Oviraptorosauria Barsbold, 1976
Definition- (Oviraptor philoceratops <- Therizinosaurus cheloniformis, Passer domesticus) (modified from Hu, Hou, Zhang and Xu, 2009)
Other definitions- (Oviraptor philoceratops <- Passer domesticus) (Maryanska et al., 2002; modified from Currie and Padian, 1997)
(Oviraptor philoceratops + Chirostenotes pergracilis) (modified from Padian et al., 1999)
(Oviraptor philoceratops <- Tyrannosaurus rex, Ornithomimus edmontonicus, Therizinosaurus cheloniformis, Troodon formosus, Passer domesticus) (Sereno, in press)
= Caenagnathiformes Sternberg, 1940
Definition- (Caenagnathus collinsi <- Passer domesticus) (Martyniuk, 2012)
= Avimimiformes Chatterjee, 1991
= Oviraptorosauria sensu Currie and Padian, 1997
Definition- (Oviraptor philoceratops <- Passer domesticus) (modified)
= Oviraptoriformes sensu Benson, 2008
Definition- (Oviraptor philoceratops <- Passer domesticus) (Sereno, in press)
= Oviraptorosauria sensu Sereno, in press
Definition- (Oviraptor philoceratops <- Tyrannosaurus rex, Ornithomimus edmontonicus, Therizinosaurus cheloniformis, Troodon formosus, Passer domesticus)
Comments- Sereno (online) proposed the name Oviraptoriformes on the Taxon Search website with a phylogenetic definition, crediting it to Sereno, 2005. Yet online citations do not count for nomenclature, rendering this a nomen nudum until recently. Benson (2008) is the first author to publish the name Oviraptoriformes, though he did such undefined in a cladogram. Thus neither proposed name for this clade has a published definition.
Sereno's (in press) definition adds a lot of non-avian specifiers to Maryanska et al.'s (2002), which was sorely needed. The only issue I have (besides O. edmontonicus vs. O. velox) is the inclusion of Tyrannosaurus as an external specifier, as I'm unaware of any suggested topology positioning it closer to Oviraptor than the other taxa. Maybe Dromaeosaurus albertensis would be better, to cover the possibility figured by Barsbold et al. (1990).
Ex-oviraptorosaurs- A femur (ZIN PH 1/13) identified as oviraptorosaurian or ornithomimid by Nessov (1995) is ornithomimid (Averianov et al., 2003).
Currie et al. (1996) identified a surabgular (NMV P186386) and dorsal (NMV P186302) from the Eumeralla Formation of Victoria as oviraptorosaurian, but these have been placed more ambiguously as theropod (Agnolin et al., 2010) and maniraptoran (Benson et al., 2012) lately.
A cervical vertebra (MACN 622) discovered with the holotype was originally identified as an oviraptorosaur (Frankfurt and Chiappe, 1999), but reidentified by Agnolin and Martinelli (2007) as a noasaurid, and probably part of the Noasaurus holotype individual.
References- 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.
Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria) specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia. Alcheringa. 20(1-2), 73-79.
Frankfurt and Chiappe, 1999. A possible oviraptorosaur from the Late Cretaceous of northwestern Argentina. Journal of Vertebrate Paleontology. 19(1), 101-105.
Averianov, Starkov and Skutschas, 2003. Dinosaurs from the Early Cretaceous Murtoi Formation in Buryatia, Eastern Russia. Journal of Vertebrate Paleontology. 23(3):586–594.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php [version 1.0, 2005 November 7]
Agnolin and Martinelli, 2007. Did oviraptorosaurs (Dinosauria; Theropoda) inhabit Argentina? Cretaceous Research. 28(5), 785-790.
Balanoff, Bever and Rowe, 2008. The endocranial morphology of oviraptorosaurs and a reinterpretation of their encephalization quotients. Journal of Vertebrate Paleontology. 28(3), 47A.
Benson, 2008. New information on Stokesosaurus, a tyrannosauroid (Dinosauria: Theropoda) from North America and the United Kingdom. Journal of Vertebrate Paleontology. 28(3), 732-750.
Agnolina, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous non-avian dinosaur faunas from Australia and New Zealand: Evidence for their Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.
Persons, Currie and Norell, 2011. Shake your feathers: The flamboyant, athletic, and possibly flirtatious caudal morphology of oviraptorosaurs. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 174.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from Southern Australia indicates high polor diversity and climate-driven dinosaur provinciality. PLOS One. 7(5), e37122.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.
Balanoff, Bever and Norell, 2013. The relationships of oviraptorosaurian dinosaurs and endocranial evolution along a morphologically bizarre lineage. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 81.
Pittman and Hutchinson, 2013. The evolution of tail joint stiffness in oviraptorosaur dinosaurs and its consequences for tail function. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 191.
Tanaka, Zelenitsky, Lu, Yi, Pu, Chang, Xu and Li, 2014. Nest type and incubation behavior in oviraptorosaurs in relation to body size. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 238-239.

unnamed possible oviraptorosaur (Frey and Martill, 1995)
Albian, Early Cretaceous
Romualdo member of Santana Formation, Brazil
Material
- (SMNS 58023) (juvenile) third sacral vertebra (63.2 mm), fourth sacral vertebra (62.5 mm), fifth sacral vertebra (71.7 mm), fragment of sixth sacral neural arch and spine, fragment of first caudal neural arch, ilial fragment
Reference- Frey and Martill, 1995. A possible oviraptorosaurid theropod from the Santana Formation (Lower Cretaceous, Albian?) of Brazil. Neues Jahrbuch Fur Geologie und Palaeontologie. 7, 397-412.

undescribed Oviraptorosauria (Suzuki and Watabe, 2000)
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia

Material- (980807 GT Coy Oviraptr) posterior skeleton (Suzuki and Watabe, 2000)
(980807 GT SZK Oviraptr) complete skeleton (Suzuki and Watabe, 2000)
(980893 BgT NAR) (Suzuki and Watabe, 2000)
(940824 BgT MAT) complete postcranial skeleton (Watabe and Suzuki, 2000)
References- Watabe and Suzuki, 2000. Report on the Japan-Mongolia Joint Paleontological Expedition to the Gobi desert, 1994. In: Results of the Hayashibara Museum of Natural Sciences, Mongolian Academy of Sciences, Mongolian Paleontological Center, Joint Paleontological Expedition, n. 1, Hayashibara Museum of Natural Sciences, Research Bulletin. 1, 30-44.
Suzuki and Watabe, 2000.

undescribed oviraptorosaur (Currie, 2002)
Aptian-Albian, Early Cretaceous
Ohshih Formation, Mongolia
Material
- (PJC.2001.10) proximal femur
Reference- Currie, 2002. Report on fieldwork in Mongolia, September 2001. In: Alberta Palaeontological Society, sixth annual symposium, “Fossils 2002’, presented by Alberta Paleontological Society, in conjunction with Canadian Society of Petroleum Geologists, Paleontological Division and Department of Earth Sciences, Mount Royal College, p. 8-12.

undescribed oviraptorosaur (Hunt-Foster and Foster, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Williams Fork Formation, Colorado, US
Material
- humerus
Comments- Hunt-Foster and Foster (2015) state this resembles Microvenator more than Anzu.
Reference- Hunt-Foster and Foster, 2015. First occurrence of an oviraptorosaur (Theropoda: Maniraptora) from the Mesaverde Group (Williams Fork Formation) of northwestern Colorado. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 148.

unnamed clade (Senter, Barsbold, Britt and Burnham, 2004)
Definition- (Incisivosaurus gauthieri + Protarchaeopteryx robustus)
Diagnosis- (modified from Senter et al., 2004) first premaxillary teeth enlarged (also in Caudipteryx); dentary with anterior margin beveled so that distal tip is a 50-60 degree point (also in Caenagnathidae).
Comments- This grouping was suggested by Senter et al. (2004), although they called the clade Protarchaeopteryx, synonymizing that genus with Incisivosaurus.
Of the six characters used to support their pairing-
1. first premaxillary teeth enlarged and anteroposteriorly compressed.
The compression seems to be the important factor, as Caudipteryx has an enlarged first pair of teeth too (though seemingly cylindrical; Zhou et al., 2000). Falcarius possesses enlarged first dentary teeth, though its premaxillae are unknown. However, labiolingual compression may be a plesiomorphy, as it's seen in carnosaurs, Dilong, Compsognathus, dromaeosaurids and Troodon. Tyrannosaurids have labiolingually compressed teeth, while those of Byronosaurus and Archaeopteryx are described as round.
2. diminutive peglike teeth immediately posterior to enlarged pair of teeth.
Again, Caudipteryx shares the size difference in posterior premaxillary teeth, though they seem to be more slender judging by the description as "needle-like" and Zhou et al.'s illustrations. Conical anterior teeth are also observed in therizinosaurs (e.g. Falcarius, Erlikosaurus), so are probably a symplesiomorphy.
3. diminutive lanceolate posterior teeth.
Small maxillary teeth are shared with therizinosaurs, Pelecanimimus, Shuvuuia, troodontids and birds, so may simply be a maniraptoriform plesiomorphy reversed in dromaeosaurs. The shape refers to constricted bases, which are also seen in these taxa (and Microraptor/Cryptovolans), and may also be plesiomorphic.
4. short, high skull.
True of Caudipteryx, Avimimus and caenagnathoids as well, so probably an oviraptorosaurian plesiomorphy.
5. tall premaxilla.
Also seen in caenagnathoids, though not Caudipteryx or Avimimus.
6. dentary with anterior margin beveled so that distal tip is a 50-60 degree point.
Also seen in caenagnathids, but not Caudipteryx, Avimimus or oviraptorids.

Incisivosaurus Xu, Cheng, Wang and Chang, 2002
I. gauthieri Xu, Cheng, Wang and Chang, 2002
= Protarchaeopteryx gauthieri (Xu, Cheng, Wang and Chang, 2002) Senter, Barsbold, Britt and Burnham, 2004
Late Valanginian-Hauterivian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China

Holotype- (IVPP V13326) skull (100 mm), incomplete mandibles, partial cervical vertebra
Diagnosis- (after Xu et al., 2002) large high-angled wear facets on the mesial margins of the teeth (unknown in Protarchaeopteryx); contact between the accessory ventral flanges of the pterygoids (unknown in Protarchaeopteryx).
(after Senter et al., 2004) nine maxillary teeth; eight or nine dentary teeth.
(proposed) fused dentary symphysis.
Comments- Xu et al. (2002) also included the following characters in their diagnosis- large incisciform first premaxillary tooth (also in Protarchaeopteryx); much smaller, subconical second to fourth premaxillary teeth (plesiomorphic for Oviraptoriformes); very small lanceolate maxillary teeth (plesiomorphic for Oviraptoriformes); triradiate palatine with very short maxillary process (plesiomorphic for Oviraptoriformes); longitudinal crest on the ventral surface of the basisphenoid (plesiomorphic for coelurosaurs). The "subsidiary ectopterygoid fenestra" they note seems to be merely a reduced subsidiary palatal fenestra shifted between the ectopterygoid and palatine as in oviraptorids. Balanoff et al. (2009) note the posterior mandibular fragment is lost.
References- Xu, Cheng, Wang and Chang, 2002. An unusual oviraptorosaurian dinosaur from China. Nature. 419, 291-293.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History. 8, 1-20.
Balanoff, Xu, Matsufune, Kobayashi and Norell, 2007. Endocranial anatomy of a primitive oviraptorosaur, Incisivosaurus gauthieri, (Theropoda: Dinosauria). Journal of Vertebrate Paleontology. 27(3), 43A.
Balanoff, Xu, Kobayashi, Matsufune and Norell, 2009. Cranial osteology of the theropod dinosaur Incisivosaurus gauthieri (Theropoda: Oviraptorosauria). American Museum Novitates. 3651, 35 pp.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.

Chuniaoia Ji and Ji, 2001
Protarchaeopteryx Ji and Ji, 1997
P. robusta Ji and Ji, 1997
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou member of Yixian Formation, Liaoning, China

Holotype- (NGMC 2125) (690 mm) incomplete skull (70 mm), mandibles, hyoid, six cervical vertebrae, five dorsal vertebrae, sacrum, fourteen caudal vertebrae, chevron, proximal scapula, partial coracoids, sternal plates (25x15 mm), partial furcula, partial humeri (87 mm), incomplete radii (73 mm), incomplete ulnae, radiale, semilunate carpal, distal carpal III, 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, two keratinous manual ungual sheaths, partial ilium (95 mm), incomplete pubes, ischial fragment(?), incomplete femora (125 mm), tibiae (160 mm), partial fibulae, astragali, calcaneum, distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual III, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, contour feathers, retrices
Diagnosis- (after Senter et al., 2004) six maxillary teeth; seven dentary teeth.
Other diagnoses- Ji et al. (1998) also included the following characters in their diagnosis- large straight premaxillary teeth (also in Incisivosaurus); short, bulbous maxillary and dentary teeth (plesiomorphic for Oviraptoriformes); all of which are primitively serrated (incorrect- Senter et al., 2004); rectrices form a fan at the end of the tail (plesiomorphic for Maniraptora).
Comments- Ji and Ji (2001) used the name Chuniaoia in a cladogram for a branch leading to Protarchaeopteryx, but it has not been defined nor has its intended purpose been published. It is similar to the name Chuniaoae used in the online supplementary information of Ji et al. (1998), which was seemingly for a Caudipteryx+Aves clade.
Relationships- Zhou (1997) classified Protarchaeopteryx as a sauriurine bird more closely related to enantiornithines than Confuciusornis due to the smaller manual digit III, but this has not been accepted since.
Xu et al. (1999) resolved Protarchaeopteryx as a paravian in a trichotomy with Caudipteryx and Troodontidae+Eumaniraptora. Xu et al. (2000) used 86 of the same characters plus three new ones, added Microraptor and removed Tyrannosauridae and Unenlagia. Their tree was slightly more resolved, as Protarchaeopteryx was found to be the sister taxon of Troodontidae+Eumaniraptora, ands thus closer to birds than Caudipteryx. However, both studies supported the paravian status of Protarchaeopteryx using the same seven characters.
- deep suborbital bar. This is actually unknown in Protarchaeopteryx, though probable given its presence in Incisivosaurus.
- unfused interdental plates. These are also present in therizinosaurs, and are recovered as derived for Paraves based only on their absence in the toothless ornithomimids and oviraptorids directly basal to this clade, plus their coded presence in Caudipteryx and Protarchaeopteryx. Notably, every other matrix has coded the latter two taxa unknown for this character.
- less than 11 caudal vertebrae with transverse processes. Protarchaeopteryx actually has more than twelve caudals with transverse processes.
- proximodistally elongate coracoid. Oviraptorids and troodontids are miscoded as lacking this character, as it is actually a maniraptoran synapomorphy.
- radius less than 70% of ulnar width. This is also present in Falcarius and Caudipteryx, suggesting it may be primitive for maniraptorans. However, there is much homoplasy.
- metacarpal I less than 33% of metacarpal II in length. Protarchaeopteryx's metacarpal I is actually 42% of metacarpal II's length.
- pubic foot projects caudally only. The pubis of Protarchaeopteryx is only visible in posterior view, making the presence or absence of an anterior expansion unknown.
Xu et al. (2000) resolved Protarchaeopteryx as a paravian more derived than Caudipteryx based on four additional characters.
- teeth with serrated mesial and distal carinae. Protarchaeopteryx was miscoded as having dental serrations.
- mid and distal caudal vertebrae at least 130% longer than proximal caudals. Protarchaeopteryx was miscoded as having elongate distal caudals.
- manus over 120% of ulnar length. Compsognathus, ornithomimids, Caudipteryx and oviraptorids are miscoded as lacking this (ornithomimids and oviraptorids are polymorphic), as it is a synapomorphy of a more inclusive clade than Paraves.
- short dorsal process of premaxilla. This is unable to be determined in Protarchaeopteryx due to poor preservation.
When these and other miscodings are corrected, the topology is changed. In Xu et al. (1999), Protarchaeopteryx is the sister taxon of Caudipteryx based on the low number of caudal vertebrae (<27) and the supposed presence of unfused interdental plates. It and Caudipteryx are oviraptorosaurs based on three characters unknown in Protarchaeopteryx. In Xu et al. (2000), Protarchaeopteryx is a paravian in a trichotomy with Caudipteryx and Troodontidae+Eumaniraptora (like in the uncorrected Xu et al., 1999 results). It's a paravian based on supposedly unfused interdental plates, the low number of caudal vertebrae (<27), and the slender radius (<70% of ulnar width). Based on the above information, this is probably due to the absence of therizinosaurs (especially Falcarius) and oviraptorosaurian synapomorphies in the matrix. Constraining Protarchaeopteryx to be an oviraptorosaur results in trees one step longer.
Holtz (2001) found Protarchaeopteryx to be either a basal maniraptoran sister to Oviraptoriformes+Paraves, a basal oviraptoriform, or a basal paravian.
Ji and Ji (2001) placed Protarchaeopteryx as a basal avialan in their cladogram, using the name Dromavialae for the node of Protarchaeopteryx+Aves. This was based on the character "real wings with symmetrical feathers of modern concept", which is vague and also occurs in dromaeosaurids and Caudipteryx.
Gishlick (2002) found Protarchaeopteryx to be in a eumaniraptoran polytomy with Deinonychus, Sinornithosaurus, Archaeopteryx and Pygostylia, though the matrix only included forelimb characters.
Paul (2002) assigned Protarchaeopteryx to the Archaeopterygidae based on several characters, none of which are convincing. For instance, the teeth are virtually identical to those of Incisivosaurus (Senter et al., 2004), so Protarchaeopteryx's "small, conical teeth" are not distinctively archaeopterygid. The sternal morphology is invalid because Archaeopteryx's supposed sternum is actually a coracoid. Other characters such as the "slender fingers", and "non-fused but tighly articulated metacarpals, tarsals and metatarsals" are symplesiomorphic within maniraptorans and vague. Only one third manual digit of Protarchaeopteryx is crossed under digit II, which is disarticulated in any case. The ilium is too incomplete to know if the preacetabular process would make it parallelogram-shaped, while the pubis is too incomplete to measure pelvic canal depth. I cannot confirm the coracoid is strongly bent as in microraptorians and avialans. The robust second manual digit is shared with caenagnathids, while the low ilium and slender pointed postacetabular process are also seen in Avimimus.
Senter (2003) combined Protarchaeopteryx and Incisivosaurus into one OTU, which he found to be the basalmost oviraptorosaur. Senter et al. (2004) came to the same conclusion.
Holtz et al. (2004) recovered Protarchaeopteryx as a paravian more derived than alvarezsaurids, excluded from Deinonychosauria and Archaeopteryx+Ornithurae.
Ji et al. (2005) added it to Clarke's bird matrix, which resulted in it being excluded from Microraptor+Aves.
Most recently, Senter (2007) found Protarchaeopteryx to be the sister group of Incisivosaurus, with both of these taxa as the most basal oviraptorosaurs.
References- Ji and Ji, 1997. A Chinese archaeopterygian, Protarchaeopteryx gen. nov.. Geological Science and Technology. 238, 38-41.
Zhou, 1997. Diversification of birds from the "Late Jurassic" of China. Journal of Vertebrate Paleontology. 17(3), 86A.
Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs from northeastern China. Nature. 393, 753-761.
Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
Xu, Zhou and Wang, 2000. The smallest known non-avian theropod dinosaur. Nature. 408, 705-708.
Holtz, 2001. Arctometatarsalia revisited: the problem of homplasy in reconstructing theropod phylogeny. pp. 99-122. 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.
Ji and Ji, 2001. How can we define a feathered dinosaur as a bird? 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. 43-46.
Padian, Ji and Ji, 2001. Feathered dinosaurs and the origin of flight. in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. 117-138.
Gishlick, 2002. The functional morphology of the forelimb of Deinonychus antirrhopus and its importance for the origin of avian flight. Unpublished PhD thesis. Yale University, 142 pp.
Paul, 2002. Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press.
Senter, 2003. Taxonomic sampling artifacts and the phylogenetic position of Aves. Unpublished PhD thesis. Northern Illonois University.
Holtz, Molnar and Currie, 2004. Basal Tetanurae. pp. 71-110, in Weishampel, Dodson and Osmólska (eds.). The Dinosauria. Second Edition. University of California Press.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae. Bulletin of Gunma Museum of Natural History 8: 1-20.
Ji, Ji, Lu, You, Chen, Liu and Liu, 2005. First avialan bird from China (Jinfengopteryx elegans gen. et sp. nov.). Geological Bulletin of China 24(3): 197-205.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.

Similicaudipteryx He, Wang and Zhou, 2008
S. yixianensis He, Wang and Zhou, 2008
= Caudipteryx yixianensis (He, Wang and Zhou, 2008) Paul, 2010
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China

Holotype- (IVPP V12556) (adult) six cervical vertebrae, cervical ribs, fourteen dorsal vertebrae, dorsal rib fragments, sacrum (85 mm), twenty-one partial caudal vertebrae, pygostyle, six chevrons, incomplete scapula, partial coracoid, incomplete sternal plates, sternal rib fragments, humerus (~130 mm), partial ulna, partial radius, ilia (one partial; 153 mm), pubes (~223 mm), ischial fragment, femora (one distal; ~220 mm), tibiae (one incomplete; ~240 mm), partial fibula (~223 mm), metatarsal I (25 mm), phalanx I-1 (28 mm), pedal ungual I (23 mm), metatarsals II (144 mm), phalanges II-1 (~43 mm), phalanx II-2 (39 mm), pedal ungual II (33 mm), metatarsals III (~183 mm), phalanges III-1 (46 mm), phalanx III-2 (34 mm), phalanx III-3 (32 mm), pedal ungual III (34 mm), metatarsals IV (153 mm), phalanges IV-1 (30 mm), phalanx IV-2 (23 mm), phalanx IV-3 (18 mm), phalanx IV-4 (21 mm), pedal ungual IV (20 mm)
Late Valanginian-Middle Aptian, Early Cretaceous
Yixian Formation, Liaoning, China

Referred- (STM4-1) (juvenile) partial skull, cervical vertebrae, dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, chevrons, scapulae, humeri (24 mm), radii (21 mm), ulnae, semilunate carpals, metacarpals I, phalanx I-1, manual ungual I, metacarpals II, phalanges II-1, phalanges II-2, manual unguals II, metacarpals III, phalanges III-1, phalanges III-2, phalanx III-3, manual ungual III, ilium, pubis, ischium, femora (38 mm), tibiae (49 mm), fibula, astragalus, metatarsals I, phalanges I-1, pedal unguals I, metatarsals II (one incomplete), phalanges II-1 (one incomplete), partial phalanx II-2, metatarsals III (one incomplete), phalanges III-1 (one incomplete), phalanx III-2, phalanx III-3 fragment, metatarsals IV (one partial), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, incomplete pedal ungual IV, metatarsal V, body feathers, remiges, retrices (Xu et al., 2010a)
(STM22-6) (subadult) skull, mandible, ten cervical vertebrae, cervical ribs, dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, pygostyle, chevrons, scapulae, coracoid, furcula, humeri (81 mm), radii (69 mm), ulnae, metacarpals I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1, phalanx II-2, manual ungual II, metacarpals III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, incomplete ilium, pubes, ischium, femur (140 mm), tibiae (182 mm), astragali, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, body feathers, remiges, retrices (Xu et al., 2010a)
Diagnosis- (modified from He et al., 2008) puboilial ratio of 1.46.
Comments- The holotype was first mentioned by Wang et al. (2007) as an unnamed caudipterid and later described and named by He et al. (2008). In its diagnosis, He et al. list a dagger-like pygostyle (also present in Nomingia and Citipati), ilium shaped like Caudipteryx (ambiguous), two large anterior dorsal hypapophyses (present in caenagnathoids) and puboilial ratio of 1.46. The latter compares with 1.04-1.12 in Caudipteryx, .96 in Nomingia and .99 in Microvenator. He et al. referred it to Caudipteridae based on several characters. Most are symplesiomorphic for maniraptorans (low number of caudal vertebrae; deep pubic peduncle; unfused metatarsus; metatarsal III longest; metatarsals II and IV subequal in width; metatarsal II slightly shorter than IV; subarctometatarsal metatarsus), the proximally placed metatarsal I is also present in Protarchaeopteryx, and the preacetabular depth is intermediate between other oviraptorosaurs. Xu et al. (2010a) briefly describe two new, younger specimens which are almost complete.
References- Wang, Jones and Evans, 2007. A juvenile anuran from the Lower Cretaceous Jiufotang Formation, Liaoning, China. Cretaceous Research. 28, 235-244.
He, Wang and Zhou, 2008. A new genus and species of caudipterid dinosaur from the Lower Cretaceous Jiufotang Formation of Western Liaoning, China. Vertebrata PalAsiatica. 46(3), 178-189.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Ptum, 2010. Moulting tail feathers in a juvenile oviraptorisaur. Nature. 468, E1.
Xu, Zheng and Yu, 2010a. Exceptional dinosaur fossils show ontogenetic development of early feathers. Nature. 464, 1338-1341.
Xu, Zheng and Yu, 2010b. Xu et al. reply. Nature. 464, 468, E2.

unnamed clade (Caudipteryx zoui + Oviraptor philoceratops)

Ganzhousaurus Wang, Sun, Sullivan and Xu, 2013
G. nankangensis Wang, Sun, Sullivan and Xu, 2013
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China

Holotype- (SDM 20090302) incomplete mandible, distal caudal vertebra, incomplete distal caudal vertebra, partial distal caudal vertebra, ilial fragment, partial tibia, distal tarsal III, metatarsal I (36 mm), phalanx I-1 (28 mm), pedal ungual I, metatarsal II (125 mm), phalanx II-1 (41 mm), phalanx II-2, partial pedal ungual II, metatarsal III (147 mm), phalanx III-1 (45 mm), partial phalanx III-2, phalanx IV-1, partial phalanx IV-2
Diagnosis- (after Wang et al., 2013) shallow dentary (ratio of maximum anteroposterior length to maximum dorsoventral depth 1.90); absence of fossa or foramen on lateral surface of dentary (also in oviraptorids; unlike Gigantoraptor, Chirostenotes and Caenagnathasia); weakly downturned anterior mandibular end (also in Khaan; unlike Nemegtomaia and Heyuannia); shallow depression immediately surrounding anterior margin of external mandibular fenestra (also in Gigantoraptor, Nemegtomaia and Heyuannia; unlike Citipati and Khaan); external mandibular fenestra subdivided by anterior process of surangular (also in oviraptorids); dentary posteroventral process slightly twisted and positioned on mandibular ventrolateral surface (also in Nemegtomaia; unlike Citipati and Heyuannia); shallow longitudinal groove along medial surface of dentary posteroventral process (also in Microvenator; unknown in other oviraptorosaurs); angular anterior process wider transversely than deep dorsoventrally (unknown in other oviraptorosaurs); sharp groove along ventrolateral surface of angular anterior process (also in Chirostenotes, Gigantoraptor and Khaan; unlike most oviraptorosaurs); ventral border of external mandibular fenestra formed mainly by angular (also in Chirostenotes and Gigantoraptor; unlike oviraptorids); ventral flange along distal half of metatarsal II (also in Avimimus; unlike Citipati, Wulatelong and Heyuannia); arctometatarsal condition absent (also in oviraptorids; unlike Avimimus, Chirostenotes and Elmisaurus).
Comments- The holotype was purchased from a fossil dealer and initially mentioned in an abstract by Wang and Xu (2012) before being officially described by Wang et al. (2013). Wang and Xu found it to be an "ingeniine" in their unpublished analysis, Wang et al. found it to be an oviraptorine most closely related to Citipati and Rinchenia in its published form, while Cau (online, 2013) found it to be in a trichotomy with caenagnathids and oviraptorids. Note the diagnosis given by Wang et al. includes numerous symplesiomorphies and will need to be revised. Initially the partial tibia was thought to be a femur by Wang and Xu.
References- Wang and Xu, 2012. A new oviraptorid specimen from the Upper Cretaceous of Southern China. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 190.
Cau, online 2013. http://theropoda.blogspot.com/2013/04/ganzhousaurus-nankangensis-wang-et-al.html
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.

Luoyanggia Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009
L. liudianensis Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009
Aptian-Albian, Early Cretaceous
Haoling Formation, Henan, China
Holotype
- (41HIII-00010) dentaries
....(41HIII-00011) ilia (one incomplete), pubes (one partial; 160 mm), ischium (100 mm), metatarsal II (175), metatarsal III (190 mm), metatarsal IV (177 mm)
Other diagnoses- Lu et al. (2009) listed three characters in their diagnosis, but all are plesiomorphic- dentary not decurved; dentary symphysis V-shaped; ischium only slightly concave dorsally.
Comments- While originally referred to the Mangchuan Formation, this was redefined to include three formations by Xu et al. in 2012.
While this is clearly a basal oviraptorosaur (albeit more derived than Incisivosaurus), it is uncertain precisely how it relates to caudipterids or caenagnathids.
Reference- Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009. A preliminary report on the new dinosaurian fauna from the Cretaceous of the Ruyang Basin, Henan province of Central China. Journal of the Paleontological Society of Korea. 25(1), 43-56.

Caudipteridae Zhou and Wang, 2000
Definition- (Caudipteryx zoui <- Caenagnathus collinsi, Oviraptor philoceratops) (Hendrickx, Hartman and Mateus, 2015)
= Caudipterygidae Osmolska, Currie and Barsbold, 2004
Comments- In 2000, Zhou and Wang proposed the family Caudipteridae for Caudipteryx. Osmolska et al. (2004) emmended this to Caudipterygidae, since Caudipteridae is formed incorrectly (ICZN Article 29.3). It has been suggested this is unecessary, since according to Article 29.4, "if after 1999 a new family-group name is based on a generic name which is or ends in a Greek or Latin word or ends in a Greek or Latin suffix, but its derivation does not follow the grammatical procedures of Articles 29.3.1 or 29.3.2, its original spelling must be maintained as the correct original spelling." However, Article 29.4.2 states this is only true provided the genus was treated as an arbitrary combination of letters (e.g. "Caudipteryxidae"), which is not the case. To complicate matters, Article 29.5 states "If a spelling of a family-group name was not formed in accordance with Article 29.3 but is in prevailing usage, that spelling is to be maintained, whether or not it is the original spelling and whether or not its derivation from the name of the type genus is in accordance with the grammatical procedures in Articles 29.3.1 and 29.3.2." According to Google, Caudipteridae has 3550 search results compared to Caudipterygidae's 685 (as of March 2010). Thus Caudipteridae should be maintained.
References- Zhou and Wang, 2000. A new species of Caudipteryx from the Yixian Formation of Liaoning, northeast China. Vertebrata PalAsiatica. 38(2), 113-130.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria (second edition). University of California Press, Berkeley. 165-183.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.

Caudipteryx Ji, Currie, Norell and Ji, 1998
Diagnosis- premaxillary teeth limited to rostral half of element; first premaxillary tooth much larger than others (also in Incisivosaurus+Protarchaeopteryx); single maxillary fenestra present; twenty-two caudal vertebrae; sternal plates oval; only two phalanges present on manual digit III.
C. zoui Ji, Currie, Norell and Ji, 1998
= Caudipteryx dongi Zhou and Wang, 2000
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou member of Yixian Formation, Liaoning, China

Holotype- (NGMC 97-4-A) (890 mm) skull (76 mm), mandibles, cervical vertebae, cervical ribs, dorsal vertebrae, dorsal ribs, gastralia, twenty-two caudal vertebrae (first caudal 12 mm), chevrons, coracoid, sternal plate (36 mm), humeri (69 mm), radii, ulnae, radiale, semilunate carpal, distal carpal III, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, ilia, pubes, ischia (77 mm), femora (147 mm), tibiae (188 mm), fibula, phalanx I-1, metatarsal II, phalanx II-1, phalanx II-2, partial pedal ungual II, metatarsal III (115 mm), fragmentary digit III, metatarsal IV, phalanx IV-1, contour feathers, remiges, retrices, gastroliths
Paratype- (NGMC 97-9-A) (725 mm) incomplete skull (79 mm), mandibles, hyoid, cervical vertebrae, cervical ribs, dorsal vertebrae, dorsal ribs, sacrum, caudal vertebrae, chevrons, scapulae (77 mm), coracoids, partial furcula, sternal plates, sternal ribs, humeri (70 mm), radii (57 mm), ulnae, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, partial phalanx III-1, incomplete ilium, partial pubes, femora (149 mm), tibiae (182 mm), astragali, calcanea, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (117 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, contour feathers, retrices, remiges, gastroliths
Referred- (IVPP V11819) specimen including femur (149 mm) (Erickson et al., 2009)
(IVPP V 12344, holotype of Caudipteryx dongi) (896 mm) frontal, pterygoid, two cervical vertebrae, six dorsal vertebrae, dorsal ribs (100 mm), three uncinate processes (30 mm), gastralia, sacrum, eleven caudal vertebrae, chevrons, partial coracoid, sternal plates (25 mm), sternal ribs (35 mm), incomplete humeri (~73 mm), radii (~58 mm), ulnae (61 mm), semilunate carpal, radiale, ulnare, metacarpal I (13 mm), phalanx I-1 (25 mm), manual ungual I (15 mm), metacarpal II (29 mm), phalanx II-1 (18.5 mm), phalanx II-2 (25 mm), manual ungual II (18 mm), metacarpal III (27 mm), ilia (115 mm), pubes, ischia (73 mm), femora (146, 152 mm), tibiae (196 mm), fibula (181 mm), astragali, calcaneum, distal tarsal III, distal tarsal IV, metatarsal I (19 mm), phalanx I-1 (12 mm), pedal ungual I (11 mm), metatarsal II (112 mm), phalanx II-1 (25 mm), phalanx II-2 (16 mm), pedal ungual II (19 mm), metatarsal III (124 mm), phalanx III-1 (27 mm), phalanx III-2 (20 mm), phalanx III-3 (17 mm), pedal ungual III (20 mm), metatarsal IV (116 mm), phalanx IV-1 (15 mm), phalanx IV-2 (9 mm), phalanx IV-3 (7 mm), phalanx IV-4 (7 mm), pedal ungual IV (16 mm), metatarsal V (~36 mm), body feathers, remiges (182 mm), gastroliths (Zhou and Wang, 2000)
(LPM 0005) skull, mandibles, cervical series, cervical ribs, dorsal series, dorsal ribs, gastralia, sacral vertebrae, caudal series, chevrons, scapulae, coracoid, furcula, humeri, radii, ulnae, semilunate carpal, metacarpals I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1 (one partial), phalanx II-2, manual ungual II, metacarpals III, phalanges III-1, phalanges III-2, ilia, ischia, femora, tibiae, fibulae, astragali, calcaneum, metatarsals I, phalanges I-1, pedal ungual I, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III, 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, metatarsals V, remiges, retrices, body feathers, gastroliths (Feduccia and Czerkas, 2015)
Diagnosis- premaxilla sharply pointed anteriorly; quadratojugal posterior process slightly developed; manual ungual II larger than manual ungual I.
C. sp. nov. (Zhou, Wang, Zhang and Xu, 2000)
Late Valanginian-Middle Aptian, Early Cretaceous
Yixian Formation, Liaoning, China

Material- (BPM 0001) (852 mm) skull, mandibles, twelve cervical vertebrae, cervical ribs, nine dorsal vertebrae, dorsal ribs (~114 mm), uncinate processes, gastralia, sacrum, twenty-two caudal vertebrae, chevrons, scapulae (80 mm), coracoids (34 mm), sternal plates (~30 mm), sternal ribs (~38 mm), humeri (72 mm), radii (59 mm), ulnae (62 mm), semilunate carpal, radiale, ulnare, metacarpal I (11 mm), phalanx I-1 (25 mm), manual ungual I (16 mm), metacarpal II (28 mm), phalanx II-1 (17 mm), phalanx II-2 (24 mm), manual ungual II (15 mm), metacarpal III (25 mm), phalanx III-1, phalanx III-2, ilia (115 mm), pubes (~124 mm), ischia (72 mm), femora (145 mm), tibiae (188 mm), fibulae (188 mm), astragali, calcanea, distal tarsal III, distal tarsal IV, metatarsal I (16 mm), phalanx I-1 (13 mm), pedal ungual I (12 mm), metatarsal II (102 mm), phalanx II-1 (23 mm), phalanx II-2 (16 mm), pedal ungual II (19 mm), metatarsal III (113 mm), phalanx III-1 (24 mm), phalanx III-2 (19 mm), phalanx III-3 (15 mm), pedal ungual III (18 mm), metatarsal IV (107 mm), phalanx IV-1 (14 mm), phalanx IV-2 (8 mm), phalanx IV-3 (6 mm), phalanx IV-4 (4 mm), pedal ungual IV (14 mm), metatarsal V (30 mm), contour feathers, remiges, retrices, gastroliths
(IVPP V 12340) (836 mm) skull, mandibles, twelve cervical vertebrae, cervical ribs, nine dorsal vertebrae (d3 ~18 mm, d6 ~18 mm, d8 ~17 mm), dorsal ribs (120 mm), uncinate processes (26 mm), gastralia, sacrum, twenty-two caudal vertebrae, chevrons, scapulae (80 mm), coracoids (35 mm), sternal ribs (36 mm), humeri (69 mm), radii (~56 mm), ulnae (61 mm), semilunate carpal, radiale, ulnare, metacarpal I (11 mm), phalanx I-1 (26 mm), manual ungual I (16 mm), metacarpal II (28 mm), phalanx II-1 (19 mm), phalanx II-2 (24 mm), manual ungual II (15 mm), metacarpal III (23 mm), phalanx III-1, phalanx III-2, ilia (112 mm), pubes (~125 mm), ischia (~72 mm), femora (145 mm), tibiae (183 mm), fibulae (175 mm), astragali, calcanea, distal tarsal III, distal tarsal IV, metatarsal I (15 mm), phalanx I-1 (12 mm), pedal ungual I (12 mm), metatarsal II (102 mm), phalanx II-1 (22 mm), phalanx II-2 (14 mm), pedal ungual II (17 mm), metatarsal III (112 mm), phalanx III-1 (23 mm), phalanx III-2 (17 mm), phalanx III-3 (13 mm), pedal ungual III (18 mm), metatarsal IV (106 mm), phalanx IV-1 (12 mm), phalanx IV-2 (8 mm), phalanx IV-3 (6 mm), phalanx IV-4 (5 mm), pedal ungual IV, metatarsal V (31 mm), contour feathers, remiges, retrices, gastroliths
Diagnosis- large premaxillary subnarial process; maxilla extended anteriorly with promaxillary fossa; external naris close to antorbital fenestra in length; jugal strongly concave posterodorsally; posterodorsal dentary process subequal in width to posteroventral process; posterodorsal dentary process longer than posteroventral process; no intramandibular joint; vomers do not extend past external nares?; ectopterygoid very thin and C-shaped; twelve cervical vertebrae?; ventral margin of coracoid irregular?; anterior margin of preacetabular process posteroventrally oriented.
Description- The various specimens referred to Caudipteryx show quite a bit of variation. They will be described together and the implications for the number of species and individual variation will be discussed afterward.
Skulls are illustrated for BPM 0001, IVPP V 12430 and NGMC 97-9-A. A complete skull is shown in the skeletal drawing of NGMC 97-4-A, but not illustrated in detail. A frontal and possible pterygoid are known in IVPP V 12344, but not illustrated in detail either. The skull is fairly short, with a snout taking up about half the length. Contra Zhou et al. (2000), the external naris is not larger than the antorbital fenestra once the specimens are articulated. It is much smaller in NGMC 97-9-A, but only slightly smaller in BPM 0001. It extends to the rostral border of the antorbital fenestra. The maxilla has a very small contribution to the external naris. The orbits are large and round, while the laterotemporal fenestrae are shorter and dorsoventrally elongate. The latter are broadest in IVPP 12430 and smallest in NGMC 97-9-A. The premaxilla is gently rounded anterodorsally, with a slightly convex ventral margin. There is an elongate narial fossa anteroventral to the external naris. The premaxilla is lower and more triangular in NGMC 97-9-A, with a narrower external naris, more vertical maxillary suture and shorter, narrower subnarial process. The naris is widest in BPM 0001, while the subnarial process is largest in IVPP V 12430. The ventral edge appears to have three notches in BPM 0001, while it is smooth in IVPP V 12430 and NGMC 97-9-A. The premaxillae are unfused and contain teeth in the anterior half. There are four procumbant teeth, the first is by far the largest. Teeth are serrationless and constricted at the base. The maxilla is reduced, with narrow ascending and posterior processes. The latter is much shorter in IVPP 12430 and thicker in BPM 0001. A single maxillary fenestra is present; there does not seem to be much of an antorbital fossa. BPM 0001 and IVPP V 12430 have an elongate anterior portion with a pneumatic fossa, while NGMC 97-9-A is blunt anteriorly without accessory fossae. Maxillae are toothless. The nasal is shorter or subequal to the frontal in length. The nasals lack rugosities and are unfused. The lacrimal is triradiate, with an elongate posterior process, suggesting the prefrontal is fused to it. While the processes are 120 degrees apart in IVPP V 12430, the anterior and posterior processes are 160 degrees apart in BPM 0001. There is a large pneumatic lacrimal foramen, but no rugosities or horns are evident. The jugal is relatively low, although not rod-like. The anterior process is more elongate in IVPP V 12430 and NGMC 97-9-A. The dorsal process is much more robust in BPM 0001 and IVPP V 12430 than in NGMC 97-9-A , but contacted the postorbital in all of them. The posterior process is shortest and not visibly bifurcated. The frontal is roughly triangular, with an elongate anterior section seemingly overlapped by the nasal. The orbital rim is raised and ventral impressions indicate a large brain. The frontals seem fused in BPM 0001, but separate in NGMC 97-9-A. The frontonasal suture appears anteriorly convex. The triradiate postorbital is much larger in BPM 0001 and IVPP V 12430 than in NGMC 97-9-A. The anterior process is expanded in the first two, while it is shorter and tapered in the latter. The posterior process is broad in the former two, but shorter and tapered in the latter. The ventral process is much more slender and elongate in NGMC 97-9-A, compared to the other two specimens. Because of these differences, I believe the unlabeled element preserved directly anterior to the detached ventral postorbital process in NGMC 97-9-A is actually the main postorbital body. It is much larger and more similar in shape to the postorbitals of BPM 0001 and IVPP V 12430, but the possibility remains it is a lacrimal or jugal. The supposed postorbital then may be the other quadratojugal. The parietal is poorly preserved in all specimens, but appears subequal to the frontal in length and quadrangular. The squamosal is also poorly preserved, but has an elongate tapered ventral process and a hooked posterior process that exposes the quadrate head laterally. The ventral process contacted the quadratojugals dorsal process. The quadratojugal is not triradiate and has a dorsal process more elongate than the anterior process. Both processes are narrow and tapered. The quadrate is single-headed, with a gently concave posterior margin and a deep notch ventrally. It is vertical and not pneumatic. The braincase is visible in BPM 0001 and NGMC 97-9-A, but shows no details besides several large foramina. The broad posterior portion of the pterygoid is preserved in BPM 0001, contacting the quadrate. Another similar element is unidentified in IVPP V 12430. An L-shaped element, tapered on one end and slightly expanded on the other, is identified as a possible pterygoid in IVPP V 12430. I can't see how this can be a pterygoid (although my grasp of three-dimensional palates is poor) and think it resembles a quadratojugal more. A large irregular element in BPM 0001 is identified as a palatine. This does not resemble any palatine I have seen, but is not triradiate. In BPM 0001, there are two pointed elements projecting posterodorsally from the anteroventral margin of the external naris that Zhou et al. identify as vomers. If that is true, they are very short compared to other theropods, as they extend only halfway past the external nares. A thin, C-shaped element in IVPP V 12430 is identified as an ectopterygoid. This is dissimilar from both the dumb-bell shaped ectopterygoids of oviraptorids or the hook-shaped ones of most theropods, although it is most similar to the latter. The ectopterygoid of NGMC 97-9-A is more robust, with the usual thickened portion seen in most theropods. Several other cranial elements are also difficult to identify. Two elongate elements preserved in the snout of IVPP V 12430 could be vomers or vomeral processes of the palatines. A vertical strap-like element, wider than the lacrimal and found in the antorbital fenestra, defies identification but is present in all three specimens. A small element in the naris of BPM 0001 is very similar to a coronoid, but its position makes this identification suspect. Curiously, two scleral plates are preserved in NGMC 97-9-A, but not in other specimens.
The dentary is toothless with two long posterior processes. The ventral process is longer than the main dentary body in BPM 0001 and IVPP 12430, but shorter in NGMC 97-9-A. The processes are subequal in length in the first two, but the dorsal process is half as long in the latter. The dorsal process is much narrower than the ventral in NGMC 97-9-A, but wider in BPM 0001 and IVPP V 12430. Anteroventrally, the dentary is concave. Medially, a shallow Mackelian groove seems to be present in NGMC 97-9-A and BPM 0001. The symphysis is well-developed, but not fused. The external mandibular fenestra is 39% of mandibular length in BPM 0001 and 33% in IVPP V 12430. The surangular and dentary may be fused in these specimens, but are loosely joined in the former. Ji et al. (1998) claim the intramandibular joint in NGMC 97-9-A was mobile, but this could not have been the case in BPM 0001 and IVPP V 12430. A ventral surangular process crosses the external mandibular fenestra in BPM 0001 and extends partway into it in NGMC 97-9-A, but not in IVPP V 12430. If a surangular foramen was present, it was very small. The angular is large, unlike oviraptorids, and cannot be distinguished from the articular. The mandibular joint is not extremely convex, unlike caenagnathoids. The retroarticular process is moderately elongate, narrower in NGMC 97-9-A. An elongate bone ventral to the surangular (and fused angular?) in IVPP V 12430 is probably the other surangular/angular in medial view, based on a posteroventral tubercle also seen in the surangular/angular of BPM 0001. A splenial is identifiable in IVPP V 12430, it is acutely triangular with a notched posterior edge. There is no indication it could be seen in lateral view. Another much larger element is identified as the splenial in BPM 0001, but appears to be a straight narrow prearticular instead. The prearticular of IVPP V 12430 is probably visible behind the dorsal dentary process. Another elongate element in BPM 0001 is more problematic. It is a bit larger than the splenial should be, triangular and has a notch posterodorsally. Perhaps it is an oddly shaped splenial. A slender, tapered hyoid is seen in NGMC 97-9-A.
An element labeled "?" by Zhou et al. located behind the skull of IVPP V 12430 looks like an atlantal neurapophysis to me. It is triangular, with a small process on one corner. Ten cervicals are reported by Ji et al., while Zhou et al. estimate twelve. Paul (2002) notes that eleven are probably present based on rib morphology. They are amphicoelous and have slender unfused ribs. The axial neural spine is prominent and expanded, the neural spine of the third cervical is tall and rectangular. More posterior cervical neural spines are low. Prominent cervicodorsal hypapophyses are said to be present. Zhou et al. report nine dorsal vertebrae, but again Paul suggests ten is a more correct number. They are procoelous and reported to lack deep pleurocoels. Currie (pers. comm., 1999) states pleurocoels are present anteriorly, but not posteriorly. Details are hard to make out, but the dorsals appear longer than Nomingia, with slightly shorter quadrangular neural spines. Nine pairs of dorsal ribs are present, the fourth the longest. Three uncinate processes are preserved in IVPP V 12344, four in BPM 0001 and six in IVPP V 12430. On the latter specimen, they are present on the first six dorsal ribs. The second, third and fourth processes are longest. They are flat, slightly curved and expanded ventrally. Gastralia are also present. There are five sacral vertebrae, unfused in IVPP V 12344. The tail contains twenty-two vertebrae, none are fused into a pygostyle. The centra lack pleurocoels and decrease in length posteriorly. Twelve to fifteen have transverse processes, the last seven have elongate prezygopophyses. It appears the centra may be grooved ventrally and are not procoelous. The last chevron is after the seventeenth caudal. Dorsoventrally elongate chevrons are present until after the tenth caudal, all but the first are distally expanded. No dromaeosaurid-like highly elongated prezygopophyses or chevrons are present.
The scapula is gently curved and gradually expanded distally to 2.4 times minimum shaft width. The acromion forms a prominent, though broad, anteriorly projecting process. The scapulocoracoid suture is broad, unfused in NGMC 97-9-A and IVPP V 12430, but fused in BPM 0001. The coracoid is subrectangular and taller than long, though not as much as in dromaeosaurids. A prominent triangular posteroventral process is present. There is a large coracoid tubercle and a foramen located near the scapulocoracoid suture. The glenoid seems to point mostly posteroventrally. The furcula is broad (interclavicular angle ~90) and U-shaped, probably without a hypocleidium. Two oval sternal plates are present, smaller than the coracoids. There is no keel or lateral processes. Several sternal ribs are preserved, they are straight, flatter than dorsal ribs and longer than the sternum.
The humerus is poorly described, but lacks a pneumatic fossa and a well-developed olecranal fossa. It is relatively straight, slender and has a low deltopectoral crest. The radius is slnder (~60-80% of ulnar width) and the ulna is bowed posteriorly. The carpus consists of a large semilunate, triangular radiale and small rounded ulnare. There are three unfused metacarpals. The first is 39-45% the length of the second. It lacks an extensor process. Phalanx I-1 is longest, so the first digit reaches well past metacarpal II. The first manual ungual is moderately curved with a well-developed flexor tubercle, but lacks a proximodorsal lip. It is slightly larger than the second manual ungual in BPM 0001 and IVPP V 12430, but smaller in NGMC 97-4-A and IVPP V 12344. Phalanx II-2 is longer than phalanx II-1. The second manual ungual is similar to the first, but has a lower flexor tubercle. Metacarpal III is very slender, straight and slightly shorter than metacarpal II (82-93%). There are only two phalanges in manual digit III. They are both very small, the second smallest, and do not reach past the midpoint of phalanx II-1. There is no ungual.
The pelvis is propubic and unfused. The ilium has a long ventrally expanded preacetabular process and rounded, posteroventrally sloped postacetabular process. The preacetabular process is 15-29% longer than the postacetabular process. The anterior margin is anterodorsally oriented in IVPP 12344, but posteroventrally oriented in IVPP V 12430. The anterodorsal margin is higher than the posterodorsal margin and is rounded anteriorly. The pubic peduncle reaches further ventrally than the ischial peduncle does. It is slightly anteroventrally oriented and notched ventrally, although this is not seen in lateral view. The m. cuppedicus fossa is shallow and reduced. The pubis is nearly straight and has an anterior foot much larger than the posterior. The pubic symphysis extends about halfway up the shaft, which is not compressed mediolaterally. The ischium is 58% of pubic length. It lacks any posterior processes, but does have a large triangular obturator process placed 60% down the shaft. It is concave posteriorly.
The greater and anterior trochantors are separated by a small notch, while the greater trochantor is well separated from the head. There is no distinct fossa for the capital ligament, and no transverse ridge bounding the popliteal fossa. The tibia is anteroposteriorly elongate when viewed proximally. The fibula is very slender, but extends distally to contact the calcaneum. The astragalus and calcaneum are not fused to the tibia, and are separate from each other as well. The ascending process of the astragalus extends 22% up the tibia and is broad and triangular. There is a shallow groove or fossa separating the process from the condyles. Two unfused distal tarsals are present. The unfused metatarsus is elongate and although the third metatarsal is very narrow, it is not arctometatarsalian. Rather, it expands proximally after reaching it's narrowest point at midlength. The first metatarsal is placed two thirds down on the posteromedial surface of the second metatarsal and has a ball-shaped distal end. The phalanx is short and stout, while the small ungual is more curved than the others. The second metatarsal is 90-91% the length of the second, the fourth is 94-95%. The second digit shows no predatory specializations- there is no proximoventral heel on phalanx II-2 and the ungual is subequal in size to the others. Unlike eumaniraptorans, the first phalanx of digit II is less than 90% of phalanx III-1. The fifth metatarsal is 27-29% of the third in length.
The body was covered with small plumulaceous feathers up to 14 mm long. At least fourteen remiges are present on metacarpal II, phalanx II-1 and phalanx II-2. They lengthen proximally (30, 63.5 and 95 mm long starting with most distal), are symmetrical and have well-developed rachis and vanes. Barbules seem not to have been present. Six retrices are present on each side of the tail (Gatesy, 2001; contra Ji et al., 1998). They are attached to the last six caudals. These are also symmetrical. Small rounded gastroliths are preserved in all specimens, measuring up to 4.5 mm in diameter, although most are less than 4 mm.
Comparison of specimens- Five specimens of Caudipteryx have been described. NGMC 97-4-A, NGMC 97-9-A and BPM 0001 are referred to the type species, C. zoui (Ji et al., 1998; Zhou et al., 2000). IVPP V 12344 was referred to a new species, C. dongi (Zhou and Wang, 2000). IVPP V 12430 was referred simply to C. sp. (Zhou et al., 2000). Zhou and Wang differetiated C. dongi from C. zoui based on the smaller sternum and longer first metacarpal. Most differences I can see between the specimens are cranial, although this may be due to the fact the skulls are well illustrated, while the postcrania is not. Are these differences real or preservational? A large amount of the variety seems to be due to crushing and distortion. For instance, there is no way the lacrimal of IVPP V 12430 could have had such a small angle between its anterior and posterior processes in life. Similarily, the posterior postorbital process of BPM 0001 is much too long, as it would extend well past the quadrate when articulated. The anterior squamosal process of that specimen is much too large and bulbous, as it would reach through the postorbital and into the orbit. More evidence that distortion has occured might come from the asymmetry in specimens. The dorsal cranial elements (nasal, frontal, parietal) are often distorted and asymmetrical. The differences least likely to be due to distortion or individual variation support BPM 0001 and IVPP V 12430 being separate from NGMC 97-9-A. Characters these two specimens share not found in the latter are- premaxilla blunt anteriorly; large premaxillary subnarial process; maxilla extended anteriorly with promaxillary fossa; external naris close to antorbital fenestra in length; jugal strongly concave posterodorsally; quadratojugal posterior process not developed; posterodorsal dentary process subequal in width to posteroventral process; posterodorsal dentary process longer than posteroventral process; no intramandibular joint. Most of the postcranium is not figured in sufficient detail to determine morphological differences in specimens. The differing number of reported cervical vertebrae might be due to misinterpretation, as Zhou et al. state "there are estimated twelve cervical vertebrae". The coracoid has a smoothly rounded ventral border in NGMC 97-9-A, unlike the irregular border of BPM 0001, although the significance of this is uncertain. The orientation of the anterior preacetabular edge differs in IVPP V 12344 and IVPP V 12430, but as the skull of the former is fragmentary, it cannot be determined if this is correlated with the cranial differences noted above. Contra Zhou and Wang, no significant differences in postcranial ratios is evident. Most ratios vary within a few percentage points of each other, so fall within the expected range of individual variation. The sternal plates are 24% of femoral length in the holotype of C. zoui, 17% in the holotype of C. dongi and an intermediate 21% in BPM 0001. A three percent difference in size does not seem to fall outside the range of individual variation. The first metacarpal of BPM 0001 and IVPP V 12430 is 39% of metacarpal II length. In IVPP V 12344, the ratio is 45%. This difference might be considered diagnostic if not for NGMC 97-4-A, which has a 42% ratio. Although stated to be "about .4" in Zhou and Wang, this figure comes from Ji et al., who only measured to the tenths place. The exact ratio, as mentioned above, is intermediate between the more divergent specimens. Once again, the 3% difference is considered insufficient to diagnose a species. The ilium is much shorter in NGMC 97-4-A (69% of femoral length) than in BPM 0001, IVPP V 12344 and IVPP V 12430 (77-79%). This is due to the broken anterior edge in the former specimen however, as can be seen in the specimen (pers. obs.). The only potentially significant proportional difference between specimens is- manual ungual I vs. manual ungual II (106% in BPM 0001 and IVPP V 12430, 84% in IVPP 12344). Although the first manual ungual of NGMC 97-4-A is incomplete, it was much smaller than manual ungual II, so seems to match IVPP V 12344 better. It therefore seems that BPM 0001 and IVPP V 12430 share several cranial characters not seen in NGMC 97-9-A; NGMC 97-9-A has a slightly different coracoid morphology than BPM 0001; BPM 0001 and IVPP V 12430 have a different preacetabular morphology than IVPP V 12344; and that BPM 0001 and IVPP V 12430 have different manual ungual ratios than IVPP V 12344 and NGMC 97-4-A. Two groups of specimens are suggested by these differences- IVPP V 12344, NGMC 97-4-A and NGMC 97-9-A are one group, while BPM 0001 and IVPP V 12430 are in the other. The inclusion of IVPP V 12344 and NGMC 97-4-A with NGMC 97-9-A is far from certain, but the fact they all differ from the other two specimens and that the latter two have similar ungual ratios suggests this may be the case. Are these differences due to ontogenetic, sexual or taxonomic variation? The minute size variation (femora vary between145-152 mm) suggests it is not ontogenetic. Settling whether two sexes or species are involved is not easily resolved with only five specimens to work with, all from different localities. One potential way to decide this would be if the groups are not sister groups in a phylogenetic analysis.
Relationships- Caudipteryx has been included in several phylogenetic analyses. Ji et al. (2005) added it to Clarke's bird matrix, which resulted in it being excluded from Microraptor+Aves.
Caudipteryx in Avialae?
Ji et al. (1998) first included Caudipteryx in a matrix of birds, alvarezsaurids and Protarchaeopteryx, with Velociraptor as an outgroup. Note this doesn't allow Caudipteryx to fall outside of Eumaniraptora. Also note it was only examined with characters that were thought to be useful for analyzing Avialae. With these caveats in mind, Ji et al. found it to be an avialan based on two characters- unserrated teeth; premaxilla reaches to anterior border of antorbital fossa. As only the premaxilla is toothed in Caudipteryx, and premaxillary teeth are plesiomorphically unserrated in maniraptoriforms (only derived dromaeosaurids and derived troodontids have serrations; Protarchaeopteryx was miscoded as having them), this doesn't support placing Caudipteryx in Avialae instead of Oviraptorosauria. The second character is also seen in Erlikosaurus and some other oviraptorosaurs (Avimimus, oviraptorids), though not all (absent in Incisivosaurus and the CMNH caenagnathid). Even Currie (1999 Ostrom Symposium) now places it as an oviraptorosaur.
Those who doubted the dinosaur-bird link have always said Caudipteryx is a bird due to its unambiguous remiges and retrices. Originally, this meant separating it from other non-avialan maniraptoriforms, which they viewed as dinosaurs. Since 2002 however, as more maniraptoriforms are discovered with remiges and retrices, workers such as Feduccia and Martin have allowed oviraptorosaurs, dromaeosaurids, troodontids, and possibly even alvarezsaurids and ornithomimosaurs to be birds as well (though they still insist therizinosaurs are sauropodomorphs). Thus their arguments for placing Caudipteryx as a bird (e.g. Martin and Czerkas, 2000; Geist and Feduccia, 2000; Ruben and Jones, 2000) are no longer valid, as they now think some taxa which lack these bird-like characters (e.g. Velociraptor) are birds anyway. Similarily, Feduccia et al. (2005) and Martin (2004) now agree Caudipteryx is a basal oviraptorosaur, though their placement of oviraptorosaurs and other maniraptorans outside of Theropoda remains incorrect.
Martin and Czerkas (2000) argued Caudipteryx was a sauriurine more closely related to Confuciusornis than to Archaeopteryx. It is uncertain whether they considered enantiornithines to be the sister taxon to Confuciusornis or to Caudipteryx+Confuciusornis. Of their supporting characters, a reduced fibula, reduced calcaneum, and "evidence of pygostyle formation" are not present in Caudipteryx (Ji et al., 1998; Zhou et al., 2000), while a reduced maxilla is not present in Confuciusornis. A reduced hypopubic cup is a completely fictional character, as no bird has a hypopubic cup. The external mandibular fenestra and ball-shaped femoral head are plesiomorphies only absent in a few basal birds (Archaeopteryx, Rahonavis, Shenzhouraptor, etc.), but present in enantiornthines in addition to confuciusornithids. On the other hand, the enlarged premaxilla, toothless maxilla and dentary, and shortened tail are shared with Confuciusornis but not Archaeopteryx. However, oviraptorosaurs besides Incisivosaurus share the enlarged premaxilla, oviraptorosaurs besides Incisivosaurus and Protarchaeopteryx also lack maxillary and dentary teeth, and all oviraptorosaurs share shortened tails. Indeed, Martin (2004) suggests Caudipteryx and other oviraptorosaurs may be most closely related to confuciusornithids. This is highly unparsimonious though, as oviraptorosaurs lack numerous paravian/eumaniraptoran, avialan, ornithurine sensu Gauthier, avebrevicaudan and pygostylian characters found in confuciusornithids; basal oviraptorosaurs such as Incisivosaurus and Caudipteryx lack most of the confuciuornithid-like characters found in derived oviraptorosaurs; and confuciusornithids in turn lack most oviraptoriform and oviraptorosaur characters.
Lu et al. (2002) found Caudipteryx to be an ornithurine (sensu Gauthier) basal to alvarezsaurids, oviraptorids and ornithothoracines. Similar to Ji et al.'s analysis, Velociraptor was the outgroup, so Caudipteryx was forced inside Eumaniraptora and only examined using characters thought to be useful for analyzing Avialae. This analysis is plagued by a large number of miscodings. When several were corrected, Caudipteryx was resolved in a trichotomy with Velociraptor and Avialae. Constraining Caudipteryx to be the sister taxon to Oviraptoridae, with both outside Aves (sensu Chiappe), only took five more steps. These are easily accounted for when you consider that Lu et al. did not include any explicitly oviraptorosaurian characters in the matrix.
There is a chance oviraptorosaurs and taxa such as confuciusornithids, omnivoropterygids and/or jeholornithids are more closely related than current phylogenies suggest, but this has yet to be supported by cladistic analyses.
Caudipteryx in Paraves?
Xu et al. (1999) resolved Caudipteryx as a paravian in a trichotomy with Protarchaeopteryx and Troodontidae+Eumaniraptora. Xu et al. (2000) used 86 of the same characters plus three new ones, added Microraptor and removed Tyrannosauridae and Unenlagia. Their tree was slightly more resolved, as Caudipteryx was found to be outside a clade consisting of Protarchaeopteryx and Troodontidae+Eumaniraptora. However, both studies supported the paravian status of Caudipteryx using the same ten characters.
- deep suborbital jugal. This also present in Harpymimus, Erlikosaurus, Incisivosaurus and the CMNH caenagnathid (none of which were included in the analysis). It is therefore probably primitive for Maniraptoriformes, and its distribution in the analysis is due to the lack of included basal taxa.
- unfused interdental plates. These are also present in therizinosaurs, and are recovered as derived for Paraves based only on their absence in the toothless ornithomimids and oviraptorids directly basal to this clade, plus their coded presence in Caudipteryx and Protarchaeopteryx. Notably, every other matrix has coded the latter two taxa unknown for this character.
- less than 26 caudal vertebrae. Xu et al.'s scoring for caudal vertebral number is flawed due to the gap between >30 vertebrae (state 0) and >26 vertebrae (state 1). Oviraptorids, troodontids and Microraptor (and possibly Protarchaeopteryx) fall within this uncoded range. Caudipteryx does have less caudals than oviraptorids (22 vs. 27-32), as do Protarchaeopteryx (~24-27), Archaeopteryx (21-23), basal troodontids (24 in Jinfengopteryx) and Microraptor (24-26). However, the long tail of scansoriopterygids and most therizinosaurs (neither included by Xu et al.) indicates it may have developed this convergently with paravians or Aves sensu Chiappe.
- less than 11 caudal vertebrae with transverse processes. Caudipteryx is miscoded and actually has thirteen caudals with transverse processes.
- mid and distal chevrons dorsoventrally flattened. Though the character is vague, Caudipteryx does not differ from oviraptorids such as Citipati in this regard.
- proximodistally elongate coracoid. Oviraptorids and troodontids are miscoded as lacking this character, as it is actually a maniraptoran synapomorphy.
- radius less than 70% of ulnar width. This is also present in Falcarius and Protarchaeopteryx, suggesting it may be primitive for maniraptorans. However, there is much homoplasy.
- metacarpal I less than 33% of metacarpal II in length. The ratio in Caudipteryx is actually 39%.
- pubic foot projects caudally only. This is miscoded for Caudipteryx, which has a larger anterior projection than posterior one.
- posterior trochanter present. This is miscoded as being present in Caudipteryx.
When these and other miscodings are corrected, the topology is changed. In Xu et al. (1999), Protarchaeopteryx is the sister taxon of Caudipteryx based on the low number of caudal vertebrae (<27) and the supposed presence of unfused interdental plates. It and Caudipteryx are oviraptorosaurs based on the pneumatic lacrimal, frontal and parietal which are subequal in length, and pubic foot which is developed anteriorly. In Xu et al. (2000), Caudipteryx is a paravian in a trichotomy with Protarchaeopteryx and Troodontidae+Eumaniraptora (like in the uncorrected Xu et al., 1999 results). It's a paravian based on supposedly unfused interdental plates, the low number of caudal vertebrae (<27), the slender radius (<70% of ulnar width), and the shortened ischium (<66% of pubic length). Regarding the latter, other non-oviraptorid oviraptorosaurs exhibit it as well (e.g. Nomingia, Avimimus, Chirostenotes). Based on the above information, this is probably due to the absence of therizinosaurs (especially Falcarius), additional oviraptorosaurs, and oviraptorosaurian synapomorphies in the matrix. Constraining Caudipteryx to be an oviraptorosaur results in trees one step longer.
Caudipteryx outside Oviraptorosauria+Paraves?
Holtz (2001) found Caudipteryx to either be a basal oviraptoriform or a basal maniraptoran sister to Oviraptoriformes+Paraves (sometimes in a clade with Protarchaeopteryx). However, later more extensive analyses of Holtz's (e.g. Holtz et al., 2004) have placed it in Oviraptorosauria.
Gishlick (2002) found Caudipteryx to be in a trichotomy with therizinosaurs and the Oviraptorosauria+Paraves clade, but the matrix only contained forelimb characters.
Caudipteryx in Oviraptorosauria?
Sereno (1999) was the first to recover Caudipteryx as an oviraptorosaur basal to Caenagnathoidea. This was based on twelve characters.
- ventral margin of external naris dorsal to maxilla. Not true in Caudipteryx, as can be clearly seen in BPM 0001.
- premaxilla participates in antorbital fossa. There does not appear to be much of an antorbital fossa in Caudipteryx. The promaxillary fossa is clearly separated from the premaxilla by a raised anterior maxillary rim.
- nasal shorter than frontal. The frontal length is difficult to determine in Caudipteryx. The nasals are shorter in NGMC 97-9-A, but longer in BPM 0001. The frontals are impossibly long in IVPP V 12430, as the orbit would be intersected by the jugal dorsal process. This suggests the long anterior portion of the frontals was overlapped by the nasals. This would also explain why the nasals are longer in BPM 0001, which is articulted, while the disarticulated NGMC 97-9-A shows the opposite condition. Thus, when articulated, the nasals would appear shorter than the frontals.
- ventral margin of maxilla inset medially for dentary and surangular. This would be very difficult to determine from crushed two-dimensional specimens such as are preserved at Liaoning. Moreover, in BPM 0001 and IVPP V 12430, the maxilla ventrally overlaps the dentary and surangular on their lateral sides, suggesting the opposite condition.
- anterior process of jugal rod-shaped in cross section. Rauhut (2000) codes this as having the opposite state, though Maryanska et al. (2002) and Holtz et al. (2004) agree with Sereno.
- palate mostly ventral to maxilla and jugal. This is also not easy to determine in Caudipteryx. However, I consider it unlikely based on a few observations. First, there is no indication of a ventrally projecting maxillary palate. Secondly, the vomers may be incredibly short and posterodorsally projected, in no position to reinforce maxillary "teeth". Finally, the C-shaped ectopterygoid would not be able to function like a strut between the maxilla and pterygoid.
- dentary dorsal margin convex. This is also present in therizinosauroids, so may be an oviraptoriform synapomorphy instead (though it is absent in Falcarius).
- external mandibular fenestra 40% of lower jaw length. Only confuciusornithids have such long mandibular fenestrae. Oviraptorids vary from 26-36%, while Chirostenotes has a 27% ratio and Caudipteryx varies between 33-39%. Change the ratio to 25% and it becomes a defensible oviraptorosaur synapomorphy.
- caudal centra twice as wide as tall. The disarticulated caudal vertebrae of NGMC 97-9-A (visible in ventral view) show this not to be the case in Caudipteryx.
- anterior caudal transverse processes (1-8) twice length of neural spines. This is hard to determine when the vertebrae are preserved in lateral view, but the eighth caudal of NGMC 97-9-A is preserved in anterior (or posterior) view and seems to have transverse processes subequal in length to the neural spine.
- acromion prong projected anteriorly or anterodorsally. This is valid, although paralleled in most eumaniraptorans.
So most of Sereno's evidence for referring Caudipteryx to the Oviraptorosauria is not valid. An external mandibular fenestra over 25% of mandibular length and an anteriorly or anterodorsally projected acromial prong do support this assignment.
Rauhut (2000, 2003) found Caudipteryx to be an oviraptorosaur more basal than Avimimus and Caenagnathoidea based on- preorbital region of the skull significantly shortened; posterior end of dentary strongly forked; maxillary and dentary teeth absent; arctometatarsalian pes. The pes is technically subarctometatarsalian, but otherwise the characters seem valid.
The many variations of the Theropod Working Group have always placed Caudipteryx within Oviraptorosauria. Specific positions include- oviraptorid sister to oviraptorines but more derived than Microvenator (Norell et al., 2001; Clark et al., 2002); basal oviraptorosaur more derived than Incisivosaurus, but outside Avimimus+Caenagnathoidea (Xu et al., 2002; Hwang et al., 2004; Lu, 2004; Xu and Norell, 2004; Novas and Pol, 2005; Xu and Zhang, 2005; Norell et al., 2006; Turner et al., 2007); basal oviraptorosaur outside Avimimus+Caenagnathoidea (Makovicky et al., 2003); oviraptorosaur with an uncertain position outside Oviraptorinae (Xu et al., 2002; Kirkland et al., 2005; Mayr et al., 2005).
Maryanska et al. (2002) found Caudipteryx to be an oviraptorosaur more derived than Avimimus, but outside of Caenagnathoidea. This was based only on a strongly concave caudal margin of the ischiadic shaft. Lu's (2004) expanded version of this dataset agreed, also placing Incisivosaurus basal to Caudipteryx.
Senter (2003) found Caudipteryx to be an oviraptorosaur more derived than Protarchaeopteryx+Incisivosaurus, but outside an Avimimus+Oviraptoridae clade. Senter et al. (2004) had the same topology but did not include Avimimus.
Holtz et al. (2004) recovered Caudipteryx as a basal oviraptorosaur outside Caenagnathoidea.
Osmolska et al. (2004) recovered Caudipteryx as an oviraptorosaurs less derived than caenagnathoids, but more derived than Avimimus.
Senter (2007) found Caudipteryx to be an oviraptorid, in a trichtomy with oviraptorines and Microvenator. Incisivosaurus, Protarchaeopteryx, Caenagnathus (excluding Chirostenotes specimens) and Avimimus were recovered as more basal.
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Xu and Zhang, 2005. A new maniraptoran dinosaur from China with long feathers on the metatarsus. Naturwissenschaften.
Norell, Clark, Turner, Makovicky, Barsbold and Rowe, 2006. A New Dromaeosaurid Theropod from Ukhaa Tolgod (Omnogov, Mongolia). American Museum Novitates. 3545, 51 pp.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
Turner, Hwang and Norell, 2007. A small derived theropod from Oosh, Early Cretaceous, Baykhangor Mongolia. American Museum Novitates. Number 3557, 27 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.
Feduccia and Czerkas, 2015. Testing the neoflightless hypothesis: Propatagium reveals flying ancestry of oviraptorosaurs. Journal of Ornithology. 156(4), 1067-1074.

Avimimidae Kurzanov, 1981
Definition- (Avimimus portentosus <- Oviraptor philoceratops, Elmisaurus rarus, Caenagnathus collinsi) (Martyniuk, 2012)
Comments- Another possible avimimid is Kol.
References- Kurzanov, 1981. On the unusual theropods from Upper Cretaceous of Mongolia. In Resetov (ed.). Iskopaemye pozvonocnye Mongolii. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 15, 39-50.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.

Microvenator Ostrom, 1970
= "Megadontosaurus" Brown vide Ostrom, 1970
M. celer Ostrom, 1970
= "Megadontosaurus ferox" Brown vide Ostrom, 1970
Late Aptian, Early Cretaceous
Cloverly Formation, Montana, US

Holotype- (AMNH 3041) (~3.7 kg; juvenile) lacrimals(?) (one lost), quadrates(?) (lost), dentary, prearticular (?) (lost), axis (13.9 mm), anterior cervical neural arch, cervical centrum (15.1 mm), cervical centrum (15.4 mm), cervical centrum (15+ mm), two cervical neural arches, two partial cervical ribs, three anterior dorsal neural arches, anterior dorsal centrum (13.5 mm), anterior dorsal centrum (13.8 mm), nine posterior dorsal neural arches, posterior dorsal centrum (14.2+ mm), posterior dorsal centrum (13.8 mm), posterior dorsal centrum (13.9 mm), posterior dorsal centrum (14.1 mm), posterior dorsal centrum (13.7 mm), posterior dorsal centrum (13.1 mm), two partial dorsal ribs, mid sacral centrum (15.1 mm), caudal centrum (11.5 mm), caudal centrum (9.6+ mm), caudal centrum (8.7 mm), caudal centrum (6.1+ mm), caudal centrum (9.9+ mm), caudal centrum (9.8 mm), caudal centrum (9.7 mm), caudal centrum (9.5 mm), caudal centrum (10.1 mm), three caudal neural arches, coracoid, humerus (81.5 mm), radius, ulna (78 mm), ulnar fragment, metacarpal I (12.5 mm), phalanx I-1 (34 mm), manual ungual I (25 mm), manual ungual III (11 mm), distal phalanx, partial ilia (~110 mm), pubes (108.6 mm), fragmentary ischia, femora (124 mm), tibia (157 mm), proximal fibula, astragalus (22.5 mm wide, 42 mm high), metatarsal I (11.8+ mm), pedal ungual I (11.9 mm)
Comments- Brown originally believed 25 large teeth to belong to this specimen, resulting in the informal name "Megadontosaurus". This was first published by Ostrom (1970), who realized the teeth belonged to Deinonychus they are still catalogued with the Microvenator holotype. The tooth (YPM 5366) questionably referred to Microvenator by Ostrom is unlikely to belong to an oviraptorid and is more probably a basal coelurosaur similar to Nedcolbertia.
This genus has a controversial placement in recent analyses, from being a caudipterid in Senter et al. (2012), avimimid in Lee et al. (2014; the position preferred here), sister to caenagnathoids in Brusatte et al. (2014) and Foth et al. (2014), and basal caenagnathid in Lu et al. (2015).
References- Ostrom, 1970. Stratigraphy and paleontology of the Cloverly Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Peabody Mus. Nat. Hist., Yale Univ., Bull. 35, 234 pp.
Makovicky and Sues, 1997. A reappraisal of the phylogenetic affinities of Microvenator celer (Theropoda: Dinosauria) from the Cloverly Formation. Journal of Vertebrate Paleontology. 17(3), 62A.
Makovicky and Sues, 1998. Anatomy and phylogenetic relationships of the theropod dinosaur Microvenator celer from the Lower Cretaceous of Montana. American Museum Novitates. 3240, 27 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.
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.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Avimimus Kurzanov, 1981
A. portentosus Kurzanov, 1981
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia

Holotype- (PIN 3907/1) partial skull, axis (17 mm), fourth cervical vertebra, sixth cervical vertebra, eighth cervical vertebra, ninth cervical vertebra (24 mm), tenth cervical vertebra, eleventh cervical vertebra, first dorsal vertebra (~22 mm), second dorsal vertebra (~23 mm), third dorsal vertebra (~24 mm), fourth dorsal vertebra (~21 mm), fifth dorsal vertebra (~24 mm), sixth dorsal vertebra (~24 mm), seventh dorsal vertebra (~24 mm), eighth dorsal vertebra (~23 mm), ninth dorsal vertebra (~24 mm), tenth dorsal vertebra (~24 mm), two sacral ribs, partial scapulocoracoid, humerus (95 mm), proximal ulna, proximal carpometacarpus, partial ilia, pubes (~166 mm), partial ischium, femur (188 mm), tibiotarsus (257 mm), fibula, tarsometatarsus (II 143 mm, III 153 mm, IV 144 mm, V 28 mm), phalanx II-1 (~19 mm), phalanx II-2 (~13 mm), pedal ungual II (~27 mm), phalanx III-1 (~30 mm), phalanx III-2 (~21 mm), phalanx III-3 (~16 mm), pedal ungual III (~29 mm), phalanx IV-1 (~14 mm), phalanx IV-2 (~9 mm), phalanx IV-3 (~8 mm), phalanx IV-4 (~7 mm), pedal ungual IV (~28 mm)
Paratype- (PIN 3906/1) postcranial fragments including ilial fragment, proximal ischium
Referred- (IGM coll.) incomplete skeleton including incomplete skull, dorsal ribs, articulated caudal vertebrae, scapulocoracoid, sternum, humerus, radius, ulna, radiale, ulnare, carpometacarpi, phalanx II-2, pelvis, hindlimb (Watabe et al., 2000)
(PIN 3907/2) ilial fragments, proximal pubis, proximal ischium (Kurzanov, 1983)
(PIN 3907/3) anterior premaxillae, posterior skull, (mandible ~85 mm) anterior dentary, posterior mandible, skeleton including axis, tenth cervical vertebra (Kurzanov, 1985)
(PIN 3907/4) fragmentary skeleton including axis, third cervical vertebrae, fourth cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, fourth dorsal vertebra (Kurzanov, 1987)
(PIN 3907/5) sacrum, ilium (~185 mm), proximal pubis, proximal ischium (Kurzanov, 1987)
(PIN 3907/6) vertebrae including sacrum (Kurzanov, 1987)
(ZPAL MgD-I/85) proximal tarsometatarsus (Osmolska, 1981)
(at least ten individuals; subadults and adults) including cranial elements, mandibular elements, femora, tibiotarsi and tarsometatarsi (Currie et al., 2008)
Late Cretaceous
Mongolia

(960822 ShT ENK) fragmentary skull, fragmentary skeleton (Watabe and Suzuki, 2000)
(960822 ShT SZK) pelvis, hindlimb elements (Watabe and Suzuki, 2000)
Comments- Watabe et al. (2006) note Kurzanov (1981) was probably mistaken regarding the type locality of Avimimus, with Yagaan Khovil in the Nemegt Formation being more probable than Udan Sayr in the Djadockta Formation.
Kurzanov (1981) reported two specimens of his new taxon Avimimus and described the holotype. He went on to describe the forelimb of the holotype (1982), the pelvis of the holotype and PIN 3907/2 (1983), and the skull of PIN 3907/3 (1985). His 1987 monograph described these specimens and additional ones (PIN 3907/4, 3907/5 and 3907/6) in detail, though it has yet to be translated from Russian.
A proximal tarsometatarsus was described as Theropoda indet. by Osmolska (1981), and later identified as Avimimus (Osmolska pers. comm. to Currie, 1987; in Currie, 1989).
Watabe et al. (2000) describe a nearly complete skeleton found in 1996 that corroborates Kurzanov's identification of several features (large narial fossa in the premaxilla; narrow scapula and large coracoid; very sharp and thin posterior ulnar ridge; completely fused carpometacarpus) in addition to revealing new anatomical information (small premaxillary teeth; anteroposteriorly elongate caudals with no evidence for a pygostyle; very narrow rod-like radius). They also attributed several thousand tracks in the same locality to Avimimus.
Currie et al. (2008) reported an Avimimus bonebed discovered in 2006, which they interpreted as evidence of gregarious behavior. Subadult remains lacked tibiotarsal and tarsometatarsal fusion.
References- Kurzanov, 1981. On the unusual theropods from Upper Cretaceous of Mongolia. In Resetov (ed.). Iskopaemye pozvonocnye Mongolii. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 15, 39-50.
Osmolska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Kurzanov, 1982. [Peculiarities of the structure of the anterior extremities of Avimimus] [in Russian]. Paleontologicheskii zhurnal. 24, 108-112.
Kurzanov, 1982. Structural characteristics of the fore limbs of Avimimus. Paleontological Journal. 16, 108-112.
Kurzanov, 1983. Avimimus and the problem of the origin of birds [in Russian]. In Resetov (ed.). Iskopaemye reptilii Mongolii. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 24, 104-109.
Kurzanov, 1983. [New data on the structure of the pelvis of Avimimus.] [in Russian]. Paleontologicheskii zhurnal. 4, 115-116.
Kurzanov, 1983. New data on the pelvic structure of Avimimus. Paleontological Journal. 17, 110-111.
Kurzanov, 1985. [The skull structure of the dinosaur Avimimus.] [in Russian]. Paleontologicheskii zhurnal. 1985, 81-89.
Kurzanov, 1985. The skull structure of the dinosaur Avimimus. Paleontological Journal. 19, 92-99.
Kurzanov, 1985. [The osteology of Avimimus portentosus and the problem of the origin of birds.] [in Russian]. Akademiya Nauk SSSR, Paleontologicheskiy Institut, Moscow. 23 pp.
Kurzanov, 1987. Avimimidae and the problem of the origin of birds [in Russian]. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 31, 1-95.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26, 1319-1324.
Norman, 1990. Problematic Theropoda: "Coelurosaurs". 280-305. in Weishampel, et al. (eds.). The Dinosauria. University of California Press, Berkeley, Los Angeles, Oxford.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Univ. Copenhagen, 311pp.
Dyke and Thorley, 1998. Reduced cladistic consensus methods and the avian affinities of Protoavis and Avimimus. Archaeopteryx. 16, 123-129.
Watabe and Suzuki, 2000. Report on the Japan-Mongolia Joint Paleontological Expedition to the Gobi desert, 1996. in Results of the Hayashibara Museum of Natural Sciences, Mongolian Academy of Sciences, Mongolian Paleontological Center, Joint Paleontological Expedition, n. 1, Hayashibara Museum of Natural Sciences, Research Bulletin. 1, 58-68.
Watabe, Weishampel, Barsbold, Tsogtbaatar and Suzuke, 2000. New nearly complete skeleton of the bird-like theropod, Avimimus, from the Upper Cretaceous of the Gobi Desert, Mongolia. Journal of Vertebrate Paleontology. 20(3), 77A.
Vickers-Rich, Chiappe and Kurzanov, 2002. The enigmatic birdlike dinosaur Avimimus portentosus: Comments and a pictorial atlas. in Chiappe and Witmer (eds.). Mesozoic Birds: Above the Heads of Dinosaurs. 65-86.
Watabe, Suzuki and Tsogtbaatar, 2006. Geological and geographical distribution of bird-like theropod, Avimimus in Mongolia. Journal of Vertebrate Paleontology. 26(3), 136A-137A.
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.
Tsuihiji, Witmer, Watabe, Barsbold and Tsogtbaatar, 2008. New information on the cranial anatomy of Avimimus portentosus (Dinosauria: Theropoda) including virtual endocasts of the brain and inner ear. Journal of Vertebrate Paleontology. 28(3), 153A.
Tsuihiji, Watabe, Gishlick, Barsbold and Tsogtbaatar, 2009. New information on the pectoral girdle and forelimb of Avimimus (Dinosauria: Theropoda) from the Gobi Desert of Mongolia. Journal of Vertebrate Paleontology. 29(3), 192A.
A. sp. (Ryan, Currie and Russell, 2001)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China

Material- (AMNH 6570) fibula (Chiappe, Norell and Clark, 2002)
(AMNH 6576) proximal caudal vertebra (Makovicky, 1995)
(AMNH coll.) dorsal vertebrae (Makovicky, 1995)
(IVPP 160788-122) sacral fragment (Makovicky, 1995)
(IVPP 160788-124) posterior dorsal vertebra (Makovicky, 1995)
(IVPP 180788-123) tenth cervical vertebra (Makovicky, 1995)
(?IVPP coll.) tarsometatarsus (Dong, 1992)
(IVPP coll.) dorsal vertebrae (Makovicky, 1995)
(PIN coll.) material (Currie and Eberth, 1993)
(RTMP 92.302.102) proximal tarsometatarsus (Ryan, Currie and Russell, 2001)
(RTMP 92.302.104) partial frontal (Ryan, Currie and Russell, 2001)
(RTMP 92.302.110) distal femur (Ryan, Currie and Russell, 2001)
(RTMP 92.302.116) partial scapulocoracoid (Ryan, Currie and Russell, 2001)
(RTMP 92.302.117) proximal humerus (Ryan, Currie and Russell, 2001)
(RTMP 92.302.119) two pedal unguals (Ryan, Currie and Russell, 2001)
(RTMP 92.302.140) anterior dorsal vertebra (Ryan, Currie and Russell, 2001)
(RTMP 92.302.149) proximal femur (Ryan, Currie and Russell, 2001)
(RTMP 92.302.150) proximal tibia (Ryan, Currie and Russell, 2001)
(RTMP 92.302.344) mid caudal vertebra (Ryan, Currie and Russell, 2001)
distal caudal vertebra (Makovicky, 1995)
Description- The orbital margin of a frontal (RTMP 92.302.104) is difficult to compare to A. portentosus, but is similarily bulbous over the orbits. There is an anterior dorsal (RTMP 92.302.140) that resembles the second of Avimimus, but has a shorter hypapophysis and more ventrally placed parapophyses. Another vertebra (RTMP 92.302.344) was not identified specifically, but appears to be a mid caudal. There seem to be two small lateral foramina, the centrum is not quadrangular in section and a low neural spine is present. A partial fused scapulocoracoid (RTMP 92.302.116) is shown, with a ventrally directed glenoid and low coracoid tubercle. It is very comparable to the holotype, but more incomplete, lacking the ventral coracoid tip, most of the anterior edge and all but the base of the scapular shaft. A proximal humerus (RTMP 92.302.117) is extremely similar to the holotype, differing only in minor proportional details. Both proximal (RTMP 92.302.149) and distal (RTMP 92.302.110) femoral ends are known. The former differs from the holotype in the more lateromedially compressed greater trochantor and less prominent anterior trochantor in proximal view. The distal femur has less extensive articular surfaces in anterior view and a less prominent lateral condyle. In distal view, it is less convex anteriorly. A proximal tibia (RTMP 92.302.150) is quite different from A. portentosus, having a bulbous lateral condyle, less dorsally projected cnemial crest, and small posterior process in proximal view. The proximal metatarsus (RTMP 92.302.102) is very similar anteriorly and posteriorly, but is differently shaped proximally, being parallelogram-like. Two pedal unguals (RTMP 92.302.119A and B) are quite odd. They are markedly asymmetrical, having smaller lateral halves with much higher grooves on that side.
Comments- Kurzanov (1987) initially reported an avimimid femur (PIN 2549-100) from the Iren Dabasu Formation, but Osmolska (1996) noted it resembled Bagaraatan more closely. Dong (1992) reported Avimimus tarsometatarsi from the Iren Dabasu Formation of China. Currie and Eberth reported numerous Iren Dabasu avimimid bones in the AMNH, IVPP and PIN collections. They noted the elements were identical to A. portentosus and under study by Currie, Zhao and Kurzanov (in prep.). Makovicky (1995) described several vertebrae from the AMNH and IVPP collections. Ryan et al. (2001) described numerous elements from the RTMP as belonging to A. portentosus. Another metatarsus (AMNH 6755) and a third metatarsal (AMNH 6754) were also referred to Avimimus by Ryan et al., but seem more similar to caenagnathids. A fibula (AMNH 6570) was illustrated and referred to Mononykinae by Chiappe et al. (2002), but Longrich and Currie (2009) found it resembled Avimimus more closely.
References- Kurzanov, 1987. Avimimidae and the problem of the origin of birds [in Russian]. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 31, 1-95.
Dong, 1992. Dinosaurian Faunas of China. China Ocean Press, Beijing.
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.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Univ. Copenhagen, 311pp.
Osmolska, 1996. An unusual theropod dinosaur from the Late Cretaceous Nemegt Formation of Mongolia. Acta Palaeontologica Polonica. 41, 1-38.
Ryan, Currie and Russell, 2001. New material of Avimimus portentosus (Theropoda) from the Iren Dabasu Formation (Upper Cretaceous) of the Erenhot Region of Inner Mongolia. Journal of Vertebrate Paleontology. 21(3), 95A.
Chiappe, Norell and Clark, 2002. The Cretaceous, short-armed Alvarezsauridae, Mononykus and its kin. pp. 87-120. in Chiappe and Witmer (eds.). Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press, Berkeley, Los Angeles, London.
Longrich and Currie, 2009. Albertonykus borealis, a new alvarezsaur (Dinosauria: Theropoda) from the Early Maastrichtian of Alberta, Canada: Implications for the systematics and ecology of the Alvarezsauridae. Cretaceous Research. 30(1), 239-252.
A? sp. (Jerzykiewicz and Russell, 1991)
Late Campanian, Late Cretaceous
Djadokhta Formation, Mongolia

Reference- Jerzykiewicz and Russell, 1991. Late Mesozoic stratigraphy and vertebrates of the Gobi Basin. Cretaceous Research. 12, 345-377.
A. sp. (Ryan, Currie and Russell, 2001)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada

Material- (RTMP 98.68.22) distal metatarsal III (Ryan, Currie and Russell, 2001)
(RTMP coll.) vertebrae, tarsometatarsi, unguals (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada

(RTMP or UALVP coll.) (unassociated) metatarsal III, distal metatarsal, ?pedal phalanx (Eberth and Currie, 2010)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada

(RTMP 98.8.28) metatarsal II (Ryan and Russell, 2001)
Comments- Ryan et al. (2001) found that although there are twelve elements in the RTMP collections labeled Avimimidae, only two can certainly be assigned to that taxon. One, RTMP 98.68.22, is a distal third metatarsal from the Dinosaur Park Formation. The other (RTMP 98.8.28) is a second metatarsal from the Scollard Formation that is unfused proximally, unlike A. portentosus. It is from the Scollard Formation. Currie (2001) wrote that a number of isolated vertebrae, tarsometatarsi and unguals have been found in Upper Cretaceous strata of North America that closely resemble those of Mongolian avimimids (RTMP coll.). Eberth and Currie (2010) listed three avimimid elements from the Albertosaurus bonebed of the Horseshoe Canyon Formation.
References- 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.
Ryan, Currie and Russell, 2001. New material of Avimimus portentosus (Theropoda) from the Iren Dabasu Formation (Upper Cretaceous) of the Erenhot Region of Inner Mongolia. Journal of Vertebrate Paleontology. 21(3), 95A.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). 279-297. in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
Ryan and Currie, 2002. Asian small theropods in North America: evidence from Avimimidae. in Alberta Palaeontological Society, sixth annual symposium, "Fossils 2002", presented by Alberta Paleontological Society, in conjunction with Canadian Society of Petroleum Geologists, Paleontological Division and Department of Earth Sciences, Mount Royal College. p. 44.
Currie, 2005. Theropod dinosaurs of Dinosaur Provincial Park. in Braman, Therrien, Koppelhus and Taylor (eds). Dinosaur Park Symposium, short papers, abstracts and program, special publication of the Royal Tyrrell Museum. p. 15-18.
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.

Caenagnathoidea Sternberg, 1940 sensu Sereno, 1999
Definition- (Caenagnathus collinsi + Oviraptor philoceratops) (Maryanska et al., 2002; modified from Sereno, 1999)
Other definitions- (Chirostenotes pergracilis + Oviraptor philoceratops) (Sereno, in press)
= Oviraptorosauria sensu Padian et al. 1999
Definition- (Oviraptor philoceratops + Chirostenotes pergracilis) (modified)
= Oviraptoroidea Barsbold, 1976 sensu Sereno, 1999
Definition- (Oviraptor philoceratops + Caenagnathus collinsi) (Maryanska et al., 2002; modified from Sereno, 1999)
Comments- Sereno's in press definition is the same as Maryanska et al.'s (2002), except it replaces Caenagnathus with Chirostenotes. This is a poor decision, as the taxa are not definitely synonymous. Chirostenotes pergracilis and Elmisaurus elegans co-occur in the same formation, and the only reason Caenagnathus is synonymized with pergracilis instead of elegans is size. Until taxonomic problems are solved for caenagnathids, it's best to associate the family with it's eponymous species, even ignoring Phylocode rules.

Gigantoraptor Xu, Tan, Wang, Zhao and Tan, 2007
G. erlianensis Xu, Tan, Wang, Zhao and Tan, 2007
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China

Holotype- (LH V0011) (8 m; ~2.1 tons; 11 year old adult) mandibles (~438 mm), posterior cervical neural arch, eight partial dorsal vertebrae, several dorsal ribs, gastralia, sacrum, twenty-seven caudal vertebrae, fourteen chevrons, incomplete scapula, furcula, sternum, humerus (735 mm), radius, ulna, radiale, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, proximal phalanx II-2, manual ungual II, distal metacarpal III, phalanx III-1, phalanx III-2, manual ungual III, partial ilium, pubes, femur (1.1 m), tibia (1.18 m), fibula, astragalus, calcaneum, metatarsus (583 mm), pedal phalanges
Diagnosis- (after Xu et al., 2007) mandible less than 45% of femoral length; fossa on the lateral surface of the dentary close to the anterior end; fossa bounded dorsally by a lateral flange anterodorsal to the external mandibular fenestra; long posteroventral process of the dentary extending to the level of the glenoid; small, posteriorly tapered retroarticular process much deeper than wide; opisthocoelous proximal caudal vertebrae; procoelous distal caudal vertebrae; pleurocoels present on most caudal vertebrae; pair of vertically arranged pneumatic openings present on the lateral surface of proximal caudal centra; large pneumatic opening present on the ventral surface of proximal and middle caudal centra; proximal caudal vertebrae with tall neural spines (about three times as tall as wide); proximal caudal vertebrae with robust and rod-like transverse processes located posteriorly; posteroventral margin of proximal caudal centra extending considerably ventrally; well-developed laminal system on the proximal caudal vertebrae (prespinal, postspinal, spinopostzygapophyseal, anterior centrodiapophyseal, posterior centrodiapophyseal, and prezygodiapophyseal laminae present); middle caudal vertebrae with vertical prezygapophyseal articular facets located proximal to the distal extremity of the process; prominent convexity ventral to the acromion process on the lateral surface of the scapula; laterally bowed humerus; humerus with prominent, spherical head; humerus with strongly medially curved deltopectoral crest; centrally constricted thick ridge running along the posterior margin of the proximal half of the humerus; ulna with a subcircular, concave proximal articular surface; radius with a subspherical distal end; metacarpal I with a slightly convex medial margin of the proximal end; medial condyle of metacarpal I three times as high as wide; medial condyle of metacarpal I extending much more distally than the lateral condyle; metacarpal II with prominent dorsolateral process on the proximal end; metacarpal II with longitudinal groove on the ventral margin of the proximal third of the shaft; manual unguals with a triangular set of lateral grooves; laterally compressed pubis; femur with straight shaft; constricted femoral neck; posteromedially oriented, spherical femoral head; anteroposteriorly wide trochanteric crest which is very robust and higher anteriorly than posteriorly; distinct narrow groove medial to the trochanteric crest extending down the posterior margin of the femoral shaft; patellar groove present on the anterior surface of the distal femur; small calcaneum obscured from anterior view by the wide astragalar main body; proximal projection on the lateral margin of distal tarsal IV; metatarsal III with ginglymoid distal end; pedal unguals with two lateral grooves; constricted proximal articular surface of pedal unguals.
Comments- This was found to be a basal oviraptorid by Xu et al. (2007) using the TWG matrix, as a basal caenagnathid in the latest and most extensive oviraptorosaur analysis (Lu et al., 2015), and as a caenagnathid in the huge analysis of Lee et al. (2014).
References- Xu, Tan, Wang, Zhao and Tan, 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature. 844-847.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

undescribed caenagnathoid (Jensen, 1970)
Cenomanian, Late Cretaceous
Mussentuchit Member of Cedar Mountain Formation, Utah, US
Material
- (BYU-E 200; holotype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU-E 201) eggshell fragment (Jensen, 1970)
(BYU VP13699; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13700; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13701; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13702; paratype of Oolithes carlylensis) eggshell fragment (Jensen, 1970)
(BYU VP13703) eggshell fragment (Jensen, 1970)
(BYU VP13704) eggshell fragment (Jensen, 1970)
(BYU VP13705) eggshell fragment (Jensen, 1970)
(BYU VP13706) eggshell fragment (Jensen, 1970)
(BYU coll.) >225 eggshell fragments (Jensen, 1970)
(RTMP 98.107.4) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(RTMP 98.107.7) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(RTMP 98.107.10) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(RTMP 98.107.15) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(RTMP 98.107.18) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(RTMP 98.107.22) eggshell fragment (Zelenitsky, Carpenter and Currie, 2000)
(RTMP 98.107 coll.) >1500 eggshell fragments (Zelenitsky, Carpenter and Currie, 2000)
(UCM 82008) eggshell fragment (Jensen, 1970)
partial skeleton including mid caudal vertebra and pygostyle (Makovicky, Shinya and Zanno, 2014)
Comments- The adult skeleton is said to be very large, "second only to ... Gigantoraptor in size." The mid caudal is "highly pneumatic" and the last four caudals "form a pygostyle-like structure." As the type material of Macroelongatoolithus carlylensis (Jensen, 1970) was found close by, it is hypothesized to belong to the species. This was originally described as Oolithes carlylensis by Jensen, then referred to Macroolithus by Zhao (1975), and named as the new genus Boletuoolithus by Bray (1998), before being referred to Macroelongatoolithus by Zelenitsky et al. (2000). M. carlylensis was emended to M. carlylei by Zelenitsky et al. (2000), but while the ICZN states species should originally be named this way (Article 31.1.3), it is not a spelling that must be corrected (Article 32.5).
References- Jensen, 1970. Fossil eggs in the Lower Cretaceous of Utah. Brigham Young University Research Studies. Geology Series. Geology Studies. 17, 51-65.
Zhao, 1975. The microstructure of the dinosaurian eggshells of Nanxiong Basin, Guangdong Province. Vertebrata PalAsiatica. 13, 105-117.
Bray, 1998. Dinosaur eggshell Boletuoolithus carlylensis, oogenus nov. from the Lower Cretaceous Cedar Mountain Formation of Utah. New Mexico Museum of Natural History and Science Bulletin. 14, 221-224.
Zelenitsky, Carpenter and Currie, 2000. First record of elongatoolithid theropod eggshell from North America: The Asian oogenus Macroelongatoolithus from the Lower Cretaceous of Utah. Journal of Vertebrate Paleontology. 20(1), 130-138.
Makovicky, Shinya and Zanno, 2014. New additions to the diversity of the Mussentuchit Member, Cedar Mountain Formation dinosaur fauna. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 175.

undescribed caenagnathoid (Li, Yin and Liu, 1995)
Maastrichtian, Late Cretaceous
Zoumagang Formation, Henan, China
Material
- (Henan Geological Museum coll.; formerly Children's Museum of Indianapolis coll.; Baby Louie) (embryo) incomplete skull, partial mandible, vertebral fragments, rib fragments, ilium, femur, partial tibia, partial fibula, limb elements, six-eight eggs (430 mm), nest (Currie, 1996)
(HX 9301) egg (Li, Yin and Liu, 1995)
(HX 9302) egg (Li, Yin and Liu, 1995)
(HX 9303) egg (Li, Yin and Liu, 1995)
(HXW 9301) egg (Li, Yin and Liu, 1995)
(HWX 9302) egg (Li, Yin and Liu, 1995)
(HWX 9303) egg (Li, Yin and Liu, 1995)
....(LACM 7477/149736b) eggshells
(LACM F.A.3818,2001-1) egg (Grellet-Tinner, 2005)
(Nanyang Museum coll.) thirteen eggs, nest (Currie, 1996)
eggs (Fang et al., 1998)
Comments- The embryo nicknamed Baby Louie was discovered in 1993 and donated to the Children's Museum of Indianapolis, though it was returned to China in 2013. Currie (1996) initially reported the specimen as a therizinosaur, since being reidentified as an oviraptorid (Norell and Clark, pers. comm. to Grellet-Tinner, 2005). Identical eggs from the same formation were described as the ootaxon Macroelongatoolithus xixiaensis (Li et al., 1995), senior synonym of Longiteresoolithus xixiaensis (Wang and Zhou, 1995) from the same formation. The former species has since been synonymized with M. carlylensis by Zelenitsky et al. (2000) and Simon (2014). Grellet-Tinner (2005) described the eggs in detail (published as Grellet-Tinner et al., 2006), though Baby Louie itself has yet to be described.
References- Li, Yin and Liu, 1995. The discovery of a new genus of dinosaur egg from Xixia, Henan, China. Journal of Wuhan Institute of Chemical Technology. 17, 38-41.
Wang and Zhou, 1995. The discovery of new typical dinosaur egg fossils in Xixia basin, Henan Province. Henan Geology. 13, 262-267.
Currie, 1996. The great dinosaur egg hunt. National Geographic. 189(May 1996), 96-111.
Fang, Lu, Cheng, Zou, Pang, Wang, Chen, Yin, Wang, Liu, Xie and Jin, 1998. On the Cretaceous fossil eggs of Xixia County, Henan Province. Geological Publishing House Beijing. 125 pp.
Zelenitsky, Carpenter and Currie, 2000. First record of elongatoolithid theropod eggshell from North America: The Asian oogenus Macroelongatoolithus from the Lower Cretaceous of Utah. Journal of Vertebrate Paleontology. 20(1), 130-138.
Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case study of saurischian dinosaur relationships and avian evolution. PhD thesis, University of Southern California. 221 pp.
Grellet-Tinner, Chiappe, Norell and Bottjer, 2006. Dinosaur eggs and nesting behaviors: A paleobiological investigation. Palaegeography, Palaeoclimatology, Palaeoecology. 232, 294-321.
Simon, 2014. Giant dinosaur (theropod) eggs of the oogenus Macroelongatoolithus (Elongatoolithidae) from southeastern Idaho: Taxonomic, paleobiogeographic and reproductive implications. Masters thesis, Montana State University. 110 pp.
Zelenitsky, Currie, Carpenter and Lu, 2015. Baby Louie: A theropod perinate from the Cretaceous of China reveals affinity of the largest known dinosaur eggs. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 244-245.

undescribed Caenagnathoidea (Buckley, 2002)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US

Material- (NS.1563.018) two partial manual unguals, partial tibia, proximal fibula, partial astragalus, (metatarsal II ~390 mm) distal metatarsal III, partial metatarsal shaft, incomplete pedal phalanx III-1, partial pedal phalanx (Buckley, 2002)
(NS.32001.077) metatarsal III, metatarsal IV (Buckley, 2002)
manual ungual (Holtz, Williams, Tremaine and Matthews, 2014)
Comments- The NS specimens were identified as Elmisaurus in the abstract, but as caenagnathid or oviraptorid in the poster.
References- Buckley, 2002. New material of Elmisaurus (Theropoda, Elmisauridae) from the Late Cretaceous Hell Creek Formation of Southeastern Montana. Journal of Vertebrate Paleontology. 22(3), 39A.
Holtz, Williams, Tremaine and Matthews, 2014. New additions to the Hell Creek Formation (Upper Maastrichtian) vertebrate fauna of Carter County, Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 149.

undescribed caenagnathoid (Iijima, Sato, Watabe, Tsogtbaatar and Ariunchimeg, 2011)
Cenomanian-Santonian, Late Cretaceous
Baynshiren Formation, Mongolia
Material
- (two hatchling individuals) articular-surangular-coronoid, two eggs, eggshell fragments
Comments- The eggs are elongatoolithid.
Reference- Iijima, Sato, Watabe, Tsogtbaatar and Ariunchimeg, 2011. Bone bed of baby oviraptorosaur and hadrosauroid dinosaurs from the Bayanshiree Formation (Late Cretaceous) in Southeastern Mongolia. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 130.

Caenagnathidae Sternberg, 1940
Definition- (Caenagnathus collinsi <- Oviraptor philoceratops) (Maryanska et al., 2002; modified from Sereno, 1998)
Other definitions- (Chirostenotes pergracilis + Chirostenotes elegans + Elmisaurus rarus + Caenagnathasia martinsoni + BHM 2033) (Sues, 1997)
(Chirostenotes pergracilis <- Oviraptor philoceratops) (Sereno, in press; modified from Padian et al., 1999)
= Elmisauridae Osmolska, 1981
= Kuszholiidae Nessov, 1992
= Caenagnathidae sensu Sues, 1997
Definition- (Chirostenotes pergracilis + Chirostenotes elegans + Elmisaurus rarus + Caenagnathasia martinsoni + BHM 2033)
= Caenagnathidae sensu Padian et al., 1999
Definition- (Chirostenotes pergracilis <- Oviraptor philoceratops) (modified)
Comments- Possible caenagnathid material from the Yalovach Formation of Tadjikistan (Ryan, 1997) is probably based on reports of oviraptorids by Nessov (1995), which probably belongs to therizinosaurs instead (Alifanov and Averianov, 2006). Reported caenagnathid forelimb elements from the Densus-Ciula Formation of Romania (Csiki and Grigorescu, 2005) has since been referred to the paravian Balaur (Csiki et al., 2010).
References- 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].
Ryan, 1997. Middle Asian Dinosaurs. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. Academic Press. p. 442-444.
Csiki and Grigorescu, 2005. A new theropod from Tustea: Are there oviraptorosaurs in the Upper Cretaceous of Europe? Kaupia. 14, 78.
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.
Csiki, Vremir, Brusatte and Norell, 2010. An aberrant island-dwelling theropod dinosaur from the Late Cretaceous of Romania. Proceedings of the National Academy of Sciences. Early Edition, 5 pp.

unnamed Caenagnathidae (Gilmore, 1924)
Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material
- (CMN 346) manual ungual I (~130 mm on curve) (Bell, Currie and Russell, 2015)
(CMN 8504) dorsal centrum (44 mm), three caudal centra (22-25 mm) (Gilmore, 1924)
(TMM 41395-1) manual ungual II (Bell, Currie and Russell, 2015)
Comments- The centra were described by Gilmore (1924) as distinct from other coelurosaurs known at the time, though possibly referrable to Chirostenotes or Dromaeosauridae (neither of which were then known from vertebrae). Currie et al. (1994) noted the caudals belonged to an oviraptorosaur, referring them to Caenagnathus sp.. Bell et al. (2015) described two manual unguals as cf. Anzu, noting they are comparable in size and age but that CMN 346 has a deeper concavity between the flexor tubercle and articular surface than that genus.
References- Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta. Canada Geological Survey, Bulletin n. 38, geological series 43, 1-13.
Currie, Godfrey and Nessov, 1994. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Bell, Currie and Russell, 2015. Large caenagnathids (Dinosauria, Oviraptorosauria) from the uppermost Cretaceous of Western Canada. Cretaceous Research. 52, 101-107.

unnamed caenagnathoid (Nessov and Khisarova, 1988)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material
- dentary
Comments- This was first described as a turtle (Nessov and Kisarova, 1988), but later identified as a caenagnathid and relative of Caenagnathasia by Currie et al. (1994).
References- Nessov and Khisarova, 1988. New data on vertebrates from the Late Cretaceous of Shakh-Shakh and Baybolat (northeastern Aral region). In Material on the history of the fauna and flora of Kazakhstan, Vol. 10. Academy of Sciences of Kazakhstan, Alma Ata. pp. 5-14. [In Russian]
Currie, Godfrey and Nessov, 1994. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.

undescribed Caenagnathidae (Fiorillo, 1989)
Late Campanian, Late Cretaceous
Judith River Formation, Montana
Material
- two specimens
Comments- Referred to Chirostenotes, but may be Elmisaurus instead.
Reference- Fiorillo, 1989. The vertebrate fauna from the Judith River Formation (Late Cretaceous) of Wheatland and Golden Valley Counties, Montana. The Mosasaur. 4, 127-142.

undescribed caenagnathid (Tokyark, 1990)
Late Cretaceous
Saskatchewan, Canada

Reference- Tokaryak, 1990. It was here a minute ago. The Saskatchewan Archaeological Society Newsletter. 11(2), 44-45.

Caenagnathidae indet. (Currie, 1992)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada

Material- (RTMP 75.11.33) cervical vertebra (Sues, 1997)
(RTMP 81.19.252) parietals (Currie, 1992)
(RTMP 84.167.44) sacrum (Makovicky, 1995)
(RTMP 89.36.109) caudal vertebra (Makovicky, 1995)
(RTMP 91.36.146) caudal vertebra (Makovicky, 1995)
(RTMP 92.36.53) proximal caudal vertebra (31.4 mm) (Currie et al., 1994)
Comments- RTMP 81.19.252 and 92.36.53 were referred to Caenagnathus sp. by Currie (1992) and Currie et al. (1994) respectively. Sues (1997) referred RTMP 75.11.33 to Chirostenotes, while Makovicky (1995) referred RTMP 84.167.44, 89.36.109 and 91.36.146 to that genus. The latter material did not have locality information listed, but is probably from the Dinosaur Park Formation of Alberta. All of this material may be Chirostenotes or Elmisaurus, as frontal and vertebral differences between the genera are not yet known.
Reference- Currie, 1992. Saurischian dinosaurs of the Late Cretaceous of Asia and North America. In N.J. Mateer and P.J. Chen (eds), Aspects of Nonmarine Cretaceous Geology. pp. 237-249. Beijing: China Ocean press.
Currie, Godfrey and Nessov, 1994. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Copenhagen University, Copenhagen, Denmark.

undescribed caenagnathid (Breithaupt, 1994)
Late Cretaceous
Wyoming

Reference- Breithaupt, 1994. Wyoming Dinosaur Diversity: Forty-Fourth Annual Field Conference-1994. Wyoming Geological Association Guidebook. 101-104.

undescribed caenagnathid (Manabe and Barrett, 2000)
Valanginian-Hauterivian, Early Cretaceous
Kuwajima Formation of the Tetori Group, Japan
Material
- (SBEI-167) manual ungual
Comments- Referred to the oviraptorosaur-therizinosaur clade by Manabe et al. (2000), it provisionally resembles caenagnathids most closely.
References- Barrett and Manabe, 2000. The dinosaur fauna from the Earliest Cretaceous Tetori Group of Central Honshu, Japan. Journal of Vertebrate Paleontology. 20(3), 28A-29A.
Manabe and Barrett, 2000. Dinosaurs. In Matsuoka (ed.). Fossils of the Kuwajima "Kaseki-kabe" (fossil-bluff). Scientific report on a Neocomian (Early Cretaceous) fossil assemblage from the Kuwajima Formation, Tetori Group, Shiramine, Ishikawa, Japan. Shiramine Village Board of Education, Japan. 93-98.
Manabe, Barrett and Isaji, 2000. A refugium for relicts? Nature. 404, 953-954.
Matsuoka, Kusuhashi, Takada and Setoguchi, 2002. A clue to the Neocomian vertebrate fauna: Initial results from the Kuwajima 'Kaseki-kabe' (Tetori Group) in Shiramine, Ishikawa, central Japan. Memoirs of the Faculty of Science, Kyoto University, Series of Geology and Mineralogy. 59(1), 33-45.

undescribed Caenagnathidae (Ryan, Currie and Russell, 2001)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material
- (AMNH 6754) distal metatarsal III
(AMNH 6755) incomplete metatarsus
Comments- AMNH 6754 and 6755 are listed on the museum's online collection database as Elmisaurus sp.. They were assigned by Ryan et al. (2001) to Avimimus, but although the metatarsus is arctometatarsalian, the third metatarsal extends up 90% of the metatarsal length in anterior view (and almost as much posteriorly). This contrasts with 45% in A. portentosus. This metatarsus is less slender than the latter, with a more reduced fourth metatarsal and no fifth metatarsal fused to it. This suggests the AMNH website's identification of these specimens as caenagnathids may be correct, while the fused metatarsals and proximal tarsals and deeply concave posterior surface may suggest a relationship with Elmisaurus. Caenagnathasia has since been identified from the Iren Debasu Formation, so this material may be referrable to that taxon.
References- Ryan, Currie and Russell, 2001. New material of Avimimus portentosus (Theropoda) from the Iren Dabasu Formation (Upper Cretaceous) of the Erenhot Region of Inner Mongolia. Journal of Vertebrate Paleontology. 21(3), 95A.

unnamed caenagnathid (Jasinski, Sullivan and Lucas, 2011)
Early Maastrichtian, Late Cretaceous
Naashoibito Member of the Ojo Alamo Formation, New Mexico, US

Material- (SMP VP-2172) incomplete pedal ungual
Reference- 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.

Hagryphus Zanno and Sampson, 2005
H. giganteus Zanno and Sampson, 2005
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US

Holotype- (UMNH VP 12765) distal radius, radiale, ulnare, semilunate carpal, distal carpal III, metacarpal I (66 mm), phalanx I-1 (87 mm), manual ungual I (94 mm on curve), metacarpal II (122 mm), phalanx II-1 (95 mm), phalanx II-2 (99 mm), metacarpal III (87 mm), phalanx III-1 (54 mm), phalanx III-2 (45 mm), phalanx III-3 (60 mm), manual ungual III (75 mm on curve), manual claw III impression, proximal pedal ungual I, distal metatarsal II, distal phalanx II-1, phalanx II-2, pedal ungual II, incomplete pedal ungual III, distal metatarsal IV, incomplete pedal ungual IV, fragmentary pedal phalanges
Referred- ?(RAM 12433) distal metatarsal II (Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013)
Diagnosis- (modified from Zanno and Sampson, 2005) (compared to Chirostenotes and Elmisaurus) manual digits I and II more robust; metacarpal I and phalanx I-1 proportionally shorter.
References- Zanno and Sampson, 2003. A new caenagnathid specimen from the Kaiprowits Formation (Late Campanian) of Utah. Journal of Vertebrate Paleontology. 23(3), 114A.
Zanno and Sampson, 2005. A new oviraptorosaur (Theropoda: Maniraptora) from the Late Cretaceous (Campanian) of Utah. Journal of Vertebrate Paleontology. 25(4), 897-904.
Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013. Late Cretaceous theropod dinosaurs of Southern Utah. In Titus and Loewen (eds.). At the Top of the Grand Staircase: The Late Cretaceous of Southern Utah. Indiana University Press. 504-525.

Ojoraptorsaurus Sullivan, Jasinski and van Tomme, 2011
O. boerei Sullivan, Jasinski and van Tomme, 2011
Early Maastrichtian, Late Cretaceous
Naashoibito Member of the Ojo Alamo Formation, New Mexico, US
Holotype
- (SMP VP-1458) (~1.8-2.1 m) incomplete pubes (~344 mm)
Diagnosis- (after Sullivan et al., 2011) enclosed pubic fossa recessed at least one cm from acetabular rim, positioned on medial surface of pubic shaft (also in Citipati); iliac peduncle articular surface of pubes sub-trapezoidal in shape (also in oviraptorids).
Other diagnoses- Sullivan also listed the "spoon-shaped" (oval) depression on the anterior dorsal surface of the pubic boot, but this is also present in Chirostenotes specimen ROM 43250. A pubic shaft which is convex anteriorly just proximal to the pubic boot is present in Avimimus and CM 78001.
Comments- The validity of this taxon is uncertain, as it is only compared to Nomingia, Chirostenotes specimen ROM 43250 and undescribed caenagnathid CM 78001, and the distribution of its diagnostic features in these taxa and other oviraptorosaurs is poorly understood. The oval depression on the dorsal pubic boot surface is absent in Nomingia, but unreported for Microvenator, oviraptorids or CM 78001. Since ROM 43250 has traces of it despite crushing, it may support a caenagnathid subclade instead. The proximomedial pubic fossa is more distally placed than Chirostenotes and CM 78001, but Sullivan et al. do not describe its position in Nomingia or oviraptorids, though it is proximal in Microvenator. Notably, Citipati sp. IGM 100/42 also has a distally placed fossa. While most oviraptorosaurs have completely anteriorly concave pubes, CM 78001 also has a slight convexity distally (contra Sullivan et al.), ROM 43250 is crushed, and a strong convexity like Ojoraptorsaurus' is present in Avimimus. Pubic convexity also varies within other coelurosaur species, such as Sinosauropteryx, Archaeornithomimus and Sinovenator. Finally, the ilial articular surface of the pubis is transversely narrower anteriorly than posteriorly, unlike ROM 43250. Yet this is also true in IGM 100/42 and Nemegtomaia, and is unreported in other caenagnathoids. Thus while not known in other caenagnathids yet, it may prove to be plesiomorphic.
Reference- Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid Ojoraptorsaurus boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria), from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 53, 418-428.

unnamed Caenagnathidae (Longrich, Barnes, Clark and Millar, 2013)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US

Material- (TMM 42920-2) incomplete manual ungual II (~80 mm)
(UTEP B38 L-3) femur (350 mm)
Comments- Longrich et al. (2013) refer these specimens to Chirostenotes sp. based on size, though the femur cannot be compared to Elmisaurus or Leptorhynchos, and the ungual has no described characters more similar to Chirostenotes than to Elmisaurus, Hagryphus or other caenagnathids. Indeed, the authors note while Longrich feels the ungual is Chirostenotes, Barnes disagrees and believes it may belong to the Leptorhynchos gaddisi type individual as it was found in proximity.
Reference- Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.

Caenagnathinae Sternberg, 1940 sensu Paul, 1988
Definition- (Caenagnathus collinsi <- Oviraptor philoceratops, Avimimus portentosus) (Martyniuk, 2012)
Other definitions- (Caenagnathus collinsi <- Elmisaurus elegans, Caenagnathasia martinsoni) (Longrich, Barnes, Clark and Millar, 2013)
(Caenagnathus collinsi <- Elmisaurus rarus) (Hendrickx, Hartman and Mateus, 2015)
Comments- Longrich et al. (2013) thought elegans was a caenagnathine, so no doubt meant to use Elmisaurus rarus as the external specifier instead.
References- Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Chirostenotes Gilmore, 1924
?= Macrophalangia Sternberg, 1932
?= Caenagnathus Sternberg, 1940
= "Steneodactylus" Gilmore vide Holtz, DML 1998
?= Epichirostenotes Sullivan, Jasinski and van Tomme, 2011
Diagnosis- (after Currie et al., 1993; compared to Caenagnathus sternbergi holotype) lower articular ridge on mandible; medial glenoid longer anteroposteriorly; mandibular ramus anterior to glenoid more robust.
(after Varricchio, 2001) chorda tympani foramen/slot present.
C. pergracilis Gilmore, 1924
?= Macrophalangia canadensis Sternberg, 1932
?= Caenagnathus collinsi Sternberg, 1940
= "Steneodactylus pergracilis" Gilmore vide Holtz, DML 1998
?= Epichirostenotes curriei Sullivan, Jasinski and van Tomme, 2011
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype
- (CMN 2367) partial metacarpal I, incomplete phalanges I-1 (63 mm), incomplete manual unguals I (44 mm), incomplete metacarpal II, phalanges II-1 (one incomplete; 65 mm), phalanges II-2 (one incomplete; 72 mm), manual ungual II (62 mm), phalanges III-3 (44 mm), manual unguals III (36 mm)
Referred- ?(CMN 8538; holotype of Macrophalangia canadensis) distal tibia, partial astragalus, distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1 (58 mm), pedal ungual I (31 mm), metatarsal II (205 mm; lost), phalanx II-1 (78 mm), phalanx II-2 (63 mm), pedal ungual II (60 mm), partial metatarsal III (230 mm; lost), phalanx III-1 (75 mm), phalanx III-2 (52 mm), phalanx III-3 (58 mm), pedal ungual III (60 mm), metatarsal IV (212 mm; lost), phalanx IV-1 (59 mm), phalanx IV-2 (33 mm), phalanx IV-3 (31 mm), phalanx IV-4 (35 mm), pedal ungual IV, metatarsal V (60 mm) (Sternberg, 1932)
?(CMN 8776; holotype of Caenagnathus collinsi) mandibles (205 mm) (Sternberg, 1940)
(CMN 12372; = CMN 12322 of Longrich et al.) manual ungual II (Longrich et al., 2013)
?(RTMP 79.14.1) manual ungual I (~90 mm on curve) (Bell, Currie and Russell, 2015)
(RTMP 79.14.499) manual ungual II (83 mm) (Currie and Russell, 1988)
(RTMP 79.20.1) (~67 kg) (sacrum- 200 mm) first sacral vertebra (36.9 mm), second sacral vertebra, third sacral vertebra, fourth sacral vertebra, fifth sacral vertebra, sixth sacral verebra (29.9 mm), proximal mid dorsal rib, three dorsal rib shafts, coracoid (58 mm tall), distal metacarpal I, phalanx I-1 (69 mm), manual ungual I (47 mm), phalanx II-1 (77 mm), phalanx II-2 (82 mm), manual ungual II (69 mm), phalanx III-1 (33 mm), phalanx III-3 (42 mm), ilium (255 mm), ischium (144 mm), femur (310 mm), tibia (367 mm), metatarsal I (39 mm), phalanx I-1 (40 mm), metatarsal II (181 mm), metatarsal III (207 mm), phalanx III-1 (58 mm), metatarsal IV (186 mm), metatarsal V (33 mm) (Currie and Russell, 1988)
?(RTMP 82.19.222) manual ungual I (91 mm on curve) (Bell, Currie and Russell, 2015)
?(RTMP 86.36.323) (~96 kg) femur (370 mm) (Bell, Currie and Russell, 2015)
?(RTMP 93.36.197) metatarsal II (261 mm) (Bell, Currie and Russell, 2015)
?(RTMP 93.36.198) metatarsal II (245 mm) (Bell, Currie and Russell, 2015)
?(RTMP 93.36.631) partial astragalocalcaneum (~75 mm wide) (Bell, Currie and Russell, 2015)
?(RTMP 93.36.475) manual ungual II (Bell, Currie and Russell, 2015)
?(RTMP 93.75.49) incomplete astragalocalcaneum (76.3 mm wide) (Bell, Currie and Russell, 2015)
?(RTMP 2009.3.29; = RTMP 82.19.222 in part) manual ungual I (Bell, Currie and Russell, 2015)
?(UA 55725) partial caudal vertebra (39.7 mm) (Bell, Currie and Russell, 2015)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada

?(CMN 9570) metatarsal II (258 mm) (Russell, 1984)
?(ROM 43250; holotype of Epichirostenotes curriei) (adult) maxilla, palatine, braincase, third cervical vertebra, sixth cervical vertebra (85 mm), seventh cervical vertebra (81 mm), two cervical ribs, anterior dorsal vertebra (40 mm), anterior dorsal vertebra, dorsal rib, gastralia fragments, sacrum (270 mm), proximal caudal vertebra, four distal caudal vertebrae, ilial fragments, pubes (422 mm), ischium (212 mm), distal tibia (Sues, 1997)
?(WL-156) multiple elements (Eberth et al., 2013)
Diagnosis- (after Currie, 1989) proximal end of metatarsal III diamond-shaped; metatarsals II and IV much shorter than III (<93%).
(after Currie et al., 1994) more elongate shallow dentary (distortion?); more elongate mandibular symphysis (distortion?); dorsal midline ridge on anterior portion of mandibular symphysis; midline anterior occlusal groove absent; lateral occlusal grooves extend almost to the tip of the dentary; lingual ridges converge and join the midline symphysial ridge.
Comments- Chirostenotes pergracilis was named in 1924 based on incomplete manus, and thought by Gilmore to be a relative of Ornitholestes. He also referred a pair of toothed dentaries to the taxon, which Currie et al. (1990) made the holotype of Richardoestesia. Holtz (DML, 1998) noted the specimen is labeled "Steneodactylus pergracilis" in the CMN's collection, showing that was an earlier proposed name for the taxon. This has not been published in the literature however. Sternberg later (1932) described Macrophalangia canadensis based on a pes from the same formation, thought to be an ornithomimid. Colbert and Russell (1969) noted the two forms may be synonymous, though this was not shown to be likely until Osmolska (1981). An alternative hypothesis, the synonymy of Chirostenotes with Dromaeosaurus, was suggested by Ostrom (1969; 1990), but is clearly incorrect. Longrich et al. (2013) proposed Macrophalangia was synonymous with Caenagnathus collinsi based on size and robusticity compared to Chirostenotes pergracilis, which they viewed as a separate taxon. While this is possible, similar or greater differences in size and robusticity are known for other coelurosaur species. Indeed, Longrich et al. felt sufficiently uncertain that they did not refer to their large taxon as Macrophalangia, which has priority over Caenagnathus. Funston (pers. comm. 2015) noted the metatarsals of Macrophalangia are lost, having gone on loan and never returned.
Caenagnathus collinsi was named for a pair of mandibles in 1940, and assigned to Aves (Sternberg, 1940). This assignment was rejected by most paleornithologists such as Wetmore (1960), who hypothesized a relationship with ornithomimids. However, Cracraft (1971) supported an avian relationship, specfically with Galloanseres. Caenagnathus' identity was finally solved by Osmolska (1976), who allied it with oviraptorids. Currie and Russell (1988) suggested Caenagnathus and Chirostenotes were synonymous, which was strengthened by Sues (1997) and shown to be true at the family level by Anzu. Senter (2007) ran Caenagnathus (based on CMN 8776) separately from Chirostenotes (based on non-mandibular remains) in his phylogenetic analysis, finding the latter to be an oviraptorid while the former was more basal. He interpreted this as supporting the genera being distinct, yet Anzu disproves this notion. As noted above, Longrich et al. (2013) separate Caenagnathus collinsi from Chirostenotes pergracilis based on differences between the type mandible of the former and sternbergi, which they synonymize with pergracilis (and refer RTMP 90.56.6 and 2001.12.12 to, but not the other mandibles here placed in elegans). However, the symphyseal angle of sternbergi jaw RTMP 2001.12.12 is more similar to Leptorhynchos gaddisi than to RTMP 1990.56.6, which is intermediate between those and collinsi. This suggests sternbergi is indeed synonymous with elegans, that the glenoid morphology of sternbergi is shared by multiple species, or that at least least four caenagnathid species lived in the Dinosaur Park Formation.
Although the basic identification of Chirostenotes, Macrophalangia and Caenagnathus as members of the same clade has been verified, the species synonymization is less certain. Cracraft named a new species of Caenagnathus, C. sternbergi, known from a posterior mandible. This differs from C. collinsi in a few characters, suggesting two species are present in the Dinosaur Park Formation. Additionally, Currie et al. (1994) described five dentaries which also differ from C. collinsi. They referred to these as Caenagnathus cf. sternbergi, as none are directly comparable to the C. sternbergi holotype. Currie (2005) illustrated a new mandible which confirms Currie et al. were correct to refer at least some of these dentaries to C. sternbergi. Finally, Currie and Russell (1988) distinguished between two kinds of metatarsus (robust CMN 8538; gracile ROM 781 and RTMP 79.20.1), tentatively believing them to be sexual morphs. Currie (1989) expanded on this, referring some specimens to Elmisaurus elegans (ROM 781, 37163 and RTMP 82.39.4) and others to Chirostenotes pergracilis (CMN 8538 and 9570, and RTMP 79.20.1). He distinguished them based on several characters. Thus there are two morphotypes for each anatomical area, one smaller than the other. Currie and Russell (1988) first proposed the synonymy of Caenagnathus collinsi with Chirostenotes pergracilis, and Caenagnathus sternbergi with Elmisaurus (then Chirostenotes) elegans. Longrich et al. (2013) have modified this to synonymize Macrophalangia with Caenagnathus collinsi, Chirostenotes with Caenagnathus sternbergi, and unnamed jaws with Elmisaurus/Leptorhynchos elegans. Yet as noted above, Macrophalangia is not notably different from Chirostenotes (RTMP 79.20.1), and sternbergi is at least as comparable to elmisaurine dentaries. This leaves many plausible alternatives. Most recently, Funston et al. proposed Caenagnathus collinsi (including RTMP 79.14.1, 82.19.222, 86.36.323, 93.36.197, 93.36.198, 93.36.631, 93.36.475, 93.75.49, 2009.3.29 and UA 55725) is distinct from Chirostenotes pergracilis (including Macrophalangia and the other Dinosaur Park specimens listed above, as well as presumedly the mandibular material listed by Longrich et al.). This was based on size, absence of a deep concavity between the flexor tubercle and articular surface of manual ungual I (unlike only the one codable ungual I of the pergracilis holotype and Frenchman Formation CMN 346), anteriorly projecting proximal surface for metatarsal IV on mt II (less developed than CMN 9570 and comparable to RTMP 79.20.1), posterior process on proximal surface (comparable in posterior extent to CMN 9570 and RTMP 79.20.1, but more abrupt distally than both) and less proximally extensive surface for metatarsal III on metatarsal II (~53% in RTMP 93.36.198 compared to ~60% in RTMP 79.20.1 and ~54% in CMN 9570). Whether these differences reflect taxonomy will be strengthened by the examination of more specimens, to determine if e.g. the manual ungual concavity and low distal angle of the posterior metatarsal II process are universal in smaller individuals.
RTMP 79.20.1 was discovered in 1979 and described by Currie and Russell in 1988 as Chirostenotes pergracilis, in which they also included the Elmisaurus elegans holotype. Ironically, Currie and Russell mentioned RTMP 82.39.4 as an American Elmisaurus specimen though he did not yet refer the elegans holotype to that genus. Currie and Russell referred RTMP 79.20.1 to their gracile morph, which included elegans, based on the slender third manual digit and gracile pes. They note the minimum width of manual phalanx III-3 is 10% of its length, compared to 11% in the Chirostenotes holotype and 10% in the Elmisaurus holotype. Sues (1997) followed this referral, but believed the gracile morph to be a separate species, which he called Chirostenotes elegans. Currie (1989) referred RTMP 79.20.1 to Chirostenotes pergracilis, not his new combination Elmisaurus elegans. Currie and Russell (1988) describe several characters consistant with this identification- unfused tarsometatarsus; metatarsals II and IV much shorter than III (87 and 90% respectively); proximal end of metatarsal III diamond-shaped; proximolateral process on metatarsal IV absent. These characters are more numerous and observable in more specimens than the 1% difference in phalangeal width, while the gracile pes may be due to the small size of RTMP 79.20.1 compared to CMN 8538. Sullivan et al. (2011) recently questioned the referral of RTMP 79.20.1 to Chirostenotes pergracilis, based on "some minor morphological differences, including a broader curvature of unguals I and II, a deeper ungual (measured from the dorsal and plantar extremity of the articular facet as per Senter, 2007), and a shorter phalanx III-3." Yet such differences are common within theropod species known from multiple specimens. Thus RTMP 79.20.1 is assigned to Chirostenotes pergracilis here, as it was by Currie (2005) and Longrich et al. (2013) as well.
ROM 43250 was discovered in 1923 and identified as an ornithomimid. Russell (1972) listed it as an undetermined ornithomimid, though Sues (1994) correctly identified it before describing it in detail (Sues, 1997). It is notable in being one of only two caenagnathid specimens from the Horseshoe Canyon Formation. Both are referred to Chirostenotes pergracilis based on large size, while the second specimen (CMN 9570) is also distinctive from Elmisaurus due to its lack of fusion and straight distal end. ROM 43250 differs from Anzu in the presence of an antorbital fossa, larger foramen magnum and more ventrally angled paroccipital processes. Sullivan et al. (2011) proposed the name Epichirostenotes curriei for ROM 43250, based on stratigraphy and several supposed ischial differences from RTMP 79.20.1. Of these, ischial length cannot be compared since Epichirostenotes doesn't preserve an ilium and Chirostenotes doesn't preserve a pubis. The deeper post-obturator body, deeper obturator process, square-tipped obturator process and longer and more ventrally projecting pubic peduncle are all minor variations comparable to those seen in Tyrannosaurus rex specimens (AMNH 5027, CM 9380 and FMNH PR2081). Other listed diagnostic characters are unknown in other Chirostenotes specimens (braincase much deeper than long, with distinctly verticalized basicranial region; otic region with deep, but anteroposteriorly narrow, lateral depression) or also present in RTMP 79.20.1 (synsacrum composed of six co-ossified vertebrae and pneumatic foramina). While ROM 43250 may indeed be a distinct species from Chirostenotes pergracilis, the known remains don't justify that anatomically, and whether it is more closely related to Chirostenotes than Elmisaurus or Hagryphus is basically unknowable. It is provisionally retained in Chirostenotes pergracilis here, since I do not support purely stratigraphy-based taxonomy. Eberth et al. (2013) also lists WL-156 as a possible Epichirostenotes specimen, though its whereabouts given the subsequent death of Wann Langston ('WL' stands for his personal collection) are uncertain.
References- Gilmore, 1924. A new coelurid dinosaur from the Belly River Cretaceous Alberta. Canada Geological Survey, Bulletin 38, geological series 43, 1-13.
Sternberg, 1932. Two new theropod dinosaurs from the Belly River Formation of Alberta. Canadian Field-Naturalist. 46(5), 99-105.
Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of Paleontology. 14(1), 81-85.
Wetmore, 1960. A classification for the birds of the world. Smithsonian Misc. Coll. 139(11), 1-37.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus. American Museum Novitiates. 2380, 1-49.
Ostrom, 1969. Osteology of Deinonychus antirrhopus, an unusual theropod from the Lower Cretaceous of Montana. Peabody Museum Bulletin. 30, i-viii + 1-165.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada. Canadian Journal of Earth Sciences. 9, 375-402.
Osmolska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25, 972-986.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26, 1319-1324.
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.
Ostrom, 1990. Dromaeosauridae. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria. Berkeley, University of California Press. 269-279.
Currie, Godfrey and Nessov, 1994. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Sues, 1994. New evidence concerning the phylogenetic position of Chirostenotes (Dinosauria: Theropoda): Journal of Vertebrate Paleontology. 14(3), 48A.
Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria: Theropoda) from Western North America. Journal of Vertebrate Paleontology. 17(4), 698-716.
http://dml.cmnh.org/1998Aug/msg00743.html
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.
Senter and Parrish, 2005. Functional analysis of the hands of the theropod dinosaur Chirostenotes pergracilis: evidence for an unusual paleoecologial role. PaleoBios. 25(2), 9-19.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid Ojoraptorsaurus boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria), from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin. 53, 418-428.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Lamanna, Sues, Schachner and Lyson, 2014. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS ONE. 9(3), e92022.
Bell, Currie and Russell, 2015. Large caenagnathids (Dinosauria, Oviraptorosauria) from the uppermost Cretaceous of Western Canada. Cretaceous Research. 52, 101-107.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
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.

Apatoraptor Funston and Currie, 2016
A. pennatus Funston and Currie, 2016
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada

Holotype- (RTMP 93.51.1) (~80 kg) palatine (~25 mm), incomplete mandible (~165 mm), hyoid (68.4 mm), (cervical series 680 mm), axial fragment, third to twelve cervical vertebrae fused to ribs (seventh 53.2, ninth 61.5 mm), nine dorsal vertebrae, nine dorsal ribs, five uncinate processes, scapulae (one proximal; 215 mm), coracoids, sternal plates (one fregmentary; 112 mm), sternal ribs, humerus (206 mm), radius, ulna (167 mm), (?)semilunate carpal, metacarpal I (52 mm), phalanx I-1 (82.4 mm), manual ungual I (50 mm on curve), metacarpal II (105 mm), phalanx II-1 (74 mm), phalanx II-2 (68.5 mm), manual ungual II (70 mm on curve), metacarpal III (75 mm), partial phalanx III-1, phalanx III-2 (32 mm), incomplete phalanx III-3, ilial fragment, partial femur (~345 mm), proximal tibia, proximal fibula
Diagnosis- (after Funston and Currie, 2016) ventral flange of angular underlying posteroventral dentary process; anterior constriction of external mandibular fenestra by posteroventral dentary process; medial fossa anterior to articular region of mandible; articular region of mandible with low articular ridge offset from dorsal margin of articular-surangular-coronoid complex; metacarpal I less than half as long as metacarpal II; manual phalanx I-1 longest in hand; manual phalanx II-1 longer than II-2.
Comments- Funston (2014) comments on RTMP 93.51.1, a specimen discovered in 1993 and first believed to be ornithomimid. It is listed as Funston et al. (2015) as "unnamed taxon", is said to be from the Horseshoe Canyon Formation by Eberth et al. (2013) and tentatively assigned to Epichirostenotes there. Currie et al. (2016) incorrectly state it's from the Dinosaur Park Formation however, when commenting on its scapulocoracoid and humerus. The specimen was officially named and described by Funston and Currie (2016), recovered as closer to Elmisaurus rarus than to elegans, Caenagnathasia or any of the larger caenagnathids.
References- Eberth, Evans, Brinkman, Therrien, Tanke and Russell, 2013. Dinosaur biostratigraphy of the Edmonton Group (Upper Cretaceous), Alberta, Canada: Evidence for climate influence. Canadian Journal of Earth Sciences. 50(7), 701-726.
Funston, 2014. Understanding Alberta’s caenagnathids: Insights on anatomy and taxonomy from a new, articulated specimen. R.E. Peter Biology Conference. Abstract 10.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osmólska, 2016. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
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.

Anzu Lamanna, Sues, Schachner and Lyson, 2014
A. wyliei Lamanna, Sues, Schachner and Lyson, 2014
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US

Holotype- (CM 78000) (~3.5 m; ~284 kg) premaxillary fragments, quadrates, pterygoids, incomplete braincase, cranial fragments, incomplete mandibles (320 mm), six cervical vertebrae, dorsal ribs, gastralia, nine caudal vertebrae, twenty-eighth caudal vertebra (33.9 mm), twenty-ninth caudal vertebra (37.2 mm), partial thirtieth caudal vertebra, six chevrons, scapulocoracoids (scap 415 mm, cor 121 mm), humerus (345 mm), radius (280 mm), ulna (280 mm), phalanx I-1 (123.5 mm), manual ungual I (89.1 mm, 130 mm on curve), metacarpal II (139 mm), phalanx II-1 (131.8 mm), phalanx III-3 (82.8 mm), femora (525 mm), tibiae (660 mm), fibulae (585, 580 mm), astragalocalcanea (98.5, 98.0 mm transversely), metatarsals I (76 mm), phalanx I-1 (101.1 mm), phalanx III-1 (114.2 mm), pedal ungual III (101.1 mm), phalanx IV-2 (63.7 mm), phalanx IV-3 (52.5 mm), phalanx IV-4 (56.5 mm), five phalanges, two pedal unguals, partial metatarsal V
Paratypes- (CM 78001) premaxillae (one incomplete, one fragmentary), maxillae, jugals, quadrates, ectopterygoid, pterygoids, braincase, cranial fragments, atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth cervical vertebra (~82.9 mm), tenth cervical vertebra, eleventh cervical vertebra (~78.5 mm), twelfth cervical vertebra, ten dorsal vertebrae (anterior 47 mm), seventeen ribs, eleven gastralia, incomplete sacrum (315 mm), twelve caudal vertebrae, eight chevrons, sternal plates (194.2 mm), ilia, pubes (450, 465 mm), ischia (305 mm), femora (505, 500 mm), tibiae (595 mm), fibulae (570 mm), astragalocalcanea (109.7 mm), phalanx III-3 (85.9 mm), pedal phalanx, four pedal unguals, metatarsal V (114.4 mm)
(FMNH PR2296; = BHM 2033) posterior mandible
Referred- ? metatarsal (Anonymous, 1997)
? material (DePalma, 2010)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Referred
- (BMRP 2013.4.1) dorsal ribs, caudal vertebra, pelvic elements, distal hindlimb including incomplete metatarsi (Holtz, Williams, Tremaine and Matthews, 2014)
?(MOR 9722) (~1.5-2 m) incomplete axial series, ribs, gastralia, partial forelimb including manual phalanges and unguals, ilia, partial pubes, ischia, partial tibia, tarsals, proximal metatarsals, pedal phalanges and pedal unguals (Flora et al., 2015)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, North Dakota, US

Paratype- (MRF 318) ninth cervical vertebra, eleventh cervical vertebra, twelfth cervical vertebra, dorsal rib, scapulocoracoid, radius (275 mm), ulna (280 mm)
Diagnosis- (after Lamanna et al., 2014) tall, crescentic cranial crest formed by posterodorsal processes of premaxillae (unknown in other caenagnathids except for Avimimus); body of maxilla lacking antorbital fossa (unknown in other caenagnathids except for ROM 43250); maxillary ascending process elongate and shaped like an inverted L (unknown in other caenagnathids); quadratojugal process of jugal dorsoventrally deep (unknown in other caenagnathids except for Avimimus); quadratojugal process of jugal bifurcated posteriorly (unknown in other caenagnathids); occipital condyle transversely wider than foramen magnum (unknown in other caenagnathids except for ROM 43250 and Avimimus); prominent lateral flange on symphyseal region of dentary; elongate retroarticular process of mandible (subequal in anteroposterior length to quadrate articulation); distal end of radius divided into two rounded processes (unknown in other caenagnathids); sulcus on ventromedial aspect of manual phalanx II-1; tubercle on anterior surface of astragalus near base of ascending process (unknown in other caenagnathids except for Nomingia and Avimimus).
Comments- Currie et al. (1994) described a posterior mandible which was larger than any Dinosaur Park Caenagnathus and differed from C. collinsi and C. sternbergi in glenoid morphology. They referred it to Caenagnathus sp.. Varricchio (2001) further noted it shared several characters with C. collinsi to the exclusion of C. sternbergi, and Lamanna et al. (2014) referred it to the present species.
Triebold et al. (2000) reported two new large oviraptorosaur specimens, identifying them as oviraptorids, though they were quickly reidentified as caenagnathid. These specimens were officially described by Lamanna et al. (2014) as the new taxon Anzu wyliei. The mandible is most similar to the Caenagnathus collinsi holotype, while the postcrania are not elmisaurine (six sacral vertebrae; preacetabular process long; preacetabular process hooked ventrally; fourth trochanter absent), but Lamanna et al.'s analysis does little to resolve the issues of American caenagnathid taxonomy.
Schachner et al. (2006; 2007) announced MRF 318 as Chirostenotes sp., which was later described as a specimen of Anzu by Lamanna et al. (2011; 2014).
Anonymous (1997) reported a Chirostenotes metatarsal associated with Tyrannosaurus specimen FMNH PR2081, which is tentatively assigned to the same species based on provenence.
DePalma (2010) referred material to Chirostenotes and Caenagnathus, which may belong to Anzu given their provenence,.
Holtz et al. (2014, 2015) announced a new specimen (BMRP 2013.4.1- Holtz pers. comm, 2015), which has only distal tarsal IV fused to the metatarsus, unfused metatarsals, and "a pair of pronounced cruciate ridges on the plantar surface of metatarsal III" which don't extend proximally as far as Elmisaurus.
References- Currie, Godfrey and Nessov, 1994. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Anonymous, 1997. Tyrannosaurus rex; A Highly Important and Virtually Complete Fossil Skeleton. Sotheby’s. 56 pp.
Triebold, Nuss and Nuss, 2000. Initial report of a new North American Oviraptor. In: The Florida Symposium on Dinosaur Bird Evolution, Presented by the Florida Institute of Paleontology at the Graves Museum of Archaeology and Natural History, Dania Beach, Florida, USA. p. 25.
Varricchio, 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from Montana. pp. 42-57. in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. New Research Inspired by the Paleontology of Philip J. Currie. Indiana Univ. Press.
Schachner, Lyson and Hanks, 2006. A preliminary report of a new specimen of Chirostenotes (Oviraptorosauria: Theropoda) from the Hell Creek Formation of North Dakota. Journal of Vertebrate Paleontology. 26, 120A.
Schachner, Lyson, Atterholt and Hanks, 2007. A preliminary report of a new specimen of Chirostenotes (Oviraptorosauria: Theropoda) from the Hell Creek Formation of North Dakota. Journal of Vertebrate Paleontology. 27(3), 141A.
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.
Lamanna, Sues, Schachner and Lyson, 2011. A new caenagnathid oviraptorosaur (Theropoda: Maniraptora) from the Upper Cretaceous (Maastrichtian) Hell Creek Formation of the Western United States. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 140.
Holtz, Williams, Tremaine and Matthews, 2014. New additions to the Hell Creek Formation (Upper Maastrichtian) vertebrate fauna of Carter County, Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 149.
Lamanna, Sues, Schachner and Lyson, 2014. A new large-bodied oviraptorosaurian theropod dinosaur from the Latest Cretaceous of Western North America. PLoS ONE. 9(3), e92022.
Flora, Wilson, Gardner and Fowler, 2015. A three-dimensionally articulated probable oviraptorosaur from the Hell Creek Formation of Montana. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 124-125.
Holtz, Williams and Tremaine, 2015. A new specimen of Anzu (Caenagnathidae, Oviraptorosauria): Implications for the proposed Caenagnathinae/Elmisaurinae division and for cursoriality in caenagnathids. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 146.

Elmisaurinae Osmolska, 1981 vide Currie, 2001
Definition- (Elmisaurus rarus <- Caenagnathus collinsi) (Hendrickx, Hartman and Mateus, 2015)
References- Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.

Kuszholia Nessov, 1992
?= Caenagnathasia Currie, Godfrey and Nessov, 1994
K. mengi Nessov, 1992
?= Caenagnathasia martinsoni Currie, Godfrey and Nessov, 1994
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan

Holotype- (ZIN PO 4602) posterior synsacrum (11, 10 mm)
Paratypes- ?(ZIN PO 4623) anterior synsacral centra (13, 10 mm)
?(ZIN PO coll.) sacral vertebrae (~20-30 mm)
Referred- ?(CCMGE 401/12457; holotype of Caenagnathasia martinsoni) anterior dentaries (Currie, Godfrey and Nessov, 1994)
?(CCMGE 402/12457; paratype of Caenagnathasia martinsoni) incomplete dentary (Currie, Godfrey and Nessov, 1994)
?(CCMGE 479/12457) proximal femur (Nessov, 1995; described by Sues and Averianov, 2015a)
?(ZIN PH 802/16) four incomplete fused sacral vertebrae (17, 14, 16.3, 16.4 mm) (Sues and Averianov, 2015a)
?(ZIN PH 932/16) incomplete posterior cervical vertebra (17 mm) (Sues and Averianov, 2015a)
?(ZIN PH 933/16) incomplete anterior cervical vertebra (19.5 mm) (Sues and Averianov, 2015a)
?(ZIN PH 934/16) incomplete dorsal vertebra (16.4 mm) (Sues and Averianov, 2015a)
?(ZIN PH 935/16) dorsal vertebra (Sues and Averianov, 2015a)
?(ZIN PH 936/16) dorsal vertebra (Sues and Averianov, 2015a)
?(ZIN PH 937/16) posterior dorsal centrum (Sues and Averianov, 2015a)
?(ZIN PH 2354/16) anterior dentaries (Sues and Averianov, 2015a)
?(ZIN PO 4603) incomplete anterior cervical vertebra (16 mm) (Nessov, 1992; described by Sues and Averianov, 2015a)
?(ZIN PO 5234) partial cervical vertebra (Sues and Averianov, 2015a)
Diagnosis- (after Currie et al., 1994; compared to Chirostenotes) fluting on the lingual margin of the occlusal edge not as distinct (ontogenetic?); first anterior occlusal groove larger; first pair of lateral occusal ridges do not meet ventrally; no tubercles on the midline or at the base of the first lateral occlusal ridge; lateral groove narrower and deeper (ontogenetic?); foramina on the floor more obvious (ontogenetic?); vascular grooves less conspicuous on the symphysial shelf; no foramina on the floor of the midline depression.
(suggested) larger sacral pleurocoels than Chirostenotes or Shixinggia (size-related?).
Other diagnoses- None of the characters listed in Nessov's (1992) diagnosis of Kuszholia are very characteristic. Dorsoventrally compressed sacral centra are common in maniraptorans. The robust second to last sacral transverse process is seen in Chirostenotes as well. Deep posterior sacral pleurocoels are common in caenagnathoids, and far from being small, Kuszholia's are large compared to other taxa. The posterior articular surface is not large, being smaller than mid sacral vertebrae, and its concavity is plesiomorphic for theropods. The large postzygapophyses are only notable compared to birds- they are normal for an oviraptorosaur. The sacrals of Chirostenotes also have a ventral median groove which is most pronounced at the junction of centra.
Currie et al. (1994) stated there was no second anterior occlusal groove in Caenagnathasia, but Sues and Averianov (2015a) reported it as likely present in new specimen ZIN PH 2354/16.
Comments- Note that while volume 30(10) of the Canadian Journal of Earth Sciences lists its date as October 1993, it was not published until February or March of 1994.
The holotype (ZIN PO 4602) consists of the last two sacral vertebrae and a fragment of the third to last, fused together. The centra are dorsoventrally compressed (anterior articular surface ~66% as tall as wide) with an oval and slightly concave posterior articular surface. The sacrum seems to be slightly concave ventrally and the centrum junctions are expanded both ventrally and laterally. There are small but deep pleurocoels present in each centrum. A median ventral groove is present, which is especially well marked at the centrum junctions. The postzygapophyses of the last sacral protrude markedly past the centrum. The second to last transverse process is long, robust and perpendicular to the sacral long axis, while the last transverse process is about half the length but otherwise similar.
Nessov referred another specimen (ZIN PO 4623) consisting of the first two fused centra of another sacrum. These are similar in being dorsoventrally compressed (posterior articular surface ~63% as tall as wide). The anterior articular surface is kidney-shaped and slightly heterocoelous. It is also similar in being slightly concave, with expanded centrum junctions and deep, oval pleurocoels in each centrum. Ventrally, there is a slight midline groove. This was later referred to Caenagnathasia by Sues and Averianov (2015a). Nessov referred to additional isolated vertebrae virtually identical to these, but 2-2.5 times larger. These may belong to larger oviraptorosaurs, therizinosaurs or dromaeosaurids, though they are impossible to evaluate without more information. Finally, he stated strongly pneumatic vertebrae with closed neurocentral sutures (unlike juvenile therizinosaurs or sauropods) could belong to Kuszholia. One is illustrated (ZIN PO 4603), which is an anterior cervical with strongly overhanging prezygapophyses, a large neural canal, an elongate centrum (2.75 times posterior height) which reaches posteriorly past the neural arch, and perhaps a large teardrop shaped pleurocoel. This was later referred to Caenagnathasia by Sues and Averianov (2015a).
Nessov and Panteleev (1993) figured and described a partial sacrum they referred to Kuszholia sp. (ZIN PO 4826). Zelenkov and Averianov (2011) stated this differs from Kuszholia in "the absence of a pleurocoel in the posterior vertebra and in the shallow slitlike pleurocoel in the penultimate vertebra", and while the pleurocoel shape appears similar, the absence of a pleurocoel in the last sacral is indeed different. They believe the specimen to be similar to Zanabazar, but I don't see any particular reseblence and refer it to Maniraptora incertae sedis here pending further study. Nessov (1995) later figured a centrum with a slit-like pleurocoel (ZIN PO 467) which he stated was "possibly from bird ?Kuszholia sp. or from a theropod or a segnosaur", and this was identified as a therizinosauroid by Sues and Averianov (2015b). He also figured proximal femur CCMGE 479/12457 as a theropod before it was referred to Caenagnathasia by Sues and Averianov (2015a).
Relationships- The holotype was first incorrectly listed as ZIN PO 3486 and identified as an ichthyornithid (Nessov, 1990). Nessov (1992) later named this as a new genus in its own family Kuszholiidae, under "suborder Theropoda + Aves." He viewed it as possibly a basal flightless bird like Patagopteryx or possibly a non-bird theropod which was convergent with birds. Nessov and Panteleev (1993) later assigned it to Patagopterygiformes, though their argument has yet to be translated from Russian. Kurochkin (2001) retained Kuszholia as Aves (sensu Chiappe) incertae sedis, and noted that patagopterygiform affinities were not yet verified. He did cite two supposed similarities though- enlarged third pair of sacral transverse processes and ventral sacrum convex. Yet Kuszholia's sacrum is ventrally concave (as in Patagopteryx and many other theropods) and the large transverse processes are on the second to last sacral. As all theropods have at least five sacrals, this corresponds to the fourth sacral or greater. All posterior sacral transverse processes are broken off in Patagopteryx in any case. Patagopteryx further differs in lacking sacral pleurocoels and having a convex posterior articular surface, as noted by Kurochkin. Kurochkin later (2006) placed Kuszholia in Ornithuromorpha (his Ornithurae), but outside Carinatae, in his phylogram, though without stated support.
The postzygapophyses of the last sacral are much larger than any avebrevicaudan, suggesting it is not a member of that clade. As noted above, dorsoventrally compressed centra are common in maniraptorans, while ventral grooves are present in the posterior sacrals of Ornitholestes and most maniraptoriforms. Very few maniraptorans have pleurocoels extending to the last sacral centrum, with examples limited to Neimongosaurus, caenagnathoids and reportedly Bambiraptor (however, personal examination of AMNH 30556 indicates pleurocoels are only present on the second, and presumably the broken first, sacral centrum). Adult therizinosaurs are far larger, with even the smallest basal members (e.g. Beipiaosaurus) being four times as big, and the fusion does indicate Kuszholia's holotype is from an adult. There are small caenagnathoids though, including Caenagnathasia from the same formation. In fact, Caenagnathasia would have comparably sized sacrals to Kuszholia if scaled from other caenagnathoids. Note Sues and Averianov (2015a) assigned the Kuszholia paratype and referred cervical to Caenagnathasia without commenting on why the holotype was also not referrable. Indeed, there seems to be no reason to refer any of the Bissekty caenagnathoid material to one genus instead of the other as the type synsacrum of Kuszholia cannot be compared to the type dentaries of Caenagnathasia. If they are synonymous, Kuszholia would have priority by two years, which is awkward as its synsacrum is less diagnostic and less phylogenetically determinable than Caenagnathasia's dentaries. The alternative is listing all material except the three dentaries and Kuszholia's holotype as Caenagnathoidea/idae indet., but tentative synonymization is chosen here.
The holotype is generally similar to Chirostenotes, but differs in having much larger pleurocoels (size related?) and narrower postzygapophyses. It also has larger pleurocoels than Shixinggia. Avimimus is unique among oviraptorosaurs in lacking sacral pleurocoels, so is quite different.
References- Nessov, 1990. Small ichthyornithiform bird and other bird remains from Bissekty Formation (Upper Cretaceous) of central Kyzylkum Desert. Proceedings of the Zoological Institute, Leningrad. 210, 59-62.
Nessov, 1992. Review of localities and remains of Mesozoic and Paleogene birds of the USSR and the description of new findings. Russkii Ornitologicheskii Zhurnal. 1(1), 7-50.
Nessov and Panteleev, 1993. On the similarity of the Late Cretaceous ornithofauna of South America and Central Asia. Trudy Zoologicheskogo Instituta, RAN. 252, 84-94.
Currie, Godfrey and Nessov, 1994. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of class Aves. Archaeopteryx. 13, 47-66.
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.
Nessov, 1997. Cretaceous nonmarine vertebrates of Northern Eurasia. Izdatelstvo Sankt-Peterburgskogo Universiteta, Saint Petersburg. 218 pp.
Kurochkin, 2001. Mesozoic birds of Mongolia and the former USSR. in Benton, Shishkin, Unwin and Kurochkin, eds. The Age of Dinosaurs in Russia and Mongolia. 533-559.
Kurochkin, 2006. Parallel evolution of theropod dinosaurs and birds. Entomological Review. 86(suppl. 1), S45-S58.
Zelenkov and Averianov, 2011. Synsacrum of a primitive bird from the Upper Cretaceous of Uzbekistan. Paleontological Journal. 45(3), 314-319.
Sues and Averianov, 2015a. New material of Caenagnathasia martinsoni (Dinosauria: Theropoda: Oviraptorosauria) from the Bissekty Formation (Upper Cretaceous: Turonian) of Uzbekistan. Cretaceous Research. 54, 50-59.
Sues and Averianov, 2015b. Therizinosauroidea (Dinosauria: Theropoda) from the Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.
K. sp. (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China

Material- (IVPP V20377) anterior dentaries
Comments- Yao et al. (2015) referred this specimen to Ceanagnathasia sp., and noted it resembles the C. martinsoni holotype more than the specimen (ZIN PH 2354/16) referred to that species by Sues and Averianov. The age difference suggests the specimen is not conspecific with the Bissekty species, and AMNH 6754 and 6755 may belong to the same taxon.
Reference- 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.

Elmisaurus Osmolska, 1981
Diagnosis- (modified from Currie, 1989) distal end of metatarsal II curves anteriorly away from the longitudinal axis of the metatarsus (unknown in Leptorhynchos).
(after Currie et al., 2015) m. tibialis cranialis tubercle on the dorsal surface of metatarsals II (unknown in Leptorhynchos); posterior medial and lateral ridges of metatarsal III midshaft close to trochlear ridges (unknown in Leptorhynchos).
Other diagnoses- Osmolska (1981) listed three diagnostic characters- tarsometatarsal fusion; posterior surface of metatarsus deeply concave; proximolateral process on metatarsal IV. Yet all are also present in Avimimus and at least the first and third in Leptorhynchos.
Currie (1989) noted metatarsals II and IV are subequal in length to III (>93%), but Avimimus' ratios are 93% and 94% respectively, making this an elmisaurine synapomorphy.
Currie et al. (2015) listed the "large, compound proximal protuberance on the posterior surfaces of metatarsals II-IV" as being diagnostic, but this is present in Chirostenotes and Leptorhynchos as well.
Comments- While Currie (1989) stated the proximal end of metatarsal III is triangular in Elmisaurus, unlike Chirostenotes, the latter does have a triangular section at some points. The only differences in cross sectional shape between caenagnathid taxa in the literature is due to proximodistal position along the metatarsal (see detailed discussion under Leptorhynchos comments).
References- Osmolska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26, 1319-1324.
Currie, 2001. Theropod dinosaurs from the Cretaceous of Mongolia. in Benton, Shishkin, Unwin and Kurochkin (eds). The Age of Dinosaurs in Russia and Mongolia. pp 434-455.
E. rarus Osmolska, 1981
= Chirostenotes rarus (Osmolska, 1981) Paul, 1988
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia

Holotype- (ZPAL MgD-I/172) (1.68 m) distal tarsals, metatarsal II (147 mm), metatarsal III (157 mm), metatarsal IV (147 mm)
Paratypes- (ZPAL MgD-I/20) distal tarsals, proximal metatarsal II, proximal metatarsal III (~173 mm), proximal metatarsal IV
(ZPAL MgD-I/98) rib fragments, gastralial fragments, two partial mid sacral centra (37 mm), proximal scapula, metacarpal I (45 mm), phalanx I-1 (65 mm), manual ungual I (44 mm), incomplete metacarpal II (~63 mm), phalanx II-1 (66 mm), phalanx II-2 (66 mm), phalanx III-1 (30 mm), phalanx III-2 (30 mm), phalanx III-3 (43 mm), proximal manual ungual, proximal pubes, partial ischia, partial femora (~308 mm), partial tibia (~340 mm), phalanx I-1 (26 mm), pedal ungual I (~23 mm), distal metatarsal II, phalanx II-1 (45 mm), partial phalanx II-2 (34 mm), fragmentary pedal ungual II (~31 mm), distal metatarsal III (~151 mm), phalanx III-1 (46 mm), phalanx III-2 (32 mm), phalanx III-3 (30 mm), distal metatarsal IV, fragmentary phalanx IV-1, phalanx IV-2 (22 mm), phalanx IV-3 (19 mm), phalanx IV-4 (18 mm), fragmentary pedal ungual IV (~26 mm), fragments
Referred- (IGM 102/6; field number PJC 2000.1) tarsometatarsus (194 mm; II 172.4, III 185, IV 175.7 mm), metatarsal V (70.3 mm) (Currie, Funston and Osmólska, 2016)
(IGM 102/7; field number PJC 2000.2) (~23 kg adult) frontal (50.5 mm), ~third to fifth cervical neural arch, partial posterior cervical vertebra (26.4 mm), incomplete anterior dorsal vertebra (27.7 mm), three proximal dorsal ribs, dorsal rib fragments, gastralial fragments, vertebral fragment, distal phalanx II-2, incomplete manual ungual II (40 mm), phalanx III-1 (28 mm), phalanx III-3 (40 mm), proximal femur (~246 mm), tibiae (323 mm), pedal phalanx I-1 (23.1 mm), incomplete pedal ungual I (30 mm), distal tarsal III, (tarsometatarsus ~176 mm) metatarsal II (161.8 mm), partial metatarsal III (~172 mm), metatarsal IV (162.2 mm) (Currie, 2001; described by Currie, Funston and Osmólska, 2016)
(IGM 102/8; field number PJC 2000.3) metatarsal IV (164 mm) (Currie, 2001; described by Currie, Funston and Osmólska, 2016)
(IGM 102/9; field number PJC 2001.8) proximal tarsometatarsus (Currie, 2001; described by Currie, Funston and Osmólska, 2016)
(IGM 102/10; field number PJC 2002.4) vertebra, tibia (Currie, Funston and Osmólska, 2016)
?(IGM 102/107) anterior dentaries (Tsuihiji, Watabe, Tsogtbaatar and Barsbold, 2016)
?(?IGM coll.) femur (Currie, 2001)
? ventral skull (www.paleofile.com)
Diagnosis- (after Osmolska, 1981) metacarpal I ~66% [69%] the length of phalanx I-1 (76% in Hagryphus; but unknown in E. elegans and other caenagnathids).
Other diagnoses- Osmolska (1981) listed several supposedly diagnostic characters- metatarsal III visible anteriorly for ~90% of its length (also in the E. elegans holotype, but not in IGM 102/7); length/width ratio of metatarsus 0.18 (also in E. elegans); digit III markedly thinner than digits I and II both of which are equally thick (untrue as digit I is more gracile than II, and conditions are the same in other caenagnathids); ventroposterior portions of manual phalanges distinctly thickened at proximal surfaces (also in Chirostenotes).
Currie et al. (2016) claimed the manual unguals of Elmisaurus were more robust and curved than Chirostenotes, with lower proximodorsal lips, they are comparable to Hagryphus.
Comments- Discovered in 1970 and described in 1981, Elmisaurus rarus was the first caenagnathid discovered with both manual and pedal remains. This allowed the synonymy between Chirostenotes and Macrophalangia to be demonstrated. Elmisaurus has been viewed as a relative of Chirostenotes (first in Elmisauridae and later in Caenagnathidae) from its discovery until the 2000s, when this was questioned by Maryanska et al. (2002). They noted it differs from Chirostenotes in having a vascular foramen between metatarsals III and IV, an m. tibialis cranialis tubercle on the dorsal surfaces of metatarsals II-IV, a deeply concave posterior side, and a proximolateral process on metatarsal IV. Yet these are all apomorphies that tell us nothing about Elmisaurus' relationships unless we find non-caenagnathid taxa that share them. Maryanska et al. stated pygostylians have the first two characters, and Avimimus the last, but Elmisaurus otherwise resembles oviraptorosaurs and is quite dissimilar to any paravian. The relationship with Avimimus is possible, as found in my modified version of Senter's (2007) matrix, but that taxon is a caenagnathid in that tree anyway. It is especially confusing that Maryanska et al. continue to refer elegans to Caenagnathidae, as it shares the characters they describe for Elmisaurus. Elmisaurus elegans is similar enough to Chirostenotes pergracilis to be synonymized by some authors (e.g. Currie and Russell, 1988), and Elmisaurus has been synonymized with Chirostenotes by others (e.g. Paul, 1988). It is completely unwarranted to widely separate the two genera, and Elmisaurus has more recently been reestablished as a caenagnathid (e.g. Longrich et al., 2013; Currie et al., 2016).
Currie (2001, 2002) first mentioned new specimens of Elmisaurus rarus, most of which were later noted in an abstract (Funston and Currie, 2014). Currie et al. (2016) described these specimens, as well as additional material of ZPAL MgD-I/98. They identified the manual ungual of the latter as "almost certainly" from digit I, which was the possible identification of Osmolska as well despite her figure 2 placing it on digit II. While Funston and Currie state the frontal "suggests the presence of a high crest", which is also assumed by Currie et al., yet no evidence is presented to support this and the frontal shows no dorsal projection. Only one specimen mentioned by Currie (2001/2002) was not described in Currie et al.- a femur referred to cf. Elmisaurus rarus. Tsuihiji et al. 92016) described a partial elmisaurine mandible that is here referred to E. rarus based on stratigraphy. Ford (www.paleofile.com) reported a partial skull photographed in "a Japanese guide book".
References- Osmolska, 1981. Coossified tarsometatarsi in theropod dinosaurs and their bearing on the problem of bird origins. Palaeontologia Polonica. 42, 79-95.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Currie, 2001. Nomadic Expeditions, Inc. report on fieldwork in Mongolia, September 2000. In: Alberta Palaeontological Society, fifth annual symposium, jointly presented by Alberta Palaeontological Society and Department of Earth Sciences, Mount Royal College. 12-16.
Currie, 2002. Report on fieldwork in Mongolia, September 2001: In: Alberta Palaeontological Society, sixth annual symposium, “Fossils 2002’, presented by Alberta Paleontological Society, in conjunction with Canadian Society of Petroleum Geologists, Paleontological Division and Department of Earth Sciences, Mount Royal College. 8-12.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian Aguja Formation of West Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Funston and Currie, 2014. New Elmisaurus (Dinosauria: Oviraptorosauria) material from Mongolia and Alberta, Canada, and its bearing on North American caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts, 2014. 134.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osmólska, 2016. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Funston, Currie and Burns, 2016. New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
Tsuihiji, Watabe, Tsogtbaatar and Barsbold, 2016. Dentaries of a caenagnathid (Dinosauria: Theropoda) from the Nemegt Formation of the Gobi Desert in Mongolia. Cretaceous Research. doi: 10.1016/j.cretres.2016.03.007
E? elegans (Parks, 1933) Currie, 1989
= Ornithomimus elegans Parks, 1933
pr= Caenagnathus sternbergi Cracraft, 1971
= Macrophalangia elegans (Parks, 1933) Koster, Currie, Eberth, Brinkman, Johnston and Braman, 1987
= Chirostenotes elegans (Parks, 1933) Currie and Russell, 1988
= Chirostenotes sternbergi (Cracraft, 1971) Snively, Currie, Brinkman, Ryan, Braman, Gardner, Lam, Spivak and Neuman, 2001
= "Leptorhynchos" elegans (Parks, 1933) Longrich, Barnes, Clark and Millar, 2013a
= Leptorhynchos elegans (Parks, 1933) Longrich, Barnes, Clark and Millar, 2013b
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada

Holotype- (ROM 781) partial distal tarsal III, distal tarsal IV, metatarsal II (155 mm), partial metatarsal III (161 mm), metatarsal IV (157 mm)
Referred- ?(CMN 2690; holotype of Caenagnathus sternbergi) posterior mandible (Cracraft, 1971)
(ROM 37163) distal metatarsal II (Currie, 1989)
?(RTMP 79.8.622) anterior dentaries (Currie et al., 1994)
(RTMP 82.16.6) tarsometatarsus (mtII 152, mtIII 172, mtIV 160, mtV 44.3 mm) (Snively, 2000)
(RTMP 82.39.4) proximal tarsometatarsus (Currie, 1989)
(RTMP 84.163.36) distal metatarsal III (Funston and Currie, 2014b; described by Funston et al., 2016)
(RTMP 86.36.186) distal metatarsal III (Funston and Currie, 2014b; described by Funston et al., 2016)
(RTMP 88.36.104) distal metatarsal II (Funston and Currie, 2014b; described by Funston et al., 2016)
?(RTMP 90.56.6) dentaries (Currie et al., 1994)
?(RTMP 91.144.1) incomplete dentaries (Currie et al., 1994)
?(RTMP 92.36.390) incomplete dentaries (Currie et al., 1994)
?(RTMP 92.40.44) anterior dentaries (Currie et al., 1994)
(RTMP 93.36.181) partial tarsometatarsus (mtII 221, mtIV 221 mm) (Funston and Currie, 2014b; described by Funston et al., 2016)
(RTMP 93.36.630) distal metatarsal III (Funston and Currie, 2014b; described by Funston et al., 2016)
(RTMP 94.12.880) tibia (280 mm) (Funston and Currie, 2014b; described by Funston et al., 2016)
(RTMP 96.12.141) incomplete tarsometatarsus (Currie, 2005; described by Funston et al., 2016)
(RTMP 2000.12.8) distal tarsal III fused to metatarsal II, phalanx II-1, pedal ungual II, distal tarsal IV fused to metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-?2 (22.3 mm) (Funston and Currie, 2014b; described by Funston et al., 2016)
?(RTMP 2001.12.12) incomplete mandibles (188 mm) (Currie, 2005; described by Funston and Currie, 2014a)
(RTMP 2005.49.190) metatarsal III (Funston and Currie, 2014b; described by Funston et al., 2016)
(UALVP 55585) (subadult) partial metatarsal III (Funston and Currie, 2014b; described by Funston et al., 2016)
Late Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US

?(MOR 1107) articular (Varricchio, 2001)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US

(MOR 752) astragalar fragment, partial metatarsal II (~131 mm), distal phalanx II-1, phalanx II-2 (26.1 mm), metatarsal fragment, phalanx III-1 (34 mm), phalanx III-2 (23.3 mm), phalanx III-3 (25.1 mm), pedal ungual III (24.3 mm), phalanx IV-1 (23.1 mm), phalanx IV-2 (16.2 mm), phalanx IV-3 (14.4 mm), phalanx IV-4 (16.5 mm), pedal ungual IV (21 mm) (Varricchio, 2001)
?(NS.31996.114H) distal metatarsal II (~216 mm) (Buckley, 2002)
?(NS.32001.017B) distal metatarsal II (Buckley, 2002)
(RTMP 96.5.12) distal metatarsal III (Funston and Currie, 2014b; described by Funston et al., 2016)
Diagnosis- (after Currie, 1989; compared to Elmisaurus rarus) metatarsal II more slender; posteromedial corner of metatarsus more deeply emarginated in dorsal view; longitudinal ridge-like posterolateral margin of metatarsal IV not as powerfully developed proximally; close to distal articular surfaces, small processes of metatarsals II and IV overlap metatarsal III.
(after Currie et al., 1994; compared to Caenagnathus collinsi holotype) higher articular ridge on mandible; medial glenoid shorter anteroposteriorly; mandibular ramus anterior to glenoid less robust.
(after Varricchio, 2001) chorda tympani foramen/slot absent.
(after Longrich, Barnes, Clark and Millar, 2013a; compared to Leptorhynchos gaddisi) tip of beak strongly upturned, with anterior occlusal margin projecting vertically; anterior margin of symphysis straight; chin squarish in lateral view.
Comments- This was originally described as a species of Ornithomimus (Parks, 1933), though Sternberg (1934) soon recognized it was not an ornithomimid. Russell (1972) synonymized it with Macrophalangia canadensis, while Currie and Russell (1988) synonymized both with Chirostenotes pergracilis. The latter authors believed the elegans specimen to be a gracile morph of the species, which could be called Chirostenotes elegans if it was in fact taxonomically distinct. Currie (1989) described two new specimens (ROM 37163 and RTMP 82.39.4), noting similarities to Elmisaurus rarus that were not seen in Chirostenotes pergracilis. He made the new combination Elmisaurus elegans. These similarities were said to be insufficient by Sues (1997) (though without justification), who called the species Chirostenotes elegans. Maryanska et al. (2002) and Osmolska et al. (2004) also assign elegans to Chirostenotes instead of Elmisaurus, though they never state their rationale. Varricchio (2001) referred a pes to E. elegans based on the distal process of metatarsal II, while Buckley (2002) referred two distal metatarsal II's to the species because of their longitudinal ridge (which would have made the posterior metatarsus surface deeply concave). However, the specimens described by Buckley are larger than MOR 752 or Dinosaur Park Formation specimens, perhaps suggesting they belong to a distinct species of Elmisaurus.
Cracraft (1971) named Caenagnathus sternbergi, known from a posterior mandible that differs from C. collinsi in several characters. Currie et al. (1994) later described five dentaries which also differ from C. collinsi. They referred to these as Caenagnathus cf. sternbergi, as none are directly comparable to the C. sternbergi holotype. Varricchio (2001) described an additional mandibular fragment from the contemporaneous Two Medicine Formation that corresponds with the C. sternbergi holotype. Currie (2005) illustrated a new mandible which confirms Currie et al. were correct to refer the dentaries to C. sternbergi, which was analyzed by Funston et al. (2013) and described in detail by Funston and Currie (2014a).
Currie and Russell (1988) first proposed the synonymy of Caenagnathus collinsi with Chirostenotes pergracilis, and Caenagnathus sternbergi with Elmisaurus (then Chirostenotes) elegans. This is followed here. Longrich et al. (2013a) place elegans in their new genus Leptorhynchos, based on the Aguja Formation L. gaddisi. While the gaddisi and elegans material is similar, their reasons for separating these from Elmisaurus are flawed (see Leptorhynchos comments), and there is no reason to refer elegans to Leptorhynchos over Elmisaurus or vice versa. This is followed by Funston and Currie (2014b), but Currie et al. (2016) refer elegans to Leptorhynchos without justification. Their referral of some mandibular material (C. sternbergi holotype, RTMP 90.56.6, 2001.12.12) to Chirostenotes pergracilis has not been evaluated here.
References- Parks, 1933. New species of dinosaurs and turtles from the Upper Cretaceous formations of Alberta. University of Toronto Studies, Geological Series. 34, 1-33.
Sternberg, 1934. Notes on certain recently described dinosaurs. Canadian Field Naturalist. 48, 7-8.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada. Canadian Journal of Earth Sciences. 9, 375-402.
Koster, Currie, Eberth, Brinkman, Johnston and Braman, 1987. Sedimentology and Palaeontology of the Upper Cretaceous Judith River/Bearpaw Formations at Dinosaur Provincial Park, Alberta, Field Trip #10. Geological Association of Canada, Mineralogical Association of Canada, Joint Annual Meeting, Saskatoon, Saskatchewan. 130 p.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25, 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26, 1319-1324.
Currie, Godfrey and Nessov, 1994. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30(10), 2255-2272.
Snively, Currie, Brinkman, Ryan, Braman, Gardner, Lam, Spivak and Neuman, 2001. Alberta's dinosaurs and other fossil vertebrates: Judith River and Edmonton groups (Campanian-Maastrichtian). In Hill (ed.). Mesozoic and Cenozoic Paleontology in the Western Plains and Rocky Mountains. Guidebook for the Field Trips of the Society of Vertebrate Paleontology 61st Annual Meeting. Museum of the Rockies Occasional Paper No. 3. 47-75.
Varricchio, 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from Montana. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University Press. 42-57
Buckley, 2002. New material of Elmisaurus (Theropoda, Elmisauridae) from the Late Cretaceous Hell Creek Formation of Southeastern Montana. Journal of Vertebrate Paleontology. 22(3), 39A.
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.
Funston, Currie and Murray, 2013. Examining the diet of a toothless dinosaur: Evidence supporting a herbivorous diet in Caenagnathus (Dinosauria: Oviraptorosauria). Journal of Vertebrate Paleontology. Program and Abstracts 2013, 131.
Longrich, Barnes, Clark and Millar, 2013a. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Longrich, Barnes, Clark and Millar, 2013b. Correction to "Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae". Bulletin of the Peabody Museum of Natural History. 54(2), 263-264.
Funston and Currie, 2014a. A previously undescribed caenagnathid mandible from the late Campanian of Alberta, and insights into the diet of Chirostenotes pergracilis (Dinosauria: Oviraptorosauria). Canadian Journal of Earth Sciences. 51(2), 156-165.
Funston and Currie, 2014b. New Elmisaurus (Dinosauria: Oviraptorosauria) material from Mongolia and Alberta, Canada, and its bearing on North American caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts, 2014. 134.
Currie, Funston and Osmólska, 2016. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Funston, Currie and Burns, 2016. New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.
E. sp. (Funston, Currie and Burns, 2016)
Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material
- (RSM P2600.1) ?scapula, proximal tibia, distal tibia, proximal fibula, astragalocalcaneum, distal metatarsal II, distal metatarsal III
(RSM P2161.1) metatarsal II
Diagnosis- (after Funston et al., 2016) distal tibia wider than E. elegans where it contacts the astragalocalcaneum; distal metatarsal II shaft straight at its distal end as in E. rarus; distal metatarsal III as mediolaterally wide as anteroposteriorly deep.
Reference- Funston, Currie and Burns, 2016. New elmisaurine specimens from North America and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica Polonica. 61(1), 159-173.

Leptorhynchos Longrich, Barnes, Clark and Millar, 2013b
= "Leptorhynchos" Longrich, Barnes, Clark and Millar, 2013a
L. gaddisi Longrich, Barnes, Clark and Millar, 2013b
= "Leptorhynchos gaddisi" Longrich, Barnes, Clark and Millar, 2013a
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Holotype
- (TMM 45920-1) dentaries
Referred- ?(TMM 42335-40) caudal centrum
?(TMM 43057-36) distal tarsal IV fused to proximal metatarsal IV
?(TMM 43057-354) distal metatarsal III
?(TMM 43057-357) pedal ungual
Diagnosis- (after Longrich et al., 2013a) differs from Elmisaurus? elegans in having more anteriorly projecting tip of beak; strongly rounded anteroventral margin of symphysis; lateral occlusal margins of dentaries not as strongly divergent in dorsal view, with tip of beak being narrower and more spoon-shaped in dorsal view.
Comments- Some of this material was first reported as Caenagnathidae indet. by Longrich et al. (2010). While Longrich et al. (2013a) intended to name Leptorhynchos gaddisi, they did not specify a type species, making the correction Longrich et al. (2013b) necessary to specify gaddisi as the type species and make the names official.
Longrich et al. (2013a) proposed this species is sister to elegans, both being closer to Chrisostenotes than to Elmisaurus rarus. Their phylogenetic analysis would seem to support this, but Elmisaurus rarus' basal position is due to their figure 14 being an Adams consensus tree. Because rarus lacks described mandibular material, it cannot be placed precisely compared to Caenagnathasia and their Caenagnathus OTUs, so will have a position at the base of the largest clade it can belong to in an Adams consensus tree. When their analysis is rerun to a posteriori exclude taxa incomparable to Elmisaurus rarus, it is in a trichotomy with Chirostenotes and elegans. Further, their analysis misacodes rarus as lacking metatarsal fusion and excludes the other five characters suggested by Currie (1989) to unite rarus and elegans to the exclusion of pergracilis.
Longrich et al. do claim elegans is more similar to Chirostenotes in one way. They state "Longrich (2008a) tentatively placed [elegans] in Chirostenotes, because the third metatarsal has an anteroposteriorly flattened shaft that is concave and broadly exposed on the posterior of the metatarsus (Currie 1989). This is a derived feature found in Chirostenotes (Currie and Russell 1988) but not Elmisaurus (Osmólska 1981)." This seems related to their new character 205- "Metatarsal III with an ovoid or subtriangular cross section (0) or anteroposteriorly flattened, with a concave posterior surface (1). Primitively in theropods the third metatarsal has an ovoid cross section, or a triangular cross section in arctometatarsalian forms. This condition is retained in most oviraptorosaurs, including the basal caenagnathid Elmisaurus rarus. In Caenagnathinae, the third metatarsal is anteroposteriorly compressed." Yet the posterior transverse exposure of proximal metatarsal III proximally seems intermediate in E. rarus' holotype compared to the two specimens of elegans. More distally, Currie's (1989) figure 2P section indicates the posterior exposure is narrow as in E. rarus. Currie's (1989) figure 2Q shows E. rarus has a concave posterior metatarsal III surface as well. As for shape, the main issue seems to be the use of different proximodistal points along the bone. Currie and Russell (1988) state in Chirostenotes pergracilis "The proximal end, viewed dorsally, is diamond shaped, tapering both anteriorly (between the contact of metatarsals II and IV) and posteriorly. Its major horizontal axis, 17.5 mm long, is anteroposterior in orientation and thins backwards." Sternberg (1932) also states the Macrophalangia holotype (possibly Caenagnathus collinsi according to Longrich et al.) has a transversely compressed proximal end. The proximal ends of E. rarus' and elegans' metatarsal III are fused too well with surrounding bones to compare. Once C. pergracilis' anterior surface is exposed, "the bone twists until the medial surface is facing anteriorly" and "the anterior edge has broadened out to 7.5 mm to separate the adjacent metatarsals and is triangular in section" (Currie and Russell, 1988). elegans' holotype is broken at about this same point and also shows a triangular section. Note a triangular section is what Longrich et al. are claiming caenagnathines don't have, though it exists in both pergracilis and elegans. Currie (1989) was also wrong in comparing the proximal diamond shape of Chirostenotes pergracilis with the more distal triangular shape of elegans, and none of these areas have been described in E. rarus. At two-thirds down in Chirostenotes, "In section, a shallow concave surface faces posteriorly at this level, while slightly concave surfaces face posteromedially and posterolaterally for contact with the adjacent metatarsals." This appears similar to E. rarus from what the anterior and posterior views suggest, and matches how Snively (2000) described an RTMP "Elmisaurus sp." metatarsus that is probably elegans. It also matches the cross section of elegans illustrated by Currie (1989- fig. 2P). While Currie's figure 2Q of Elmisaurus rarus would suggest a slightly different shape where the articular surfaces are smaller and that for metatarsal IV doesn't angle posteromedially, the narrow anterior exposure of metatarsal III means it must have been taken more proximally, probably about halfway down considering the ratio between anterior and posterior exposure of metatarsal III. This leaves anteroposterior compression, which varies throughout the bone in elegans at least. Distally it's transversely compressed but proximally it's anteroposteriorly compressed. The E. rarus section which is probably intermediate in position is also intermediate in compression, being slightly transversely compressed. If Chirostenotes/Caenagnathus specimens are any indication, the bone switches back to transversely compressed at its proximal tip. So there are actually no valid described differences in metatarsal III sectional shape between Chirostenotes and any Elmisaurus species in the primary literature.
Correcting the coding for Elmisaurus rarus having fused metatarsals and adding the 5 other characters (one as an additional ordered state of the arctometatarsaly character) results in elegans and gaddisi being in a trichotomy with E. rarus. This elmisaur clade is sister to a caenagnath clade containing Chirostenotes, Hagryphus, Caenagnathus collinsi and C. sp. from Hell Creek. The only elements described in rarus, elegans and gaddisi are distal metatarsal III and proximal metatarsal IV. Neither of these seem particularly distinctive in any of the species, and rarus' limited illustrations and description makes them difficult to compare in depth. Thus there's no obvious reason to refer elegans to either Elmisaurus or Leptorhynchos, so it would have been easier and more accurate to just keep elegans in Elmisaurus and not name a new genus. Longrich et al.'s diagnoses for Leptorhynchos and each of its species are purely mandibular, so we can't even evaluate Elmisaurus rarus or the elegans holotype for them.
References- Sternberg, 1932. Two new theropod dinosaurs from the Belly River Formation of Alberta. Canadian Field-Naturalist. 46(5), 99-105.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis (Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta, Canada. Canadian Journal of Earth Sciences. 25, 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda) from North America. Canadian Journal of Earth Sciences. 26, 1319-1324.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Unpublished Masters Thesis. 273 pp.
Longrich, Sankey and Tanke, 2010. Texacephale langstoni, a new genus of pachycephalosaurid (Dinosauria: Ornithischia) from the upper Campanian Aguja Formation, southern Texas, USA. Cretaceous Research. 31, 274-284.
Longrich, Barnes, Clark and Millar, 2013a. Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Longrich, Barnes, Clark and Millar, 2013b. Correction to "Caenagnathidae from the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae". Bulletin of the Peabody Museum of Natural History. 54(2), 263-264.

Oviraptoridae Barsbold, 1976
Definition- (Oviraptor philoceratops <- Caenagnathus collinsi) (Maryanska et al., 2002; modified from Sereno, 1998)
Other definitions- (Oviraptor philoceratops <- Chirostenotes pergracilis) (Sereno, in press; modified from Padian et al., 1999)
= Ingeniidae Barsbold, 1981
= Oviraptoridae sensu Padian et al., 1999
definition- (Oviraptor philoceratops <- Chirostenotes pergracilis) (modified)
Comments- Supposed oviraptorid material from the Yalovach Formation of Tadjikistan (Nessov, 1995) probably belongs to therizinosaurs instead (Alifanov and Averianov, 2006).
References- Barsbold, 1976. On a new Late Cretaceous family of small theropods (Oviraptoridae fam. n.) of Mongolia. Doklady Akademia Nauk SSSR. 226, 685-688. [in Russian]
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].
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.
Smith and Molnar, 2006. Jaw musculature and function in oviraptorosaurs. Journal of Vertebrate Paleontology. 26(3), 126A.

Oviraptorinae Barsbold, 1976 sensu Barsbold, 1981
Definition- (Oviraptor philoceratops + Citipati osmolskae) (Osmolska, Currie and Barsbold, 2004)
Comments- Barsbold created this taxon to separate Oviraptor (to which he referred specimens now distinguished as Conchoraptor and Citipati sp.) from "Ingenia". Once he named Conchoraptor and Rinchenia (originally Oviraptor mongoliensis), these were placed in Oviraptorinae as well. Barsbold et al. (1990) retain this taxonomy, though their cladogram shows oviraptorines to be paraphyletic to "ingeniines", with Conchoraptor closer to "Ingenia" than to Oviraptor (still including Citipati sp.). Similarly, Maryanska et al. (2002) recovered Rinchenia, Citipati and Conchoraptor as paraphyletic to "Ingenia". The definition of Osmolska et al. (2004) functions in their phylogeny, where Rinchenia and Citipati form a clade exclusive of "Ingenia", Conchoraptor and Khaan. Oviraptor was not included in the analysis, but was assumed to be part of this clade, perhaps due to its cranial crest. Heyuannia was assigned to Oviraptorinae in their taxon list, but not included in the analysis, nor was the rationale for the assignment discussed. In Lu's (2004) based on Maryanska et al.'s matrix, Shixinggia and all caenagnathoids are oviraptorines. In his tree based on the TWG data, Oviraptorinae contains Avimimus, Shixinggia and all caenagnathoids except Conchoraptor. The situation is similar in Senter's (2007) analysis, where Oviraptor is placed basally as suggested by Clark et al. (2002), and Oviraptorinae thus includes all oviraptorosaurs except Microvenator and Caudipteryx. The exact content of Oviraptorinae cannot be determined from Lu et al.'s (2015) figured phylogeny, as Oviraptor is in a polytomy with Citipati and "ingeniines" (though a posteriori removal of fragmentary taxa in this polytomy, such as Shixinggia, may show hidden resolution). However, it would definitely exclude the oviraptorid Nankangia, Nomingia and Yulong. It's apparent the original concept of Oviraptorinae is probably not monophyletic with respect to "Ingeniinae", and using Osmolska et al.'s definition leads to situations where useful similarity to their or Barsbold's concept of the taxon is lost. A more useful definition would be (Oviraptor philoceratops <- "Ingenia" yanshini).
References- Barsbold, 1976. On a new Late Cretaceous family of small theropods (Oviraptoridae fam. n.) of Mongolia. Doklady Akademia Nauk SSSR. 226, 685-688. [in Russian]
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press, Berkeley. pp. 249-258.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. Southrern Methodist University. unpublished PhD dissertation.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
Persons, Currie and Norell, 2014. Oviraptorosaur tail forms and functions. Acta Palaeontologica Polonica. 59(3), 553-567.

Shanyangosaurus Xue, Zhang and Bi, 1996
S. niupanggouensis Xue, Zhang and Bi, 1996
Middle-Late Maastrichtian, Late Cretaceous
Shanyang Formation, Shaanxi, China

Holotype- (NWUV 1111) (~1.7 m) uncinate processes(?), partial synsacrum (centra 32 mm), proximal scapula, humeri (116 mm), femur (258 mm), tibia (327 mm), metatarsals IV (137 mm; one proximal), partial phalanx, pedal ungual
Comments- Xue et al. (1996) report "ribs with horizontal hooks", but no ribs are mentioned in the material list, nor are any shown in the plates. They would support a pennaraptoran identity if present however.
While Shanyangosaurus was originally identified only to the level of Theropoda, it is most similar to oviraptorids (Mortimer, DML 2000; followed by Holtz et al., 2004) in the low acromion, absent fourth trochanter, cnemial crest shape, unfused metatarsus whose fourth metatarsal is wider than deep, and an elevated femoral head.
References- Xue, Zhang, Bi, Yue and Chen, 1996. The development and environmental changes of the intermontane basins in the Eastern part of Qinling Mountains. Geological Publishing House, Beijing. ISBN 7-116-02125-6. 179 pages.
Mortimer, DML 2000. http://dml.cmnh.org/2000Sep/msg00125.html
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second Edition. University of California Press. 71-110.

unnamed possible Oviraptoridae (Riabinin, 1938)
Santonian, Late Cretaceous
Syuk Syuk Formation, Kazakhstan

Material- unguals
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.
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.

unnamed oviraptorid (Osmolska, 1976)
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia

Material- (ZPAL MgD-I/96) fragmentary skull
Comments- Osmolska (1976) mentioned a fragmentary skull from the Nemegt Formation of Mongolia housed in the ZPAL. She referred it to Oviraptor sp. along with (?)Conchoraptor specimen ZPAL MgD-I/95 and another specimen. Maryanska and Osmolska (1997) later described a fragmentary skull from the same locality, referring to it as ZPAL MgD-I/96. The identical locality and preserved material, along with the closeness in specimen number to ZPAL MgD-I/95 lead me to believe this is the same specimen Osmolska mentioned. It is described as crested, so may belong to Rinchenia or Nemegtomaia, both also from the Nemegt Formation.
References- Osmolska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Maryanska and Osmolska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.

unnamed Oviraptoridae (Erben, 1995)
Late Cretaceous
Hugang Formation, Henan, China
Material
- (STIPB coll.) eggshells
Comments- Wiemann et al. (2015) report several eggshells they refer to Macroolithus yaotuensis and Heyuannia, but the latter is based on the flawed reasoning of Cheng et al.. The latter assigned shells to Heyuannia "or an oviraptorosaurian of similar kind" based on its subarctometatarsaly. Yet this is true in most other oviraptorids as well except for "Ingenia".
References- Erben, 1995. The Cretaceous/Tertiary boundary in the Nanxiong-Basin (continental facies, SE-China). Stuttgart: Franz Steiner Verlag. 245 pp.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. https://dx.doi.org/10.7287/peerj.preprints.1080v1

unnamed Oviraptoridae (Dong and Currie, 1996)
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia

Material- (IVPP V9608) vertebral fragments, proximal scapula, partial furcula, humerus (168 mm), incomplete radius, incomplete ulna, metacarpal I (32 mm), phalanx I-1 (79 mm), partial manual ungual I, metacarpal II (82+ mm), phalanx II-1 (60 mm), phalanx II-2 (66 mm), manual ungual II, incomplete metacarpal III, phalanx III-1 (38 mm), phalanx III-2 (35 mm), phalanx III-3 (40 mm), manual ungual III, partial femur, partial tibia, phalanx II-2 (33 mm), pedal ungual II (28 mm), phalanx III-1, phalanx III-2 (36 mm), phalanx III-3 (28 mm), pedal ungual III, phalanx IV-1 (30 mm), phalanx IV-2 (27 mm), six eggs, egg fragments
two skeletons (Dong, 1992)
Comments- IVPP V9608 was referred to Oviraptor philoceratops by Dong and Currie (1996), but may be Citipati instead. Further study is necessary, especially of manual proportions. Longrich et al. (2010) referred it to Oviraptorinae indet.. Dong (1992) earlier referred to three partial Oviraptor philoceratops skeletons from the Djadochta Formation, the other two of which are undescribed.
Reference- Dong, 1992. Dinosaurian Faunas of China. China Ocean Press, Beijing.
Dong and Currie, 1995. On the discovery of an oviraptorid skeleton on a nest of eggs. Journal of Vertebrate Paleontology. 15(3), 26A.
Dong and Currie, 1996. On the discovery of an oviraptorid skeleton on a nest of eggs at Bayan Mandahu, Inner Mongolia, People's Republic of China. Canadian Journal of Earth Sciences. 33, 631-636.
Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5), 945-960.

unnamed Oviraptoridae (Maryanska and Osmolska, 1997)
Late Campanian, Late Cretaceous
Red Beds of Khermeen Tsav, Mongolia

Material- (AMNH coll.) skull (Norell, 1992)
(IGM 100/30A) fragmentary skull (Maryanska and Osmolska, 1997)
(IGM coll.; "GIN A") quadrates (Maryanska and Osmolska, 1997)
(IGM coll.; "GIN B") partial quadratojugal, distal quadrate (Maryanska and Osmolska, 1997)
Comments- The skull preliminarily reported by Clark (1992) as Oviraptor sp. may be the Citipati holotype (which was at the AMNH for a time), but the locality is different. This may have been due to incomplete stratigraphic work at Ukhaa Tolgod at the time however. The IGM material represents crestless oviraptorids, perhaps Conchoraptor or "Ingenia".
References- Clark, 1992. The Mongolian-American Museum Expeditions to the Gobi Desert, 1990-1992. Journal of Vertebrate Paleontology. 12(3), 24A.
Maryanska and Osmolska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.

undescribed oviraptorid (Kirkland, Hernandez-Rivera, Aguillon-Martínez, de Jesus, Gomez-Nunez and Vallejo, 2000)
Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico

Reference- Kirkland, Hernandez-Rivera, Aguillon-Martínez, de Jesus, Gomez-Nunez and Vallejo, 2000. The Late Cretaceous Difunta Group of the Parras Basin, Coahuila, Mexico and its vertebrate fauna. Universidad Autónoma del Estado de Hidalgo, Avances en Investigación. 3, 133-172.

undescribed oviraptorid (Norton, DML 2000)
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia

Material- (IGM 100/1009) (juvenile) skull
Comments- Norton (DML, 2000) noted this specimen was discovered in 1993 and on display at the AMNH Fighting Dinosaurs exhibit. It is probably Citipati or Khaan based on the fact it was discovered at Ukhaa Tolgod.
Reference- http://dml.cmnh.org/2000Jun/msg00082.html

undescribed oviraptorid (Lee, Barsbold, Jacobs and Currie, 2008)
Cenomanian-Early Maastrichtian, Late Cretaceous
Baynshiren, Baruungoyot or Nemegt Formation, Mongolia

Material- embryos, nest
Comments- Listed as 'oviraptorosaurid'.
Reference- Lee, Barsbold, Jacobs and Currie, 2008. A short report of Korea-Mongolia International Dinosaur Project (1st and 2nd year). Journal of Vertebrate Paleontology. 28(3), 104A-105A.

undescribed Oviraptoridae (Weishampel, Fastovsky, Watabe, Barsbold and Tsogtbaatar, 2000)
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Material
- (IGM 100/1017) (embryo) femur, fragmentary bones, partial egg (Weishampel et al., 2008)
(IGM 100/1018) (embryo) partial skull, surangular?, six cervical vertebrae, few cervical ribs, three dorsal vertebrae, dorsal ribs, scapula, coracoid, humerus (17 mm), radius, ulna, metacarpal I, metacarpal II, metacarpal III, ilia, femora (30 mm), tibia (42 mm), distal fibula, astragalus, metatarsal II, metatarsal III, pedal ungual III, metatarsal IV, several pedal phalanges, eggshell (Weishampel et al., 2008)
(IGM 100/1019-1) (embryo) partial skull, surangular or articular, three cervical vertebrae, humerus (22 mm), eggshell (Weishampel et al., 2008)
(IGM 100/1019-2) (embryo) dorsal vertebrae, ilium, femur (39 mm), tibia (55 mm), fibula, astragalus (Weishampel et al., 2008)
(PJC.2001.2) partial skeleton including metatarsal I, phalanx I-1, metatarsals II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-4 (Currie, 2002)
partial skeleton (Currie, 2001)
three dorsal vertebrae, three caudal vertebrae (Badamkhatan, 2008)
elements (Currie et al., 2008)
Comments- These may be Rinchenia or Nemegtomaia based on their provenence. PJC.2001.2 was referred to cf. Ingenia sp. by Currie (2002), though "Ingenia" is seemingly unknown from the Nemegt Formation. Weishampel et al. (2008) described a nest of four embryos which they stated resembled Nemegtomaia slightly more. Currie et al.'s (2008) material is from an Avimimus bonebed.
References- Weishampel, Fastovsky, Watabe, Barsbold and Tsogtbaatar, 2000. New embryonic and hatchling dinosaur remains from the Late Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 20(3), 78A.
Currie, 2001. Nomadic expeditions, Inc. report on fieldwork in Mongolia, September 2000. In: Alberta Palaeontological Society, fifth annual symposium, jointly presented by Alberta Palaeontological Society and Department of Earth Sciences, Mount Royal College, p. 12-16.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. In: Alberta Palaeontological Society, sixth annual symposium, “Fossils 2002’, presented by Alberta Paleontological Society, in conjunction with Canadian Society of Petroleum Geologists, Paleontological Division and Department of Earth Sciences, Mount Royal College, p. 8-12.
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.
Weishampel, Fastovsky, Watabe, Varricchio, Jackson, Tsogtbaatar and Barsbold, 2008. New oviraptorid embryos from Bugin-Tsav, Nemegt Formation (Upper Cretaceous), Mongolia, with insights into their habitat and growth. Journal of Vertebrate Paleontology. 28(4), 1110-1119.

unnamed oviraptorid (Lu, 2004)
Maastrichtian, Late Cretaceous
Pingling Formation, Guangdong, China
Material
- (E-1) partial skull, mandible (150 mm)
....(K2-1) two posterior cervical vertebrae (30 mm), nine dorsal vertebrae (20 mm), dorsal rib fragment
....(K2-12) incomplete pes
Comments- These specimens probably belong to the same individual, called NXMV by Lu (2004). Wang et al. (2013) believes it may belong to Ganzhousaurus.
References- Lu, 2004. Oviraptorid dinosaurs from Southern China. Southrern Methodist University. unpublished PhD dissertation.
Lu, 2005. Oviraptorid dinosaurs from Southern China. Geological Publishing House, Beijing. ISBN 7-116-04368-3. 200 pages + 8 plates.
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.

unnamed Oviraptoridae (Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005)
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China
Material
- (Chimei Museum 41) (embryo) centra, distal femora, tibia, phalanges, unguals, fragments, partial egg (169x83.8 mm) (Cheng, Ji, Wu and Shan, 2008)
(IVPP V20182) (embryo) partial skull (46.8 mm), partial mandible (38.7 mm), penultimate and last sacral vertebrae, sacral ribs, first to fourth caudal vertebrae, fifth caudal neural arch, proximal metatarsal II, proximal metatarsal III, proximal metatarsal IV, egg (198.3x88 mm) (Wang, Zhang, Sullivan and Xu, 2016)
(IVPP V20183) (embryo) incomplete nasal, lacrimal, frontal, anterior dentaries, three posterior sacral centra, mid sacral rib, first to sixth caudal centra, incomplete ilium (45.5 mm), incomplete pubes, partial ischium, femur (49.2 mm), tibia (56.2 mm), incomplete metatarsal II, phalanges II-?, incomplete metatarsal III, phalanges III-?, egg (163.5x74.8 mm) (Wang, Zhang, Sullivan and Xu, 2016)
(IVPP V20184) (embryo) partial parietal, incomplete squamosal, quadratojugal, quadrate, cervical centrum, two cervical neural arches, few dorsal centra, few fragmentary dorsal ribs, furcula, (?)femur, egg (179.5x92.1 mm) (Wang, Zhang, Sullivan and Xu, 2016)
(NMNS-0015726-F02-embryo-01) (embryo) centra, neural arches, partial femora, tibiae (one incomplete; one partial), incomplete metatarsal II (~29.6 mm), metatarsal III (~32 mm), phalanx III-1, metatarsal IV (22.5 mm), phalanx IV-1, fragments, incomplete egg (173.5x76.3 mm) (Cheng, Ji, Wu and Shan, 2008)
(NMNS-CYN-2004-DINO-05/I) egg (Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015)
(NMNS-VPDINO-2002-0901) six sacral vertebrae, first caudal vertebra, second caudal vertebra, incomplete ilia, proximal pubes, ischia, proximal femora, incomplete tibia, incomplete fibula, tarsus, two partial metatarsals, two eggs (175 mm) (Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005)
gastralia, posterior sacrum, over twenty-six caudal vertebrae, pelvis including ischia, hindlimb including femur, tibia and metatarsals, two eggs (193 mm) (He, Varricchio, Jackson, Jin and Pust, 2012)
specimen including skull and mandible (Lu, Chen, Kobayshi and Lee, 2015)
Comments- The eggs are preserved inside the pelvic cavity of NMNS-VPDINO-2002-0901 and resemble Macroolithus yaotunensis and M. rugustus (Sato et al., 2005).
Cheng et al. (2008) found Chimei Museum 41 and NMNS-0015726-F02-embryo-01 were assignable to Macroolithus yaotunensis. They assigned these to Heyuannia "or an oviraptorosaurian of similar kind" based on its subarctometatarsaly. Yet this is true in most other oviraptorids as well except for "Ingenia", including potentially Banji, Jiangxisaurus and Nankangia from the same formation.
He et al. (2012) remark on an additional specimen with two internal eggs, assignable only to Elongatoolithidae.
Wang et al. (2013) note embryos from the same formation inside of Macroolithus yaotunensis eggs. These are detailed in Wang et al. (2016), which only identifies the embryos as Oviraptoridae.
Lu et al. (2015) report a new taxon, diagnosed by- external naris above line running from quadratic and premaxillary ventral edges (also in Rinchenia and Nemegtomaia); dome-like skull roof; dentary symphysis not decurved (also in Nankangia).
Wiemann et al. (2015) report several eggshells they refer to Macroolithus yaotuensis and Heyuannia, but the latter is based on the flawed reasoning of Cheng et al..
These remains may belong to the contemporaneous Banji, Jiangxisaurus, Ganzhousaurus and/or Nankangia.
References- Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005. A pair of shelled eggs inside a female dinosaur. Science. 308, 375.
Cheng, Ji, Wu and Shan, 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic skeletons discovered for the first time in China. Acta Geologica Sinica. 82(6), 1089-1094.
He, Varricchio, Jackson, Jin and Pust, 2012. An oviraptorid adult-egg association and the origin of avialan reproductive strategies. Journal of Vertebrate Paleontology. Program and Abstracts 2012, 108.
Wang, Zhang and Xu, 2013. New oviraptorid (Theropoda, Oviraptorosauria) embryos from the Upper Cretaceous of Southern China. Journal of Vertebrate Paleontology. Program and Abstracts 2013, 234.
Lu, Chen, Kobayshi and Lee, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 168.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. | https://dx.doi.org/10.7287/peerj.preprints.1080v1
Wang, Zhang, Sullivan and Xu, 2016. Elongatoolithid eggs containing oviraptorid (Theropoda, Oviraptorosauria) embryos from the Upper Cretaceous of southern China. BMC Evolutionary Biology. 16:67.

undescribed oviraptorid (Jensen, 2008)
Late Campanian, Late Cretaceous
Red Beds of Khermeen Tsav, Mongolia
Material
- (private coll.; PMO X678 cast; UALVP 49394 cast) skull (112 mm), mandibles (99.8 mm), hyoids, eight partial cervical vertebrae, dorsal rib fragments (Knutsen, 2008)
(UALVL 49393 cast) skull, sclerotic ring, mandibles (139 mm), hyoids, phalanx I-1 (57.5 mm), manual ungual I (47 mm), incomplete metacarpal II, phalanx II-1 (56 mm), phalanx II-2 (55 mm), manual ungual II (47 mm), incomplete metacarpal III, phalanx III-1 (38 mm), phalanx III-2, phalanx III-3, manual ungual III (Fanti et al., 2012)
Comments- This taxon is known from two skulls with similar high, pointed crests. Casts are common, and at least one is definitely privately owned according to www.prehistoricstore.com. The latter site also lists it as being from Khermeen Tsav. This specimen was analyzed by Jensen (2008), who called it Oviraptor sp.. The other specimen includes a manus preserved next to the skull and is listed as Oviraptor philoceratops on the Witmer Lab website. Fanti et al. (2012) used UALVP casts to include measurements of both specimens in their dataset as "oviraptorid incertae sedis".
References- Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
http://www.oucom.ohiou.edu/dbms-witmer/collections/Theropods/oviraptor.htm

unnamed Oviraptoridae (Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015)
Maastrichtian, Late Cretaceous
Yuanpu or Pingling Formations, Guangdong, China
Material
- eggshells
Comments- Wiemann et al. (2015) report several eggshells they refer to Macroolithus yaotuensis and Heyuannia, but the latter is based on the flawed reasoning of Cheng et al. (2008).
References- Cheng, Ji, Wu and Shan, 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic skeletons discovered for the first time in China. Acta Geologica Sinica. 82(6), 1089-1094.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. | https://dx.doi.org/10.7287/peerj.preprints.1080v1

undescribed Oviraptoridae (Fanti et al., 2012)
Late Cretaceous(?)
Asia(?)
Material
- (IGM coll.) specimen including skull (147.3 mm), mandible (120 mm), metacarpal I (24 mm), phalanx I-1 (30.1 mm), manual ungual I (31.7 mm), phalanx II-2 (17.8 mm), manual ungual II (81.8 mm), femur (238 mm), tibia (275 mm), phalanx III-3 (17 mm), phalanx IV-2 (13.3 mm), phalanx IV-3 (10 mm), phalanx IV-4 (9.3 mm), pedal ungual IV (30 mm) (Fanti et al., 2012)
(IGM coll.) specimen including skull (162 mm), humerus (113 mm), radius (90 mm), ulna (~98 mm), metacarpal I (22.4 mm), phalanx I-1 (29.2 mm), manual ungual I (35.2 mm), metacarpal II (40.6 mm), phalanx II-1 (19.6 mm), phalanx II-2 (15 mm), manual ungual II (18 mm), femur (220 mm), tibia (265 mm), phalanx III-3 (15.8 mm), phalanx IV-2 (13.6 mm), phalanx IV-3 (9.9 mm), phalanx IV-4 (8.9 mm), pedal ungual IV (27.3 mm) (Fanti et al., 2012)
skull, mandible, five anterior cervical vertebrae (online)
skull, mandibles (online)
Comments- Fanti et al. (2012) listed measurements for two "oviraptorid incertae sedis" specimens at the IGM. Analyzing their proportions may enable identification. Numerous unpublished oviraptorid specimens are known and have been photographed online. Most are referred to Oviraptor, Conchoraptor or "Ingenia", but this is generally based on the pre-1980's convention of calling all oviraptorids Oviraptor or the pre-2000 convention of calling all crestless oviraptorids Conchoraptor or "Ingenia". The more complete skeletons, especially when mounted, often contain faked portions.
References- Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.

Banji Xu and Han, 2010
B. long Xu and Han, 2010
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China
Holotype-
(IVPP V16896) (juvenile) skull (65 mm), mandibles (one partial, one incomplete)
Diagnosis- (after Xu and Han, 2010) premaxillonasal crest with stepped posterior end; premaxillonasal crest with two longitudinal grooves and numerous oblique striations on lateral surface; elongate external naris placed posteriorly; deep fossa on dorsal surface of palatal process of pterygoid; several longitudinal grooves on the posterodorsal dentary; several tubercles on the dorsal surangular shelf.
Comments- Xu and Han (2010) initially recovered Banji as a basal oviraptorid based on a Maryanska et al. analysis, a position mirrored by the latest and largest oviraptorosaur analysis (Lu et al., 2015).
Reference- Xu and Han, 2010. A new oviraptorid dinosaur (Theropoda: Oviraptorosauria) from the Upper Cretaceous of China. Vertebrata PalAsiatica. 48(1), 11-18.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Citipati Clark, Norell and Barsbold, 2001
Diagnosis- (after Clark et al., 2002) accessory opening on the lateral surface of the ascending process of the premaxilla.
C. osmolskae Clark, Norell and Barsbold, 2001
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia

Holotype- (IGM 100/978) (~105 kg) complete skeleton including skull (160 mm), sclerotic ossicles, mandibles (151 mm), hyoids, twelve cervical vertebrae, cervical ribs, seven dorsal vertebrae, thirty caudal vertebrae (first caudal 38.2 mm), pygostyle, chevrons, scapulocoracoid, furcula, sternal plates, humerus (230 mm), radius (189 mm), ulna (200 mm), femur (345 mm), tibia (397 mm), astragalus, metatarsal I (37 mm), tarsometatarsus (II 176 mm; III 192 mm; IV 188 mm)
Paratypes- (IGM 100/971) (embryo) partial skull (~40 mm), mandibles, cervical vertebra, three fragmentary dorsal vertebrae, several dorsal rib fragments, three sacral centra, few distal caudal vertebrae, scapulae, coracoids, furcula, humeri (20 mm), proximal radius, proximal ulna, partial ilia, femur (36 mm), tibia (50 mm), fibula, astragali, partial metatarsal II, partial metatarsal III, partial metatarsal IV, pedal phalanges, nest, eggshell fragments (Norell et al., 1994)
?(IGM 100/979; Big Mama) distal cervical rib, anterior dorsal vertebra, distal dorsal ribs 1-6, four uncinate processes, gastralia, incomplete furcula, sternal plates, four sternal ribs, partial scapulae, coracoids, humeri (215 mm; one proximal), radii (198 mm; one distal), ulnae (214 mm; one distal), proximal carpal, semilunate carpal, metacarpal I (~45 mm), phalanx I-1 (88.4 mm), manual ungual I (91.1 mm), metacarpal II (89.5 mm), phalanx II-1 (65.5 mm), phalanx II-2 (61.5 mm), manual ungual II (76 mm), metacarpal III (89.4 mm), phalanx III-1 (33.8 mm), phalanx III-2 (~31 mm), phalanx III-3 (~45 mm), manual ungual III, distal pubes, distal ischia, distal femur, incomplete tibia, fibulae (~415 mm), astragalus, calcaneum, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II (~173 mm), phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (~190 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (~177 mm), phalanx IV-1, phalanx IV-2 (24.9 mm), phalanx IV-3 (20.1 mm), phalanx IV-4 (19 mm), pedal ungual IV (48.4 mm), nest, fifteen eggs (Clark et al., 1999)
Referred- ?(IGM 100/1004; Big Auntie) twelve cervical vertebrae, cervical ribs, several dorsal vertebrae, eight dorsal ribs, four uncinate processes, partial sacrum, scapulae, partial coracoid, furcula, sternal fragment, three sternal ribs, humeri, radii, ulnae, phalanx I-1, manual ungual I, phalanx II-1, phalanx II-2, manual ungual II, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, fragmentary ilium, proximal pubis, proximal ischium, partial femora (405 mm), incomplete tibia, incomplete fibula, metatarsal I, phalanx I-1, pedal ungual I, distal metatarsal II, distal metatarsal III, distal metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, nest, seven eggs (Webster, 1996)
?(IGM 100/1125) two eggs (Grellet-Tinner, 2005)
Diagnosis- (after Clark et al., 2002) anterodorsally sloping occiput and quadrate; parietal much longer along the midline than the frontal and reaching nearly to the level of the anterior end of the orbit; ascending process of jugal perpendicular to the horizontal ramus (rather than extending posterodorsally); external naris nearly circular; nasal process of premaxilla vertical rather than sloping posterodorsally; cervical vertebrae elongate (approximately twice as long as they are wide).
Comments- The first published specimen of this species was an embryo discovered in 1993 in a nest of eggs (Norell et al., 1994), later described in detail by Norell et al. (2001). Originally assigned to Oviraptoridae indet., it was tentatively referred to the then unnamed Citipati osmolskae by Norell et al. (2001) and formally referred to that taxon by Clark et al. (2001). Two partial juvenile paravian skulls were found with the specimen, originally identified as Velociraptor (Norell et al., 1994) and later troodontids (Norell and Makovicky, 1999).
Norell et al. (1995) announced a brooding oviraptorid on a nest found in 1993, which was initially referred to Oviraptor based on manual resemblences. It was mentioned in the popular press as "Big Mama". The specimen was later described in detail (Clark et al., 1999) and said to be an oviraptorid most closely related to Oviraptor, before being officially referred to Citipati osmolskae by Clark et al. (2001).
A second brooding Citipati specimen found in 1995 is known as "Big Auntie" (Clark pers. comm. to Auditore). It was mentioned by Webster (1996) and Clark et al. (1999). A photograph of this specimen is present in Codd (2004), incorrectly identified as IGM 100/42. Similarly, it is photographed and incorrectly identified as Oviraptor philoceratops holotype AMNH 6517 by Codd et al. (2007). Erickson et al. (2009) examined it histologically.
Grellet-Tinner (2005) described the eggs in detail in his thesis.
The holotype of the genus, a virtually complete specimen, was initially identified as Oviraptor philoceratops (Webster, 1996) until it was preliminarily described and officially named by Clark et al. (2001). The skull and mandibles were later described in detail by Clark et al. (2002), though the postcrania remains largely undescribed and unillustrated, with only Nesbitt et al. (2009) and Balanoff and Norell (2012) showing the furcula and describing some details. More recently, Persons et al. (2014) describe the tail and pygostyle.
References- 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.
Clark, 1995. The egg thief exonerated. Natural History. 6/95, 56-56.
Norell, Clark, Chiappe, and Dashzeveg, 1995. A nesting dinosaur. Nature. 378, 774-776.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Norell and Clark, 1997. Birds are dinosaurs. Sci. Spectrum. 8, 28-34.
Clark, Norell and Chiappe, 1998. A "brooding" oviraptorid from the Late Cretaceous of Mongolia and its avian characters. JVP 18(3) 34A.
Clark, Norell and Chiappe, 1999. An oviraptorid skeleton from the Late Cretaceous of Ukhaa Tolgod, Mongolia, preserved in an avianlike brooding position over an oviraptorid nest. American Museum Novitates. 3265, 1-36.
Norell and Makovicky, 1999. Important features of the dromaeosaur skeleton II: information from newly collected specimens of Velociraptor mongoliensis. American Museum Novitates. 3282, 1-45
Norell, Clark and Chiappe, 2001. An embryonic oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Mongolia. American Museum Novitates. 3315 1-17.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Codd, 2004. The uncinate processes in birds and their implications for the breathing mechanics of maniraptoran dinosaurs. Dissertation zur Erlangung des Doktotgrades der Mathematisch-Naturwissenschaftlichen Fakultat der Rheinischen Friedrich-Wilhelms-Universitat Bonn. 108 pp.
Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case study of saurischian dinosaur relationships and avian evolution. PhD thesis, University of Southern California. 221 pp.
Grellet-Tinner, Chiappe, Norell and Bottjer, 2006. Dinosaur eggs and nesting behaviors: A paleobiological investigation. Palaegeography, Palaeoclimatology, Palaeoecology. 232, 294-321.
Codd, Manning, Norell and Perry, 2007. Avian-like breathing mechanics in maniraptoran dinosaurs. Proceedings of the Royal Society B. 275(1631), 157-161.
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.
Nesbitt, Turner, Spaulding, Conrad and Norell, 2009. The theropod furcula. Journal of Morphology. 270, 856-879.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.
Pittman and Mallison, 2014. Tail function in oviraptorosaur dinosaurs: Insights from a 3D tail model of Citipati osmolskae (Theropoda: Oviraptorosauria). Journal of Vertebrate Paleontology. Program and Abstracts 2014, 205-106.
Moyer, Zheng, Norell and Schweitzer, 2015. Microscopic and immunohistochemical analyses of the claw of the nesting dinosaur, Citipati osmolskae. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 186.
Persons, Currie and Norell, 2014. Oviraptorosaur tail forms and functions. Acta Palaeontologica Polonica. 59(3), 553-567.
C. sp. nov. (Barsbold, 1981)
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia

Material- (IGM 100/42; Zamyn Khondt oviraptorid) incomplete skull (180mm), mandibles (153, ~156 mm), fiftteen cervical vertebrae, cervical ribs, seven dorsal vertebrae, three dorsal centra, eighteen dorsal ribs, sacrum, thirty caudal vertebrae (first caudal 32 mm), twenty-three chevrons, scapulacoracoids (scap ~238 mm), furcula, sternal plates (~79, ~84 mm), humeri (205 mm), radius (180 mm), ulna (188 mm), semilunate carpal, metacarpal I (47 mm), phalanx I-1 (95 mm), manual ungual I (73 mm), metacarpal II (104 mm), phalanx II-1 (58 mm), phalanx II-2 (73 mm), manual ungual II (64 mm), metacarpal III (98 mm), phalanx III-1 (42 mm), phalanx III-2 (40 mm), phalanx III-3 (56 mm), manual ungual III (54 mm), ilia (290 mm), pubes (320 mm), ischia, femora (305 mm), tibiae (380 mm), fibula, astragalus, calcaneum, metatarsal I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (168 mm), phalanx III-1 (39 mm), phalanx III-2, phalanx III-3 (31 mm), pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2 (24 mm), phalanx IV-3 (30.5 mm), phalanx IV-4 (26 mm), pedal ungual IV (45 mm), metatarsal V
Diagnosis- (after Clark et al., 2001) anterior edge of the premaxilla is concave in lateral view; vomers unfused; cervical vertebrae are not elongate.
Comments- This specimen was originally identified as Oviraptor philoceratops (Barsbold, 1981), an identification which persisted unquestioned through the 80's and 90's (e.g. Barsbold, 1983, 1986; Barsbold et al., 1990; Maryanska et al., 2002). The completeness relative to the O. philoceratops holotype resulted in IGM 100/42 becoming the standard example of the species. In 2001, Clark et al. realized the specimen was more similar to the recently named Citipati osmolskae, and may represent another species of that genus. Clark et al. (2002) reiterated this, while Osmolska et al. (2004) found it to be the sister taxon of Citipati osmolskae in their cladistic analysis. However, Lu (2004) found that IGM 100/42 was the sister taxon to Oviraptor or Conchoraptor (depending on taxa included) using Maryanska et al.'s (2002) characters. This and the resemblence of the more recently described Nemegtomaia (which was more closely related to Citipati than to IGM 100/42 or Oviraptor in Lu's analysis) suggests we use caution when assigning IGM 100/42 to a particular genus.
The specimen has never been described in detail, though many elements have been illustrated in varied sources. Barsbold (1981) illustrated the skull, mandible, furcula, sternal plates and manual phalanges. He later (1983) illustrated the skull in ventral view, mandible in medial view and scapulocoracoid. Barsbold et al. (1990) illustrated the skull in dorsal view, humerus, complete manus, and metatarsus. Jensen (2008) analyzed the skull for his thesis and presented detailed photographs. The almost complete skeleton is mounted, but publically available photos are too small to be useful. The data matrices of Norell et al. (2001), Maryanska et al. (2002) and Osmolska et al. (2004) are the most useful published sources of information besides the figures. Fanti et al. (2012) and Currie et al. (2016) provide measurements.
References- Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120. [in Russian]
Barsbold, 1986. Raubdinosaurier Oviraptoren. In Vorobyeva (editor). Herpetologische Untersuchungen in der Mongolischen Volksrepublik. 210-223. Akademia Nauk SSSR Institut Evolyucionnoy Morfologii i Ekologii Zhivotnikhim. Moskva: A.M. Severtsova. [in Russian, German summary]
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press, Berkeley. pp. 249-258.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Norell, Clark and Makovicky, 2001. Relationships among Maniraptora: problems and prospects. 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. New Haven: Peabody Mus. Nat. Hist., Yale Univ. pp. 49-67.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. Southrern Methodist University. unpublished PhD dissertation.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Currie, Funston and Osmólska, 2016. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.

Huanansaurus Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015
H. ganzhouensis Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China

Holotype- (HGM41HIII-0443) incomplete skull (206.1 mm), mandible (182.8 mm), atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, incomplete sixth cervical vertebra, partial seventh cervical vertebra, gastralia, incomplete humerus (195 mm), fragmentary radius (190 mm), fragmentary ulna, fragmentary carpals, fragmentary metacarpal I (40 mm), phalanx I-1 (80 mm), incomplete manual unguals I (70 mm), incomplete metacarpal II (85 mm), phalanges II-1 (one partial; 55 mm), phalanges II-2 (65 mm), manual unguals II (65 mm), incomplete metacarpal III (84 mm), phalanx III-1 (32 mm), phalanx III-2 (30 mm), phalanges III-3 (44 mm), manual unguals III (52 mm), distal femur, proximal tibia, incomplete metatarsal II, phalanx II-1 (45 mm), phalanx II-2 (35 mm), pedal ungual II (55 mm), incomplete metatarsal III, phalanx III-1 (45 mm), phalanx III-2 (39 mm), phalanx III-3 (35 mm), pedal ungual III (50 mm), incomplete metatarsal IV, phalanx IV-1 (33 mm), phalanx IV-2 (30 mm), phalanx IV-3 (21 mm), phalanx IV-4 (20 mm), pedal ungual IV (50 mm)
Diagnosis- (after Lu et al., 2015) posterodorsal process of the premaxillae contact lacrimals (also in Citipati and Conchoraptor); distinct opening near posteroventral corner of distal end of posterodorsal premaxillary process; circular supratemporal fenestra much smaller than laterotemporal fenestra; nuchal crest pronounced; mandibular condyles of quadrate posterior to occipital condyle; pneumatized dentaries; anterodorsal tip of dentary projecting anterodorsally at an angle of 45 degrees or less relative to the ventral margin of symphysis; length of dentary symphysis between 20% and 25% of mandible length; dentary portion of dorsal external mandibular fenestra margin strongly concave; posteroventral dentary process twisted so that lateral surface faces somewhat ventrally; angular contributes extensively to border of external mandibular fenestra; metacarpal I long and slender, diameter 20% of length; proximodorsal lip on all manual unguals prominent.
Comments- Lu et al. (2014) announced this specimen in an abstract as a sister taxon to Citipati. It was fully described and named by Lu et al. (2015), and emerged in a trichotomy with Citipati osmolskae and IGM 100/42 in their analysis derived from Maryanska et al.'s oviraptorosaur matrix.
References- Lu, Kobayashi, Pu, Chang, Zhang, Shang and Liu, 2014. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleogeographical implications. Journal of Vertebrate Paleontology. Program and Abstracts 2014, 164.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Nankangia Lu, Yi, Zhong and Wei, 2013
N. jiangxiensis Lu, Yi, Zhong and Wei, 2013
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China

Holotype- (GMNH F10003) anterior mandibles, five mid dorsal vertebrae (38, 37, 38, 39, 40 mm), partial penultimate sacral vertebra, last sacral vertebra (43 mm), three proximal caudal vertebrae (38, 44, 42 mm), proximal caudal neural arch, six mid caudal centra (41, 39, 38, 41, 38, 37 mm), incomplete mid caudal vertebra, two mid chevrons, scapulocoracoids (scapula 260 mm), furcular fragments, incomplete humerus (240 mm), ilia (one incomplete; 330 mm), pubes (one incomplete; 380 mm), ischia (205 mm), femora (350, 380 mm), tibia (400 mm), fibular fragment, astragalus, calcaneum
Diagnosis- (after Lu et al., 2013) rostral end of mandibular symphyseal region not downturned (also in caenagnathids, Incisivosaurus, Luoyanggia and Ganzhousaurus); two infradiapophyseal fossae on ventral surface near base of transverse process of dorsal vertebrae; pneumatic fossae on sacral vertebrae slit-like; neural spines of proximal caudal vertebrae wider transversely than anteroposteriorly, forming a large posterior fossa with rugose central area; large fossa on anterior surface (infraprezygapophyseal fossa) and another (infradiapophyseal fossa) on ventral surface of base of transverse process of the proximal caudals; femur longer than ilium (also in Yulong and Khaan); ratio of height to length of ilium 0.36; femoral neck extending dorsomedially at about an angle of 90 degrees to the shaft; femur and tibia approximately the same length.
Comments- The holotype was donated to the GMNH by a farmer in 2010. Lu et al. (2013) added Nankangia to a version of the Maryanska et al. analysis and found it to be a basal oviraptorid, a similar position to that in the latest and most extensive oviraptorosaur analysis (Lu et al., 2015).
References- Lu, Yi, Zhong and Wei, 2013. A new oviraptorosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleoecological implications. PLoS ONE. 8(11), e80557.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Nomingia Barsbold, Osmolska, Watabe, Currie and Tsogtbaatar, 2000
N. gobiensis Barsbold, Osmolska, Watabe, Currie and Tsogtbaatar, 2000
= "Nomingia brevicaudia" Skrepnick, DML 2000
Early Maastrichtian, Late Cretaceous
Nemegt Svita (=Beds of Bugeen Tsav), Mongolia

Holotype- (IGM 100/119; incorrectly listed as GIN 940824 by Barsbold et al. 2000) (1.8 m; ~44 kg) three cervical vertebrae, ten dorsal vertebrae, ten fragmentary dorsal ribs, several gastralia, five sacral vertebrae, nineteen caudal vertebrae (460 mm; first caudal 28.3 mm), pygostyle (56 mm), fifteen chevrons, ilia (252 mm), pubes (243 mm), ischia (145 mm), femur (285 mm), tibiae (355 mm), fibulae
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia

Referred- (IGM 102/5; = PJC.2001.5) pygostyle (Currie, 2002)
Diagnosis- seven sacral vertebrae sutured/fused together; pleurocoels in proximal caudal vertebrae; last five vertebrae fused into pygostyle; mid-dorsal ischial process(?); prominent tibiofibular crest on distal femur.
Description- The specimen was a subadult based on the unfused sutures between presacral neural arches and centra. Based on comparison to oviraptorids and adjusting for the shorter tail, it may have been about 1.8 meters long.
The first three preserved vertebrae are cervicals. They have a single pair of pleurocoels and low neural spines, although not as low as Microvenator. The last has a moderate-sized epipophysis overhanging the postzygopophysis, which are much shorter than Microvenator. The second to last cervical centrum has a ventral ridge, while the last has a large hypapophysis. Cervical ribs were not fused to centra.
The ten dorsal vertebrae also have pairs of pleurocoels and neural spines grading from moderate to tall (about twice as tall as centra). The first two dorsals have large hypapophyses, the next next five have ventral ridges and the last three are flat ventrally. The centra are short compared to Microvenator. The ninth dorsal vertebra has a bipartite transverse process; one part contacts the rib tuberculum, the other contacts the ilium's preacetabular process. The tenth dorsal vertebra is sutured to the sacrum and the neural spine contacts the first sacral neural spine. The transverse process does not contact the ilium however. The ribs and gastralia are not described or illustrated.
Five sacral vertebrae are fused and have transverse processes in contact with the ilia. Their neural spines are in contact and project slightly above the ilium. Ventrally, the centra are subequal in width and the second through fourth centra are grooved.
Twenty-four caudal vertebrae are present, the last five fused into a pygostyle. The first caudal vertebra is sutured to the sacrum. The centra gradually decrease in length, until the last ten before the pygostyle are 60-65% the length of the first. Pleurocoels are present in the first ten centra. The first two and the eleventh and twelfth centra are flat ventrally, but the others have a pair of ridges bounding a median groove. Sixteen caudals have neural spines and eighteen have transverse processes. The fifteenth through nineteenth have elongate prezygopophyses, though not comparable to dromaeosaurs. The postzygopophyses join to form a continuous midline crest from the fourteenth onward and the prezygopophyses join it after the eighteenth caudal. The last five caudals are fused indistinguishably. Seventeen caudals have chevrons (only fifteen are preserved), ending after the eighteenth. The first fourteen chevrons are dorsoventrally elongate, none are distally expanded. Instead, the last several arectangular, while the proximal six are tapered distally.
The pelvis is slightly propubic (~20 degrees to the vertical). The ilium has an expanded preacetabular process, with a blunt anteroventral corner. The dorsal margin is convex until the acetabular midpoint, where it becomes straight. It is then angled posteroventrally to take part in the rounded postacetabular process. The preacetabular process is about 15% longer than the postacetabular process. The pubic peduncle is vertically oriented, extends ventrally as far as the ischial peduncle and has a concave ventral edge. There is a shallow elongate cuppedicus fossa, no supracetabular crest and a short shallow brevis fossa. A low antitrochantor is present on the ischial peduncle. Dorsally, the iliac blades converge medially to contact the sacral neural spines. The pubis is very slightly concave anteriorly and lacks any proximal foramina or processes. The foot is slightly larger anterior than posteriorly and acutely pointed in both directions. The symphysis extends for half of the pubic length. The ischium is 60% of the pubis in length and has a triangular obturator process placed halfway down the shaft. There may be a small mid-dorsal ischial process, although this could just be a broken area.
The femur has a horizontal head that is separated from the greater trochantor. There is a slight neck. The lesser trochantor is either fused to the greater trochantor or separated by a small groove. No fourth trochantor is visible, but a posterior trochantor may be present. A pronounced tibiofibular crest is present. The tibia is similar to Ingenia. The fibula reaches the calcaneum, is weakly concave proximomedially and has a craniolaterally projecting tibiofibularis tubercle. The astragalocalcaneum is unfused and the ascending process is 20% of tibial height.
Comments- The holotype was discovered in 1994, illustrated in Sloan (1999) and described briefly by Barsbold et al. (2000) before its official publication. The species was originally called "Nomingia brevicaudia", but was changed shortly before publication (Skrepnick, DML 2000). The latter species name has never been published however.
Relationships- Barsbold et al. (2000) did not place Nomingia in a family, but think the strong posterior curve to the ischium, long preacetabular process and straight pubis might suggest caenagnathid affinities. Maryanska et al. (2002) found Nomingia to be weakly placed in the Caenagnathidae due to sharing two characters (dorsal margin of the ilium arched along the central portion of the blade; preacetabular process of the ilium longer than the postacetabular process) with Chirostenotes, though they noted Rinchenia has the first character as well. This assignment has been followed by Osmolska et al. (2004) and Lu and Zhang (2005) without comment. Lu (2004) found Nomingia to be outside a clade containing caenagnathoids and Avimimus based on a modified version of the Theropod Working Group matrix, and as a derived oviraptorid sister to Rinchenia in a modified version of Maryanska et al.'s matrix. The latter result was later published by Lu et al. (2004). The most recent and extensive oviraptorosaur analysis (Lu et al., 2015) found Nomingia to be a basal oviraptorid sister to Yulong.
References- Sloan, 1999. Feathers for T. rex? National Geographic. 196(5), 98-107.
Barsbold, Currie, Myhrvold, Osmolska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 6766, 155
Barsbold, Osmolska, Watabe, Currie and Tsogtbaatar, 2000. A new oviraptorosaur (Dinosauria, Theropoda) from Mongolia: The first dinosaur with a pygostyle. Acta Paleontologica Polonica. 45(2), 97-106.
http://dml.cmnh.org/2000Oct/msg00074.html
Currie, 2002. Report on fieldwork in Mongolia, September 2001. in Alberta Palaeontological Society, sixth annual symposium, “Fossils 2002’, presented by Alberta Paleontological Society, in conjunction with Canadian Society of Petroleum Geologists, Paleontological Division and Department of Earth Sciences, Mount Royal College. 8-12.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from Southern China. Southern Methodist University. unpublished PhD dissertation.
Lu, Tomida, Azuma, Dong and Lee, 2004. New Oviraptorid Dinosaur (Dinosauria: Oviraptorosauria) from the Nemegt Formation of Southwestern Mongolia. Bull. Natn. Sci. Mus.. Tokyo, Ser. C. 30, 95-130.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Lu and Zhang, 2005. A new oviraptorid (Theropoda: Oviraptorosauria) from the Upper Cretaceous of the Nanxiong Basin, Guangdong Province of southern China. Acta Palaeontologica Sinica 44(3): 412-422.
Persons, Currie and Norell, 2011. Shake your feathers: The flamboyant, athletic, and possibly flirtatious caudal morphology of oviraptorosaurs. Journal of Vertebrate Paleontology. Program and Abstracts 2011, 174.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.
Persons, Currie and Norell, 2014. Oviraptorosaur tail forms and functions. Acta Palaeontologica Polonica. 59(3), 553-567.

Oviraptor Osborn, 1924b
= "Fenestrosaurus" Osborn, 1924a
O. philoceratops Osborn, 1924b
= "Fenestrosaurus philoceratops" Osborn, 1924a
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia
Holotype
- (AMNH 6517) incomplete skull (~179 mm), mandibles (195 mm), cervical vertebrae 4-12, dorsal vertebrae 1-7, several dorsal ribs, four uncinate processes, scapula (231 mm), partial coracoid, furcula, sternal ribs, humerus (185 mm), radius (163 mm), ulna (147 mm), phalanx I-1 (79 mm), carpometacarpus (107 mm; mcII 95 mm), phalanx II-1 (54 mm), phalanx II-2 (68 mm), manual ungual II (~47 mm), metacarpal III (99 mm), phalanx III-1 (39 mm), phalanx III-2 (37 mm), phalanx III-3 (28 mm), manual ungual III, partial ilium
Referred- (AMNH 6508) ~15 eggs, nest (Osborn, 1924b)
Comments- The first oviraptorid discovered, this taxon was originally referred to the Ornithomimidae by Osborn (1924b). Barsbold (1976) erected Oviraptoridae for the it and referred six individuals including IGM 100/20 and 100/21 to Oviraptor philoceratops. They were later (Barsbold, 1986) made the holotype and paratype of a new genus, Conchoraptor. Osmolska (1976) believed these and additional material (three skulls including ZPAL MgD-I/95 and MgD-I/96 and fragmentary postcrania) belonged to a new species, referring to them as Oviraptor sp.. None of these are referred to Oviraptor currently, with ZPAL MgD-I/95 most often being assigned to Conchoraptor (Maryanska et al., 2002; Kundrat, 2007) or "Ingenia" (Paul, 1998), though it may be Khaan instead. ZPAL MgD-I/96 is crested and from the Nemegt Formation, so may be Rinchenia or Nemegtomaia. Barsbold (1981) referred IGM 100/42 to O. philoceratops, which resulted in this specimen being the standard example of the species through the 80's and 90's (e.g. Barsbold, 1983; Barsbold, 1986; Barsbold et al., 1990; Maryanska et al., 2002). This was finally shown to be incorrect by Clark et al. (2002), who noted a greater resemblence to Citipati osmolskae, and assigned it to that genus. Citipati specimens, including IGM 100/979 (Clark et al., 1995) and the holotype IGM 100/978 (Webster, 1996) were referred to Oviraptor based on cranial and manual resemblences to IGM 100/42 until the distinctness of Citipati was recognized by Clark et al. (2001). In addition, Rinchenia (IGM 100/32) was originally described as Oviraptor mongoliensis (Barsbold, 1986) before being renamed unofficially by Barsbold in 1997 and officially by Osmolska et al. in 2004. Paul (1988) lumped "Ingenia" into Oviraptor as O. yanshini, but this would result in all oviraptorids being Oviraptor in modern phylogenies. Dong and Currie (1996) referred a fragmentary skeleton (IVPP V9608) to Oviraptor philoceratops, but this may be Citipati instead and was referred to Oviraptorinae indet. by Longrich et al. (2010). Morell (1997) labeled IGM 100/1002 Oviraptor before it was made a paratype of Khaan by Clark et al. (2001). Jensen (2008) calls the tall-crested cast PMO X678 Oviraptor sp., which may be the same taxon as a skull and manus on the Witmer Lab website labeled Oviraptor philoceratops, but how taxon this related to Oviraptor requires further study. Numerous other oviraptorid specimens, have been called Oviraptor, but only the holotype can be properly referred to the genus.
Oviraptor was originally hypothesized to be an egg-eater (Osborn, 1924) based on close association with a supposed Protoceratops nest with eggs (AMNH 6508). However, Norell et al. (1994) discovered a brooding oviraptorid specimen which showed the nest and eggs belonged to Oviraptor itself. The diet of Oviraptor is still debated although a lizard preserved in the holotype's body cavity (Norell et al., 1995) suggests it was at least partially carnivorous.
The holotype was only briefly illustrated and described by Osborn (1924b), but a detailed redescription has not yet appeared. Smith's (1992) attempt made many errors, which were corrected by Clark et al. (2002) in their description of the skull. Makovicky (1995) added information about the preserved vertebrae, though the appendicular remains remain largely ignored.
Osborn (1924b) initially assigned Oviraptor to Ornithomimidae, and even after the naming of Oviraptoridae and discovery of more genera, was not placed in a cladistic context until Barsbold et al. (1990). The latter authors positioned it outside a Conchoraptor+"Ingenia" clade. This is found in nearly all analyses since, despite the massive increase in taxon number. The remaining unknown is whether Oviraptor is basal to Citipati+"Ingenia", or closer to some taxa with short dentaries.
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.
Brown and Schlaikjer, 1940. The structure and relationships of Protoceratops. Annals of the New York academy of Sciences. 40(3), 133-266.
Barsbold, 1976. On a new Late Cretaceous family of small theropods (Oviraptoridae fam. n.) of Mongolia. Doklady Akademia Nauk SSSR. 226, 685-688. [in Russian]
Osmolska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120. [in Russian]
Barsbold, 1986. Raubdinosaurier Oviraptoren. In Vorobyeva (editor). Herpetologische Untersuchungen in der Mongolischen Volksrepublik. 210-223. Akademia Nauk SSSR Institut Evolyucionnoy Morfologii i Ekologii Zhivotnikhim. Moskva: A.M. Severtsova. [in Russian, German summary]
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press, Berkeley. pp. 249-258.
Smith, 1990. Osteology of Oviraptor philoceratops, a possible herbivorous theropod from the Upper Cretaceous of Mongolia. Journal of Vertebrate Paleontology. 3(supplement), 42A.
Smith, 1992. The type specimen of Oviraptor philoceratops, a theropod dinosaur from the Upper Cretaceous of Mongolia. Neues Jahrbuch für Geologie und Palaontologie, Abhandlungen. 186, 365-388.
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.
Norell, Clark, Chiappe, and Dashzeveg, 1995. A nesting dinosaur. Nature. 378, 774-776.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria (Dinosauria: Theropoda). M.S. thesis, Copenhagen University, Copenhagen, Denmark.
Norell, Dingus and Gaffney, 1995. Discovering Dinosaurs. E.J. Knopf. 225 pp.
Dong and Currie, 1996. On the discovery of an oviraptorid skeleton on a nest of eggs at Bayan Mandahu, Inner Mongolia, People's Republic of China. Canadian Journal of Earth Sciences. 33, 631-636.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Barsbold, 1997. Oviraptorosauria. In Currie and Padian (editors). Encyclopedia of dinosaurs. 505-509. New York: Academic Press.
Morell, 1997. The Origin of Birds: the Dinosaur Debate. Audubon. March-April, 36-45.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5), 945-960.
http://www.oucom.ohiou.edu/dbms-witmer/collections/Theropods/oviraptor.htm

Rinchenia Osmolska, Currie and Barsbold, 2004
= "Rinchenia" Barsbold, 1997
R. mongoliensis (Barsbold, 1986) Osmolska, Currie and Barsbold, 2004
= Oviraptor mongoliensis Barsbold, 1986
= "Rinchenia" mongoliensis (Barsbold, 1986) Barsbold, 1997
= Citipati mongoliensis (Barsbold, 1986) Paul, 2010
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia

Holotype- (GI 100/32A) skull (156 mm), mandibles (150 mm), twelve cervical vertebrae, cervical ribs, nine dorsal vertebrae, seven sacral vertebrae (217 mm), twenty-three caudal vertebrae, five chevrons, scapulocoracoids (scap ~203 mm), furcula, sternal plate, humerus (163 mm), radius (~103.3 mm), ulna (~116 mm), phalanx I-1 (~72 mm), proximal manual ungual I, three metacarpal or phalangeal fragments, ilium (225 mm), femur (287 mm), tibia (~370 mm), proximal fibula, phalanx I-1 (~38 mm), pedal ungual I (~25 mm), proximal metatarsal II, phalanx II-1 (~48 mm), phalanx II-2 (~36 mm), pedal ungual II (~40 mm), phalanx III-3 (23.8 mm), proximal metatarsal IV, phalanx IV-1 (~40 mm), phalanx IV-2 (17 mm), phalanx IV-3 (12 mm), phalanx IV-4 (11.9 mm), pedal ungual IV (33 mm)
Comments- Barsbold (1986) originally described this specimen as a new species of Oviraptor, but later (1997) assigned it to a new genus, "Rinchenia". The absence of particular details in the latter description kept Rinchenia a nomen nudum until Osmolska et al. (2004).
The skull and mandible were first illustrated by Barsbold (1986), the ilium by Barsbold et al. (1990), and the twenty-sixth caudal vertebra by Barsbold et al. (1990). Nothing else has been publically illustrated of this specimen, although a photo of part of the skull was published by Clark et al. (2002) and the first few cervical vertebrae are visible in an online photo of the skull and mandible. Snively (2000) studies and illustrates a metatarsus as Rinchenia mongoliensis, but the specimen number (IGM 100/42) shows this is actually Citipati sp. nov.. Available descriptions are also sparse, with far more information available in the data matrices of Norell et al. (2001), Maryanska et al. (2002), Lu (2004) and Osmolska et al. (2004) than in any written work. Fanti et al. (2012) provide measurements.
References- Barsbold, 1986. Raubdinosaurier Oviraptoren. In Vorobyeva (editor). Herpetologische Untersuchungen in der Mongolischen Volksrepublik. 210-223. Akademia Nauk SSSR Institut Evolyucionnoy Morfologii i Ekologii Zhivotnikhim. Moskva: A.M. Severtsova. [in Russian, German summary]
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press, Berkeley. pp. 249-258.
Barsbold, 1997. Oviraptorosauria. In Currie and Padian (editors). Encyclopedia of dinosaurs. 505-509. New York: Academic Press.
Barsbold, Currie, Myhrvold, Osmolska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 403, 155-156.
Snively, 2000. Functional morphology of the tyrannosaurid arctometatarsus. Unpublished Masters Thesis. 273 pp.
Norell, Clark and Makovicky, 2001. Relationships among Maniraptora: Problems and prospects. 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. New Haven. 49-67.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. Southern Methodist University. unpublished PhD dissertation.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.

Shixinggia Lu and Zhang, 2005
S. oblita Lu and Zhang, 2005
Maastrichtian, Late Cretaceous
Pingling Formation, Guangdong, China

Holotype- (BPV-112) eighth dorsal vertebra (25 mm), ninth dorsal vertebra (25 mm), tenth dorsal vertebra (25 mm), two incomplete dorsal ribs, sacrum (27, 27, 27, 30, 30, 30, 30 mm), first caudal vertebra, second caudal vertebra, third caudal vertebra, ilia (242 mm), proximal pubis, partial femur, proximal tibia, proximal fibula, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-3, phalanx IV-4, pedal ungual IV
....(Shixing Museum coll.) fragments
Diagnosis- (modified from Lu and Zhang, 2005) preacetabular process lacking anteroventral process; anterioposteriorly shortened preacetabular process; large (pneumatic) foramen in the anterolateral surface of the proximal femur; small (pneumatic?) foramen in the proximomedial tibial surface.
Comments- This was first named and described in Lu's (2004) thesis, then officially by Lu and Zhang (2005). Lu and Zhang assign it to Oviraptoridae, though Lu (2004) found it to be a caenagnathid in a modified version of Maryanska et al.'s (2002) matrix, and sister taxon to Heyuannia within Oviraptoridae in a modified version of the Theropod Working Group matrix. It is a basal oviraptorid of Oviraptor- or Rinchenia-grade in the most recent and extensive oviraptorosaur analysis, that of Lu et al. (2015).
References- Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, Zhang and Li, 2003. A new oviraptorid dinosaur from the Late Cretaceous of Shixing, Nanxiong Basin of Guangdong Province, Southern China. JVP 23(3), 73A.
Lu, 2004. Oviraptorid dinosaurs from Southern China. Southern Methodist University. unpublished PhD dissertation.
Lu, 2005. Oviraptorid dinosaurs from Southern China. Geological Publishing House, Beijing. ISBN 7-116-04368-3. 200 pages + 8 plates.
Lu and Zhang, 2005. A new oviraptorid (Theropoda: Oviraptorosauria) from the Upper Cretaceous of the Nanxiong Basin, Guangdong Province of southern China. Acta Palaeontologica Sinica. 44(3), 412-422.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Tongtianlong Lu, Chen, Brusatte, Zhu and Shen, 2016
T. limosus Lu, Chen, Brusatte, Zhu and Shen, 2016
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China

Holotype- (DYM-2013-8) skull (~189 mm), mandible (119 mm), (cervical series 410 mm) atlas, axis (17 mm), third cervical vertebra ( mm), fourth cervical vertebra ( mm), fifth cervical vertebra ( mm), sixth cervical vertebra ( mm), seventh cervical vertebra ( mm), eighth cervical vertebra ( mm), ninth cervical vertebra ( mm), tenth cervical vertebra ( mm), eleventh cervical vertebra ( mm), (dorsal series 350 mm) several partial dorsal vertebrae, dorsal ribs, sacrum, nineteen caudal vertebrae, chevrons, incomplete scapulae (185 mm), coracoids (72 mm), furcula, partial sternum, humeri (131 mm), incomplete radius (102 mm), fragmentary ilium, partial pubes, partial ischia, partial femur (250 mm), incomplete tibia (320 mm), incomplete fibula, astragalus, calcaneum, partial tarsometatarsus (II 125, III 135, IV 110 mm), phalanx II-1, phalanx II-2, pedal ungual II, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal V (38 mm)
Diagnosis- (after Lu et al., 2016) dome-like skull roof with highest point located above posterodorsal corner of orbit; anterior margin of premaxilla highly convex in lateral view; distinct process at middle of anterior parietal margin; lacrimal shaft anteroposteriorly long in lateral view, with flat lateral surface; foramen magnum smaller than occipital condyle (also in Incisivosaurus and Anzu); absence of dentary symphyseal ventral process; absence of posterolateral sternal process.
Comments- Lu et al. (2016) recovered this as an "ingeniine" sister to Wulatelong+Banji in their analysis using a modified version of Maryanska et al.'s oviraptorosaur matrix.
Reference- Lu, Chen, Brusatte, Zhu and Shen, 2016. A Late Cretaceous diversification of Asian oviraptorid dinosaurs: Evidence from a new species preserved in an unusual posture. Scientific Reports. 6, 35780.

Wulatelong Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao, 2013
W. gobiensis Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao, 2013
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype
- (IVPP V18409) (29 kg; adult) incomplete skull, partial mandible, eleven partial dorsal vertebrae, four partial dorsal ribs, first sacral centrum, sixteen partial caudal vertebrae, incomplete scapulocoracoids, sternal plates (one partial, one incomplete), partial humerus, metacarpal I (~35 mm), incomplete phalanx I-1 (78 mm), manual ungual I, metacarpal II (~76 mm), phalanx II-1 (35 mm), phalanx II-2 (33 mm), partial manual ungual II, phalanx III-1 (51 mm), partial phalanx III-2, phalanx III-3 fragment, manual ungual III fragment, incomplete ilium (~225 mm), incomplete pubis (~250 mm), partial ischium, incomplete femur (255 mm), incomplete tibia (325 mm), incomplete fibula, proximal tarsus, distal tarsal IV, incomplete metatarsal II, phalanx II-2 (27 mm), pedal ungual II (45 mm), incomplete metatarsal III (143 mm), phalanx III-1 (43 mm), phalanx III-2 (30 mm), phalanx III-3 (24 mm), pedal ungual III, metatarsal IV (139 mm), phalanx IV-1 (30 mm), phalanx IV-2 (23 mm), phalanx IV-3 (20 mm), phalanx IV-4 (16 mm), pedal ungual IV (35 mm), metatarsal V
Diagnosis- (after Xu et al., 2013) ventral extremity of external naris located below midheight of premaxilla; strap-like jugal process of maxilla extends well beyond preorbital bar posteriorly and overlaps jugal; anterodorsal process of surangular basally constricted in lateral view.
Comments- The holotype was discovered in 2009. This entry switches the identification of manual digits II and III (III and IV of the authors) based on figure 2A, as there is no room for a third non-ungual phalanx in the uppermost digit. The length of the complete phalanx in the other digit and fragmentary nature of more distal phalanges makes this uncertain, however. Xu et al. (2013) proposed Wulatelong was a basal oviraptorid, which was corroborated by the analysis of Lu et al. (2015).
References- Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao, 2013. A new oviraptorid from the Upper Cretaceous of Nei Mongol, China, and its stratigraphic implications. Vertebrata PalAsiatica. 51(2), 85-101.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Yulong Lu, Currie, Xu, Zhang, Pu and Jia, 2013
Y. mini Lu, Currie, Xu, Zhang, Pu and Jia, 2013
Late Cretaceous
Qiupa Formation, Henan, China
Holotype
- (HGM 41HIII-0107) (<1 year old young juvenile) skull (48.16 mm), mandibles (51.97 mm), atlas, axis, five cervical vertebrae, posterior dorsal vertebrae, dorsal ribs, about twenty proximal and mid caudal vertebrae, scapula (42 mm), clavicles, humeri (44.6 mm), radii (44.5 mm), ulnae (44.7 mm), distal phalanx I-1, manual ungual I, metacarpal II (22.8 mm), phalanx II-1, phalanx II-2, manual ungual II, metacarpal III, phalanx III-2, phalanx III-3, manual ungual III, incomplete ilium (55 mm), partial femur (72 mm), tibiae (88.8 mm), incomplete fibulae, astragali, metatarsal I, partial phalanx I-1, metatarsals II, metatarsals III (43.3 mm), metatarsals IV, pedal phalanges including III-3 (11.7 mm), pedal unguals
Paratypes- (HGM 41HIII-0301) (embryo) skeleton including radius (22.6 mm), ulna (21 mm), ilium (39.3 mm), femur (46 mm), 26 eggs (19 not purchased), nest
(HGM 41HIII-0108) (young juvenile) posterior skull, posterior mandibles
(HGM 41HIII-0109) (young juvenile) skull, mandibles (48.54 mm), partial postcranial skeleton including scapula (50.26 mm), humerus (35.9 mm)
(HGM 41HIII-0110) (young juvenile) incomplete skull, incomplete mandibles, few cervical centra
(HGM 41HIII-0111) (young juvenile) ilium
Diagnosis- (after Lu et al., 2013) posterodorsal corner of antorbital fenestra and anteroventral corner of external naris at the same level (also in Citipati); distinct opening in premaxilla anteroventral to the external naris; antorbital fossa partly bordered by premaxilla anterodorsally; subnarial process of premaxilla does not contact anterior process of lacrimal; parietal almost as long as frontal; in dorsal view, posterior margin forms a straight line between postzygapophyses in fourth and fifth cervical vertebrae; femur longer than ilium.
Comments- Kobayashi et al. (2008) first mentioned Qiupa oviraptorids. While the holotype and HGM 41HIII-0108 were found associated, the others were found in separate quarries up to 4 km apart. The diagnostic characters may merely be characteristic of juvenile oviraptorids, and Yulong's recovered poisition in Lu et al.'s (2013) Maryanksa et al.-based analysis (between Gigantoraptor and more derived oviraptorids) may be basal due to ontogenetically variable characters, as juvenile theropods commonly end up more basally than adults. However, Lu et al.'s (2015) most recent and extensive oviraptorosaurian analysis still found it to be a basal oviraptorid, this time in a clade with taxa known only from older material (Nomingia, Nankangia).
References- Kobayashi, Lu, Lee, Xu and Zhang, 2008. A new basal ornithomimid (Dinosauria: Theropoda) from the Late Cretaceous in Henan province of China. Journal of Vertebrate Paleontology. 28(3), 101A.
Lu, Currie, Xu, Zhang, Pu and Jia, 2013. Chicken-sized oviraptorid dinosaurs from central China and their ontogenetic implications. Naturwissenschaften. 100(2), 165-175.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

"Ingeniinae" Barsbold, 1981
Definition- ("Ingenia" yanshini <- Oviraptor philoceratops) (suggested)
Other definitions- ("Ingenia" yanshini + Conchoraptor gracilis) (Osmolska, Currie and Barsbold, 2004)
Comments- Barsbold created this taxon to separate "Ingenia" from Oviraptor (to which he referred specimens now distinguished as Conchoraptor and Citipati sp.). Once he named Conchoraptor and Rinchenia (originally Oviraptor mongoliensis), these were excluded from "Ingeniinae" as well. Barsbold et al. (1990) retain this taxonomy, though their cladogram shows oviraptorines to be paraphyletic to "ingeniines", with Conchoraptor closer to "Ingenia" than to Oviraptor. Maryanska et al. (2002) found Conchoraptor and Ingenia to be in a clade exclusive of Citipati and Rinchenia, noting this was equivalent to "Ingeniinae" but poorly supported by data. The definition of Osmolska et al. (2004) functions in their phylogeny, where "Ingenia", Conchoraptor and Khaan form a clade sister to a Citipati + Rinchenia clade (which also presumably includes Oviraptor due merely to its crest). In Lu's (2004) topology based on the Theropod Working Group matrix, Avimimus, Chirostenotes and all oviraptorids are "ingeniines", thus losing any resemblence to Barsbold's or Osmolska et al.'s intent. In his phylogeny based on Maryanska et al.'s matrix, all oviraptorids except Khaan and Oviraptor are "ingeniines". Senter's (2007) phylogeny finds Conchoraptor, Khaan, Heyuannia and "Ingenia" to be "ingeniines". The most recent and extensive analysis (Lu et al., 2005) finds Conchoraptor, Machairasaurus, Jiangxisaurus, Ganzhousaurus, Nemegtomaia, "Ingenia" and Heyuannia to be in the clade, though they place the name at the more basal node containing Khaan too.
The quotations around the name reflect the fact that Ingenia is preoccupied by a nematode genus.
References- Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press, Berkeley. pp. 249-258.
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati osmolskae (Theropoda, Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops. American Museum Novitates. 3364, 1-24.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. Southrern Methodist University. unpublished PhD dissertation.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Conchoraptor Barsbold, 1986
C. gracilis Barsbold, 1986
= Citipati gracilis (Barsbold, 1986) Paul, 2010
Late Campanian, Late Cretaceous
Red Beds of Khermeen Tsav, Mongolia

Holotype- (IGM 100/20) skull (~108 mm), mandible
Paratype- (IGM 100/38) metacarpal I (24.6 mm), phalanx I-1 (32.9 mm), manual ungual I (~26 mm), metacarpal II (42.8 mm), distal phalanx II-1, phalanx II-2 (26.3 mm), manual ungual II (~23 mm), metacarpal III (41.4 mm), phalanx III-1 (17.6 mm), phalanx III-2 (~19 mm), phalanx III-3 (~25 mm), manual ungual III (~22 mm)
Referred- (IGM 100/21) mandible (93 mm) (Barsbold, 1976)
(IGM 100/36) material including skull, furcula, ilium (176 mm), pubis (145 mm), femur (182 mm), tibia (215 mm), metatarsal III (102.2 mm), phalanx III-1 (31.1 mm), phalanx III-3 (14.6 mm), phalanx IV-2 (13.5 mm), phalanx IV-3 (9.6 mm), phalanx IV-4 (7.3 mm), pedal ungual IV (18.9 mm) (Barsbold 1981)
(IGM 100/39) material including metacarpal I (~23 mm), phalanx I-1 (41 mm), manual ungual I (~27 mm), phalanx II-2 (~31 mm), manual ungual II (~31 mm), metacarpal III (44 mm), phalanx III-3 (~24 mm), manual ungual III (20 mm) (Maryanska et al., 2002)
(IGM 100/46) material including (?)metatarsal II, (?)metatarsal III, pedal phalanx III-3 (18.2 mm), (?)metatarsal IV, phalanx IV-2 (15.9 mm), phalanx IV-3 (11.5 mm), phalanx IV-4 (11.2 mm), pedal ungual IV (24.9 mm), (?)two pedal unguals, (?)metatarsal V (Maryanska et al., 2002)
(IGM 100/47) materal including ilium (173 mm), femur (195 mm) and tibia (258 mm) (Maryanska et al., 2002)
(ZPAL MgD-I/95) incomplete skull (98 mm), mandibles (86 mm), metatarsus (141 mm) (Osmolska, 1976)
(ZPAL MgD-I/101) skull, postcranial fragments (ZPAL online)
Maastrichtian, Late Cretaceous
Baruungoyot Formation, Mongolia

(IGM 100/1275) partial skeleton including sacrum, twenty-seven caudal vertebrae (first caudal 24.5 mm), pygostyle, femur (212 mm), tibia (250 mm), metatarsal III (107.5) and phalanx III-1 (35.3 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 1B) material including ilium (74.5 mm) and femur (81.2 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 2B) material including ilium (75 mm), femur (80 mm), metatarsal III (52.2 mm) and phalanx III-1 (14.1 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 7A) material including ilium (77.7 mm) and femur (82.2 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 7B) material including femur (87.3 mm) and tibia (110 mm) (Lu et al., 2013)
Late Cretaceous
Mongolia

(FPDM-V6232) skull, skeleton (Azuma, 2005)
(FPDM-V6234) skull, skeleton (Azuma, 2005)
(IGM 97/212) specimen including femur (250 mm) (Erickson et al., 2009)
(IGM 100/42; note not the same as the Citipati sp. complete skeleton) specimen including tibia (225 mm), phalanx III-3 (17.4 mm), phalanx IV-2 (18.2 mm), phalanx IV-3 (13.1 mm), phalanx IV-4 (11.9 mm), phalanx IV-5 (25.5 mm) (Fanti et al., 2012)
(IGM 100/97) material including femur (185 mm) and tibia (258 mm) (Fanti et al., 2012)
(IGM 100/1203) material including scapula, ilium (158 mm), ischium, femur (154 mm), tibia (184 mm), metatarsal III (83.5 mm) and phalanx III-1 (24 mm) (Lu et al., 2013)
(IGM 100/3006) material including partial skull, proximal caudal vertebrae, chevrons, scapula, coracoid, femur (152.7 mm), metatarsal III (87.5 mm) and phalanx III-1 (24.8 mm) (Balanoff and Norell, 2012; braincase described by Balanoff et al., 2014)
(IGM 102/01) material including partial tibia, phalanx IV-2 (15.2 mm) (Fanti et al., 2012)
(IGM 102/03) material including partial skull, partial scapula, partial humerus, radius (99 mm), ulna (100.1 mm), metacarpal I (24 mm), phalanx I-1 (31 mm), manual ungual I (42 mm), metacarpal II (~44 mm), phalanx II-1 (18.3 mm), phalanx II-2 (15.2 mm), manual ungual II (22 mm), ilium (226 mm), femur (240 mm), tibia (277 mm), metatarsal III (121 mm), phalanx III-1 (31 mm), phalanx III-3 (17.2 mm), phalanx IV-2 (14.4 mm), phalanx IV-3 (10.9 mm), phalanx IV-4 (8.7 mm), pedal ungual IV (33 mm) (Fanti et al., 2012)
(IGM 110/07) material including skull (94 mm), mandible (87.2 mm) (Fanti et al., 2012)
(IGM 110/10) material including mandible (~55 mm) (Fanti et al., 2012)
(IGM 110/11) material including partial skull (Fanti et al., 2012)
(IGM 110/12) material including skull (~101 mm), partial dentary (Fanti et al., 2012)
(IGM 110/18; actually two individuals, as listed below) material including distal caudal vertebrae, pubis and ischium (Lu, 2004)
----(IGM 110/18a) material including humerus (91.1 mm), radius (87.3 mm), ulna (88.2 mm) (Fanti et al., 2012)
----(IGM 110/18b) material including radius (~78.9 mm), ulna (75.1 mm) (Fanti et al., 2012)
(IGM 110/19) material including thirty-two caudal vertebrae, ilium (~115 mm) (Barsbold et al., 2000)
(IGM 110/20) material including partial skull (Fanti et al., 2012)
(IGM 110/21) material including mandible (68.2 mm), humerus (61.3 mm), radius (~54.6 mm), ulna (55 mm), metacarpal I (22.8 mm), partial metacarpal II, tibia (164 mm) (Fanti et al., 2012)
(IGM 110/22) material including partial skull (Fanti et al., 2012)
(IGM 110/25) material including scapula (Fanti et al., 2012)
(IGM 110/26) material including pedal phalanx III-3 (10.6 mm), phalanx IV-2 (13.9 mm), pedal ungual IV (17.9 mm) (Fanti et al., 2012)
(IGM 110/30) material including metacarpal I (26 mm), phalanx I-1 (38 mm), manual ungual I (~30 mm), phalanx II-1 (~27.5 mm), phalanx II-2 (33 mm), manual ungual II (25 mm) (Fanti et al., 2012)
(PIN coll.) skull (92 mm), mandibles (68 mm) (Maryanska et al., 2002)
(PMO X677; cast) skull, mandibles (Jensen, 2008)
(UALVP 49391; cast) skull (118 mm), mandible (89 mm) (Fanti et al., 2012)
(UALVP 49392; cast) skull (99 mm), mandible (86.6 mm) (Fanti et al., 2012)
(UALVP 54983) material including first caudal vertebra (23.9 mm) and femur (213 mm) (Funston et al., 2015)
---- material including first caudal vertebra (22.9 mm) and femur (212 mm) (Funston et al., 2015)
(ZPAL MgD-I/99) material including scapula and ilium (270 mm) (Fanti et al., 2012)
(ZPAL MgD-I/100) material including partial scapula, humerus (108 mm), phalanx I-1 (18 mm), manual ungual I (19 mm), phalanx II-1 (21 mm), phalanx II-2 (22 mm), manual ungual II (24 mm), ilium (210 mm), partial femur (Maryanska et al., 2002)
(ZPAL MgD-I/106) material including partial scapula and partial femur (Maryanska et al., 2002)
(private coll.) (1.1 m) skull, mandible, eight cervical vertebrae, cervical ribs, thirteen dorsal vertebrae, dorsal ribs, uncinate processes, over thirty caudal vertebrae, several chevrons, scapulae, sternal plates, sternal ribs, ilia, proximal pubis, proximal ischium, femur, tibia, fibula, proximal tarsus, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V, eggs?
(private coll.) skull, mandible, several cervical vertebrae, dorsal ribs, caudal vertebrae, chevrons, scapulae, humerus, proximal radius, ulna, manus, ilia, pubes, ischia, femora, tibiae, proximal fibulae, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
(private coll.) skull, eleven cervical vertebrae, twelve dorsal vertebrae, twenty-three dorsal ribs, uncinate process, nine caudal vertebrae, nine chevrons, scapula, sternal plates, humeri, radius, ulna, manus, ilia, femora, tibia, fibula, metatarsal II, phalanx II-1, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, pedal digit IV
(private coll.) skull, mandible, nine cervical vertebrae, two cervical ribs, thirteen dorsal vertebrae, eleven dorsal ribs, four uncinate processes, thirty-one caudal vertebrae, scapula, coracoid, furcula, humerus, radius, ulna, manus? (fake?), ilium, femora, tibia, proximal fibula, metatarsal II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
Comments- The holotype of Conchoraptor was discovered in 1971 and photographed by Barsbold (1976). At the time, it was referred to Oviraptor philoceratops. Additionally, a mandible of IGM 100/21 was photographed, while the morphology of these and four other specimens was briefly described. Barsbold (1977) described cranial characterics of Khermeen Tsav oviraptorids, probably including IGM 100/20 as well as "Ingenia" material. Barsbold (1981) illustrated a furcula of IGM 100/36, referring it and three other "supposedly young" specimens to O. philoceratops as well. These are probably IGM 100/38, 100/39, 100/46 and 100/47 listed in Maryanska et al. (2002) and may be the four specimens noted by Barsbold (1976). In 1986 Barsbold named Conchoraptor gracilis based on IGM 100/20, illustrating the skull of 100/20 and manus of 100/38 (as shown by Fanti et al., 2012). Barsbold et al. (1990) illustrated the skull in dorsal view and also a metatarsus and two pedal unguals, which may belong to IGM 100/46 since it is the only original specimen given pedal measurements by Fanti et al.. The skull was photographed incorrectly mounted on the "Ingenia" holotype in Psihoyos (1994). This same mount (with a mandible that may belong to Conchoraptor's holotype, "Ingenia"'s holotype or neither) is displayed at the Nakasato Dinosaur Center as Ingenia. Fanti et al. listed IGM 100/36 among Ingenia specimens and incorrectly listed IGM 100/80-D as the holotype skull, which is more likely the "Ingenia" skull noted by Lu (2004).
Osmolska (1976) described a skull and mandibles (ZPAL MgD-I/95) as Oviraptor sp.. She also noted an additional skull and postcranial fragments from the same formation, and a fragmentary skull from the Nemegt Formation. She believed they and the IGM specimens noted by Barsbold (1976) belonged to a new species. The Nemegt skull (probably ZPAL MgD-I/96) is crested and thus not Conchoraptor, but the other specimens may be. Possible specimen numbers for the second Khermeen Tsav skull are MgD-I/101 (listed on the ZPAL website as oviraptorid), and MgD-I/100 or MgD-I/106 (listed as Conchoraptor by Maryanska et al., 2002; but not given cranial measurements by Fanti et al., making them less likely candidates). ZPAL MgD-I/95 was illustrated as Oviraptor yanshini by Paul (1988), while Elzanowski (1999) described its palate in detail and called it Oviraptor sp.. Maryanska and Osmolska (1997) stated it was probably Conchoraptor or "Ingenia", along with a few additional fragmentary skulls (GIN 100/30A and two unnumbered GIN specimens called GIN A and B). Most recently, it was listed as Conchoraptor by Maryanska et al. (2002) and had its braincase and cranial pneumaticity described by Kundrat (2007) and Kundrat and Janacek (2007), who referred to it as Conchoraptor. Holtz (1994) has been the only publication since Osmolska (1976) to reference the postcrania, listing the metatarsus of ZPAL MgD-I/95 (as Conchoraptor) in a measurement table.
Additional material has also been referred to Conchoraptor. Barsbold et al. (2000) listed a tail (IGM 110/19) as belonging to the taxon, while Lu (2004) noted distal caudal vertebrae and pelvic elements from Conchoraptor specimen IGM 110/18. Besides ZPAL MgD-I/100 and 106, Maryanska et al. (2002) refer a PIN specimen and multiple unnumbered IGM specimens to it. The PIN specimen is quite possibly a skull and mandibles referred to "Ingenia" by Glut (1997) and Witmer (his website), as Witmer notes it is from the PIN. It was also photographed in Currie (2001). Osmolska et al. (2004) illustrate a skull as Conchoraptor that appears to be neither the holotype, ZPAL MgD-I/95 or the PIN specimen, and may therefore be one of the other specimens mentioned above. Jensen (2008) analyzed a cast for his thesis (PMO X677) which he referred to Conchoraptor gracilis. Erickson et al. (2009) examined the histology of IGM 97/212, which they referred to Conchoraptor gracilis. Balanoff and Norell (2012) mention many postcranial characters, largely from IGM 100/1203 and 100/3006, noting an in prep. paper. Fanti et al. and Lu et al. (2013) listed numerous specimens as belonging to this species and provided measurements. Among these is IGM 100/1275, which Balanoff and Norell (2012) mention numerous details of and Persons et al. (in press) describe the tail of. The latter authors note this specimen is one of many partial skeletons found together and catalogued as IGM MAE 97-212. Other specimens such as IGM 100/1203 and 3006 may also belong here, and Balanoff et al. (2014) have described the braincase of IGM 100/3006
It is especially common to find crestless privately held oviraptorid specimens being referred to Conchoraptor. One such incomplete skeleton (whose cast I viewed at SVP 2001) is listed as being from the Red Beds of Khermeen Tsav, but has manual proportions more similar to Heyuannia, so is provisionally referred to that taxon here. Another specimen is more difficult to identify as it doesn't preserve arms, though the constricted third metatarsal is unlike "Ingenia". Additional mounted skeletons have also been referred to Conchoraptor, but it is uncertain how much real and/or correctly identified material they are based on.
One notable issue surrounding Conchoraptor is the possibility some referred specimens belong to other taxa, as no referred specimen has ever been assigned to the genus based on shared derived characters with the holotype. This is particularily worrisome given the recent discovery of additional uncrested oviraptorids such as Khaan and Heyuannia, and the possibility young specimens of crested oviraptorids may lack crests. Indeed, the only supposed Conchoraptor specimen to be described in detail is ZPAL MgD-I/95. Although some information can be found in Barsbold (1976), Barsbold et al. (1990), Lu (2004) and Osmolska et al. (2004), the specimens examined are not specified (a recent exception is Balanoff and Norell, 2012). Similarly, the matrices of Norell et al. (2001), Maryanska et al. (2002), Lu (2004), Fanti et al. (2012) and Lu et al. (2013) are extremely informative and indicate basically every element is represented in at least one specimen, but obviously don't indicate which specimens were used to code each character. Maryanska et al. do list several specimens they used to code their Conchoraptor OTU, but ironically the holotype was not among them. The measurement tables in Fanti et al. and Currie et al. (2016) will prove useful to refer specimens based on ratios once they are examined.
References- Barsbold, 1976. On a new Late Cretaceous family of small theropods (Oviraptoridae fam. n.) of Mongolia. Doklady Akademia Nauk SSSR. 226, 685-688. [in Russian]
Osmolska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
Barsbold, 1977. Kinetism and peculiarities of the jaw apparatus of oviraptors (Theropoda, Saurischia). Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 4, 34–47.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, 1986. Raubdinosaurier Oviraptoren. In Vorobyeva (editor). Herpetologische Untersuchungen in der Mongolischen Volksrepublik. 210-223. Akademia Nauk SSSR Institut Evolyucionnoy Morfologii i Ekologii Zhivotnikhim. Moskva: A.M. Severtsova. [in Russian, German summary]
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press, Berkeley. pp. 249-258.
Holtz, 1994. The arctometatarsalian pes, an unusual structure of the metatarsus of Cretaceous Theropoda (Dinosauria: Saurischia). Journal of Vertebrate Paleontology. 14, 480-519.
Psihoyos, 1994. Hunting Dinosaurs. Random House. 288 pp.
Glut, 1997. Dinosaurs, the Encyclopedia: Mcfarland & Company, Inc., Publishers, 1076pp.
Maryanska and Osmolska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.
Elzanowski, 1999. A comparison of the jaw skeleton in theropods and birds, with a description of the palate in the Oviraptoridae. Smithsonian Contributions to Paleobiology. 89, 311-323.
Barsbold, Currie, Myhrvold, Osmolska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 403, 155-156.
Currie, 2001. Theropod dinosaurs from the Cretaceous of Mongolia. in Benton, Shishkin, Unwin and Kurochkin (eds). The Age of Dinosaurs in Russia and Mongolia. pp 434-455.
Norell, Clark and Makovicky, 2001. Relationships among Maniraptora: problems and prospects. 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. New Haven: Peabody Mus. Nat. Hist., Yale Univ. pp. 49-67.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. Southrern Methodist University. unpublished PhD dissertation.
Azuma, 2005. The Flying Dinosaurs: Fukui Prefectural Dinosaur Museum. 118pp.
Kundrat, 2007. Avian-like attributes of a virtual brain model of the oviraptorid theropod Conchoraptor gracilis. Naturwissenschaften. 94, 499-504.
Kundrat and Janacek, 2007. Cranial pneumatization and auditory perceptions of the oviraptorid dinosaur Conchoraptor gracilis (Theropoda, Maniraptora) from the Late Cretaceous of Mongolia. Naturwissenschaften. 94(9), 769-778.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles. Masters Thesis. University of Oslo. 48 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.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Balanoff, Bever and Norell, 2014. Reconsidering the avian nature of the oviraptorosaur brain (Dinosauria: Theropoda). PLoS ONE. 9(12), e113559.
Persons, Currie and Norell, 2014. Oviraptorosaur tail forms and functions. Acta Palaeontologica Polonica. 59(3), 553-567.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osmólska, 2016. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.
Balanoff and Norell, in prep.
http://www.oucom.ohiou.edu/dbms-witmer/dinoskulls02.htm
http://www.paleo.pan.pl/collect.htm#Mon-reptilia

Heyuannia Lu, 2002
H. huangi Lu, 2002
= Citipati huangi (Lu, 2002) Paul, 2010
Late Santonian-Early Campanian, Late Cretaceous
Zhutian (=Dalangshan) Formation, Guangdong, China
Holotype
- (HYMV1-1) partial skull (~150 mm), mandible (~150 mm), hyoid (50 mm), thirteen cervical vertebrae (520 mm; fourth 30 mm; sixth ~45 mm), twelve dorsal vertebrae (320 mm), dorsal ribs (30-160 mm), uncnate processes, first sacral vertebra (27 mm), second sacral vertebra (30 mm), third sacral vertebra (30 mm), fourth sacral vertebra (~30 mm), fifth sacral vertebra (~30 mm), sixth sacral vertebra (30 mm), seventh sacral vertebra (25 mm), eighth sacral vertebra (25 mm), first caudal vertebra (26 mm), second caudal vertebra (26 mm), third caudal vertebra (26 mm), fourth caudal vertebra (26 mm), fifth caudal vertebra (26 mm), sixth caudal vertebra (25 mm), seventh caudal vertebra (25 mm), first chevron (60 mm), second chevron (100 mm), third chevron (100 mm), fourth chevron (90 mm), fifth chevron, sixth chevron, proximal scapula, coracoid fragment, furcula, partial ilia (260 mm), pubes (260 mm), ischia (195 mm), femora (255 mm), tibiae (320 mm), proximal fibula, astragalus, calcaneum, metatarsal I (30 mm), phalanx I-1 (19 mm), pedal ungual I (30 mm), metatarsal II (110 mm), phalanx II-1 (35 mm), phalanx II-2 (20 mm), pedal ungual II (45 mm), metatarsal III (135 mm), phalanx III-1 (37 mm), phalanx III-2 (27 mm), metatarsal IV (120 mm), phalanx IV-1 (25 mm), phalanx IV-2 (20 mm), phalanx IV-3 (10 mm), phalanx IV-4 (10 mm)
Paratypes- (HYMV1-2) two dorsal ribs, gastralia, scapula (~175 mm), incomplete coracoid, furcula, partial sternum, sternal ribs, humerus (130 mm), radius (110 mm), ulna (127 mm), radiale, semilunate carpal, metacarpal I (32 mm), phalanx I-1, proximal metacarpal II, partial phalanx II-1, phalanx II-2, proximal metacarpal III
(HYMV1-3) phalanx I-1, manual ungual I, distal metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, distal metacarpal III, phalanx III-1
(HYMV1-4) partial pubis, distal femur, proximal tibia, proximal fibula
(HYMV1-5) incomplete manus
Referred- (HYMV1-6) partial skull, mandibular fragment, posterior cervical vertebrae, anterior dorsal vertebrae, dorsal ribs, uncinate processes, gastralia, last ten caudal vertebrae, partial scapulae (~165 mm), coracoids (50 mm; one partial), incomplete furcula, partial sternum, three partial sternal ribs, humeri (one proximal), partial radius, partial ulna, proximal femur, tibia (Lu, 2004)
(HYMV1-7) last three caudal vertebrae, distal metacarpal I, phalanx I-1, manual ungual I, metacarpal II fragment (Lu, 2004)
(HYMV2-1) humerus (105 mm), incomplete radius, ulna (103 mm), radiale, ulnare, semilunate carpal, metacarpal I (29 mm), phalanx I-1 (32 mm), manual ungual I (35 mm), metacarpal II (57 mm), phalanx II-1 (17 mm), phalanx II-2 (16 mm), manual ungual II (21 mm), metacarpal III (52 mm) (Lu, 2004)
(HYMV2-2) partial humerus, partial radius, partial ulna (Lu, 2004)
(HYMV2-3) femur, tibia, fibula, astragalus (Lu, 2004)
(HYMV2-4) pectoral girdle (Lu, 2004)
(HYMV2-5) pelvis, partial hindlimb including metatarsal V fragment (Lu, 2004)
(HYMV2-6) ten mid caudal vertebrae (Lu, 2004)
(HYMV2-7) partial pelvis (Lu, 2004)
(HYMV2-8) radius, ulna, metacarpal I (Lu, 2004)
? bones and eggs (Qiu and Huang, 2001)
Late Campanian, Late Cretaceous
Red Beds of Khermeen Tsav, Mongolia

Referred- ?(private coll.) (840 mm) skull, mandibles, several cervical vertebrae, dorsal vertebra, dorsal ribs, twenty-six caudal vertebrae, eighteen chevrons, scapulae, coracoids, furcula, humeri, radii, ulna, 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-1, phalanx III-2, phalanx III-3, manual ungual III, ilia, femora, tibia, fibula, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, metatarsal V (pers. obs.)
Diagnosis- (after Lu, 2004) quadratojugal articular surface of the quadrate groove-like; quadrate diverticulum enters the quadrate anterolaterally; pneumatic foramina present on the neural arches and ribs of cervical vertebrae; angle of the fused scapula and coracoid approximately 145º; ratio of coracoid length to the scapular length approximately 0.35; metacarpal I wraps metacarpal II proximally; pubis as long as ischium; ratio of femur length to tibia length 0.8.
Comments- Lu (2002) briefly described Heyuannia, then described it in more depth including additional specimens in his 2004 thesis (published in 2005). The forelimb and pectoral girdle were described further by Lu et al. (2005), though they incorrectly illustrate HYMV1-6 as 1-4. This makes Heyuannia one of the best described oviraptorids. Qiu and Huang (2001) previously mentioned "Ingenia" bones and supposed oviraptorid eggs from the same locality, which are therefore probably Heyuannia instead. A nearly complete privately owned skeleton has been casted and widely distributed as Conchoraptor, but its manual proportions more nearly resemble Heyuannia. This it is provisionally referred to Heyuannia here, though it is supposedly from the Red Beds of Khermeen Tsav like Conchoraptor and "Ingenia".
Cheng et al. (2008) referred two Macroolithus yaotunensis eggs with embryos (Chimei Museum 41 and NMNS-0015726-F02-embryo-01) from the Nanxiong Formation of Jiangxi to Heyuannia "or an oviraptorosaurian of similar kind" based on subarctometatarsaly. Yet this is true in most other oviraptorids as well except for "Ingenia". Wiemann et al. (2015a, b) used Cheng et al.'s paper to justify assigning all M. yaotunensis eggs to Heyuannia, including specimens from the Hugang Formation of Henan, the Yuanpu or Pingling Formation of Guangdong and the Nanxiong Formation of Jiangxi.
References- Qiu and Huang, 2001. Dinosaur fossils from the Heyuan Basin in Guangdong Province, China. in Deng and Wang (eds). Proceedings of the Eighth Annual Meeting of the Chinese Society of Vertebrate Paleontology, China Ocean Press, Beijing. 59-63.
Lu, 2002. A new oviraptorosaurid (Theropoda: Oviraptorosauria) from the Late Cretaceous of Southern China. Journal of Vertebrate Paleontology. 22(4), 871-875.
Lu, 2004. Oviraptorid dinosaurs from Southern China. Southrern Methodist University. unpublished PhD dissertation.
Lu, 2005. Oviraptorid dinosaurs from Southern China. Geological Publishing House, Beijing. ISBN 7-116-04368-3. 200 pages + 8 plates.
Lu, Huang and Qiu, 2005. The pectoral girdle and the forelimb of Heyuannia (Dinosauria: Oviraptorosauria). in Carpenter (ed.). The Carnivorous Dinosaurs. Indiana University Press. 256-273.
Cheng, Ji, Wu and Shan, 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic skeletons discovered for the first time in China. Acta Geologica Sinica. 82(6), 1089-1094.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, Müller and Sander, 2015a. The blue-green eggs of dinosaurs: How fossil metabolites provide insights into the evolution of bird reproduction. PeerJ PrePrints. https://dx.doi.org/10.7287/peerj.preprints.1080v1
Wiemann, Yang and Sander, 2015b. The colorful eggs of dinosaurs: How fossil metabolites reveal nesting behavior. Journal of Vertebrate Paleontology. Program and Abstracts 2015, 237.

"Ingenia" Barsbold, 1981 preoccupied Gerlach, 1957
"I." yanshini Barsbold, 1981
= Oviraptor yanshini Paul, 1988
= Ajancingenia yanshini (Barsbold, 1981) Easter, 2013a
Late Campanian, Late Cretaceous
Red Beds of Khermeen Tsav, Mongolia

Holotype- (IGM 100/30) parietal, braincase, mandible, nine cervical vertebrae, fourteen dorsal ribs, sacrum, thirty caudal vertebrae (first caudal 25.3 mm), twelve chevrons, scapulacoracoids (scap ~145 mm), furcula, sternum (~72 mm), humeri (141 mm), radii, ulnae (116 mm), semilunate carpal, metacarpal I (31.6 mm), phalanx I-1 (42 mm), manual ungual I (49.2 mm), metacarpal II (50.3 mm), phalanx II-1 (21.3 mm), phalanx II-2 (19.8 mm), manual ungual II (25.8 mm), metacarpal III (46.5 mm), phalanx III-1 (12.8 mm), phalanx III-2 (11 mm), phalanx III-3 (10 mm), manual ungual III (17 mm), ilia (~194 mm), pubes, ischia, femora (~228 mm), tibiae (290 mm), fibulae, astragali, calcanea, metatarsal I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (125 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
Paratypes- (IGM 100/31) frontal fragment, parietals, mandible (105 mm), postcrania including eight sacral vertebrae, radius (102.6 mm), metacarpal I (30.3 mm), phalanx I-1 (33.5 mm), manual ungual I (46.9 mm), metacarpal II (46.2 mm), phalanx II-1 (18.7 mm), phalanx II-2 (16.2 mm), manual ungual II (25.7 mm), metacarpal III (44 mm), phalanx III-1 (7.2 mm), phalanx III-2 (8 mm), phalanx III-3 (3 mm), ilium (228 mm), pubis (244 mm), ischium, femur (241 mm), tibia, fibula, metatarsal III (119.7 mm), phalanx III-3 (18.7 mm), phalanx IV-2 (14.5 mm), phalanx IV-3 (11.5 mm), phalanx IV-4 (10.6 mm), pedal ungual IV (33.3 mm)
(IGM 100/32) mandible (~120 mm), postcrania including cervical vertebrae, twenty-seven caudal vertebrae, partial scapula, humerus (140 mm), radius (110.5 mm), ulna (115.2 mm), semilunate carpal, phalanx I-1 (34.9 mm), manual ungual I (48.9 mm), metacarpal II (50.5 mm), phalanx II-1 (21.9 mm), phalanx II-2 (18.9 mm), manual ungual II (25.7 mm), metacarpal III (41.6 mm), phalanx III-1 (12.6 mm), ilium (242 mm), pubis (220 mm), femur (254 mm), tibia (294 mm), metatarsus (132 mm), phalanx III-1 (33.1 mm), phalanx III-3 (18.9 mm), phalanx IV-2 (16 mm), phalanx IV-3 (1.6 mm), phalanx IV-4 (11.2 mm), pedal ungual IV (32 mm)
(IGM 100/33) postcrania including sacrum, nineteen caudal vertebrae, scapula, sternal plates (~60 mm), furcula, humerus (128 mm), radius (94.7 mm), ulna (99 mm), metacarpal I (23.8 mm), phalanx I-1 (27.7 mm), manual ungual I (39.5 mm), metacarpal II (41.7 mm), phalanx II-1 (17.5 mm), phalanx II-2 (14.5 mm), manual ungual II (20 mm), partial metacarpal III, ilium, pubis (210 mm), ischium, femur (233 mm), tibia (267 mm), fibula, metatarsal III (124.5 mm), phalanx III-1 (34.3 mm), phalanx III-3 (17.4 mm), phalanx IV-2 (13.9 mm), phalanx IV-3 (9.8 mm), phalanx IV-4 (9.2 mm), pedal ungual IV (30.2 mm)
Late Cretaceous
Mongolia

Referred- (BHI coll.) skull, mandible (Fanti et al., 2012)
(FDPM-V6240) skull, skeleton (Azuma, 2005)
(IGM 100/34) material including scapula, partial humerus (69 mm), ilium (119 mm), femur (135 mm), tibia (162 mm), metatarsus (75.8 mm), phalanx III-1 (20.6 mm), phalanx III-3 (11.8 mm), phalanx IV-2 (10.8 mm), phalanx IV-3 (6.4 mm), phalanx IV-4 (6.2 mm), pedal ungual IV (18 mm) (Snively, 2000)
(IGM 100/35) material including femur (135 mm), partial tibia and phalanx III-1 (20.4 mm) (Maryanska et al., 2002)
(IGM 100/80-1) material including skull (115.6 mm), eleven cervical vertebrae (Lu, 2004)
?(IGM 102/11) material including first caudal vertebra (20.6 mm) and femur (242 mm) (Funston et al., 2015)
?(IGM 102/12) material including first caudal vertebra (29.3 mm) and femur (288 mm) (Funston et al., 2015)
(IGM 110/02) (Fanti et al., 2012)
(IGM 110/03) material including humerus (73.5 mm), radius (54.7 mm), ulna (55.3 mm), metacarpal I (15.4 mm), phalanx I-1 (16.8 mm), manual ungual I (19 mm), metacarpal II (23.8 mm), phalanx II-1 (9.5 mm) (Fanti et al., 2012)
(IGM PJC2002.17) skull, mandible (Fanti et al., 2012)
?(IGM coll.) material including skull (147.3 mm), mandible (120 mm), femur (238 mm), tibia (275 mm), metatarsal III (127.5 mm) and phalanx III-1 (34.5 mm) (Lu et al., 2013)
?(IGM coll.) material including skull (162 mm), humerus (113 mm), radius (90 mm), ulna (98 mm), metacarpal II (40.6 mm), femur (220 mm), tibia (265 mm), metatarsal III (118 mm) and phalanx III-1 (34.3 mm) (Lu et al., 2013)
Comments- The genus Ingenia is preoccupied by a nematode, as noted by Taylor (DML, 2004). Barsbold is aware of the situation, but Easter (2013a) did not consult him, proposing his own replacement name in a publication which largely consisted of copied and reworded information from this website used without my permission. While the initial objection to citing The Theropod Database was editorial, Easter did not object to this, lied about consulting with Barsbold in an online forum and took credit for my work in that same forum. Based on my objection, an erratum was published (Easter, 2013b), and while his name remains technically valid according to the ICZN, I do not support its use due to Easter's unprofessional and unethical actions.
Barsbold (1977) described cranial characterics of unspecified Khermeen Tsav oviraptorids, including "Ingenia" paratype IGM 100/31 and probably Conchoraptor material. He illustrated a mandible which was later labeled Ingenia by Barsbold et al. (1990). A mandible was listed as being present in the holotype by Barsbold (1981), so this is tentatively assumed to be it. "Ingenia" was first described by Barsbold (1981), who illustrated the manual digits of the holotype and mentioned three other specimens in addition to the holotype. Barsbold (1984) didn't expand on the description, but did illustrate the holotype's braincase, furcula and sternum, and the furcula and sternal plates of paratype IGM 100/33. Barsbold (1986) later illustrated the humerus, femur and metatarsus of the holotype. Barsbold et al. (1990) added some further information and illustrated the humerus in different views, radius, ulna, complete manus, pelvis and complete hindlimb missing only metatarsal V, pedal phalanx II-2 and digit I. These may all be from the holotype as well, but the specimen numbers were not listed. Psihoyos (1994) includes a photograph of the holotype (based on Dyke and Norell, 2005), which is largely complete except for the dorsal series, though the skull of the Conchoraptor holotype is mounted on it. This mount (with a mandible that is not the same as the one illustrated by Barsbold, 1977) is displayed at the Nakasato Dinosaur Center as Ingenia. Barsbold et al. (2000) illustrated the twenty-seventh caudal vertebra of IGM 100/32, and the nineteenth caudal vertebra and pelvis of IGM 100/33. The latter appears slightly different from the figure in Barsbold et al. (1990), perhaps indicating they are different specimens. Lu (2004) noted many anatomical details, including noting some of the elements preserved in paratype specimens. Osmolska et al. (2004) illustrated a scapulocoracoid and described several anatomical details. Osmolska (2003, 2004) described the skull roof of IGM 100/31, which she probably incorrectly stated was from the White Beds of Khermeen Tsav, which are slightly younger than the Red Beds. Fanti et al. (2012) and Currie et al. (2016) provide measurements of numerous elements.
Snively (2000) studied and illustrated the metatarsus of IGM 100/32 and 100/34. Maryanska et al. (2002) lists the latter specimen and IGM 100/35, while Lu (2004) notes cervical and cranial characters of IGM 100/80-1. The latter is notable as previously the only "Ingenia" cranial material thought to exist was the braincase of the holotype, whereas Lu's statements imply much of the skull is present in IGM 100/80-1. Confusingly, Fanti et al. list IGM 100/80-D as a skull and the holotype of Conchoraptor, but this is probably the same cranial specimen of "Ingenia" Lu notes. As with Conchoraptor, no evidence has ever been published defending the placement of paratypes or referred material in "Ingenia". Indeed, the sternum of IGM 100/33 differs from the holotype in lacking fusion between sternal plates (Barsbold, 1983), IGM 100/31 is said to lack fibulocalcanear contact (Lu, 2004) unlike the holotype, IGM 100/33 has ventrally keeled sacrals while the holotype has ventrally grooved sacrals, and varying amounts of sacral vertebrae have been described (7 by Barsbold, 1983; 8 in IGM 100/31 by Lu, 2004). Multiple taxa may be represented, or these may simply be ontogenetic/individual variation and/or illustration or descriptive error. The measurement table in Fanti et al. will prove useful to refer specimens based on ratios once it is examined.
Not "Ingenia"- Paul (1988) illustrated ZPAL MgD-I/95 as Oviraptor yanshini, a combination no other workers follow as it would result in placing all oviraptorids in Oviraptor given recent phylogenies. ZPAL MgD-I/95 is currently assigned to Conchoraptor, though it has not been compared with "Ingenia", which comes from the same formation. It may belong to either genus, or even a new taxon. Khaan paratype IGM 100/973 was first photographed in Dashzeveg et al. (1995) as cf. Ingenia, and labeled Ingenia yanshini in Webster (1996). This is the source of the Djadochta Formation listing for cf. Ingenia sp. in Weishampel et al. (2004). A skull and mandibles in the PIN collections are referred to Ingenia yanshini by Glut (1997) and Ingenia sp. by Witmer on his website. This is possibly the PIN coll. specimen referred to Conchoraptor by Maryanska et al. (2002) and is assigned to that genus here. Maryanska and Osmolska (1997) note ZPAL MgD-I/95 and a few additional fragmentary skulls (GIN 100/30A and two unnumbered GIN specimens called GIN A and B) from the Red Beds of Khermeen Tsav may belong to "Ingenia" or Conchoraptor. The holotype of Nemegtomaia was originally called Ingenia sp. by Lu (1999) and Lu et al. (2002), which is the source of the Nemegt Formation listing for Ingenia sp. in Weishampel et al. (2004). Qiu and Huang (2001) mentioned Ingenia bones and supposed oviraptorid eggs from the locality Heyuannia was later described from in Guangdong, China. They are thus probably Heyuannia instead of "Ingenia". Currie (2002) noted a new partial skeleton (PJC.2001.2) from the Nemegt Formation which he referred to cf. Ingenia sp.. It may end up to be referrable to Rinchenia or Nemegtomaia which are present in that formation.
References- Barsbold, 1977. Kinetism and peculiarities of the jaw apparatus of oviraptors (Theropoda, Saurischia). Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 4, 34–47.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological Expedition Transactions. 15, 28-39. [in Russian]
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120. [in Russian]
Barsbold, 1986. Raubdinosaurier Oviraptoren. In Vorobyeva (editor). Herpetologische Untersuchungen in der Mongolischen Volksrepublik. 210-223. Akademia Nauk SSSR Institut Evolyucionnoy Morfologii i Ekologii Zhivotnikhim. Moskva: A.M. Severtsova. [in Russian, German summary]
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster Co., New York. 464 pp.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. in Weishampel, Dodson and Osmolska (eds). The Dinosauria. University of California Press, Berkeley. pp. 249-258.
Psihoyos, 1994. Hunting Dinosaurs. Random House. 288 pp.
Dashzeveg, Novacek, Norell, Clark, Chiappe, Davidson, McKenna, Dingus, Swisher III and Perle, 1995. Unusual preservation in a new vertebrate assemblage from the Late Cretaceous of Mongolia. Nature. 374, 446-449.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Glut, 1997. Dinosaurs, the Encyclopedia: Mcfarland & Company, Inc., Publishers, 1076pp.
Maryanska and Osmolska, 1997. The quadrate of oviraptorid dinosaurs. Acta Palaeontologia Polonica. 42, 377-387.
Lu, 1999. New material of Ingenia (Barsbold, 1981) from the Nemegt Formation of southwestern Mongolia and its phylogenetic relationships among Oviraptorosauria. Unpublished Masters Thesis, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 50 pp.
Barsbold, Currie, Myhrvold, Osmolska, Tsogtbaatar and Watabe, 2000. A pygostyle from a non-avian theropod. Nature. 403, 155-156.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Unpublished Masters Thesis. 273 pp.
Norell, Clark and Makovicky, 2001. Relationships among Maniraptora: problems and prospects. 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. New Haven: Peabody Mus. Nat. Hist., Yale Univ. pp. 49-67.
Qiu and Huang, 2001. Dinosaur fossils from the Heyuan Basin in Guangdong Province, China. in Deng and Wang (eds). Proceedings of the Eighth Annual Meeting of the Chinese Society of Vertebrate Paleontology, China Ocean Press, Beijing. 59-63.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. In: Alberta Palaeontological Society, sixth annual symposium, “Fossils 2002’, presented by Alberta Paleontological Society, in conjunction with Canadian Society of Petroleum Geologists, Paleontological Division and Department of Earth Sciences, Mount Royal College, p. 8-12.
Lu, Dong, Azuma, Barsbold and Tomida, 2002. Oviraptorosaurs compared to birds. In Zhou and Zhang (eds.). Proceedings of'the 5th Symposium of the Society of Avian Paleontology and Evolution. Science Press. Beijing China. 175- 189.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Osmolska, 2003. Some aspects of the oviraptorosaur (Dinosauria, Theropoda) braincase. 1st EAVP Meeting, Basel Switzerland 15th–19th July 2003, 33. Natural History Museum, Basel.
Lu, 2004. Oviraptorid dinosaurs from Southern China. Southrern Methodist University. unpublished PhD dissertation.
Osmolska, 2004. Evidence on relation of brain to endocranial cavity in oviraptorid dinosaurs. Acta Palaeontologica Polonica. 49(2), 321-324.
http://dml.cmnh.org/2004Sep/msg00022.html
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004. Dinosaur Distribution. In Weishampel, Dodson and Osmolska. The Dinosauria Second Edition. University of California Press. 861 pp.
Azuma, 2005. The Flying Dinosaurs: Fukui Prefectural Dinosaur Museum. 118pp.
Dyke and Norell, 2005. Caudipteryx as a non-avialan theropod rather than a flightless bird. Acta Palaeontologica Polonica. 50(1), 101-116.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
http://www.oucom.ohiou.edu/dbms-witmer/dinoskulls02.htm
Easter, 2013a. A new name for the oviraptorid dinosaur "Ingenia" yanshini (Barsbold, 1981; preoccupied by Gerlach, 1957). Zootaxa. 3737(2), 184-190.
Easter, 2013b. Erratum. Zootaxa. 3750(1), 100.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osmólska, 2016. New specimens of the crested theropod dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica. 61(1), 143-157.

Jiangxisaurus Wei, Pu, Xu, Liu and Lu, 2013
J. ganzhouensis Wei, Pu, Xu, Liu and Lu, 2013
Campanian-Maastrichtian, Late Cretaceous
Nanxiong Formation, Jiangxi, China

Holotype- (HGM41HIII0421) (subadult) skull (150 mm), mandibles (130 mm), hyoid (60 mm), partial atlas, axis (31 mm), third cervical vertebra (~32 mm), fourth cervical vertebra (33 mm), fifth cervical vertebra (33 mm), sixth cervical vertebra (32 mm), seventh cervical vertebra (28 mm), eighth cervical vertebra (25 mm), three dorsal vertebrae (25, 26, 28 mm), nine partial dorsal ribs, first caudal vertebra (25 mm), second caudal vertebra (25 mm), third caudal vertebra (25 mm), fourth caudal vertebra (25 mm), fifth caudal vertebra (25 mm), sixth caudal vertebra (22 mm), seventh caudal vertebra (20 mm), eighth caudal vertebra (20 mm), ninth caudal vertebra (16 mm), three chevrons, scapula, coracoids, incomplete furcula, partial sternal plates, four sternal ribs, humerus (136 mm), radius (96 mm), ulna (95 mm), radiale, semilunate carpal, metacarpal I (25 mm), phalanx I-1 (30 mm), manual ungual I, metacarpal II (45 mm), phalanx II-?, metacarpal III (35 mm), phalanx III-?, ilial fragment, pubic fragment, ischial fragment
Diagnosis- (after Wei et al., 2013) elongated mandible (height 20% of length); weakly downturned mandibular symphysis; surangular with elongate and concave lateral surface; radiohumeral radio ~70%.
Comments- Wei et al. (2013) described this taxon as an oviraptorid of uncertain phylogenetic placement, but Lu et al (2015) recovered it as an "ingeniine."
References- Wei, Pu, Xu, Liu and Lu, 2013. A new oviraptorid dinosaur (Theropoda: Oviraptorosauria) from the Late Cretaceous of Jiangxi Province, Southern China. Acta Geologica Sinica (English Edition). 87(4), 899-904.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous of southern China and its paleobiogeographical implications. Scientific Reports. 5, 11490.

Khaan Clark, Norell and Barsbold, 2001
K. mckennai Clark, Norell and Barsbold, 2001
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia

Holotype- (IGM 100/1127) skull (~127 mm), mandible (108.4 mm), hyoids, twelve cervical vertebrae, cervical ribs, several dorsal vertebrae, dorsal ribs, uncinate process, gastralia, sacrum, twenty-seven caudal vertebrae (first caudal 18.3 mm), chevrons, scapulacoracoids (154.1 mm; scap 124.7 mm), furcula, sternal plates (58.8 mm), humeri (110.5, 108.6 mm), radii (~102 mm), ulnae (96.5 mm), radiale, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II (47.3 mm), phalanx II-1, phalanx II-2, manual ungual II, metacarpal III (44.4 mm), phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ilia (182 mm), pubis, ischium, femur (185 mm), tibia (212 mm), fibula (201 mm), metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal IV (~95 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
Paratypes- (IGM 100/973) skull (118.8 mm), mandibles (104 mm), hyoids, atlas, axis, ilia (193.5, 187.5 mm), pubis (156.8 mm), ischium (133.1 mm), femora (188.8, 199 mm), tibiae (221.4, 224 mm), fibula, astragali, calcanea, distal tarsal III, metatarsals I (26.7, ~23 mm), phalanges I-1, pedal ungual I, metatarsals II (82.8, 89.9 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (97.7, 98.8 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV (87.3, 93.9 mm), phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV, metatarsal V (35.8 mm)
(IGM 100/1002) incomplete skull (113.8 mm), mandible (101.6 mm), hyoids, twelve cervical vertebrae, cervical ribs, ten dorsal vertebrae, dorsal ribs, gastralia, sacrum, seven caudal vertebrae, four chevrons, scapulocoracoids (167 mm; scap 138.6 mm), furcula, sternal plates (55.1 mm), sternal ribs, humerus (117.5 mm), radii (95.3 mm), ulnae (~101.6 mm), radiale, semilunate carpal, metacarpal I (24.7 mm), phalanx I-1 (38 mm), manual ungual I (44.5 mm), metacarpal II (50.4 mm), phalanx II-1 (26 mm), phalanx II-2 (28.3 mm), manual ungual II (37 mm), metacarpal III (47.6 mm), phalanx III-1 (16.2 mm), phalanx III-2, phalanx III-3, manual ungual III, ilium (186 mm), pubis (~167 mm), ischium (~124 mm), femur (196 mm), tibia (232 mm), fibula (214 mm), astragalus, distal tarsal IV, metatarsal I (23.3 mm), phalanx I-1, pedal ungual I, metatarsals II (91.2 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals III (105.6 mm), phalanges III-1, phalanges III-2, phalanges III-3 (15.3 mm), pedal unguals III, metatarsals IV (98.3 mm), phalanges IV-1, phalanges IV-2 (14 mm), phalanges IV-3 (11.5 mm), phalanges IV-4 (10.7 mm), pedal unguals IV (31 mm), metatarsal V (35.9 mm)
Comments- IGM 100/973 was discovered in 1993 and first photographed in Novacek et al. (1994). It was photographed in Dashzeveg et al. (1995) as cf. Ingenia, and labeled Ingenia yanshini in Webster (1996). IGM 100/1127 and 100/1002 were discovered in 1995, and the latter was photographed in Morell (1997) and labeled Oviraptor. Clark et al. (2001) made these their new taxon Khaan mckennai. IGM 100/1002 and 100/1127 were found in close proximity to each other, and have been called "Romeo and Juliet." Balanoff and Norell (2012) recently described the taxon in depth, based on Balanoff's (2011) thesis.
Gatesy and Middleton (1997) published hindlimb measurements for an "undescribed oviraptorid (Norell pers. comm.)" which is a Khaan specimen based on nearly identical measurements listed by Dyke and Norell (2005).
References- Novacek, Norell, McKenna and Clark, 1994. Fossils of the Flaming Cliffs. Scientific American. 271(6), 60-69.
Dashzeveg, Novacek, Norell, Clark, Chiappe, Davidson, McKenna, Dingus, Swisher III and Perle, 1995. Unusual preservation in a new vertebrate assemblage from the Late Cretaceous of Mongolia. Nature. 374, 446-449.
Webster, 1996. Dinosaurs of the Gobi. National Geographic. 190(1), 70-89.
Gatesy and Middleton, 1997. Bipedalism, flight, and the evolution of theropod locomotor diversity. Journal of Vertebrate Paleontology. 17(2), 308-329.
Morell, 1997. The Origin of Birds: the Dinosaur Debate. Audubon. March-April, 36-45.
Clark, Norell and Barsbold, 2001. Two new oviraptorids (Theropoda: Oviraptorosauria) from the Late Cretaceous Djadokta Formation, Ukhaa Tolgod. Journal of Vertebrate Paleontology. 21(2), 209-213.
Lu, 2004. Oviraptorid dinosaurs from southern China. Southrern Methodist University. unpublished PhD dissertation.
Osmólska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel, Dodson and Osmólska, (eds). The Dinosauria, Second Edition. University of California Press (Berkeley). pp. 165-183.
Dyke and Norell, 2005. Caudipteryx as a non-avialan theropod rather than a flightless bird. Acta Palaeontologica Polonica. 50(1), 101-116.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.
Balanoff and Norell, 2009. Adult morphology and variation within the oviraptorid Khaan mckennai (Theropoda: Oviraptorosauria). Journal of Vertebrate Paleontology. 29(3), 57A.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution. PhD thesis. Columbia University. 522 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria: Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.

Machairasaurus Longrich, Currie and Dong, 2010
M. leptonychus Longrich, Currie and Dong, 2010
Campanian, Late Cretaceous
Bayan Mandahu Formation, Inner Mongolia, China
Holotype
- (IVPP V15979) distal radius, distal ulna, radiale, semilunate carpal, metacarpal I (24 mm), phalanx I-1 (30 mm), manual ungual I (29 mm), metacarpal II (38 mm), phalanx II-1 (21 mm), phalanx II-2 (22 mm), manual ungual II (24 mm), metacarpal III (34 mm), phalanx III-1 (12 mm), phalanx III-2, phalanx III-3, manual ungual III, manual fragments, several pedal phalangeal fragments, pedal ungual I, pedal ungual IV
Referred- (IVPP V15980) dorsal ribs, caudal vertebrae, chevrons, metacarpal I (32 mm), phalanx I-1 (79 mm), manual ungual I (50 mm), phalanx II-1 (60 mm), phalanx II-2 (66 mm), phalanx III-1 (38 mm), partial tibia, fragmentary metatarsal II, phalanx III-3 (28 mm), fragmentary metatarsal IV, phalanx IV-2 (27 mm) (Longrich, Currie and Dong, 2010)
Diagnosis- (after Longrich et al., 2010) manual unguals I–III elongate and bladelike in lateral view (length of claw approximately 400 per cent the height of the proximal articular surface).
Comments- Discovered in 1988 and 1990, the type specimens of Machairasaurus leptonychus were described by Longrich et al. in 2010 as a new taxon. They used a version of Osmolska et al.'s oviraptorosaur matrix which placed Machairasaurus in "Ingeniinae", more derived than Khaan and Conchoraptor but more basal than Heyuannia and "Ingenia". Fanti et al. (1012) provide measurements.
References- Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5), 945-960.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE. 7(2), e31330.

Nemegtomaia Lu, Tomida, Azuma, Dong and Lee, 2005
= Nemegtia Lu, Tomida, Azuma, Dong and Lee, 2004 preoccupied Szczechura, 1978
N. barsboldi (Lu, Tomida, Azuma, Dong and Lee, 2004) Lu, Tomida, Azuma, Dong and Lee, 2005
= Nemegtia barsboldi Lu, Tomida, Azuma, Dong and Lee, 2004
= Citipati barsboldi (Lu, Tomida, Azuma, Dong and Lee, 2004) Paul, 2010
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia

Holotype- (IGM 100/2112) skull (179 mm), mandibles (153 mm), partial axis, third cervical vertebra (33 mm), fourth cervical vertebra (29 mm), fifth cervical vertebra (32 mm), sixth cervical vertebra (31 mm), seventh cervical vertebra (31 mm), eighth cervical vertebra (33 mm), ninth cervical vertebra (35 mm), tenth cervical vertebra (37 mm), eleventh cervical vertebra (37 mm), twelfth cervical vertebra (36 mm), thirteenth cervical vertebra (33 mm), most cervical ribs, first dorsal vertebrae (31 mm), partial second dorsal vertebra, sixth dorsal neural arch, seventh dorsal neural arch, eighth dorsal vertebra, ninth dorsal vertebra, tenth dorsal vertebra, sacrum (32, 33, 29, 29, 29, 32, 40, ? mm), first caudal neural arch, second caudal neural arch, proximal scapula, distal humeri, radius, ilia (290 mm), proximal pubes, proximal ischia, proximal femur
Referred- (IGM 107/16) several fragmentary dorsal ribs, distal radius (~99 mm), distal ulna, semilunate carpals, two carpals, metacarpals I (17.7 mm), phalanges I-1 (24.9 mm), manual unguals I (34 mm), metacarpals II (34 mm), phalanges II-1 (16 mm), phalanges II-2 (13.5 mm), manual unguals II (16 mm), metacarpals III (31 mm), phalanges III-1 (one fragmentary; 9.7 mm), proximal phalanx III-2, ilial fragment, incomplete femora (~222 mm) (Fanti et al., 2012)
Maastrichtian, Late Cretaceous
Baruungoyot Formation, Mongolia

Referred- (IGM 107/15) partial skull (172 mm), partial mandibles (152 mm), cervical vertebral fragments, dorsal vertebral fragments, rib fragments, partial scapulae (~185 mm), partial humeri (~152 mm), partial radius (~144 mm), partial ulna, partial phalanx I-1, partial manual ungual I, fragmentary metacarpal II, partial phalanx II-2 (18 mm), incomplete manual ungual II (32 mm), metacarpal III fragment, partial pubes, incomplete femora (~286 mm), partial tibiae (~317 mm), partial fibulae, phalanges I-1, pedal unguals I, phalanx II-1, phalanx II-2, phalanx III-3 (25 mm), phalanx IV-2 (23 mm), phalanx IV-3 (16 mm), phalanx IV-4 (18.8 mm), pedal ungual IV (33 mm), nest, egg fragments (Fanti et al., 2012)
Diagnosis- (modified from Lu et al., 2004) prefrontal present; quadrate articulates with quadratojugal with a convex condyle.
Comments- The holotype was originally called Ingenia sp. by Lu (1999) and Lu et al. (2002), then Oviraptor sp. in Lu (2004). Lu et al. (2004) described it in detail and named it Nemegtia, though it was renamed by Lu et al. (2005) because Nemegtia is a genus of ostracod. Mounted skeletons incorporating the Nemegtomaia holotype exist but are misleading, as most of the appendicular and caudal material is faked. Note Fanti et al. (2012) claimed IGM 107/16 was from the Nemegt and Baruungoyot Formations in different parts of their paper, but personal communication (1-11-2017) from Fanti indicates the original collection information is incomplete and the associated sediment leaves Fanti favoring a source in the Nemegt Formation.
References- Lu, 1999. New material of Ingenia (Barsbold, 1981) from the Nemegt Formation of southwestern Mongolia and its phylogenetic relationships among Oviraptorosauria. Unpublished Masters Thesis, Institute of Vertebrate Paleontology and Paleoanthropology, Chinese Academy of Sciences, Beijing, 50 pp.
Lu, 2000. Oviraptorosaurs compared to birds. in Shi and Zhang (eds.). Fifth International Meeting of the Society of Avian Paleontology and Evolution and the Symposium on Jehol Biota. Vertebrata PalAsiatica. 38 (suppl.), 18.
Lu, Dong, Azuma, Barsbold and Tomida, 2002. Oviraptorosaurs compared to birds. In Zhou and Zhang (eds.). Proceedings of'the 5th Symposium of the Society of Avian Paleontology and Evolution. Science Press. Beijing China. 175- 189.
Lu, 2004. Oviraptorid dinosaurs from southern China. Southrern Methodist University. unpublished PhD dissertation.
Lu, Tomida, Azuma, Dong and Lee, 2004. New Oviraptorid Dinosaur (Dinosauria: Oviraptorosauria) from the Nemegt Formation of Southwestern Mongolia. Bulletin of the National Science Museum, Tokyo, Series C. 30, 95-130.
Lu, 2005. Oviraptorid dinosaurs from Southern China. Geological Publishing House, Beijing. ISBN 7-116-04368-3. 200 pages + 8 plates.
Lu, Tomida, Azuma, Dong and Lee, 2005. Nemegtomaia gen. nov., a replacement name for the oviraptorosaurian dinosaur Nemegtia Lu et al., 2004, a preoccupied name. Bulletin of the National Science Museum, Tokyo, Series C. 31, 51.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press. 320 pp.
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