatarsal V (33.2
mm)
Paratypes- (FIP 002-136) (at least two adults) thirty-four elements including
caudal vertebrae, humerus (145 mm), femur (170 mm), tibia (225 mm), metatarsal
III (105 mm)
Referred- (MOR 553S-7-30-91-274) maxilla (~94 mm) (Currie and Varricchio, 2004)
Comments- The holotype was discovered in 1993 and initially believed
to be Saurornitholestes, then reported as Velociraptor sp. (Burnham
et al., 1997). Buchholz (DML 1997) used the name "Linsterosaurus"
for this taxon on the Dinosaur Mailing List between 1997 and 2000, while the
Oxford Museum of Natural History labeled their cast of the specimen Velociraptor
feinbergi (Taylor, DML 2003). Neither of these names have been published
however. It was later described as a new taxon of dromaeosaurid (Burnham et
al., 2000), though some believe it should be synonymized with Saurornitholestes. However, Hohman and Varricchio (2021) described juvenile Saurornitholestes maxilla MOR 9753 which is smaller than the Bambiraptor holotype, but shares characters with adult Saurornitholestes maxillae. Similarly, they find larger maxilla MOR 553S-7-30-91-274 groups with Bambiraptor in Currie and Varracchio's dromaeosaurid analysis and differs from Saurornitholestes. Thus the taxa are unlikely to be synonymous. Olshevsky (DML, 2000) noted Bambiraptor feinbergi was named after multiple
people, so suggested it be emended to B. feinbergorum, which is followed
on several websites and publications. However, the Fourth Edition of the ICZN
no longer requires emendations based on this reasoning (Article 31.1.3), as
Creisler pointed out later that day (DML, 2000). Norell and Makovicky (2004)
first published the name Bambiraptor feinbergorum, which according to
ICZN Article 32.2.3 is an available name with its own authorship, though an
objective junior synonym of B. feinbergi.
Currie and Evans (2020) identified the first and second premaxillary teeth, the latter of which is catalogued as "Paronychodon?" due to the ridges also found in Saurornitholestes ('Zapsalis') teeth.
Longrich and Currie (2009) found Bambiraptor to be a saurornitholestiine
in their analysis, and Brusatte et al. (2014) found the genera to be sister
taxa within Velociraptorinae. Foth et al. (2014) and Senter et al. (2012) have
both found Bambiraptor to be outside the Velociraptorinae+Dromaeosaurinae
clade, though neither included Saurornitholestes.
References- Buchholz, DML 1997. https://web.archive.org/web/20180115091825/http://dml.cmnh.org/1997Jun/msg00518.html
Burnham, Derstler and Linster, 1997. A new specimen of Velociraptor
(Dinosauria: Theropoda) from the Two Medicine Formation of Montana. Dinofest
International Proceedings. 73-75.
Burnham, Derstler, Currie, Bakker, Zhou and Ostrom, 2000. Remarkable new birdlike
dinosaur (Theropoda: Maniraptora) from the Upper Cretaceous of Montana. The
University of Kansas Paleontological Contributions. 13, 1-14.
Burnham and Durstler, 2000. Bambiraptor feinbergi (Dinosauria, Theropoda,
Dromaeosauridae). The Florida Symposium on Dinosaur Bird Evolution. Publications
in Paleontology No.2, Graves Museum of Archaeology and Natural History. 10.
Cooley, 2000. The Florida Symposium on Dinosaur Bird Evolution. Publications
in Paleontology No.2, Graves Museum of Archaeology and Natural History. 13.
Creisler, DML 2000. https://web.archive.org/web/20180115091851/http://dml.cmnh.org/2000Mar/msg00361.html
Derstler and Burnham, 2000. Phylogenetic Context of Bambiraptor feinbergi.
The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology
No.2, Graves Museum of Archaeology and Natural History. 15.
Garstka, Marsic, Carroll, Heffelfinger, Lyson and Ng, 2000. Analysis of the
structure and articulation of the forelimb bones of the maniraptoran dinosaur,
Bambiraptor feinbergi, support predation as the pre-adaptation of flight.
The Florida Symposium on Dinosaur Bird Evolution. Publications in Paleontology
No.2, Graves Museum of Archaeology and Natural History. 16.
Olshevsky, DML 2000. https://web.archive.org/web/20180115091846/http://dml.cmnh.org/2000Mar/msg00342.html
Taylor, DML 2003. https://web.archive.org/web/20180115091925/http://dml.cmnh.org/2003Oct/msg00210.html
Burnham, 2004. New information on Bambiraptor feinbergi (Theropoda: Dromaeosauridae)
from the Late Cretaceous of Montana. in Currie, Koppelhus, Shugar and Wright
(eds). Feathered Dragons. Studies on the transition from dinosaurs to birds.
Indiana University Press. 67-111.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation
(Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright
(eds). Feathered Dragons. Studies on the transition from dinosaurs to birds.
Indiana University Press. 112-132.
Norell and Makovicky, 2004. Dromaeosauridae. in Weishampel, Dodson and Osm�lska
(eds.). The Dinosauria (second edition). University of California Press, Berkeley.
196-209.
Senter, 2006. Comparison of forelimb function between Deinonychus and
Bambiraptor (Theropoda: Dromaeosauridae). Journal of Vertebrate Paleontology.
26(4), 897-906.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
from the Late Cretaceous of North America. Proceedings of the National Academy
of Sciences. 106(13), 5002-5007.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Currie and Evans, 2020 (online 2019). Cranial anatomy of new specimens of Saurornitholestes langstoni
(Dinosauria, Theropoda, Dromaeosauridae) from the Dinosaur Park
Formation (Campanian) of Alberta. The Anatomical Record. 303(4),
691-715.
Hohman and Varricchio, 2021. Ontogenetic implications of two distinct
dromaeosaurid maxillae from the Cretaceous (Campanian) Two Medicine
Formation of Montana, U.S.A. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 141-142.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as
2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist
in testing and constructing morphological characters. Journal of
Vertebrate Paleontology. 41(5), 2010087.
Variraptor Le Loeuff and Buffetaut,
1998
V. mechinorum Le Loeuff and Buffetaut, 1998
Late Campanian-Early Maastrichtian, Late Cretaceous
Gres a Reptiles Formation, France
Holotype- (MDE-D168) thirteenth dorsal vertebra
....(MDE-D169) (sacrum 160 mm) first sacral vertebra (25 mm), second sacral
vertebra (27 mm), third sacral vertebra (32 mm), fourth sacral vertebra (32
mm), fifth sacral vertebra (30 mm)
....(CM-645) ilium (230 mm) (Chanthasit and Buffetaut, 2009)
Paratypes- ?(MDE-D01) anterior dorsal vertebra (29 mm) (Le Loeuff, Buffetaut,
Mechin and Mechin-Salessy, 1992)
?(MDE-D158) humerus (195 mm)
?(MDE-D203; cast is MDE-D49 and TMP 1991.185.0001) incomplete femur (~230 mm) (Le Loeuff, Buffetaut,
Mechin and Mechin-Salessy, 1992)
(MDE coll.) fifth sacral vertebra (Le Loeuff, Buffetaut, Mechin and Mechin-Salessy,
1992)
Referred- ?(CM-186) tooth (FABL 4.6 mm) (Chanthasit and Buffetaut, 2009)
?(CM-218) tooth (FABL 7.1 mm) (Chanthasit and Buffetaut, 2009)
?(CM-307) ulna (145 mm) (Chanthasit and Buffetaut, 2009)
?(CM-537) tooth (FABL 5.9 mm) (Chanthasit and Buffetaut, 2009)
?(CM-684) tooth (FABL 6.7 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M187) femur (214 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M228) pedal ungual III or IV (46.6 mm on curve) (Chanthasit and Buffetaut,
2009)
?(MC-M324) tooth (FABL 5 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M561) tooth (FABL 4.3 mm) (Chanthasit and Buffetaut, 2009)
?(MC-M938) tooth (Chanthasit and Buffetaut, 2009)
(MC-PSP6) (sacrum 134 mm) first sacral vertebra (20.4 mm), second sacral vertebra
(26 mm), third sacral vertebra (28.4 mm), fourth sacral vertebra (28.6 mm),
fifth sacral vertebra (22.6 mm) (Chanthasit and Buffetaut, 2009)
?(MC-VC1) tooth (Chanthasit and Buffetaut, 2009)
?(MC-VC2) tooth (FABL 6 mm) (Chanthasit and Buffetaut, 2009)
?(MC coll.) proximal manual ungual (Chanthasit and Buffetaut, 2009)
? several fragmentary dorsal ribs (Le Loeuff, Buffetaut, Mechin and Mechin-Salessy,
1992)
Diagnosis- (proposed) sacral centra three and four transversely compressed
(transverse width <60% of first sacral centrum).
Comments- Le Loeuff et al. (1992) described a femur (MDE-D203) from Metisson
(Fox-Amphoux, Var), and an anterior dorsal vertebra (MDE-D01), posterior sacral
vertebra (MDE coll.) and several fragmentary dorsal ribs from Roques-Hautes
(Bouches-du-Rhone). They believed these were congeneric or at least related
to Elopteryx, most closely related to dromaeosaurids, though perhaps
deserving their own family or subfamily (Elopterygidae or Elopteryginae). The
femur was only stated to share general characteristics with Elopteryx
(reduced fourth trochanter, posterior trochanter, "shape and size")
while differing in having a linear capital ligament fossa and absent fourth
trochanter. The other remains are not comparable to Elopteryx. It is
here tentatively assigned to Variraptor as it is from the type locality.
Buffetaut et al. (1997) first mention the sacrum (MDE-D169) and humerus (MDE-D158)
from Bastide Neuve (Fox-Amphoux, Var) as a new taxon of dromaeosaurid. Le Loeuf
and Buffetaut (1998) described and named this taxon Variraptor, with
the sacrum and an articulated dorsal vertebra as the holotype. They referred
the anterior dorsal and posterior sacral described by Le Loeuff et al. (1992),
as well as a femur (MDE-D49) and the aforementioned humerus. The fifth sacral
was said to be identical to that of the holotype, though the other remains are
not comparable. The femur was not mentioned further, and seems to be a cast
of MDE-D203.
Buffetaut et al. (1999) first mentioned teeth (MC-M324, M561 and M938) and a
femur (MC-M187) from Massecaps (Cruzy, Herualt) and referred it to Variraptor.
Chanthasit and Buffetaut (2009) later described this material, as well as a
partial manual ungual (MC coll.) and pedal ungual (MC-M228) from the same locality.
Chanthasit and Buffetaut also described two teeth (MC-VC1 and VC2) from Combebelle
and a sacrum (MC-PSP6) from Plo Saint Pons, both also in Cruzy. Finally, they
describe five teeth (CM-186, 218, 307, 537 and 684), an ulna (CM-307) and an
ilium (CM-645) from Bastide Neuve. They noted the ilium matches the holotype
sacrum exactly, so probably belongs to the same individual, while the sacrum
has Variraptor's apomorphic proportions. Unfortunately, the femur was
not compared to MDE-D203, but the ulna does differ from Pyroraptor in
lacking a deep brachial fossa. Chanthasit and Buffetaut do not refer most of
their material to a particular genus, though it is all provisionally referred
to Variraptor here, given its presence in Bastide Neuve, Roques-Hautes
and Plo Saint Pons.
Allain and Taquet (2000) correctly noted no diagnostic dorsal or sacral characters
were described by Le Loeuff and Buffetaut, so they considered Variraptor
a nomen dubium. Yet as noted by Chanthasit and Buffetaut, Variraptor
does differ from all other comparable dromaeosaurids. The presence of posterior
dorsal pleurocoels distinguishes it from Microraptor and Velociraptor,
while it has one less pair of dorsal pleurocoels than Achillobator. The
pleurocoels are smaller than in Deinonychus however. The dorsal vertebra
is anteroposteriorly compressed, unlike microraptorians. Indeed, the compression
is a dromaeosaurine synapomorphy (Senter, 2007). Variraptor has less
sacrals than Deinonychus, Ornithodesmus and unenlagiines, and
its sacrum is not dorsally arched like those of Saurornitholestes and
Ornithodesmus. Variraptor differs from Microraptor, Sinornithosaurus,
Saurornitholestes, Ornithodesmus and Velociraptor in having
highly compressed mid sacral centra. It may be a synonym of Pyroraptor,
but it would be a senior synonym if so. The newly described ilium will probably
provide further diagnostic characters.
Rauhut (2000) refers Variraptor to Coelurosauria indet. since some Archaeopteryx
specimens and perhaps juvenile individuals of other maniraptoriform clades have
five sacral vertebrae, while he finds the rectangular posterior central face
of the last sacral (and presumably proximal caudals) to be potentially present
in other maniraptorans. Indeed, basal members of Therizinosauria, Oviraptorosauria
and Troodontidae all have five sacral vertebrae as well. Also, most non-pygostylian
maniraptorans have rectangular proximal caudal centra. Yet troodontids and basal
oviraptorosaurs lack posterior dorsal and sacral pleurocoels, and the high degree
of fusion in Variraptor suggests it was not a juvenile. The only non-dromaeosaurid
taxon with five sacral vertebrae, pleurocoelous posterior dorsals and anterior
sacrals, and rectangular proximal caudal vertebrae is Falcarius. Variraptor
is more similar to dromaeosaurids in having a hyposphene separated by the posterior
ligament groove, and shorter dorsal centra.
Rauhut further suggested the paratype anterior dorsal vertebra is a caenagnathid,
based on resemblence to Chirostenotes, especially in having two pairs
of pleurocoels. Yet Achillobator and Utahraptor both also have
two pairs of pleurocoels in some dorsals, and Variraptor resembles Deinonychus
more than Chirostenotes in having a longer infraprezygopophyseal fossa,
more poorly developed inradiapophyseal fossa, anteroposteriorly narrower neural
spine, and basally restricted posterior interspinous ligament groove. The prominent
epipophyses also suggest anterior cervical epipophyses would be well developed,
as in dromaeosaurids but not oviraptorosaurs. Rauhut's hypothesis is unsupported,
and Variraptor is not demonstrably a chimaera.
Senter et al. (2004) included Variraptor in a phylogenetic analysis and
found only that it was a coelurosaur closer to birds than tyrannosauroids, and
not a Jehol microraptorian or member of the Achillobator+Utahraptor+Dromaeosaurus
clade. However, their analysis lacked many relevent characters (precise sacral
number; sacral pleurocoel presence; posterior dorsal pleurocoel presence; proximal
caudal centrum shape; etc.), so this result is meaningless.
Hartman et al. (2019) recover the taxon sister to Bambiraptor,
while Longrich et al. (2021) suggest several characters are shared with
unenlagiines ("enlarged cuppedicus fossa, arched dorsal margin of the
ilium and deep anterior blade, short posterior blade, concave
posterodorsal margin of the blade, and a brevis fossa that is reduced
anteriorly").
References- Le Loeuff, Buffetaut, Mechin and Mechin-Salessy, 1992. The
first record of dromaeosaurid dinosaur (Saurichia, Theropoda) in the Maastrichtian
of Southern Europe: palaeobiogeographical implications. Bulletin de la Societe
Geologique de France. 163(3), 337-343.
Buffetaut, Le Loeuff, Cavin, Duffaud, Gheerbrant, Laurent, Martin, Rage, Tong
and Vasse, 1997. Late Cretaceous non-marine vertebrates from southern France:
a review of recent finds. Geobios. 20, 101-108.
Le Loeuff and Buffetaut, 1998. A new dromaeosaurid theropod from the Upper Cretaceous
of Southern France. Oryctos. 1, 105-112.
Buffetaut, Le Loeuff, Tong, Duffaud, Cavin, Garcia, Ward and L'association Culturelle,
Archeologique Et Paleontologique De Cruzy, 1999. A new Late Cretaceous vertebrate
locality at Crazy (Harault, southern France). Les Comptes rendus de l'Acad�mie
des sciences. 328, 203-208.
Allain and Taquet, 2000. A new genus of Dromaeosauridae (Dinosauria, Theropoda)
from the Upper Cretaceous of France. Journal of Vertebrate Paleontology. 20(2),
404-407.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). PhD thesis, University of Bristol. 583 pp.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae.
Bulletin of Gunma Museum of Natural History. 8, 1-20.
Chanthasit and Buffetaut, 2009. New data on the Dromaeosauridae (Dinosauria:
Theropoda) from the Late Cretaceous of southern France. Bulletin de la Societe
Geologique de France. 180(2), 145-154.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247.
Longrich, Martill and Jacobs, 2021. A new dromaeosaurid dinosaur from
the Wessex Formation (Lower Cretaceous, Barremian) of the Isle of
Wight, and implications for European palaeobiogeography. Cretaceous
Research. Journal Pre-proof. DOI: 10.1016/j.cretres.2021.105123
Eudromaeosauria Longrich and Currie, 2009
Definition- (Saurornitholestes langstoni + Velociraptor mongoliensis
+ Deinonychus antirrhopus + Dromaeosaurus albertensis) (Longrich
and Currie, 2009)
= Dromaeosauridae sensu Padian et al., 1999
Definition- (Dromaeosaurus albertensis + Velociraptor mongoliensis)
(modified)
"Dromaeosaurus" gracilis
(Marsh, 1888) Matthew and Brown, 1922
= Coelurus gracilis Marsh, 1888
= Chirostenotes gracilis (Marsh, 1888) Gilmore, 1924
Late Aptian-Early 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 Belly River Group of Canada at the AMNH (including AMNH
5387) were nearly identical to the holotype. Matthew and Brown (1922) provisionally
referred the taxon to their new genus Dromaeosaurus, as (?)Dromaeosaurus
gracilis. This was done without justification. Gilmore (1924) believed the
AMNH unguals belonged to Chirostenotes, and stated Coelurus gracilis "should now be removed to this genus" but didn't explicitly use the combination Chirostenotes gracilis. Ostrom (1969) notes a strong resemblence of the holotype to the first
manual ungual of Deinonychus, but considered it indeterminate. Lipka
(1998) reported recovering teeth similar to those described by Lull, which remain
undescribed. He noted Coelurus gracilis may relate to the Arundel Deinonychus-like
teeth he described.
The extremely large flexor tubercle and proximodorsal lip suggest the ungual
is from digit III as suggested by Lull, contra Ostrom. The holotype ungual differs
from Chirostenotes in being more curved, having a smaller proximodorsal
lip, more proximally placed flexor tubercle and a proximally undivided longitudinal
groove. Microvenator's unguals resemble Chirostenotes in being
less curved with more distally placed flexor tubercles. Hagryphus is
more similar in being more strongly curved with a more proximally placed flexor
tubercle than Chirostenotes, but still has a divided groove and larger
proximodorsal lip. These oviraptorosaurs also all have lower flexor tubercles
than USNM 4973. Among oviraptorids, only Citipati is similar in having
strongly curved third manual unguals with a proximodorsal lip, but differs in
having a lower flexor tubercle. This also seems to be the case for Yixianosaurus
and Tanycolagreus. Archaeopteryx and Confuciusornis unguals
are more slender and less curved with much lower flexor tubercles. This leaves
dromaeosaurids, among which microraptorians and Saurornitholestes seem
less similar than derived taxa like Velociraptor and Deinonychus
in having lower flexor tubercles. This suggests USNM 4973 is a derived dromaeosaurid,
perhaps a synonym of Deinonychus based on its provenence. Thus Matthew
and Brown's generic assignment is more accurate than Marsh's, and is followed
here. It should be noted that based on its early age, USNM 4973 is near certainly
not Dromaeosaurus however. The unguals mentioned by Gilmore are presumably
also dromaeosaurid, perhaps belonging to Saurornitholestes and/or Dromaeosaurus.
Lull's and Lipka's referred teeth may also be dromaeosaurid, though basal coelurosaurs
may also lack mesial serrations. More information is needed.
References- Marsh, 1888. Notice of a new genus of Sauropoda and other
dinosaurs from the Potomac Formation. American Journal of Science, 3rd Series.
35, 89-94.
Lull, 1911. The Reptilia of the Arundel Formation. Maryland Geological Survey,
Lower Cretaceous. 174-178.
Gilmore, 1920. Osteology of the carnivorous Dinosauria in the United States
National Museum with special reference to the genera Antrodemus (Allosaurus)
and Ceratosaurus. United States National Museum Bulletin. 110, l-154.
Matthew and Brown, 1922. The family Deinodontidae, with notice of a new genus
from Cretaceous of Alberta. Bulletin of the American Museum of Natural History.
66, 367-385.
Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta.
Canada Geological Survey, Bulletin 38, geological series 43, 1-13.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod
from the Lower Cretaceous of Montana. Peabody Museum of Natural History Bulletin.
30, 1-165.
Lipka, 1998. The affinities of the enigmatic theropods of the Arundel Clay facies
(Aptian), Potomac Formation, Atlantic Coastal Plain of Maryland. in Kirkland,
Lucas and Estep (eds). Lower to Middle Cretaceous Terrestrial Ecosystems. New
Mexico Museum of Natural History and Sciences Bulletin. 14, 229-234.
Vectiraptor Longrich, Martill and Jacobs, 2022
= "Vectiraptor" Longrich, Martill and Jacobs, 2021 online
V. greeni Longrich, Martill and Jacobs, 2022
= "Vectiraptor greeni" Longrich, Martill and Jacobs, 2021 online
Barremian, Early Cretaceous
Wessex Formation, England
Material- (IWCMS 2021.31.1) (~7-9 year old adult) incomplete posterior dorsal vertebra (~39 mm)
....(IWCMS 2021.31.2) partial anterior dorsal vertebra (~48 mm)
....(IWCMS 2021.31.3) partial synsacrum
Diagnosis- (after Longrich et
al., 2021) small anterior dorsal pleurocoels and large, elliptical
posterior
dorsal pleurocoels; camellate internal architecture in dorsal
vertebrae (also in Unenlagiinae); dorsal vertebrae with large neural
canals (also in other paravians); dorsal vertebrae with deep,
triangular infradiapophyseal fossae (also in Deinonychus, Utahraptor and ?Itemirus);
infradiapophyseal fossae divided by accessory lamina in anterior dorsal
vertebrae; deep dorsal infrapostzygapophyseal fossae; massive dorsal
neural spines with broad ligament scars (also in Saurornitholestes);
sacral centra not pneumatized; ventral median groove not extending the
full length of sacrum; sacrum with massive neural canal (also in Saurornitholestes and Rahonavis).
Comments- The holotype was
collected in 2004 and described in an online Journal Pre-proof on December 17 2021 by
Longrich et al. (2021). As the paper has no mention of ZooBank, according to ICZN Article 8.5.3 (an electronic work
must "be registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Vectiraptor greeni" Longrich et
al., 2021 was a nomen nudum that became valid in the June
2022 issue.
Longrich et al. proposed "affinities with Eudromaeosauria,
perhaps as a primitive member of the group or sister to it" based on
broad interspinous ligament scars that differ from Unenlagia,
and three characters shared with that genus and Eudromaeosauria (short
dorsal centra, posterior dorsal pleurocoels, tall and anteroposteriorly
narrow dorsal neural spines). Based on this, they suggested several
teeth from the Wessex Formation (IWCMS 2002.1, 3, 4 and NHMUK R 16510)
may be referrable as they are the correct size, although the
cooccurance of dromaeosaurid species in other formations makes this
speculative. Adding it to Hartman et al.'s maniraptoromorph matrix
results in it being a velociraptorine sister taxon to IGM 100/22-23.
Reference- Longrich, Martill
and Jacobs, 2022 (online 2021). A new dromaeosaurid dinosaur from the Wessex
Formation (Lower Cretaceous, Barremian) of the Isle of Wight, and
implications for European palaeobiogeography. Cretaceous Research.
134, 105123.
Yurgovuchia Senter, Kirkland,
DeBlieux, Madsen and Toth, 2012
Y. doellingi Senter, Kirkland, DeBlieux, Madsen and Toth, 2012
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Holotype- (UMNH VP 20211) (adult) atlas, axis, incomplete third cervical
vertebra (45.1 mm), incomplete posterior cervical vertebra (~24.89 mm), incomplete
posterior cervical vertebra (~40.95 mm), two partial cervical neural arches,
first dorsal vertebra (~26.42 mm), partial dorsal vertebra (32.04 mm), partial
dorsal neural arch, incomplete proximal caudal vertebra (40.17 mm), incomplete
proximal caudal vertebra (45.95 mm), proximal caudal vertebra (48.03 mm), mid
caudal vertebra (52.91 mm), mid caudal vertebra, mid caudal vertebra (55 mm),
distal caudal centrum (~48.34 mm), proximal pubis
Diagnosis- (after Senter et al., 2012) centrum of axis with single pneumatopore
on each side; cranial end of centrum of third cervical vertebra not beveled;
cervical prezygapophyses flexed; epipophyses of cervical vertebrae above postzygapophyseal
facets; anterior dorsal vertebrae with hypapophyses and without pneumatopores;
anterior faces of centra of proximal caudal vertebrae round; caudal prezygapophyses
elongated distal to transition point, but not over the length of a centrum;
pubis without pubic tubercle.
Comments- The holotype was discovered in 2005 and initially mentioned
by Senter et al. (2010). The latter noted a lacrimal, radius and ilium among
the material, which were not in the final description. They state the pubis
has a marked pubic tubercle, unlike the holotype. This indicates Senter et al.
included UMNH VP 21751 and 21752 in the hypodigm, though they later (2012) described
these as a velociraptorine instead. As the supposed lacrimal was stated to lack
an anterodorsal process, it's possible this ended up being the holotype proximal
pubis. The identity of the supposed ilium is unknown. Senter et al. (2012) found
Yurgovuchia to be a dromaeosaurine more derived than Deinonychus
in their analysis, using a modified form of Senter's TWiG-based matrix.
References- Senter, Kirkland, Deblieux and Madsen, 2010. Three new theropods
from the Cedar Mountain Formation (Lower Cretaceous) of Utah. Journal of Vertebrate
Paleontology. Program and Abstracts 2010, 162A.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Dromaeosaurinae Matthew and Brown, 1922
Definition- (Dromaeosaurus albertensis <- Velociraptor mongoliensis)
(Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017; modified from Sereno, 1998)
Other definitions- (Dromaeosaurus albertensis <- Microraptor
zhaoianus, Velociraptor mongoliensis, Unenlagia comahuensis, Passer domesticus)
(Turner et al., 2012)
= Dromaeosaurinae sensu Turner et al., 2012
Definition- (Dromaeosaurus albertensis <- Microraptor zhaoianus,
Velociraptor mongoliensis, Unenlagia comahuensis, Passer domesticus)
Comments- The comments for Velociraptorinae apply here as well, though
I consider Passer even less necessary in Turner et al.'s definition.
References-
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Late Cenomanian, Late Cretaceous
Dakota Formation, Utah, US
Material- teeth
Comments- These remains were listed as Dromaeosaurinae indet. by Kirkland
et al. (1998) and Eaton et al. (1999). They are presumably one of the two Dromaeosauridae
indet. gen. et sp. teeth listed by Kirkland et al. (1997).
References- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton,
Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the
Central Colorado Plateau: a key to understanding 35 million years of tectonics,
sedimentology, evolution, and biogeography. Brigham Young University Geology
Studies. 42, 69-103.
Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the Colorado Plateau.
in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous Terrestrial
Ecosystems. New Mexico Museum of Natural History and Science Bulletin. 14, 79-89.
Eaton, Cifelli, Hutchison, Kirkland and Parrish, 1999. Cretaceous vertebrate
faunas from the Kaiparowits Plateau, south central Utah. in Gillette (ed.).
Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous Publication.
99-1, 345-353.
Dromaeosaurinae indet. (Fiorillo, 1999)
Cenomanian-Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 71598) teeth and fragments (Fiorillo, 1999)
partial tooth (Garrison et al., 2007)
Comments- Fiorillo (1999) notes laterally compressed teeth which possess
labiolingually broad serrations which are not hooked apically. He refers these
to Dromaeosaurinae. Garrison et al. (2007) describe and illustrate a partial
tooth with flat serrations (6/mm) on the distal carinae with prominent blood
pits, which they refer to Dromaeosaurinae indet.. Cifelli et al. (1999) also
list Dromaeosaurinae indet. teeth.
References- Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999.
Medial Cretaceous vertebrates from the Cedar Mountain Formation, Emery County,
Utah: the Mussentuchit Local Fauna. in Gillette (ed.). Vertebrate Paleontology
in Utah. Utah Geological Survey, Miscellaneous Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison Eggshell
site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. in Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 259-268.
Garrison, Brinkman, Nichols, Layer, Burge and Thayn, 2007. A multidisciplinary
study of the Lower Cretaceous Cedar Mountain Formation, Mussentuchit Wash, Utah:
a determination of the paleoenvironment and paleoecology of the Eolambia caroljonesa
dinosaur quarry. Cretaceous Research. 28, 461-494.
Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Middle-Late Turonian, Late Cretaceous
Smoky Hollow Member of the Straight Cliffs Formation, Utah, US
Material- (OMNH 23712) tooth (Parrish, 1999)
(OMNH 25422) tooth (Parrish, 1999)
(OMNH 24437) tooth (Parrish, 1999)
(OMNH 24438) tooth (Parrish, 1999)
(OMNH 25420) tooth (Parrish, 1999)
Comments- These were listed as Dromaeosaurinae by Kirkland et al. (1998)
and Parrish (1999).
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian)
of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the
Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous
Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin.
14, 79-89.
Dromaeosaurinae indet. (Kirkland, Lucas and Estep, 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah, US
Material- (MNA NB-9) tooth (Parrish, 1999)
(OMNH 21222, 21318, 21230, 21281, 21335, 21340, 21395, 21868, 21991, 23636,
24307, 24308, 24327) thirteen teeth (Parrish, 1999)
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian)
of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
Dromaeosaurinae indet. (Parrish, 1999)
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Material- (UCM 83240) tooth (Parrish, 1999)
(UCM 8659; in part) tooth (Parrish, 1999)
(OMNH 21117, 23178, 23527, 23595, 23608, 23854, 23875, 23969-23975, 24157, 24159,
24160, 23851, 23565 (in part), 24381) nineteen teeth (Parrish, 1999)
Reference- Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-.
Judithian) of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah.
Utah Geological Survey, Miscellaneous Publication. 99-1, 319-321.
unnamed dromaeosaurine (Rauhut and Zinke, 1995)
Barremian, Early Cretaceous
Una Formation, Spain
Material- (IPFUB Una Th 37, 39, 48, 49, 67, 71-80) thirty-seven teeth
(<5 mm)
Comments- These teeth are moderately to strongly recurved, and some have
a very slight basal constriction. Most specimens have bot mesial and distal
serrations, with the mesial serrations restricted apically. Some have a reduced
number of distal serrations, and only weakly developed mesial serrations. One
lacks serrations entirely. The DSDI varies between .86 and 1.17, with an average
of 1.03. Serrations are chisel-shaped. The mesial carina twists lingually in
the basal half.
References- Rauhut and Zunke, 1995. A description of the Barremian dinosaur
fauna from Una with a comparison of that of Las Hoyas. II. International Symposium
of Lithographic Limestone, Extended Abstracts. 123-126.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca,
Spain. Cretaceous Research. 23, 255-263.
undescribed Dromaeosaurinae (Ortega, Escaso, Perez Garcia, Torices and
Sanz, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- teeth
Comments- Ortega et al. (2009) state teeth of Dromaeosaurinae are present.
Reference- Ortega, Escaso, Perez Garcia, Torices and Sanz, 2009. The
vertebrate diversity of the Upper Campanian-Lower Maastrichtian "Lo Hueco"
fossil-site (Cuenca, Spain). Journal of Vertebrate Paleontology. 29(3), 159A-160A.
undescribed Dromaeosaurinae (Averianov, 2007)
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- teeth
Comments- Averianov states these are similar to dromaeosaurine teeth
from the Bostobe Formation (ZIN PH 27/49 - 31/49). They probably belong to Itemirus.
Reference- Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits
in the northeastern Aral Sea region, Kazakhstan. Cretaceous Research.
unnamed Dromaeosaurinae (Nessov and Khisorova, 1988)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- (ZIN PH 27/49) tooth (Averianov, 2007)
(ZIN PH 28/49) tooth (Averianov, 2007)
(ZIN PH 29/49) tooth (Averianov, 2007)
(ZIN PH 30/49) tooth (Averianov, 2007)
(ZIN PH 31/49) tooth (Averianov, 2007)
(ZIN PH 33/49) (juvenile) tooth (FABL 3.1 mm) (Averianov, 2007)
two teeth (Dyke and Malakhov, 2004)
Comments- ZIN PH 27/49 - 31/49 have FABLs between 9.1 and 16.7 mm, BW/FABLs
of .54-.86, lingually shifted mesial carinae, 14 mesial serrations per 5 mm
and 15-17 distal serrations per 5 mm. Those described by Dyke and Malakhov (2004)
as dromaeosaurids were said to be similar. They are probably the same specimens
that were mentioned as dromaeosaurids by Nessov and Khisorova (1988) and Nessov
(1995). Averianov (2007) noted they had low DSDIs for dromaeosaurids and considered
the possibility they are anterior maxillary/dentary teeth from juvenile tyrannosauroids.
ZIN PH 33/49 has a BW/FABL of .48, a lingually displaced mesial
carina which may have originally had small serrations, 6 distal serrations per
mm, wrinkled enamel and two vertical ridges lingually. Averianov referred it
to Dromaeosauridae based on its small serrations, and to Dromaeosaurinae based
on its relative thickness and lingually displaced mesial carina. He viewed Paronychodon
as juvenile dromaeosaurid teeth, with this specimen as an intemediate age.
References- Nessov and Khisorova, 1988. [New data on vertebrates of the
Late Cretaceous in the localities Shakh-Shakh and Baibolat (North-Eastern Aral
Region)]. Matyerialy po istorii fauny i flory Kazakhstana. 10, 5-14.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
Dyke and Malakhov, 2004. Abundance and taphonomy of dinosaur teeth and other
vertebrate remains from the Bostobynskaya Formation, north-east Aral Sea region,
Republic of Kazakhstan. Cretaceous Research.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern
Aral Sea region, Kazakhstan. Cretaceous Research.
unnamed dromaeosaurine (Alifanov and Bolotsky, 2002)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- (1/314) tooth (19x7.5x2.9 mm) (Bolotsky, 2011)
(1/806) tooth (20.2x9.9x3.2 mm) (Bolotsky, 2011)
(1/1079) tooth (23.2x8.8x5.8 mm) (Bolotsky, 2011)
(1/1080) tooth (18x8.8x1.1 mm) (Bolotsky, 2011)
Comments- Alifanov and Bolotsky (2002) referred these remains to cf.
Dromaeosaurus sp., while Bolotsky (2011) described them as Dromaeosaurinae.
Reference- Alifanov and Bolotsky, 2002. New data about the assemblages
of the Upper Cretaceous carnivorous dinosaurs (Theropoda) from the Amur region.
In Kirillova (ed.). Fourth International Symposium of IGCP 434. Cretaceous continental
margin of East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
Bolotsky, 2011. On paleoecology of carnivorous dinosaurs (Tyrannosauridae, Dromaeosauridae)
from Late Cretaceous fossil deposits of Amur region, Russian far East. Global
Geology. 14(1), 1-6.
unnamed possible dromaeosaurine (Maisch and Matzke, 2020)
Oxfordian, Late Jurassic
Qigu Formation, Xinjiang, China
Material- (SGP 2002/6) tooth (10.3x5.15x3.3 mm)
(SGP 2002/7) tooth (9.0x4.8x2.5 mm)
Comments- Maisch and Matzke (2020) refer these to ?Dromaeosauridae indet. based largely on the twisted mesial carina.
Reference- Maisch and Matzke,
2020. Small theropod teeth (Dinosauria) from the Upper Jurassic Qigu
Formation of the southern Junggar Basin, NW China. Neues Jahrbuch f�r
Geologie und Pal�ontologie Abhandlungen. 295(1), 91-100.
unnamed possible dromaeosaurine (Han, Clark, Xu, Sullivan, Choiniere and Hone, 2011)
Late Oxfordian, Late Jurassic
Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Material- (IVPP V15857) tooth (7.1x3.7x2.1 mm)
Comments- Discovered by the Sino-American expeditions between 2001 and 2010, this is called
Morphotype 5 by Han et al. (2011), who refer it to a
dromaeosaurid. It is placed in that clade here based on the
twisted mesial carina.
Reference- Han, Clark, Xu,
Sullivan, Choiniere and Hone, 2011. Theropod teeth from the
Middle-Upper Jurassic Shishugou Formation of northwest Xinjiang, China.
Journal of Vertebrate Paleontology. 31(1), 111-126.
undescribed dromaeosaurine (Watanabe and Sereno, 2010)
Cenomanian, Late Cretaceous
Ulansuhai Formation, Inner Mongolia, China
Material- (juvenile or subadult) premaxilla, maxilla, lacrimal, frontal,
postorbital, quadrate, ectopterygoid, atlantal intercentrum, mid dorsal neural
arch, ischium
Diagnosis- (after Watanabe and Sereno, 2010) deep sagittal crest extending
onto frontals; hourglass-shaped hyposphene-hypantrum articulation in dorsal
vertebrae.
Comments- This is described as having a short snout and deep obturator
process. It was found to group with Dromaeosaurus, Utahraptor
and Achillobator.
Reference- Watanabe and Sereno, 2010. A large short-snouted dromaeosaurid
(Theropoda: Maniraptora) from Inner Mongolia. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 184A.
undescribed possible dromaeosaurine (Currie and Eberth, 1993)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- teeth
Comments- Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in
the Iren Dabasu" and that "Most of these can be attributed to Velociraptor,
although some of the teeth suggest that there was a second, larger
species of an indeterminate dromaeosaurine dromaeosaurid." A
pedal phalanx II-1 (AMNH 6572) mentioned by Ostrom (1969) as being 20%
larger than Deinonychus may belong
to the same taxon, based on size.
References- Ostrom, 1969. Osteology of Deinonychus antirrhopus,
an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum of
Natural History Bulletin. 30, 1-165.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic
of China. Cretaceous Research. 14, 127-144.
unnamed possible dromaeosaurine (Goodwin, Clemens, Hutchinson, Wood,
Zavada, Kemp, Duffin and Schaff, 1999)
Tithonian, Late Jurassic
Mugher Mudstone, Ethiopia
Material- (JN-96-2B/UCMP 170803) tooth
several teeth (Hall and Goodwin, 2011)
Comments- JN-96-2B/UCMP 170803 is slightly recurved, with a FABL of 9.3
mm and a BW of 7.3 mm. It has both mesial and distal serrations, but no further
details are given or visible in the photograph. The authors state the morphology
"compares well with the dentary teeth of Dromaeosaurus albertensis"
and describe it as having possible dromaeosaurine affinity. The lack of specific
similarities and wide temporal and spatial gulf between this specimen and verified
dromaeosaurines makes the assignment questionable.
References- Goodwin, Clemens, Hutchinson, Wood, Zavada, Kemp, Duffin
and Schaff, 1999. Mesozoic continental vertebrates with associated palynostratigraphic
dates from the northwestern Ethiopian Plateau. Journal of Vertebrate Paleontology.
19(4), 728-741.
Hall and Goodwin, 2011. A diverse dinosaur tooth assemblage from the Upper Jurassic
of Ethiopia: Implications for Gondwanan dinosaur biogeography. Journal of Vertebrate
Paleontology. Program and Abstracts 2011, 121.
Atrociraptor Currie and Varricchio,
2004
A. marshalli Currie and Varricchio, 2004
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
Holotype- (RTMP 95.166.1) (~1.7 m) (skull ~210 mm) premaxillae, maxilla,
dentaries (one partial), ?third premaxillary tooth (10x5x3.5 mm); anterior maxillary
tooth (12x5.5x3.5 mm), anterior dentary tooth (?x5.2x2.8 mm), posterior dentary
tooth (7x5.2x2.6 mm), fragments
Referred- ?(AMNH coll.) tooth (Bell, 2007)
(RTMP 65.16.150) tooth (7.2x4.7x2.6 mm) (Larson and Currie, 2013)
(RTMP 84.64.3) tooth (5.5x4.3x1.9 mm) (Larson and Currie, 2013)
(RTMP 84.64.12) tooth (7.3x4.5x2.3 mm) (Larson and Currie, 2013)
(RTMP 84.79.4) tooth (8x5.6x2.6 mm) (Larson and Currie, 2013)
(RTMP 85.12.2) tooth (5.1x3.2x1.7 mm) (Larson and Currie, 2013)
(RTMP 85.98.2) tooth (9.5x5.5x2.6 mm) (Larson and Currie, 2013)
(RTMP 86.64.2) tooth (4.1x3.2x1.6 mm) (Larson and Currie, 2013)
(RTMP 86.64.3) tooth (9.6x5.1x2.3 mm) (Larson and Currie, 2013)
(RTMP 90.82.17) tooth (7.6x4.7x2.4 mm) (Larson and Currie, 2013)
(RTMP 90.82.18) tooth (9.1x5.8x2.3 mm) (Larson and Currie, 2013)
(RTMP 90.82.21) tooth (6.6x4.7x2.4 mm) (Larson and Currie, 2013)
(RTMP 90.82.108) tooth (9.4x5.2x2.8 mm) (Larson and Currie, 2013)
(RTMP 93.12.21) tooth (7.7x4.4x2.5 mm) (Larson and Currie, 2013)
?(RTMP 97.39.4) tooth (Ryan et al., 2000)
(RTMP 98.63.4) tooth (Larson et al., 2010)
(RTMP 99.50.117) tooth (6.4x4.1x1.3 mm) (Larson et al., 2010)
?(RTMP 1005-1008, 1033, 1034) six teeth (Baszio, 1997)
?(RTMP 1009) premaxillary tooth (Baszio, 1997)
(RTMP 2000.45.35) tooth (5.8x3.9x2.1 mm) (Eberth and Currie, 2010)
(RTMP 2000.45.40) tooth (Eberth and Currie, 2010)
(RTMP 2000.45.103) tooth (10.2x4.3x2.1 mm) (Larson et al., 2010)
(RTMP 2003.45.49) tooth (6.6x4.2x2.2 mm) (Larson et al., 2010)
(RTMP 2003.45.52) tooth (7.1x4.7x? mm) (Larson et al., 2010)
(RTMP 2003.45.60) tooth (Larson et al., 2010)
(RTMP 2005.7.3) tooth (8.4x5.2x2.4 mm) (Larson and Currie, 2013)
?(RTMP coll.) seven teeth (Ryan et al., 2000)
(RTMP or UALVP coll.) (unassociated) five teeth (one uncollected) (Eberth and
Currie, 2010)
?(RTMP or UALVP coll.) (unassociated) manual phalanx, manual ungual (Eberth
and Currie, 2010)
?(RTMP coll.; lost) tooth (Torices et al., 2014)
?(UALVP 52075) manual ungual III (Eberth and Currie, 2010)
?(UALVP 55571) tooth (Torices et al., 2014)
? several premaxillary teeth (Baszio, 1997)
Diagnosis- (after Currie and Varricchio, 2004) snout short and deep;
premaxilla taller than long (also in Deinonychus); subnarial and nasal
processes of premaxilla inclined more than 45 degrees dorsally; large maxillary
fenestra; maxillary fenestra directly dorsal to promaxillary fenestra; maxillary
teeth isodont.
(after Larson et al., 2010) differs from Saurornitholestes langstoni
in- generally larger basal width and serration size; lower crown height compared
to FABL.
Other diagnoses- Currie and Varricchio (2004) also listed highly posteriorly
inclined maxillary teeth in their diagnosis, supposedly shared with Bambiraptor
and Deinonychus, but Senter (2010) showed this was due to taphonomy.
Comments- Discovered in 1995, Atrociraptor was announced in an
abstract by Currie and Varricchio (2000), then named and fully described by
Currie and Varricchio (2004). Powers et al. (2022) describes the specimen in more detail using CT scanning.
Baszio (1997) noted Horseshoe Canyon teeth he referred to Saurornitholestes
sp. differ from Dinosaur Park specimens in being smaller with less variable
serration shape. Ryan et al. (1997) listed seven teeth from this formation as
Saurornitholestes sp.. Ryan et al. (2000) later described these teeth
as cf. Saurornitholestes sp., as they noted the then unnamed Atrociraptor
had teeth nearly identical to Saurornitholestes'. Larson and Currie (2013)
found that Atrociraptor teeth only partially overlapped Saurornitholestes
teeth in morphometric analyses. These teeth are all provisionally referred to
Atrociraptor here, as no definite Saurornitholestes specimens
have been found in this formation. Eberth and Currie (2010) refer three manual
elements to Atrociraptor, but these cannot be compared to the type material
and may belong to the dromaeosaurine or Richardoestesia also present
in the bonebed, if not the troodontids.
RTMP 1009 was figured by Baszio as a Richardoestesia Paronychodon-like
tooth. It has three faint ridges, is moderately recurved, and has a high DSDI
(10 mesial serrations per mm vs. 5.7 distal serrations per mm). Distal serrations
are taller than apicobasally wide and rounded, while mesial serrations are shorter
than wide. Though the ridges and distal serration shape are similar to Saurornitholestes premaxillary teeth,
it differs in the curvature, fewer and weaker ridges, small serrations, and high DSDI. The basal constriction,
serration size, DSDI and curvature are similar to Richardoestesia gilmorei,
but the ridges and serration shape differ. It may be a posterior
premaxillary tooth, which have fewer and weaker grooves than anterior
teeth in Saurornitholestes.
However, Currie and Evans (2020) find in the holotype's second
premaxillary tooth "there do not appear to be any apicobasal
longitudinal ridges and flutes. This could be a function of wear and/or
poor preservation, however." Whether this means Saurornitholestes and Atrociraptor coexisted in the Horseshoe Canyon Formation, Atrociraptor exhibited variation in dental ridges, or taphonomy is responsible is uncertain.
References- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan, Currie, Gardner and Livigne, 1997. Baby hadrosaurid material associated
with an unusually high abundance of troodontid teeth from the Horseshoe Canyon
Formation (Early Maastrichtian), Alberta, Canada. Journal of Vertebrate Paleontology.
17(3), 72A.
Currie and Varricchio, 2000. New dromaeosaurids from the Late
Cretaceous of western North America. The Florida Symposium on Dinosaur
Bird Evolution. Publications in Paleontology No.2, Graves Museum of
Archaeology and Natural History. 14.
Ryan, Currie, Gardner, Vickaryous and Lavigne, 2000. Baby hadrosaurid material
associated with an unusually high abundance of Troodon teeth from the
Horseshoe Canyon Formation, Upper Cretaceous, Alberta, Canada. Gaia. 15, 123-133.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation
(Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright
(eds). Feathered Dragons. Studies on the transition from dinosaurs to birds.
Indiana University Press. 112-132.
Bell, 2007. The Danek bonebed: An unusual dinosaur assemblage from the Horseshoe
Canyon Formation, Edmonton, Alberta. Journal of Vertebrate Paleontology. 27(3),
46A.
Eberth and Currie, 2010. Stratigraphy, sedimentology, and taphonomy of the Albertosaurus
bonebed (upper Horseshoe Canyon Formation; Maastrichtian), southern Alberta,
Canada. Canadian Journal of Earth Sciences. 47(9), 1119-1143.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe
Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta,
with special reference to the Albertosaurus sarcophagus bonebed. Canadian
Journal of Earth Sciences. 47(9), 1159-1181.
Senter, 2010. Using creation science to demonstrate evolution: Application of
a creationist method for visualizing gaps in the fossil record to a phylogenetic
study of coelurosaurian dinosaurs. Journal of Evolutionary Biology. 23(8), 1732-1743.
Larson and Currie, 2013. Multivariate analyses of small theropod dinosaur teeth
and implications for paleoecological turnover through time. PloS ONE. 8(1),
e54329.
Torices, Funston, Kraichy and Currie, 2014. The first appearance of Troodon
in the Upper Cretaceous site of Danek Bonebed, and a reevaluation of troodontid
quantitative tooth morphotypes. Canadian Journal of Earth Sciences. 51, 1039-1044.
Currie and Evans, 2020 (online 2019). Cranial anatomy of new specimens of Saurornitholestes langstoni
(Dinosauria, Theropoda, Dromaeosauridae) from the Dinosaur Park
Formation (Campanian) of Alberta. The Anatomical Record. 303(4),
691-715.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as
2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist
in testing and constructing morphological characters. Journal of
Vertebrate Paleontology. 41(5), 2010087.
Saurornitholestiinae Longrich and Currie, 2009
Definition- (Saurornitholestes langstoni <- Dromaeosaurus albertensis,
Velociraptor mongoliensis, Microraptor zhaoianus) (Longrich and Currie,
2009)
Saurornitholestes Sues, 1978
= Zapsalis Cope, 1876b
S. langstoni Sues, 1978
= Laelaps explanatus Cope, 1876a
= Laelaps laevifrons Cope, 1876b
= Zapsalis abradens Cope, 1876b
= Dryptosaurus explanatus (Cope, 1876a) Hay, 1902
= Dryptosaurus laevifrons (Cope, 1876b) Hay, 1902
= Deinodon explanatus (Cope, 1876a) Lambe, 1902
= Deinodon laevifrons (Cope, 1876b) Osborn, 1902
= Aublysodon explanatus (Cope, 1876a) Hatcher, 1903
= Dromaeosaurus laevifrons (Cope, 1876b) Matthew and Brown, 1922
= Dromaeosaurus explanatus (Cope, 1876a) Kuhn, 1939
= Dromaeosaurus abradens (Cope, 1876b) Matthew and Brown, 1922 vide Kuhn, 1939
= Velociraptor langstoni (Sues, 1978) Paul, 1988
= Paronychodon explanatus (Cope, 1876a) Olshevsky, 1995
= Saurornitholestes explanatus (Cope, 1876a) Martyniuk, 2012
Diagnosis- (after Sullivan, 2006) ratio of the length (measured along
the midline) to the thickness (posterior part of the frontal) is 10:1.
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
Holotype- (RTMP 74.10.5) maxillary tooth (9.2 mm), lacrimal, jugal, postorbital, frontals,
quadrate, ectopterygoid, angular, anterior dentary tooth (8.9 mm), two vertebrae, dorsal rib fragments,
gastralia, caudal prezygopophyses, distal phalanx I-1, manual ungual I, distal
phalanx II-2, manual ungual II, distal metacarpal III, phalanx III-1 (17 mm),
distal phalanx III-3, manual ungual III
Paratypes- (CMN 12343) frontal
(CMN 12354) frontal
(UA 5283) posterior frontal
Referred- (CMN 2664) second or third premaxillary tooth (Currie, Rigby
and Sloan, 1990)
(ROM 57164) incomplete frontal (Currie and Evans, 2020)
(RTMP 67.20.36) last dorsal vertebra, sacrum (Norell and Makovicky, 1997)
(RTMP 70.37.1) first premaxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 78.9.96) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 79.8.643) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 79.15.3) second premaxillary tooth (11.8x6.4x? mm) (Currie, Rigby and Sloan, 1990)
(RTMP 80.16.312) frontal (Currie, 1987)
(RTMP 80.16.833) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 80.16.996) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.14.34) cervical vertebra (Rauhut, 2003)
(RTMP 81.20.259) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.23.7) frontal (Currie, 1987)
(RTMP 81.26.37) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.14.45) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.16.43) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.7) premaxillary tooth (15 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.180) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.336) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.19.458) premaxillary tooth (12.5 mm), tooth (13 mm) (Sankey, Brinkman, Guenther
and Currie, 2002)
(RTMP 82.24.16) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 82.26.1) ribs, incomplete caudal series, distal tarsal,
metatarsal II, pedal ungual II (distal caudal vertebra 31.1 mm)
(Currie, 1995; Makovicky, 1995)
(RTMP 82.112.10) tooth (8.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.91.39) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 84.91.40) premaxillary tooth (12.5 mm), tooth (12 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 84.94.6) frontal (Currie, 1987)
(RTMP 84.163.80) premaxillary tooth (10 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.2) premaxillary tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.131) premaxillary tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.132) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.6.133) premaxillary tooth (~11.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.43.69) frontal (Currie, 1987)
(RTMP 85.56.48) (adult) posterior cervical vertebra (Makovicky, 1995)
(RTMP 85.58.65) premaxillary tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.9.92) premaxillary tooth (~14.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.23.104) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.36.117) incomplete premaxilla (Currie, Rigby and Sloan, 1990)
(RTMP 86.36.425) premaxillary tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
?(RTMP 86.77.2) ischium (Hutchinson, 2001)
(RTMP 86.77.57) frontal (Currie, 1987)
(RTMP 86.77.112) premaxillary tooth (~14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.78.1) basioccipital (Rauhut, 2003)
(RTMP 86.130.219) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.198.46) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.4.47) tooth (4.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.19.68) tooth (4.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.31.14) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.31.52) tooth (12.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.31.54) tooth (4.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.33.55) tooth (9.9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.5) premaxillary tooth (15.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.11) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.68) tooth (~11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.70) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.184) tooth (7.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.300) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.383) premaxillary tooth (Baszio, 1997)
(RTMP 87.36.392) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.36.418) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.39.93) tooth (11.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.43.5) tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.46.53) tooth (9.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.48.77) tooth (6.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.48.86) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.50.8) second premaxillary tooth (13.3x6.8x? mm) (Currie and Evans, 2020)
(RTMP 87.50.38) tooth (14.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.50.100) tooth (8.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.51.23) tooth (8.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.62.87) tooth ( mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.72.4) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.72.23) tooth (~9.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.72.26) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.120) tooth (~11.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.125) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.126) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.79.89) tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.79.90) tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.83.1) tooth (12.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.112.9) tooth (8.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.153.12) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.153.55) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.62) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.63) tooth (8.1 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.158.78) tooth (4.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.158.80) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.158.81) tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.112.21) tooth (5.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.112.28) tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.154.64) tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.50.123) tooth (Baszio, 1997)
(RTMP 88.50.126) tooth (Baszio, 1997)
(RTMP 88.87.89) tooth (Baszio, 1997)
(RTMP 88.116.42) tooth (Baszio, 1997)
(RTMP 88.121.39) (128 kg) quadratojugal, quadrate, mandible
(dentary 120 mm), teeth (6.2-12.3 mm), three dorsal ribs, gastralia, distal caudal
series, scapula (~155 mm), coracoid (~65 mm prox-dist), humerus (~177 mm), femur
(214 mm), tibia (283 mm), fibula, astragalus, calcaneum, metatarsal II (101 mm), phalanx II-2 (31 mm),
pedal ungual II (69 mm straight), metatarsal III (115 mm), metatarsal IV (107
mm) (Currie, Rigby and Sloan, 1990; Currie and Varricchio in prep.)
(RTMP 88.162.1) tooth (Baszio, 1997)
(RTMP 88.215.72) tooth (Baszio, 1997)
(RTMP 89.36.312) premaxillary tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.353) tooth (Baszio, 1997)
(RTMP 89.36.400) tooth (Baszio, 1997)
(RTMP 89.36.403) tooth (Baszio, 1997)
(RTMP 89.36.410) tooth (Baszio, 1997)
(RTMP 89.50.8) second premaxillary tooth (13.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.50.202) premaxillary tooth (9.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.6.15) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.50.208) premaxillary tooth (~11.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 90.53.21) premaxillary tooth (~11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 91.36.112) dentary, tooth (Rauhut, 2003)
(RTMP 91.50.60) premaxillary tooth (14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 92.36.129) sacrum (Rauhut, 2003)
(RTMP 92.36.333) sternal plate (Godfrey and Currie, 2004)
(RTMP 92.50.23) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 92.83.2) tooth (Currie and Jacobsen, 1995)
(RTMP 93.36.98) tarsal (Rauhut, 2003)
(RTMP 94.12.268) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.844 maxilla (Currie and Varricchio, 2004)
(RTMP 95.2.18) tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.31) tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.33) tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.36) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.38) tooth (11 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.40) tooth (7.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.42) tooth (7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.43) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.12.74) tooth (10.2 mm), tooth (6.3 mm), tooth (9.4 mm) (Sankey, Brinkman,
Guenther and Currie, 2002)
(RTMP 95.12.109) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.19.4) tooth (8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.21.5) tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.16) tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.25) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.27) tooth (6.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.28) tooth (~5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.92.54) tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.124.4) tooth (6.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.126.29) tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.127.25) tooth (11.8 mm), tooth (~7.8 mm), tooth (6.7 mm), tooth (7.5
mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.129.2) tooth (9.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.131.12) tooth (9.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.137.2) tooth (9.2 mm), tooth (10.5 mm) (Sankey, Brinkman, Guenther
and Currie, 2002)
(RTMP 95.147.26) tooth (2.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.151.10) tooth (3.9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.177.48) tooth (6 mm), tooth (4.7 mm), tooth (~3.4 mm) (Sankey, Brinkman,
Guenther and Currie, 2002)
(RTMP 95.179.3) tooth (10.4 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.180.4) tooth (~3.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.181.11) tooth (~5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.182.21) tooth (~7.6 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.184.23) tooth (8.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.406.5) tooth (10.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.34) tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.38) tooth (11.7 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.102) tooth (11.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.104) tooth (10.8 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.112) tooth (16 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.115) tooth (14 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.118) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.359) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.360) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.361) tooth (13 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.363) tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.364) tooth (9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.366) tooth (6.3 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 96.12.422) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 97.80.36) tooth (3.9 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 97.133.2) premaxillary tooth (12 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP coll.; lost) premaxillary tooth (Baszio, 1997)
(UALVP 55425) surangular, articular (Currie and Evans, 2020)
(UALVP 55700) (~2 m, 8 year old subadult) skull (~210 mm), sclerotic
plates, mandibles, hyoids, axis, third cervical vertebra, fourth-tenth
cervical vertebrae, cervical ribs, dorsal series, dorsal ribs, uncinate
processes, gastralia, sacrum, incomplete caudal series, chevrons,
scapulae, coracoids, furcula, sternum, sternal ribs, forelimbs missing
a few phalanges, pelves, hindlimbs missing a few phalanges (femur ~228
mm), pedal ungual II claw sheath (Currie and Evans, 2015)
thirty premaxillary teeth (Baszio, 1997)
Late Campanian, Late Cretaceous
Oldman or Dinosaur Park Formation of the Belly River Group, Alberta, Canada
(NHMUK R4463) sacrum (Howse and Milner, 1993)
tooth (12 mm) (Lambe, 1902)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
?(CMN 12071; = CMN 12072 in Colbert and Russell?) metatarsal IV (Colbert and Russell, 1969)
?(CMN 12240) pedal ungual II (Colbert and Russell, 1969)
(CMN 12410) tooth (CMN online)
(RTMP 94.144.105) tooth (Ryan and Russell, 2001)
(UA 12091) dentary (Sues, 1977)
(UA 12339) dentary (Sues, 1977)
Middle Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
(RTMP 88.86.29) tooth (Ryan and Russell, 2001)
(RTMP 96.62 coll.) twenty-seven teeth (Peng, Russell and Brinkman, 2001)
(RTMP 2009.37.109) tooth (Cullen, Fanti, Capobianco, Ryan and Evans, 2016)
(RTMP coll.) three teeth (Cullen, Fanti, Capobianco, Ryan and Evans, 2016)
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
(AMNH 3953; holotype of Zapsalis abradens) second premaxillary tooth (12 mm) (Cope, 1876b)
(AMNH 3958; holotype of Laelaps explanatus) tooth (11 mm), twenty-five
teeth (Cope, 1876a)
(AMNH 3961; holotype of Laelaps laevifrons) tooth (15 mm) (Cope, 1876b)
(AMNH 8518) tooth (Sahni, 1972)
(ANSP 15808) tooth (Fiorillo and Currie, 1994)
(ANSP 15810) tooth (Fiorillo and Currie, 1994)
(ANSP 15811) tooth (Fiorillo and Currie, 1994)
(ANSP 15814) tooth (Fiorillo and Currie, 1994)
(ANSP 15815) tooth (Fiorillo and Currie, 1994)
(ANSP 15817) tooth (Fiorillo and Currie, 1994)
(ANSP 15818) tooth (Fiorillo and Currie, 1994)
(ANSP 15819) tooth (Fiorillo and Currie, 1994)
(ANSP 15823) tooth (Fiorillo and Currie, 1994)
(ANSP 15824) tooth (Fiorillo and Currie, 1994)
(ANSP 15825) tooth (Fiorillo and Currie, 1994)
(ANSP 15826) tooth (Fiorillo and Currie, 1994)
(ANSP 15828) tooth (Fiorillo and Currie, 1994)
(ANSP 15829) tooth (Fiorillo and Currie, 1994)
(ANSP 15831) tooth (Fiorillo and Currie, 1994)
(ANSP 15833) tooth (Fiorillo and Currie, 1994)
(ANSP 15838) tooth (Fiorillo and Currie, 1994)
(ANSP 15939) tooth (Fiorillo and Currie, 1994)
(ANSP 15943) tooth (Fiorillo and Currie, 1994)
(ANSP 15945) tooth (Fiorillo and Currie, 1994)
(ANSP 15946) tooth (Fiorillo and Currie, 1994)
(ANSP 15948) tooth (Fiorillo and Currie, 1994)
(ANSP 15949) tooth (Fiorillo and Currie, 1994)
(ANSP 15951) tooth (Fiorillo and Currie, 1994)
(ANSP 15953) tooth (Fiorillo and Currie, 1994)
(ANSP 15954) tooth (Fiorillo and Currie, 1994)
(ANSP 15955) tooth (Fiorillo and Currie, 1994)
(ANSP 15956) tooth (Fiorillo and Currie, 1994)
(ANSP 15957) tooth (Fiorillo and Currie, 1994)
(ANSP 15958) tooth (Fiorillo and Currie, 1994)
(ANSP 15960) tooth (Fiorillo and Currie, 1994)
(ANSP 15975) tooth (Fiorillo and Currie, 1994)
(ANSP 16195) tooth (Fiorillo and Currie, 1994)
(ANSP 16197) tooth (Fiorillo and Currie, 1994)
(ANSP 16198) tooth (Fiorillo and Currie, 1994)
(ANSP 16199) tooth (Fiorillo and Currie, 1994)
(ANSP 16225) tooth (Fiorillo and Currie, 1994)
(ANSP 16226) tooth (Fiorillo and Currie, 1994)
(ANSP 16227) tooth (Fiorillo and Currie, 1994)
(ANSP 16228) tooth (Fiorillo and Currie, 1994)
(ANSP 16229) tooth (Fiorillo and Currie, 1994)
(ANSP 17646) tooth (Fiorillo and Currie, 1994)
(ANSP 17648) tooth (Fiorillo and Currie, 1994)
(ANSP 17649) tooth (Fiorillo and Currie, 1994)
(ANSP 17773) tooth (Fiorillo and Currie, 1994)
(ANSP 17774) tooth (Fiorillo and Currie, 1994)
(ANSP 17775) tooth (Fiorillo and Currie, 1994)
(ANSP 17776) tooth (Fiorillo and Currie, 1994)
(ANSP 17777) tooth (Fiorillo and Currie, 1994)
(ANSP 17778) tooth (Fiorillo and Currie, 1994)
(ANSP 17781) tooth (Fiorillo and Currie, 1994)
(ANSP 17783) tooth (Fiorillo and Currie, 1994)
(ANSP 17785) tooth (Fiorillo and Currie, 1994)
(ANSP 17786) tooth (Fiorillo and Currie, 1994)
(ANSP 17787) tooth (Fiorillo and Currie, 1994)
(ANSP 17788) tooth (Fiorillo and Currie, 1994)
(ANSP 17789) tooth (Fiorillo and Currie, 1994)
(ANSP 17801) tooth (Fiorillo and Currie, 1994)
(ANSP 17802) tooth (Fiorillo and Currie, 1994)
(ANSP 17803) tooth (Fiorillo and Currie, 1994)
(ANSP 17962) tooth (Fiorillo and Currie, 1994)
(ANSP 17963) tooth (Fiorillo and Currie, 1994)
(ANSP 18002) tooth (Fiorillo and Currie, 1994)
(ANSP 18006) tooth (Fiorillo and Currie, 1994)
(ANSP 18100) tooth (Fiorillo and Currie, 1994)
(ANSP 18103) tooth (Fiorillo and Currie, 1994)
(ANSP 18105) tooth (Fiorillo and Currie, 1994)
(ANSP 18109) tooth (Fiorillo and Currie, 1994)
(ANSP 18112) tooth (Fiorillo and Currie, 1994)
(ANSP 18118) tooth (Fiorillo and Currie, 1994)
(ANSP 18122) tooth (Fiorillo and Currie, 1994)
(BDM 005) frontal (Wilson and Fowler, 2020)
(MOR 125) phalanx (MOR online)
(UCMP 137559) tooth (UCMP online)
(UCMP 137560) tooth (UCMP online)
(UCMP 150584) tooth (UCMP online)
(UCMP 150585) teeth (UCMP online)
(UCMP 150586) teeth (UCMP online)
(UCMP 150587) teeth (UCMP online)
(UCMP 150588) ungual fragment (UCMP online)
(YPM PU 24968) (YPM online)
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
(MOR 484) ungual (MOR online)
(MOR 553E-6-26-91-104) metatarsal IV (MOR online)
(MOR 553S-7-10-9-197) pedal phalanx IV-4 (MOR online)
(MOR 660) (19.5 kg) partial skeleton including nine cervical vertebrae, thirteen
dorsal vertebrae, gastralia, sacrum, first caudal vertebra, distal caudal vertebrae,
chevrons, humerus (178 mm), radius, ulna (135 mm), metacarpal I (30 mm), phalanx
I-1 (61 mm), manual ungual I (55 straight), phalanx II-1 (49 mm), phalanx II-2
(43 mm), manual ungual II (35 mm), metacarpal III (67 mm), phalanx III-1 (13
mm), phalanx III-2 (29 mm), phalanx III-3 (40 mm), ilium, femur (225 mm), tibia
(257 or 301 mm), metatarsal II (112 mm), phalanx II-1 (32 mm), phalanx II-2
(30 mm), pedal ungual II (74 mm straight), metatarsal III (136 mm) and metatarsal
IV (123 mm), (Britt, 1993)
(MOR 666) tibia (256 mm), fibula (225 mm) (Christiansen, 1999)
(MOR 9753) (juvenile) maxilla (55 mm) (Hohman and Varricchio, 2021)
teeth (Redman, Moore and Varricchio, 2015)
Diagnosis-
(after Currie and Evans, 2020) relatively short skull (skull/femur
length ratio less than 1.0); large maxillary fenestra that is
positioned near the dorsal margin of the antorbital fossa; maxillary
sinuses that interconnect with those inside the nasal and lacrimal;
enlarged second premaxillary tooth has a flat lingual surface and has
longitudinal ridges and grooves on both labial and lingual surfaces.
Comments- Unfortunately, besides the original description of the fragmentary
holotype (Sues, 1978), Currie et al.'s (1990) description of teeth (see also
Sankey et al., 2002), Makovicky's (1995) as yet unpublished description
of vertebrae, and very recent cranial description of Currie and Evans (2020), Saurornitholestes has not received a detailed description
and much information must be pieced together from sporadic references throughout
the literature. Currie and Varricchio (2004) note both RTMP 88.121.39 and MOR
660 are being studied by them, which had been occurring for over a decade as
of that date.
Saurornitholestes was historically assigned to Velociraptorinae, but
was recently given its own subfamily outside the Velociraptorinae+Dromaeosaurinae
clade (Longrich and Currie, 2009; Currie and Evans, 2020). Brusatte et al. (2014) still recovered it
in Velociraptorinae however, and here it is placed in Dromaeosaurinae based
on Hartman et al. (2019).
Laelaps explanatus- Cope (1876a) described this tooth as a new
species of Laelaps (in which he placed all Judithian theropods). Once
Marsh provided the replacement name Dryptosaurus for the preoccupied
Laelaps, Hay (1902) moved explanatus to that genus. Lambe (1902)
meanwhile referred it to Deinodon, while Hatcher referred it to Aublysodon,
both contemporary tyrannosaurids. Matthew and Brown (1922) questionably referred
the species to their new genus Dromaeosaurus as Dromaeosaurus(?)
sp., probably based on size, for it was the smallest of their 'deinodontids'.
Kuhn (1939) formalized this as D. explanatus. Ostrom (1969) noted these
have a high DSDI, and assigned the species to Dromaeosauridae. Olshevsky's (1995)
referral of the species to Paronychodon is almost certainly incorrect,
as it is serrated and lacks grooves.
Of Judith River theropods, only Saurornitholestes and Richardoestesia
have a comparable DSDI. However, Richardoestesia is smaller (FABL up
to 4.7 vs. 6.6 mm) and has more serrations per mm (6+ vs. 4.5). The compression
(BW/FABL = 2.8/6.6 mm = .42) is comparable to Saurornitholestes, and
while the measured tooth is lower than most sampled by Sankey et al. (2002),
it is still within the range of variation (lower 5%). The short mesial carina,
which is only serrated apically, is also common in dromaeosaurids and differs
from Dromaeosaurus in lacking a lingual twist. As Saurornitholestes
langstoni is common in the equivalent Dinosaur Park Formation further north,
explanatus is here referred to that species. It should be noted that
the name explanatus has over a century of priority over langstoni
though, so would technically have precedence as a senior synonym. This is not
followed here because from the limited description, explanatus is indistinguishable
from not only Saurornitholestes, but Bambiraptor and Velociraptor
as well. It is only referred to Saurornitholestes due to provenance.
Lambe (1902) referred another tooth to this species, from the Belly River Group
of Alberta. It is also said to have a high DSDI and seems to be Saurornitholestes
as well. It has a FABL of 6 mm and a BW of 3 mm.
Laelaps laevifrons- Cope (1876b) described this tooth as a new
species of Laelaps (in which he placed all Judithian theropods). Once
Marsh provided the replacement name Dryptosaurus for the preoccupied
Laelaps, Hay (1902) moved laevifrons to that genus. Osborn (1902)
meanwhile referred it to Deinodon. Matthew and Brown (1922) referred
the species to their new genus Dromaeosaurus, probably based on size,
for it was the smallest of their 'deinodontids'.
Of Judith River theropod taxa, Zapsalis and Paronychodon can be
eliminated due to the lack of grooves, while Richardoestesia can be eliminated
due to the low serration count and troodontids due to the high serration count.
This leaves tyrannosaurids, Dromaeosaurus and Saurornitholestes
as potential candidates. Tooth size is within the range of juvenile tyrannosaurids,
in the upper 10% of Dromaeosaurus, and the upper 1% of Saurornitholestes.
The lack of mesial serrations is known in Saurornitholestes and juvenile
tyrannosaurids, but not Dromaeosaurus. Crown compression (BW of 4 mm
and FABL of 7 mm) is comparable to Dromaeosaurus and tyrannosaurids when
size is taken into account, though hypothetically possible for a Saurornitholestes
that size. Crown elongation is similar to all three taxa when compared to FABL.
Crown elongation vs. crown compression is however withion the range of Saurornitholestes
and Dromaeosaurus, but not tyrannosaurids (which usually have stouter
crowns when they are that thick). The serration density (5/mm) compared to crown
compression is comparable to tyrannosaurids, and in the outer range of Dromaeosaurus
and Saurornitholestes. Serration density compared to crown elongation
is comparable to Saurornitholestes, and outside either Dromaeosaurus
or tyrannosaurids. When all components are analyzed together, laevifrons
falls out within Saurornitholestes. It is here viewed as a particularily
large example of that genus. As Saurornitholestes langstoni is common
in the equivalent Dinosaur Park Formation further north, laevifrons is
here referred to that species. It should be noted that the name laevifrons
has over a century of priority over langstoni though, so would technically
have precedence as a senior synonym (though explanatus has even more
priority). This is not followed here because from the limited description, laevifrons
is indistinguishable from not only Saurornitholestes, but Bambiraptor
and Velociraptor as well. It is only referred to Saurornitholestes
due to provenance.
Zapsalis-
The holotype of Zapsalis abradens (AMNH 3953) is a tooth which is flat lingually,
with no mesial serrations and 3 distal serrations per mm (Cope, 1876b). There
are three lingual ridges and four labial ones. The length is 12 mm, the FABL
6.5 mm, and the BW 3 mm. It was synonymized with Paronychodon lacustris
by Estes (1964) based on its flat lingual side and ridges, which was followed
in the literature for decades. Currie et al. (1990) later restricted Paronychodon to the unserrated
forms, and believed the serrated ones were growth abnormalities of named genera-
Dromaeosaurus, Saurornitholestes and Troodon. Baszio (1997)
included these in his Paronychodon discussion, referred to them as Paronychodon-like
teeth. Sankey et al. (2002) separated the Paronychodon-like teeth with
Dromaeosaurus-like and Saurornitholestes-like serrations as ?Dromaeosaurus
morphotype A. In addition to their ridges and flat lingual side, they differ
from Dromaeosaurus albertensis in being shorter and labiolingually narrower,
and having a straight distal edge. They argued complete D. albertensis
tooth rows are known, so the morphology of morphotype A teeth is not due to
positional variation. Furthermore, they have a different stratigraphic distribution
than D. albertensis, being present in the Late Maastrichtian unlike the
latter species. In 2007, Mortimer (DML, 2007) noticed that Zapsalis matched ?Dromaeosaurus
morphotype A, although which kind of dromaeosaurid it was was still a
mystery. The question was solved by the description of a complete
Saurornitholestes langstoni skull UALVP 55700 (Currie and Evans, 2020) which featured a tooth nearly identical to Zapsalis' holotype in the second premaxillary position. Its other premaxillary teeth are also ridged, showing Zapsalis consists of premaxillary teeth of Saurornitholestes or closely related taxa. Currie and Evans proposed Zapsalis' holotype differed from Saurornitholestes langstoni
(RTMP 79.15.3 and UALVP 55700 at least) in lacking mesial serrations
(seemingly not taphonomic) and being concave apicodistally, and thus
"recommended that the two genera be kept separate." Given the
large amount of dental variation within theropod species and
correlation between their formations, this seems flimsy unless more
Judith River teeth are discovered that share those characters with AMNH
3953. Indeed, Baszio figures two Dinosaur Park teeth with convex
apicodistal margins and no mesial serrations (RTMP 87.36.383, RTMP
coll.), blending the morphotypes. It would also lead to such a
narrow concept of Saurornitholestes
that even the Two Medicine material couldn't justifiably be
referred. One motivation for keeping the genera separate is the
obvious priority Zapsalis abradens has over Saurornitholestes langstoni, and unlike explanatus or laevifrons, it can be distinguished from all other named taxa (even Atrociraptor and Bambiraptor). Yet it does not seem distinguishable from 'Zapsalis' teeth from the later Scollard and Lance Formations (Baszio, 1997: pl. VI, fig. 87, 88), suggesting Zapsalis' holotype is indeterminate at the species level. While one could argue for a genus Zapsalis containing indeterminate Z. abradens, and diagnostic 'Z.langstoni' and 'Z. sullivani', a petition to the ICZN to suppress Zapsalis abradens would cause far less disruption.
Early finds- Colbert and Russell (1969) tentatively referred a fourth
metatarsal (as CMN 12072) and a pedal ungual II (CMN 12240) to Dromaeosaurus,
as did Ostrom (1969) for the latter element. Paul (1988) referred the ungual
to Saurornitholestes instead, as he hypothesized Dromaeosaurus
to have a reduced ungual as in Adasaurus. Both elements were referred
to Saurornitholestes by Currie (1995), but as those bones remain undescribed
for known Saurornitholestes specimens (and Dromaeosaurus may not
be closer to Adasaurus), such referrals are only provisional. The CMN online catalogue lists 12072 as a cast of the Dromaeosaurus holotype, but 12071 as a real Dromaeosaurus fossil, so 12072 may have been a typo.
Russell (1969) listed CMN 12343, 12354 (both found in 1968) and UA 5283 as frontals different from
Dromaeosaurus, Troodon and a therizinosaur frontal (CMN 12355),
but these were later made paratypes of Saurornitholestes when that genus
was described.
Sahni (1972) identified AMNH 8518 as Troodon, but the large DSDI matches
Saurornitholestes better. Howse and Milner (1993) identified NHMUK R4463
as a troodontid, but Makovicky (1995) recognized it as Saurornitholestes.
Sues (1977) described two dentaries (UA 12091 and 12339) discovered in 1969
and 1974 respectively. He referred these to Dromaeosaurus sp., but they
were later realized to be Saurornitholestes (Paul, 1988).
The holotype (RTMP 74.10.5) was discovered in 1974, and described by
Sues (1978) as a new genus of dromaeosaurid. Currie and Evans
(2020) report "the quarry was reopened in 2002 and additional cranial
material was recovered" including a lacrimal, jugal, postorbital and
angular described in that paper.
Recent discoveries- Howse and Milner (1993) illustrated a sacrum (NHMUK
R4463) and identified it as a troodontid, comparing it favorably to Ornithodesmus.
Makovicky (1995) reidentified it as Saurornitholestes, which was first
published publically in Norell and Makovicky (1997).
Makovicky (1995) describes the caudal vertebrae of RTMP 82.26.1 in depth. It
is also mentioned by Currie (1995, 2005).
RTMP 88.121.39 is a specimen discovered in 1988 and first mentioned by
Currie et al. (1990). The dorsal ribs were noted to be apneumatic by
Britt (1993). Makovicky (1995) described the elongate distal caudal
prezygapophyseal anatomy. Currie (1995) noted it has a T-shaped
quadratojugal, fifteen dentary teeth, fused interdental plates, a
posterior splenial notch, a collumnar articular process and elongated
distal caudal prezygapophyses. Christiansen (1999) lists measurements
for the femur and tibia. Xu et al. (1999) noted the glenoid faced
laterally and the coracoid was similar to other dromaeosaurids.
Jacobsen (2001) describes and illustrates the dentary, noting tooth
marks on the bone (these were previously mentioned by Tanke and Currie,
2000). He stated the specimen is under study by Currie and Varricchio.
Rauhut (2003) lists the scapular dimensions, and illustrates the
splenial, scapulocoracoid and fibula. Codd (2004) notes that there are
scars for uncinate processes on three preserved ribs. Jasinowski et al.
(2006) illustrate and describe the scapulocoracoid and humerus.
Brusatte et al. (2013) list various limb element lengths. Funston
and Currie (2018) figure the tibiotarsus in different views.
Currie and Evans (2020) describe and figure the cranial material.
MOR 660 is a partial skeleton discovered in 1990. Britt (1993) describes the
pneumaticity of MOR 660 and notes it is being studied by Currie and Varricchio.
Makovicky (1995) describes the presacral and sacral morphology in depth. Novas
(1998) mentions the ulna has a feeble olecranon process and lacks the proximoanterior
process for articulation with the radius. Currie and Dong (2001) note that the
anterior dorsals have very large hyapophyses. Hutchinson (2001) illustrates
the ilium, and notes it has a supratrochanteric process and a reduced but present
cuppedicus fossa. Rauhut (2003) notes it has a concave anterior pubic peduncle
edge, while he notes the posterior dorsal centra are subcircular, all dorsal
centra are pleurocoelous and the presence of five sacral vertebrae in his theropod
phylogeny study from that same year. He also illustrates manual ungual II. Claessens
(2004) studied the medial gastralia of the specimen. Organ and Adams (2005)
studied the histology of elongate prezygapophyses and chevrons from this specimen.
Novas and Pol (2005) note the radius has a triangular proximal articular surface
and the femur is robust. Martinez and Novas (2006) note the distal ulna has
an anteroposteriorly expanded articular surface. Turner et al. (2007) lists
the femoral length in their supplementary information table. Norell et al. (2006)
note the posterior cervicals lack carotid processes, but have a raised lip instead.
Csiki et al. (2010) list the hindlimb measurements. Brusatte et al. (2013) list
various limb element lengths.
Norell and Makovicky (1997) note the last dorsal centrum and sacral centra of
RTMP 67.20.36 are pleurocoelous.
Christiansen (1999) lists measurements for the tibia and fibula of MOR 666.
Xu et al. (1999) mention a sternal plate found in 1992 (RTMP 92.36.333) that
was later described in detail by Godfrey and Currie (2004).
Hutchinson (2001) identified an ischium (RTMP 86.77.2) as Troodon, but
it is more likely Saurornitholestes based on comparison to Bambiraptor
and Troodon specimen MOR 553L.
Rauhut (2003) mentions there are five sacral vertebrae in RTMP 92.36.129.
Rauhut (2004) lists RTMP 94.12.844 (a maxilla), but it was not described and
illustrated until Currie and Varricchio's (2004) paper. Powers et al. (2022) provide CT scans of the element.
Currie and Evans (2015) reported a new incomplete skeleton (UALVP 55700- McFeeters,
pers. comm. 2015) found on June 20 2014. It preserves uncinate processes, furcula, sternum and sternal
ribs, with a shorter and broader snout then Velociraptor
and a larger pedal ungual II. The skull was described in depth by
Currie and Evans (2020), who state the mandibular CT scans and
postcrania "will be described in a separate paper."
UALVP 55425 is a fused surangular and articular found on May 25 2009 (Currie and Evans, 2020).
Hohman and Varricchio (2021) described juvenile maxilla MOR 9753 found
in the 1990 from Bob's Vacation Site, which is smaller than Bambiraptor's holotype but shares characters with Saurornitholestes suggesting the former is not a juvenile of the latter.
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Currie, 1987. Theropods of the Judith River Formation. Occasional Paper of the
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Mart�nez and Novas, 2006. Aniksosaurus darwini gen. et sp. nov.,
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Mortimer, DML 2007. https://web.archive.org/web/20160806072151/http://dml.cmnh.org/2007Jul/msg00354.html
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Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
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Csiki, Vremir, Brusatte and Norell, 2010. An aberrant island-dwelling theropod
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the Upper Two Medicine Formation in the vicinity of Egg Mountain. Journal of
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S? sullivani Jasinski,
2015
Late Campanian, Late Cretaceous
De-na-zin Member of Kirtland Formation, New Mexico, US
Holotype- (SMP VP-1270) frontal (42.8 mm) (Sullivan and Lucas, 2000)
Referred- ?(NMMNH P-22578) tooth (8.4x4.4x1.9 mm) (Williamson and Brusatte,
2014)
?(NMMNH P-26201) tooth (Williamson and Brusatte, 2014)
?(NMMNH P-26205) tooth (8.7x4.9x2.2 mm) (Williamson and Brusatte, 2014)
?(NMMNH P-32752) tooth (8.9x4.4x2.4 mm) (Williamson and Brusatte, 2014)
?(NMMNH P-68395) tooth (4.2x3.7x2.1 mm) (Williamson and Brusatte, 2014)
?(SMP VP-1741) partial pedal ungual II (Sullivan, 2006)
?(SMP VP-1901) tooth (15 mm) (Sullivan, 2006)
Diagnosis- (after Jasinski, 2015) differs from S. langstoni in-
more constricted anteriorly; less prominent nasal facets; less prominent anterior
projection between the nasal and lacrimal facet regions; deeper and less strongly
demarcated orbital rim; deeper and more prominent olfactory bulb surface; more
pronounced and longer ventrally-directed ridge between the olfactory bulb surface
and the cerebral hemisphere surface; more robust frontal median sutural surface;
while slightly smaller, still being more robust.
Comments- Sullivan and Lucas (2000) described SMP VP-1270 as an individual
of Saurornitholestes langstoni. Later, Sullivan (2006) described another
frontal as the new species S. robustus, referring the original frontal
and two more elements to this taxon. Evans et al. (2014) later reidentified
S. robustus as a troodontid, though the other remains are still dromaeosaurid.
Williamson and Brusatte (2014) described more teeth as Dromaeosauridae morphotype
A, which they found fall within the range of variation of Dinosaur Park Saurornitholestes.
Jasinski (2015) used SMP VP-1270 as the holotype of his new species Saurornitholestes
sullivani, and kept other De-na-zin dromaeosaurid remains as indeterminate.
However, only the S. langstoni holotype was compared, not other specimens
of that species to check for variation which might explain sullivani's
differences. Furthermore, most other North American dromaeosaurids from the
Late Cretaceous (Acheroraptor, Atrociraptor)
don't preserve frontals, making proposed Saurornitholestes frontal synapomorphies
difficult to test.
References- Sullivan and Lucas, 2000. First occurrence of Saurornitholestes
(Theropoda: Dromaeosauridae) from the Upper Cretaceous of New Mexico. in Lucas
and Heckert (eds.). Dinosaurs of New Mexico. New Mexico Museum of Natural History
and Science, Bulletin 17. pp. 105-108.
Sullivan, 2006. Saurornitholestes robustus, n. sp. (Theropoda: Dromaeosauridae)
from the Upper Cretaceous Kirtland Formation (De-na-zin Member), San Juan Basin,
New Mexico. In Lucas and Sullivan (eds.). Late Cretaceous vertebrates from the
Western Interior. New Mexico Museum of Natural History and Science Bulletin.
35, 253-256.
Evans, Larson, Cullen and Sullivan, 2014. 'Saurornitholestes' robustus
is a troodontid (Dinosauria: Theropoda). Canadian Journal of Earth Sciences.
51(7), 730-734.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Jasinski, 2015.
S? sp. (Gangloff, 1998)
Campanian, Late Cretaceous
Prince Creek Formation, Alaska, US
(AK249-V-034) tooth (Fiorillo and Currie, 2000)
(AK249-V-035) tooth (Fiorillo and Currie, 2000)
vertebra (Gangloff, 1998)
References- Gangloff, 1998. Arctic Dinosaurs with Emphasis on the Cretaceous
Record of Alaska and the Eurasian-North American Connection. Kirkland, Lucas
and Estep (eds). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico
Museum of Natural History and Science, Bulletin. 14, 211-220.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation
(Cretaceous) of Northern Alaska, with speculations on Arctic dinosaur paleoecology.
Journal of Vertebrate Paleontology. 20(4), 675-682.
S? sp. nov. (Baszio, 1997)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
(CMN coll.) teeth (Russell, 1935)
(UA MR-4:1-3, 21-32) fifteen teeth (Baszio, 1997)
Comments- These teeth are highly variable, but have smaller serrations
than S. langstoni. Mesial serrations are usually absent, and the teeth
are only distinguishable from Richardoestesia based on serration count.
References- Russell, 1935. Fauna of the Upper Milk River beds, Southern
Alberta. Transactions of the Royal Society of Canada, series 3, section 4. 29,
115-127.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
S? sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Material- (RTMP 86.55.7) tooth (Bell and Currie, 2015)
(RTMP 86.55.9) tooth (Bell and Currie, 2015)
(RTMP 86.55.27) tooth (Bell and Currie, 2015)
(RTMP 86.55.38) tooth (Bell and Currie, 2015)
(RTMP 88.55.129; paratype of Boreonykus certekorum) incomplete distal
caudal vertebra (Bell and Currie, 2015)
(RTMP 87.55.148) tooth (Bell and Currie, 2015)
(RTMP 87.55.149) tooth (Bell and Currie, 2015)
(RTMP 86.55.184; paratype of Boreonykus certekorum) pedal ungual II (Bell
and Currie, 2015)
(RTMP 87.55.329) tooth (Bell and Currie, 2015)
(RTMP 87.55.333) tooth (Bell and Currie, 2015)
(RTMP 87.55.1523) tooth (Ryan and Russell, 2001)
(RTMP 89.55.707) tooth (Bell and Currie, 2015)
(RTMP 89.55.1005) tooth (Bell and Currie, 2015)
(RTMP 89.55.1141) tooth (Bell and Currie, 2015)
(RTMP 89.55.1162) tooth (Bell and Currie, 2015)
(RTMP 2004.23.4) tooth (Fanti and Miyashita, 2009)
(UALVP 53597; paratype of Boreonykus certekorum) incomplete manual ungual
II (Bell and Currie, 2015)
two teeth (Fanti and Miyashita, 2009)
Late Campanian-Early Maastrichtian, Late Cretaceous
Wapiti Formation, British Columbia, Canada
Material- (PRPRC coll.) tooth (McCrea and Buckley, 2004)
Comments- Ryan and Russell (2001) listed the frontal RTMP 89.55.47 as
"Saurornitholestes undescribed n. sp. (Currie pers. comm.)".
Bell and Currie (2015) described this and other material (distal caudal RTMP
88.55.129, manual ungual UALVP 53597, and pedal ungual II RTMP 86.55.184) from
the Pipestone Creek Pachyrhinosaurus lakustai bonebed as Boreonykus
certekorum, noting that preserved elements are not duplicated and are properly
proportioned to belong to one individual. The fourteen referred teeth are shed
so were admitted to at least belong to another individual. While the authors
assigned Boreonykus to Velociraptorinae, Cau (online, 2015) noted the
frontal differs from dromaeosaurids and that its association with the postcrania
cannot be substantiated. Boreonykus is here placed as ?Paraves incertae
sedis, though it may be Paronychodon.
McCrea and Buckley (2004) first reported a theropod tooth excavated
with a hadrosaur from British Columbia in 2002-2003. Reid (2016)
notes McCrea assigned it to Saurornitholestes online, and he refers it to cf. Boreonykus certekorum based on the formation and lack of distal inclination.
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press.
279-297.
McCrea and Buckley, 2004. Excavating British Columbia's first dinosaurs
and other palaeontological projects in the Tumbler Ridge area. Eighth
Alberta Palaeontological Society Symposium, Abstracts. 24-33.
Buckley, McCrea and Currie, 2005. Theropod teeth from the Upper Cretaceous Kaskapau
(Middle Turonian) and the Wapiti (Upper Campanian - Lower Maastrichtian) formations
of north-eastern British Columbia, Canada. Journal of Vertebrate Paleontology.
25(3), 40A-41A.
Fanti and Miyashita, 2009. A high latitude vertebrate fossil assemblage from
the Late Cretaceous of west-central Alberta, Canada: Evidence for dinosaur nesting
and vertebrate latitudinal gradient. Palaeogeography, Palaeoclimatology, Palaeoecology.
275, 37-53.
Bell and Currie, 2015. A high-latitude dromaeosaurid, Boreonykus certekorum,
gen. et sp. nov. (Theropoda), from the upper Campanian Wapiti Formation, west-central
Alberta. Journal of Vertebrate Paleontology. e1034359. DOI: 10.1080/02724634.2015.1034359.
Cau, online 2015. http://theropoda.blogspot.com/2015/12/boreonykus-e-un-dromaeosauridae.html
Reid, 2016. A review of dinosaurian body fossils from British Columbia, Canada. PeerJ Preprints. 4:e1369v3.
S. sp. nov? (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
Material- (RTMP 87.16.18) tooth (Ryan and Russell, 2001)
(UA 103) premaxillary tooth (Baszio, 1997)
(UA 121) premaxillary tooth (Baszio, 1997)
(UA JLE 64:109) lateral tooth (Baszio, 1997)
several premaxillary teeth (Baszio, 1997)
teeth (Ryan and Russell, 2001)
Comments- Baszio (1997) reported UA JLE 64:109 was similar to teeth from the Lance
Formation, perhaps indicating they are the same species. He referred it to Saurornitholestes, which may have merit as he also describes several Zapsalis-style premaxillary teeth that are characteristic of that genus. Those teeth have numerous, strong ridges unlike Bambiraptor or Atrociraptor, and may have more ridges than S. langstoni (6-8).
References- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
S. sp. (Carpenter, 1982)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material-
(UCMP 124983) tooth (2.1 mm) (Carpenter, 1982)
(UCMP 124984) tooth (3 mm) (Carpenter, 1982)
(UCMP 124985) tooth (2.7 mm) (Carpenter, 1982)
(UCMP 125238) dentary fragment (Carpenter, 1982)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
(FMNH PR2893) tooth (4.4x3.1x1.2 mm) (Gates, Zanno and Makovicky,
2015)
(FMNH PR2895) tooth (4.5x3x1.3 mm) (Gates, Zanno and Makovicky, 2015)
(FMNH PR2897) premaxillary tooth (Gates, Zanno and Makovicky, 2015)
(FMNH PR2898) tooth (5.4x3.7x1.3 mm) (Gates, Zanno and Makovicky, 2015)
teeth and elements (DePalma, 2010)
Comments- Carpenter (1982) considered UCMP 125238 to be probably conspecific
with Hell Creek specimen UW 13684. Both have fused interdental plates, subrectangular
alveoli and elongate external foramina, which Carpenter compares to Velociraptor
(in which he includes Saurornitholestes). These characters are also present
in more basal dromaeosaurids such as Sinornithosaurus and Bambiraptor,
though the elongate foramina does distinguish the dentary fragments from Dromaeosaurus.
The former preserves a short, curved tooth with poorly developed mesial serrations
and 5.5 distal serrations per mm. Referred teeth are short, strongly curved
and highly compressed, with well developed distal serrations and variably developed
mesial ones. Carpenter states they resemble Velociraptor, which may indicate
they belong to Saurornitholestes.
FMNH PR2897 is a Zapsalis-style premaxillary tooth, more similar to the Zapsalis holotype than some Saurornitholestes langstoni in lacking mesial serrations and having a concave apicodistal edge. The numerous, strong ridges are unlike Bambiraptor or Atrociraptor.
References-
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous
Lance and Hell Creek formations and a description of a new species of theropod.
Contributions to Geology, University of Wyoming. 20(2), 123-134.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous
bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis,
University of Kansas. 227 pp.
Gates, Zanno and Makovicky, 2015. Theropod teeth from the upper Maastrichtian
Hell Creek Formation "Sue" Quarry: New morphotypes and faunal comparisons.
Acta Palaeontologica Polonica. 60(1), 131-139.
S. sp. nov. (Estes, 1964)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- (AMNH 22670) fourteen teeth (AMNH online)
(AMNH 27122; in part) premaxillary tooth (AMNH online)
(AMNH coll.) premaxillary teeth (Estes, 1964)
(UA BTB 132) premaxillary tooth (Baszio, 1997)
(UA BTB 123, 129-131, 161-164) seven lateral teeth (Baszio, 1997)
(UCM 39502) tooth (1.7 mm) (Carpenter, 1982)
(UCM 45055) tooth (2.6 mm) (Carpenter, 1982)
(UW 13684) dentary fragment (Carpenter, 1982)
(YPM PU 55007) (YPM online)
(YPM PU 55502) (YPM online)
(YPM PU 55522) (YPM online)
(YPM PU 55535) (YPM online)
(YPM PU 55560) (YPM online)
(YPM PU 55561) (YPM online)
Comments- Carpenter (1982) considered UW 13684 to be probably conspecific with Hell Creek
specimen UCMP 125238. Both have fused interdental plates, subrectangular alveoli
and elongate external foramina, which Carpenter compares to Velociraptor
(in which he includes Saurornitholestes). These characters are also present
in more basal dromaeosaurids such as Sinornithosaurus and Bambiraptor,
though the elongate foramina does distinguish the dentary fragments from Dromaeosaurus.
Referred teeth are short, strongly curved and highly compressed, with well developed
distal serrations and variably developed mesial ones. Carpenter states they
resemble Velociraptor, which may indicate they belong to basal dromaeosaurids.
AMNH 22670 is listed as Deinonychus sp. on the AMNH website, but this
is unlikely due to the age difference from the Cloverly Formation.
Baszio (1997) refers Lance Formation teeth (UA BTB coll.) to Saurornitholestes
sp., but notes they differ from S. langstoni in being smaller, with
rounded basal serrations and mesial serrations which are developed as a slight
serrated structure if present. The premaxillary tooth UA BTB 132 has numerous, strong ridges unlike Bambiraptor or Atrociraptor.
The YPM specimens are listed as Dromaeosauridae, but photos indicate they are 'velociraptorine'-type teeth with high DSDIs.
References-
Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern
Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek
formations and a description of a new species of theropod. Contributions to
Geology, University of Wyoming. 20(2), 123-134.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
S? sp. (Williamson, 1997)
Late Santonian-Early Campanian, Late Cretaceous
Allison Member of the Menefee Formation, New Mexico, US
Material- (NMMNH P-25054) tooth (?x3x1.4 mm)
Comments- Williamson (1997) referred a tooth to cf. Saurornitholestes
sp. and Williamson and Brusatte (2014) found it fell within the range of
variation of Dinosaur Park Saurornitholestes teeth.
References- Williamson, 1997. A new Late Cretaceous (Early Campanian)
vertebrate fauna from the Allison Member, Menefee Formation, San Juan Basin,
New Mexico. New Mexico Museum of Natural Hisory and Science Bulletin. 11, 51-59.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
S? sp. (Armstrong-Ziegler, 1980)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (NMMNH P-26240) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27486) tooth (Williamson and Brusatte, 2014)
(NMMNH P-27487) tooth (?x2.7x1.3 mm) (Williamson and Brusatte, 2014)
(NMMNH P-27559) tooth (8.6x5.5x2.5 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30001) tooth (7.1x4x2.1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30002) tooth (?x3.4x1.6 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30003) tooth (3.8x2.9x1.4 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30330) tooth (3.6x3x1.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38027) tooth (?x4.5x2.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38425) tooth (?x2.5x1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38431) tooth (Williamson and Brusatte, 2014)
(NMMNH P-38432) tooth (Williamson and Brusatte, 2014)
(NMMNH P-49808) tooth (?x5.8x2.5 mm) (Williamson and Brusatte, 2014)
(NMMNH P-66896) tooth (9.1x4.3x2.3 mm) (Williamson and Brusatte, 2014)
teeth (Armstrong-Ziegler, 1980)
three teeth (Hall, 1991)
Comments- Armstrong-Ziegler (1978) reported dromaeosaurid teeth, while
three others were referred to Saurornitholestes langstoni by Hall (1991).
Sullivan and Lucas (2006) could not confirm the latter identification. Williamson
and Brusatte (2014) describe several teeth as falling within the range of variation
of Dinosaur Park Saurornitholestes teeth, including some identified by
Hall as troodontids.
References- Armstrong-Ziegler, 1980. Amphibia and Reptilia from the Campanian
of New Mexico. Fieldiana Geology (new series). 4, 39 pp.
Hall, 1991. Lower vertebrate paleontology of the upper Fruitland Formation,
Fossil Forest area, New Mexico, and implications for Late Cretaceous terrestrial
biostratigraphy. Masters Thesis. University of Kansas. 126 pp.
Sullivan and Lucas, 2006. The Kirtlandian land-vertebrate "age" -
faunal composition, temporal position and biostratigraphic correlation in the
nonmarine Upper Cretaceous of western North America. New Mexico Museum of Natural
History and Science Bulletin. 35, 7-29.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
S? sp. (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- ?(NMMNH P-07189) tooth (8.3x?x3.1 mm)
(NMMNH P-30220) tooth (?x?x.5 mm)
(NMMNH P-30222) tooth
(NMMNH P-30223) tooth (?x4.1x2.2 mm)
(NMMNH P-30225) tooth (?x4x2 mm)
(NMMNH P-30226) tooth
(NMMNH P-30227) tooth (?x2.6x1.2 mm)
(NMMNH P-30228) tooth (?x3.7x1.7 mm)
(NMMNH P-30229) tooth (?x3.3x1.5 mm)
(NMMNH P-30230) tooth
(NMMNH P-30231) tooth
(NMMNH P-30232) tooth (?x4.7x2.3 mm)
(NMMNH P-32831) tooth (?x4.3x? mm)
(NMMNH P-33144) tooth (?x~4.3x2 mm)
(NMMNH P-33145) tooth (?x3.1x1.5 mm)
(NMMNH P-33146) tooth (?x4.1x~1.7 mm)
(NMMNH P-33149) tooth
(NMMNH P-33473) tooth (6.5x3.2x1.7 mm)
(NMMNH P-33474) tooth
(NMMNH P-33475) tooth (?x4.6x2.1 mm)
(NMMNH P-33476) tooth (3.2x2.5x1.1 mm)
(NMMNH P-33477) tooth (?x?x1.4 mm)
(NMMNH P-33484) tooth (?x3.8x1.8 mm)
(NMMNH P-33485) tooth
(NMMNH P-33486) tooth (6.7x5.1x2.6 mm)
(NMMNH P-33487) tooth
Comments- Williamson and Brusatte (2014) described these as Dromaeosauridae
morphotype A, which they found fall within the range of variation of Dinosaur
Park Saurornitholestes.
Reference- Williamson and Brusatte, 2014. Small theropod teeth from the
Late Cretaceous of the San Juan Basin, Northwestern New Mexico and their implications
for understanding Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
S? sp. (Sankey et al., 2005)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Material- (LSUMG 5158) tooth (Sankey, 2001)
(LSUMG 5659) tooth (2.8 mm) (Sankey, 2001)
(LSUMG 5923) tooth (5.3 mm) (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 5924) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 5928) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 5950) tooth (~2.2 mm) (Sankey, 2001)
(LSUMG 5980) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6132) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6139) tooth (~5.2 mm) (Sankey, 2001)
(LSUMG 6183) tooth (Sankey, 2001)
(LSUMG 6184) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6185) tooth (Sankey, 2001)
(LSUMG 6204) tooth (6 mm) (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6229) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6234) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6270) tooth (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6280) tooth (~7.5 mm) (Sankey, Standhardt and Shiebout, 2005)
(LSUMG 6281) tooth (7.2 mm) (Sankey, Standhardt and Shiebout, 2005)
? metacarpal, phalanx (Jasinski, Sullivan and Dodson, 2015)
Late Campanian, Late Cretaceous
Aguja Formation, Mexico
teeth (Montellano, Monroy, Hernandez-Rivera and Torres, 2009)
Comments- Montellano et al. (2009) reported teeth from S. langstoni,
plus two unnamed taxa they called Saurornitholestes n. sp. A? and S.
n. sp. C.. Evaluation of these awaits their description.
References- Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation,
Big Bend, Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous
(Campanian-Maastrichtian), Big Bend National Park, Texas. in Carpenter (ed).
The Carnivorous Dinosaurs. 127-152.
Montellano, Monroy, Hernandez-Rivera and Torres, 2009. Late Cretaceous microvertebrate
fauna from the Northern state of Coahuila, Mexico. Journal of Vertebrate Paleontology.
29(3), 151A.
Jasinski, Sullivan and Dodson, 2015. Late Cretaceous dromaeosaurid theropod
dinosaurs (Dinosauria: Dromaeosauridae) from southern Laramidia and implications
for dinosaur faunal provinciality in North America. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 150.
S? sp. (Standhardt, 1986)
Early Maastrichtian, Late Cretaceous
Upper Aguja Formation, Texas, US
Material- (LSUMG 113:coll.) several teeth
Reference- Standhardt, 1986. Vertebrate paleontology of the Cretaceous/Tertiary
transition of Big Bend National Park, Texas. Unpublished Ph.D. dissertation,
Louisiana State University, Baton Rouge. 298 pp.
S? sp. (Schwimmer, Sanders, Erickson and Weems, 2015)
Middle Campanian, Late Cretaceous
Coachman Formation, South Carolina, US
Material- (ChM PV8674) tooth
(ChM PV8675) tooth
(ChM PV8679) incomplete pedal ungual II
(SCSM 2005.11.1) tooth
Comments- Schwimmer et al. (2015) referred these to Saurornitholestes
langstoni.
Reference- Schwimmer, Sanders, Erickson and Weems, 2015. A Late Cretaceous
dinosaur and reptile assemblage from South Carolina, USA. Transactions of the
American Philosophical Society. 105(2), 157 pp.
S? sp. (Schwimmer, Sanders, Erickson and Weems, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Donaho Creek or Peedee Formation, South Carolina, US
Material- (SCSM 99.55.1) partial pedal ungual II
Comments- Schwimmer et al. (2015) referred this to Saurornitholestes
langstoni.
Reference- Schwimmer, Sanders, Erickson and Weems, 2015. A Late Cretaceous
dinosaur and reptile assemblage from South Carolina, USA. Transactions of the
American Philosophical Society. 105(2), 157 pp.
S? sp. (Alifanov and Bolotsky, 2002)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- (1/1081) tooth (15x10x4 mm) (Bolotsky, 2011)
(1/1082) tooth (15x7x2 mm) (Bolotsky, 2011)
(1/1083) tooth (13x8x1.8 mm) (Bolotsky, 2011)
Reference- Alifanov and Bolotsky, 2002. New data about the assemblages
of the Upper Cretaceous carnivorous dinosaurs (Theropoda) from the Amur region.
In Kirillova (ed.). Fourth International Symposium of IGCP 434. Cretaceous continental
margin of East Asia: Stratigraphy, sedimentation, and tectonics. 25-26.
Bolotsky, 2011. On paleoecology of carnivorous dinosaurs (Tyrannosauridae, Dromaeosauridae)
from Late Cretaceous fossil deposits of Amur region, Russian far East. Global
Geology. 14(1), 1-6.
Tsaagan Norell, Clark, Turner, Makovicky,
Barsbold and Rowe, 2006
= Linheraptor Xu, Choiniere, Pittman, Tan, Xiao, Li, Tan, Clark, Norell,
Hone and Sullivan, 2010
T. mangas Norell, Clark, Turner, Makovicky, Barsbold and Rowe,
2006
= Velociraptor mangas (Norell, Clark, Turner, Makovicky, Barsbold and
Rowe, 2006) Paul, 2010
= Linheraptor exquisitus Xu, Choiniere, Pittman, Tan, Xiao, Li, Tan,
Clark, Norell, Hone and Sullivan, 2010
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadokhta Formation, Mongolia
Holotype- (IGM 100/1015) (adult) skull (201 mm), sclerotic rings, mandibles,
ten cervical vertebrae, cervical ribs, proximal scapula, partial coracoid
Late Campanian, Late Cretaceous
Zos Wash, Djadokhta Formation, Mongolia
Referred- (IGM 100/3503; "IGM 100/981" of Turner et al., 2007) (~1.5 m, ~15
kg) frontal, jugal fragment, ectopterygoids, pterygoid, incomplete
dentary, dorsal vertebra, few proximal dorsal ribs, gastralia, seven
distal caudal vertebrae, several chevrons, scapulocoracoid, humeral
fragment, radius, ulna (~113 mm), scapholunare, semilunate carpal,
metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx
II-1, phalanx II-2, manual ungual II, phalanx III-1, phalanx III-2,
phalanx III-3, manual ungual III, distal pubis, ischium, proximal
fibula, phalanges I-1, pedal unguals I, propximal metatarsal II,
proximal metatarsal III, phalanges III-3, pedal unguals III,
metatarsals IV (one proximal), phalanx IV-4, pedal ungual IV,
metatarsals V (Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006)
Late Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Referred- (IVPP V16923; holotype of Linheraptor exquisitus) (~1.8 m; adult)
skull (225 mm), sclerotic rings, incomplete mandibles, hyoids (one partial),
(cervical series 320 mm) atlas, axis, third cervical vertebra, fourth cervical
vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical
vertebra, eighth cervical vertebra, ninth cervical vertebra, tenth cervical
vertebra, cervical ribs 2-10, first dorsal vertebra, dorsal ribs 1-8, tenth
dorsal rib, uncinate processes, gastralia, seven proximal caudal vertebrae (first
caudal 22.2 mm), eleven distal caudal vertebrae (longest 30 mm), seven proximal
chevrons, distal chevrons, scapula (~160 mm), sternum (80 mm), humerus (~155
mm), radius (110 mm), metacarpal I (25.3 mm), phalanx I-1 (47.6 mm), metacarpal
II (64.4 mm), phalanx II-1 (32.3 mm), phalanx II-2 (53.5 mm), manual ungual
II (41 mm), metacarpal III (53.1 mm), phalanx III-1 (15.5 mm), phalanx III-2
(12 mm), phalanx III-3 (33 mm), pubis (240 mm), femur (220 mm), tibiotarsi (252
mm), fibula, distal tarsal IV, phalanx I-1 (15.7 mm), pedal ungual ! (17.5 mm
straight), metatarsal II (103.2 mm), phalanx II-2 (39.9 mm), pedal unguals II
(55.6 mm), metatarsal III (127.9 mm), phalanx III-1 (54.2 mm), phalanx III-2
(32.9 mm), phalanx III-3 (19.3 mm), pedal ungual III (48.5 mm), metatarsal IV
(112.8 mm), phalanx IV-1 (40 mm), phalanx IV-2 (28 mm), phalanx IV-3 (26.1 mm),
phalanx IV-4 (26.4 mm), pedal ungual IV (18 mm) (Xu et al., 2010)
Diagnosis- (after Norell et al., 2006) paroccipital process pendulous;
maxillary fenestra large; maxillary fenestra located at anterior edge of antorbital
fossa (also in Atrociraptor); jugal meets the squamosal to exclude the
postorbital from the margin of the infratemporal fenestra.
(after Xu et al., 2010) Xu et al. listed two characters supposedly shared between
Linheraptor and Tsaagan, which would be diagnostic of the latter
if they are synonymous- lacrimal with relatively broad medial lamina; sharp
angle between anterior and dorsal processes of quadratojugal.
(after Turner et al., 2012) oval-shaped foramen magnum; low coracoid tuber;
weak subglenoid shelf; dorsoventrally oriented path of supracoracoid nerve through
coracoid.
(after Xu et al., 2015) Note Xu et al. viewed Tsaagan and Linheraptor
as taxonomically distinct, but listed the following characters as shared between
them. Promaxillary fenestra not visible in lateral view; maxilla without ridge
dorsal to supralabial foramina (also probably in Achillobator); longitudinal
fossa along midline of nasals (also in Velociraptor); jugal without ridge
parallel to orbital rim; frontal supraorbital rim rugose (also in Dromaeosaurus);
parietal with prominent nuchal crest (also in Adasaurus); parietal forms
entire nuchal crest (also in Sinornithosaurus); pterygoid flange small
(also in Dromaeosaurus); all teeth without mesial serrations (also in
some microraptorians).
Other diagnoses- Norell et al. (2006) originally listed paroccipital
process not twisted distally in their diagnosis, but this is absent in the Linheraptor
holotype (Xu et al., 2015). Turner et al. (2012) suggested it was taphonomic
in the Tsaagan holotype, though Xu et al. view it as a character supporting
taxonomic separation of the specimens. Similarly, Norell et al. listed 'basipterygoid
process elongate and anteroventrally directed' as being diagnostic, but the
Linheraptor holotype has shorter processes which are only ventrally directed.
Again, Xu et al. view it as having taxonomic implications, thouigh it could
also be individual variation.
Xu et al. (2010) listed two supposed autapomorphies of Linheraptor- greatly
enlarged maxillary fenestra subequal in size to external naris; several large
foramina on lateral surface of jugal. The former would be individual variation
if the genus is a synonym of Tsaagan, while Xu et al. (2015) later noted
the jugal foramina are also present in Velociraptor and "might be
present in T. mangas but are simply obscured by specimen damage."
They also listed 'lacrimal lacking lateral flange over descending process' as
shared between Linheraptor and Tsaagan, but this is actually untrue
for former's holotype (Xu et al., 2015).
Xu et al. (2015) describe 61 differences between Linheraptor and Tsaagan,
interpreting the following as autapomorphies of the former- nasal process of
premaxilla transversely compressed along entire length; premaxillary process
of nasal terminates at anterior border of external naris; lacrimal with several
anteriorly opening foramina on dorsal surface (possibly obscured by poor preservation
in Tsaagan); pyramidal projection on anterior border of supratemporal
fossa; supraoccipital crest short and sharp; quadratojugal dorsal process distally
expanded; basipterygoid process is not inclined anteriorly. If the taxa are
synonymous, these are merely examples of individual variation, as seen in other
theropods (e.g. Allosaurus, Tyrannosaurus, Microraptor,
Archaeopteryx).
Comments- The holotype was discovered in 1993 and originally identified
as Velociraptor (e.g. Webster, 1996) before being identified as a new
genus (Turner et al., 2006) then described by Norell et al. (2006). Tsaagan
(under its specimen number) was used as an OTU in the Theropod Working Group
analyses since 2001 however. Powers et al. (2022) provide CT scan data on the maxilla. Linheraptor's holotype was discovered in
2008 and described by Xu et al. (2010) as a new taxon of dromaeosaurid. Both
Senter (2011) and Turner et al. (2012) synonymized Linheraptor with Tsaagan,
though only the latter justified it. This is provisionally accepted here despite
a recent paper by Xu et al. (2015) defending Linheraptor's validity based
on 61 differences from Tsaagan. While these differences are real, Xu
et al. do not establish they are outside the range of individual variation expected
for a dromaeosaurid species. To the contrary, they dismiss the idea of utilizing
individual variation present in Velociraptor mongoliensis until authors
"determine how observed morphological variations relate to inter- or possibly
intraspecific factors" ... "by including all Velociraptor specimens
in a specimen-level phylogenetic analysis, by using morphometric methods to
quantify the variation present and by deepening our understanding of the biological
significance of the variations observed." Yet while ideal, such studies
have not been completed for any Mesozoic theropod. Thus insisting "noticeable
variations between L. exquisitus and T. mangas are grounds for
taxonomic separation" until those studies are done on Velociraptor
would logically result in the numerous differences present between almost all
described individuals of e.g. Allosaurus and Tyrannosaurus being
viewed as taxonomically significant until the studies are done on them as well.
Such a conclusion is highly unrealistic, and given the large amount of individual
variation present in all theropod species including V. mongoliensis,
Tsaagan and Linheraptor are accepted as synonymous here until
either further specimens show their differences covary or phylogenetic analyses
find them to not be sister taxa even when their shared characters are included.
Tsaagan has been recovered as a velociraptorine (Senter et al., 2012;
Brusatte et al., 2014) and a saurornitholestiine (Longrich and Currie, 2009).
IGM 100/3503- Discovered in 1998, Norell et al. (2006) mention "A second, as yet undescribed species of dromaeosaurid, which differs from Tsaagan mangas
in frontal morphology [that] is known from the nearby locality of Zos
Wash." Turner et al. (2007) describe an ulna as IGM 100/981, referring
it to Velociraptor mongoliensis
based on sharing several unlisted characteristics. This specimen was
said to be from "the Gilvent Wash locality near Ukhaa Tolgod", but IGM
100/981 is a specimen from Khulsan later described as Kuru.
Napoli et al. (2021) clarified the matter, stating "Turner et al.
(2007a) erroneously identified the Zos Wash specimen as IGM 100/981.
The correct specimen number for the Zos Wash specimen is IGM
100/3503." Ironically Ruebenstahl et al.'s (2021) abstract also
switches 100/981 and 100/3503, but they do say the Zos Wash specimen
"appears referrable to Velociraptor, although it differs from other Velociraptor mongoliensis
specimens in some respects such as the absence of a dorsal
ectopterygoid recess." Napoli et al. state "This individual is
currently referred to Velociraptor mongoliensis
and is under renewed study." Based on data presented in Ruebenstahl et
al.'s SVP presentation and preliminary inclusion in the Hartman et al.
maniraptoromorph analysis, I believe it is closest to Tsaagan and falls within the range of variation of the latter and Linheraptor.
References- Webster, 1996. Dinosaurs of the Gobi: Unearthing a fossil
trove. National Geographic. 190(1), 70-89.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid
theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545,
51 pp.
Turner, Pol, Norell and Hwang, 2006. Resolving dromaeosaurid phylogeny: New
information and additions to the tree. Journal of Vertebrate Paleontology. 26(3),
133A.
Turner, Makovicky and Norell, 2007. Feather quill knobs in the dinosaur Velociraptor.
Science. 317, 1721.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
from the Late Cretaceous of North America. Proceedings of the National Academy
of Sciences. 106(13), 5002-5007.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Xu, Choiniere, Pittman, Tan, Xiao, Li, Tan, Clark, Norell, Hone and Sullivan,
2010. A new dromaeosaurid (Dinosauria: Theropoda) from the Upper Cretaceous
Wulansuhai Formation of Inner Mongolia, China. Zootaxa. 2403, 1-9.
Senter, 2011. Using creation science to demonstrate evolution 2: Morphological
continuity within Dinosauria. Journal of Evolutionary Biology. 24, 2197-2216.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2012 online. Tsaagan mangas, Digital Morphology. http://digimorph.org/specimens/Tsaagan_mangas/
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Brusatte, Vremir, Csiki-Sava, Turner, Watanabe, Erickson and Norell, 2013. The
osteology of Balaur bondoc, an island-dwelling dromaeosaurid (Dinosauria:
Theropoda) from the Late Cretaceous of Romania. Bulletin of the American Museum
of Natural History. 374, 1-100.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Xu, Pittman, Sullivan, Choiniere, Tan, Clark, Norell and Wang, 2015. The taxonomic
status of the Late Cretaceous dromaeosaurid Linheraptor exquisitus and
its implications for dromaeosaurid systematics. Vertebrata PalAsiatica. 53(1),
29-62.
Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021. A new
dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central
Mongolia. American Museum Novitates. 3982, 47 pp.
Ruebenstahl, Napoli, Bhullar, Turner and Norell, 2021. Two new
eudromaeosaurs from Khulsan (central Mongolia) reveal modern-like
faunal predatory structure amoung non-avian dinosaurs. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 222-223.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as
2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist
in testing and constructing morphological characters. Journal of
Vertebrate Paleontology. 41(5), 2010087.
undescribed dromaeosaurine (Powers, Sullivan and Currie, 2020)
Late Campanian, Late Cretaceous
Djadokhta Formation, Mongolia
Material- (private coll.; cast UALVP 49389) skull (~193 mm), sclerotic plates, mandibles, pes
Comments- Powers et al. (2020)
included in their analysis "a cast of a complete skull belonging to an
undescribed eudromaeosaurian", which "is of a specimen in private
collections and has no available locality data. The likeness of this
specimen is akin to that of an Asian dromaeosaurid. and has been
frequently labeled as Velociraptor,
however, its morphology is quite different from other specimens
referred to this genus." They report that "In the cluster
analysis UALVP 49389 was found to be most like [Tsaagan and Linheraptor]
and often plotted close to them but toward the extreme of variation
along each PC." The maxilla is illustrated in their figure 1M and
differs from most other dromaeosaurids in lacking an externally obvious
promaxillary fenestra and having a slender maxillary fenestra along the
ventral edge of the ascending process, and is similar to Tsaagan and Velociraptor in having an elongated anterior ramus. Powers' (2020) thesis this paper is based on listed the specimen as Velociraptor
sp. and noted it consists of "Cast of articulated skull and foot",
while the UA online catalogue lists it being from the Djadokhta
Formation of Mongolia. Powers et al.'s figure allows one to
identify the cast as one labeled Velociraptor mongoliensis
in the WitmerLab Collections (Witmer, online 2012) and ultimately sold
by Gaston Design Inc. (2022 online; but available since at least 2004- https://web.archive.org/web/20040609002120/http://gastondesign.com/dinoskulls.htm), although no pes is associated with either cast.
References- Witmer, online 2012. https://people.ohio.edu/witmerl/collections/Theropods/Velociraptor_gaston.htm
Powers, 2020. The evolution of snout shape in eudromaeosaurians and its
ecological significance. Masters thesis, University of Alberta. 437 pp.
Powers, Sullivan and Currie, 2020. Re-examining ratio based
premaxillary and maxillary characters in Eudromaeosauria (Dinosauria :
Theropoda): Divergent trends in snout morphology between Asian and
North American taxa. Palaeogeography, Palaeoclimatology, Palaeoecology.
547, 109704.
Gaston Design Inc., 2022 online. https://gastondesign.com/product/velociraptor-skull/
undescribed dromaeosaurine (Currie and Varricchio, 2004)
Cenomanian-Turonian, Late Cretaceous
Teel Ulaan Jalzai, Bayanshiree Formation, Mongolia
Material-
(IGM 100/22) premaxillae, partial maxillae, four dorsal vertebrae (25,
25 mm), five proximal dorsal ribs, first caudal vertebra, second caudal
vertebra (29 mm), third caudal vertebra (29 mm), fourth caudal vertebra
(31 mm), fifth caudal vertebra (32 mm), sixth caudal vertebra (32 mm),
seventh caudal vertebra, eighth caudal vertebra (35 mm), ninth caudal
vertebra (37 mm), tenth caudal vertebra (37 mm), eleventh caudal
vertebra (37 mm), twelfth caudal vertebra (36 mm), thirteenth caudal
vertebra (34 mm), fourteenth caudal vertebra (35 mm), fifteenth caudal
vertebra (34 mm), sixteenth caudal vertebra (33 mm), seventeenth caudal
vertebra, eighteenth caudal vertebra (30 mm), nineteenth caudal
vertebra (29 mm), twentieth caudal vertebra (26 mm), twenty-first
caudal vertebra, four chevrons, proximal scapulae, proximal coracoids,
incomplete humeri (~186 mm), radii (one partial; 134 mm), ulnae (one
incomplete; 146 mm), scapholunares, semilunate carpals, metacarpals I
(34, 34 mm), phalanges I-1 (59, 61 mm), incomplete manual unguals I,
metacarpals II (one incomplete; 72 mm), phalanges II-1 (one incomplete;
45 mm), phalanges II-2 (one incomplete; 59 mm), incomplete manual
unguals II, metacarpals III (one incomplete; 59 mm), phalanges III-1
(one incomplete; 24 mm), phalanges III-2 (17, 17 mm), phalanges III-3
(one incomplete; 43 mm), incomplete manual unguals III, ischium (130
mm), distal femur, tibia (279 mm), incomplete fibula (249 mm),
incomplete astragalus, metatarsals I (one incomplete; 39 mm), phalanx
I-1 (26 mm), pedal ungual I (~28 mm), distal tarsal III fused to
metatarsal II and III (mtII 111, mtIII 134 mm), incomplete pedal ungual
II, phalanges III-1 (one partial; 52 mm), partial phalanges III-2,
proximal phalanx III-3, pedal ungual III (~44 mm), metatarsal IV (121
mm), partial phalanges IV-1, phalanges IV-2 (31, 30 mm), phalanges IV-3
(24 mm), phalanges IV-4 (22 mm), pedal unguals IV, pedal phalanx x-?
(33 mm)
Cenomanian-Turonian, Late Cretaceous
Shine Us Khuduk, Bayanshiree Formation, Mongolia
(IGM 100/23; intended holotype) partial skull, incomplete mandibles,
posterior synsacrum, first caudal vertebra, second caudal vertebra (28
mm), third caudal vertebra (33 mm), fourth caudal vertebra (35 mm), two
proximal chevrons, posterior ilium, incomplete ischium, proximal
metatarsal II, proximal metatarsal III, proximal metatarsal IV
Diagnosis (after Kubota and Barsbold, 2007) convex posterodorsal edge
of ilium with posteriorly curved distal end; transversely wide distal end of
pedal phalanx II-1.
Other diagnoses- Turner et al. (2012) noted other dromaeosaurids like
Velociraptor and Dromaeosaurus also possess two shallow subalveolar
grooves on the labial surface of the dentary, which Kubota and Barsbold considered
diagnostic.
Comments- These specimens were discovered in the 1970s and must have been referred to Adasaurus
early on, as the ischium of IGM 100/22 was used in Barsbold's (1983)
pelvic illustration. Similarly, scorings indicate Norell et al.
(2001) and Senter (2007) used the specimens when scoring Adasaurus
into their TWiG analyses. However, the fact these were new
specimens besides the Adasaurus holotype and paratype was not
published until Currie and Varricchio (2004), who provided some dental
information and specimen numbers. Shortly after in an SVP
abstract, Kubota and Barsbold (2007) determined they are from a
different taxon, with Adasaurus being a velociraptorine which
lived later. They find it to group with Dromaeosaurus and Achillobator
based on the low DSDI, and differ from Dromaeosaurus in having less anterior
maxillary teeth with lingually twisted mesial carinae, and from Achillobator
in lacking double dorsal pleurocoels, lacking caudal pleurocoels, having a low
ilium with tapered postacetabular process and a distally placed obturator process.
Kubota (2015) describes and names the taxon in his thesis.
Currie and Varricchio (2004) report there are four premaxillary teeth,
eleven maxillary teeth and thirteen dentary teeth in IGM 100/23 and
100/22, but 100/22 doesn't preserve a dentary and only has fragmentary
maxillae (Kubota, 2015). They also state teeth have four serrations per
mm in both mesial and distal carinae, but the actual values are 3.2-5
and 3.5-5.5 respectively (average 4.25 and 4.19). Jasinowski et al.
(2006) list humeri and an ulna for IGM 100/22 (which is only labeled
'dromaeosaur').
References-
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions
of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian
dinosaurs. In Gauthier and Gall (eds.). New Perspectives on the
Origin and Early Evolution of Birds: Proceedings of the International Symposium
in Honor of John H. Ostrom. Yale University Press. 49-67.
Currie and Varricchio, 2004. A new dromaeosaurid from the
Horseshoe Canyon Formation (Upper Cretaceous) of Alberta, Canada. in Currie,
Koppelhus, Shugar and Wright (eds). Feathered Dragons. Studies on the transition
from dinosaurs to birds. Indiana University Press. 112-132.
Jasinowski, Russell and Currie, 2006. An integrative phylogenetic and extrapolatory
approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder
musculature. Zoological Journal of the Linnean Society. 146, 301-344.
Kubota and Barsbold, 2007. New dromaeosaurid (Dinosauria Theropoda) from the
Upper Cretaceous Bayanshiree Formation of Mongolia. Journal of Vertebrate Paleontology.
27(3), 102A.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 5(4), 429-463.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Kubota, 2015. Descriptions of Mongolian dromaeosaurids (Dinosauria:
Theropoda) and phylogeny of Dromaeosauridae. PhD thesis, University of
Tsubaka. 397 pp.
Dromaeosaurus Matthew
and Brown, 1922
Not Dromaeosaurus- The teeth referred to Dromaeosaurus
by Russell (1935) from the Milk River Formation of Alberta seem to have a large
DSDI, so are more probably Saurornitholestes. Colbert and Russell (1969)
tentatively referred a fourth metatarsal (CMN 12072) and a pedal ungual II (CMN
12240) to Dromaeosaurus, as did Ostrom (1969) for the latter element.
Paul (1988b) referred the ungual to Saurornitholestes instead, as he
hypothesized Dromaeosaurus to have a reduced ungual as in Adasaurus.
Both elements were referred to Saurornitholestes by Currie (1995).
Sues (1977) described two dentaries (UA 12091 and 12339) discovered in 1969
and 1974 respectively. He referred these to Dromaeosaurus sp., but they
were later realized to be Saurornitholestes (Paul, 1988b).
Rowe et al. (1992) identified cf. Dromaeosaurus teeth (including TMM
43057-314) from the Aguja Formation of Texas. Sankey (1998) later identified
Dromaeosaurus teeth from another area of that formation, but these and
Rowe et al.'s specimens were referred to Theropoda "family and genus undetermined."
They consisted of two tooth fragments (LSUMG 5483 and 6239) which were similar
to Dromaeosaurus except in lacking a lingually twisted mesial carina,
and were reidentified as tyrannosaurid teeth by Sankey et al. (2005).
Matthew and Brown (1922) referred Lalaeps laevifrons to Dromaeosaurus
as D. laevifrons, while Sahni (1972) synonymized it with D. albertensis.
It is properly referred to Saurornitholestes langstoni. Matthew and Brown
also tentatively referred Laelaps cristatus to the genus as D? cristatus,
but this is a Troodon specimen as recognized by Olshevsky (1995). They
referred Laelaps explanatus to Dromaeosaurus? sp., which Kuhn
(1939) formalized as Dromaeosaurus explanatus. Sahni synonymized it with
D. albertensis, but is is a Saurornitholestes specimen. Matthew
and Brown also referred Laelaps falculus to Dromaeosaurus? sp.,
which Olshevsky (1978) formalized as Dromaeosaurus falculus. Sahni also
synonymized this species with D. albertensis, but is it probably a juvenile
Gorgosaurus or Daspletosaurus. Zapsalis abradens was considered
comparable to Dromaeosaurus by Matthew and Brown, and Tracy Ford's taxonomic
lists include the combination Dromaeosaurus abradens. More recently,
Zapsalis teeth were called ?Dromaeosaurus morphotype A by Sankey
et al. (2002), but Zapsalis is now known to be Saurornitholestes premaxillary teeth (Currie and Evans, 2020). Matthew and Brown referred "Coelurus"
gracilis to Dromaeosaurus as D? gracilis, but it cannot be
compared to Dromaeosaurus though it does seem to be dromaeosaurid. Tracy
Ford's lists also include the combination Dromaeosaurus minutus for "Ornithomimus"
minutus, an alvarezsaurid. Finally, Paul (1988a) referred Adasaurus
to Dromaeosaurus as D. mongoliensis, but now that the genus is
better understood it appears to be a more basal dromaeosaurid.
References- Matthew and Brown, 1922. The family Deinodontidae, with notice
of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum
of Natural History. 46(6), 367-385.
Russell, 1935. Fauna of the Upper Milk River beds, Southern Alberta. Transactions
of the Royal Society of Canada, series 3, section 4. 29, 115-127.
Kuhn, 1939. Fossillium Catalogus.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus.
American Museum Novitates. 2380, 49 pp.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod
from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural
History. 30, 165 pp.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin
of the AMNH. 147.
Sues, 1977. Dentaries of small theropods from the Judith River Formation (Campanian)
of Alberta, Canada. Canadian Journal of Earth Sciences. 14, 587-592.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster Co., New York.
464 pp.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods
of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus
- and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River
- Deinonychus, Velociraptor and Saurornitholestes. Hunteria.
2(4), 1-9.
Rowe, Ciffelli, Lehman and Weil, 1992. The Campanian Terlingua local fauna,
with a summary of other vertebrates from the Aguja Formation, Trans-Pecos, Texas.
Journal of Vertebrate Paleontology. 12, 472-493.
Currie, 1995. New information on the anatomy and relationships of Dromaeosaurus
albertensis (Dinosauria: Theropoda). Journal of Vertebrate Paleontology.
15(3), 576-591.
Olshevsky, 1995. The origin and evolution of the tyrannosaurids. Kyoryugaku
Saizensen [Dino-Frontline]. 9, 92-119 (part 1); 10, 75-99 (part 2) [in Japanese].
Sankey, 1998. Vertebrate paleontology and magnetostratigraphy of the upper Aguja
Formation (Late Campanian), Talley Mountain area, Big Bend National Park, Texas.
Unpublished Ph.D. dissertation, Louisiana State University, Baton Rouge. 263
pp.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend,
Texas. Journal of Paleontology. 75(1), 208-215.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous
(Campanian-Maastrichtian), Big Bend National Park, Texas. in Carpenter (ed).
The Carnivorous Dinosaurs. 127-152.
Currie and Evans, 2020 (online 2019). Cranial anatomy of new specimens of Saurornitholestes langstoni
(Dinosauria, Theropoda, Dromaeosauridae) from the Dinosaur Park
Formation (Campanian) of Alberta. The Anatomical Record. 303(4),
691-715.
D. albertensis Matthew and Brown, 1922
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Belly River Group, Alberta, Canada
Holotype- (AMNH 5356) (adult) incomplete skull (240 mm) partial sclerotic
ring, mandibles (202.5 mm), hyoids, metacarpal I, distal metatarsal I, phalanx
I-1 (27 mm), distal metatarsal II, phalanx II-1 (37 mm), phalanx II-2 (39 mm),
distal metatarsal III, distal phalanx III-1, phalanx III-2 (32 mm), partial
pedal ungual III, distal phalanx IV-2, phalanx IV-3 (24 mm)
Referred- (ANSP 15827) tooth (Fiorillo and Currie, 1994)
(ANSP 15959) premaxillary tooth (Fiorillo and Currie, 1994)
(ANSP 15963) anterior maxillary tooth (Fiorillo and Currie, 1994)
(ANSP 17639) tooth (Fiorillo and Currie, 1994)
(ANSP 17805) tooth (Fiorillo and Currie, 1994)
(ANSP 17960) tooth (Fiorillo and Currie, 1994)
(ANSP 17798) tooth (Fiorillo and Currie, 1994)
(ANSP 18108) tooth (Fiorillo and Currie, 1994)
(ANSP 18110) tooth (Fiorillo and Currie, 1994)
(CMN 12349) frontal (Sues, 1978)
(RTMP 66.25.16) lateral tooth (14 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 66.31.117) tooth (Baszio, 1997)
(RTMP 74.10.91) lateral tooth (Baszio, 1997)
(RTMP 78.9.41) tooth (Baszio, 1997)
(RTMP 79.8.732) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 79.11.165) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 79.14.100) premaxillary tooth (8.8 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 79.14.1007) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 79.15.2) premaxillary tooth (13 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 80.8.298) lateral tooth (9.8 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 80.8.308) lateral tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 80.13.35) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 80.16.2094) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 81.14.60) lateral tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 81.16.161) premaxillary tooth (10.5 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 81.16.281) lateral tooth (10 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 81.16.461) premaxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.22.93) premaxillary tooth (7.8 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 81.26.48) anterior dentary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 81.26.175) lateral tooth (11 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 81.27.66) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 82.11.2) lateral tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 82.16.361) lateral tooth (Baszio, 1997)
(RTMP 82.18.137) lateral tooth (11 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 82.18.250) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 82.19.185) partial dentary (Currie, 1987a)
(RTMP 83.36.8) maxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 83.67.38) lateral tooth (13.2 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 84.67.115) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 84.89.47) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 84.89.48) lateral tooth (14.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 84.92.267) premaxillary tooth (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 85.6.135) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.36.332) lateral tooth (6.4 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 85.36.336) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 85.36.337) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.36.338) premaxillary tooth (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 85.43.4) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.52.10) premaxillary tooth (6.5 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 85.56.200) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.59.11) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 85.59.81) premaxillary tooth (12 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 85.66.56) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 85.68.32) premaxillary tooth (14.5 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 85.68.47) premaxillary tooth (9.8 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 86.5.39) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.11.20) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.18.99) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 86.33.52) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 86.54.67) premaxillary tooth (8 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 86.76.11) lateral tooth (17.2 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 86.76.13) premaxillary tooth (13 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 86.130.211) lateral tooth (9.5 mm) (Baszio, 1997)
(RTMP 86.130.218) lateral tooth (10.8 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 86.184.44) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.77.136) premaxillary tooth (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 87.153.56) premaxillary tooth (9.5 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 87.157.30) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.50.45) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.50.66) lateral tooth (Baszio, 1997)
(RTMP 88.50.127) premaxillary tooth (10 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 88.77.46) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.86.30) premaxillary tooth (7.3 mm) (Baszio, 1997)
(RTMP 88.87.87) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.215.73) lateral tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 89.36.354) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 89.36.404) lateral tooth (Baszio, 1997)
(RTMP 89.77.6) lateral tooth (11 mm), lateral tooth (11.5 mm) (Sankey, Brinkman,
Guenther and Currie, 2002)
(RTMP 89.103.11) lateral tooth (15.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 89.155.2) lateral tooth (14.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 90.145.1) premaxillary tooth (10.7 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 92.77.2) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 93.36.460) lateral tooth (16.2), lateral tooth (16.5 mm) (Sankey, Brinkman,
Guenther and Currie, 2002)
(RTMP 93.36.462) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 93.36.472) lateral tooth (~13 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 94.12.241) lateral tooth (~15.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 94.12.243) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 94.12.247) lateral tooth (15.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 94.12.266) premaxillary tooth (9.2 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 94.12.652) lateral tooth (12.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 94.92.2) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.99.2) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.99.3) premaxillary tooth (9.2 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 94.142.4) lateral tooth (18 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 94.172.39) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.127.26a) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.127.26b) lateral tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.134.5) premaxillary tooth (11 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.137.1) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.143.45) lateral tooth (9 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.151.8) premaxillary tooth (10.5 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 95.151.10) jaw bone with lateral tooth (13 mm) (Sankey, Brinkman, Guenther
and Currie, 2002)
(RTMP 95.171.40) jaw bone with two lateral teeth (14, 12.3 mm) (Sankey, Brinkman,
Guenther and Currie, 2002)
(RTMP 95.181.9) premaxillary tooth (7.3 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP 95.406.4) lateral tooth (11.4 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.665.59) lateral tooth (12 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 96.12.362) lateral tooth (13.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 98.93.172) lateral tooth (10.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 99.55.328) lateral tooth (8.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP coll.; Bob's site) lateral tooth (13 mm) (Sankey, Brinkman, Guenther and
Currie, 2002)
(RTMP coll.) three teeth (Ryan et al., 2001)
(RTMP coll.) almost a dozen postcranial elements (Currie, 2005)
thirteen specimens (Brinkman et al., 1998)
Late Campanian, Late Cretaceous
Judith River Formation, Montana
Material- (AMNH 8516) lateral tooth (Sahni, 1972)
(AMNH 8517) sixteen teeth (AMNH online)
?(AMNH 8519) tooth (AMNH online)
?(AMNH 8520) posterior dentary tooth (AMNH online)
(AMNH 8521) thirty-four teeth (AMNH online)
(UCMP 137558) tooth (UCMP online)
?(UCMP 154575) manual ungual III (UCMP online)
(YPM PU 21847) (YPM online)
(YPM PU 22245) (YPM online)
Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- (AMNH 12884) ten teeth (AMNH online)
(AMNH 12885) seven teeth (AMNH online)
(AMNH 12886) fourteen teeth (AMNH online)
(AMNH 12887) six teeth (AMNH online)
(AMNH 12888) tooth (AMNH online)
(UCMP 120850) two teeth (UCMP online)
material (Demar and Breithaupt, 2006)
Campanian-Maastrichtian, Late Cretaceous
Prince Creek Formation, Alaska, US
Material- (AK232-V-073) tooth (Fiorillo and Gangloff, 2000)
(AK238-V-009) tooth (Fiorillo and Gangloff, 2000)
(AK292-V-03) tooth (Fiorillo and Gangloff, 2000)
(AK381-V-052) tooth (Fiorillo and Gangloff, 2000)
(AK306-V-035) tooth (Fiorillo and Gangloff, 2000)
(AK335-V-013) tooth (Fiorillo and Gangloff, 2000)
(AK383-V-180) tooth (Fiorillo and Gangloff, 2000)
(AK385-V-002) tooth (Fiorillo and Gangloff, 2000)
(AK390-V-041) tooth (Fiorillo and Gangloff, 2000)
(AK456-V-024) tooth (Fiorillo and Gangloff, 2000)
(AK456-V-025) tooth (Fiorillo and Gangloff, 2000)
(AK490-V-170) tooth (Fiorillo and Gangloff, 2000)
(AK497-V-001FT) tooth (Fiorillo and Gangloff, 2000)
(UAM-AK83.V-090) tooth (Fiorillo and Gangloff, 2000)
Diagnosis- (after Turner et al., 2012) premaxilla deeper and thicker
than other dromaeosaurids; presence of either an enlarged promaxillary fenestra
or an extremely anteroventrally placed maxillary fenestra with no promaxillary
fenestra depending on interpretation of identity of opening; anteroposterior
short lateral lamina of maxilla anterior to antorbital fossa; nine maxillary
teeth; nasals with V-shaped suture posteriorly between frontals; deep notches
anteriorly on frontal for articulation with lacrimal; tip of frontal flatter
and margin of supratemporal fossa less pronounced; postorbital process of frontal
sharply demarcated from the dorsomedial orbital margin; quadratojugal stout;
dorsal tympanic recess very weakly expressed; moderately developed preotic pendant;
expression of anterior tympanic recess and/or basipterygoid recess absent on
the basisphenoid or basipterygoid processes; posteromedial process of palatine
slender; anterior and posterior tooth denticles subequal in size; anterior carina
of maxillary or dentary tooth close to midline of tooth near tip, twists toward
lingual surface.
Comments- The holotype was discovered in 1914 but not described until
1922 (Matthew and Brown, 1922) as a new subfamily of tyrannosaurs. Currie (1987a)
stated that Dinosaur Park Dromaeosaurus teeth suggest more than one species
was present, citing the in prep. at the time Currie et al. (1990). He noted
CMN 12349 is more robust than the holotype and that the supratemporal fossa
ridge is more strongly curved, implying these could be reason for referring
it to the other Dromaeosaurus species. However, Currie et al. never proposed
more than one morphotype of Dromaeosaurus once it was published, though
they did refer to teeth with Dromaeosaurus-like serrations which lack
a twisted mesial carina that "may represent a distinct species of dromaeosaurid,
or a gracile, small form of tyrannosaurid." Similar teeth from the Aguja
Formation have been referred to juvenile tyrannosaurids.
Is CMN 12349 a therizinosaur frontal?- Sues (1978) identified frontal CMN 12349 as Dromaeosaurus
and frontal CMN 12355 as Theropoda indet., although note his plates 7
and 8 are switched so that they are given each others' captions. Currie
(1987b) accepted CMN 12349 as Dromaeosaurus, but with regard to CMN 12355 stated that
"comparison with Mongolian specimens suggests that it may represent Erlicosaurus (Currie, in preparation)." Currie (1992) again stated CMN 12355 "may represent the segnosaurid Erlicosaurus"
(although note that as in Sues' paper it is mislabeled 12349 in his
Figure 2) and also referred TMP 1981.016.0231 to Segnosauridae. Currie
(2005) listed CMN 12349 as a tentative "therizinosauroid similar to Erlikosaurus", but figured CMN 12355 so probably meant that specimen. Indeed, it seems CMN 12349 was correctly identified as Dromaeosaurus
in the first place as it is similar to the holotype and has only been
referred to Therizinosauroidea accidentally due to Sues' original plate
caption mistake.
References- Matthew and Brown, 1922. The family Deinodontidae, with notice
of a new genus from the Cretaceous of Alberta. Bulletin of the American Museum
of Natural History. 46(6), 367-385.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus.
American Museum Novitates. 2380, 49 pp.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin
of the AMNH. 147.
Sues, 1978. A new small theropod dinosaur from the Judith River Formation (Campanian)
of Alberta, Canada. Journal of the Linnean Society: Zoology. 62, 381-400.
Currie, 1987a. Bird-like characteristics of the jaws and teeth of troodontid
theropods (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. 7, 72-81.
Currie, 1987b. Theropods of the Judith River Formation of Dinosaur Frovincial
Park, Alberta. In Currie and Koster (eds). 4th Symposium of Mesozoic Terrestrial
Ecosystems Short Papers. Tyrell Museum of Palaeontology, Drumheller, Alberta.
52-60.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press. 107-125.
Currie, 1992. Saurischian dinosaurs of the Late Cretaceous of Asia and North
America. In Mateer and Chen (eds.). Aspects of Nonmarine Cretaceous Geology. China Ocean Press. 237-249.
Fiorillo and Currie, 1994. Theropod teeth from the Judith River Formation (Upper
Cretaceous) of south-central Montana. Journal of Vertebrate Paleontology. 14(1),
74-80.
Currie, 1995. New information on the anatomy and relationships of Dromaeosaurus
albertensis (Dinosauria: Theropoda). Journal of Vertebrate Paleontology.
15(3), 576-591.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Brinkman, Ryan, and Eberth, 1998. The paleogeographic and stratigraphic distribution
of ceratopsids (Ornithischia) in the Upper Judith River Group of western Canada.
Palaios. 13, 160-169.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation
(Cretaceous) of northern Alaska, with speculations on Arctic dinosaur paleoecology.
Journal of Vertebrate Paleontology. 20(4), 675-682.
Ryan, Russell, Eberth and Currie, 2001. The taphonomy of a Centrosaurus
(Ornithischia: Ceratopsidae) bone bed from th Dinosaur Park Formation (Upper
Campanian), Alberta, Canada, with comments on cranial ontogeny. Palaios. 16,
482-506.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Currie, 2005. Theropods, including birds. In Currie and Koppelhus (eds.). Dinosaur
Provincial Park, a spectacular ecosystem revealed. Part Two, Flora and Fauna
from the park. Indiana University Press. 367-397.
Demar and Breithaupt, 2006. The nonmammalian vertebrate microfossil assemblages
of the Mesaverde Formation (Upper Cretaceous, Campanian) of the Wind River and
Bighorn Basin, Wyoming. in Lucas and Sullivan (eds). Late Cretaceous Vertbrates
from the Western Interior. New Mexico Museum of Natural History & Science.
Bulletin 35, 33-53.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
D? sp. (Gangloff, 1998)
Albian-Cenomanian, Early-Late Cretaceous
Chandler Formation, Alaska, US
Material- (AK211-V-001) tooth
Comments- This was originally referred to Alectrosaurus (Gangloff,
1998), and later to Dromaeosaurus albertensis (Fiorillo and Gangloff,
2000) based on strong lateral compression and a lack of blood grooves. However,
it seems to early to belong to D. albertensis itself and is probably
another genus of dromaeosaurine.
References- Gangloff, 1998. Arctic dinosaurs with emphasis on the Cretaceous
record of Alaska and the Eurasian-North American connection. in Lucas, Kirkland
and Estep (eds.). Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico
Museum of Natural History and Science Bulletin. 14, 211-220.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation
(Cretaceous) of northern Alaska, with speculations on Arctic dinosaur paleoecology.
Journal of Vertebrate Paleontology. 20(4), 675-682.
D. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Foremost Formation of the Belly River Group, Alberta, Canada
Material- (RTMP 97.99.4) phalanx
Reference- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press. 279-297.
D. sp. nov. (Peng, Russell and Brinkman, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Belly River Group, Alberta, Canada
Material- (RTMP 96.62.1) tooth (Peng et al., 2001)
(RTMP 96.62.2) tooth (Peng et al., 2001)
(RTMP 96.62.3) tooth (Peng et al., 2001)
(RTMP 96.62.69) tooth (Peng et al., 2001)
(RTMP 96.62.70) tooth (Peng et al., 2001)
(RTMP 96.103.1) tooth (Ryan and Russell, 2001)
four specimens (Brinkman et al., 1998)
Comments- These were referred to D. albertensis by Peng et al.
(2001), but the teeth have less distinctively displaced mesial carinae than
that species.
References- Brinkman, Ryan, and Eberth, 1998. The paleogeographic and
stratigraphic distribution of ceratopsids (Ornithischia) in the Upper Judith
River Group of western Canada. Palaios. 13, 160-169.
Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages (exclusive
of mammals) from the Foremost and Oldman Formations of the Judith River Group
(Campanian) of southeastern Alberta: An illustrated guide. Provincial Museum
of Alberta, Natural History Occasional Paper. 25, 1-546.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
D. spp. nov. (Russell, 1933)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
Material- (RTMP 1010) tooth (Baszio, 1997)
(RTMP 1011) lateral tooth (Baszio, 1997)
(RTMP 1012) tooth (Baszio, 1997)
(RTMP 1013) lateral tooth (Baszio, 1997)
(RTMP 1029) lateral tooth (Baszio, 1997)
(RTMP 1035) tooth (Baszio, 1997)
(RTMP 83.45.3) tooth (11.2x4.8x4 mm) (Larson and Currie, 2013)
(RTMP 85.98.5) tooth (9.3x4.7x3 mm) (Larson and Currie, 2013)
(RTMP 98.63.32) tooth (11.8x7.4x4.1 mm) (Larson et al., 2010)
(RTMP 98.63.71) tooth (6.6x4.6x2 mm) (Larson et al., 2010)
(RTMP 98.64.17) tooth (9.6x6x3.2 mm) (Larson et al., 2010)
(RTMP 98.84.3) tooth (8.9x7.4x3.4 mm) (Larson and Currie, 2013)
(RTMP 99.50.116) tooth (11.7x7.7x4 mm) (Larson et al., 2010)
(RTMP 2000.45.40) tooth (10x6.6x2.8 mm) (Larson et al., 2010)
(RTMP 2000.45.82) tooth (11x6.1x3.4 mm) (Larson et al., 2010)
(RTMP 2000.45.102) tooth (9.6x6.5x2.9 mm) (Larson et al., 2010)
(RTMP 2001.45.83) tooth (16.6x7.9x4.7 mm) (Larson et al., 2010)
(RTMP 2002.45.50) tooth (11.2x6.7x3.3 mm) (Larson et al., 2010)
(RTMP 2003.45.60) tooth (9.6x5.4x2.4 mm) (Larson et al., 2010)
(RTMP 2005.7.5) tooth (8.2x3.6x2.4 mm) (Larson and Currie, 2013)
(RTMP 2009.122.2) tooth (8.8x5.4x3 mm) (Larson and Currie, 2013)
(RTMP 2009.122.4) tooth (9.9x6.6x3.2 mm) (Larson and Currie, 2013)
(RTMP 2009.136.8) tooth (Larson et al., 2010)
(RTMP coll.) two teeth (Baszio, 1997)
tooth (Russell, 1933)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- tooth (Baszio, 1997)
three specimens (Tokaryk and Bryant, 2004)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta
Material- (RTMP 81.31.99) tooth (Ryan and Russell, 2001)
(UA Alberta 1:98) tooth (Baszio, 1997)
(UA Alberta 1:99) tooth (Baszio, 1997)
(UA Alberta 3:103) tooth (Baszio, 1997)
(UA Alberta 3:104) lateral tooth (Baszio, 1997)
(UA JLE 62:113) tooth (Baszio, 1997)
(UA KUA-1:106) lateral tooth (Baszio, 1997)
(UA KUA-1:119) tooth (Baszio, 1997)
(UA KUA-2:110) tooth (Baszio, 1997)
Comments- Baszio (1997) notes these teeth are smaller than Dinosaur Park
specimens and differ in that they are slightly more convex labially than lingually.
Larson and Currie (2013) found that upper Horseshoe Canyon teeth are quantitatively
different from Dinosaur Park teeth, and that lower Horseshoe Canyon teeth are
not but lack the twisted mesial carina present in D. albertensis.
References- Russell, 1933. The Cretaceous-Tertiary transition of Alberta.
Transactions of the Royal Society of Canada, series 3. 26(4), 121-156.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
Tokaryk and Bryant, 2004. The fauna from the Tyrannosaurus rex excavation,
Frenchman Formation (Late Maastrichtian), Saskatchewan. Summary of Investigations
2004, Volume 1. Saskatchewan Geological Survey, Saskatchewan Industry Resources,
Miscellaneous Report 2004-4, 1-12.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe
Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta,
with special reference to the Albertosaurus sarcophagus bonebed. Canadian
Journal of Earth Sciences. 47(9), 1159-1181.
Larson and Currie, 2013. Multivariate analyses of small theropod dinosaur teeth
and implications for paleoecological turnover through time. PloS ONE. 8(1),
e54329.
D. sp. (Mongelli and Varricchio, 1998)
Early Campanian, Late Cretaceous
Lower Two Medicine Formation, Montana, US
Material- teeth
Reference- Mongelli and Varricchio, 1998. Theropod teeth of the Lower
Two Medicine Formation (Campanian) of northwestern Montana. Journal of Vertebrate
Paleontology. 18(3), 64A.
D. sp. (Wilson, 2008)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, South Dakota, US
Material- two teeth (Wilson, 2008)
teeth, elements (DePalma, 2010)
References- Wilson, 2008. Comparative taphonomy and paleoecological reconstruction
of two microvertebrate accumulations from the Late Cretaceous Hell Creek Formation
(Maastrichtian), eastern Montana. Palaios. 23, 289-297.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous
bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis,
University of Kansas. 227 pp.
D. sp. (Stokosa, 2005)
Maastrichtian, Late Cretaceous
Fox Hills Formation, South Dakota, US
Material- (SDSM 14516) tooth fragment (Stokosa, 2005)
Reference- Stokosa, 2005. Enamel microstructure variation within the
Theropoda. in Carpenter (ed). The Carnivorous Dinosaurs. 163-178.
D. sp. (Lillegraven and Eberle, 1999)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
Material- (UW 26254)
(UW 26580)
(UW 27203)
Reference- Lillegraven and Eberle, 1999. Vertebrate faunal changes through
Lancian and Puercan time in southern Wyoming. Journal of Paleontology. 73(4),
691-710.
D. sp. nov. (Breithaupt, 1982)
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Material- ?(AMNH 25268) astragalus (AMNH online)
(UA 125) lateral tooth (Baszio, 1997)
(UA 126) lateral tooth (Baszio, 1997)
(UA 128) lateral tooth (Baszio, 1997)
(YPM PU 22312) (YPM online)
two specimens (Breithaupt, 1982)
?specimen (Derstler, 1995)
Comments- Baszio (1997) notes these teeth are much smaller than D.
albertensis with more (6) serrations per mm.
References- Breithaupt, 1982. Paleontology and paleoecology of the Lance
Formation (Maastrichtian), east flank of Rock Springs Uplift, Sweetwater County,
Wyoming. Contributions to Geology, University of Wyoming. 21(2), 123-151.
Derstler, 1995. The Dragons' Grave: An Edmontosaurus bonebed containing
theropod egg shells and juveniles, Lance Formation (uppermost Cretaceous), Niobrara
County, Wyoming. Journal of Vertebrate Paleontology. 15(3), 26A.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
D. sp. (Williamson, 2001; described by Williamson and Brusatte,
2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
Material- (NMMNH P-33148) anterior tooth (6.5x3.8x2.5 mm) (Williamson
and Brusatte, 2014)
(NMMNH P-33489) tooth (Williamson and Brusatte, 2014)
References- Williamson, 2001. Dinosaurs from microvertebrate sites in
the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New
Mexico. 2001 GSA abstracts.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Dromaeosauroides
Christiansen and Bonde, 2003
D. bornholmensis Christiansen and Bonde, 2003
Late Berriasian, Early Cretaceous
Jydegaard Formation, Denmark
Holotype- (MGUH DK 315) anterior dentary tooth (21.7x9.7x6.6 mm)
Referred- (MGUH DK 559) tooth (15 mm) (Bonde, 2012)
Diagnosis- 25% larger than Dromaeosaurus albertensis; 6 distal
serrations per mm compared to 3-3.5/mm in D. albertensis.
Comments- The holotype was discovered in 2000 and mentioned as Dromaeosauridae
indet. by Bonde (2001). The BW/FABL is .68, while the crown is fairly tall and
recurved. Both carinae are lingually displaced, the mesial carina only extending
down 1/3 of the crown. Mesial serrations decrease in size toard the base, with
6.1/mm present apically. Distal serrations are square, labiolingually broad,
lack blood grooves and have a density of 6/mm.
References- Bonde, 2001. A Berriasian "Wealden fauna" from
Bornholm, Denmark. Palaeontological Association 45th annual meeting. 4.
Christiansen and Bonde, 2003. The first dinosaur from Denmark. Neues Jahrbuch
der Geologie und Palaontologie Abhandlungen. 227(2), 287-299.
Bonde and Christiansen, 2003. New dinosaurs from Denmark. Comptes Rendus Palevol.
2, 13-26.
Bonde, 2012. Danish dinosaurs: A review. In Godefroit (ed.). Bernissart Dinosaurs
and Early Cretaceous Terrestrial Ecosystems. Indiana University Press. 434-451.
Velociraptorinae Barsbold, 1983
Definition- (Velociraptor mongoliensis <- Dromaeosaurus
albertensis) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017; modified from Sereno, 1998)
Other definitions- (Velociraptor mongoliensis <- Microraptor
zhaoianus, Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus)
(Turner et al., 2012)
= Itemirinae Kurzanov, 1976 vide Martyniuk, 2012
= Velociraptorinae sensu Turner et al., 2012
Definition- (Velociraptor mongoliensis <- Microraptor zhaoianus,
Dromaeosaurus albertensis, Unenlagia comahuensis, Passer domesticus)
Comments- Since Turner et al. (2012) uses a stem-based definition for
Dromaeosauridae, the node stem triplet advocated by Padian et al. (1999) no
longer functions. Turner et al.'s definition is slightly superior to Sereno's
1998 one by adding Microraptor, Unenlagia and Passer, but
I think the chance of a (Dromaeosaurus (Velociraptor, Passer))
topology is very low, and wouldn't have a problem with microraptorians being
velociraptorines. But since Unenlagiinae is active, I suppose it's best to keep
subfamilies out of subfamilies.
Kurzanov (1976) erected Itemiridae for Itemirus only, sister to tyrannosaurids
than to dromaeosaurids. Martynuik (2012) both defined Itemiridae as excluding
Dromaeosaurus, and created an undefined Itemirinae within Dromaeosauridae
with no explicit content (besides Itemirus). Here (as in Longrich and
Currie, 2009), Itemirus is recovered as closer to Velociraptor
than to Dromaeosaurus. This would technically make Itemirinae a senior
synonym of Velociraptorinae, as the latter family-level taxon was named 7 years
later. Yet the Itemirus holotype has only been found to have this position
in one published analysis, while it was recovered closer to Dromaeosaurus
by Sues and Averianov (2014). Thus until there is a consensus on the position
of Itemirus, the subfamily Velociraptorinae is used here.
References- Kurzanov, 1976. Braincase structure in the carnosaur Itemirus
n. gen., and some aspects of the cranial anatomy of dinosaurs. Paleontological
Journal. 1976, 361-369.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
from the Late Cretaceous of North America. Proceedings of the National Academy
of Sciences. 106(13), 5002-5007.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Vernon, New Jersey. Pan Aves. 189 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Sues and Averianov, 2014. Dromaeosauridae (Dinosauria: Theropoda) from the Bissekty
Formation (Upper Cretaceous: Turonian) of Uzbekistan and the phylogenetic position
of Itemirus medullaris Kurzanov, 1976. Cretaceous Research. 51, 225-240.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399.
Shri Turner, Montanari and Norell, 2021
= "Ichabodcraniosaurus" Novacek,
1996
S. devi Turner, Montanari and Norell, 2021
Late Campanian, Late Cretaceous
Khulsan, Baron Goyot Formation, Mongolia
Material-
(IGM 100/980) third cervical vertebra (28 mm), fourth cervical vertebra
(28.3 mm), fifth cervical vertebra (30.6 mm), sixth cervical vertebra
(24.2 mm), seventh cervical vertebra (19.55 mm), eighth cervical
vertebra (18.5 mm), ninth cervical vertebra (19.6 mm), tenth cervical
vertebra (19.1 mm), few cervical ribs, first dorsal vertebra (21.5 mm),
second dorsal vertebra (19.5 mm), third dorsal vertebra (17.5 mm),
fourth dorsal vertebra (16.3 mm), fifth dorsal vertebra (17.25 mm),
sixth dorsal vertebra (17.5 mm), seventh dorsal vertebra (17 mm),
eighth dorsal vertebra (16.9 mm), ninth dorsal vertebra, tenth dorsal
vertebra, eleventh dorsal vertebra, twelfth dorsal vertebra, thirteenth
dorsal vertebra (15.1 mm), five proximal dorsal ribs, fragmentary
sacrum, first caudal vertebra (18.8 mm), second caudal vertebra (18.8
mm), third caudal vertebra (20 mm), fourth caudal vertebra (21.05 mm),
fifth caudal vertebra (23.4 mm), sixth caudal vertebra (~21 mm),
seventh caudal vertebra, eighth caudal vertebra, five chevrons, ilia
(183.8 mm), pubes (~245 mm; one partial), ischia (~118 mm), femur (205
mm), incomplete tibiae (246 mm), incomplete fibula, partial astragali,
partial calcaneum, metatarsal I (30.9 mm), phalanx I-1 (22.5 mm), pedal
ungual I, incomplete metatarsal II (~90 mm), phalanx II-1 (25.8 mm),
phalanx II-2, pedal ungual II (91.3 mm on curve), incomplete metatarsal
III (~117 mm), distal phalanx III-1 (~47 mm), phalanx III-2 (30 mm),
phalanx III-3 (28.4 mm), pedal ungual III (49.4 mm on curve),
incomplete metatarsal IV (~124 mm), fragmentary phalanx IV-1 (~41.5
mm), phalanx IV-2 (27.1 mm), phalanx IV-3 (23.4 mm), phalanx IV-4 (22.4
mm), pedal ungual IV (36.5 mm on curve), partial metatarsal V
Diagnosis- (after Turner et al., 2021) first dorsal epipophysis large and overhangs
posterior margin of postzygapophysis; two dorsoventrally aligned pleurocoels
on first and second dorsals; posteriorly inclined scar on lateral surface of
neural arch.
Comments-
This specimen was discovered by Norell in July 1991 and mentioned by
Norell et al. (1992) as "a very well preserved, articulated postcranial
skeleton". Novacek (1996) wrote in a popular book ""It lacked a
precious head. Although Mark [Norell] was understandably upset,
it was still an important find. Informally called "Ichabodcraniosaurus,"
the skeleton remains under investigation in New York to this
day." Norell and Makovicky (1999) mention IGM 100/980 several
times, stating it can be referred to Dromaeosauridae based on caudal
and pedal characters, but that insufficient elements are preserved to
refer it to Velociraptor. A photo
of the pelvis and adjacent vertebrae is included as figure 24. Norell (DML,
2002) confirmed that "Ichabodcraniosaurus" and IGM 100/980 are the
same specimen. Turner (2008) described its anatomy as a new taxon of dromaeosaurid,
though he did not name it. While IGM 100/980 is partially scored in his matrix,
Turner didn't include the specimen in his analysis. It was officially described and named Shri devi by Turner et al. (2021), who included it in a version of the TWiG analysis which recovered it as a velociraptorine sister to Velociraptor.
References- Norell, Clark and Perle, 1992. New dromaeosaur material from
the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 12(3),
45A.
Novacek, 1996. Dinosaurs of the Flaming Cliffs. Anchor Books. 367 pp.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton
II: Information from newly collected specimens of Velociraptor mongoliensis.
American Museum Novitates. 3282, 45 pp.
Norell, DML 2002. https://web.archive.org/web/20210120042909/http://dml.cmnh.org/2002Feb/msg00648.html
Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis.
Columbia University. 666 pp.
Turner, Montanari and Norell, 2021. A new dromaeosaurid from the Late
Cretaceous Khulsan locality of Mongolia. American Museum Novitates.
3965, 46 pp.
Velociraptor Osborn, 1924b
= "Ovoraptor" Osborn, 1924a
Diagnosis- (after Turner et al., 2012; for Velociraptor mongoliensis, at the time including IGM 100/982) premaxilla with long maxillary
process reaching well beyond caudal margin of external nares; first and second
premaxillary teeth larger than third and fourth; anterior border of internal
antorbital fenestra broadly rounded; maxillary fenestra not located in a posteriorly
open depression; frontal long, almost four times longer than wide across orbital
portion, and almost four times as long as parietal; dentary very shallow, its
depth constituting 1/8 to 1/7 of its length, ventral margin convex; flange-like m.
ambiens tubercle located proximally on anterior face of pubis.
Not Velociraptor- Besides the remains listed below, V. osmolskae,
and Bohlin's (1953) and Young's (1958) questionably referred specimens, many
remains have been referred to Velociraptor that probably do not belong
there. In 1982, Carpenter stated that he and Paul (in prep.) would show Velociraptor
occurred in North America. Though the collaborative publication never surfaced,
Paul (1984, 1988, 1988) did synonymize Deinonychus and Saurornitholestes
with Velociraptor. Recent analyses indicate these do not form a monophyletic
clade with respect to Dromaeosaurus however (Currie and Varricchio, 2004;
Senter, 2007), and thus almost all non-microraptorian, non-unenlagiine dromaeosaurids
would be a single genus under this scheme. Still, this has led to the occassional
identification of Velociraptor in American sediments (eg. AMNH website). Dong et al. (1989) state Velociraptor material was discovered in the Iren Dabasu Formation of Inner Mongolia, China, and Dong (1992) specifies "teeth of Velociraptor". Currie and Eberth (1993) state "Isolated dromaeosaurid teeth and bones are common in
the Iren Dabasu" and that "Most of these can be attributed to Velociraptor..." Yet no
rationale was presented, and the only two specified Iren Dabasu
dromaeosaurid elements in the literature are clearly not Velociraptor (AMNH 6572 is twice the size,
while IVPP 270790-4 is different from most dromaeosaurid teeth).
Matsukawa and Obata (1994) report through personal communication with Mateer
(1992) that cf. Velociraptor mongoliensis was discovered in the Zouyun
Formation, though this is much too early to actually be that genus. Norell et
al. (1994) identified two juvenile skulls as Velociraptor, but these
seem more likely to be Byronosaurus (Norell and Makovicky, 1999). Nessov
(1995) notes that unguals attributed to Velociraptor are known in the
Santonian Syuk-Syuk Formation of Kazakhstan, but these reports have not been
verified. The Tsaagan holotype was originally referred to Velociraptor
(e.g. Webster, 1996) before it was identified as a new genus by Norell et al.
(2006). By 2004 a skull cast was in circulation from Gaston Design Inc., generally referred to Velociraptor (e.g. Witmer, online 2012), but Powers et al. (2020) used maxillary proportions to find it is closer to Tsaagan. Turner et al. (2007) describe an ulna as IGM 100/981, referring it to
Velociraptor mongoliensis based on sharing several unlisted
characteristics, but this specimen is actually IGM 100/3503 (Napoli et al., 2021) and is here placed in Tsaagan.
References- Osborn, 1924a. The discovery of an unknown continent. Natural
History. 24(2), 133-149.
Osborn, 1924b. Three new Theropoda, Protoceratops zone, central Mongolia.
American Museum Novitates. 144, 1-12.
Bohlin, 1953. Fossil reptiles from Mongolia and Kansu. Sino-Swedish
Expedition Publication. 37, 1-105.
Young, 1958. The first record of dinosaurian remains from Shansi. Vertebrata
PalAsiatica. 2(4), 231-236.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek
formations and a description of a new species of theropod. Contributions to
Geology, University of Wyoming. 20(2), 123-134.
Paul, 1984. The archosaurs: A phylogenetic study. In Reif and Westphal (eds.).
Third Symposium on Mesozoic Terrestrial Ecosystems. Tubingen. 175-180.
Paul, 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide.
Simon and Schuster, New York. 464 pp.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods
of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus
- and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River
- Deinonychus, Velociraptor and Saurornitholestes. Hunteria.
2(4), 1-9.
Dong, Currie and Russell, 1989. The 1988 field program of The Dinosaur Project. Vertebrata PalAsiatica. 27(3), 233-236.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Currie and Eberth, 1993. Palaeontology, sedimentology and palaeoecology of the
Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia, People s Republic
of China. Cretaceous Research. 14, 127-144.
Matsukawa and Obata, 1994. Cretaceous, a contribution to dinosaur facies in
Asia based on molluscan paleontology and stratigraphy. Cretaceous Research.
15, 101-125.
Norell, Clark, Dashzeveg, Barsbold, Chiappe, Davidson, McKenna and Novacek,
1994. A theropod dinosaur embryo, and the affinities of the Flaming Cliffs dinosaur
eggs. Science 266, 779-782.
Nessov, 1995. Dinosaurs of Northern Eurasia: new data about assemblages, ecology
and paleobiogeography. Scientific Research Institute of the Earth's Crust, St.
Petersburg State University, St. Petersburg, Russia. 156 pp. + 14 pl. [in Russian
with short English, German, and French abstracts].
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton
II: Information from newly collected specimens of Velociraptor mongoliensis.
American Museum Novitates. 3282, 45 pp.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation
(Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright
(eds). Feathered Dragons. Studies on the transition from dinosaurs to birds.
Indiana University Press. 112-132.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid
theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545,
51 pp.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 5(4), 429-463.
Turner, Makovicky and Norell, 2007. Feather quill knobs in the dinosaur Velociraptor.
Science. 317, 1721.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Witmer, online 2012. https://people.ohio.edu/witmerl/collections/Theropods/Velociraptor_gaston.htm
Powers, Sullivan and Currie, 2020. Re-examining ratio based
premaxillary and maxillary characters in Eudromaeosauria (Dinosauria :
Theropoda): Divergent trends in snout morphology between Asian and
North American taxa. Palaeogeography, Palaeoclimatology, Palaeoecology.
547, 109704.
Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021. A new
dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central
Mongolia. American Museum Novitates. 3982, 47 pp.
V. mongoliensis Osborn,
1924b
= "Ovoraptor djadochtari" Osborn, 1924a
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Holotype- (AMNH 6515) skull (176 mm), mandibles (175 mm), teeth (4.15-7.91
mm), manual phalanx III-3 (39 mm), manual ungual III (45 mm on curve)
Referred- (IGM coll.; 980826 BDz Velocira ENKH) specimen including dentary,
radius, ulna, semilunate carpal, metacarpal I, distal phalanx I-1,
manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual
ungual II, femur, tibia, fibula and fragments (Suzuki and Watabe, 2000)
Late Campanian, Late Cretaceous
Chimney Buttes, Djadokhta Formation, Mongolia
(IGM 100/986) (~25 kg) cranial fragments, cervical vertebrae, three anterior
dorsal vertebrae, dorsal vertebrae 8-13, dorsal ribs, sacrum, caudal vertebrae
1-26, chevrons, proximal scapulacoracoid, sternal plate, distal humerus, proximal
radius, proximal ulnae, metacarpal I, phalanx I-1 (47.1 mm), manual ungual I,
phalanx II-1 (33.2 mm), manual ungual II (48.8 mm curve), phalanx III-1 (19
mm), phalanx III-2 (11.1 mm), phalanx III-3 (33.2 mm), manual ungual III, ilia
(145.8 mm), pubes (213, 208 mm), ischia (115.1, 118.8 mm), femur (238 mm), tibia
(255 mm), fibula, astragalocalcaneum, distal tarsal III, distal tarsal IV, metatarsal
I (27.9 mm), metatarsal II (84.8 mm), phalanx II-1 (26.6 mm), phalanx II-2 (28
mm), pedal ungual II, metatarsal III (99.1 mm), phalanx III-1 (44 mm), phalanx
III-2 (27.4 mm), phalanx III-3 (12.2 mm), metatarsal IV (91.6 mm), phalanx IV-1
(30.1 mm), phalanx IV-2 (23 mm), phalanx IV-3 (18.6 mm), phalanx IV-4 (19.6,
18.6 mm), pedal ungual IV (25.6 mm straight), metatarsal V (40.1 mm) (Norell
and Mackovicky, 1999)
Late Campanian, Late Cretaceous
Tugrikin Shire, Djadokhta Formation, Mongolia
(IGM 100/24) incomplete skull, mandibles, teeth (7.35-9.44 mm), few fragmentary
postcranial elements (Barsbold, 1983)
(IGM 100/25; fighting dinosaurs specimen) (2.07 m, 15 kg) skull (230
mm), mandibles, cervical vertebrae 1-10, cervical ribs, dorsal
vertebrae 1-13, dorsal ribs, five uncinate processes, gastralia,
sacrum, caudal vertebrae 1-43, chevrons, scapulae, coracoids, sternal
plates (95 mm), humeri (~147 mm), radii, ulnae, semilunate carpal,
metacarpal I (25 mm), phalanx I-1 (44 mm), manual ungual I (47 mm),
metacarpal II (58 mm), phalanx II-1 (35 mm), phalanx II-2 (48 mm),
manual ungual II (~51 mm), metacarpal III (50 mm), phalanx III-1 (20
mm), phalanx III-2 (12 mm), phalanx III-3 (35 mm), manual ungual III
(33 mm), ilium (~188 mm), pubis, ischium, femur (208 mm), tibia (243
mm), fibula, metatarsus (101 mm), pes including phalanx II-1 (21 mm),
phalanx II-2 (24 mm), pedal ungual II (52 mm) (Kielan-Jaworowska and
Barsbold, 1972; described by Barsbold, 1974)
(IGM 100/54; 940728 TS-V NAR) (subadult) skull (~230 mm), mandibles,
first-ninth cervical vertebrae, twelve dorsal vertebrae, sixteen dorsal
ribs, five uncinate processes, gastralia, sacrum, first-third caudal
vertebrae, scapula, partial coracoids, sternal plates, humerus, radius,
ulna, semilunate carpal, metacarpal I (26 mm), phalanx I-1 (49 mm),
manual ungual I (46 mm), metacarpal II (56 mm), phalanx II-1 (37 mm),
phalanx II-2 (52 mm), manual ungual II (~37 mm), metacarpal III (50
mm), phalanx III-1 (20 mm), phalanx III-2 (14 mm), phalanx III-3 (36
mm), manual ungual III (30 mm), ilia, proximal pubis, femora (192, 194
mm), tibiae, fibulae, astragalocalcanea, metatarsal I, proximal pedal
ungual I, distal tarsal III fused with metatarsals II and metatarsals
III (one incomplete), phalanx II-1, phalanges II-2, pedal ungual II,
phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III,
metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3,
phalanges IV-4, pedal unguals IV, metatarsal V (Watabe and Suzuki, 2000)
(IGM 100/976) (adult) partial skull including premaxillae, incomplete
maxillae, partial nasals, frontals, partial parietals, braincase,
incomplete mandible, atlas, axis, third cervical vertebra, fourth
cervical vertebra, seventh cervical vertebra (23.6 mm), eighth cervical
vertebra, ninth cervical vertebra (26.3 mm), cervical ribs, third
dorsal vertebra (15.2 mm), scapulae, coracoids, furcula, sternal
plates, sternal ribs, proximal humeri (Norell et al., 1992)
(IGM 100/985) (subadult) skull fragments, vertebral fragments, dorsal rib fragments,
twelve rows of gastralia, sacrum (89.2 mm), first caudal vertebra (17 mm), first
chevron (33.6 mm), second caudal vertebra (19.1 mm), third caudal vertebra (20.3
mm), fourth caudal vertebra (23.4 mm), fifth caudal vertebra (24.5 mm), sixth
caudal vertebra (24.1 mm), sixth chevron (26.4 mm), seventh caudal vertebra
(23.9 mm), eighth caudal vertebra (25.9 mm), sternal plates (71 mm), proximal
humeri, manual elements including unguals, ilia (126.8, 132.8 mm), pubis (167
mm), ischia (97.7 mm), femur, partial tibia, partial fibula, partial astragalus,
distal tarsal III, metatarsi I (22, 21.4 mm), phalanges I-1 (16.9 mm), pedal
ungual I (20.3 mm), metatarsi II (71, 71.2 mm), phalanges II-1 (23.6 mm), phalanges
II-2 (24.2, 24 mm), pedal ungual II (64.8 mm on curve), metatarsi III (85.6,
86.2 mm), phalanges III-1 (37.6 mm), phalanges III-2 (24.7, 24.9 mm), pedal
ungual III (32.3 mm), metatarsi IV (79, 78.2 mm), phalanges IV-1 (26.8 mm),
phalanges IV-2 (21, 21.3 mm), phalanges IV-3 (17, 16.8 mm), phalanges IV-4 (16,
16.1 mm), pedal unguals IV (32.3, 31 mm on curve), metatarsal V (Norell and
Mackovicky, 1997)
(PIN 3143/8; Moscow specimen) incomplete skull, mandible (Barsbold and Osmolska, 1999)
Diagnosis- (after Kundrat, 2004) differs from V. sp.
nov. in- triangle-shaped auricular fossa; the utriculo-saccular and
anterior semicircular canal prominence does not project too far
medially; hollowed area behind posterior internal acoustic fossa
absent; minor expansion of medulla oblongata either anteriorly between
the roots of III-VI and VII, and ventrolaterally below the metotic
fissure; endoneurocranial base not clearly corrugated longitudinally;
transverse convexity of endoneurocranial base at anteriormost point of
metotic fissure absent; dorsal tympanic recess less expanded
anteriorly; absence of crista intertuberalis; shallow parabasisphenoid
recess is separated by low median ridge; paracondylar recesses
projected ventrally; foramen magnum considerably compressed
dorsoventrally.
(after Turner et al., 2012) supratemporal fossa and
fenestra subcircular, bound by laterally convex supratemporal arcade (unknown in IGM 100/982); lateral
wall of braincase possesses a deep prootic recess (unknown in IGM 100/982); V-shaped
furcula with reduced and asymmetrically developed hypocledium (unknown in IGM 100/982); well-developed
obturator tuberosity (unknown in IGM 100/982); rounded longitudinal ischial ridge (also in Deinonychus; unknown in IGM 100/982).
Comments- The holotype was discovered on August 11 1923 and originally
named Ovoraptor djadochtari by Osborn (1924a) in a magazine article. Osborn merely describes it as "a wonderfully alert little
egg-snatcher, hence Ovoraptor of the Djadochta formation", a "swift-moving, carnivorous dinosaur." However, this publication
lacked a proper description, and Osborn (1924b) decided to name the taxon Velociraptor
mongoliensis in his official description later that year. The skull was
redescribed by Sues in 1977.
IGM 100/25 was discovered in early August 1971 (Kielan-Jaworowska
and Barsbold, 1972), initially mentioned as "the skeleton of a Velociraptor mongoliensis preserved together with the skeleton of Protoceratops andrewsi"
and photographed as plate II figure 2. It was described briefly
by Barsbold (1974), with the skull, sterna, pedal digit II illustrated
by him in 1983. Now known as the "fighting dinosaurs" specimen, it has
been analyzed by several authors (Osmolska, 1993; Unwin et al., 1995;
Carpenter, 2000). Barsbold and Osmolska (1999) described the
skull in detail, while Currie et al. (2016) provide manual measurements
for it. Paul (1984, 1988) first noticed unicinate processes, which were
described in detail by Codd (2004).
Barsbold (1983) also mentioned another specimen, IGM 100/24, a
"partially destroyed skull and remains of postcranial skeleton". He
illustrates the medial mandible. Barsbold and Osmolska (1999)
note it was discovered by a Mongolian expedition, and illustrate the
skull and braincase.
PIN 3143/8 was discovered at Tugrikin Shire by the Soviet-Mongolian
expedition, who also found material at Ikh Shunkht (Ivakhnenko and
Kurzanov, 1988 misstyped as 1982 in Watabe et al., 2010).
Barsbold and Osmolska (1999) state the postcrania of IGM 100/24 and 100/25 "will be published at a later date (Barsbold &
Osmolska in preparation)."
IGM 100/976 was discovered in 1991 and mentioned by Norell et al.
(1992). Norell et al. (1997) briefly described the furcula, while the
other postcrania were described in depth by Norell and Makovicky
(1999), the braincase was described by Norell et al. (2004) and the
endocast by King et al. (2020). IGM 100/985 and 100/986 were discovered
in 1993, and described by Norell and Makovicky in 1997 and 1999
respectively. Though IGM 100/985 was originally only referred to
Dromaeosauridae indet., it was referred to Velociraptor specifically by 1999.
Three Velociraptor specimens
were listed by Watabe and Suzuki (2000) as being from Tugrikin Shire
found between July 21 and August 1 1994. The best described is
IGM 100/54, with the field number 940728 TS-V NAR from Tugrikin Shire
V, photographed by them as "complete skeleton (fore limb shown) of Velociraptor
in Tugrikin Shire". This is complete except the distal tail and
left forelimb, with the skull and mandible photographed as plate 1
figure 3 in Tsogtbaatar (2004), the skeleton as figure 2 in that
publication and in higher resolution in Saneyoshi et al. (2011).
Hone et al. (2012) and Codd (2004) provide good photographs of the
torso, while Hattori (2016) describes and illustrates a
metatarsus. Hone et al. described a pterosaur (?azhdarchid) long
bone fragment eaten by the specimen, while Saneyoshi et al. described
taphonomy. Jasinoski et al. (2006) used this specimen (listed as
IGM 94.07.28) for pectoral and forelimb information. Currie et
al. (2016) lists manual measurements for MPC-D unnumbered specimen,
which are here guessed to be IGM 100/54 given the <10% length
difference using the un-ideal scale in Senoyoshi et al.'s photo (scale
bar inserted digitally over an oblique photograph).
A specimen collected in August 26 1998 and reported by Suzuki and Watabe (2000), "Velociraptor forelimbs" with field number 980826 BDz Velocira ENKH discovered at Bayn Dzak, was listed as "Velociraptor lower jaw and partial postcranial bones." The in situ specimen 980826 BDz Velocira ENKH is figured as "Velociraptor skeleton in Bayn Dzak (central Gobi region)."
References- Osborn, 1924a. The discovery of an unknown continent. Natural
History. 24(2), 133-149.
Osborn, 1924b. Three new Theropoda, Protoceratops zone, central Mongolia.
American Museum Novitates. 144, 1-12.
Kielan-Jaworowska and Barsbold, 1972. Narrative of the Polish-Mongolian Palaeontological
Expeditions, 1967-1972. Palaeontologia Polonica. 27, 1-12.
Barsbold, 1974. Dueling dinosaurs. Priroda. 2, 81-83. [in Russian]
Sues, 1977. The skull of Velociraptor mongoliensis, a small Cretaceous
theropod dinosaur from Mongolia. Palontologische Zeitschrift. 51, 173-184.
Barsbold, 1983. Avian features in the morphology of predatory dinosaurs. Transactions
of the Joint Soviet Mongolian Paleontological Expedition. 24, 96-103.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions
of the Joint Soviet-Mongolian Paleontological Expedition. 19, 5-119.
Paul, 1984. The archosaurs: A phylogenetic study. In Reif and Westphal (eds.).
Third Symposium on Mesozoic Terrestrial Ecosystems. Tubingen. 175-180.
Ivakhnenko and Kurzanov, 1988. Geological structure and age of
Udan-Sair and Shara-Tsav localities. Trudy Sovmestnoi
Sovetsko-Mongolʼkoi Paleontologicheskoi Ekspeditsii. 34, 100105.
Paul, 1988. Predatory Dinosaurs of the World: A Complete Illustrated Guide.
Simon and Schuster, New York. 464 pp.
Paul, 1988. The small predatory dinosaurs of the mid-Mesozoic: the horned theropods
of the Morrison and Great Oolite - Ornitholestes and Proceratosaurus
- and the sickle-claw theropods of the Cloverly, Djadokhta and Judith River
- Deinonychus, Velociraptor and Saurornitholestes. Hunteria.
2(4), 1-9.
Norell, Clark and Perle, 1992. New dromaeosaur material from the Late Cretaceous
of Mongolia. Journal of Vertebrate Paleontology. 12(3), 45A.
Osmolska, 1993. Were the Mongolian "fighting dinosaurs" really fighting?
Revue de Paleobiologie. 7, 161-162.
Unwin, Perle and Trueman, 1995. Protoceratops and Velociraptor
preserved in association: Evidence from predatory behavior in predatory dinosaurs?
Journal of Vertebrate Paleontology. 15(3), 57A.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton:
Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Norell, Makovicky and Clark, 1997. A Velociraptor wishbone. Nature. 389,
447.
Feduccia and Martin, 1998. Theropod-bird link reconsidered. Nature. 391, 754.
Norell and Makovicky, 1998. A revised look at the osteology of dromaeosaurs:
evidence from new specimens of Velociraptor. Journal of Vertebrate Paleontology.
18(3), 66A.
Barsbold and Osm�lska, 1999. The skull of Velociraptor (Theropoda)
from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica. 44(2),
189-219.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton
II: Information from newly collected specimens of Velociraptor mongoliensis.
American Museum Novitates. 3282, 45 pp.
Carpenter, 2000. Evidence of predatory behavior by carnivorous dinosaurs. GAIA.
15, 135-144.
Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1998. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 83-98.
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 30-44.
Smith, 2002. An examination of dental morphology and variation in theropod dinosaurs:
Implications for the identification of shed teeth. PhD dissertation. University
of Pennsylvania.
Codd, 2004. The uncinate processes in birds and their implications for the breathing
mechanics of maniraptoran dinosaurs. PhD thesis, Rheinischen Friedrich-Wilhelms-Universit�t. 108 pp.
Kundrat, 2004. Two morphotypes of the Velociraptor neurocranium. ICVM-7 Abstracts. Journal
of Morphology. 260(3), 305.
Norell, Makovicky and Clark, 2004. The braincase of Velociraptor. In
Currie, Koppelhus, Shugar and Wright (eds.). Feathered Dragons: Studies on the
Transition from Dinosaurs to Birds. Indiana University Press. 133-143.
Tsogtbaatar, 2004. Fossil specimens prepared in Mongolian Paleontological Center 1993-2001. Hayashibara Museum of Natural
Sciences Research Bulletin. 2, 123-128.
Parsons and Parsons, 2005. A comparison of postcranial features found within
the ontogenies of the maniraptoran theropod dinosaurs Deinonychus antirrhopus
(Saurischia, Theropoda) and Velociraptor mongoliensis (Saurischia, Theropoda).
Journal of Vertebrate Paleontology. 25(3), 99A.
Jasinowski, Russell and Currie, 2006. An integrative phylogenetic and extrapolatory
approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder
musculature. Zoological Journal of the Linnean Society. 146, 301-344.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid
theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545,
51 pp.
Garcia, Erickson, Curry Rogers and Norell, 2007. Longevity and growth patterns
in the dromaeosaurid Velociraptor mongoliensis inferred from long bone
histology. Journal of Vertebrate Paleontology. 27(3), 79A.
Gombert, 2008. Craniofacial ontogeny in Velociraptor mongoliensis. Journal
of Vertebrate Paleontology. 28(3), 85A.
Watabe,
Tsogtbaatar, Suzuki and Saneyoshi, 2010. Geology of
dinosaur-fossil-bearing localities (Jurassic and Cretaceous: Mesozoic)
in the Gobi Desert: Results of the HMNS-MPC Joint Paleontological
Expedition. Hayashibara Museum of Natural Sciences Research Bulletin.
3, 41-118.
Saneyoshi, Watabe, Suzuki and Tsogtbaatar, 2011. Trace fossils on
dinosaur bones from Upper Cretaceous eolian deposits in Mongolia:
Taphonomic interpretation of paleoecosystems in ancient desert
environments. Palaeogeography, Palaeoclimatology, Palaeoecology. 311,
38-47.
Hone, Tsuihiji, Watabe and Tsogtbaatar, 2012. Pterosaurs as a food
source for small dromaeosaurs. Palaeogeography, Palaeoclimatology,
Palaeoecology. 331-332, 27-30.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Barsbold, 2016. "The Fighting Dinosaurs": The position of their bodies
before and after death. Paleontological Journal. 50(12), 1412-1417.
Currie, Funston and Osm�lska, 2016. New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
King, Sipla, Georgi, Balanoff and Neenan, 2020. The endocranium and trophic ecology of Velociraptor mongoliensis. Journal of Anatomy. 237(5), 861-869.
V. sp. nov. (Norell and Makovicky, 1997)
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Material- (IGM 100/982) (adult) incomplete skull (200 mm), mandible
(190 mm), five or six incomplete cervical vertebrae, cervical ribs,
four incomplete anterior dorsal vertebrae, two partial posterior dorsal
vertebrae, partial dorsal ribs, gastralia, synsacrum, incomplete
first-sixth caudal vertebrae, proximal chevrons, incomplete scapulae,
incomplete coracoids, partial furcula, partial sternal plates,
incomplete humeri (119 mm), incomplete radii, incomplete ulnae,
metacarpal I (19.2 mm), phalanx I-1 (39.9 mm), manual ungual I (30.2 mm
straight), metacarpal II (50.8 mm), phalanx II-1 (31.3 mm), phalanx
II-2 (45.8 mm), manual ungual II (33 mm), metacarpal III (45 mm),
phalanx III-1 (17.6 mm), phalanx III-2 (10.1 mm), phalanx III-3 (32.2
mm), manual ungual III (25 mm), incomplete ilia (131.7, 124.6 mm),
incomplete pubes (132 mm), ischial fragment, incomplete femora (184
mm), incomplete tibiae (182 mm), incomplete fibulae,
astragalocalcaneum, distal tarsal IV, metatarsal I (22.5 mm), phalanges
I-1 (19.8, 19.3 mm), pedal ungual I (16 mm straight), metatarsals II
(73.7 mm), phalanges II-1 (23.9 mm), phalanges II-2 (26.1 mm), pedal
unguals II (66.3 mm curve), metatarsals III (88.1 mm), phalanges III-1
(39.6 mm), phalanx III-2 (24.7 mm), phalanx III-1 fragment, metatarsals
IV (83.4 mm), phalanx IV-1 (27.1 mm), phalanx IV-2 (21 mm), proximal
phalanx IV-3, metatarsal V
Diagnosis- (after Norell and Makovicky, 1999) vertical ridge dorsal to acetabulum on ilium.
(after Powers et al., 2020) elongate antorbital fossa.
(after Powers et al., 2021a) anteriorly abbreviated cerebellar fossa
with elongate olfactory canal; lacrimal notch of the frontal that is
more vertically oriented as opposed to transversely in V. mongoliensis.
(after Powers et al., 2022) maxillary fenestra within a broad, shallow
maxillary fossa that extends anterior to the maxillary fenestra.
Comments- IGM 100/982 was
discovered in 1995 and first mentioned by Norell and Makovicky (1997)
as an undescribed dromaeosaur specimen collected by the MAS-AMNH
expeditions possessing a wide flat brevis shelf on its ilium. The
postcranium was described by Norell and Makovicky in 1999, while the
braincase was described by Norell et al. (2004). The skull was
first figured in figure 6C of Norell et al. (2006). Norell and
Makovicky referred it "to Velociraptor mongoliensis based on
comparisons of cranial material with the holotype (AMNH 6515)",
although even this early on they did note "A poorly defined vertical
crest above the acetabulum that divides the anterior and posterior
muscle fossae on the lateral surface of the pelvis is present on IGM
100/982 but is absent on other specimens." In an abstract, Kundrat
(2004) noted several braincase characters which differed between it and
IGM 100/976, concluding "IGM 100/982 represents a more advanced
velociraptorine morphotype than does IGM 100/976" although he wrongly
thought it was 100/982 that was confirmed to be V. mongoliensis and
100/976 that "might be referable to a new velociraptorine taxon."
Powers (2020) names and describes the new species in his thesis, which
also has sections which have been published since. Powers et al.
(2020) discovered that using maxillary morphometrics "MPC-D 100/982
plotted far away from all other specimens referred to the genus
Velociraptor, the separation being greater than that seen among many
specimens representing different species" and states "A description of
this specimen and review of variation within the genus Velociraptor is
currently underway." Similarly, Powers et al. (2022)
provides CT data on the maxilla and mentions it as "a possible third
[species of Velociraptor] based on a currently undescribed specimen
(MPC-D 100/982, description in prep.)", although as noted above many
aspects of the specimen have been described in the literature.
Two abstracts devoted to the new species were published in 2021 (Powers
et al., 2021a, b).
References-
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton:
Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton
II: Information from newly collected specimens of Velociraptor mongoliensis.
American Museum Novitates. 3282, 45 pp.
Kundrat, 2004. Two morphotypes of the Velociraptor neurocranium. ICVM-7 Abstracts. Journal
of Morphology. 260(3), 305.
Norell, Makovicky and Clark, 2004. The braincase of Velociraptor. In
Currie, Koppelhus, Shugar and Wright (eds.). Feathered Dragons: Studies on the
Transition from Dinosaurs to Birds. Indiana University Press. 133-143.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid
theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545,
51 pp.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian
physiology inherited by birds? Reconciling slow growth in Archaeopteryx.
PLoS ONE. 4(10), e7390.
Powers, 2020. The evolution of snout shape in eudromaeosaurians and its
ecological significance. Masters thesis, University of Alberta. 437 pp.
Powers, Sullivan and Currie, 2020. Re-examining ratio based
premaxillary and maxillary characters in Eudromaeosauria (Dinosauria :
Theropoda): Divergent trends in snout morphology between Asian and
North American taxa. Palaeogeography, Palaeoclimatology, Palaeoecology.
547, 109704.
Powers, Norell and Currie, 2021a. New shallow snouted species of Velociraptor sheds light on intraspecific variation in Velociraptor mongoliensis
and possible niche partitioning between species. 9th Annual Meeting
Canadian Society of Vertebrate Palaeontology Online Abstracts. 31.
Powers, Norell and Currie, 2021b. Examination of morphological variation across Velociraptor mongoliensis specimens reveals a new species with possible ecomorphological variation in snout dimensions. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 210-211.
Powers, Fabbri, Doschak, Bhullar, Evans, Norell and Currie, 2022 (as
2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist
in testing and constructing morphological characters. Journal of
Vertebrate Paleontology. 41(5), 2010087.
V. sp. indet. (Bohlin, 1953)
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Material- ?(AMNH 6518) partial maxilla, distal tarsal III, distal tarsal
IV, metatarsal I, phalanx I-1 (16.2 mm), pedal ungual I, incomplete metatarsal
II, phalanx II-1 (28 mm), phalanx II-2 (36 mm), pedal ungual II (56 mm), incomplete
metatarsal III (~250 mm), phalanx III-?, metatarsal IV, phalanx IV-?, metatarsal
V (40.1 mm) (Ostrom, 1969)
numerous teeth (9-14 mm) (Barsbold and Osmolska, 1999)
Late Campanian, Late Cretaceous
Tugrikin Shire, Djadokhta Formation, Mongolia
?(IGM 100/2000; 940725 TS-V OTGN or 940801 TS-I WTB) (juvenile) skull, mandible, postcranium (Barsbold and Osmolska,
1999)
?(IGM coll.; 940725 TS-V OTGN or 940801 TS-I WTB) skeleton (Watabe and Suzuki, 2000)
?(IGM coll.; 980725 TS-I Velocira NAR) dorsal vertebrae, ribs, caudal series, partial pelvis, hindlimb (Suzuki and Watabe, 2000)
Late Campanian, Late Cretaceous
Udyn Sayr, Djadokhta Formation, Mongolia
?(uncollected?) forelimbs (Suzuki and Watabe, 2000)
?(uncollected?) teeth (Watabe and Tsogtbaatar, 2004)
Late Campanian, Late Cretaceous
Ikh Shunkht, Baron Goyot Formation, Mongolia
?(IGM or PIN coll.) (Ivakhnenko and Kurzanov, 1988)
Late Campanian, Late Cretaceous
Khulsan, Baron Goyot Formation, Mongolia
?(ZPAL MgD-I/97) partial skull, partial mandibles, distal tibia, astragalus,
metatarsal I, phalanx I-1, pedal ungual I, metatarsus (~80 mm), phalanx II-1,
phalanx II-2, pedal ungual II, phalanx III-1, phalanx III-2, phalanx III-3,
phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV (Osmolska,
1982)
? several incomplete skeletons (Osmolska, 1982)
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
?(IVPP V16138) premaxillary tooth, incomplete lateral tooth (Hone, Choiniere, Sullivan, Xu, Pittman and Tan, 2010)
Campanian-Maastrichtian, Late Cretaceous
Minhe Formation, Inner Mongolia, China
? two teeth (Bohlin, 1953)
Late Cretaceous
Tzoyun, Shanxi, China
?(IVPP V965) tooth (Young, 1958)
Late Cretaceous?
Mongolia?
?(IGM 100/200) third premaxillary tooth (6.17x4.03x1.63 mm), partial maxilla, maxillary teeth (7.35-9.93 mm) (Smith, 2002)
?(University of Manchester LL.12392) manual ungual (Johnson et al., 2009)
Comments- These are specimens referred to Velociraptor which have not been described, figured or preserved well enough to refer to either V. mongoliensis or V. sp. nov.. Indeed, basically none of them are complete enough or have been described well enough to even refer to Velociraptor, and could easily be Tsaagan, Shri, Kuru or another eudromaeosaur.
Bohlin (1953) referred two teeth, and more questionably a penultimate
phalanx and ungual, to Velociraptor mongoliensis (mispelled V. mongoliense).
The teeth are indeed probably dromaeosaurid, and roughly similar to Velociraptor.
While they lack mesial serrations like the geographically close V. osmolskae,
V. mongoliensis teeth often lack mesial serrations as well (perhaps due
to wear in some cases). They are thus indeterminate at least within Velociraptor,
and probably within basal Dromaeosauridae. The phalanx is said to be broader
than Velociraptor's III-3 and the illustrated ungual is clearly a parvicursorine
manual ungual I however.
Young (1958) referred IVPP V965 to cf. Velociraptor mongoliensis,
although stated it is "somewhat larger than the type". As it is based
on a single tooth, referral to any particular dromaeosaurid species is
questionable.
AMNH 6518 was discovered on August 11 1923, but not published until
Ostrom (1969) who illustrated several phalanges. Images of the
entire specimen are available at the AMNH online catalogue.
ZPAL MgD-I/97 was found between 1963 and 1971 on the Polish-Mongolian
expeditions, and is mentioned briefly by Osmolska (1982) in her description of
Hulsanpes.
The crania are figured by Barsbold and Osmolska (1999), who say the
postcrania "will be published at a later date (Barsbold &
Osmolska in preparation)."
IGM 100/2000
was listed as "the complete skeleton of a young individual. It includes
skull with mandible and postcranium" by Barsbold and Osmolska (1999),
who also wrote it was found "by the Mongolian-Japanaese
Palaeontological Expeditions" at Tugrikin Shire. It is possibly
the "Velociraptor
skull and small skeleton" reported by Watabe and SuzukI (2000).
Those authors list 940725 TS-V OTGN and 940801 TS-I WTB as "Velociraptor skeleton" from Tugrikin Shire V and "Velociraptor
skeleton" from Tugrikin Shire I respectively, but which specimen became
IGM 100/2000 is unknown and no description or illustration of either
has been published.
Suzuki and Watabe (2000) reported two new Velociraptor specimens collected in 1998. "The postcranial skeleton of Velociraptor"
with field number 980725 TS-I Velocira NAR discovered on July 25 at
Tugrikin Shire I, described as consisting of "dorsal vertebrae, ribs,
pelvic part, hind limb, and complete tail"; and "Velociraptor forelimbs" discovered between August 16-23 at Udyn Sayr.
Smith (2002) uses IGM 100/200 as one of his Velociraptor
dental specimens used to measure tooth shape in his gigantic database
of theropods, which includes a premaxillary tooth and a maxillary
fragment that includes at least six tooth positions.
Watabe and Tsogtbaatar (2004) report "teeth of Velociraptor" discovered on July 23 2000 at Udyn Sayr.
Hone et al. (2010) describe two teeth found in 2008-2009 from the Wulansuhai Formation of Inner Mongolia as Velociraptor, associated with a cf. Protoceratops skeleton exhibiting tooth-marked bones. These teeth have mesial serrations, so are unlikely to be Tsaagan or "Velociraptor" osmolskae which are both known from this formation. Yet no characters are described that would refer these to Velociraptor over another dromaeosaurid with mesial serrations and a high DSDI, so they are only tentatively referred here.
References-
Bohlin, 1953. Fossil reptiles from Mongolia and Kansu. Sino-Swedish
Expedition Publication. 37, 1-105.
Young, 1958. The first record of dinosaurian remains from
Shansi. Vertebrata PalAsiatica. 2(4), 231-236.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod
from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural
History. 30, 165 pp.
Osmolska, 1982. Hulsanpes perlei n. g. n. sp. (Deinonychosauria, Saurischia,
Dinosauria) from the Upper Cretaceous Barun Goyot Formation of Mongolia. Neues
Jahrbuch fur Geologie und Palaeontologie, Monatshefte. 1982(7), 440-448.
Ivakhnenko and Kurzanov, 1988. Geological structure and age of
Udan-Sair and Shara-Tsav localities. Trudy Sovmestnoi
Sovetsko-Mongolʼkoi Paleontologicheskoi Ekspeditsii. 34, 100105.
Barsbold and Osm�lska, 1999. The skull of Velociraptor (Theropoda)
from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica. 44(2),
189-219.
Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1998. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 83-98.
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 30-44.
Smith, 2002. An examination of dental morphology and variation in theropod dinosaurs:
Implications for the identification of shed teeth. PhD dissertation. University
of Pennsylvania.
Watabe and Tsogtbaatar, 2004. Report on the Japan - Mongolia Joint Paleontological Expedition to the
Gobi desert, 2000. Hayashibara Museum of Natural Sciences Research
Bulletin. 2, 45-67.
Johnson, Mustansar, Manning, Margetts and Mummery, 2009. Virtual repair of fossil
CT scan data. Journal of Vertebrate Paleontology. 29(3), 123A.
Hone, Choiniere, Sullivan, Xu, Pittman and Tan, 2010. New evidence for a trophic relationship between the dinosaurs Velociraptor and Protoceratops. Palaeogeography, Palaeoclimatology, Palaeoecology. 291, 488-492.
Luanchuanraptor Lu, Xu,
Zhang, Ji, Jia, Hu, Zhang and Wu, 2007
L. henanensis Lu, Xu, Zhang, Ji, Jia, Hu, Zhang and Wu, 2007
Late Cretaceous
Qiupa Formation, Henan, China
Holotype- (41HIII-0100) incomplete frontal, premaxillary tooth, three lateral
teeth, posterior cervical vertebra, anterior cervical rib, posterior cervical
rib, first dorsal vertebra, anterior dorsal rib, two proximal caudal vertebrae,
two middle caudal vertebrae, thirteen distal caudal vertebrae, three proximal
chevrons, middle chevron, two distal chevrons, scapulocoracoid, humerus, manual
phalanx I-1, incomplete manual ungual, ilium, pubis, femoral shaft, fragments
Diagnosis- (after Lu et al., 2007) distal caudal prezygapophyses equal
to centra in length; distal caudal vertebrae with neural spine or dorsal groove;
elongated proximodorsal processes on distal chevrons; elongate median posterior
process on distal chevrons (also in some Deinonychus chevrons); deep
concavity present on medial coracoid; deltopectoral crest extends over half
of humerus.
Comments- High resolution photos are available in Xu's (2007) thesis,
as compared to the low quality of Lu et al.'s (2007) officially published pdf.
This taxon has yet to be included in a published cladistic analysis, but when
run in a version of the TWiG matrix, it emerges as the sister group to Velociraptor.
The plesiomorphic characters may suggest a more basal placement, however.
References- Lu, Xu, Zhang, Ji, Jia, Hu, Zhang and Wu, 2007. New dromaeosaurid
dinosaur from the Late Cretaceous Qiupa Formation of Luanchuan area, western
Henan, China. Geological Bulletin of China. 26(7), 777-786.
Xu, 2007. The scientific characters and protection of dinosaurs from Henan Province.
China University of Geosciences. 120 pp.
Daurlong Wang, Cau, Guo, Ma, Qing and Liu, 2022
D. wangi Wang, Cau, Guo, Ma, Qing and Liu, 2022
Early Aptian, Early Cretaceous
Pigeon Hill, Longjiang Formation, Inner Mongolia, China
Holotype- (IMMNH-PV00731) (1.5
m) skull (170 mm), sclerotic ring, mandible, ten cervical vertebrae,
twelve dorsal vertebrae, dorsal ribs, gastralia, synsacrum, caudal
series (800 mm), chevrons, scapulocoracoids (one partial; scapula 112
mm), sternum (75 mm), four sternal ribs, incomplete humerus (120 mm),
radius (90 mm), ulna (96 mm), scapholunare, semilunate carpal,
metacarpal I (20 mm), phalanges I-1 (one incomplete; 36 mm), manual
unguals I (30 mm), metacarpal II (50 mm), phalanges II-1 (one
incomplete; 30 mm), phalanges II-2 (43 mm), manual unguals II (37 mm),
metacarpal III (48 mm), phalanges III-1 (one incomplete; 17 mm),
phalanges III-2 (10 mm), phalanges III-3 (20 mm), manual unguals III
(17 mm), partial ilium (~115 mm), incomplete pubis (~147 mm), ischium
(~92 mm), femora (one incomplete; 180 mm), tibiae (198 mm), incomplete
fibula, incomplete metatarsal III (110 mm), phalanx III-1 (35 mm),
phalanges III-2 (30 mm), phalanges III-3 (23 mm), pedal unguals III (30
mm), partial metatarsal IV (95 mm), phalanx IV-1 (30 mm), phalanx IV-2
(20 mm), phalanx IV-3 (20 mm), phalanges IV-4 (13 mm), pedal unguals IV
(25 mm), pedal claw sheaths, body feathers, intestine
Diagnosis- (after Wang et al.,
2022) large, trapezoid promaxillary fenestra placed at anteroventral
corner of antorbital fossa; maxillary fenestra large, shallow and
posterodorsally located, so that the pila promaxillaris is wider than
the pila interfenestralis; fan-shaped distal end of first sternal rib.
Comments- Discovered between
2018 and 2019 (Cau, pers. comm. 4-23), Wang et al. (2023) included this
in a version of Cau's megamatrix and recovered it sister to
"Zhenyuanlong", then Tianyuraptor
in a clade sister to Eudromaeosauria. Added to Hartman et al.'s
maniraptoromorph matrix it resolves as a velociraptorine closest to Kuru and Adasaurus.
Reference- Wang, Cau, Guo, Ma,
Qing and Liu, 2022. Intestinal preservation in a birdlike dinosaur
supports conservatism in digestive canal evolution among theropods.
Scientific Reports. 12:19965.
Kuru Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021
= "Airakoraptor" Perle, Norell and Clark, 1999
K. kulla Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021
Late Campanian, Late Cretaceous
Khulsan, Baron Goyot Formation, Mongolia
Holotype-
(IGM 100/981) premaxilla, partial lacrimal, incomplete dentary,
incomplete surangular, three lateral teeth, incomplete ~third/fourth
cervical vertebra, partial ~fourth/fifth cervical vertebra, fragmentary
~fifth/sixth cervical vertebra, tenth cervical vertebra, incomplete
first dorsal vertebra, second dorsal vertebra, third dorsal vertebra,
incomplete fourth dorsal vertebra, partial fifth dorsal vertebra, five
incomplete posterior dorsal vertebrae, ~sixth caudal vertebra, two
distal caudal vertebrae (one lost), proximal humerus, distal humeri,
distal radius, proximal ulna, semilunate carpal, metacarpal I, phalanx
?I-1, incomplete manual ungual I, proximal metacarpal II, phalanx II-2,
manual ungual II, proximal metacarpal III, phalanx ?III-1, phalanx
?III-2, posterior ilium, partial pubes, femora (one incomplete; ~220
mm), proximal tibia, partial astragalus, metatarsal I, pedal ungual I,
partial metatarsals II, phalanx II-1, phalanges II-2, partial pedal
unguals II, partial metatarsal III, phalanges III-1 (one incomplete),
phalanx III-2, partial pedal ungual III, incomplete metatarsal IV (~113
mm), phalanx IV-1, five pedal phalanges
Diagnosis- (after Napoli et
al., 2021) deeply incised groove framing anterior and ventral margins
of premaxillary narial fossa; lacrimal with posterolateral hornlet
arising from posterodorsal process; lacrimal with poorly developed boss
at intersection of ventral, posterodorsal, and anterior processes;
dentary lacking ventral secondary row of nutrient foramina; dentary
nutrient foramina set in shallow groove posteriorly; two posterior
surangular foramina; proportionally deep surangular; dentary teeth
serrated on both mesial and distal carinae; anteriorly displaced dorsal
pleurocoels; dorsal hyposphenes widely separated and joined by web of
bone; distal tarsals unfused to metatarsus; mediolaterally narrow
metatarsal II; reduced pedal digit II.
Comments- While the description merely states that in the manus
"most phalanges are not confidently assignable", the reconstruction in
Ruebenstahl et al.'s (2021) SVP presentation shows phalanges I-1, III-3
and III-2 outlined.
"Airakoraptor" was originally listed in
a bibliographic entry in Perle et al.'s (1999) description of Achillobator.
This supposed paper is listed as authored by Norell, Clark and Perle in
a 1992 issue of the Journal of Vertebrate Paleontology. No pages
numbers or issue are listed, though it does say "Vol. 11 Toronto.
Canada", which indicates it refers to an abstract from the Toronto
Society of Vertebrate paleontology meeting in 1992 (volume 11 covered
the 1991 meeting in San Diego, so must be a mistake). There is an
abstract in that issue by those authors, but it has a different title
which does not mention "Airakoraptor" by name. Instead of "Morphology
dromaeosaurian dinosaur- Airakoraptor from the Upper Cretaceous of
Mongolia", it is simply "New dromaeosaur material from the Late
Cretaceous of Mongolia". So Perle et al.'s listing must refer to an
earlier version of the abstract. Napoli et al. (2021) confirm that of
the specimens noted in the abstract, IGM 100/981 was "familiar
(erroneously) under the provisional name "Airakoraptor," and that "the
name is a nomen nudum."
IGM 100/981 was initially mentioned by Norell et al. (1992) as "a
partial skeleton including portions of the skull, mandible, vertebral
column, manus and pes" and said to "possess features unknown in described
dromaeosaurs" such as a deep mandible and dual posterior
surangular foramina. This specimen is also probably that described by
Novacek (1996) as discovered by Perle on July 4th - "a fine skeleton of a
dromaeosaur, perhaps not Velociraptor
but something rare, and new", since Shri was found by Norell. It was later mentioned
as an "unnamed dromaeosaur from Khulsan collected by the Mongolian American
Expeditions" by Norell and Makovicky (1997), stated to have a wide
and flat brevis shelf and share an anterior tubercle on metatarsal III with
Velociraptor and Deinonychus. In Norell and Makovicky (1999) it
is noted to lack a fourth trochanter, unlike Velociraptor.
They state it "can be referred to the Dromaeosauridae on the basis of
pedal and tail morphology, but insufficient elements are present to
refer these specimens to Velociraptor mongoliensis."
Norell et al. (2006) illustrate the surangular as "an undescribed
dromaeosaur from Khulson (IGM 100/981)" and it clearly shows dual
surangular foramina, matching figure 6A of Napoli et al. (2021) who
finally described the specimen as Kuru kulla. Confusingly, Turner et
al. (2007) described an ulna of a supposed Velociraptor
specimen they called IGM 100/981. This specimen was said to be from the
Gilvent Wash locality near Ukhaa Tolgod, and Napoli et al. state the
specimen number was erroneous and is actually IGM 100/3503 which is
here referred to Tsaagan.
References-
Norell, Clark and Perle, 1992. New dromaeosaur material from the Late Cretaceous
of Mongolia. Journal of Vertebrate Paleontology. 12(3), 45A.
"Norell, Clark and Perle, 1992. Morphology dromaeosaurian dinosaur- Airakoraptor
from the Upper Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 11.
Toronto, Canada." [not a real paper]
Novacek, 1996. Dinosaurs of the Flaming Cliffs. Anchor Books. 367 pp.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton:
Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton
II: Information from newly collected specimens of Velociraptor mongoliensis.
American Museum Novitates. 3282, 45 pp.
Perle, Norell and Clark, 1999. A new maniraptoran Theropod - Achillobator
giganticus (Dromaeosauridae) - from the Upper Cretaceous of Burkhant, Mongolia.
Contribution no. 101 of the Mongolian-American Paleontological Project. 1-105.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A new dromaeosaurid
theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545,
51 pp.
Turner, Makovicky and Norell, 2007. Feather quill knobs in the dinosaur Velociraptor.
Science. 317, 1721.
Napoli, Ruebenstahl, Bhullar, Turner and Norell, 2021. A new
dromaeosaurid (Dinosauria: Coelurosauria) from Khulsan, central
Mongolia. American Museum Novitates. 3982, 47 pp.
Ruebenstahl, Napoli, Bhullar, Turner and Norell, 2021. Two new
eudromaeosaurs from Khulsan (central Mongolia) reveal modern-like
faunal predatory structure amoung non-avian dinosaurs. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 222-223.
Turner, Montanari and Norell, 2021. A new dromaeosaurid from the Late
Cretaceous Khulsan locality of Mongolia. American Museum Novitates.
3965, 46 pp.
Adasaurus Barsbold, 1983
= "Adasaurus" Barsbold, 1977
A. mongoliensis Barsbold, 1983
= Dromaeosaurus mongoliensis (Barsbold, 1983) Paul, 1988
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
Holotype-
(IGM 100/20) (~2.7 m) (adult) posterior skull, posterior mandibles,
atlantal neurapophyses, atlantal intercentrum, axis, third cervical
vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth
cervical vertebra, seventh cervical vertebra, eighth cervical vertebra,
ninth cervical vertebra (38 mm), tenth cervical vertebra (35 mm), first
dorsal vertebra (31 mm), second dorsal vertebra (31 mm), third dorsal
vertebra (28 mm), fourth dorsal vertebra (28 mm), fifth dorsal vertebra
(27 mm), sixth dorsal vertebra (27 mm), seventh dorsal vertebra (27
mm), eighth dorsal vertebra (27 mm), ninth dorsal vertebra (27 mm),
tenth dorsal vertebra (26 mm), eleventh dorsal vertebra (27 mm),
synsacrum (217 mm; 29, 31, ?, ?, 31, 29 mm), proximal caudal vertebra
(36 mm), sixth-eleventh caudal vertebrae (39, 42, 42, 43, 43, 43 mm),
six chevrons, incomplete scapulacoracoid, fragmentary sternal plate,
incomplete ilia, incomplete pubes, proximal ischium, femora (267, 263
mm), tibiae (one partial, one incomplete; 296 mm), fibulae, incomplete
astragalocalcaneum, fused distal tarsals III and IV, incomplete
metatarsus (III 146, IV 130 mm), phalanx III-1, phalanx IV-1 (40 mm),
phalanx IV-2 (31 mm), phalanx IV-3 (26 mm), phalanx IV-4 (23 mm), pedal
ungual IV, metatarsal V
Paratype- (IGM 100/21) (~2.7 m) two mid caudal vertebrae, metatarsal I (34
mm), phalanx I-1 (27 mm), pedal ungual I, tarsometatarsus
(II 122, III 145, IV 130 mm), phalanx II-1 (34 mm), phalanx II-2 (32 mm),
phalanx III-1 (55 mm), phalanx III-2 (36 mm), phalanx III-3, pedal
ungual III, phalanx IV-1 (39 mm), phalanx IV-2 (30 mm), phalanx
IV-3 (25 mm), phalanx IV-4 (32 mm), pedal ungual IV
Diagnosis- (after Kubota and Barsbold, 2006) dorsoventrally expanded
anterior process of jugal; low dorsal ridge on median frontals (continuous with
sagittal crest on parietals); untwisted paroccipital process; large surangular
foramen; pleurocoels only on anterior sacrals; preacetabular process strongly
notched anteriorly.
(after Turner et al., 2012) ventral process of lacrimal strongly curved anteriorly;
dorsally displaced triangular process along lateral edge of quadrate shaft.
(proposed) metatarsal II reduced in width.
Other diagnoses- Contra Barsbold (1983) and most later references, the
supposed small pedal ungual II does not belong to digit II (Senter,
2010).
Comments- This genus was discovered between 1964 and 1966, and first mentioned by Barsbold (1977) as a "latest
(Maastrichtian) Mongolian Dromaeosauridae" with a reduced pedal ungual
II, with the pelvis illustrated and labeled as Adasaurus. This does not
count as a proper description however, due to the lack of a type species or
holotype. Osmolska (1980) lists one dromaeosaurid taxon and individual from the Nemegt Formation (as ?Velociraptor sp.), which may be based on Adasaurus
and may be the source of Glut's (1982) reference to "a possible
dromaeosaurid, mentioned by Osmolska (1978), from the Nemegt Formation
of Mongolia." Note no Osmolska reference from 1978 exists and Adasaurus
was already listed by Glut on page 40 however. The taxon was
later described extremely briefly by Barsbold (1983) and officially
named Adasaurus mongoliensis, though the paratype was mistyped as IGM
100/51. While remaining poorly described and virtually unillustrated in the
literature for decades due to an embargo, Adasaurus
has finally been partially illustrated and described by Turner et al.
(2012) with a full description in Kubota's (2015) thesis. Although
Currie and Varricchio (2004) referred snout and forelimb remains to Adasaurus based on IGM 100/22
and 100/23, Kubota and Barsbold (2007) have determined these belong to a new
taxon from the earlier Bayanshiree Formation.
The holotype specimen was approximately 2.7 meters long if the femur is scaled
from Velociraptor. It was from an old individual, so this is probably
close to the maximum size Adasaurus got. Contra Norell and Makovicky
(1997), the specimen is not pathological (Turner et al., 2012).
Jasinowski et al. (2006) list a scapulocoracoid and sternal plate for
the holotype. The pelvis was illustrated by Barsbold (1983) after the
initial schematic drawing in Barsbold (1977), but this is quite
different from the actual material as it is a composite with the
holotype's ilium and pubis and IGM 100/22's ischium.. The femoral
morphology was briefly noted by Perle et al. (1999) and Kim et al.
(2005). The latter also illustrate the proximal femur in two views.
While Adasaurus
is perhaps most famous due to its supposedly reduced second pedal ungual, Senter
(2010 as a 2006 pers. comm. from Kubota) notes that ungual does not actually
pertain to the holotype's digit II.
Relationships- Adasaurus was originally identified as a dromaeosaurid
(Barsbold, 1977) and later specified to be a dromaeosaurine (Barsbold, 1983;
Paul, 1988). More recently, Longrich
and Currie (2009) found it to be a velociraptorine, which was also found by
Kubota and Barsbold (2006, 2007), Senter et al. (2012), Turner et al. (2012),
Brusatte et al. (2014) and Foth et al. (2014). There is certainly no reason
to synonymize Adasaurus with Dromaeosaurus as Paul (1988) did,
as many other dromaeosaurid genera seem to be more closely related to Dromaeosaurus.
References- Barsbold, 1977. [On the evolution of the carnivorous dinosaurs].
Transactions of the Joint Soviet Mongolian Paleontological Expedition. 4, 48-56.
Osmolska, 1980. The Late Cretaceous vertebrate assemblages of the Gobi
Desert, Mongolia. Memoires de la Societe Geologique de France. 139,
145-150.
Glut, 1982. The New Dinosaur Dictionary. Citadel Press. 288 pp.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions
of the Joint Soviet-Mongolian Palaeontological Expedition. 19, 117 pp.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster, New York.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton:
Information from a new specimen. American Museum Novitates. 3215, 1-28.
Perle, Norell and Clark, 1999. A new maniraptoran Theropod - Achillobator
giganticus (Dromaeosauridae) - from the Upper Cretaceous of Burkhant, Mongolia.
Contribution no. 101 of the Mongolian-American Paleontological Project. 1-105.
Currie and Varricchio, 2004. A new dromaeosaurid from the Horseshoe Canyon Formation
(Upper Cretaceous) of Alberta, Canada. in Currie, Koppelhus, Shugar and Wright
(eds). Feathered Dragons. Studies on the transition from dinosaurs to birds.
Indiana University Press. 112-132.
Kim, Gishlick and Tsuihiji, 2005. The first non-avian maniraptoran skeletal
remains from the Lower Cretaceous of Korea. Cretaceous Research. 26, 299-306.
Jasinowski, Russell and Currie, 2006. An integrative phylogenetic and extrapolatory
approach to the reconstruction of dromaeosaur (Theropoda: Eumaniraptora) shoulder
musculature. Zoological Journal of the Linnean Society. 146, 301-344.
Kubota and Barsbold, 2006. Reexamination of Adasaurus mongoliensis (Dinosauria:
Theropoda) from the Upper Cretaceous Nemegt Formation of Mongolia. Journal of
Vertebrate Paleontology. 26(3), 88A.
Kubota and Barsbold, 2007. New dromaeosaurid (Dinosauria Theropoda) from the
Upper Cretaceous Bayanshiree Formation of Mongolia. Journal of Vertebrate Paleontology.
27(3), 102A.
Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis.
Columbia University. 666 pp.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
from the Late Cretaceous of North America. Proceedings of the National Academy
of Sciences. 106(13), 5002-5007.
Senter, 2010. Using creation science to demonstrate evolution: Application of
a creationist method for visualizing gaps in the fossil record to a phylogenetic
study of coelurosaurian dinosaurs. Journal of Evolutionary Biology. 23(8), 1732-1743.
Nesbitt, Clarke, Turner and Norell, 2011. A small alvarezsaurid from the Eastern
Gobi Desert offers insight into evolutionary patterns in the Alvarezsauroidea.
Journal of Vertebrate Paleontology. 31(1), 144-153.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Brusatte, Vremir, Csiki-Sava, Turner, Watanabe, Erickson and Norell, 2013. The
osteology of Balaur bondoc, an island-dwelling dromaeosaurid (Dinosauria:
Theropoda) from the Late Cretaceous of Romania. Bulletin of the American Museum
of Natural History. 374, 1-100.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Kubota, 2015. Descriptions of Mongolian dromaeosaurids (Dinosauria:
Theropoda) and phylogeny of Dromaeosauridae. PhD thesis, University of
Tsubaka. 397 pp.
"Koreanosaurus" Kim,
1979
"K. koreanensis" (Kim, 1993) new comb.
= Deinonychus "koreanensis" Kim, 1993
Aptian-Early Albian, Early Cretaceous
Lower Gugyedong Formation, South Korea
Material- (DGBU-78) femur (~400 mm)
Comments- Originally referred to the Deinodontidae (Kim, 1979), and after
that Hypsilophodontidae or Hadrosauridae (Kim, 1983). By 1986, Kim had decided
it was perhaps synonymous with Deinonychus (pers. comm. to Olshevsky).
In 1993, the name Deinonychus "koreanensis" was used in a photo
caption and faunal list. Lee et al. (2001) remove the specimen from Deinonychus,
based on the presence of a fourth trochanter. However, Deinonychus varies
in this character. Kim et al. (2005) refer the specimen to Eumaniraptora based
on a proximolateral ridge, shelf-like posterior trochanter, and absence of an
accessory trochanter and mediodistal crest. The presence of a large fourth trochanter
was noted to be similar to Adasaurus and Velociraptor.
References- Kim, 1979. [Dinosaur and volcano discovered from Tabri, Euiseong,
Korea].
Kim, 1983. Cretaceous dinosaurs from Korea. Journal of the Geological Society
of Korea. 19(3), 115-126.
Kim, 1993. Journal of Natural History and Environments Vol 1 #1, June 1993.
Published by the World Society of Natural History and Environments, Pusan University,
Pusan, Korea. ISSN 1225-6404.
Lee, Yu and Wood. 2001. A review of vertebrate faunas from the Gyeongsang Supergroup
(Cretaceous) in South Korea. Palaeogeography, Palaeoclimatology, Palaeoecology.
165, 357-373.
Kim, Gishlick and Tsuihiji, 2005. The first non-avian maniraptoran skeletal
remains from the Lower Cretaceous of Korea. Cretaceous Research. 26, 299-306.
unnamed clade (Deinonychus antirrhopus + Utahraptor ostrommaysi)
= Megaraptora Miller, 2004 (non Benson, Carrano and Brusatte,
2009)
Comments- This clade is based on Hartman et al.'s maniraptoromorph analysis, notable for excluding Dromaeosaurus unlike e.g. Senter's coelurosaur analyses. Miller (2004) had previously suggested a pairing of Deinonychus and Utahraptor, which he called Megaraptora despite it not including Megaraptor which was named six years earlier and which he had to be aware of. Benson et al.'s Megaraptora defined to include Megaraptor wasn't proposed until five years later, but is used here based on being universally accepted and including the eponymous genus.
References- Miller, 2004. A new phylogeny of the Dromaeosauridae. 2004 Student Showcase Journal. 20, 123-158.
Benson, Carrano and Brusatte, 2010 (2009 online). A new clade of archaic large-bodied predatory
dinosaurs (Theropoda: Allosauroidea) that survived to the latest Mesozoic. Naturwissenschaften.
97(1), 71-78.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Pneumatoraptor Osi, Apesteguia
and Kowalewski, 2010
P. fodori Osi, Apesteguia and Kowalewski, 2010
Santonian, Late Cretaceous
Csehbanya Formation, Hungary
Holotype- (MTM V.2008.38.1) (adult) scapulocoracoid
Diagnosis- (after Osi et al., 2010) large, circular pneumatic foramen
ventral to the coracoid foramen (2 mm in diameter) that opens towards the coracoid
tubercle and is also in connection with the hollow scapular blade.
Comments- While only assigned to Paraves by Osi et al. (2010), unpublished
analyses using TWiG characters suggest Pneumatoraptor may be a dromaeosaurid.
Reference- Osi, Apesteguia and Kowalewski, 2010. Non-avian theropod dinosaurs
from the early Late Cretaceous of Central Europe. Cretaceous Research. 31(3),
304-320.
Yixianosaurus Xu and Wang,
2003
Y. longimanus Xu and Wang, 2003
Late Valanginian-Early Aptian, Early Cretaceous
Lujiatun or Jianshangou Beds of the Yixian Formation, Liaoning, China
Holotype- (IVPP V12638) (subadult or adult) (~1.3-2 kg) four partial dorsal
ribs, seven gastralia or sternal ribs, furcula, scapulae (65 mm), coracoids,
humeri (89 mm), radii (63 mm), ulnae (62, 64 mm), scapholunare, semilunate
carpal, distal carpal III, metacarpal I (14.5 mm), phalanx I-1 (33, 34 mm), manual ungual I (26,
25 mm), metacarpal II (36, 35 mm), phalanx II-1 (26, 25 mm), phalanx II-2 (38,
36 mm), manual ungual II (28 mm), metacarpal III (34 mm), phalanx III-1 (~9
mm), phalanx III-2 (8 mm), phalanx III-3 (22 mm), manual ungual III (22.5, 23
mm), manual claw sheaths, fragments, feathers
Comments- This specimen was discovered in 2001 and described in 2003
as a maniraptoran perhaps most closely related to scansoriopterygids. This was
based on the elongate manus (compared to humerus) and phalanx II-2 (compared
to metacarpal II). These are correlated with each other however, as an elongate
phalanx II-2 will lead to a long manus. In addition to scansoriopterygids, tyrannosauroids,
ornithomimosaurs, troodontids, and some dromaeosaurids, basal avialans and basal
coelurosaurs have similarly elongate II-2 (compared to II-1). Dececchi et al.
(2010) restudied the specimen for an SVP abstract and while they were secretive
regarding its phylogenetic position, do indicate it is not a paravian. Dececchi
et al. (2012) later published a full paper, noting the presence of a furcula,
and finding it to be a maniraptoran in a polytomy with alvarezsaurids, therizinosaurs
and pennaraptorans. Xu et al. (2013) replied, describing misinterpretations
of anatomy by Dececchi et al., including the supposed intermedium being a broken
part of the semilunate carpal. They noted Dececchi et al. coded Yixianosaurus
for many characters that could not be determined from the specimen, and their
recoded matrix found it to be a deinonychosaur outside of Unenlagiinae, Eudromaeosauria
and Troodontidae. Foth et al. (2014) recovered the genus as a basal paravian.
References- Xu and Wang, 2003. A new maniraptoran dinosaur from the Early
Cretaceous Yixian Formation of Western Liaoning. Vertebrata PalAsiatica. 41(3),
195-202.
Dececchi, Hone, Sullivan and Xu, 2010. A re-analysis of the "coeluriasaurian
pit-bull" Yixianosaurus longimanus with implications for the theropod
dinosaur diversity of the Jehol Biota. Journal of Vertebrate Paleontology. Program
and Abstracts 2010, 81A.
Dececchi, Larsson and Hone, 2012. Yixianosaurus longimanus (Theropoda:
Dinosauria) and its bearing on the evolution of Maniraptora and ecology of the
Jehol fauna. Vertebrata PalAsiatica. 59(2), 111-139.
Xu, Sullivan and Wang, 2013. The systematic position of the enigmatic theropod
dinosaur Yixianosaurus longimanus. Vertebrata PalAsiatica. 51(3), 169-183.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Deinonychus Ostrom, 1969a
= "Daptosaurus" Brown vide Chure and McIntosh, 1989
D. antirrhopus Ostrom, 1969a
= Velociraptor antirrhopus (Ostrom, 1969a) Paul, 1988
= "Daptosaurus agilis" Brown vide Chure and McIntosh, 1989
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Montana, US
Holotype- (YPM 5205) distal tarsal III, distal tarsal IV, metatarsal
I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal
ungual II, metatarsal III (146.5 mm), phalanx III-1, phalanx III-2, phalanx
III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx
IV-3, pedal ungual IV
?...
(YPM 5203) seventh caudal vertebra (45.5 mm), eighth caudal vertebra (47.6 mm),
ninth caudal vertebra (50.2 mm), eleventh caudal vertebra (53 mm), twelfth caudal
vertebra (53.1 mm), thirteenth caudal vertebra (53.2 mm), fourteenth caudal
vertebra (53 mm), fifteenth caudal vertebra (~53.5 mm), sixteenth caudal vertebra
(52.6 mm), seventeenth caudal vertebra (51.7 mm), eighteenth caudal vertebra
(50 mm), nineteenth caudal vertebra (50.2 mm), twentieth caudal vertebra (48.3
mm), twenty-first caudal vertebra (47.5 mm), twenty-second caudal vertebra (46.9
mm), twenty-third caudal vertebra (45.2 mm), twenty-fourth caudal vertebra (42.8
mm), twenty-fifth caudal vertebra (42 mm), twenty-sixth caudal vertebra (41.5
mm), twenty-eighth caudal vertebra (37.8 mm), twenty-ninth caudal vertebra (37.5
mm), thirtieth caudal vertebra (33.6 mm), thirty-first caudal vertebra (32.1
mm), thirty-second caudal vertebra (32.3 mm), thirty-third caudal vertebra (28.7
mm), thirty-fourth caudal vertebra (24.8 mm), thirty-fifth caudal vertebra (23.3
mm), thirty-sixth caudal vertebra (20.1 mm), chevrons
?...
(YPM 5206) radii (one partial) (176.5 mm), ulna, metacarpal I (45.8, 43.2 mm),
phalanx I-1 (74.1 mm), manual ungual I, metacarpal II (93.7, ~88.3 mm), phalanx
II-1 (54 mm), phalanx II-2 (76.5 mm), manual ungual II, metacarpal III (82 mm),
phalanx III-1 (29.9, 30.6 mm), phalanx III-2 (20.5 mm), manual ungual III
?...
(YPM 5210) (partial skull 364 mm) jugal, postorbital, quadratojugals,
squamosals, ectopterygoid, pterygoids, vomer, dentary, splenial, surangular,
angular, prearticulars, articular, 39 teeth, atlas intercentrum, partial atlantic
neural arch, axis, seventh cervical vertebra (41.8 mm), tenth cervical vertebra
(29.5 mm), cervical ribs, ninth dorsal vertebra (30 mm), fourth caudal vertebra
(39.6 mm)
Paratypes-
(YPM 5201) thirteenth caudal vertebra (48.3 mm), fifteenth caudal vertebra (46.6
mm), sixteenth caudal vertebra (46.4 mm), seventeenth caudal vertebra (47.5
mm), eighteenth caudal vertebra (46 mm), nineteenth caudal vertebra (45 mm),
twentieth caudal vertebra (41.6 mm), twenty-first caudal vertebra (42.6 mm),
twenty-second caudal vertebra (43.2 mm), twenty-third caudal vertebra (~41.6
mm), twenty-fourth caudal vertebra (~40 mm), twenty-fifth caudal vertebra (39.4
mm), twenty-sixth caudal vertebra (~35 mm), twenty-eighth caudal vertebra (35
mm), twenty-ninth caudal vertebra (34.3 mm), chevrons
(YPM 5202) ~twenty caudal vertebrae, chevrons
(YPM 5204) partial atlas neural arch and odontoid, axis (31.5 mm), fourth cervical
vertebra (33 mm), fifth cervical vertebra (32 mm), third dorsal vertebra (28.4
mm), fifth dorsal vertebra (28.6 mm), sixth dorsal vertebra (28.5 mm), eighth
dorsal vertebra (28.6 mm)
(YPM 5207) distal tarsal III, distal tarsal IV, phalanx I-1 (33.1 mm), phalanx
III-2 (39 mm), phalanx III-3 (37.4 mm)
(YPM 5208) scapholunare, semilunate carpal
(YPM 5209) manual phalanx II-2 (78 mm), manual phalanx III-3 (52 mm), manual
ungual III
(YPM 5211) semilunate carpal
(YPM 5212) manual ungual I
(YPM 5213) manual phalanx I-1 (72 mm)
(YPM 5214) metacarpal III
(YPM 5215) manual phalanx III-3
(YPM 5216) manual phalanx II-1 (55.3 mm)
(YPM 5217) semilunate carpal, distal tarsal III, metatarsal I (39.1 mm), phalanx
I-1, pedal ungual I, phalanx II-1 (43.3, 42.8 mm), metatarsal III (150.5 mm),
phalanx III-1, phalanx III-3 (35.4 mm), pedal ungual III, phalanx IV-1 (46.5
mm), phalanx IV-2 (33.6 mm), phalanx IV-3, phalanx IV-4 (28 mm), pedal ungual
IV, metatarsal V (69.1 mm)
(YPM 5218) pedal ungual II, phalanx IV-1 (46.6 mm), phalanx IV-2 (36.9 mm),
phalanx IV-3 (27.8 mm), pedal ungual IV
(YPM 5219) pedal ungual I or II
(YPM 5220) radius (172 mm), ulnae (174.2, 180 mm), manual phalanx I-1 (67.2
mm), manual ungual I
(YPM 5221) manual ungual II
(YPM 5222) manual ungual II
(YPM 5223) distal tarsal IV
(YPM 5224) manual ungual I
(YPM 5225) calcaneum
(YPM 5226) astragalus, calcaneum
(YPM 5227) pedal ungual IV
(YPM 5228) scapholunare
(YPM 5229) distal tarsal
(YPM 5230) ulna
(YPM 5231) twelve teeth
(YPM 5232) (3.06 m, 45 kg) partial skull (332 mm) including premaxillae, maxilla,
nasals, lacrimal, jugals, postorbitals, squamosal, quadratojugal, palatine,
vomer, dentaries, angular, articulars, 33 teeth
(YPM 5233) ectopterygoid, pterygoid
(YPM 5234) surangular
(YPM 5235) ischium (161 mm)
(YPM 5236) coracoid
(YPM 5237) splenial
(YPM 5238) splenial
(YPM 5239) pterygoid
(YPM 5240) proximal metatarsal I (45.5 mm), metatarsal I (40 mm)
(YPM 5241) dorsal ribs
(YPM 5242) scapholunare
(YPM 5243) manual phalanx III-3 (48.7 mm), manual ungual III
(YPM 5244) chevron
(YPM 5245) anterior dorsal rib
(YPM 5246) sternal ribs
(YPM 5247) gastralia
(YPM 5248) gastralia
(YPM 5249) mid dorsal rib, posterior dorsal rib
(YPM 5250) dorsal ribs, sternal ribs
(YPM 5251) gastralia
(YPM 5252) gastralia?
(YPM 5253) fragmentary caudal vertebra
(YPM 5254) cervical ribs?
(YPM 5255) rib
(YPM 5256) ribs
(YPM 5257) anterior dorsal rib, posterior dorsal rib
(YPM 5258) dorsal rib
(YPM 5259) two anterior dorsal ribs
(YPM 5260) dorsal rib
(YPM 5261) epipterygoid, gastralia
(YPM 5262) rib
(YPM 5263) metatarsal V
(YPM 5264) ribs, gastralial fragments
(YPM 5265) ribs, gastralial fragments
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Montana, US
(AMNH 3015; intended holotype of "Daptosaurus agilis") (3.06 m, 45 kg, >6 year old adult) incomplete
atlas intercentrum, axial fragment, fourth cervical vertebra, fifth cervical
vertebra (36.7 mm), sixth cervical vertebra (38.5 mm), seventh cervical vertebra
(~39 mm), eighth cervical vertebra (~29 mm), ninth cervical vertebra (~31 mm),
tenth cervical vertebra (~30 mm), (dorsal series ~374.5 mm) first dorsal vertebra
(~31 mm), second dorsal vertebra (~30 mm), third dorsal vertebra (~30 mm), fourth
dorsal vertebra (~30 mm), fifth dorsal vertebra (~27 mm), sixth dorsal vertebra
(~28.5 mm), seventh dorsal vertebra (~27.5 mm), eighth dorsal vertebra (~31.5
mm), ninth dorsal vertebra (~30 mm), tenth dorsal vertebra (~28 mm), eleventh
dorsal vertebra (~27 mm), twelfth dorsal vertebra, thirteenth dorsal vertebra,
gastralia, second caudal vertebra (33.1 mm), third caudal vertebra (37.4 mm),
fourth caudal vertebra (38.3 mm), fifth caudal vertebra (39 mm), sixth caudal
vertebra (41.4 mm), seventh caudal vertebra (43.6 mm), eighth caudal vertebra
(44.5 mm), ninth caudal vertebra (47.6 mm), tenth caudal vertebra (~49.4 mm),
eleventh caudal vertebra (~50.6 mm), twelfth caudal vertebra, thirteenth caudal
vertebra, fourteenth caudal vertebra, fifteenth caudal vertebra (~47.2 mm),
sixteenth caudal vertebra (48 mm), seventeenth caudal vertebra (47.8 mm), eighteenth
caudal vertebra (46.2 mm), nineteenth caudal vertebra (47.1 mm), twentieth caudal
vertebra (46.6 mm), twenty-first caudal vertebra, twenty-second caudal vertebra,
twenty-third caudal vertebra (43.5 mm), twenty-fourth caudal vertebra (43.2
mm), twenty-fifth caudal vertebra (43 mm), twenty-sixth caudal vertebra (41.8
mm), chevrons, scapulae (~190 mm), coracoid fragment, humeri (~237, 227 mm),
radii (~172 mm), ulnae (186), scapholunares, semilunate carpal, metacarpal I (35.5
mm), manual ungual I (90 mm), phalanx II-1 (62.2 mm), phalanx II-2 (70.7 mm),
metacarpal III (73.4 mm), phalanx III-1 (21.7 mm), phalanx III-2 (~15.5 mm),
phalanx III-3 (47.3 mm), ilium (~245 mm), ischia (158, ~161 mm), femur (284
mm), tibia with astragalus (324 mm, 312 without), fibula (~297 mm), astragalus
(62 mm wide, 77 mm high), calcaneum, distal tarsal III, distal tarsal IV, metatarsal
II (129 mm), phalanx II-1 (37.7 mm), phalanx II-2 (42.2 mm), pedal ungual II
(92 mm), metatarsal III (151 mm), phalanx III-1 (52.5 mm), phalanx III-2 (33
mm), phalanx III-3 (28 mm), pedal ungual III (62 mm), metatarsal IV (134 mm),
phalanx IV-1 (44.7 mm), phalanx IV-2 (35.9 mm), phalanx IV-3 (32.2 mm), phalanx
IV-4 (26.3 mm), pedal ungual IV, metatarsal V, eggshells
(AMNH 3037) teeth, manual elements, pedal elements
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Montana, US
Referred-
(AMNH 3041 in part) twenty-five teeth
(Ostrom, 1970)
(AMNH 30832) teeth (AMNH online)
(MCZ 8791) (one year old juvenile; ~7 kg) partial maxilla, articular, partial
?seventh cervical vertebra, five dorsal centra (~eleventh 17.4 mm), three proximal
caudal centra, three mid caudal centra, incomplete coracoids, proximal radius,
partial ulna, phalanx II-2 (56.2 mm), ilial fragment, distal femur, partial
tibia, proximal fibula, proximal phalanx II-1, partial pedal ungual II, phalanx
IV-4 (14.5 mm), gut contents (Parsons and Parsons, 2004; described by Parson
and Parsons, 2015)
(MOR 682) eleven teeth (Maxwell and Ostrom, 1995)
(MOR 747) (at least four individuals) ~320 elements including maxilla, nasal,
lacrimal, jugal, prefrontal, frontal, parietal, quadrate, laterosphenoid, mesethmoid,
braincase, ectopterygoid, palatine, anterior dentary, teeth, other skull material,
cervical vertebra, several caudal vertebrae, radius, ulna, manus, pes (Maxwell
and Witmer, 1996)
(MOR 1178) (~2.32 m) maxillary tooth fragment, squamosal
fragment, (?)ectopterygoid, dentary fragment, axis, sixth cervical vertebra,
anterior dorsal vertebra, mid dorsal vertebra, sacral neural arch fragment,
proximal caudal vertebra, two mid caudal vertebrae, distal caudal vertebra,
scapula, coracoid, sternal(?) fragment, partial humerus, semilunate carpal,
manual ungual I, ilium fragment, partial femora, proximal fibula, tarsal, pedal
phalanx I-1, proximal metatarsal II, pedal ungual II, peal ungual III, phalanx
III-1, metatarsal (?)IV, phalanx IV-4, pedal ungual IV, gut contents (Parsons
and Parsons, 2002)
(MOR coll.) two teeth (Maxwell, 1993)
(YPM 5266) gastralia (Maxwell and Ostrom, 1995)
(YPM 5267) two ?anterior dorsal ribs (Maxwell and Ostrom, 1995)
(YPM 5268) dorsal rib fragments (Maxwell and Ostrom, 1995)
(YPM 5269) dorsal rib fragments (Maxwell and Ostrom, 1995)
(YPM 5270) metacarpal II (~93.5 mm) (Maxwell and Ostrom, 1995)
(YPM 5272) five teeth (Ostrom, 1970)
(YPM 5273) tooth (Ostrom, 1970)
(YPM 5278) tooth (Ostrom, 1970)
(YPM 5279) tooth (Ostrom, 1970)
(YPM 5280) tooth, partial tooth (Ostrom, 1970)
(YPM 5281) caudal vertebra, pedal ungual (Ostrom, 1970)
(YPM 5283) two partial teeth (Ostrom, 1970)
(YPM 5287) caudal vertebra (Ostrom, 1970)
(YPM 5288) four partial dorsal and caudal vertebrae, fragments (Ostrom, 1970)
?(YPM 5289) tooth (Ostrom, 1970)
?(YPM 5291) two tooth fragments (Ostrom, 1970)
(YPM 5397) tooth (Ostrom, 1970)
(YPM 5420) eleven teeth (Ostrom, 1970)
(YPM 5538 in part) proximal fibula, fibular fragment? (YPM online)
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Wyoming, US
(YPM 5271) partial tooth (Ostrom, 1970)
(YPM 5274) tooth (Ostrom, 1970)
?(YPM 5290) cervical centrum (Ostrom, 1970)
(YPM 5356) tooth fragment, fragment (Ostrom, 1970)
(YPM 5371) incomplete tooth (Ostrom, 1970)
(YPM 5376) tooth (Ostrom, 1970)
(YPM 5379) tooth (Ostrom, 1970)
(YPM 5441) two teeth (Ostrom, 1970)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Montana, US
(MCZ 4371; field number 74M #7) (3.43 m, 73 kg) snout fragments, mandibular fragments, teeth, nearly
complete cervical series, nearly complete dorsal series, rib fragments, gastralia
fragments, sacrum, complete caudal series, humerus (254 mm), radius (192 mm),
ulna (208 mm), phalanx I-1 (77.3 mm), manual ungual I, metacarpal II, phalanx
II-1 (64.2 mm), phalanx II-2 (83.4 mm), manual ungual II, metacarpal III (~90
mm), phalanx III-1 (35 mm), phalanx III-2 (23.2 mm), ilia (325 mm), pubes (380+
mm), ischium (175 mm), femora (336 mm), tibia with astragalus (382 mm, 368 without),
metatarsal I (45.6 mm), phalanx I-1 (34.8 mm), pedal ungual I, metatarsal II
(144.3 mm), phalanx II-1 (47 mm), phalanx II-2 (49.9 mm), pedal phalanx II,
metatarsal III (164.4 mm), phalanx III-1 (64.4 mm), phalanx III-2 (44 mm), phalanx
III-3 (41.3 mm), pedal ungual III, metatarsal IV (150.4 mm), phalanx IV-1 (55.5
mm), phalanx IV-2 (41.8 mm), phalanx IV-3 (35.2 mm), phalanx IV-4 (32.6 mm),
pedal ungual IV, metatarsal V (78 mm) (Ostrom, 1976)
Aptian-Early Albian, Early Cretaceous
Little Sheep Mudstone Member, Cloverly Formation, Wyoming, US
(FMNH
2262) (juvenile) premaxillary tooth (?x4.4x? mm; three lateral
teeth (~12x5.5x? mm; 8.1x4.3x? mm; ?x4x? mm) (Gignac, Makovicky,
Erickson and Walsh, 2010)
(YPM 4886) distal metatarsal IV (Ostrom, 1970)
(YPM 4887) distal femur (Ostrom, 1970)
(YPM 5275) tooth (Ostrom, 1970)
Aptian-Albian, Early Cretaceous
Cloverly Formation, Montana, US
(AMNH 21609; = AMNH uncatalogued in Ostrom, 1970 p 209?) 3 teeth (AMNH online)
(MOR 947) frontal (MOR online)
(MOR 1182) (adult) specimen including caudal series, radius and manual phalanx
(Parsons and Parsons, 2006)
(YPM 64996) bone fragments (YPM online)
Early Cretaceous
Arizona, US
? teeth (Thayer and Ratkevich, 1996)
Aptian-Middle Albian, Early Cretaceous
Trinity Group, Texas
(FMNH 2-51#1) premaxillary tooth (9 mm), seven lateral teeth (4, 5, 12.2, 16.2,
21.3, 21.8, 22.5 mm), four tooth fragments (one juvenile) (Gallup, 1975)
(FMNH 3-51#1) eight teeth (4.6, 4.7, 5, 5.6, 6.9, 6.9, 7, 11.9 mm), eight tooth
fragments (Gallup, 1975)
(FMNH 8s-52#1) three teeth (6.1, 12.3, 13.5 mm) (Gallup, 1975)
(FMNH 9s-51#1) two teeth (7.3, 10 mm) (Gallup, 1975)
(FMNH 11s-52#1) three teeth (6.8, 9, 9 mm), tooth fragment (Gallup, 1975)
(FMNH 121-50) tooth (2 mm) (Gallup, 1975)
(FMNH 123-50/124-50) juvenile tooth (8.4 mm), two teeth (13.6, 19.5 mm) (Gallup,
1975)
(FMNH 202-50) tooth (17.9 mm), tooth fragment (Gallup, 1975)
(FMNH 219-50) tooth (Gallup, 1975)
(FMNH 222-50) tooth fragment (Gallup, 1975)
(FMNH Turtle Gully) tooth (8.5 mm), tooth fragment (Gallup, 1975)
(FMNH UvZ) tooth (17 mm), incomplete pedal phalanx III-3 (36 mm) (Gallup, 1975)
(FMNH coll.) dorsal zygapophyseal fragment (Gallup, 1975)
? seven teeth (Brinkman, Cifelli and Czaplewski, 1998)
Cenomanian, Early Cretaceous
Woodbine Formation, Texas, US
Material- ? teeth, limb fragments, manual ungual and/or pedal ungual (Main,
Noto and Scotese, 2011)
? teeth (Bennett et al., 2012)
Aptian-Albian, Early Cretaceous
Antlers Formation, Oklahoma, US
(OMNH 16564) tooth (FABL 6.65 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 16565) tooth (FABL 7.09 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 16566) tooth (FABL 3.64 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 17709) tooth (FABL 9.06 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 34203) tooth (FABL 6.19 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49410) tooth (FABL 9.56 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49411) tooth (FABL 9.24 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49412) tooth (FABL 7.32 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 49415) tooth (FABL 7.26 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 50268) (subadult) two teeth, partial prootic, occiput, two partial mid
dorsal ribs, fifth sacral vertebra (22.5 mm), incomplete proximal chevron, coracoid
(73.27 mm), humeral fragments, partial metacarpals I (31.17, 31.04 mm), partial
phalanges I-1 (~54.04 mm), incomplete manual ungual I, partial metacarpals II,
partial phalanx II-1, partial phalanx II-2, phalanx III-2 (14.56 mm), phalanx
III-3 (39.34 mm), femoral fragments, partial metatarsal I, incomplete phalanx
II-2 (33.7 mm), partial pedal unguals II, partial phalanx III-3, partial phalanx
IV-3 (27.26 mm), phalanx IV-4 (23.94 mm) (Brinkman, Cifelli and Czaplewski,
1998)
(OMNH 52711) tooth (FABL 6.51 mm) (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 53492) tooth (Brinkman, Cifelli and Czaplewski, 1998)
(OMNH 54004) three teeth (FABL 8.15, 6.47, 4.75 mm) (Brinkman, Cifelli and Czaplewski,
1998)
(OMNH 63061) material including femur and tibia (Pascucci, D'Emic and Turner, 2020)
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?
US?
(YPM 64979) vertebra (YPM online)
Diagnosis- (after Turner et al., 2012) four asymmetrical, subincisiform
premaxillary teeth; maxillary fenestra located dorsal to level of promaxillary
fenestra; 15 maxillary teeth; lobate anteriormost process of jugal beneath antorbital
fenestra; lacrimal boss prominent; prefrontal greatly reduced; 16 nearly isodont
dentary teeth; all teeth with anterior and posterior serrations; denticles of
posterior serrations nearly twice as large as denticles of anterior serrations
on all teeth; large, wide calcaneum.
Comments-
Brown discovered AMNH 3015 in 1931, and AMNH 3037 in 1932. As
Chure and McIntosh (1989) stated, Brown proposed "Daptosaurus agilis"
with the intended holotype AMNH 3015, but "never got around to
describing these animals, and it was left to John Ostrom to describe
and name them many years later in 1970, based in many cases on much
better and more diagnostic material which he had collected."
"None of Brown's names were ever published (until Glut 1972 [for
"Peltosaurus"]) and it does no harm to mention them here, as Ostrom's
names are well established taxa having full priority. Brown's names,
however, were used in public lectures, and for a time on some exhibited
specimens. In addition, they were written on photographs and
preliminary skeletal reconstructions made under Brown's
direction." In his 1931 field report, Brown first mentioned Deinonychus in saying "closely associated with one Tentontosaurus skeleton is a small carnivorous dinosaur but encased in lime difficult to prepare" (Grellet-Tinner and Makovicky, 2006).
The
YPM paratypes were found in 1964-1966 (YPM online), with the
holotype being discovered in late August 1964. The YPM quarry was
reopened in 1993 by the MOR, resulting in abundant new material (MOR
747; Maxwell and Witmer, 1996)- "one cervical vertebra, several caudal
vertebrae, a radius and ulna, two manus, two pes, and portions of the
skull, including ... the prefrontal, frontal, parietal, quadrate,
laterosphenoid, and portions of the braincase. Other cranial elements
in the new sample, such as the nasal, lacrimal, and ectopterygoid are
the best samples collected to date." Powers et al. (2022)
describe the maxilla of YPM 5232 in detail based on CT scans.
MCZ 4371 was found in July 1974 and described by Ostrom (1976).
Ostrom (1969b) believed that "YPM 5203 (a caudal sequence), 5205 (left
and right pes), 5206 (left and right manus) and 52010 (several
vertebrae and an incomplete skull and jaws) belong to a single
individual because of distinct preservation common to these alone and
the fact that they are all slightly larger than other comparable
lements obtained at the Yale site" but could not prove it. The
three articulated caudal series (YPM 5201-5203) show at least three
individuals were present among YPM 5201-5265, and Maxwell and Witmer
report four right splenials showing at least four were present.
Ostrom (1969a, b) misidentified the scapholunare as an ulnare, the
semilunate carpal as the radiale (= scapholunare in maniraptoriforms), and the coracoid as the pubis.
Parsons and Parsons (2002) initially identified MOR 1178 as a new troodontid-like
taxon based on an apparent parietal sagittal crest, dorsal groove on distal
caudal vertebrae, robust metatarsal IV and pedal ungual ratios similar to Troodon.
However, in the period between abstract submission and conference poster, they
had reidentified it as a subadult Deinonychus. This was published in
their 2003 paper, with no mention of the ectopterygoid or metatarsal IV that
were noted in 2002. They may have been reidentified as the squamosal fragment
and metatarsal II. The supposed parietal fragment was reidentified as a sacral
fragment. Some features such as highly elongated forelimbs and curved manual
ungual III are seen as ontogenetic differences from adult specimens.
Parsons and Parsons (2004) describe another young Deinonychus (MCZ 8791)
which was discovered in 1982. It is smaller than MOR 1178 and has even longer
forelimbs. Parsons and Parsons (2013) describe further details of this specimen.
Parsons and Parsons (2006) report on "newly discovered specimen" MOR 1182
Makovicky and Grellet-Tinner (2000) and Grellet-Tinner's (2005) thesis described
Deinonychus eggshells preserved with AMNH 3015 but left unidentified
for decades, which were officially published as Grellet-Tinner and Makovicky
(2006).
Main et al. (2011) reported material of dromaeosaurids and basal tetanurines
from the Woodbine Formation of Texas, the former of which are provisionally
listed as Deinonychus here.
Deinonychus was traditionally identified as a velociraptorine, but has
most recently been recovered as a dromaeosaurine (Senter et al., 2012; Foth
et al., 2014; Lee et al., 2014), sister taxon to Velociraptorinae+Dromaeosaurinae
(Longrich and Currie, 2009), or a velociraptorine after all (Brusatte et al.,
2014).
References- Ostrom, 1969a. A new theropod dinosaur from the Lower Cretaceous
of Montana. Postilla. 128, 17 pp.
Ostrom, 1969b. Osteology of Deinonychus antirrhopus, an unusual theropod
from the Lower Cretaceous of Montana. Bulletin of the Peabody Museum of Natural
History. 30, 165 pp.
Ostrom, 1970. Stratigraphy and paleontology of the Cloverly Formation (lower
Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Bulletin of the
Peabody Museum of Natural History. 35, 234 pp.
Ostrom, 1974. The pectoral girdle and forelimb function of Deinonychus
(Reptilia: Saurischia): a correction. Postilla. 165, 11 pp.
Gallup, 1975. Lower Cretaceous dinosaurs and associated vertebrates from north-central
Texas in the Field Museum of Natural History. MS thesis, University of Texas
at Austin. 159 pp.
Ostrom, 1976. On a new specimen of the Lower Cretaceous theropod dinosaur Deinonychus
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Paul. 1988. The small predatory dinosaurs of the mid-Mesozoic: The horned theropods
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Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves)
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Chiappe and Grellet-Tinner, 2000. Dinosaur eggshells and the origin of birds.
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Gishlick, 2001. The function of the manus and forelimb of Deinonychus antirrhopus
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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
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Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case
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of Vertebrate Paleontology. 25(3), 99A.
Grellet-Tinner and Makovicky, 2006. A possible egg of the dromaeosaur Deinonychus
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26(3), 109A.
Gignac, Makovicky, Erickson and Walsh, 2007. The effect of tooth morphology
on indentation force in non-avian theropod dinosaurs. Journal of Vertebrate
Paleontology. 27(3), 81A.
Parsons and Parsons, 2007. Avian-like manual phalanx found within gut contents
of Lower Cretaceous dromaeosaurid: New data on the feeding behavior of Deinonychus
antirrhopus (Saurischia: Theropoda). Journal of Vertebrate Paleontology.
27(3), 128A.
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in Deinonychus antirrhopus and other nonavian theropod dinosaurs. Bulletin
of the Peabody Museum of Natural History. 48(1), 103-138.
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antirrhopus using tooth indentation simulations. Journal of Vertebrate Paleontology.
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wildfires and biodiversity of coastal Appalachia in the Cretaceous (Cenomanian)
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antirrhopus and the consequent re-interpretation of dromaeosaurid features
that enhanced the evolution of avian flight. Journal of Vertebrate Paleontology.
Program and Abstracts 2012, 155.
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Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
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insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Parsons and Parsons, 2015. Morphological variations within the ontogeny of Deinonychus
antirrhopus (Theropoda, Dromaeosauridae). PLoS ONE. 10(4), e0121476.
Pascucci, 2019. Bone Histology of the theropod dinosaur Deinonychus antirrhopus. Masters thesis, Adelphi University. 38 pp.
Pascucci, D'Emic and Turner, 2020. Osteohistology of the Early Cretaceous theropod Deinonychus anitirrhopus. The Society of Vertebrate Paleontology 80th Annual Meeting, Conference Program. 265.
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2021). A new hypothesis of eudromaeosaurian evolution: CT scans assist
in testing and constructing morphological characters. Journal of
Vertebrate Paleontology. 41(5), 2010087.
Itemirus Kurzanov, 1976a
I. medullaris Kurzanov, 1976a
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (PIN 327/699) (adult) braincase
Referred- ?(CCMGE 456/12457) anterior dorsal centrum (Nessov, 1995)
?(CCMGE 461/12457) tooth (Nessov, 1995)
?(CCMGE? ?/12457; lost?) (juvenile) maxilla (Nessov, 1995)
?(CCMGE? ?/12457) teeth (Nessov, 1995)
?(ZIN PH 11/16) pedal phalanx II-2 (96.4 mm) (Sues and Averianov, 2014)
?(ZIN PH 12/16) pedal ungual III (Sues and Averianov, 2014)
?(ZIN PH 13/16) incomplete first dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 73/16) incomplete posterior dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 74/16) anterior dorsal centrum (Sues and Averianov, 2014)
?(ZIN PH 75/16) posterior dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 89/16) anterior cervical vertebra (Sues and Averianov, 2014)
?(ZIN PH 90/16) incomplete posterior cervical vertebra (Sues and Averianov,
2014)
?(ZIN PH 92/16) incomplete anterior cervical vertebra (Sues and Averianov, 2014)
?(ZIN PH 96/16) first dorsal vertebra (Sues and Averianov, 2014)
?(ZIN PH 182/16) metacarpal I (39 mm) (Sues and Averianov, 2014)
?(ZIN PH 183/16) partial metacarpal I (~76 mm) (Sues and Averianov, 2014)
?(ZIN PH 967/16) frontal (Sues and Averianov, 2014)
?(ZIN PH 2263/16) posterior cervical vertebra (Sues and Averianov, 2014)
?(ZIN PH 2269/16) incomplete posterior dorsal vertebra (Sues and Averianov,
2014)
?(ZIN PH 2273/16) proximal caudal vertebra (Sues and Averianov, 2014)
?(ZIN PH 2338/16) dentary fragment (Sues and Averianov, 2014)
?(ZIN PH 2344/16) premaxillary tooth (Sues and Averianov, 2014)
?(ZIN PH 2346/16) incomplete distal caudal vertebra (Sues and Averianov, 2014)
?(ZIN PH 2351/16) lateral tooth (Sues and Averianov, 2014)
?(ZIN PH 2352/16) lateral tooth (Sues and Averianov, 2014)
?(ZIN PH 2353/16) lateral tooth (Sues and Averianov, 2014)
?(ZIN PH? ?/16) teeth (Sues and Averianov, 2014)
?(ZIN PH? ?/16) pedal phalanx II-2 fragment (Sues and Averianov, 2014)
Diagnosis- (after Sues and Averianov, 2014) differs from Dromaeosaurus
in- basipterygoid recesses on dorsolateral surfaces of basipterygoid processes
present (also in Velociraptor, Tsaagan and Bambiraptor);
mesial serrations of teeth significantly smaller than distal serrations (also
in most dromaeosaurids except Utahraptor and Achillobator; teeth
tentatively referred). Differs from Utahraptor in- pleurocoels present
in anterior dorsals (also in other dromaeosaurids except Yurgovuchia;
vertebrae tentatively referred). Differs from both Utahraptor and Achillobator
in lateral surfaces of dorsal centra have deep, emarginated fossae (also in
Deinonychus; vertebrae tentatively referred).
(suggested) occipital condyle wider than foramen magnum.
Comments- The holotype braincase was discovered in 1958, and described
by Kurzanov as the new genus Itemirus in 1976(a,b). Kurzanov assigned Itemirus
to its own family, Itemiridae, which he placed sister to Tyrannosauridae, with
these both sister to Dromaeosauridae (including Stenonychosaurus but
not Saurornithoides) within the Carnosauria. Theropod braincases were
poorly known at the time so there was little to compare Itemirus to and
much of the description is outdated. Some aspects were described in a more modern
perspective by Currie and Zhao (1994), in comparison with Troodon. Chure
and Madsen (1998) found the deep basipterygoid recesses to be similar to their
cf. Stokesosaurus braincase, but noted several characters were less derived
than Stokesosaurus and tyrannosaurids. This feature led Holtz (2004)
to refer to Itemirus as a possible tyrannosauroid, though it has been
since noted in most dromaeosaurids and basal paravians. Longrich and Currie
(2009) found Itemirus to clade in Velociraptorinae, based on a paravian
matrix. Miyashita (2011) used a coelurosaur matrix which grouped Itemirus
with tyrannosauroids, but noted (DML, 2011) that few steps are needed to constrain
it to Dromaeosauridae.
Sues and Averianov (2004) referred additional material to Itemirus, describing
it as a dromaeosaurid and stating it reaches sizes comparable to Utahraptor.
Nessov (1995) referred the maxilla CCMGE 600/12457 to Alectrosaurus,
but it was reidentified as dromaeosaurid by Averianov and Sues (2012) and described
as cf. Itemirus by Sues and Averianov (2014). Tanaka et al. (2021) referred it to their new carcharodontosaur Ulughbegsaurus along with maxillary fragment ZIN PH 357/16 that was described as cf. Itemirus by Sues and Averianov. CCMGE 456/12457 was figured
by Nessov as a segnosaur, but later reidentified as cf. Itemirus by Sues
and Averianov (2014). The unnumbered maxilla and teeth listed above were identified
as Deinonychosauria and Dromaeosauridae by Nessov. The maxilla is described
as shorter and wider than Deinonychus, though not mentioned by Sues and
Averianov (2014) as no obviously corresponding specimen was found, while the
teeth (including that in plate 1 figure 4 of Nessov, assigned to cf. Itemirus
by Sues and Averianov, 2014) were described as small. While Sues and Averianov
(2004) refer "several braincases" to Itemirus in their abstract,
only the holotype was noted in the resulting publication. Sues (pers. comm.
2014) stated these were fragments that showed few diagnostic features once prepared,
so their identification remains uncertain. ZIN PH 73/16 was mislabeled ZIN PH
2273/16 in Sues and Averianov's (2014) figure. The final publication of the
Bissekty dromaeosaurid was Sues and Averianov's (2014), which tentatively referred
all material to Itemirus because "there are no morphological differences
among the few comparable remains." Based on the limited materal, this must
remain provisional. Several of the characters in this publication are miscoded
(dorsal tympanic recess is present; supratemporal fossa border is simply curved;
frontal's postorbital process not sharply demarcated; dentary has a labial groove;
posterior dorsal parapophyses are not on prominent pedicles; hyposphenes are
not separated), most favoring a dromaeosaurid identity. Given these miscodings,
Sues and Averianov's recovery of Itemirus as a dromaeosaurine more derived
than Tsaagan may be questioned.
References- Kurzanov, 1976a. Stroyeniye mozgovoy korobki karnozavra Itemirus gen. nov. i nekotoryye voprosy kranial-noy anatomii dinozavrov. Paleontologicheskii Zhurnal. 1976(3), 127-137.
Kurzanov, 1976b. Brain-case structure in the carnosaur Itemirus
n. gen. and some aspects of the cranial anatomy of dinosaurs. Paleontological Journal. 1976(3), 361-369.
Martinson, Nessov and Starobogatov, 1986. Unique find of gill apparatus of bivalve
molluscs Trigonioidoidea from the Cretaceous deposits of Kyzylkum. Byuleten'
Moskovskogo Obschestva Ispytatelei Prirody, Otdel Geologicheskii. 61, 94-98.
Currie and Zhao, 1994. A new troodontid (Dinosauria, Theropoda) braincase from
the Dinosaur Park Formation (Campanian) of Alberta. Canadian Journal of Earth
Sciences. 30(10-11), 2234-2247.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust. 1-156.
Chure and Madsen, 1998. An unusual braincase (?Stokesosaurus clevelandi)
from the Cleveland-Lloyd Dinosaur Quarry, Utah (Morrison Formation; Late Jurassic).
Journal of Vertebrate Paleontology. 18(1), 115-125.
Holtz, 2004. Tyrannosauroidea. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria Second Edition. University of California Press. 111-136.
Sues and Averianov, 2004. Dinosaurs from the Upper Cretaceous (Turonian) of
Dzharakuduk, Kyzylkum Desert, Uzbekistan. Journal of Vertebrate Paleontology.
24(3), 51A-52A.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern
Aral Sea region, Kazakhstan. Cretaceous Research. 28(3), 532-544.
Longrich and Currie, 2009. A microraptorine (Dinosauria–Dromaeosauridae)
from the Late Cretaceous of North America. Proceedings of the National Academy
of Sciences. 106(13), 5002-5007.
Miyashita, 2011. Cranial morphology of the basal tyrannosauroid Itemirus
medullaris and evolution of the braincase pneumaticity in non-avian coelurosaurs.
Journal of Vertebrate Paleontology. Program and Abstracts 2011, 159.
Miyashita, DML 2011. https://web.archive.org/web/20180115091932/http://dml.cmnh.org/2011Nov/msg00368.html
Averianov and Sues, 2012. Skeletal remains of Tyrannosauroidea (Dinosauria:
Theropoda) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan.
34, 284-297.
Sues and Averianov, 2014. Dromaeosauridae (Dinosauria: Theropoda) from the Bissekty
Formation (Upper Cretaceous: Turonian) of Uzbekistan and the phylogenetic position
of Itemirus medullaris Kurzanov, 1976. Cretaceous Research. 51, 225-240.
Tanaka, Anvarov, Zelenitsky, Ahmedshaev and Kobayashi, 2021. A new
carcharodontosaurian theropod dinosaur occupies apex predator niche in
the early Late Cretaceous of Uzbekistan. Royal Society Open Science. 8:
210923.
Achillobator Perle, Norell
and Clark, 1999
A. giganticus Perle, Norell and Clark, 1999
Cenomanian-Turonian, Late Cretaceous
Burkhant, Bayanshiree 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), phalanx I-1 (78 mm), metacarpal
III (71 mm), two manual unguals (112 mm), ilium (531 mm), pubes (548 mm), ischium
(378 mm), femur (505 mm), tibia (490.9 mm), phalanx II-2 (56.4 mm), metatarsal
III (234.4 mm), phalanx III-2 (55 mm), metatarsal IV (209.6 mm)
Referred- ? pedal phalanx II-2 (Barsbold et al., 2007)
Diagnosis- (modified from Perle et al., 1999) pelvis propubic; anterior
pubic foot slightly longer than posterior foot; very stout pedal phalanx II-2.
(after Turner et al., 2012) promaxillary fenestra completely exposed; promaxillary
and maxillary fenestra elongate and vertically oriented at same level in maxilla;
large triangular obturator process on ischium situated on proximal half of ischial
shaft; femur longer than tibia; metatarsal III wide proximally.
(proposed) 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 based on a draft left in Mongolia
in 1997 (Turner et al., 2012). 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 semiarticulated 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 its 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.
Barsbold, Kobayashi and Kubota, 2007. New discovery of dinosaur fossils from
the Upper Cretaceous Bayanshiree Formation of Mongolia. Journal of Vertebrate
Paleontology. 27(3), 44A.
Senter, 2007. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 5(4), 429-463.
Turner, Hwang and Norell, 2007. A small derived theropod from Oosh, Early Cretaceous,
Baykhangor Mongolia. American Museum Novitates. 3557, 27 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Utahraptor Kirkland, Gaston
and Burge, 1993
= "Utahraptor" Kirkland, 1992
U. ostrommaysi Kirkland, Gaston and Burge, 1993
= "Utahraptor spielbergi" Bonar, Lassieur, McCafferty and Voci, 1993
Barremian, Early Cretaceous
Yellow Cat Member of the Cedar Mountain Formation, Utah, US
Holotype- (CEU 184v.86; = CEUM 1430) pedal ungual II
Paratypes- ?...(CEU 184v.260) (5.03 m) tibia (503 mm)
?...(CEU 184v.294) pedal ungual II
?...(CEU 184v.400) premaxilla
Referred- ?...(CEU 184v.667; = CEUM 3528) quadratojugal (Britt et al.,
2001)
?...(CEU 184v coll.; = CEUM 1370) premaxilla (Britt et al., 2001)
?...(CEU 184v coll.; = CEUM 4023) partial palatine (Britt et al., 2001)
Paratypes- (BYU 9429) (3.89 m) mid caudal vertebrae (67 mm)
(BYU 9435) distal caudal vertebra
(BYU 9436) distal caudal vertebra
(BYU 9438) pedal ungual II
(BYU 13068) pedal ungual II
Referred- ?(BYU 10978) incomplete ischium (Britt et al., 2001)
(BYU 14281) femur (Britt et al., 2001)
(BYU 14389) partial ilium (Britt et al., 2001)
(BYU 14585) premaxilla (Britt et al., 2001)
(BYU 15417) femur (Britt et al., 2001)
(BYU 15465) (~5.9 m) femur (600 mm) (Erickson et al., 2009)
(BYU coll.) teeth (Kirkland et al., 2005)
(BYU and CEU coll.) (at least nine individuals, CEU material probably belongs
to holotype individual; 3 juveniles, 2 subadult and 4 adults; ~3-5.6 m, some
~11 m?) 360 elements including nasal, cervical vertebrae, dorsal vertebrae,
sacral vertebrae, proximal caudal vertebrae, coracoid, femora (120-565 mm),
tibia, astragalus, metatarsal II, phalanx II-1, metatarsal III, metatarsal IV
(Britt et al., 2001)
? material including dentary (Kirkland, Loewen, Deblieux, Madsen and Choiniere,
2011)
? material including premaxillary tooth, braincase, dentary, surangular and
astragalus (Kirkland, Loewen, Deblieux, Madsen and Choiniere, 2011)
Diagnosis- (after Turner et al., 2012) elongate nasal process of premaxilla;
quadratojugal L-shaped; dorsal vertebrae lack pleurocoels; well developed notch
present between anterior and greater trochanter; distal end of metatarsal III
smooth, not ginglymoid.
Comments- The name Utahraptor was first used in a press release
from Kirkland and Dinamation International Society (Olshevsky, 2000) and subsequently
published in several articles in July 1992 (e.g. Browne, 1992). Its species
name was originally going to be U. spielbergi, but Steven Spielberg's
lawyers apparently objected (Bakker pers. comm. to Tegowski, 1996). That name
has only been published in a children's magazine and merchandise, which of course
lacked a proper description. Once the taxon was properly published, Kirkland
et al. (1993) spelled the species name ostrommaysi, but this was emmended
by Olshevsky (1993) to ostrommaysorum
"because it honors two persons" (Olshevsky, 2000). However, Costa and
David (2019) noted that the ICZN actually says (Article 31.1.2) "A
species-group name, if a noun in the genitive case, formed directly
from a modern personal name is to be formed by adding to the stem of that name -i if the personal name is that of a man, -orum if of men ...", and ostrommaysi
was technically formed from two modern personal names (Ostrom and
Mays), not a modern personal name. Thus according to Costa and
David "Under Art. 11.3, it has to be regarded as an arbitrary
combination of letters, and not a correctly formed name in the genitive
case according to Arts. 31.1.1 or 31.1.2. For this reason, it is deemed
to be preserved unaltered and it is not subject to a justified
emendation." This makes ostrommaysorum
an unjustified emendation, which could be preserved if it were in
prevailing usage (ICZN Articles 33.2.3.1 and 33.3.1). But the
authors found it was only "used in 28 of the 57 works that mention the
taxon, representing a percentage of ca. 49%, which is far from
representing a "substantial majority"", cementing ostrommaysi as the correct name for now. The
name Utahraptor "oweni" has also appeared online in various
places but is probably a mistake for Valdoraptor oweni.
The type material was discovered in 1975 (BYU material) and 1991-1992 (CEU material),
but only described in 1993. The other CEU type material may belong to the same
individual as the holotype (Kirkland et al., 2008). A supposed lacrimal (CEU
184v.83) was also a paratype, but was later found to be a postorbital from the
ankylosaur Gastonia (Britt et al., 2001). Britt et al. also state a previously
identified surangular is actually a long bone fragment, but no surangular was
identified in the original description. Turner et al. (2012) consider this bone
to be a surangular or partial splenial. Britt et al. further determined that
the supposed manual unguals BYU 9438, BYU 13068 and CEU 184v.294 are actually
pedal unguals, which was confirmed by Senter (2007a).
New material was announced by Britt et al. (2001) and Chure et al. (2007) in
abstracts, but has yet to be described in detail. This consists of one individual
from the Yellow Cat (=Gaston) Quarry (CEU coll.) and at least eight from the
Dalton Wells Quarry (>240 elements in the BYU coll.). Anatomical information
on these specimens can be gleaned from the matrices of Senter (2007b), Longrich
and Currie (2009) and Turner et al. (2012), the latter of which also illustrate
a few elements. Of particular interest are caudal vertebrae about twice as long
as those belonging to a specimen with a 565 mm long femur. If this turns out
to be correct and not due to misidentification or unusual proportions, it could
indicate individuals over ten meters long. Turner et al. considered the ischium
BYU 10978 to be from an ornithomimid instead, based on the proximally placed
obturator process, lack of a lateral ridge, rodlike shaft and semicircular proximolateral
scar. Yet Achillobator also has a proximally placed obturator process
and the proximolateral scar is common among theropods. It is here provisionally
retained in Utahraptor pending its description.
Kirkland et al. (2011) mention two specimens, of which at least the large one
resembles Utahraptor in its premaxillary teeth, surangular and astragalus.
The small individual has serrationless dentary teeth that are procumbant anteriorly.
The large individual has large nuchal and sagittal crests on the braincase,
deep and pendant paroccipital processes, larghe olfactory bulbs and serrations
on the first dentary tooth.
References- Browne, 1992. A creature to make T. rex tremble. The
New York Times. July 21st.
Bonar, Lassieur, McCafferty and Voci, 1993. Disney Adventures. 3(9), 27-37.
Kirkland, Burge and Gaston, 1993. A large dromaeosaur (Theropoda) from the Lower
Cretaceous of eastern Utah. Hunteria. 2(10), 1-16.
Olshevsky, 1993. The name game. The Dinosaur Report. 1993, 5-6.
Tegowski, DML 1997. https://web.archive.org/web/20180115091831/http://dml.cmnh.org/1997Nov/msg00113.html
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic
Meanderings. 3, 1-157.
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. Journal of Vertebrate Paleontology. 21(3), 36A.
Kirkland, Scheetz and Foster, 2005. Jurassic and Lower Cretaceous dinosaur quarries
of Western Colorado and Eastern Utah. in Rishard (compiler). 2005 Rocky Mountain
Section of the Geological Society of America Field Trip Guidebook, Grand Junction
Geological Society. Field Trip 402, 1-26.
Chure, Britt and Scheetz, 2007. Sickle-claw theropod dinosaurs of the Lower
Cretaceous Cedar Mountain Formation from the Dalton Wells quarry and Dinosaur
National Monument, Utah. Journal of Vertebrate Paleontology. 27(3), 59A.
Senter, 2007a. A method for distinguishing dromaeosaurid manual unguals from
pedal "sickle claws". Bulletin of the Gunma Museum of Natural History.
11, 1-6.
Senter, 2007b. A new look at the phylogeny of Coelurosauria (Dinosauria: Theropoda).
Journal of Systematic Palaeontology. 5(4), 429-463.
Kirkland, Barrick, Bartlett, Bird and Burge, 2008. Taphonomy of the Gaston Quarry
(Early Cretaceous, Yellow Cat Member, Cedar Mountain Formation), East-Central
Utah: The case for expanding the hypodigm of Utahraptor beyond the sickle-claw.
Journal of Vertebrate Paleontology. 28(3), 100A.
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.
Longrich and Currie, 2009. A microraptorine (Dinosauria-Dromaeosauridae) from
the Late Cretaceous of North America. Proceedings of the National Academy of
Sciences. 106(13), 5002-5007.
Kirkland, Loewen, Deblieux, Madsen and Choiniere, 2011. New theropod cranial
material from the Yellow Cat Member, Cedar Mountain Formation (Barremian-Basal
Aptian, Cretaceous), Stikes Quarry, North of Arches National Park, East-Central
Utah. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 137.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Costa and David, 2019. Commentaries on different uses of the specific epithet of the large dromaeosaurid Utahraptor Kirkland et al., 1993 (Dinosauria, Theropoda). Bulletin of Zoological Nomenclature. 76, 90-96.
U? sp. (Tidwell, Carpenter and Meyer, 2001)
Aptian, Early Cretaceous
Tony's Bonebed, Poison Strip Member of Cedar Mountain Formation, Utah, US
Material- (DMNH coll.) ischium
Comments- Tidwell et al. (2001) wrote "A large deinonychid-type ischium found in the DMNH quarry may indicate the presence of Utahraptor."
References-
Tidwell, Carpenter and Meyer, 2001. New titanosauriform (Sauropoda)
from the Poison Strip Member of the Cedar Mountain Formation (Lower
Cretaceous), Utah. In Tanke and Carpenter (eds.). Mesozoic Vertebrate
Life. Indiana University Press. 139-165.