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
= Avimimiformes Chatterjee, 1991
= Oviraptorosauria sensu Sereno, in press
Definition- (Oviraptor philoceratops <- Tyrannosaurus rex, Ornithomimus edmontonicus, Therizinosaurus cheloniformis, Troodon formosus, Passer domesticus)
Comments- 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).
A femur (ZIN PH 1/13) identified as oviraptorosaurian or ornithomimid by Nessov (1995) is ornithomimid (Averianov et al., 2003).
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.
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.
Agnolin and Martinelli, 2007. Did oviraptorosaurs (Dinosauria; Theropoda) inhabit Argentina? Cretaceous Research. doi: 10.1016/j.cretres.2006.10.006

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.

unnamed oviraptorosaur (Currie, Vickers-Rich and Rich, 1996)
Early Albian, Early Cretaceous
Eumeralla Formation, Victoria, Australia
Material- (NMV P186386) surangular
?(NMV P186302) dorsal vertebra (23 mm)
Comments- Britt (1993) mentions NMV 186303 (the holotype femur of Timimus) as a dromaeosaurid dorsal vertebra. This may be a typo for NMV 186302.
References- Britt, 1993. Pneumatic postcranial bones in dinosaurs and other archosaurs. PhD Thesis, University of Calgary (Canada), Alberta.
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.

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.

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, Kobayashi, Matsufune and Norell, 2009. Cranial osteology of the theropod dinosaur Incisivosaurus gauthieri (Theropoda: Oviraptorosauria). American Museum Novitates. 3651, 35 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.
Comments- 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).
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.

Caudipteridae Zhou and Wang, 2000

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, 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)
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, 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.
References- Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs from northeastern China. Nature. 393, 753-761.
Sereno, 1999. The evolution of dinosaurs. Science. 284, 2137-2147.
Xu, Wang and Wu, 1999. A dromaeosaurid dinosaur with filamentous integument from the Yixian Formation of China. Nature. 401, 262-266.
Jones, Farlow, Ruben, Henderson and Hillenius, 2000. Cursoriality in bipedal archosaurs. Nature. 406, 716-718.
Geist and Feduccia, 2000. Gravity defying behaviors: Identifying models for protoaves. American Zoologist. 40(4), 664-675.
Martin and Czerkas, 2000. The fossil record of feather evolution in the Mesozoic. American Zoologist. 40(4), 687-694.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria, Saurischia). Ph.D. dissertation, University of Bristol, Bristol. 583 pp.
Ruben and Jones, 2000. Selective factors associated with the origin of fur and feathers. American Zoologist. 40(4), 585-596.
Xu, Zhou and Wang, 2000. The smallest known non-avian theropod dinosaur. Nature. 408, 705-708.
Zhou and Wang, 2000. A new species of Caudipteryx from the Yixian Formation of Liaoning, northeast China. Vertebrata PalAsiatica. 38(2), 113-130.
Zhou, Wang, Zhang and Xu, 2000. Important features of Caudipteryx - evidence from two nearly complete new specimens. Vertebrata PalAsiatica. 38(4), 241-254.
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.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian dinosaurs. pp. 49–67 in Gauthier and Gall (eds.). New Perspectives on the Origin and Early Evolution of Birds: Proceedings of the International Symposium in Honor of John H. Ostrom. Yale Univ. Press.
Ruben and Jones, 2001. Feathered dinosaurs and other myths: a cold, hard look at reality. Journal of Morphology. 248(3), 278.
Christiansen and Bonde, 2002. Limb proportions and avian terrestrial locomotion. Journal of Ornithology. 143, 356-371.
Clark, Norell and Makovicky, 2002. Cladistic approaches to the relationships of birds to other theropod dinosaurs. pp. 31–60. in Chiappe and Witmer (eds.). Mesozoic Birds: Above the Heads of Dinosaurs. University of California Press, Berkeley.
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.
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. Beijing Science Press, pg. 175-189.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
Paul, 2002. Dinosaurs of the Air: The Evolution and Loss of Flight in Dinosaurs and Birds. Baltimore: Johns Hopkins University Press.
Xu, Cheng, Wang and Chang, 2002. An unusual oviraptorosaurian dinosaur from China. Nature. 419, 291-293.
Xu, Norell, Wang, Makovicky and Wu, 2002. A basal troodontid from the Early Cretaceous of China. Nature. 415, 780-784.
Makovicky, Norell, Clark and Rowe, 2003. Osteology and relationships of Byronosaurus jaffei (Theropoda: Troodontidae). American Museum Novitates. 3402, 1-32.
Rauhut, 2003. The interrelationships and evolution of basal theropod dinosaurs. Special Papers in Paleontology. 69, 1-215.
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.
Hwang, Norell, Ji and Gao, 2004. A large compsognathid from the Early Cretaceous Yixian Formation of China. Journal of Systematic Palaeontology. 2(1), 13-30.
Lu, 2004. Oviraptorid dinosaurs from Southern China. Southern Methodist University. unpublished PhD dissertation.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica. 50(6), 978-990.
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, Barsold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae (Dinosauria, Theropoda). Bulletin of the Gunma Museum of Natural History. 8, 1-20.
Xu and Norell, 2004. A new troodontid dinosaur from China with avian-like sleeping posture. Nature. 431, 838-841.
Dyke and Norell, 2005. Caudipteryx as a non-avialan theropod rather than a flightless bird. Acta Palaeontologica Polonica. 50(1), 101-116.
Feduccia, Lingham-Soliar and Hinchliffe, 2005. Do feathered dinosaurs exist? Testing the hypothesis on neontological and paleontological evidence. J. Morphology.
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.
Kirkland, Zanno, Sampson, Clark and DeBlieux, 2005. A primitive therizinosauroid dinosaur from the Early Cretaceous of Utah. Nature. 435, 84-87.
Mayr, Pohl and Peters, 2005. A well-preserved Archaeopteryx specimen with theropod features. Science. 310, 1483-1486.
Novas and Pol, 2005. New evidence on deinonychosaurian dinosaurs from the Late Cretaceous of Patagonia. Nature. 433, 858-861.
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. doi:10.1371/journal.pone.0007390

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)
= Caenagnathidae sensu Sues, 1997
Definition- (Chirostenotes pergracilis + Chirostenotes elegans + Elmisaurus rarus + Caenagnathasia martinsoni + BHM 2033)
= 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.

undescribed caenagnathoid (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
(NS.32001.077) metatarsal III, metatarsal IV
Comments- These were identified as Elmisaurus in the abstract, but as caenagnathid or oviraptorid in the poster.
Reference- 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.

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
= Avimimidae Kurzanov, 1981
= Kuszholiidae Nessov, 1992
= 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).
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.
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.

unnamed caenagnathid (Gilmore, 1924)
Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- (CMN 8504) dorsal centrum (44 mm), three caudal centra (22-25 mm)
Comments- These 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 known from vertebrae at the time). Currie et al. (1993) noted the caudals belonged to an oviraptorosaur, referring them to Caenagnathus sp.. They may belong to Chirostenotes or Elmisaurus.
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, 1993. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 2255-2272.

unnamed possible caenagnathid (Lapparent, 1960)
Cenomanian, Late Cretaceous
Kem Kem Formation, Morocco
Material- ?(CMN 41800; bone taxon I) distal manual phalanx (Russell, 1996)
(CMN 41820; bone taxon J) manual ungual (81 mm) (Russell, 1996)
(CMN 41977; cast of material in Eberharde private coll.; bone taxon J) manual ungual (189 mm) (Russell, 1996)
?(CMN 50794; bone taxon I) manual phalanx I-1 (250 mm) (Russell, 1996)
?(CMN 50805; bone taxon I) manual phalanx II-? (111 mm) (Russell, 1996)
Albian, Early Cretaceous
Continental Intercalaire, Algeria
?(MNNHN coll.; from Alrar) manual phalanx (300+ mm) (Lapparent, 1960)
(MNNHN coll.; from Dijoua) manual ungual (85 mm) (Lapparent, 1960)
Comments- The MNNHN manual ungual from Dijoua was referred to Carcharodontosaurus saharicus by Lapparent (1960), but matches unguals called bone taxon J by Russell (1996). They are similar to those of caenagnathids. The MNNHN manual phalanx from Alrar was referred to Carcharodontosaurus saharicus by Lapparent (1960), and identified as a metatarsal, but matches phalanges called bone taxon I by Russell (1996).
References- Lapparent, 1960. Les dinosauriens du "Continental intercalaire" du Sahara central. Mem. Soc. Geol. France 88A: 1-57.
Russell, 1996. Isolated dinosaur bones from the Middle Cretaceous of the Tafilalt, Morocco. Bulletin du Muse'um national d'Histoire naturelle (4e se'r.) 18, 349-402.

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. (1993).
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, 1993. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 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., 1993)
Comments- RTMP 81.19.252 and 92.36.53 were referred to Caenagnathus sp. by Currie (1992) and Currie et al. (1993) 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 cranial 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, 1993. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 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, Ishikawa, 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: 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. edited by Matsuoka H., published by Shiramine Village Board of Education, Ishikawa Prefecture, Japan, p. 93-98.
Manabe, Barrett and Isaji, 2000, A refugium for relicts? Nature. 404, 953-954.

undescribed caenagnathid (Csiki and Grigorescu, 2005)
Late Maastrichtian, Late Cretaceous
Densus Ciula Formation, Romania
Material- fragmentary humerus, ulna, metacarpal, several manual phalanges
Reference- Csiki and Grigorescu, 2005. A new theropod from Tustea: are there oviraptorosaurs in the Upper Cretaceous of Europe? Kaupia. 14, 78.

undescribed caenagnathidae (Ryan, Currie and Russell, 2001)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Debasu 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.
References- Ryan, Currie and Russell, 2001. New material of Avimimus portentosus (Theropoda) from the Iren Debasu Formation (Upper Cretaceous) of the Erenhot Region of Inner Mongolia. Journal of Vertebrate Paleontology. 21(3), 95A.
AMNH fossil collections.

Caenagnathasia Currie, Godfrey and Nessov, 1993
C. martinsoni Currie, Godfrey and Nessov, 1993
Late Turonian-Coniacian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (N 401/12457) (adult) anterior dentaries
Paratype- (N 402/12457) (adult) incomplete dentary
Diagnosis- (after Currie et al., 1993; compared to Chirostenotes) fluting on the lingual margin of the occlusal edge not as distinct; first anterior occlusal groove larger; first pair of lateral occusal ridges do not meet ventrally; no second anterior occlusal groove; no tubercles on the midline or at the base of the first lateral occlusal ridge; lateral groove narrower and deeper; foramina on the floor more obvious; vascular grooves less conspicuous on the symphysial shelf; no foramina on the floor of the midline depression.
Reference- Currie, Godfrey and Nessov, 1993. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 2255-2272.

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, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, 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
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.

Kuszholia Nessov, 1992
K. mengi Nessov, 1992
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (PO 4602) (~530 mm) posterior synsacrum (~63 mm)
Paratypes- ?(PO 403) anterior cervical vertebra (16 mm)
?(PO 4623) anterior sacral centra
?(PO coll.) sacral vertebrae (~20-30 mm)
Referred- ?(PO 467) centrum (Nessov, 1995)
?(PO 4826) partial sacrum (Nessov and Panteleev, 1993)
Diagnosis- larger sacral pleurocoels than Chirostenotes or Shixinggia (size-related?).
Comments- None of the characters listed in Nessov's (1992) diagnosis 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.
The holotype (PO 4602) consists of the last two sacral vertebrae and a fragment of the third to last, fused together. The last centrum is 10 mm long and the second to last is 11 mm long. 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 (PO 4623) consisting of the first two centra of another sacrum. The first centrum is 13 mm long, the second is 10 mm and there is a fragment of a third centrum as well. 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. Nessov referred to additional isolated vertebrae virtually identical to these, but 2-2.5 times larger. Finally, he stated strongly pneumatic vertebrae with closed neurocentral sutures (unlike juvenile therizinosaurs or sauropods) could belong to Kuszholia. One is illustrated, which appears to be 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. Nessov and Panteleev (1993) figured and described a partial sacrum they referred to Kuszholia sp.. Nessov (1995) later figured a centrum with a slit-like pleurcoel which he stated was "possibly from bird ?Kuszholia sp. or from a theropod or a segnosaur."
Nessov (1992) erected this taxon 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. 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. While few details are available for Bambiraptor, deinonychosaurs differ from Kuszholia in having a rectangular posterior articular surface to match their proximal caudals. This sacral/caudal morphology is also found in oviraptorids (e.g. Microvenator, Gigantoraptor, Rinchenia, Conchoraptor, "Ingenia"). Yet Neimingosaurus and caenagnathids (e.g. Shixinggia, RTMP 92.36.53) have rounded articular surfaces like Kuszholia. 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 caenagnathids though, including Caenagnathasia from the same formation. In fact, Caenagnathasia would have comparably sized sacrals to Kuszholia if scaled from other caenagnathoids. If they are synonymous, Kuszholia would have priority. 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. Kuszholia is thus provisionally referred to the Caenagnathidae.
As for the other material, the paratype anterior sacrum seems similar enough in size and characteristics to belong to Kuszholia, though the slightly heterocoelous anterior articular surface is odd. Nessov describes it as "slightly
saddle-shaped, with a little hollow, stretching from the top to the bottom along the middle line." Yet heterocoelous sacrals are only known for hesperornithines and neornithines among theropods, the former which are apneumatic and the latter of which lack sacral pleurocoels. Further study of the specimen will be necessary. The referred sacral vertebrae may belong to larger oviraptorosaurs, therizinosaurs or dromaeosaurids, though they are impossible to evaluate without more information. The cervical is of the right size to belong to Kuszholia and matches oviraptorosaurs in general morphology, but could also be from a small troodontid or large bird.
References- 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.
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.
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.

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- (GIN 100/119; incorrectly listed as GIN 940824 by Barsbold et al. 2000) (1.8 m) three cervical vertebrae, ten dorsal vertebrae, ten fragmentary dorsal ribs, several gastralia, five sacral vertebrae, twenty-four caudal vertebrae (516 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- (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), though they inappropriately excluded tyrannosaurids and alvarezsaurids from the analysis a priori.
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.

Similicaudipteryx He, Wang and Zhou, 2008
S. yixianensis He, Wang and Zhou, 2008
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)
Diagnosis- (modified from He et al., 2008) puboilial ratio of 1.46.
Comments- This specimen 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), ilium shaped like Caudipteryx (ambiguous), two large anterior dorsal hypapophyses (present in all 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. In fact, Similicaudipteryx shares large anterior dorsal hypapophyses with caenagnathoids, and a pygostyle with Nomingia specifically (unknown in other caenagnathids).
Reference- Wang, Jones and Evans, 2007. A juvenile anuran from the Lower Cretaceous Jiufotang Formation, Liaoning, China. Cretaceous Research. 28, 235-244.

Caenagnathinae Sternberg, 1940 sensu Paul, 1988
Chirostenotes Gilmore, 1924
pr= Macrophalangia Sternberg, 1932
pr= Caenagnathus Sternberg, 1940
= "Steneodactylus" Gilmore vide Holtz, DML 1998
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
pr= Macrophalangia canadensis Sternberg, 1932
pr= Caenagnathus collinsi Sternberg, 1940
= "Steneodactylus pergracilis" Gilmore vide Holtz, DML 1998
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (CMN 2367) distal metacarpal I, phalanx I-1 (63 mm), manual ungual I (44 mm), distal metacarpal II, phalanx II-1 (65 mm), phalanx II-2 (72 mm), manual ungual II (62 mm), phalanx III-3 (44 mm), manual ungual 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), phalanx II-1 (78 mm), phalanx II-2 (63 mm), pedal ungual II (60 mm), partial metatarsal III (230 mm), phalanx III-1 (75 mm), phalanx III-2 (52 mm), phalanx III-3 (58 mm), pedal ungual III (60 mm), metatarsal IV (212 mm), 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)
(RTMP 79.14.499) manual ungual II (83 mm) (Currie and Russell, 1988)
?(RTMP 79.20.1) (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)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
?(CMN 9570) metatarsal II (258 mm) (Russell, 1984)
?(ROM 43250) (adult) maxilla, palatine, braincase, anterior cervical vertebra, cervical vertebra (85 mm), cervical vertebra, 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)
Diagnosis- (after Currie, 1989) proximal end of metatarsal III diamond-shaped; metatarsals II and IV much shorter than III (<93%).
(after Currie et al., 1993) 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.
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 basically proven by the undescribed CMNH specimens. 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 the undescribed CMNH specimens have Caenagnathus-like mandibles on Chirostenotes-like skeletons, disproving this notion.
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. (1993) 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 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. This is followed here.
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. Thus it is assigned to Chirostenotes pergracilis here, as it was by Currie (2005) 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. These may end up being a distinct species however, perhaps related to the Hell Creek species or Hagryphus, once more comparable material (mandibles, metatarsi, etc.) is described.
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, 1993. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 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.
C? sp. nov. (Currie, Godfrey and Nessov, 1993)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Material- (BHM 2033) posterior mandible (Currie et al., 1993)
(CMNH coll.) skull fragments, twelve cervical vertebrae, ten cervical ribs, ten dorsal vertebrae, eighteen dorsal ribs, six sacral vertebrae, twelve caudal vertebrae, five chevrons, sternal plates, ilia, pubes, ischia, femora, tibiae, astragali, calcanea, metatarsal V, two phalanges, four unguals (Triebold, Nuss and Nuss, 2000)
(CMNH coll.) incomplete skull, mandible, five cervical vertebrae, four cervical ribs, ten dorsal ribs, gastralia, seven caudal vertebrae, five chevrons, scapulocoracoids, partial sternal plate, humerus, radius, ulna, femora, tibiae, astragali, calcanea, metatarsal I, partial metatarsal V, eight phalanges, four unguals (Triebold, Nuss and Nuss, 2000)
? metatarsal (anonymous, 1997)
Comments- Currie et al. (1993) 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. Triebold et al. (2000) reported two new large oviraptorosaur specimens, identifying them as oviraptorids. Examination of the material indicates they are caenagnathids however. Although privately held for several years, these specimens are now under study at the CMNH. It's currently unknown if they share BHM 2033's mandibular morphology, but they are provisionally assigned to the same species based on their provenence and very large size. While these are the most complete caenagnathid specimens known, they lack metatarsals II-IV which are key in distinguishing Elmisaurus from Chirostenotes. Anonymous (1997) reported a Chirostenotes metatarsal associated with Tyrannosaurus specimen FMNH PR2081, which is tentatively assigned to the same species based on provenence.
References- Currie, Godfrey and Nessov, 1993. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 2255-2272.
Anonymous, 1997. Tyrannosaurus rex; A Highly Important and Virtually Complete Fossil Skeleton. Sotheby’s. 56pp.
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.

Elmisaurinae Osmolska, 1981 sensu Currie, 2001
Diagnosis- seven sacral vertebrae; preacetabular process short (<64% of postacetabular process); preacetabular process not hooked ventrally; fourth trochanter present; arctometatarsus; proximolateral process on distal tarsal IV; proximally fused tarsometatarsus.
Comments- The first four characters are shared by Avimimus and Shixinggia, while the second three are shared by Avimimus and Elmisaurus. Unfortunately, Shixinggia and Elmisaurus are not comparable using their described materials. It is possible they are synonymous.

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 caenagnahid in my modified version of Senter's (2007) analysis.
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.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.

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, nine dorsal vertebrae, two sacral ribs, partial scapulocoracoid, humerus (95 mm), proximal ulna, proximal carpometacarpus, partial ilia, pubes, partial ischium, femur (188 mm), tibiotarsus (257 mm), fibula, tarsometatarsus (153 mm), phalanx II-1, phalanx II-2, pedal ungual II, phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
Paratype- (PIN 3906/1) postcranial fragments including ilial fragment, proximal ischium
Referred- (IGM coll.) incomplete skeleton including incomplete skull with premaxilla, dorsal ribs, articulated caudal vertebrae, pectoral girdle, scapulacoracoid, humerus, radius, ulna, carpometacarpus, 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, 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 Russian]. 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.
A. sp. (Ryan, Currie and Russell, 2001)
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Debasu 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 (2008) 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 Debasu 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, 2008. 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. doi: 10.1016/j.cretres.2008.07.005
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)
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.).
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 Debasu 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.

Elmisaurus Osmolska, 1981
Diagnosis- (modified from Currie, 1989) tarsometatarsal fusion; posterior surface of metatarsus deeply concave; metatarsals II and IV subequal in length to III (>93%); distal end of metatarsal II curves anteriorly away from the longitudinal axis of the metatarsus; proximal end of metatarsal III triangular; proximolateral process on metatarsal IV.
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) metacarpal I (45 mm), phalanx I-1 (65 mm), incomplete metacarpal II (~63 mm), phalanx II-1 (66 mm), phalanx II-2 (66 mm), manual ungual II (44 mm), phalanx III-1 (30 mm), phalanx III-2 (30 mm), phalanx III-3 (43 mm), proximal manual ungual, pedal phalanx I-1 (26 mm), distal metatarsal II, phalanx II-1 (44 mm), partial phalanx II-2 (33 mm), distal metatarsal III (~151 mm), phalanx III-1 (43 mm), phalanx III-2 (32 mm), proximal phalanx III-3, distal metatarsal IV, two fragmentary phalanges IV-?, two fragmentary pedal unguals (II and IV), indeterminate fragments of long bones
Referred- (PJC. 2001.8) proximal tarsometarsus (Currie, 2001)
eight dorsal vertebrae, manual phalanx III-2, phalanx III-3, manual ungual III, partial femur, tibiae, partial metatarsal III, partial metatarsal IV, pedal phalanx I-1, pedal ungual I, pedal ungual II (Currie, 2001)
metatarsal IV (Currie, 2001)
? femur (Currie, 2001)
? ventral skull (www.paleofile.com)
Diagnosis- m. tibialis cranialis tubercle on the dorsal surfaces of metatarsals II–IV.
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 recently, 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 except for the m. tibialis cranialis tubercle. 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. Further comparison with Chirostenotes may be possible when new Elmisaurus remains mentioned by Currie (2001, 2002) are described. 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.
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
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., 1993)
(RTMP 82.16.6) tarsometatarsus (Snively, 2000)
(RTMP 82.39.4) proximal tarsometatarsus (Currie, 1989)
?(RTMP 90.56.6) dentaries (Currie et al., 1993)
?(RTMP 91.144.1) incomplete dentaries (Currie et al., 1993)
?(RTMP 92.36.390) incomplete dentaries (Currie et al., 1993)
?(RTMP 92.40.44) anterior dentaries (Currie et al., 1993)
(RTMP 96.12.141) (Currie, 2005)
?(RTMP 2001.12.12) mandible (Currie, 2005)
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)
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., 1993; 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.
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. (1993) 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.
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.
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, 1993. New caenagnathid (Dinosauria: Theropoda) specimens from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth Sciences. 30, 2255-2272.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Unpublished Masters Thesis. 273 pp.
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.
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.

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.

Gigantoraptor Xu, Tan, Wang, Zhao and Tan, 2007
G. erlianensis Xu, Tan, Wang, Zhao and Tan, 2007
Late Campanian-Early Maastrichtian, Late Cretaceous
Iren Debasu Formation, Inner Mongolia, China
Holotype- (LH V0011) (8 m; 1.4 tons; 11 year old adult) mandibles, posterior cervical neural arch, eight partial dorsal vertebrae, several dorsal ribs, gastralia, sacrum, twenty-seven caudal vertebrae, fourteen chevrons, incomplete scapula, furcula, sternum, humerus, 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.
Reference- Xu, Tan, Wang, Zhao and Tan, 2007. A gigantic bird-like dinosaur from the Late Cretaceous of China. Nature. 844-847.

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 et al.'s (2004) topology, Oviraptorinae would include all oviraptorids except for Khaan. In Lu's (2004) based on Maryanska et al.'s (2002) matrix, Shixinggia and all caenagnathoids are oviraptorines. In his tree based on the TWG's 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 oviraptorids except Microvenator and Caudipteryx. The phylogeny on this website is similar except Caudipteryx is placed more basally. Oviraptorinae as used here is therefore equivalent to the Oviraptoridae of most authors. 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.
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.
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.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.

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 (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. 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.

unnamed Oviraptoridae (Maryanska and Osmolska, 1997)
Late Campanian, Late Cretaceous
Red Beds of Khermeen Tsav, Mongolia
Material- (AMNH coll.) skull (Norell, 1992)
(GIN 100/30A) fragmentary skull (Maryanska and Osmolska, 1997)
(GIN coll.; "GIN A") quadrates (Maryanska and Osmolska, 1997)
(GIN coll.; "GIN B") partial quadratojugal, distal quadrate (Maryanska and Osmolska, 1997)
(ZPAL coll.) skull, postcranial fragments (Osmolska, 1976)
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 GIN material represents crestless oviraptorids, perhaps Conchoraptor or "Ingenia". The ZPAL material may be MgD-I/101 (listed on the ZPAL website as oviraptorid), MgD-I/100 or MgD-I/106 (both listed as Conchoraptor by Maryanska et al., 2002). Osmolska (1976) listed it as Oviraptor sp., but this was before the diversity of oviraptorid genera was known.
References- Osmolska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
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.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta Palaeontologica Polonica. 47 (1), 97-116.
http://www.paleo.pan.pl/collect.htm#Mon-reptilia

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 Oviraptoridae (Norton, DML 2000)
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia
Material- (IGM 100/1009) (juvenile) skull
(IGM 100/1018) (adult) skeleton, nest
Comments- Norton (DML, 2000) noted these specimens were discovered in 1993 and on display at the AMNH Fighting Dinosaurs exhibit. They are probably Citipati or Khaan based on the fact they were discovered at Ukhaa Tolgod.
Reference- http://dml.cmnh.org/2000Jun/msg00082.html

undescribed Oviraptoridae (Weishampel, Fastovsky, Watabe, Barsbold and Tsogtbaatar, 2000)
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Material- (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)
three partial embryos, eggs (Weishampel, Fastovsky, Watabe, Barsbold and Tsogtbaatar, 2000)
partial skeleton (Currie, 2001)
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 and neither Rinchenia nor Nemegtomaia preserve pes.
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.

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).
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.

unnamed oviraptorid (Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005)
Campanian, Late Cretaceous
Yuanpu (=Nanxiong) Formation, Jiangxi, China
Material- (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)
Comments- The eggs are preserved inside the pelvic cavity and resemble Macroolithus yaotunensis and M. rugustus.
Reference- Sato, Cheng, Wu, Zelenitsky and Hsiao, 2005. A pair of shelled eggs inside a female dinosaur. Science. 308, 375.

unnamed oviraptorid (ZPAL online)
Late Campanian-Maastrichtian, Late Cretaceous
Red Beds of Khermeen Tsav or Nemegt Formation, Mongolia
Material- (ZPAL MgD-I/101) (ZPAL online)
Comments- This specimen may be the skull and/or postcranial fragments listed by Osmolska (1976) as Oviraptor sp.. As no further information is available, it may be Rinchenia, Nemegtomaia, Conchoraptor, "Ingenia" or a new taxon.
References- Osmolska, 1976. New light on skull anatomy and systematic position of Oviraptor. Nature. 262, 683-684.
http://www.paleo.pan.pl/collect.htm#Mon-reptilia

undescribed Oviraptoridae (online)
Late Cretaceous(?)
Asia(?)
Material- skull, mandible, five anterior cervical vertebrae
skull, mandibles
Comments- 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.

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) (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.
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, 1-27.

Oviraptor Osborn, 1924
= "Fenestrosaurus" Osborn, 1924
O. philoceratops Osborn, 1924
= "Fenestrosaurus philoceratops" Osborn, 1924
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 (78 mm), carpometacarpus (107 mm), phalanx II-1 (55 mm), phalanx II-2 (68 mm), manual ungual II, metacarpal III (99 mm), phalanx III-1 (37 mm), phalanx III-2 (29 mm), phalanx III-3 (28 mm), manual ungual III, partial ilium
Referred- (AMNH 6508) ~15 eggs, nest
Comments- The first oviraptorid discovered, this taxon was originally referred to the Ornithomimidae by Osborn (1924). 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. Further study is necessary. Morell (1997) labeled IGM 100/1002 Oviraptor before it was made a paratype of Khaan by Clark et al. (2001). 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 (1924), 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.
References- Osborn, 1924. The discovery of an unknown continent. Natural History. 24(2), 133-149.
Osborn, 1924. Three new Theropoda, Protoceratops zone, central Mongolia. American Museum Novitates. 144, 1-12.
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.

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) complete skeleton including skull, sclerotic ossicles, mandibles, hyoids, cervical vertebrae, cervical ribs and pectoral girdle
Referred- (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, proximal radius, proximal ulna, partial ilia, femur, tibia, fibula, astragali, partial metatarsal II, partial metatarsal III, partial metatarsal IV, pedal phalanges, nest, eggshell fragments (Clark et al., 2001)
?(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 (227 mm; one proximal), radii (201 mm; one distal), ulnae (222 mm; one distal), proximal carpal, semilunate carpal, metacarpal I, phalanx I-1 (81 mm), manual ungual I, metacarpal II (100 mm), phalanx II-1 (55 mm), phalanx II-2 (55 mm), manual ungual II, metacarpal III, phalanx III-1 (34 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, 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, nest, fifteen eggs (Clark et al., 1999)
?(IGM 100/1004; Big Auntie) twelve cervical vertebrae, cervical ribs, several dorsal vertebrae, eight dorsal ribs, four uncinate processes, partial sacrum, scapulae, partial coracoid, 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)
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". It 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.
The holotype of the genus, a virtually complete specimen, was initially identified as Oviraptor philoceratops (Webster, 1996) until 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 undescribed and unillustrated.
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.
Codd, Manning, Norell and Perry, 2007. Avian-like breathing mechanics in maniraptoran dinosaurs. Proc. R. Soc. B (doi:10.1098/rspb.2007.1233).
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS ONE. 4(10), e7390. doi:10.1371/journal.pone.0007390
C. sp. nov. (Barsbold, 1981)
Late Campanian, Late Cretaceous
Djadochta Formation, Mongolia
Material- (IGM 100/42; Zamyn Khondt oviraptorid) incomplete skull (175, 197 mm), mandibles (153, 156 mm), fiftteen cervical vertebrae, cervical ribs, seven dorsal vertebrae, three dorsal centra, eighteen dorsal ribs, sacrum, thirty caudal vertebrae, twenty-three chevrons, scapulacoracoids, furcula, sternal plates (79-84 mm), humeri, radius, ulna, phalanx I-1 (87 mm), manual ungual I, phalanx II-1 (53 mm), phalanx II-2 (66 mm), manual ungual II, phalanx III-1 (38 mm), phalanx III-2 (34 mm), phalanx III-3 (51 mm), manual ungual III, ilia, pubes, ischia, femora, tibiae, fibula, astragalus, calcaneum, metatarsal I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (168 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
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. 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.
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.

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
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (GIN 100/2112) skull, mandibles, 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, proximal pubes, proximal ischia, proximal femur
Diagnosis- (modified from Lu et al., 2004) prefrontal present; quadrate articulates with quadratojugal with a convex condyle.
Comments- This specimen 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.
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. Bull. Natn. Sci. Mus.. Tokyo, Ser. 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.

"Ingeniinae" Barsbold, 1981
Definition- ("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 et al.'s (2004) topology, Nomingia and Rinchenia would also be "ingeniines", while Oviraptor, Citipati, Nemegtomaia, Khaan and Heyuannia would not. 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 (2002) matrix, all oviraptorids except Khaan and Oviraptor are "ingeniines". Senter's (2007) phylogeny finds Conchoraptor, Khaan, Heyuannia and "Ingenia" to be "ingeniines"..
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.
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.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology.

Conchoraptor Barsbold, 1986
C. gracilis Barsbold, 1986
Late Campanian, Late Cretaceous
Red Beds of Khermeen Tsav, Mongolia
Holotype- (IGM 100/20) skull (99 mm), (?)mandible, postcrania including metacarpal I (26 mm), phalanx I-1 (36 mm), manual ungual I, metacarpal II (40 mm), distal phalanx II-1, phalanx II-2 (27 mm), manual ungual II, metacarpal III (41 mm), phalanx III-1 (19 mm), phalanx III-2 (19 mm), phalanx III-3 (25 mm), manual ungual III, (?)metatarsal II, (?)metatarsal III, (?)metatarsal IV, (?)two pedal unguals, (?)metatarsal V
Referred- ?(IGM 100/21) mandible (93 mm) (Barsbold, 1976)
?(IGM 100/36) material including skull, furcula (Barsbold 1981)
?(IGM 100/38) (Maryanska et al., 2002)
?(IGM 100/39) (Maryanska et al., 2002)
?(IGM 100/46) (Maryanska et al., 2002)
?(IGM 100/47) (Maryanska et al., 2002)
?(ZPAL MgD-I/95) incomplete skull (98 mm), mandibles (86 mm), metatarsus (141 mm) (Osmolska, 1976)
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 110/18) material including distal caudal vertebrae, pubis and ischium (Lu, 2004)
?(IGM 110/19) material including thirty-two caudal vertebrae (Barsbold et al., 2000)
?(IGM coll.) (multiple specimens) (Maryanska et al., 2002)
?(PIN coll.) skull, mandibles (Maryanska et al., 2002)
?(ZPAL MgD-I/100) (Maryanska et al., 2002)
?(ZPAL MgD-I/106) (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 and manus. Barsbold et al. (1990) illustrated the skull in dorsal view and also a metatarsus and two pedal unguals, which may belong to the holotype as well (no specimen number was listed). 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.
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). 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. Erickson et al. (2009) examined the histology of IGM 97/212, which they referred to Conchoraptor gracilis.
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. Similarly, the matrices of Norell et al. (2001), Maryanska et al. (2002) and Lu (2004) 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.
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.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian physiology inherited by birds? Reconciling slow growth in Archaeopteryx. PLoS ONE. 4(10), e7390. doi:10.1371/journal.pone.0007390
http://www.oucom.ohiou.edu/dbms-witmer/dinoskulls02.htm
http://www.paleo.pan.pl/collect.htm#Mon-reptilia

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, mandible, hyoids, twelve cervical vertebrae, several dorsal vertebrae, dorsal ribs, uncinate process, gastralia, twenty-seven caudal vertebrae, chevrons, scapulacoracoids, furcula, sternal plates, humeri, radii, ulnae, two carpals, 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, pubis, ischium, femur, tibia, fibula, 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
Paratypes- (IGM 100/973) skull (135 mm), mandible, atlas, axis, incomplete skeleton including femur (374 mm), tibia (453 mm) and metatarsus (212 mm)
(IGM 100/1002) incomplete skull, mandible, hyoids, twelve cervical vertebrae, ten dorsal vertebrae, dorsal ribs, gastralia, sacrum, six caudal vertebrae, four chevrons, scapulocoracoids, furcula, sternal plates, sternal ribs, humerus, radii, ulnae, radiale, 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, ilium, pubis, ischium, femur (193 mm), tibia (231 mm), fibula, astragalus, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (108 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
Comments- IGM 100/973 was 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 100/1002 was photographed in Morell (1997) and labeled Oviraptor. Clark et al. (2001) made these paratypes of their new taxon Khaan mckennai, using IGM 100/1127 as the holotype. IGM 100/1002 and 100/1127 were found in close proximity to each other, and have been called "Romeo and Juliet." They have been described extremely briefly and published photos are small, but the data matrices of Lu (2004), Osmolska et al. (2004) and Senter (2007) are informative.
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.

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
Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (GI 100/32A) skull (148 mm), mandibles (144 mm), thirteen cervical vertebrae, cervical ribs, dorsal vertebrae, six or seven sacral vertebrae, twenty-three caudal vertebrae, scapulocoracoids, furcula, sternal plate, humerus, radius, ulna, manual ungual, ilium, femur
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 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.
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 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.
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.

Heyuannia Lu, 2002
H. huangi Lu, 2002
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, incomplete coracoid, furcula, partial sternum, sternal ribs, humerus, radius, ulna, radiale, semilunate carpal, metacarpal I, 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, 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, incomplete radius, ulna, radiale, ulnare, semilunate carpal, metacarpal I, phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, metacarpal III (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). 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".
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. pp 256-273.

"Ingenia" Barsbold, 1981 preoccupied Gerlach, 1957
"I." yanshini Barsbold, 1981
= Oviraptor yanshini Paul, 1988
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, twelve chevrons, scapulacoracoids, furcula, sternum (72 mm), humeri, radii, ulnae, 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, pubes, ischia, femora (228 mm), tibiae (281 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, postcrania including eight sacral vertebrae, ilium, ischium, femur, tibia, fibula
?(IGM 100/32) postcrania including cervical vertebrae, twenty-seven caudal vertebrae, semilunate carpal, metacarpals, metatarsus
?(IGM 100/33) postcrania including nineteen caudal vertebrae, sternal plates (60 mm), furcula, ilium, pubis, ischium, femur, tibia, fibula
Late Cretaceous
Mongolia
Referred- ?(FDPM-V6240) skull, skeleton (Azuma, 2005)
?(IGM 100/34) material including metatarsus (Snively, 2000)
?(IGM 100/35) (Maryanska et al., 2002)
?(IGM 100/80-1) material including skull, eleven cervical vertebrae (Lu, 2004)
Comments- The genus Ingenia is preoccupied by a nematode, as noted by Taylor (DML, 2004). Barsbold is apparently aware of the situation, though no replacement name has been suggested so far.
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 sme 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.
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. 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 specimen illustrated by Barsbold et al. (1990), 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.
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.
http://www.oucom.ohiou.edu/dbms-witmer/dinoskulls02.htm