Oviraptorosauria Barsbold, 1976
Definition- (Oviraptor philoceratops <- Therizinosaurus
cheloniformis, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019; modified from Hu, Hou, Zhang and
Xu, 2009)
Other definitions- (Oviraptor philoceratops <- Passer domesticus)
(Maryanska, Osmolska and Wolsan, 2002; modified from Currie and Padian in Barsbold, 1997)
(Oviraptor philoceratops + Chirostenotes pergracilis) (modified
from Padian, Hutchinson and Holtz, 1999)
(Oviraptor philoceratops <- Tyrannosaurus rex, Ornithomimus edmontonicus,
Therizinosaurus cheloniformis, Troodon formosus, Passer domesticus)
(Sereno, online 2005)
= Caenagnathiformes Sternberg, 1940
Definition- (Caenagnathus collinsi <- Passer domesticus)
(Martyniuk, 2012)
= Avimimiformes Chatterjee, 1991
= Oviraptorosauria sensu Currie and Padian in Barsbold, 1997
Definition- (Oviraptor philoceratops <- Passer domesticus)
(modified)
= "Oviraptoriformes" Sereno, online 2005
Definition- (Oviraptor philoceratops <- Passer domesticus)
(Sereno, online 2005)
= Oviraptorosauria sensu Sereno, online 2005
Definition- (Oviraptor philoceratops <- Tyrannosaurus rex, Ornithomimus
edmontonicus, Therizinosaurus cheloniformis, Troodon formosus,
Passer domesticus)
= Oviraptoriformes Benson, 2008
Comments- Osborn (1924) described Oviraptor as an ornithomimid based on the toothless jaws, while Caenagnathus
was originally (Sternberg, 1940) assigned to its own order
Caenagnathiformes believed to be closer to crown Aves than
hesperornithines or Ichthyornis.
Cracraft (1971) further placed caenagnathiforms closest to
galloanserines based on the sliding mandibular glenoid. Osmolska
(1976) first noticed the similarity between her 'Oviraptor sp.' (actually Conchoraptor) and Caenagnathus,
assigning both to Caenagnathidae within Theropoda. "The
caenagnathids may represent a Cretaceous continuation of the same
dinosaur lineage that gave rise to the birds", she said, and "a new
suprafamilial taxon will have to be established as soon as the new
material of Oviraptor has
been described." This was done that same year when Barsbold
(1976) erected Oviraptorosauria (incorrectly attributed in his paper to
Barsbold, 1974) for oviraptorids. He wrote "the difference in
the type of [edentulous] structure itself and undoubtedly the ecology
of the oviraptorids and the ornithomimids are so great that their
separation is presently fully valid." Barsbold viewed
oviraptorosaurs as potentially diverging early from 'coelurosaurs'
(coelophysoids and coelurid/compsognathid-grade taxa) due to supposedly
primitive characters like the 'interclavicle' (actually a furcula
unrecognized in other theropods at the time). By 1981 Barsbold
had read Osmolska's paper and agreed with her the genera were closely
related but wrote "considerable differences in the structure of the
lower jaw of Caenagnathus and Oviraptor
make it necessary to assign them to different families, which enter the
infra-order Oviraptorosauria." Currie and Russell (1988) were the
first authors to recognize Microvenator
and the 'elmisaurids' (caenagnathid manus and pedes) belonged to
Oviraptorosauria, but in 1994 Russell and Dong tried to expand the
concept to include therizinosaurs, troodontids and ornithomimosaurs as
well. Besides a few suggestions therizinosaurs might belong, this
more inclusive Oviraptorosauria was not followed, and with the addition
of basal taxa like Caudipteryx
the group's content remains similar to this day. Note references
to an "Oviraptorosauridae" as in Norell et al. (2000) are in error as
there is no genus "Oviraptorosaurus" to base it on.
Avimimus was described in 1981
as closer to Aves than other theropods known at the time, then in the
1990s was associated with arctometatarsalian coelurosaurs. Currie
(1989) was prescient in saying "similarities in the foot suggest that Avimimus may have been derived from elmisaurid dinosaurs" but Avimimus
was first assigned to Oviraptorosauria in the first TWiG analysis
(Norell et al., 2001). Chatterjee (1991) created a classification
for Mesozoic birds where "each Mesozoic avian genus has been assigned
to a corresponding higher level taxon or order" and so proposed an
Avimimiformes for Avimimus, which has been ignored since then for being monotypic.
Sereno (online 2005) proposed the name Oviraptoriformes "to accommodate
clades most closely related to Oviraptorosauria", with the phylogenetic
definition "the most inclusive clade containing Oviraptor philoceratops Osborn 1924 but not Passer domesticus
(Linnaeus 1758)." Yet online citations do not count for
nomenclature. Benson (2008) is the first author to publish the name
Oviraptoriformes, though he did such undefined in a phylogram without
comment or definition. Holtz (2012) stated "Therizinosauria and Oviraptorosauria may together form
a clade Oviraptoriformes" but again provided no definition.
Martyniuk (2012) created a stem-based definition for Sternberg's (1940)
Caenagnathiformes, generally unused since the 1970s, as all taxa closer to Caenagnathus collinsi than to Passer domesticus,
which seemed more like an attempt to replace the newer Oviraptorosauria
since he placed therizinosaurs further from birds. Yet it would
work for a topology in which therizinosaurs were closer to
oviraptorosaurs than to birds, is published with a proper definition
and is an older name than Oviraptoriformes.
Paul (2016) used the informal term oviraptorosauriforms for
"omnivoropterygids, oviraptorosaurs, and avians and their common
ancestor, operative only if three groups form a clade that excludes all
other dinosaurs." This would imply an 'Oviraptorosauriformes'
with a partial definition (Omnivoropteryx sinousaorum + Oviraptor philoceratops + Passer domesticus,
- XXX), but such a clade is poorly supported, taking 37 extra steps in
Hartman et al.'s TWiG maniraptoromorph analysis for instance.
Oviraptorosauria defined-
The first proposed phylogenetic definition of Oviraptorosauria was
written in "A Note Added by the Editors" in Barsbold's (1997)
encyclopedia entry, so is properly cited Currie and Padian in Barsbold,
1997. Their definition was "Oviraptoridae and all taxa closer to Oviraptor
than to birds" which would have included therizinosaurs in most
topologies over the following decade. Padian et al. (1999) noted
this was inconsistant with common usage and so proposed a node-based
definition instead- "Oviraptor and Chirostenotes (=Caenagnathus) and all descendants of their most recent common ancestor." However in the early 2000s taxa like Caudipteryx and Incisivosaurus
were consistantly recognized as related to but outside this node and
called oviraptorosaurs, so Hu et al. (2009) finally proposed a
stem-based definition excluding therizinosaurs.
Ex-oviraptorosaurs-
A partial sacrum and ilium (SMNS 58023) described by Frey and Martill
(1995) as a possible oviraptorosaur, but was reidentified as a
megaraptoran (Aranciaga Rolando et al., 2018).
A femur (ZIN PH 1/13) identified as oviraptorosaurian
or ornithomimid by Nessov (1995) is ornithomimid (Averianov et al., 2003).
Currie et al. (1996) identified a surangular (NMV P186386) and dorsal (NMV P186302)
from the Eumeralla Formation of Victoria as oviraptorosaurian, but these have
been placed more ambiguously as theropod (Agnolin et al., 2010) and maniraptoran
(Benson et al., 2012) lately.
A cervical vertebra (MACN 622) discovered with the Noasaurus 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
holotype individual.
Paul (2010) placed Sapeornis
in Oviraptorosauria, but that takes 37 more steps in Hartman et al.'s
(2019) matrix so is highly unlikely. Paul also placed Epidexipteryx there (but not Scansoriopteryx), and Brusatte et al. (2014) recovered an Epidexipteryx plus Pedopenna
pair as the basalmost oviraptorosaurs. Cau (2018) recovered all
scansoriopterygids as oviraptorosaurs in his Bayesian analysis.
In a modified version of the Hartman et al. matrix, scansoriopterygids
move to Oviraptorosauria in 9 steps, but getting Epidexipteryx or Pedopenna there without Scansoriopteryx takes 13 steps and 4 steps respectively.
References- Osborn, 1924. Three new Theropoda, Protoceratops zone, central Mongolia.
American Museum Novitates. 144, 1-12.
Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of
Paleontology. 14(1), 81-85.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism
to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Barsbold, 1976. The evolution and systematics of late Mesozoic
carnivorous dinosaurs. In Kramarenko, Luvsandansan, Voronin, Barsbold,
Rozhdestvensky, Trofimov and Reshetov (Eds.). Paleontology and
Biostratigraphy of Mongolia. The Joint Soviet-Mongolian Paleontological
Expedition, Transactions. 3, 68-75.
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.
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(7), 972-986.
Currie, 1989. Theropod dinosaurs of the Cretaceous. In Padian and Chure (Eds.). The Age of Dinosaurs.
Short Courses in Paleontology. 2, 113-120.
Chatterjee, 1991. Cranial anatomy and relationships of a new Triassic bird from
Texas. Philosophical Transactions of the Royal Society of London Series B. 332(1265),
277-342.
Russell and Dong, 1994. The affinities of a new theropod
from the Alxa Desert, Inner Mongolia, People’s Republic of China. Canadian
Journal of Earth Sciences. 30(10), 2107-2127.
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.
Nessov, 1995. Dinosaurs of nothern Eurasia: new data about
assemblages, ecology, and paleobiogeography. Institute for Scientific Research
on the Earth's Crust, St. Petersburg State University, St. Petersburg. 1-156.
Currie, Vickers-Rich and Rich, 1996. Possible oviraptorosaur (Theropoda, Dinosauria)
specimens from the Early Cretaceous Otway Group of Dinosaur Cove, Australia.
Alcheringa. 20(1-2), 73-79.
Barsbold, 1997. Oviraptorosauria. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. 505-509.
Frankfurt and Chiappe, 1999. A possible oviraptorosaur from the Late Cretaceous
of northwestern Argentina. Journal of Vertebrate Paleontology. 19(1), 101-105.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions
and nomenclature of the major taxonomic categories of the carnivorous Dinosauria
(Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Norell, Makovicky and Clark, 2000. A new troodontid theropod from Ukhaa Tolgod,
Mongolia. Journal of Vertebrate Paleontology. 20(1), 7-11.
Norell, Clark and Makovicky, 2001. Phylogenetic relationships among coelurosaurian
theropods. In Gauthier and Gall (eds.). New Perspectives on the Origin and Early
Evolution of Birds: Proceedings of the International Symposium in Honor of John
H. Ostrom. 49-67.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47(1), 97-116.
Averianov, Starkov and Skutschas, 2003. Dinosaurs from the Early Cretaceous
Murtoi Formation in Buryatia, eastern Russia. Journal of Vertebrate Paleontology.
23(3), 586-594.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php
[version 1.0, 2005 November 7]
Agnolin and Martinelli, 2007. Did oviraptorosaurs (Dinosauria; Theropoda) inhabit
Argentina? Cretaceous Research. 28(5), 785-790.
Balanoff, Bever and Rowe, 2008. The endocranial morphology of oviraptorosaurs
and a reinterpretation of their encephalization quotients. Journal of Vertebrate
Paleontology. 28(3), 47A.
Benson, 2008. New information on Stokesosaurus, a tyrannosauroid (Dinosauria:
Theropoda) from North America and the United Kingdom. Journal of Vertebrate
Paleontology. 28(3), 732-750.
Hu, Hou, Zhang and Xu, 2009. A pre-Archaeopteryx troodontid theropod
from China with long feathers on the metatarsus. Nature. 461, 640-643.
Agnolin, Ezcurra, Pais and Salisbury, 2010. A reappraisal of the Cretaceous
non-avian dinosaur faunas from Australia and New Zealand: Evidence for their
Gondwanan affinities. Journal of Systematic Palaeontology. 8(2), 257-300.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution.
PhD thesis. Columbia University. 522 pp.
Persons, Currie and Norell, 2011. Shake your feathers: The flamboyant, athletic,
and possibly flirtatious caudal morphology of oviraptorosaurs. Journal of Vertebrate
Paleontology. Program and Abstracts 2011, 174.
Benson, Rich, Vickers-Rich and Hall, 2012. Theropod fauna from southern Australia
indicates high polor diversity and climate-driven dinosaur provinciality. PLOS
One. 7(5), e37122.
Holtz, 2012. Theropods. In Brett-Surman, Holtz and Farlow (Eds.). The
Complete Dinosaur. Second edition. Indiana University Press. 347-378.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Vernon, New Jersey. Pan Aves. 189 pp.
Balanoff, Bever and Norell, 2013. The relationships of oviraptorosaurian dinosaurs
and endocranial evolution along a morphologically bizarre lineage. Journal of
Vertebrate Paleontology. Program and Abstracts 2013, 81.
Pittman and Hutchinson, 2013. The evolution of tail joint stiffness in oviraptorosaur
dinosaurs and its consequences for tail function. Journal of Vertebrate Paleontology.
Program and Abstracts 2013, 191.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Tanaka, Zelenitsky, Lu, Yi, Pu, Chang, Xu and Li, 2014. Nest type and incubation
behavior in oviraptorosaurs in relation to body size. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 238-239.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd edition. Princeton University Press.
360 pp.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25. DOI: 10.4435/BSPI.2018.01
Aranciaga Rolando, Brisson Egli, Sales, Martinelli,
Canale and Ezcurra, 2018 (online 2017). A supposed Gondwanan oviraptorosaur from the
Albian of Brazil represents the oldest South American megaraptoran.
Cretaceous Research. 84, 107-119.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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. Alberta Palaeontological Society, sixth annual symposium. 8-12.
undescribed Oviraptorosauria (Watabe and Suzuki, 2000)
Late Campanian, Late Cretaceous
Dzamin Khond, Djadochta Formation, Mongolia
Material- elements
Comments- Watabe and Suzuki (2000) state "isolated bones of oviraptorosauria" were found in 1993, 1994 and/or 1998.
Reference- Watabe and Suzuki,
2000. Cretaceous fossil localities and a list of fossils collected by
the Hayashibara Museum of Natural Sciences and Mongolian
Paleontological Center Joint Paleontological Expedition (JMJPE) from
1993 through 1998. Hayashibara Museum of Natural Sciences Research
Bulletin. 1, 99-108.
undescribed Oviraptorosauria (Suzuki and Watabe, 2000)
Early Maastrichtian, Late Cretaceous
Altan Uul, Nemegt Formation, Mongolia
Material- ?(IGM coll.; 060908 AU-IV BON) partial skeleton
including pelvic elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and
Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
(IGM coll.; 980803 BgT NAR) dorsal vertebrae, caudal vertebrae, hindlimb elements (Suzuki and Watabe, 2000)
(IGM coll.; 060811 BgT-II KHTB)
partial skeleton (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(uncollected?) elements
Early Maastrichtian?, Late Cretaceous
Khermeen Tsav, Nemegt Formation?, Mongolia
(IGM coll.; 060904 KmT KHTB Ovi) partial skull, dorsal vertebrae,
sternal, forelimb, pelvis, hindlimb (Matsumoto, Hashimoto, Sonoda,
Fujiyama, Mifune, Kawahara and Saneyoshi,
2010)
Early Maastrichtian, Late Cretaceous
Upper White Beds of Khermeen Tsav, Nemegt Formation, Mongolia
(IGM coll.; 060906 KmT KHTB) (four individuals)
Early Maastrichtian, Late Cretaceous
Tsagaan Khushuu, Nemegt Formation, Mongolia
(uncollected?) elements (Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010)
Comments- These are field numbers for a joint HMNS-IGM collection.
980803 BgT NAR was found in 1994 at Bugin Tsav with a small Tarbosaurus specimen, preserves cervicodorsal hypapophyses and was at the IGM as of 2000 (Suzuki and Watabe, 2000).
Matsumoto et al. (2010) list 060904 KmT KHTB Ovi as Oviraptorosauria
found on September 4 2004, but this would indicate the field number
would be 040904 and be from Cenozoic locality Baga Dzan. It's
more likely this was found on September 4 2006. Prepared contents
are listed as "dorsal vertebrae, pelvic part and hindlimb" and "part of
skull, forelimb and sternal."
Watabe et al. (2010) wrote "isolated bones and partial skeletons of
Oviraptorosauria (Theropoda)" were discovered in 2006 at Bugin Tsav, of
which 060811 BgT-II KHTB was labeled Oviraptorosauria and found on June
8. They also stated "isolated bones of Oviraptorosauria" were
found in 2006 at Gurilin Tsav but did not list collected
specimens. "A partial skeleton
of small theropod" was said to be found that year at Altan Uul, which
may correspond to 060908 AU-IV BON discovered on September 8 at Altan
Uul IV, labeled as "?Oviraptorosauria" with material listed as
"pelvic." Watabe et al. say "isolated and articulated bones of
... Oviraptorosauria" were found at Tsaagan Khushuu in 2006, but did not
list collected material. They stated "the associated bones of
Oviraptorosauria including cranial elements were found" in the Upper
White Beds of Khermeen Tsav in 2006, and that "those bones are from
more than four individuals of mass-burial in a single layer."
Watabe et al.'s list indicates the specimen number 060906 KmT KHTB,
found on September 6.
References- Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1998. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 83-98.
Matsumoto, Hashimoto, Sonoda, Fujiyama, Mifune, Kawahara and Saneyoshi,
2010. Report of the preparation works for Mongolian specimens in
Hayashibara Museum of Natural Sciences: 1999-2008. Hayashibara Museum
of Natural Sciences Research Bulletin. 3, 167-185.
Watabe, Suzuki, Tsogtbaatar, Tsubamoto and Saneyoshi, 2010. Report of
the HMNS-MPC Joint Paleontological Expedition in 2006. Hayashibara
Museum of Natural Sciences Research Bulletin. 3, 11-18.
Chuniaoia Ji and Ji, 2001
Comments- Ji and Ji (2001) used the name Chuniaoia in a cladogram for
a branch leading to Protarchaeopteryx, but it has not been defined nor
has its intended purpose been published. It is similar to the name Chuniaoae
used in the online supplementary information of Ji et al. (1998), which was
seemingly for a Caudipteryx+Aves clade.
References-
Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs from northeastern China.
Nature. 393, 753-761.
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.
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 (16 mm), five dorsal vertebrae (13 mm), 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 (74 mm),
scapholunare, semilunate carpal, distal carpal III, metacarpal I (17
mm), phalanx I-1, manual ungual I, metacarpal II (44 mm), phalanx II-1,
phalanx II-2, manual ungual II, metacarpal III (45 mm), phalanx III-1,
phalanx III-2, phalanx III-3, manual ungual III, two keratinous manual
ungual sheaths, partial ilium (95 mm), incomplete pubes (80 mm),
ischial fragment(?), incomplete femora (125 mm), tibiae (160 mm),
partial fibulae, astragali, calcaneum, distal tarsal III, distal tarsal
IV, metatarsal I (13 mm), phalanx I-1, pedal ungual I, metatarsal II
(77 mm), phalanx II-1, phalanx II-2, pedal ungual III, metatarsal III
(86 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III,
metatarsal IV (81 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal ungual IV, metatarsal V, contour feathers, retrices
Diagnosis- (after Senter et al., 2004) six maxillary teeth; seven dentary
teeth.
Other diagnoses-
Ji and Ji's (1997) initial diagnosis is formed entirely of
plesiomorphies (claviform and unserrated dentition; sternum is thin and
flat; tail is long; forelimb/hindlimb index is 0.7; forelimb resembles Archaeopteryx
in morphology with three unguals, the second of which is enlarged;
ilium is large and elongated; pubes are robust and distally fused; hind
limb is long and robust with digit I reduced and dorsally migrated;
torso feathers are 50 mm in length with short and robust shafts; tail
fan is extremely well developed; retrices are as long as 150 mm with a
slender and elongated shaft and slender and gracile barbs) and untrue statements (proximal
metatarsals are fused; pedal digit I lies in opposition to digit III
and forms a grasping apparatus).
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 Maniraptora);
all of which are primitively serrated (incorrect- Senter et al., 2004); rectrices
form a fan at the end of the tail (plesiomorphic for Pennaraptora).
Comments- The holotype was discovered in 1996.
Phylogenetic relationships- Ji and Ji (1997) initially referred Protarchaeopteryx
to Archaeopterygidae based on constricted tooth roots ("claviform"
teeth; primitive for maniraptorans) and "elongated forelimb or
primitive wing with three talons" (primitive for maniraptoriforms) and
"extremely well developed tail fan, and feather morphology" (primitive
for pennaraptorans).
Zhou (1997) classified Protarchaeopteryx as a sauriurine
bird more closely related to enantiornithines than Confuciusornis due
to the smaller manual ungual III, but Sauriurae is near universally rejected and subsequent analyses have shown Confuciusornis is far more similar to enantiornithines than Protarchaeopteryx.
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,
and thus closer to birds than Caudipteryx. However, both studies supported
the paravian status of Protarchaeopteryx
using the same seven characters. Of these, three (deep suborbital
bar; unfused interdental plates; pubic foot projects posteriorly only)
are unknown in Protarchaeopteryx, and two others (less than 11 caudal
vertebrae with transverse processes; metacarpal I less than 33% of
metacarpal II in length) are not present in the taxon.
Oviraptorids and troodontids are miscoded as lacking a proximodistally
elongate coracoid, which is actually a pennaraptoran
synapomorphy. A radius less than 70% of ulnar width is shared
with e.g. Caudipteryx, Similicaudipteryx and Anzu and is highly homoplasious in basal Paraves.
Xu et al. (2000) resolved Protarchaeopteryx as a paravian more derived
than Caudipteryx
based on four additional characters. Two (teeth with serrated
mesial and distal carinae; mid and distal caudal vertebrae at least
130% longer than proximal caudals) arn't present in Protarchaeopteryx , while a short dorsal process of the
premaxilla cannot be determined. The remaining character (manus
over 120% of ulnar length) was misscored as lacking in Compsognathus, ornithomimids, Caudipteryx
and oviraptorids (ornithomimids and oviraptorids
are polymorphic), but is also a composite character involving lengths of different manual elements.
Holtz (2001) found Protarchaeopteryx to be either a basal member of the therizinosaur-oviraptorosaur clade, sister taxon to Pennaraptora, 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 basal pennaraptorans.
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,
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 taphonomic instead of structural 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 avepectorans. The robust second manual
digit is shared with caenagnathids, while the low ilium and slender pointed
postacetabular process are unquantified but also present in e.g. Similicaudipteryx.
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.
Senter (2007) and Cau (2018) found Protarchaeopteryx to be the sister
group of Incisivosaurus, with both of these taxa as the most basal oviraptorosaurs.
Funston and Currie (2016) added the taxon to a version of Maryanska's
oviraptorosaur matrix and found it emerged as the sistergroup of Archaeopteryx, albeit with Velociraptor and Herrerasaurus as the only other included non-oviraptorosaurs.
Hartman et al. (2019) recovered Protarchaeopteryx
as the sister taxon to Pennaraptora, but after adding more taxa it is
the most basal oviraptorosaur. In that updated matrix it takes 13
steps to be an archaeopterygid, 34 steps to be between Confuciusornis and enantiornithines, 5 steps to be a paravian,
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 Osm�lska (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.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic Palaeontology. 5(4), 429-463.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25. DOI: 10.4435/BSPI.2018.01
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Similicaudipteryx
He, Wang and Zhou, 2008
S. yixianensis He, Wang and Zhou, 2008
= Caudipteryx yixianensis (He, Wang and Zhou, 2008) Paul, 2010
Early Albian, Early Cretaceous
Jiufotang Formation, Liaoning, China
Holotype- (IVPP V12556) (adult) six cervical vertebrae, cervical ribs,
fourteen dorsal vertebrae, dorsal rib fragments, sacrum (85 mm), twenty-one
partial caudal vertebrae, pygostyle, six chevrons, incomplete scapula, partial
coracoid, incomplete sternal plates, sternal rib fragments, humerus (~130 mm),
partial ulna, partial radius, ilia (one partial; 153 mm), pubes (~223 mm), ischial
fragment, femora (one distal; ~220 mm), tibiae (one incomplete; ~240 mm), partial
fibula (~223 mm), metatarsal I (25 mm), phalanx I-1 (28 mm), pedal ungual I
(23 mm), metatarsals II (144 mm), phalanges II-1 (~43 mm), phalanx II-2 (39
mm), pedal ungual II (33 mm), metatarsals III (~183 mm), phalanges III-1 (46
mm), phalanx III-2 (34 mm), phalanx III-3 (32 mm), pedal ungual III (34 mm),
metatarsals IV (153 mm), phalanges IV-1 (30 mm), phalanx IV-2 (23 mm), phalanx
IV-3 (18 mm), phalanx IV-4 (21 mm), pedal ungual IV (20 mm)
Late Valanginian-Middle Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Referred- (STM4-1) (juvenile) partial skull, cervical vertebrae, dorsal
vertebrae, dorsal ribs, gastralia, caudal vertebrae, chevrons, scapulae, humeri
(24 mm), radii (21 mm), ulnae, semilunate carpals, metacarpals I, phalanx I-1,
manual ungual I, metacarpals II, phalanges II-1, phalanges II-2, manual unguals
II, metacarpals III, phalanges III-1, phalanges III-2, phalanx III-3, manual
ungual III, ilium, pubis, ischium, femora (38 mm), tibiae (49 mm), fibula, astragalus,
metatarsals I, phalanges I-1, pedal unguals I, metatarsals II (one incomplete),
phalanges II-1 (one incomplete), partial phalanx II-2, metatarsals III (one
incomplete), phalanges III-1 (one incomplete), phalanx III-2, phalanx III-3
fragment, metatarsals IV (one partial), phalanx IV-1, phalanx IV-2, phalanx
IV-3, phalanx IV-4, incomplete pedal ungual IV, metatarsal V, body feathers,
remiges, retrices (Xu et al., 2010a)
(STM22-6) (subadult) skull, mandible, ten cervical vertebrae, cervical ribs,
dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, pygostyle, chevrons,
scapulae, coracoid, furcula, humeri (81 mm), radii (69 mm), ulnae, metacarpals
I, phalanges I-1, manual unguals I, metacarpals II, phalanges II-1, phalanx
II-2, manual ungual II, metacarpals III, phalanx III-1, phalanx III-2, phalanx
III-3, manual ungual III, incomplete ilium, pubes, ischium, femur (140 mm),
tibiae (182 mm), astragali, phalanx I-1, pedal ungual I, metatarsal II, phalanx
II-1, phalanx II-2, pedal ungual II, metatarsal III, phalanx III-1, phalanx
III-2, phalanx III-3, metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3,
body feathers, remiges, retrices (Xu et al., 2010a)
Diagnosis- (modified from He et al., 2008) puboilial ratio of 1.46.
Other diagnoses- In its
diagnosis, He et al. list a dagger-like pygostyle (also present in Nomingia
and Citipati), ilium shaped like Caudipteryx (ambiguous), two
large anterior dorsal hypapophyses (present in caenagnathoids) and puboilial
ratio of 1.46. The latter compares with 1.04-1.12 in Caudipteryx, .96
in Nomingia and .99 in Microvenator.
Comments- The holotype was first mentioned by Wang et al. (2007) as an
unnamed caudipterid and later described and named by He et al. (2008). Xu et al. (2010a) briefly describe
two new, younger specimens which are almost complete.
He et al. referred it to
Caudipteridae based on several characters. Most are symplesiomorphic for maniraptorans
(low number of caudal vertebrae; deep pubic peduncle; unfused metatarsus; metatarsal
III longest; metatarsals II and IV subequal in width; metatarsal II slightly
shorter than IV; subarctometatarsal metatarsus), the proximally placed metatarsal
I is also present in Protarchaeopteryx, and the preacetabular depth is
intermediate between other oviraptorosaurs.
Xu et al. (2011) added Similicaudipteryx to Senter's TWiG matrix and recovered it as an oviraptorosaur more derived than Protarchaeopteryx.
Lamanna et al. (2014) added this to Maryanska's oviraptorosaur matrix and recovered it as a caudipterid.
Cau (2018) recovered it as more derived than Protarchaeopteryx and Incisivosaurus, but less than Caudipteryx.
Most recently, Hartman et al. (2019) recovered it as the most basal oviraptorosaur, but after adding several taxa Protarchaeopteryx moves to be the basalmost member instead.
References- Wang, Jones and Evans, 2007. A juvenile anuran from the Lower
Cretaceous Jiufotang Formation, Liaoning, China. Cretaceous Research. 28, 235-244.
He, Wang and Zhou, 2008. A new genus and species of caudipterid dinosaur from
the Lower Cretaceous Jiufotang Formation of Western Liaoning, China. Vertebrata
PalAsiatica. 46(3), 178-189.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Ptum, 2010. Moulting tail feathers in a juvenile oviraptorisaur. Nature. 468,
E1.
Xu, Zheng and Yu, 2010a. Exceptional dinosaur fossils show ontogenetic development
of early feathers. Nature. 464, 1338-1341.
Xu, Zheng and Yu, 2010b. Xu et al. reply. Nature. 464, 468, E2.
Xu, You, Du and Han, 2011. An Archaeopteryx-like theropod
from China and the origin of Avialae. Nature. 475, 465-470.
Lamanna, Sues, Schachner and Lyson, 2014. A new large-bodied oviraptorosaurian
theropod dinosaur from the Latest Cretaceous of Western North America. PLoS
ONE. 9(3), e92022.
Cau,
2018. The assembly of the avian body plan: A 160-million-year long
process. Bollettino della Societ� Paleontologica Italiana. 57(1),
1-25. DOI: 10.4435/BSPI.2018.01
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Ganzhousaurus Wang, Sun,
Sullivan and Xu, 2013
G. nankangensis Wang, Sun, Sullivan and Xu, 2013
Early Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Holotype- (SDM 20090302) incomplete mandible, distal caudal vertebra,
incomplete distal caudal vertebra, partial distal caudal vertebra, ilial fragment,
partial tibia, distal tarsal III, metatarsal I (36 mm), phalanx I-1 (28 mm),
pedal ungual I, metatarsal II (125 mm), phalanx II-1 (41 mm), phalanx II-2,
partial pedal ungual II, metatarsal III (147 mm), phalanx III-1 (45 mm), partial
phalanx III-2, phalanx IV-1, partial phalanx IV-2
Diagnosis- (after Wang et al., 2013) shallow dentary (ratio of maximum
anteroposterior length to maximum dorsoventral depth 1.90); absence of fossa
or foramen on lateral surface of dentary (also in oviraptorids; unlike Gigantoraptor,
Chirostenotes and Caenagnathasia); weakly downturned anterior
mandibular end (also in Khaan; unlike Nemegtomaia and Heyuannia);
shallow depression immediately surrounding anterior margin of external mandibular
fenestra (also in Gigantoraptor, Nemegtomaia and Heyuannia;
unlike Citipati and Khaan); external mandibular fenestra subdivided
by anterior process of surangular (also in oviraptorids); dentary posteroventral
process slightly twisted and positioned on mandibular ventrolateral surface
(also in Nemegtomaia; unlike Citipati and Heyuannia); shallow
longitudinal groove along medial surface of dentary posteroventral process (also
in Microvenator; unknown in other oviraptorosaurs); angular anterior
process wider transversely than deep dorsoventrally (unknown in other oviraptorosaurs);
sharp groove along ventrolateral surface of angular anterior process (also in
Chirostenotes, Gigantoraptor and Khaan; unlike most oviraptorosaurs);
ventral border of external mandibular fenestra formed mainly by angular (also
in Chirostenotes and Gigantoraptor; unlike oviraptorids); ventral
flange along distal half of metatarsal II (also in Avimimus; unlike Citipati,
Wulatelong and Heyuannia); arctometatarsal condition absent (also
in oviraptorids; unlike Avimimus, Chirostenotes and Elmisaurus).
Comments- The holotype was purchased from a fossil dealer and initially
mentioned in an abstract by Wang and Xu (2012) before being officially described
by Wang et al. (2013). Initially the partial tibia was thought to be a
femur by Wang and Xu. Note the diagnosis given by Wang et al. includes numerous symplesiomorphies
and will need to be revised.
Wang and Xu found it to be a heyuannine in
their unpublished analysis, while Wang et al. found it to be most
closely related to Citipati and Rinchenia in its published form,
while Cau (online, 2013) found it to be in a trichotomy with caenagnathids and
oviraptorids. Hartman et al. (2019) found it to be between Similicaudipteryx and caudipterids.
References- Wang and Xu, 2012. A new oviraptorid specimen from the Upper
Cretaceous of southern China. Journal of Vertebrate Paleontology. Program and
Abstracts 2012, 190.
Cau, online 2013. http://theropoda.blogspot.com/2013/04/ganzhousaurus-nankangensis-wang-et-al.html
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda)
from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
undescribed oviraptorosaur (Zhou, O'Connor and Wang, 2014)
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Comments- Zhou et al. (2014)
reported "one caudipteryid (Maniraptora: Oviraptorosauria)" from
Sihedang. This may be "Lingyuanosaurus", which was described in
2019 as a therizinosaur, but has oviraptorosaurian characters as well.
Reference- Zhou, O'Connor and Wang, 2014. A new species from an ornithuromorph (Aves: Ornithothoraces)
dominated locality of the Jehol Biota. Chinese Science Bulletin. 59(36), 5366-5378.
Caudipteridae Zhou and Wang, 2000
Definition- (Caudipteryx zoui <- Caenagnathus collinsi,
Oviraptor philoceratops) (Hendrickx, Hartman and Mateus, 2015)
= Caudipterygidae Osmolska, Currie and Barsbold, 2004
Comments- In 2000, Zhou and Wang proposed the family Caudipteridae for
Caudipteryx.
Osmolska et al. (2004) emmended this to Caudipterygidae, since
Caudipteridae is formed incorrectly (ICZN Article 29.3). It has been
suggested this is unecessary, since according to Article 29.4, "if
after 1999 a new family-group name is based on a generic name which is
or ends in a Greek or Latin word or ends in a Greek or Latin suffix,
but its derivation does not follow the grammatical procedures of
Articles 29.3.1 or 29.3.2, its original spelling must be maintained as
the correct original spelling." However, Article 29.4.2 states this is
only true provided the genus was treated as an arbitrary combination of
letters (e.g. "Caudipteryxidae"), which is not the case. To complicate
matters, Article 29.5 states "If a spelling of a family-group name was
not formed in accordance with Article 29.3 but is in prevailing usage,
that spelling is to be maintained, whether or not it is the original
spelling and whether or not its derivation from the name of the type
genus is in accordance with the grammatical procedures in Articles
29.3.1 and 29.3.2." Caudipteridae has 6420 search results
compared to Caudipterygidae's 402 on Google, and 15 vs. 14 on Google
Scholar (as of 8-26-19). Thus Caudipteridae should be maintained.
References- Zhou and Wang, 2000. A new species of Caudipteryx
from the Yixian Formation of Liaoning, northeast China. Vertebrata PalAsiatica.
38(2), 113-130.
Osm�lska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel,
Dodson and Osm�lska, (eds.). The Dinosauria, Second Edition. University
of California Press. 165-183.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Caudipteryx Ji, Currie, Norell and Ji, 1998
Diagnosis- premaxillary teeth limited to rostral half of element; first
premaxillary tooth much larger than others (also in Incisivosaurus and Protarchaeopteryx);
single maxillary fenestra present; twenty-two caudal vertebrae; sternal plates
oval; only two phalanges present on manual digit III.
Other diagnoses- (after Ji et
al., 1998) elongate, hooked premaxillary teeth with broad roots;
maxilla and dentary edentulous; tail short (one-quarter of the length
of the body); arm long for a non-avian theropod; short manual unguals;
Leg-to-arm ratio, 2.5.
Reference- Ji, Currie, Norell and Ji, 1998. Two feathered dinosaurs
from northeastern China. Nature. 393, 753-761.
C. zoui Ji, Currie, Norell and Ji,
1998
= Caudipteryx dongi Zhou and Wang, 2000
Late Barremian-Early Aptian, Early Cretaceous
Jianshangou member of Yixian Formation, Liaoning, China
Holotype- (NGMC 97-4-A) (890 mm) skull (76 mm), mandibles, cervical vertebae,
cervical ribs, dorsal vertebrae, dorsal ribs, gastralia, twenty-two caudal vertebrae
(first caudal 12 mm), chevrons, coracoid, sternal plate (36 mm), humeri (69
mm), radii, ulnae, scapholunare, 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, scapholunare, ulnare, metacarpal
I (13 mm), phalanx I-1 (25 mm), manual ungual I (15 mm), metacarpal II (29 mm),
phalanx II-1 (18.5 mm), phalanx II-2 (25 mm), manual ungual II (18 mm), metacarpal
III (27 mm), ilia (115 mm), pubes, ischia (73 mm), femora (146, 152 mm), tibiae
(196 mm), fibula (181 mm), astragali, calcaneum, distal tarsal III, distal tarsal
IV, metatarsal I (19 mm), phalanx I-1 (12 mm), pedal ungual I (11 mm), metatarsal
II (112 mm), phalanx II-1 (25 mm), phalanx II-2 (16 mm), pedal ungual II (19
mm), metatarsal III (124 mm), phalanx III-1 (27 mm), phalanx III-2 (20 mm),
phalanx III-3 (17 mm), pedal ungual III (20 mm), metatarsal IV (116 mm), phalanx
IV-1 (15 mm), phalanx IV-2 (9 mm), phalanx IV-3 (7 mm), phalanx IV-4 (7 mm),
pedal ungual IV (16 mm), metatarsal V (~36 mm), body feathers, remiges (182
mm), gastroliths (Zhou and Wang, 2000)
(LPM 0005) skull, mandibles, cervical series, cervical ribs, dorsal series,
dorsal ribs, gastralia, sacral vertebrae, caudal series, chevrons, scapulae,
coracoid, furcula, humeri, radii, ulnae, semilunate carpal, metacarpals I, phalanges
I-1, manual unguals I, metacarpals II, phalanges II-1 (one partial), phalanx
II-2, manual ungual II, metacarpals III, phalanges III-1, phalanges III-2, ilia,
ischia, femora, tibiae, fibulae, astragali, calcaneum, metatarsals I, phalanges
I-1, pedal ungual I, metatarsals II, phalanges II-1, phalanges II-2, pedal unguals
II, metatarsals III, phalanges III-1, phalanges III-2, phalanges III-3, pedal
unguals III, metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3,
phalanges IV-4, pedal unguals IV, metatarsals V, remiges, retrices, body feathers,
gastroliths (Feduccia and Czerkas, 2015)
(STM 4-3) specimen including dorsal vertebrae, dorsal ribs, scapula and coracoids (Zheng et al., 2014)
Diagnosis- (suggested) 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, scapholunare, ulnare, metacarpal
I (11 mm), phalanx I-1 (25 mm), manual ungual I (16 mm), metacarpal II (28 mm),
phalanx II-1 (17 mm), phalanx II-2 (24 mm), manual ungual II (15 mm), metacarpal
III (25 mm), phalanx III-1, phalanx III-2, ilia (115 mm), pubes (~124 mm), ischia
(72 mm), femora (145 mm), tibiae (188 mm), fibulae (188 mm), astragali, calcanea,
distal tarsal III, distal tarsal IV, metatarsal I (16 mm), phalanx I-1 (13 mm),
pedal ungual I (12 mm), metatarsal II (102 mm), phalanx II-1 (23 mm), phalanx
II-2 (16 mm), pedal ungual II (19 mm), metatarsal III (113 mm), phalanx III-1
(24 mm), phalanx III-2 (19 mm), phalanx III-3 (15 mm), pedal ungual III (18
mm), metatarsal IV (107 mm), phalanx IV-1 (14 mm), phalanx IV-2 (8 mm), phalanx
IV-3 (6 mm), phalanx IV-4 (4 mm), pedal ungual IV (14 mm), metatarsal V (30
mm), contour feathers, remiges, retrices, gastroliths
(IVPP V 12340) (836 mm) skull, mandibles, twelve cervical vertebrae, cervical
ribs, nine dorsal vertebrae (d3 ~18 mm, d6 ~18 mm, d8 ~17 mm), dorsal ribs (120
mm), uncinate processes (26 mm), gastralia, sacrum, twenty-two caudal vertebrae,
chevrons, scapulae (80 mm), coracoids (35 mm), sternal ribs (36 mm), humeri
(69 mm), radii (~56 mm), ulnae (61 mm), semilunate carpal, scapholunare, 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- (suggested) 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.
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 between 145-152
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.
Phylogenetic relationships-
Sereno (1999) was the first to recover Caudipteryx as an oviraptorosaur which has been the consensus since 2002, but a few other possibilities have been proposed over the years.
Caudipteryx in Avialae? Ji et al. (1998) first included Caudipteryx in Chiappe's bird matrix, also including 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; dorsal premaxillary process 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 misscored as having them), this doesn't support placing Caudipteryx
in Avialae instead of Oviraptorosauria. The second character is also seen in caenagnathoids (Anzu, Avimimus, oviraptorids),
though absent in Incisivosaurus.
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. A reduced hypopubic
cup is a 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 paravians like Archaeopteryx, but present in at least some enantiornithines in addition to confuciusornithids. On
the other hand, the enlarged premaxilla, reduced maxilla, toothless maxilla and dentary and
shortened tail are indeed shared with Confuciusornis but not Archaeopteryx, but are also found in caenagnathoids. Martin (2004) even suggested Caudipteryx
and other oviraptorosaurs may be most closely related to
confuciusornithids. This is highly unparsimonious though, as
oviraptorosaurs lack numerous paravian/eumaniraptoran, avialan,
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 closer to birds than Archaeopteryx but basal to paraphyletic
alvarezsaurids, oviraptorids and ornithothoracines. Similar to Ji et
al.'s analysis, this was based on Chiappe's bird matrix so has the same
issues, except this time almost all alvarezsaurid characters were
excluded and Oviraptorosauria was added as an OTU though without many
characters linking Caudipteryx to it. So this matrix again forces Caudipteryx, alvarezsaurids and now oviraptorosaurs to be avialans, which is unparsimonious when non-paravian outgroups are used.
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. Of these, a deep jugal, unfused
interdental plates, less than 11 caudal vertebrae with transverse
processes, metacarpal I less than 33% of metacarpal II in length, pubic
foot projects posteriorly only and posterior trochanter present are
misscored in Caudipteryx.
Oviraptorids and troodontids are miscoded as lacking a proximodistally
elongate coracoid, which is actually a pennaraptoran
synapomorphy. Most deinonychosaurs (except Archaeopteryx, Ningyuanosaurus, Eosinopteryx and Jinfengopteryx) and jeholornithids have more than 25 caudal vertebrae, so the low number in Caudipteryx is not a paravian character (and is shared with e.g. Similicaudipteryx). A radius less than 70% of ulnar width is shared with e.g. Protarchaeopteryx, Similicaudipteryx and Anzu
and is highly homoplasious in basal Paraves. However, mid and
distal chevrons dorsoventrally flattened (defined as in Hartman et al.
2019 as 'Chevrons - anteroposterior length >75% of dorsoventral
depth by chevron 10') is shared with almost all paravians to the
exclusion of oviraptorosaurs.
Caudipteryx outside Pennaraptora? Holtz (2001) found Caudipteryx to either be sister to a therizinosaur plus oviraptorosaur clade or a basal pennaraptoran. Gishlick (2002) found Caudipteryx to be in a trichotomy with therizinosaurs
and Pennaraptora, but the matrix only contained forelimb
characters.
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.
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. In Gauthier and Gall (eds.). New Perspectives
on the Origin and Early Evolution of Birds: Proceedings of the International
Symposium in Honor of John H. Ostrom. Yale University Press. 99-122.
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.
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. 175-189.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica.
50(6), 978-990.
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. Journal of Morphology. 266(2), 125-166.
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.
Zheng, O'Connor, Wang, Wang, Zhang and Zhou, 2014. On the absence of sternal
elements in Anchiornis (Paraves) and Sapeornis (Aves) and the
complex early evolution of the avian sternum. Proceedings of the National Academy
of Sciences. 111(38), 13900-13905.
Feduccia and Czerkas, 2015. Testing the neoflightless hypothesis: Propatagium
reveals flying ancestry of oviraptorosaurs. Journal of Ornithology. 156(4),
1067-1074.
"Xingtianosaurus" Qiu, Wang, Wang, Li, Zhang and Ma, 2019
"X. ganqi" Qiu, Wang, Wang, Li, Zhang and Ma, 2019
Early Aptian, Early Cretaceous
Dawangzhangzi Beds of Yixian Formation, Liaoning, China
Material- (IVPP V13390) four dorsal centra, dorsal rib fragments,
(?)uncinate processes, gastralia, anterior sacral centrum, about
eighteen caudal vertebrae and centra, nine chevrons, partial scapulae
(~56.2 mm), (?)sternal plate (~32.7 mm), humeri (one incomplete, one
partial; 72.4 mm), radii (69.9, 63.3 mm), ulnae (one incomplete; 73.5,
69.5 mm), scapholunare, (?)ulnare, semilunate carpals, metacarpals I (16.6,
15.4 mm), phalanges I-1 (one partial; 25.5 mm), manual ungual I
(impression), metacarpals II (one incomplete; 40.5 mm), proximal
phalanx II-1, distal phalanx II-2, partial manual unguals II (21.2,
~19.7 mm), metacarpals III (one incomplete; 37.2 mm), proximal phalanx
III-1, distal phalanx III-3, manual unguals III (15.4, 15.6 mm),
partial ilium (~89.8 mm), pubes (~107 mm), ischium (~54.6 mm), femora
(127, 120.4 mm), incomplete tibiae (171.3 mm), incomplete fibulae,
astragalus, two distal tarsals, metatarsals I (9.8, 10.2 mm), phalanx
I-1 (19 mm), pedal ungual I (14.6 mm), metatarsals II (80.4, 78.5 mm),
phalanges II-1 (28.3, 27.8 mm), phalanges II-2 (22.6, 23.8 mm), pedal
unguals II, metatarsals III (89.7, 90.2 mm), phalanges III-1 (27, 27
mm), phalanges III-2 (20.9, 20.7 mm), phalanges III-3 (20.3, 21.4 mm),
pedal unguals III (17.2, 15 mm), metatarsals IV (82.8, 82.9 mm),
phalanges IV-1 (17.3, 17.6 mm), phalanges IV-2 (14.5, 14.3 mm),
phalanges IV-3 (12.7, 12.8 mm), phalanges IV-4 (16, 14.7 mm), pedal
unguals IV (16, 15.8 mm), metatarsal V (25.6 mm), remiges
Other diagnoses- Of the listed characters in Qiu et al.'s (2019) diagnosis, none definitely distinguish this taxon from Protarchaeopteryx.
- "small pleurocoel close to the dorsal edge of the lateral surface of
the dorsal vertebral centrum", pleurocoels are also present in Protarchaeopteryx's posterior dorsals (unlike Caudipteryx) although their size and position are unreported.
- "humerus longer than the scapula" is unknown in Protarchaeopteryx as only the proximal scapula is preserved.
- "proportionally long ulna (as long as humerus)" with an ulnohumeral ratio of 101%, which is larger than ~86% in Protarchaeopteryx. This is actually due to the longer humerus in Protarchaeopteryx,
as the femur and ulna are about the same length in each specimen.
One interesting possibility is that previous authors were mistaken in
identifying the proximal humerus of Protarchaeopteryx, as the purported fragment of is indistinct and even has a hole which could be a coracoid foramen.
- "relatively small radiale [= scapholunare] angle (39�, compared to >48� in other
oviraptorosaurs with known radiale angle)" compared to 26� in Protarchaeopteryx.
- "extremely short metacarpal I (<40% length of the metacarpal II)" is actually 41%, compared to ~39% in Protarchaeopteryx.
- "small ligament pits on the manual phalanges" doesn't seem distinct from some Caudipteryx (NGMC 97-9-A) in the one phalanx illustrated as preserving them (II-2), but does seem larger in at least I-1 and III-3 of Protarchaeopteryx. Qiu et al.'s Caudipteryx example (their Fig. 7c) is IVPP V12430, which does have a large pit on II-2.
Comments- This was described by Qiu et al. on April 25 2019 as a new taxon
of caudipterid. However, this paper has no mention of ZooBank
and as of February 6 2020
"Xingtianosaurus" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Xingtianosaurus ganqi" Qiu
et al., 2019 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
If Protarchaeopteryx's humerus has been correctly interpreted, it's much longer than "Xingtianosaurus"',
but the altter's greater scapholunare angle and smaller manual ligament pits
on at least two phalanges may be outside of individual variation.
Yet the sternal plates would certainly seem to be very different.
Protarchaeopteryx's are a rounded rectangle, while "Xingtianosaurus"'
are illustrated as being drop-shaped with a notch that creates
anterolateral and posterolateral processes. Yet the medial edge
is along a crack in the matrix, with similar texture to the sternal
bone on the other side in some areas, and the entire anterior edge is
hidden by ribs. Only a short section representing about a fourth
of the supposed lateral notch is exposed, and this could be
breakage. Indeed, with only a short curved portion of the edge
obviously visible, the whole bone could easily be one of the otherwise
missing coracoids. Thus definitely distinguishing the taxa may
involve the proper identification of elements beyond what published
photos allow.
Qiu et al. (2019) used a version of Maryanska's oviraptorosaur matrix
and recovered "Xingtianosaurus" as a caudipterid. Adding it to
Hartman et al.'s analysis also recovers it as a caudipterid.
Reference- Qiu, Wang, Wang, Li,
Zhang and Ma, 2019. A new caudipterid from the Lower Cretaceous of
China with information on the evolution of the manus of
Oviraptorosauria. Scientific Reports. 9:6431. DOI:
10.1038/s41598-019-42547-6
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; contact between
the accessory ventral flanges of the pterygoids.
(after Senter et al., 2004) nine maxillary teeth; eight or nine dentary teeth.
Other diagnoses- 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 Oviraptorosauria); very small lanceolate maxillary teeth (plesiomorphic
for Oviraptorosauria); 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.
Comments- Balanoff
et al. (2009) note the posterior mandibular fragment is lost.
References- Xu, Cheng, Wang and Chang, 2002. An unusual oviraptorosaurian
dinosaur from China. Nature. 419, 291-293.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae.
Bulletin of Gunma Museum of Natural History. 8, 1-20.
Balanoff, Xu, Matsufune, Kobayashi and Norell, 2007. Endocranial anatomy of
a primitive oviraptorosaur, Incisivosaurus gauthieri, (Theropoda: Dinosauria).
Journal of Vertebrate Paleontology. 27(3), 43A.
Balanoff, Xu, Kobayashi, Matsufune and Norell, 2009 online. Incisivosaurus gauthieri, Digital Morphology. http://digimorph.org/specimens/Incisivosaurus_gauthieri/
Balanoff, Xu, Kobayashi, Matsufune and Norell, 2009. Cranial osteology of the
theropod dinosaur Incisivosaurus gauthieri (Theropoda: Oviraptorosauria).
American Museum Novitates. 3651, 35 pp.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution.
PhD thesis. Columbia University. 522 pp.
Caenagnathoidea Sternberg, 1940 sensu Sereno,
1999b
Definition- (Caenagnathus collinsi + Oviraptor philoceratops)
(Maryanska, Osmolska and Wolsan, 2002; modified from Sereno, 1999b)
Other definitions- (Chirostenotes pergracilis + Oviraptor philoceratops)
(Sereno, online 2005)
(Caenagnathus collinsi + Oviraptor philoceratops + Avimimus portentosus) (Hendrickx,
Mateus, Ara�jo and Choiniere, 2019)
= Oviraptorosauria sensu Padian, Hutchinson and Holtz, 1999
Definition- (Oviraptor philoceratops + Chirostenotes pergracilis)
(modified)
= Oviraptoroidea Barsbold, 1976 sensu Sereno, 1999a
Definition- (Oviraptor philoceratops + Caenagnathus collinsi)
(modified)
= Caenagnathoidea sensu Sereno, online 2005
Definition- (Chirostenotes pergracilis + Oviraptor philoceratops)
= Caenagnathoidea sensu Hendrickx,
Mateus, Ara�jo and Choiniere, 2019
Definition- (Caenagnathus collinsi + Oviraptor philoceratops + Avimimus portentosus)
= Edentoraptora Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
Comments- This node covered known oviraptorosaurs until Sereno (1999a) identified Caudipteryx
as a basal oviraptorosaurian and proposed Oviraptoroidea as a
name. However, according to ICZN rules Caenagnathoidea was
implicitly named in 1940 when Sternberg erected Caenagnathidae, so it
must have priority over Oviraptoroidea which was implicitly created in
1976 by Barsbold. Thus (though without explanation in the text)
Sereno (1999b) next used Caenagnathoidea for this node, with its
eponymous genera implied by the families it contains in the
cladogram.
Funston et al. (2020) noted "a well-supported (decay index 2) group of Avimimus
and Caenagnathoidea, referred to here as Edentoraptora based on the
ubiquitous absence of teeth in these animals." No phylogenetic
definition was provided and the group has the same content as
Caenagnathoidea in this site's topology based on Hartman et al. where Avimimus is a caenagnathid. Forcing Avimimus outside Caenagnathoidea in that matrix and thus making Edentoraptora a unique clade takes ten additional steps.
Caenagnathoidea defined- Maryanska et al. (2002) specified the defining genera and species, but Sereno (online 2005) tried to replace 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 its eponymous species. Hendrickx et al. (2019) added Avimimus to the definition as an internal specifier, but this is unneccesary and indeed whether Avimimus is a caenagnathoid is an active problem in oviraptorosaur phylogeny.
References-
Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of
Paleontology. 14(1), 81-85.
Barsbold, 1976. On a new Late Cretaceous family of small
theropods (Oviraptoridae fam. n.) of Mongolia. Doklady Akademia Nauk
SSSR. 226, 685-688.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions
and nomenclature of the major taxonomic categories of the carnivorous Dinosauria
(Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Sereno, 1999a. The evolution of dinosaurs. Science. 284, 2137-2147.
Sereno, 1999b. A rationale for dinosaurian taxonomy. Journal of Vertebrate Paleontology. 19(4), 788-790.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php
[version 1.0, 2005 November 7]
Hendrickx,
Mateus, Ara�jo and Choiniere, 2019. The distribution of dental features
in non-avian theropod dinosaurs: Taxonomic potential, degree of
homoplasy, and major evolutionary trends. Palaeontologia Electronica.
22.3.74, 1-110.
Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new
two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.
Royal Society Open Science. 7: 201184.
undescribed Caenagnathoidea (Buckley, 2002)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (NS.1563.018) two partial manual unguals, partial tibia, proximal
fibula, partial astragalus, (metatarsal II ~390 mm) distal metatarsal III, partial
metatarsal shaft, incomplete pedal phalanx III-1, partial pedal phalanx (Buckley,
2002)
(NS.32001.077) metatarsal III, metatarsal IV (Buckley, 2002)
manual ungual (Holtz, Williams, Tremaine and Matthews, 2014)
Comments- The NS specimens were identified as Elmisaurus in the
abstract, but as caenagnathid or oviraptorid in the poster. They may belong to Anzu or Elmisaurus? sp. both known from the formation.
References- Buckley, 2002. New material of Elmisaurus (Theropoda,
Elmisauridae) from the Late Cretaceous Hell Creek Formation of southeastern
Montana. Journal of Vertebrate Paleontology. 22(3), 39A.
Holtz, Williams, Tremaine and Matthews, 2014. New additions to the Hell Creek
Formation (Upper Maastrichtian) vertebrate fauna of Carter County, Montana.
Journal of Vertebrate Paleontology. Program and Abstracts 2014, 149.
Luoyanggia Lu, Xu, Jiang,
Jia, Li, Yuan, Zhang and Ji, 2009
L. liudianensis Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009
Aptian-Albian, Early Cretaceous
Haoling Formation, Henan, China
Holotype- (41HIII-00010) dentaries
....(41HIII-00011; field number KLR 07-62-44a) incomplete ilium, ischium
....(field number KLR 07-62-28a-16) metatarsal II (175), metatarsal III (190 mm), metatarsal IV (177 mm)
....(field number KLR 07-62-49-1) ilium, incomplete pubis (160 mm), ischium
(100 mm)
Other diagnoses- Lu et al. (2009) listed three characters in their diagnosis,
but all are plesiomorphic- dentary not decurved; dentary symphysis V-shaped;
ischium only slightly concave dorsally.
Comments- While originally referred to the Mangchuan Formation, this
was redefined to include three formations by Xu et al. in 2012.
While Lu et al. (2009) referred Luoyanggia to Oviraptoridae, they seemingly mean Oviraptorosauria as they refer Chirostenotes to that family and state "Luoyanggia may have a certain relationship with Caudiptery"x. Funston and Currie (2016) added it to Maryanska's oviraptorosaur analysis and found it resolve sister to Similicaudipteryx
as the most basal oviraptorosaurs (note their published results did not
find their shortest trees). Hartman et al. (2019) recovered it as
a caenagnathine caenagnathid sister to Anzu, but adding more taxa places it less specifically as a caenagnathid outside the Elmisaurus+Avimimus clade.
Reference- Lu, Xu, Jiang, Jia, Li, Yuan, Zhang and Ji, 2009. A preliminary
report on the new dinosaurian fauna from the Cretaceous of the Ruyang Basin,
Henan province of Central China. Journal of the Paleontological Society of Korea.
25(1), 43-56.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
undescribed caenagnathid (Britt, Chure, Currie, Holmes, Theurer and Scheetz, 2021)
Late Albian, Early Cretaceous
DNM-16, Mussentuchit Member, Cedar Mountain Formation, Uhat, US
Material- (MCZ 3040) (~3 m, 4
year old subadult) posterior dorsal vertebra, rib fragments, incomplete
femur, tibia, astragalocalcaneum, couple partial metatarsals, phalanx
II-1, pedal phalanges
Comments- Found in the mid
1970s, this was presented by Britt et al. at SVP 2021. They wrote
"1) camellate pneumatization of posterior dorsal vertebrae, 2)
extremely high, parallel sided astragalar ascending process, 3)
diminutive calcaneum fused to the astragalus, and 4) adaptations for
extreme retraction and flexion in pedal Digit II" ..." suggests
affinities with the Troodontidae." However, the characters of
pedal phalanx II-1 are also present in Elmisaurus,
the other characters are also present in oviraptorosaurs, and e.g. the
pleurocoelous posterior dorsal is unknown in troodontids but consistant
with caenagnathoids. Inclusion in Hartman et al.'s
maniraptoromorph matrix results in a placement within Caenagnathidae,
conssistant with geography.
Reference- Britt, Chure,
Currie, Holmes, Theurer and Scheetz, 2021. A new deinonychosaurian
theropod from the Mid-Cretaceous (Albian) Mussentuchit Member of the
Cedar Mountain Formation in Dinosaur National Monument, northeastern
Utah, USA. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 65-66
undescribed Caenagnathidae (Fiorillo, 1989)
Late Campanian, Late Cretaceous
Judith River Formation, Montana
Material- two specimens
Comments- Referred to Chirostenotes, but may be Caenagnathus or Citipes
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.
Caenagnathidae indet. (Longrich, Barnes, Clark and Millar, 2013)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Material- (TMM 42335-40) caudal centrum
(TMM 42920-2) incomplete manual ungual II (~80 mm)
(TMM 43057-354) distal metatarsal III (~15.5 mm trans)
(TMM 43057-357) pedal ungual
(UTEP B38 L-3) femur (350 mm)
Comments- Longrich et al. (2013) refer TMM 42920-2 and UTEP B38 L-3 to Chirostenotes
sp. based on medium size, but they might also be juvenile Caenagnathus
so are assigned to Caenagnathidae indet. here. They assign TMM
42335-40, TMM 43057-354 and TMM 43057-357 to their new taxon Leptorhynchos gaddisi
based on small size, but the latter is here considered indeterminate
and possibly juvenile. These isolated small postcrania could be
juvenile Chirostenotes or adult Citipes (provisionally known from the formation based on tarsometatarsal fragment TMM 43057-36).
Reference- Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from
the Upper Campanian Aguja Formation of west Texas, and a revision of the Caenagnathinae.
Bulletin of the Peabody Museum of Natural History. 54(1), 23-49.
Caenagnathidae indet. (Yun and Funston, 2021)
Middle-Late Campanian, Late Cretaceous
Fales Rocks Locality (UW V-81006), Mesaverde Formation, Wyoming, US
Material- (UW 44439) incomplete metatarsal I (~32.7 mm)
Comments- Yun and Funston (2021) described this and concluded "UW 44439 is particularly similar to Anzu wyliei and CMN 8538, and contrasts in several features with Chirostenotes pergracilis",
namely "the metatarsal is teardrop-shaped in extensor (dorsal) view,
owing to the proximally-tapering shaft and a rounded distal condyle"
and "the distal articulation surface of UW 44439 is nearly spherical in
dorsal view." They placed it in Caenagnathidae indet..
Reference- Yun and Funston,
2021. A caenagnathid oviraptorosaur metatarsal from the Mesaverde
Formation (Campanian) of Wyoming, USA. Vertebrate Anatomy Morphology
Palaeontology. 9, 105-115.
undescribed caenagnathid (Hunt-Foster and Foster, 2015)
Late Campanian-Early Maastrichtian, Late Cretaceous
Williams Fork Formation, Colorado, US
Material- humerus
Comments- Hunt-Foster and Foster (2015) state this resembles Microvenator
more than Anzu.
Reference- Hunt-Foster and Foster, 2015. First occurrence of an oviraptorosaur
(Theropoda: Maniraptora) from the Mesaverde Group (Williams Fork Formation)
of northwestern Colorado. Journal of Vertebrate Paleontology. Program and Abstracts
2015, 148.
Caenagnathidae indet. (Jasinski, Sullivan and Lucas, 2011)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (SMP VP-2172) incomplete pedal ungual
Reference- Jasinski, Sullivan and Lucas, 2011. Taxonomic composition
of the Alamo Wash local fauna from the Upper Cretaceous Ojo Alamo Formation
(Naashoibito Member), San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann
(eds.). Fossil Record 3. New Mexico Museum of Natural History and Science Bulletin.
53, 216-271.
Caenagnathidae indet. (Buckley, 2002)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material-
(NS.31996.114H) distal metatarsal II (~216 mm) (Buckley, 2002)
(NS.32001.017B) distal metatarsal II (Buckley, 2002)
(TMP 1996.005.0012) distal metatarsal III (Funston et al., 2016)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
metatarsal (Anonymous, 1997)
material (DePalma, 2010)
Comments- Anonymous (1997) reported a Chirostenotes metatarsal associated with
Tyrannosaurus specimen FMNH PR2081.
Buckley (2002) referred two
distal metatarsals II to Citipes (her Elmisaurus elegans) because of their longitudinal
ridge which would have made the posterior metatarsus surface deeply
concave as in Elmisaurus and Citipes. However, these are the size of large Chirostenotes specimens.
DePalma (2010) referred material to Chirostenotes and Caenagnathus.
Funston et al. (2016) figured a tiny distal metatarsal III (TMP 1996.005.0012) as Leptorhynchos elegans, claiming well-defined cruciate ridges as in Elmisaurus and Citipes, and that the distal articular surface was taller than wide as in Citipes. However, all verified specimens of Citipes have surfaces wider than tall as in Elmisaurus.
While this and the other three figured metatarsals (TMP 1995.403.0010,
1984.163.0036 and 1986.036.0186) have surfaces taller than wide, that
has not been shown to be taxonomically informative. Note while the
figure caption states it and the other figured metatarsals are "from
the Upper Campanian Dinosaur Park Formation, Dinosaur Provincial Park,
Alberta, Canada" the TMP online catalogue shows this specimen is from
the Hell Creek Formation of Montana. While these metatarsi may be Elmisaurus or Citipes, the large coeval Anzu has not had its distal metatarsals described yet (though TMP 1996.005.0012 would have to be a juvenile if referred to it).
References-
Anonymous, 1997. Tyrannosaurus rex; A Highly Important and Virtually
Complete Fossil Skeleton. Sotheby’s. 56 pp.
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.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous
bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis,
University of Kansas. 227 pp.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
undescribed caenagnathid (Breithaupt, 1994)
Late Maastrichtian, Late Cretaceous
Lance Formation,
Wyoming
Comments- This is in a faunal list as Chirostenotes.
Reference- Breithaupt, 1994. Wyoming Dinosaur Diversity. Forty-Fourth
Annual Field Conference-1994. Wyoming Geological Association Guidebook. 101-104.
Caenagnathidae indet. (TMP online)
Late Cretaceous?
Alberta?, Canada?
Material- (TMP 1978.040.0001) metacarpal
Caenagnathidae indet. (TMP online)
Middle-Late Campanian, Late Cretaceous
Belly River Group or Bearpaw Formation, Alberta, Canada
Material- (TMP 1979.014.0799) phalanx
(TMP 1980.016.2095) incomplete pubis (Rhodes, Funston and Currie, 2020)
Caenagnathidae indet. (TMP online)
Middle-Late Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
Material- (TMP 1965.025.0036) phalanx (TMP online)
(TMP 1978.019.0002) vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0137) dorsal vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0138) sacral vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0339) manual phalanx (TMP online)
(TMP 1979.008.0366) vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0743) vertebra (TMP online, as Chirostenotes)
(TMP 1979.008.0823) metatarsal (TMP online, as Chirostenotes)
(TMP 1980.016.0810) cranial element (TMP online)
(UALVP 49064) dorsal vertebra (UALVP online)
Caenagnathidae indet. (Currie, 1992)
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Material- (TMP 1966.011.0051) vertebra (TMP online, as Chirostenotes)
(TMP 1967.013.0040) phalanx (TMP online, as Chirostenotes)
(TMP 1967.014.0031) ungual (TMP online, as Chirostenotes)
(TMP 1967.016.0038) ungual (TMP online, as Chirostenotes)
(TMP 1967.017.0035) metatarsal (TMP online, as Chirostenotes)
(TMP 1967.017.0038) ungual (TMP online, as Chirostenotes)
(TMP 1967.019.0147) pedal ungual (TMP online, as Chirostenotes)
(TMP 1967.020.0251) caudal vertebra (TMP online, as Chirostenotes)
(TMP 1973.023.0002) ungual (TMP online, as Chirostenotes)
(TMP 1975.011.0033) cervical vertebra (Sues, 1997)
(TMP 1979.008.0799) caudal vertebra (TMP online, as Chirostenotes)
(TMP 1979.014.0508) pedal phalanx (TMP online, as Chirostenotes)
(TMP 1979.014.0902) pedal phalanx (TMP online, as Chirostenotes)
(TMP 1979.014.0923) caudal vertebra (TMP online, as Chirostenotes)
(TMP 1980.008.0281) pedal ungual (TMP online, as Chirostenotes)
(TMP 1980.016.0556) dorsal vertebra (TMP online, as Chirostenotes)
(TMP 1980.016.0557) sacral vertebra (TMP online, as Chirostenotes)
(TMP 1980.016.0578) pedal phalanx (TMP online, as Chirostenotes)
(TMP 1980.016.0994) metatarsal (TMP online, as Chirostenotes)
(TMP 1980.016.1095) pedal phalanx (TMP online, as Caenagnathus)
(TMP 1980.016.1130) sacral vertebra (TMP online, as Chirostenotes)
(TMP 1980.016.1133) ungual (TMP online, as Chirostenotes)
(TMP 1980.016.1481) metatarsal (TMP online, as Chirostenotes)
(TMP 1980.016.1503) sacrum (Funston, 2020)
(TMP 1980.020.0184) ungual (TMP online, as Chirostenotes)
(TMP 1981.019.0252) parietals (Currie, 1992)
(TMP 1981.019.0285) sacrum (Funston, 2020)
(TMP 1982.016.0275) distal pubis, phalanx (Rhodes, Funston and Currie, 2020)
(TMP 1984.163.0102) sacrum (Funston, 2020)
(TMP 1984.167.0044) sacrum (Makovicky, 1995)
(TMP 1989.036.0109) caudal vertebra (Makovicky, 1995)
(TMP 1991.036.0146) caudal vertebra (Makovicky, 1995)
(TMP 1992.036.0053) proximal caudal vertebra (31.4 mm) (Currie et al., 1993)
(TMP 1994.012.0603) pubes (Rhodes, Funston and Currie, 2020)
(TMP 1995.403.0010) distal metatarsal III (Funston, Currie and Burns, 2016)
(TMP 1998.093.0013) posterior ilium (Rhodes, Funston and Currie, 2020)
(TMP 2001.012.0216) distal quadrate (Funston, 2020)
(TMP 2002.012.0103) partial ilium (Rhodes, Funston and Currie, 2020)
Comments- Only specimens through 1980 have been added from the TMP online catalogue so far. These are often referred to Caenagnathus or Chirostenotes
in the catalogue, but are retained as Caenagnathidae here as
differences between these geneera have been contentious and only
recently resolved.
TMP 1981.019.0252 and 1992.036.0053 were referred to Caenagnathus
sp. by Currie (1992) and Currie et al. (1993) respectively. Sues (1997)
referred TMP 1975.011.0033 (misidentified as a caudal on the TMP online catalogue) to Chirostenotes, while Makovicky (1995) referred TMP 1984.167.0044, 1989.036.0109 and 1991.036.0146 to that genus. The latter material did
not have locality information listed, but is probably from the Dinosaur Park
Formation of Alberta. TMP 1995.403.0010 is figured by Funston et al. (2016) as Leptorhynchus elegans (now Citipes), but seems too large. It may be Chirostenotes or young Caenagnathus
instead. Funston (2019) describes and figures partial ilium TMP
1998.093.0013, pubes TMP 1994.012.0603 and pubis TMP
1980.016.2095. The pubes differ from the larger one referred to Caenagnathus, so are believed by Funston to be either Chirostenotes or Citipes.
Reference- Currie, 1992. Saurischian dinosaurs of the Late Cretaceous
of Asia and North America. In Mateer and Chen (eds.). Aspects of Nonmarine
Cretaceous Geology. China Ocean Press. 237-249.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). Masters thesis, Copenhagen University. [pp]
Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria:
Theropoda) from western North America. Journal of Vertebrate Paleontology. 17(4),
698-716.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian)
of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution.
Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Rhodes, Funston and Currie, 2020. New material reveals the pelvic
morphology of Caenagnathidae (Theropoda, Oviraptorosauria). Cretaceous
Research. 114, 104521.
Caenagnathidae indet. (Eberth, Evans, Brinkman, Therrien, Tanke and Russell, 2013)
Middle Maastrichtian, Late Cretaceous
Tolman Member of Horseshoe Canyon Formation, Alberta, Canada
(UALVP 57349) (1 year old juvenile, ~8.0 kg) incomplete tibia (~210 mm) (Funston and Currie, 2018)
Early Maastrichtian, Late Cretaceous
boundary of Morrin and Tolman Members of Horseshoe Canyon Formation, Alberta, Canada
(WL-156) multiple elements (Eberth et al., 2013)
Comments- Eberth et al. (2013) lists WL-156 as a possible Epichirostenotes
specimen, though further details including its whereabouts given the subsequent death of Wann Langston
('WL' stands for his personal collection) are uncertain.
UALVP 57349 is a juvenile tibia found on July 23 2016 and described by
Funston and Currie (2018) as Caenagnathidae indet.. While the two
recognized Horseshow Canyon caenagnathid taxa preserve tibiae, only the
proximal end is exposed in Apatoraptor while only the distal end is preserved in Epichirostenotes. UALVP 57349 is indistinguishable from either, given its mostly broken proximal end and Epichirostenotes' unillustrated and briefly described distal end. The widely expanded distal end (236% of shaft width) is most similar to Elmisaurus (184%) as opposed to Anzu (158%), Chirostenotes (161%) or Citipes (132%).
References- Eberth, Evans, Brinkman, Therrien, Tanke and Russell, 2013.
Dinosaur biostratigraphy of the Edmonton Group (Upper Cretaceous), Alberta,
Canada: Evidence for climate influence. Canadian Journal of Earth Sciences.
50(7), 701-726.
Funston and Currie, 2018. A small caenagnathid tibia from the Horseshoe
Canyon Formation (Maastrichtian): Implications for growth and lifestyle
in oviraptorosaurs. Cretaceous Research. 92, 220-230.
undescribed caenagnathid (Ryan and Russell, 2001)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
Material- (RTMP 86.207.17) partial cervical vertebra
Comments- This was cited as Segnosauridae indet. by Ryan and
Russell (2001) in a faunal list, but Cullen et al. (2020) found it "to
be most similar in morphology to caenagnathids."
Reference- Ryan and Russell, 2001. Dinosaurs of Alberta (exclusive of
Aves). In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University
Press. 279-297.
Cullen, Larson, Zanno, Currie and Evans, 2020. Theropod biodiversity
patterns in the Dinosaur Park Formation (Late Cretaceous: Campanian) of
Alberta revealed through morphometrics and biostratigraphy. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 115.
unnamed Caenagnathidae (Gilmore, 1924)
Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- (CMN 346) manual ungual I (~130 mm on curve) (Bell, Currie and
Russell, 2015)
(CMN 8504) dorsal centrum (44 mm), three caudal centra (22-25 mm) (Gilmore,
1924)
(RSM P2161.1) metatarsal II (~211 mm) (Funston, Currie and Burns, 2016)
(TMM 41395-1) manual ungual II (Bell, Currie and Russell, 2015)
Comments- The centra were described by Gilmore (1924) as distinct from
other coelurosaurs known at the time, though possibly referrable to Chirostenotes
or Dromaeosauridae (neither of which were then known from vertebrae). Currie
et al. (1993) noted the caudals belonged to an oviraptorosaur, referring them
to Caenagnathus sp.. Bell et al. (2015) described two manual unguals
as cf. Anzu, noting they are comparable in size and age but that CMN
346 has a deeper concavity between the flexor tubercle and articular surface
than that genus. Elmisaurus sp. is also present in this formation based on RSM P2600.1. Funston et al. (2016) describe metatarsal II RSM P2161.1 as Leptorhynchos sp., but its large size and unfused distal tarsal are more like Chirostenotes or Caenagnathus. It lacks the large posterior protuberance on the proximal end seen in Caenagnathus, and has a more prominent posteromedial ridge as in Elmisaurus and Citipes.
References- Gilmore, 1924. A new coelurid dinosaur from the Belly River
Cretaceous Alberta. Canada Geological Survey, Bulletin n. 38, geological series
43, 1-13.
Tokaryak, 1990. It was here a minute ago. The Saskatchewan
Archaeological Society Newsletter. 11(2), 44-45.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
Bell, Currie and Russell, 2015 (online 2014). Large caenagnathids (Dinosauria, Oviraptorosauria)
from the uppermost Cretaceous of western Canada. Cretaceous Research. 52, 101-107.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
unnamed caenagnathid (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. 5-14.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
undescribed caenagnathid (Manabe and Barrett, 2000)
Valanginian-Hauterivian, Early Cretaceous
Kuwajima Formation of the Tetori Group, Japan
Material- (SBEI-167) manual ungual
Comments- Referred to the oviraptorosaur-therizinosaur clade by Manabe
et al. (2000), it provisionally resembles caenagnathids most closely.
References- Barrett and Manabe, 2000. The dinosaur fauna from the Earliest
Cretaceous Tetori Group of Central Honshu, Japan. Journal of Vertebrate Paleontology.
20(3), 28A-29A.
Manabe and Barrett, 2000. Dinosaurs. In Matsuoka (ed.). Fossils of the Kuwajima
"Kaseki-kabe" (fossil-bluff). Scientific report on a Neocomian (Early
Cretaceous) fossil assemblage from the Kuwajima Formation, Tetori Group, Shiramine,
Ishikawa, Japan. Shiramine Village Board of Education, Japan. 93-98.
Manabe, Barrett and Isaji, 2000. A refugium for relicts? Nature. 404, 953-954.
Matsuoka, Kusuhashi, Takada and Setoguchi, 2002. A clue to the Neocomian vertebrate
fauna: Initial results from the Kuwajima 'Kaseki-kabe' (Tetori Group) in Shiramine,
Ishikawa, central Japan. Memoirs of the Faculty of Science, Kyoto University,
Series of Geology and Mineralogy. 59(1), 33-45.
Microvenator Ostrom, 1970
= "Megadontosaurus" Ostrom, 1970
M. celer Ostrom, 1970
= "Megadontosaurus ferox" Brown vide Chure and McIntosh, 1989
Mid-Late Albian, Early Cretaceous
Himes Member, Cloverly Formation, Montana, US
Holotype- (AMNH 3041 in part) (~3.7 kg; juvenile) lacrimals(?) (one lost), quadrates(?)
(lost), dentary, prearticular (?) (lost), axis (13.9 mm), anterior cervical
neural arch, cervical centrum (15.1 mm), cervical centrum (15.4 mm), cervical
centrum (15+ mm), two cervical neural arches, two partial cervical ribs, three
anterior dorsal neural arches, anterior dorsal centrum (13.5 mm), anterior dorsal
centrum (13.8 mm), nine posterior dorsal neural arches, posterior dorsal centrum
(14.2+ mm), posterior dorsal centrum (13.8 mm), posterior dorsal centrum (13.9
mm), posterior dorsal centrum (14.1 mm), posterior dorsal centrum (13.7 mm),
posterior dorsal centrum (13.1 mm), two partial dorsal ribs, mid sacral centrum
(15.1 mm), caudal centrum (11.5 mm), caudal centrum (9.6+ mm), caudal centrum
(8.7 mm), caudal centrum (6.1+ mm), caudal centrum (9.9+ mm), caudal centrum
(9.8 mm), caudal centrum (9.7 mm), caudal centrum (9.5 mm), caudal centrum (10.1
mm), three caudal neural arches, coracoid, humerus (81.5 mm), radius, ulna (78
mm), ulnar fragment, metacarpal I (12.5 mm), phalanx I-1 (34 mm), manual ungual
I (25 mm), manual ungual III (11 mm), distal phalanx, partial ilia (~110 mm),
pubes (108.6 mm), fragmentary ischia, femora (124 mm), tibia (157 mm), proximal
fibula, astragalus (22.5 mm wide, 42 mm high), metatarsal I (11.8+ mm), pedal
ungual I (11.9 mm)
Diagnosis- (after Makovicky and
Sues, 1998) dorsal and caudal centra distinctly wider than high;
accessory trochanter at base of the anterior trochanter of the femur.
Other diagnoses- (after Ostrom, 1970) very small (approximately one half to two thirds the size of Ornitholestes or Coelurus),
delicately built; hollow, thin-walled vertebrae and limb bones.
cervicals without neural spines and with double pleurocels; dorsal
neural arches low and highly sculpted, neural spines low and
rectangular, postzygapophyses far behind posterior border of centrum;
pubis profile concave anteriorly, distal extremities only moderately
expanded; femur with short but prominent anterior trochanter and a
depression at site of fourth trochanter; astragalus with very high and
broad ascending process.
Comments-
The holotype was collected in 1933 but not described until 1970
(Ostrom, 1970). As noted by Ostrom, "associated with the type
specimen were 25 teeth of the Deinonychus
type which Barnum Brown believed belonged to this specimen."
"Because of the extremely large size of the associated teeth, this
specimen has been known informally in conversation by the name
"Megadontosaurus"" As Chure and McIntosh (1989) stated, Brown
proposed "Megadontosaurus ferox" with the intended holotype AMNH 3041,
but "never got around to
describing these animals, and it was left to John Ostrom to describe
and name them many years later in 1970, based in many cases on much
better and more diagnostic material which he had collected."
"None of
Brown's names were ever published (until Glut 1972 [for "Peltosaurus"])
and it does no harm to mention them here, as Ostrom's names are well
established taxa having full priority. Brown's names, however, were
used in public lectures, and for a time on some exhibited specimens. In
addition, they were written on photographs and preliminary skeletal
reconstructions made under Brown's direction." Ostrom writes
"without excveption, the 25 associated teeth duplicate in size and form
the teeth found with the type of Deinonychus antirrhopus", including the high DSDI. He referred them to that taxon, and noted "they are still catalogued with the Microvenator remains under AMNH 3041."
The tooth (YPM
5366) questionably referred to Microvenator
by Ostrom is unlikely to belong to an oviraptorosaur, as it is
plesiomorphically recurved and ziphodont, so is referred to Tetanurae
indet. here. The same is true of another tooth (MOR coll.)
referred to Microvenator by Maxwell (1993). Yun and Funston (2021) figured the first metatarsal in detail.
This genus has a controversial placement in recent analyses, from being
a caudipterid in Senter et al. (2012), avimimid in Lee et al. (2014),
sister to caenagnathoids in Brusatte et al. (2014) and Foth et al.
(2014), and basal caenagnathid in Funston and Currie (2016).
Hartman et al. (2019) recovered it as a basal oviraptorid, but with the
addition of more taxa it moved to the first branching caenagnathid.
References- Ostrom, 1970. Stratigraphy and paleontology of the Cloverly
Formation (Lower Cretaceous) of the Bighorn Basin area, Wyoming and Montana. Peabody Museum Bulletin. 35, 234 pp.
Chure and McIntosh, 1989. A Bibliography of the Dinosauria (Exclusive of the Aves)
1677-1986. Museum of Western Colorado Paleontology Series #1. 226 pp.
Maxwell, 1993. Neonate dinosaur remains and dinosaur eggshell from the Lower
Cretaceous Cloverly Formation of Montana. Journal of Vertebrate Paleontology.
13(3), 48A.
Makovicky and Sues, 1997. A reappraisal of the phylogenetic affinities of Microvenator
celer (Theropoda: Dinosauria) from the Cloverly Formation. Journal of Vertebrate
Paleontology. 17(3), 62A.
Makovicky and Sues, 1998. Anatomy and phylogenetic relationships of the theropod
dinosaur Microvenator celer from the Lower Cretaceous of Montana. American
Museum Novitates. 3240, 27 pp.
Senter, Kirkland, DeBlieux, Madsen and Toth, 2012. New dromaeosaurids (Dinosauria:
Theropoda) from the Lower Cretaceous of Utah, and the evolution of the dromaeosaurid
tail. PLoS ONE. 7(5), e36790.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247.
Yun and Funston, 2021. A caenagnathid oviraptorosaur metatarsal from
the Mesaverde Formation (Campanian) of Wyoming, USA. Vertebrate Anatomy
Morphology Palaeontology. 9, 105-115.
Caenagnathidae sensu Sues, 1997
Definition- (Chirostenotes pergracilis + Chirostenotes elegans
+ Elmisaurus rarus + Caenagnathasia martinsoni + BHM 2033)
Comments- Phylogeny and taxonomy within this group have been incredibly
contentious, along with which elements can be referred to each
species. The listed diagnoses are an exception to the norm for
this website in that they only compare to this group of caenagnathids
and not other taxa on this page like Microvenator or oviraptorids, even if avimimids are correctly nested within the clade.
Leptorhynchos Longrich, Barnes,
Clark and Millar, 2013b
= "Leptorhynchos" Longrich, Barnes, Clark and Millar, 2013a
L. gaddisi Longrich, Barnes, Clark and Millar, 2013b
= "Leptorhynchos gaddisi" Longrich, Barnes, Clark and Millar, 2013a
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
Holotype- (TMM 45920-1) (juvenile?) dentaries (symph 24.45 mm)
Diagnosis- Provisionally indeterminate relative to Chirostenotes and Elmisaurus.
Other diagnoses- Longrich et al. (2013a) diagnosed Leptorhynchos gaddisi based on comparison with TMP 1992.036.0390, a specimen they referred to elegans (as Leptorhynchos elegans, now separated as Citipes elegans). However, that specimen is here considered Chirostenotes pergracilis based on its large size and juvenile features. In turn, they diagnosed the genus Leptorhynchos based on comparison with TMP 1990.056.0006, another mandible of Chirostenotes pergracilis. Of characters listed as diagnostic for Leptorhynchos,
the shortness or depth of the dentary and dentary contribution to the
dorsal margin of the external mandibular fenestra cannot be evaluated
in the type as the posterior section is missing. An upturned dentary
tip and four symphyseal ridges are also present in Chirostenotes
(e.g. complete mandibles TMP 2001.012.0012). "Dentaries strongly
divergent, with posterior rami of dentaries forming an angle of roughly
35 degrees in dorsal view" is not true in the L. gaddisi holotype. Indeed, the diagnosis of L. gaddisi
lists "lateral occlusal margins of the dentaries not as strongly
divergent in dorsal view, with the tip of the beak being narrower and
more spoon-shaped in dorsal view", which is true as they show an angle
of about 20 degrees. Both this and the "short caudal symphyseal shelf"
fall within the range of variation of Chirostenotes (again compare to TMP 2001.012.0012 which has a ~20 degree angle and a shorter symphysis than gaddisi's holotype). Of the remaining two listed characters for L. gaddisi, "a strongly rounded anteroventral margin of the symphysis" is indistinguishable from some Chirostenotes (TMP 1992.036.0390) and "a more anteriorly projecting tip of the beak" than 1992.036.0390 is true in other Chirostenotes jaws like 2001.012.0012. Thus all suggested characters fall within the range of variation of Chirostenotes pergracilis. The size is relatively small but within the range of known juvenile Chirostenotes,
plus Funston et al. (2020) showed dentary fusion occurs even in those
juveniles, so this does not suggest "that the animal was mature" as
Longrich et al. claim. Intriguingly, the Leptorhynchos
holotype seems to show fibrous external bone texture (Longrich et al.'s
Fig. 8D below the label) and a poorly developed m. genioglossus
attachment, both of which Funston et al. suggested could be associated
with immaturity. Thus it could easily be a juvenile of a Chirostenotes-sized taxon. As it cannot presently be distinguished from Chirostenotes or the cf. Elmisaurus rarus dentaries IGM 102/107, it is here considered an indeterminate caenagnathid.
Comments- Some of this material was first reported as Caenagnathidae
indet. by Longrich et al. (2010). While Longrich et al. (2013a) intended to
name Leptorhynchos gaddisi, they did not specify a type species, making
the correction Longrich et al. (2013b) necessary to specify gaddisi as
the type species and make the names official.
Longrich et al. (2013a) proposed this species is sister to elegans, both
being closer to Chirostenotes than to Elmisaurus rarus. Their
phylogenetic analysis would seem to support this, but Elmisaurus rarus'
basal position is due to their figure 14 being an Adams consensus tree. Because
rarus lacks described mandibular material, it cannot be placed precisely
compared to Caenagnathasia and their Caenagnathus OTUs, so will
have a position at the base of the largest clade it can belong to in an Adams
consensus tree. When their analysis is rerun to a posteriori exclude taxa incomparable
to Elmisaurus rarus, it is in a trichotomy with Chirostenotes
and elegans. Further, their analysis miscodes rarus as lacking
metatarsal fusion and excludes the other five characters suggested by Currie
(1989) to unite rarus and elegans to the exclusion of pergracilis.
Longrich et al. do claim elegans is more similar to Chirostenotes
in one way. They state "Longrich (2008a) tentatively placed [elegans]
in Chirostenotes, because the third metatarsal has an anteroposteriorly
flattened shaft that is concave and broadly exposed on the posterior of the
metatarsus (Currie 1989). This is a derived feature found in Chirostenotes
(Currie and Russell 1988) but not Elmisaurus (Osm�lska 1981)."
This seems related to their new character 205- "Metatarsal III with an
ovoid or subtriangular cross section (0) or anteroposteriorly flattened, with
a concave posterior surface (1). Primitively in theropods the third metatarsal
has an ovoid cross section, or a triangular cross section in arctometatarsalian
forms. This condition is retained in most oviraptorosaurs, including the basal
caenagnathid Elmisaurus rarus. In Caenagnathinae, the third metatarsal
is anteroposteriorly compressed." Yet the posterior transverse exposure
of proximal metatarsal III proximally seems intermediate in E. rarus'
holotype compared to the two specimens of elegans. More distally, Currie's
(1989) figure 2P section indicates the posterior exposure is narrow as in E.
rarus. Currie's (1989) figure 2Q shows E. rarus has a concave posterior
metatarsal III surface as well. As for shape, the main issue seems to be the
use of different proximodistal points along the bone. Currie and Russell (1988)
state in Chirostenotes pergracilis "The proximal end, viewed dorsally,
is diamond shaped, tapering both anteriorly (between the contact of metatarsals
II and IV) and posteriorly. Its major horizontal axis, 17.5 mm long, is anteroposterior
in orientation and thins backwards." Sternberg (1932) also states the Macrophalangia
holotype (possibly Caenagnathus collinsi according to Longrich et al.)
has a transversely compressed proximal end. The proximal ends of E. rarus'
and elegans' metatarsal III are fused too well with surrounding bones
to compare. Once C. pergracilis' anterior surface is exposed, "the
bone twists until the medial surface is facing anteriorly" and "the
anterior edge has broadened out to 7.5 mm to separate the adjacent metatarsals
and is triangular in section" (Currie and Russell, 1988). elegans'
holotype is broken at about this same point and also shows a triangular section.
Note a triangular section is what Longrich et al. are claiming caenagnathines
don't have, though it exists in both pergracilis and elegans.
Currie (1989) was also wrong in comparing the proximal diamond shape of Chirostenotes
pergracilis with the more distal triangular shape of elegans, and
none of these areas have been described in E. rarus. At two-thirds down
in Chirostenotes, "In section, a shallow concave surface faces posteriorly
at this level, while slightly concave surfaces face posteromedially and posterolaterally
for contact with the adjacent metatarsals." This appears similar to E.
rarus from what the anterior and posterior views suggest, and matches how
Snively (2000) described the specimen. It also matches the cross section of elegans
illustrated by Currie (1989- fig. 2P). While Currie's figure 2Q of Elmisaurus
rarus would suggest a slightly different shape where the articular surfaces
are smaller and that for metatarsal IV doesn't angle posteromedially, the narrow
anterior exposure of metatarsal III means it must have been taken more proximally,
probably about halfway down considering the ratio between anterior and posterior
exposure of metatarsal III. This leaves anteroposterior compression, which varies
throughout the bone in elegans at least. Distally it's transversely compressed
but proximally it's anteroposteriorly compressed. The E. rarus section
which is probably intermediate in position is also intermediate in compression,
being slightly transversely compressed. If Chirostenotes
specimens are any indication, the bone switches back to transversely compressed
at its proximal tip. So there are actually no valid described differences in
metatarsal III sectional shape between Chirostenotes and any Elmisaurus
species in the primary literature.
As discussed above under Other diagnoses, the Leptorhynchos gaddisi
holotype is indeterminate and possibly juvenile. None of the referred
material is diagnostic, but the small and fused distal tarsal and
metatarsal IV (TMM 43057-36) are like Citipes and Elmisaurus.
If anything, this suggests the large Aguja ungual and femur (TMM
42920-2 and UTEP B38 L-3) described by Longrich et al. might belong to
adult Leptorhynchos/Chirostenotes, while TMM 43057-36 is a separate smaller species, perhaps Citipes.
The small caudal centrum (TMM 42335-40), distal metatarsal III (TMM
43057-354) and pedal ungual (TMM 43057-357) are here placed in
Caenagnathidae indet., as they could be juveniles, while the proximal
partial tarsometatarsus is provisionally referred to the
contemporeneous Citipes elegans.
References- Sternberg, 1932. Two new theropod dinosaurs from the Belly
River Formation of Alberta. Canadian Field-Naturalist. 46(5), 99-105.
Currie and Russell, 1988. Osteology and relationships of Chirostenotes pergracilis
(Saurischia, Theropoda) from the Judith River (Oldman) Formation of Alberta,
Canada. Canadian Journal of Earth Sciences. 25(7), 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda)
from North America. Canadian Journal of Earth Sciences. 26(6), 1319-1324.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Masters Thesis, University of Calgary. 273 pp.
Longrich, Sankey and Tanke, 2010. Texacephale langstoni, a new genus
of pachycephalosaurid (Dinosauria: Ornithischia) from the upper Campanian Aguja
Formation, southern Texas, USA. Cretaceous Research. 31, 274-284.
Longrich, Barnes, Clark and Millar, 2013a. Caenagnathidae from the Upper Campanian
Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin
of the Peabody Museum of Natural History. 54(1), 23-49.
Longrich, Barnes, Clark and Millar, 2013b. Correction to "Caenagnathidae
from the Upper Campanian Aguja Formation of west Texas, and a revision of the
Caenagnathinae". Bulletin of the Peabody Museum of Natural History. 54(2),
263-264.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020 (online
2019). Histology of caenagnathid (Theropoda, Oviraptorosauria)
dentaries and implications for development, ontogenetic edentulism, and
taxonomy. The Anatomical Record. 303(4), 918-934.
Ojoraptorsaurus Sullivan,
Jasinski and van Tomme, 2011
O. boerei Sullivan, Jasinski and van Tomme, 2011
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Holotype- (SMP VP-1458) (~1.8-2.1 m) incomplete pubes (~344 mm)
Diagnosis-
(after Sullivan et al., 2011) iliac peduncle articular surface of pubes narrower anteriorly than posteriorly (also in Microvenator, Caenagnathus, Chirostenotes and Chirostenotes/Citipes TMP 1994.012.0603; unknown in other caenagnathids except Epichirostenotes);
proximomedial pubic fossa recessed distally from acetabular rim at
least one third the narrowest anteroposterior width of the ilial
peduncle (unknown in other caenagnathids except Microvenator, Anzu, Epichirostenotes, Chirostenotes/Citipes TMP 1980.016.2095 and Elmisaurus?); oval
depression on the anterodorsal surface of pubic boot (also in Microvenator and Epichirostenotes; unknown in other caenagnathids except Anzu and Chirostenotes/Citipes TMP 1994.012.0603); pubic shaft strongly convex
anteriorly just proximal to pubic boot (unknown in other caenagnathids except Microvenator, Anzu, Caenagnathus, Epichirostenotes?, Chirostenotes/Citipes TMP 1980.016.2095 and TMP 1994.012.0603 and Elmisaurus?).
Other diagnoses- Sullivan et al. (2011) also listed differences from Epichirostenotes
"in being 20% smaller and relatively more robust, lacking a dorsal
suture on the pubic boot, and having a significantly lower pubic
apron/pubis length ratio." However, all other caenagnathid pubes
are more robust than Epichirostenotes'
seems to be, perhaps due to the amount of crushing in that taxon's
holotype. The small size difference with undescribed histology in
each specimen means little, and that "there is a trace of a suture" on
the dorsal surface of Epichirostenotes'
pubis is not a strong indicator of a less mature specimen. The
apron/pubis ratio (supposedly 38% vs. 49%) actually refers to the
distance from the distal interpubic foramen tip to the proximomedial
corner of the apron, based on its stated length in Ojoraptorsaurus. Yet this is 39% in Epichirostenotes according to Sues' figure, comparable to Anzu's 38% and Ojoraptorsaurus' estimated 38% (Caenagnathus and Chirostenotes or Citipes
TMP 1980.016.2095 are now known to be shorter at 33% and 30%
respectively). So it seems Sullivan et al. overestimated the
length in Epichirostenotes.
Comments- The holotype was discovered in summer 2002.
The validity of this taxon was initially uncertain, as it was only compared
to Nomingia (here an oviraptorid), Epichirostenotes and Anzu by Sullivan et al. (2011) and the distribution of its diagnostic features in these
taxa and other oviraptorosaurs was poorly understood. Since then, pubes of Caenagnathus, Chirostenotes or Citipes and Elmisaurus have been described. These show Ojoraptorsaurus to be diagnosable, although the sub-trapezoidal ilial articulation would seem to be a symplesiomorphic difference from Epichirostenotes. The amount of separation of the proximomedial fossa from the acetabulum should be determinable in Caenagnathus and Elmisaurus as well, but is currently undescribed and unfigured.
Reference- Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid
Ojoraptorsaurus boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria),
from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member), San Juan
Basin, New Mexico. New Mexico Museum of Natural History and Science Bulletin.
53, 418-428.
Kuszholia Nessov, 1992
?= Caenagnathasia Currie, Godfrey and Nessov, 1993
K. mengi Nessov, 1992
?= Caenagnathasia martinsoni Currie, Godfrey and Nessov, 1993
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (ZIN PO 4602) posterior synsacrum (11, 10 mm)
Paratypes- ?(ZIN PO 4623) anterior synsacral centra (13, 10 mm)
?(ZIN PO coll.) sacral vertebrae (~20-30 mm)
Referred- ?(CCMGE 401/12457; holotype of Caenagnathasia martinsoni)
anterior dentaries (symph 11.2 mm) (Currie, Godfrey and Nessov, 1993)
?(CCMGE 402/12457; paratype of Caenagnathasia martinsoni) incomplete
dentary (symph 8.94 mm) (Currie, Godfrey and Nessov, 1993)
?(CCMGE 479/12457) proximal femur (Nessov, 1995; described by Sues and Averianov,
2015)
?(ZIN PH 802/16) four incomplete fused sacral vertebrae (17, 14, 16.3, 16.4
mm) (Sues and Averianov, 2015)
?(ZIN PH 932/16) incomplete posterior cervical vertebra (17 mm) (Sues and Averianov,
2015)
?(ZIN PH 933/16) incomplete anterior cervical vertebra (19.5 mm) (Sues and Averianov,
2015)
?(ZIN PH 934/16) incomplete dorsal vertebra (16.4 mm) (Sues and Averianov, 2015)
?(ZIN PH 935/16) dorsal vertebra (Sues and Averianov, 2015)
?(ZIN PH 936/16) dorsal vertebra (Sues and Averianov, 2015)
?(ZIN PH 937/16) posterior dorsal centrum (Sues and Averianov, 2015)
?(ZIN PH 2354/16) anterior dentaries (symph 7.5 mm) (Sues and Averianov, 2015)
?(ZIN PO 4603) incomplete anterior cervical vertebra (16 mm) (Nessov, 1992;
described by Sues and Averianov, 2015)
?(ZIN PO 5234) partial cervical vertebra (Sues and Averianov, 2015)
? (lost) mandible with teeth(?) (Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020)
Diagnosis- (after Currie et al., 1993)
first anterior occlusal groove much larger than second grooves (when present) (unknown in other caenagnathids except Caenagnathus, Chirostenotes and Elmisaurus); lateral occusal ridges
do not meet anteriorly (also in Anzu, Chirostenotes, Elmisaurus and Citipes; unknown in other caenagnathids except Caenagnathus); lingual ridge with taller and sharper edge (the cause of their "lateral groove is narrower and deeper") (also in Caenagnathus; unknown in other caenagnathids except Anzu, Chirostenotes, Elmisaurus and Citipes); defined medial edges to fossae between lingual grooves (unknown in other caenagnathids except Anzu, Caenagnathus, Chirostenotes, Elmisaurus and Citipes); vascular grooves not conspicuous
on symphysial shelf (also in Caenagnathus and Elmisaurus; unknown in other caenagnathids except Anzu, Chirostenotes and Citipes);
no foramina on the floor of median depression posterior to first
anterior occlusal groove (unknown in other caenagnathids except Anzu?, Caenagnathus, Chirostenotes, Elmisaurus and Citipes).
(suggested) large sacral pleurocoels (unknown in other caenagnathids except Chirostenotes, Epichirostenotes and Citipes).
Other diagnoses- None of the characters listed in Nessov's (1992) diagnosis
of Kuszholia are very characteristic. Dorsoventrally compressed sacral
centra are common in maniraptorans. The robust second to last sacral transverse
process is seen in Chirostenotes and Epichirostenotes as well. Deep posterior sacral pleurocoels
are common in caenagnathoids, and far from being small, Kuszholia's are
large compared to other taxa. The posterior articular surface is not large,
being smaller than mid sacral vertebrae, and its concavity is plesiomorphic
for theropods. The large postzygapophyses are only notable compared to birds-
they are normal for an oviraptorosaur. The sacrals of Chirostenotes also
have a ventral median groove which is most pronounced at the junction of centra.
Currie et al. (1993) stated there was no second anterior occlusal groove in
Caenagnathasia,
but Sues and Averianov (2015) reported it as present in new specimen
ZIN PH 2354/16 and showed it is present although small on the right
side of the holotype as well. Currie et al. also stated the "fluting on
the lingual margin of the occlusal edge is not as distinct in the Asian
jaws, and there are no toothlike apical projections on the ridges"
unlike Caenagnathus (including Chirostenotes), but this is untrue for ZIN PH 2354/16. They said "there are no tubercles on
the midline or at the base of the first lateral occlusal ridge", but Funston et al. (2020a) found that in Chirostenotes
"some dentaries lack these nodules altogether (TMP 1990.056.0006; TMP
2001.012. 0012). Therefore, the presence or absence of these rugose
patches is probably best explained by individual variation." Indeedd,
a tubercle is on the midline in ZIN PH 2354/16.
Comments- Note that while volume 30(10) of the Canadian Journal of Earth
Sciences lists its date as October 1993, it was not published until February
or March of 1994.
The holotype (ZIN PO 4602) consists of the last two sacral vertebrae and a fragment
of the third to last, fused together. The centra are dorsoventrally compressed
(anterior articular surface ~66% as tall as wide) with an oval and slightly
concave posterior articular surface. The sacrum seems to be slightly concave
ventrally and the centrum junctions are expanded both ventrally and laterally.
There are small but deep pleurocoels present in each centrum. A median ventral
groove is present, which is especially well marked at the centrum junctions.
The postzygapophyses of the last sacral protrude markedly past the centrum.
The second to last transverse process is long, robust and perpendicular to the
sacral long axis, while the last transverse process is about half the length
but otherwise similar.
Nessov referred another specimen (ZIN PO 4623) consisting of the first two fused
centra of another sacrum. These are similar in being dorsoventrally compressed
(posterior articular surface ~63% as tall as wide). The anterior articular surface
is kidney-shaped and slightly heterocoelous. It is also similar in being slightly
concave, with expanded centrum junctions and deep, oval pleurocoels in each
centrum. Ventrally, there is a slight midline groove. This was later referred
to Caenagnathasia by Sues and Averianov (2015). Nessov referred to additional
isolated vertebrae virtually identical to these, but 2-2.5 times larger. These
may belong to larger oviraptorosaurs, therizinosaurs or dromaeosaurids, though
they are impossible to evaluate without more information. Finally, he stated
strongly pneumatic vertebrae with closed neurocentral sutures (unlike juvenile
therizinosaurs or sauropods) could belong to Kuszholia. One is illustrated
(ZIN PO 4603), which is an anterior cervical with strongly overhanging prezygapophyses,
a large neural canal, an elongate centrum (2.75 times posterior height) which
reaches posteriorly past the neural arch, and perhaps a large teardrop shaped
pleurocoel. This was later referred to Caenagnathasia by Sues and Averianov
(2015).
Nessov and Panteleev (1993) figured and described a partial sacrum they referred
to Kuszholia sp. (ZIN PO 4826). Zelenkov and Averianov (2011) stated
this differs from Kuszholia in "the absence of a pleurocoel in the
posterior vertebra and in the shallow slitlike pleurocoel in the penultimate
vertebra", and while the pleurocoel shape appears similar, the absence
of a pleurocoel in the last sacral is indeed different. They believe the specimen
to be similar to Zanabazar, but I don't see any particular resemblence
and refer it to Maniraptora incertae sedis here pending further study. Nessov
(1995) later figured a centrum with a slit-like pleurocoel (ZIN PO 467) which
he stated was "possibly from bird ?Kuszholia sp. or from a theropod
or a segnosaur", and this was identified as a therizinosauroid by Sues
and Averianov (2016). He also figured proximal femur CCMGE 479/12457 as a theropod
before it was referred to Caenagnathasia by Sues and Averianov (2015).
Funston et al. (2020b) report "a specimen of Caenagnathasia martinsoni
that bore teeth was allegedly collected (J. Stiegler pers. comm.), but
was lost in transit", which would be interesting as Funston et al.
(2020a) concluded "the occlusal structures in caenagnathids cannot be
homologous to alveoli, because they fail Patterson’s (1982) test of
similarity by differing in developmental and histological nature."
Relationships- The holotype was first incorrectly listed as ZIN PO 3486
and identified as an ichthyornithid (Nessov, 1990). Nessov (1992) later named
this as a new genus in its own family Kuszholiidae, under "suborder Theropoda
+ Aves." He viewed it as possibly a basal flightless bird like Patagopteryx
or possibly a non-bird theropod which was convergent with birds. Nessov and
Panteleev (1993) later assigned it to Patagopterygiformes, though their argument
has yet to be translated from Russian. Kurochkin (2001) retained Kuszholia
as Aves (sensu Chiappe) incertae sedis, and noted that patagopterygiform affinities
were not yet verified. He did cite two supposed similarities though- enlarged
third pair of sacral transverse processes and ventral sacrum convex. Yet Kuszholia's
sacrum is ventrally concave (as in Patagopteryx and many other theropods)
and the large transverse processes are on the second to last sacral. As all
theropods have at least five sacrals, this corresponds to the fourth sacral
or greater. All posterior sacral transverse processes are broken off in Patagopteryx
in any case. Patagopteryx further differs in lacking sacral pleurocoels
and having a convex posterior articular surface, as noted by Kurochkin. Kurochkin
later (2006) placed Kuszholia in Euornithes (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 and caenagnathoids.
Adult therizinosaurs are far larger, with even the smallest basal members (e.g.
Beipiaosaurus) being four times as big, and the fusion does indicate
Kuszholia's holotype is from an adult. There are small caenagnathoids
though, including Caenagnathasia from the same formation. In fact, Caenagnathasia
would have comparably sized sacrals to Kuszholia if scaled from other
caenagnathoids. Note Sues and Averianov (2015) assigned the Kuszholia
paratype and referred cervical to Caenagnathasia without commenting on
why the holotype was also not referrable. Indeed, there seems to be no reason
to refer any of the Bissekty caenagnathoid material to one genus instead of
the other as the type synsacrum of Kuszholia cannot be compared to the
type dentaries of Caenagnathasia. If they are synonymous, Kuszholia
would have priority by two years, which is awkward as its synsacrum is less
diagnostic and less phylogenetically determinable than Caenagnathasia's
dentaries. The alternative is listing all material except the three dentaries
and Kuszholia's holotype as Caenagnathoidea/idae indet., but tentative
synonymization is chosen here.
The holotype is generally similar to Chirostenotes, but differs in having
much larger pleurocoels and narrower postzygapophyses. It also
has larger pleurocoels than Epichirostenotes, Citipes and Shixinggia. Avimimus
is unique among oviraptorosaurs in lacking sacral pleurocoels, so is
quite different. Funston and Currie (2016) found the composite OTU
(not including the Kuszholia holotype) to be an elmisaurine caenagnathid between Caenagnathus and Chirostenotes. Adding taxa to Hartman et al.'s matrix recovers the Caenanagathasia dentaries in Avimimus, but the Kuszholia composite in Elmisaurinae as derived as Elmisaurus but outside Shixinggia+Avimimus.
References- Nessov, 1990. Small ichthyornithiform bird and other bird
remains from Bissekty Formation (Upper Cretaceous) of central Kyzylkum Desert.
Proceedings of the Zoological Institute, Leningrad. 210, 59-62.
Nessov, 1992. Review of localities and remains of Mesozoic and Paleogene birds
of the USSR and the description of new findings. Russkii Ornitologicheskii Zhurnal.
1(1), 7-50.
Nessov and Panteleev, 1993. On the similarity of the Late Cretaceous ornithofauna
of South America and Central Asia. Trudy Zoologicheskogo Instituta, RAN. 252,
84-94.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of class Aves.
Archaeopteryx. 13, 47-66.
Nessov, 1995. Dinosaurs of nothern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
Nessov, 1997. Cretaceous nonmarine vertebrates of Northern Eurasia. Izdatelstvo
Sankt-Peterburgskogo Universiteta, Saint Petersburg. 218 pp.
Kurochkin, 2001. Mesozoic birds of Mongolia and the former USSR. In Benton,
Shishkin, Unwin and Kurochkin (eds.). The Age of Dinosaurs in Russia and Mongolia.
533-559.
Kurochkin, 2006. Parallel evolution of theropod dinosaurs and birds. Entomological
Review. 86(suppl. 1), S45-S58.
Zelenkov and Averianov, 2011. Synsacrum of a primitive bird from the Upper Cretaceous
of Uzbekistan. Paleontological Journal. 45(3), 314-319.
Sues and Averianov, 2015 (online 2014). New material of Caenagnathasia martinsoni
(Dinosauria: Theropoda: Oviraptorosauria) from the Bissekty Formation (Upper
Cretaceous: Turonian) of Uzbekistan. Cretaceous Research. 54, 50-59.
Sues and Averianov, 2016 (online 2015). Therizinosauroidea (Dinosauria: Theropoda) from the
Upper Cretaceous of Uzbekistan. Cretaceous Research. 59, 155-178.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020a (online
2019). Histology of caenagnathid (Theropoda, Oviraptorosauria)
dentaries and implications for development, ontogenetic edentulism, and
taxonomy. The Anatomical Record. 303(4), 918-934.
Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020b. A new
two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.
Royal Society Open Science. 7: 201184.
K. sp. (Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- (IVPP V20377) anterior dentaries (symph 8.0 mm)
Comments-
This was discovered in 2012 at "a rare microvertebrate locality within
the Iren Dabasu Formation, about 16 km northeast of Erenhot City",
which would put it in localities Q-T of Xing et al. (2012). Yao
et al. (2015) referred IVPP V20377 to Caenagnathasia
sp., and it does possess all characters here listed as diagnostic for
that taxon. Of their characters listed as varying between Caenagnathasia specimens, most also vary between Chirostenotes
specimens (posterior surface of symphysis with tubercle; chin-like
eminence between anterior and ventral surfaces; pneumatic foramen in
front of mandibular fenestra on lateral surface of dentary; depression
on posteroventral margin of symphysis [ontogenetic?]), while paired
second anterior occlusal grooves flanking first anterior occlusal
groove is polymorphic in the holotype, and lateral projections on
lingual ridges may be absent in ZIN PH 2354/16 due to preservation.
This leaves presence of a median symphyseal groove on the posterodorsal
depression in the holotype and IVPP V20377 but not ZIN PH 2354/16
(ontogenetic?) and on the posteroventral symphysis of IVPP V20377 but
not the holotype or ZIN PH 2354/16 (taxonomic?). The age difference
suggests the specimen is not conspecific with the Bissekty species.
References-
Xing, He, Li and Xi, 2012. A review on the study of the stratigraphy,
sedimentology, and paleontology of the Iren Dabasu Formation, Inner
Mongolia. In Dong (ed.). Proceedings of the Thirteenth Annual Meeting
of the Chinese Society of Vertebrate Paleontology. China Ocean Press.
1-44.
Yao, Wang, Sullivan, Wang, Stidham and Xu, 2015. Caenagnathasia
sp. (Theropoda: Oviraptorosauria) from the Iren Dabasu Formation (Upper Cretaceous:
Campanian) of Erenhot, Nei Mongol, China. Vertebrata PalAsiatica. 53(4), 291-298.
Wang, Zhang and Yang, 2018. Reevaluation of the dentary structures of
caenagnathid oviraptorosaurs (Dinosauria, Theropoda). Scientific
Reports. 8:391.
Caenagnathinae Sternberg, 1940 sensu Paul, 1988
Definition- (Caenagnathus collinsi <- Oviraptor philoceratops,
Avimimus portentosus) (Martyniuk, 2012)
Other definitions- (Caenagnathus collinsi <- Elmisaurus elegans,
Caenagnathasia martinsoni) (Longrich, Barnes, Clark and Millar, 2013)
(Caenagnathus collinsi <- Elmisaurus rarus) (Hendrickx, Hartman
and Mateus, 2015)
Comments- Longrich et al. (2013) thought elegans was a caenagnathine,
so no doubt meant to use Elmisaurus rarus as the external specifier instead.
References- Sternberg, 1940. A toothless bird from the Cretaceous of Alberta. Journal of
Paleontology. 14(1), 81-85.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster: New York
464 pp.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged
Dinosaurs. Vernon, New Jersey. Pan Aves. 189 pp.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian
Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin
of the Peabody Museum of Natural History. 54(1), 23-49.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian theropod discoveries
and classification. PalArch's Journal of Vertebrate Palaeontology. 12(1), 1-73.
Anzu Lamanna, Sues, Schachner and Lyson,
2014
A. wyliei Lamanna, Sues, Schachner and Lyson, 2014
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Holotype- (CM 78000) (~3.5 m; ~284 kg) premaxillary fragments, quadrates,
pterygoids, incomplete braincase, cranial fragments, incomplete mandibles (320
mm, symph 80 mm), six cervical vertebrae, dorsal ribs, gastralia, nine caudal vertebrae,
twenty-eighth caudal vertebra (33.9 mm), twenty-ninth caudal vertebra (37.2
mm), partial thirtieth caudal vertebra, six chevrons, scapulocoracoids (scap
415 mm, cor 121 mm), humerus (345 mm), radius (280 mm), ulna (280 mm), phalanx
I-1 (123.5 mm), manual ungual I (89.1 mm, 130 mm on curve), metacarpal II (139
mm), phalanx II-1 (131.8 mm), phalanx III-3 (82.8 mm), femora (525 mm), tibiae
(660 mm), fibulae (585, 580 mm), astragalocalcanea (98.5, 98.0 mm transversely),
metatarsals I (76 mm), phalanx I-1 (101.1 mm), phalanx III-1 (114.2 mm), pedal
ungual III (101.1 mm), phalanx IV-2 (63.7 mm), phalanx IV-3 (52.5 mm), phalanx
IV-4 (56.5 mm), five phalanges, two pedal unguals, partial metatarsal V
Paratypes- (CM 78001) premaxillae (one incomplete, one fragmentary),
maxillae, jugals, quadrates, ectopterygoid, pterygoids, braincase, cranial fragments,
atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical
vertebra, sixth cervical vertebra, seventh cervical vertebra, eighth cervical
vertebra, ninth cervical vertebra (~82.9 mm), tenth cervical vertebra, eleventh
cervical vertebra (~78.5 mm), twelfth cervical vertebra, ten dorsal vertebrae
(anterior 47 mm), seventeen ribs, eleven gastralia, incomplete sacrum (315 mm),
twelve caudal vertebrae, eight chevrons, sternal plates (194.2 mm), ilia, pubes
(450, 465 mm), ischia (305 mm), femora (505, 500 mm), tibiae (595 mm), fibulae
(570 mm), astragalocalcanea (109.7 mm), phalanx III-3 (85.9 mm), pedal phalanx,
four pedal unguals, metatarsal V (114.4 mm)
(FMNH PR2296 = BHM 2033) posterior mandible
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Referred- (BMRP 2013.4.1) dorsal ribs, caudal vertebra, pelvic elements,
distal hindlimb including incomplete metatarsi (Holtz, Williams, Tremaine and
Matthews, 2014)
?(MOR 9722) (~1.5-2 m) incomplete axial series, ribs, gastralia, partial forelimb
including manual phalanges and unguals, ilia, partial pubes, ischia, partial
tibia, tarsals, proximal metatarsals, pedal phalanges and pedal unguals (Flora, Wilson, Gardner and Fowler, 2015)
(ROM 65884) (8 year old subadult) three dorsal vertebral fragments,
several dorsal rib fragments, several gastralial fragments, incomplete
~twenty-seventh caudal vertebrae (30.9 mm), ~twenty-eighth caudal
vertebra (31.4 mm), ~twenty-ninth caudal vertebra (35.2 mm), distal
manual phalanx II-1, pubic fragments, tibial fragments, incomplete
fibula, incomplete pes (sold), partial metatarsal II, partial
metatarsal III, partial metatarsal IV, metatarsal V, metatarsal
fragments (Benner, Cullen and Evans, 2016)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, North Dakota, US
Paratype- (MRF 318) ninth cervical vertebra, eleventh cervical vertebra,
twelfth cervical vertebra, dorsal rib, scapulocoracoid, radius (275 mm), ulna
(280 mm)
Late Cretaceous?
USA?
Referred- (BDM uncatalogued) metatarsal I (Yun and Funston, 2021)
Diagnosis-
(after Lamanna et al., 2014) tall, crescentic cranial crest formed by
posterodorsal processes of premaxillae (unknown in other
caenagnathids); body of maxilla lacking antorbital fossa (unknown in
other caenagnathids except Epichirostenotes); maxillary ascending process elongate
and shaped like an inverted L (unknown in other caenagnathids); quadratojugal
process of jugal dorsoventrally deep (unknown in other caenagnathids); quadratojugal process of jugal bifurcated posteriorly
(unknown in other caenagnathids); occipital condyle transversely wider than
foramen magnum (unknown in other caenagnathids except Epichirostenotes);
prominent lateral flange on symphyseal region of dentary (unknown in other caenagnathids
except Caenagnathus, Chirostenotes, Elmisaurus, Citipes and Kuszholia); elongate retroarticular
process of mandible (subequal in anteroposterior length to quadrate articulation) (unknown in other caenagnathids except
for Caenagnathus, Chirostenotes and Apatoraptor) ;
distal end of radius divided into two rounded processes (unknown in other caenagnathids);
sulcus on ventromedial aspect of manual phalanx II-1 (unknown in other caenagnathids except Hagryphus, Chirostenotes and Elmisaurus); tubercle on anterior surface
of astragalus near base of ascending process (unknown in other caenagnathids
except Chirostenotes).
(after Funston and Currie, 2016) rounded hypapophyses in anterior dorsal vertebrae (unknown in other caenagnathids except Apatoraptor and Epichirostenotes); fenestra in posterolateral sternum (unknown in other caenagnathids except Apatoraptor).
Comments- Currie et al. (1993) described a posterior mandible (their BHM 2033) which was
larger than any Dinosaur Park Caenagnathus and differed from C. collinsi
and C. sternbergi in glenoid morphology. They referred it to Caenagnathus
sp.. Varricchio (2001) further noted it shared several characters with C.
collinsi to the exclusion of C. sternbergi, and Lamanna et al. (2014)
referred it to the present species.
Triebold et al. (2000) reported two new large oviraptorosaur specimens discovered in 1998, identifying
them as oviraptorids, though they were quickly reidentified as caenagnathid.
These specimens were officially described by Lamanna et al. (2014) as the new
taxon Anzu wyliei. The mandible is most similar to the Caenagnathus
collinsi holotype.
Schachner et al. (2006; 2007) announced MRF 318 as Chirostenotes sp.,
which was later described as a specimen of Anzu by Lamanna et al. (2011;
2014).
Holtz et al. (2014, 2015) announced a new specimen (BMRP 2013.4.1- Holtz pers.
comm, 2015) found in June 2013, which has only distal tarsal IV fused to the metatarsus, unfused
metatarsals, and "a pair of pronounced cruciate ridges on the plantar surface
of metatarsal III" which don't extend proximally as far as Elmisaurus.
ROM 65884 was found in 2012, and unfortunately a "nearly complete right
foot was also collected, assembled into a display mount, and sold to a
private collector" before the ROM bought the rest of the material
(Cullen et al., 2020). Initially reported in an SVP abstract by
Benner et al. (2016) as "a large-bodied taxon similar in size to the
recently described, coeval taxon Anzu wyliei", Cullen et al. described the specimen as
cf. Anzu wyliei, as it is
from the same formation and almost identical in anatomy and size ("c.
3% difference in linear size, on average"). Indeed, the slightly
curved metatarsal V is shared with Anzu but not Chirostenotes, Elmisaurus or seemingly Macrophalangia (+ Caenagnathus?).
While this specimen was found in the same state as BMRP 2013.4.1 and
preserves almost the same elements, they were discovered in different
years and different counties and both metatarsi of BMRP 2013.4.1 were
recovered by the Burpee Museum.
Yun and Funston (2021) figure the metatarsal I of the holotype in
detail, as well as an additional Anzu metatarsal I called "BDM
uncatalogued specimen," for which no other information is available in
the literature.
Funston and Currie (2016) recover Anzu as basal to the caenagnathine-elmisaurine split, sister to Epichirostenotes.
References-
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
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. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life.
New Research Inspired by the Paleontology of Philip J. Currie. Indiana University
Press. 42-57.
Schachner, Lyson and Hanks, 2006. A preliminary report of a new specimen of
Chirostenotes (Oviraptorosauria: Theropoda) from the Hell Creek Formation
of North Dakota. Journal of Vertebrate Paleontology. 26, 120A.
Schachner, Lyson, Atterholt and Hanks, 2007. A preliminary report of a new specimen
of Chirostenotes (Oviraptorosauria: Theropoda) from the Hell Creek Formation
of North Dakota. Journal of Vertebrate Paleontology. 27(3), 141A.
Lamanna, Sues, Schachner and Lyson, 2011. A new caenagnathid oviraptorosaur
(Theropoda: Maniraptora) from the Upper Cretaceous (Maastrichtian) Hell Creek
Formation of the western United States. Journal of Vertebrate Paleontology.
Program and Abstracts 2011, 140.
Holtz, Williams, Tremaine and Matthews, 2014. New additions to the Hell Creek
Formation (Upper Maastrichtian) vertebrate fauna of Carter County, Montana.
Journal of Vertebrate Paleontology. Program and Abstracts 2014, 149.
Lamanna, Sues, Schachner and Lyson, 2014. A new large-bodied oviraptorosaurian
theropod dinosaur from the Latest Cretaceous of western North America. PLoS
ONE. 9(3), e92022.
Flora, Wilson, Gardner and Fowler, 2015. A three-dimensionally articulated probable
oviraptorosaur from the Hell Creek Formation of Montana. Journal of Vertebrate
Paleontology. Program and Abstracts 2015, 124-125.
Holtz, Williams and Tremaine, 2015. A new specimen of Anzu (Caenagnathidae,
Oviraptorosauria): Implications for the proposed Caenagnathinae/Elmisaurinae
division and for cursoriality in caenagnathids. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 146.
Benner, Cullen and Evans, 2016. A new large-bodied caenagnathid
specimen (Theropoda, Oviraptorosauria) from the Hell Creek Formation
(Late Cretaceous) of Montana, with implications for osteohistological
variability in caenagnathids. Journal of Vertebrate Paleontology.
Program and Abstracts, 96.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Cullen, Simon, Benner and Evans, online 2020. Morphology and osteohistology of a large-bodied caenagnathid (Theropoda:
Oviraptorosauria) from the Hell Creek Formation (Montana): Implications
for size-based classifications and growth reconstruction in theropods. Papers in Palaeontology. Early View. DOI: 10.1002/spp2.1302
Yun and Funston, 2021. A caenagnathid oviraptorosaur metatarsal from
the Mesaverde Formation (Campanian) of Wyoming, USA. Vertebrate Anatomy
Morphology Palaeontology. 9, 105-115.
Caenagnathus Sternberg, 1940
?= Macrophalangia Sternberg, 1932
C. collinsi Sternberg, 1940
?= Macrophalangia canadensis Sternberg, 1932
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (CMN 8776) incomplete mandibles (230 mm, symph 61.9 mm) (femur ~377 mm)
Referred-
?(CMN 8538; holotype of Macrophalangia canadensis) distal
tibia, partial astragalus, distal tarsal III, distal tarsal IV, metatarsal I (~47 mm),
phalanx I-1 (58 mm), pedal ungual I (31 mm), metatarsal II (205 mm; lost), phalanx
II-1 (78 mm), phalanx II-2 (63 mm), pedal ungual II (60 mm), partial metatarsal
III (230 mm; lost), phalanx III-1 (75 mm), phalanx III-2 (52 mm), phalanx III-3
(58 mm), pedal ungual III (60 mm), metatarsal IV (212 mm; lost), phalanx IV-1
(59 mm), phalanx IV-2 (33 mm), phalanx IV-3 (31 mm), phalanx IV-4 (35 mm), pedal
ungual IV, metatarsal V (60 mm) (Sternberg, 1932)
(CMN 12372; = CMN 12322 of Longrich et al.) manual ungual II (Longrich et al.,
2013)
(TMP 1979.014.0001) manual ungual I (~90 mm on curve) (femur ~364 mm) (Funston, Persons, Bradley and Currie, 2015)
(TMP 1979.015.0001) manual ungual I (~69 mm) (Currie, 1992)
(TMP 1982.019.0222) manual ungual I (91 mm on curve) (femur ~369 mm) (Funston, Persons, Bradley and Currie, 2015)
?(TMP 1986.036.0323) (~96 kg) femur (370 mm) (Funston, Persons, Bradley and Currie, 2015)
?(TMP 1993.036.0197) (femur ~438 mm) metatarsal II (261 mm) (Funston, Persons, Bradley and Currie, 2015)
?(TMP 1993.036.0198) (femur ~411 mm) metatarsal II (245 mm) (Funston, Persons, Bradley and Currie, 2015)
(TMP 1993.0360.475) manual ungual II (108 mm on curve) (femur ~403 mm) (Bell, Currie and Russell, 2015)
(TMP 2009.003.0029; = TMP 1982.019.0222 in part) manual ungual I (85 mm on curve) (femur ~343 mm) (Bell, Currie and
Russell, 2015)
(UALVP 55725) partial caudal vertebra (39.7 mm) (femur ~393 mm) (Funston, Persons, Bradley and Currie, 2015)
(UALVP 56638) (12 year old adult) pubes (416 mm) (Funston, 2020)
(UALVP 59791) partial ilium (Funston, 2020)
?(UALVP 59921) manual phalanx I-1 (102 mm) (Funston, 2020)
Diagnosis- (after Sternberg, 1940) elongate dentary symphysis (also in Anzu; unknown in other caenagnathids
except Chirostenotes, Elmisaurus and Kuszholia).
(after Currie et al., 1993) dorsal midline ridge on anterior
portion of mandibular symphysis (unknown in other caenagnathids
except Chirostenotes, Elmisaurus and Kuszholia); lingual ridges converge to meet (unknown in other caenagnathids
except Anzu, Chirostenotes, Elmisaurus, Citipes and Kuszholia); lateral
occlusal grooves confluent with anterior occlusal grooves (also in Anzu; unknown in other caenagnathids
except Chirostenotes, Elmisaurus and Kuszholia); lower articular ridge on mandible (also in Anzu and Apatoraptor; unknown in other caenagnathids
except Chirostenotes); medial glenoid longer anteroposteriorly (also in Anzu and Apatoraptor; unknown in other caenagnathids
except Chirostenotes).
(after Funston et al., 2015) slight concavity between proximal articular surface
and flexor tubercle in manual ungual I (also in Anzu; unknown in other caenagnathids
except Hagryphus, Chirostenotes and Elmisaurus); short and deep groove between proximal articular surface
and flexor tubercle in manual ungual II (unknown in other caenagnathids
except Chirostenotes and Elmisaurus); distally abrupt large
posterior protuberance on proximal end of metatarsal II (unknown in other caenagnathids
except Chirostenotes, Elmisaurus and Citipes); facet for metatarsal III on metatarsal II extends only about halfway up element (also in Citipes; unknown in other caenagnathids
except Chirostenotes).
(after Funston and Currie, 2016) dentary-angular bar below external
mandibular fenestra not bowed ventrally (unknown in other caenagnathids
except Anzu, Chirostenotes and Apatoraptor).
(after Funston, 2020) low ilium above acetabulum (unknown in other caenagnathids
except Chirostenotes and Citipes); rounded ventral edge of preacetabular blade (unknown in other caenagnathids
except Anzu and Apatoraptor); inclined ventral edge of pubic peduncle of ilium (also in Chirostenotes; unknown in other caenagnathids
except Anzu? and Citipes);
pubic peduncle of ilium with anteroposterior ridge on ventral surface
for contact with pubis, forming concavo-convex contact (unknown in other caenagnathids
except Chirostenotes); pubes relatively straight in anterior view, producing transversely narrow proximal end (also in Epichirostenotes; unknown in other caenagnathids
except Chirostenotes/Citipes TMP 1994.012.0603);
iliac contact of pubis with anteroposterior concavity for ridge on the
pubic peduncle of the ilium, forming concavo-convex contact (unknown in other caenagnathids
except Chirostenotes/Citipes TMP 1994.012.0603).
(proposed) dentary posterodorsal process extends posteriorly past coronoid process (unknown in other caenagnathids
except Anzu, Chirostenotes, Apatoraptor and Kuszholia); pedal phalanx III-3 longer than III-2 (unknown in other caenagnathids except Elmisaurus and Citipes).
Other diagnoses- Being the first described caenagnathid mandible and without relationships with Oviraptor
being recognized for 36 more years, Sternberg's (1940) initial
diagnosis included mostly characters plesiomorphic for caenagnathoids
(no teeth; symphysis relatively broad; mandibular rami parallel;
external mandibular fenestra large; articular surface large, oval and
convex, with a ridged axis parallel to the direction of the ramus) and
caenagnathids (dentaries fused at symphysis). The final character
(retroarticular, and lateral and medial articular processes short) is
relative to Aves and true of most Mesozoic theropods.
Funston et al. (2015) also listed "the anterior projection of the facet
for metatarsal IV on the proximal end" of metatarsal II as being
different from Chirostenotes, but this is comparable to TMP 1979.020.001.
Comments- Caenagnathus collinsi
was named for a pair of mandibles found in Summer 1936, 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, specifically with Galloanseres. In the same
paper he named a new species of Caenagnathus,
C. sternbergi, known from a posterior mandible. This differs from C.
collinsi in a few characters, suggesting early that two species were present in the
Dinosaur Park Formation. Currie et al. (1993) later described five dentaries
which also differ from C. collinsi. They referred to these as Caenagnathus
cf. sternbergi, as none were directly comparable to the C. sternbergi
holotype. A mandible discovered in 2001 (TMP 2001.012.0012) and
described by Funston and Currie (2014) eventually verified Currie et
al.'s dentaries went with sternbergi articulars.
Caenagnathus' identity was finally solved by Osmolska (1976), who allied
it with oviraptorids. Currie and Russell (1988) first suggested Caenagnathus
and Chirostenotes (then only known from postcrania) were synonymous, which was strengthened by Sues' (1994)
Horseshoe Canyon specimen with Chirostenotes-like postcrania and an edentulous maxilla and shown to be true at the family level by Anzu (Lamanna et al., 2011) which preserved skeletons with Caenagnathus-like mandibles. Longrich
et al. (2013) separated Caenagnathus collinsi from Chirostenotes pergracilis
based on differences between the type mandible of the former and sternbergi,
which they synonymize with pergracilis
(as partial skeleton UALVP 59400 later confirmed). They referred
manual ungual CMN 12372 ("erroneously reported as CMN 12322 by those
authors" as stated by Bell et al., 2015) based on "its large size and
robust construction." Bell et al. (2015) described two additional
manual unguals (TMP 1993.036.0475 and 2009.003.0029), noting Caenagnathus "based on size alone poses a conceivable match for the Dinosaur Park unguals described here." Funston et
al. (2015) agreed Caenagnathus collinsi is distinct from Chirostenotes pergracilis, describing several new specimens as Caenagnathus (TMP 79.14.1, 82.19.222,
86.36.323, 93.36.197, 93.36.198, 93.36.631, 93.75.49 and
UALVP 55725). Funston (2020) later considered referral of femora (especially
TMP 1986.036.0323) to be more ambiguous due to his histological
analyses of Chirostenotes
indicating it may have grown larger than previously expected.
Funston also redescribed the holotype mandible and referred a large
ilium (UALVP 59791) and pubis (UALVP 56638), the former of which is
lower than Chirostenotes or Citipes and has a derived pubic contact not present in Chirostenotes.
The ilium and pubis are described in detail and figured in Funston's
(2019) thesis, where he notes regarding UALVP 56638 "no locality data
accompanies the specimen because it was mislabelled, but the excellent
preservation and associated matrix are consistent with the DPF."
Of the new UALVP specimens, 55725 was found on May 31 2014, 59791 May
27 2018 and 59921 July 8 2019 (UALVP online).
Sternberg (1932) described
Macrophalangia canadensis based on a pes found in 1928, then thought
to be an ornithomimid. Colbert and Russell (1969) noted it may be
synonymous with Chirostenotes, though this was not shown to be likely until Osmolska (1981) described Elmisaurus which preserved a manus and pes of similar morphology. Longrich et
al. (2013) proposed Macrophalangia was synonymous with Caenagnathus
collinsi based on size and robusticity compared to Chirostenotes pergracilis,
which they viewed as a separate taxon. Similar or greater differences
in size and robusticity are known for other coelurosaur species so this
is not very convincing evidence. Funston (2020) referred Macrophalangia to Chirostenotes, but noted it differed from
TMP 1979.020.0001 in having a less constricted metatarsal I (waist 95% of proximal depth vs. 76%), pedal
phalanx I-1 longer than metatarsal I (123% vs. 94%) and a straight metatarsal V
(unillustrated in Macrophalangia but described as a "small round splint"). The former two characters were said to be
shared with Anzu, but while phalanx I-1 length (133%) is even greater than Macrophalangia,
metatarsal I constriction (82%) and metatarsal V curvature are both
intermediate in Anzu. Funston (pers. comm. 2015) noted the metatarsals of Macrophalangia
are lost, having gone on loan and never being returned. It is here provisionally referred to Caenagnathus but as in Longrich et al. is not officially considered a senior synonym pending more information (e.g. histology of CMN 8538).
Currie (1992) reported "an isolated ungual from the Judith River
Formation of Alberta [that] is unusual in that the flexor tubercle is
well developed and extends posteriorly below the articulation with the
penultimate phalanx. The only Cretaceous animal that I am aware of with
this unusual articulation is Segnosaurus (Perle, 1981)." This last reference is actually for Erlikosaurus, whose pedal unguals match this description, while the manual unguals of Segnosaurus
are unfigured and its preserved pedal ungual is only figured in dorsal
view. TMP 1979.015.0001 was figured as "possible segnosaurid
ungual." Cullen et al. (2020) found this specimen "to be most
similar in morphology to caenagnathids", which was predicted by Currie
in the initial 1992 publication, where he wrote "the ungual resembles
the manual ungual of Chirostenotes
if the articulation is ignored, and it is possible that the distinctive
nature of the articulation may have been produced by disease or
injury." Given the presence of three Dinosaur Park caenagnathids,
I refer TMP 1979.015.0001 to Caenagnathus based on its large size.
References-
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 Miscellaneous Collections. 139(11), 1-37.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus.
American Museum Novitiates. 2380, 1-49.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism
to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Osmolska, 1976. New light on skull anatomy and systematic position of Oviraptor.
Nature. 262, 683-684.
Osmolska, 1981. Coossified tarsometatarsi in theropod dinosaurs
and their bearing on the problem of bird origins. Palaeontologia Polonica. 42,
79-95.
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(7), 972-986.
Currie, 1992. Saurischian dinosaurs of the Late Cretaceous of Asia and North
America. In Mateer and Chen (eds.). Aspects of Nonmarine Cretaceous Geology. China Ocean Press. 237-249.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
Sues, 1994. New evidence concerning the phylogenetic position of Chirostenotes
(Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 14(3), 48A.
Lamanna, Sues, Schachner and Lyson, 2011. A new caenagnathid oviraptorosaur
(Theropoda: Maniraptora) from the Upper Cretaceous (Maastrichtian) Hell Creek
Formation of the western United States. Journal of Vertebrate Paleontology.
Program and Abstracts 2011, 140.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian
Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin
of the Peabody Museum of Natural History. 54(1), 23-49.
Funston and Currie, 2014. A previously undescribed caenagnathid mandible from
the late Campanian of Alberta, and insights into the diet of Chirostenotes
pergracilis (Dinosauria: Oviraptorosauria). Canadian Journal of Earth Sciences.
51(2), 156-165.
Bell, Currie and Russell, 2015 (online 2014). Large caenagnathids (Dinosauria, Oviraptorosauria)
from the uppermost Cretaceous of western Canada. Cretaceous Research. 52, 101-107.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied
caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of
Alberta, Canada. Cretaceous Research. 54, 179-187.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Cullen, Larson, Zanno, Currie and Evans, 2020. Theropod biodiversity
patterns in the Dinosaur Park Formation (Late Cretaceous: Campanian) of
Alberta revealed through morphometrics and biostratigraphy. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 115.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian)
of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution.
Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Rhodes, Funston and Currie, 2020. New material reveals the pelvic
morphology of Caenagnathidae (Theropoda, Oviraptorosauria). Cretaceous
Research. 114, 104521.
Epichirostenotes Sullivan, Jasinski and van Tomme, 2011
E. curriei Sullivan, Jasinski and van Tomme, 2011
Early Maastrichtian, Late Cretaceous
Horsethief Member of Horseshoe Canyon Formation, Alberta, Canada
Holotype- (ROM 43250) (adult) maxilla, palatine,
braincase, third cervical vertebra, sixth cervical vertebra (85 mm), seventh
cervical vertebra (81 mm), two cervical ribs, anterior dorsal vertebra (40 mm),
anterior dorsal vertebra, dorsal rib, gastralia fragments, sacrum (270 mm),
proximal caudal vertebra, four distal caudal vertebrae, ilial fragments, pubes
(422 mm), ischium (212 mm), distal tibia
Middle Maastrichtian, Late Cretaceous
Tolman Member of Horseshoe Canyon Formation, Alberta, Canada
Referred- ?(CMN 9570) metatarsal II (258 mm) (Russell, 1984)
Diagnosis- (after Sullivan et al., 2011) ischium tapers gradually after obturator process (also in Anzu and Elmisaurus; unknown in other caenagnathids except Chirostenotes); ventral edge of ischium distal to obturator process shallowly concave (also in Anzu; unknown in other caenagnathids except Chirostenotes and Elmisaurus); triangular obturator process (distal edge <20% of obturator depth) (also in Anzu and Elmisaurus; unknown in other caenagnathids except Chirostenotes); ventrodistally curved pubic peduncle on ischium (also in Anzu; unknown in other caenagnathids except Chirostenotes).
(proposed) presence of an antorbital fossa (unknown in other caenagnathids except Anzu); larger foramen magnum than occipital condyle (unknown in other caenagnathids except Anzu); ventrally directed paroccipital processes (unknown in other caenagnathids except Anzu).
Other diagnoses- Sullivan et
al. (2011) also listed "braincase much deeper than long, with
distinctly verticalized basicranial region" which seems to be shared
with Anzu and unknown in
other caenagnathids. Similarly, "otic region with deep, but
anteroposteriorly narrow, lateral depression" has not been evalauated
in Anzu, and is not preserved in other taxa. "Synsacrum composed of six co-ossified vertebrae and pneumatic
foramina" is also true in Anzu and Chirostenotes in regard to sacral number, and these two taxa, Citipes and Kuszholia in regard to all sacrals being pleurocoelous.
Comments- ROM 43250 was
discovered on June 12 1923 and identified as an ornithomimid. Russell
(1972) listed it as an undetermined ornithomimid, though Sues (1994)
correctly identified it as caenagnathid in an SVP abstract before
describing it in detail (Sues, 1997). Sullivan et al. (2011) proposed
the name Epichirostenotes
curriei
for ROM 43250, based on stratigraphy and several ischial differences
from TMP 1979.020.0001. These ischial characters may prove to be
diagnostic (in particular, the more abrupt tapering is present in two Chirostenotes specimens), but they are comparable to interspecific variation in Tyrannosaurus
rex
specimens (AMNH 5027, CM 9380 and FMNH PR2081) so could prove to be
questionable as more specimens are discovered. It is largely
incomparable to the similarly sized Caenagnathus,
although the pubic shaft is much more slender in side view. This may
be due to the extensive taphonomic crushing described by Sues (1997)
however.
CMN 9570 was discovered in 1926 and "was referred to Macrophalangia in the records of the National Museum of Natural Sciences (Ottawa) by Wann Langston, Jr." before being described as cf. Macrophalangia by Russell (1984). Currie and Russell (1988) illustrated it and referred the element to Chirostenotes sp.. Two caenagnathid taxa have since been identified from the Horseshoe Canyon Formation- Apatoraptor and Epichirostenotes, both based on partial skeletons lacking metatarsi. Funston (2019) tentaively refers it to the latter based on size.
References-
Russell, 1972. Ostrich dinosaurs from the Late Cretaceous of western Canada.
Canadian Journal of Earth Sciences. 9, 375-402.
Russell, 1984. A check list of the families and genera of North American dinosaurs. Syllogeus. 53, 1-35.
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(7), 972-986.
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.
Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid Ojoraptorsaurus
boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria), from the Upper Cretaceous
Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico
Museum of Natural History and Science Bulletin. 53, 418-428.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
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, scapholunare, ulnare, semilunate
carpal, distal carpal III, metacarpal I (66 mm), phalanx I-1 (87 mm), manual
ungual I (94 mm on curve), metacarpal II (122 mm), phalanx II-1 (95 mm), phalanx
II-2 (99 mm), metacarpal III (87 mm), phalanx III-1 (54 mm), phalanx III-2 (45
mm), phalanx III-3 (60 mm), manual ungual III (75 mm on curve), manual claw
III impression, proximal pedal ungual I, distal metatarsal II, distal phalanx
II-1, phalanx II-2, pedal ungual II, incomplete pedal ungual III, distal metatarsal
IV, incomplete pedal ungual IV, fragmentary pedal phalanges
Referred- ?(RAM 12433) distal metatarsal II (Zanno, Loewen, Farke, Kim,
Claessens and McGarrity, 2013)
Diagnosis- (after Zanno and Sampson, 2005) larger than Chirostenotes (also in Caenagnathus and Anzu); robust metacarpal I (transverse shaft width ~20% of length) (unknown in other caenagnathids
except Chirostenotes, Apatoraptor and Elmisaurus); robust manual phalanx I-1 (transverse shaft width ~14% of length) (unknown in other caenagnathids
except Anzu, Caenagnathus, Chirostenotes and Elmisaurus);
interphalangeal joint of manual digit I extending just distal to the
metacarpophalangeal joint of digit II (unknown in other caenagnathids
except Chirostenotes, Apatoraptor and Elmisaurus).
Comments- The holotype was discovered in 2002 (Zanno et al., 2013). Being of similar size to the contemporaneous Caenagnathus collinsi,
it differs in having a much more robust manual phalanx I-1 and a groove
between the articular facet and flexor tubercle of manual ungual I.
Zanno and Sampson (2005) conservatively avoided the Chirostenotes/Caenagnathus versus Elmisaurus controversy and merely referred Hagryphus
to Oviraptorosauria, though believed it to be most similar to those two
genera. Funston and Currie (2016) recovered it more basal in
Caenagnathidae than other Late Cretaceous American taxa, though closer
than Microvenator and Gigantoraptor.
References- Zanno and Sampson, 2003. A new caenagnathid specimen from
the Kaiprowits Formation (Late Campanian) of Utah. Journal of Vertebrate Paleontology.
23(3), 114A.
Zanno and Sampson, 2005. A new oviraptorosaur (Theropoda: Maniraptora) from
the Late Cretaceous (Campanian) of Utah. Journal of Vertebrate Paleontology.
25(4), 897-904.
Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013. Late Cretaceous theropod
dinosaurs of southern Utah. In Titus and Loewen (eds.). At the Top of the Grand
Staircase: The Late Cretaceous of Southern Utah. Indiana University Press. 504-525.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Apatoraptor Funston and Currie,
2016
A. pennatus Funston and Currie, 2016
Early Maastrichtian, Late Cretaceous
Horsethief Member of Horseshoe Canyon Formation, Alberta, Canada
Holotype- (TMP 1993.051.0001) (~80 kg, 2 year old subadult) palatine (~25 mm), incomplete mandible
(~165 mm), hyoid (68.4 mm), (cervical series 680 mm), axial fragment, third
to twelve cervical vertebrae fused to ribs (seventh 53.2, ninth 61.5 mm), nine
dorsal vertebrae, nine dorsal ribs, five uncinate processes, scapulae (one proximal;
215 mm), coracoids, sternal plates (one fragmentary; 112 mm), sternal ribs,
humerus (206 mm), radius, ulna (167 mm), (?)semilunate carpal, metacarpal I
(52 mm), phalanx I-1 (82.4 mm), manual ungual I (50 mm on curve), metacarpal
II (105 mm), phalanx II-1 (74 mm), phalanx II-2 (68.5 mm), manual ungual II
(70 mm on curve), metacarpal III (75 mm), partial phalanx III-1, phalanx III-2
(32 mm), incomplete phalanx III-3, ilial fragment, partial femur (~345 mm),
proximal tibia, proximal fibula
Diagnosis- (after Funston and Currie, 2016) anterior constriction of external
mandibular fenestra by posteroventral dentary process (unknown in other caenagnathids except Anzu, Caenagnathus, Chirostenotes, Elmisaurus and Kuszholia); articular region of mandible with low articular
ridge offset from dorsal margin of articular-surangular-coronoid complex (also in Anzu; unknown in other caenagnathids except Caenagnathus and Chirostenotes); small transverse processes on third cervical (also in Kuszholia?; unknown in other caenagnathids except Epichirostenotes); no postzygodiapophyseal lamina on sixth cervical (unknown in other caenagnathids except Epichirostenotes);
parapophyses of sixth and seventh cervicals extend far anteriorly and
are separated by a deep depression (unknown in other caenagnathids
except Epichirostenotes); deep infraprezygapophyseal fossa in sixth and seventh cervicals? (also in Kuszholia?; unknown in other caenagnathids except Epichirostenotes); infradiapophyseal fossa present in fifth-eighth cervicals (unknown in other caenagnathids except Epichirostenotes);
lamina connecting tuberculum and capitulum with corresponding pneumatic
foramen in mid dorsal ribs (unknown in other caenagnathids except Chirostenotes?, Epichirostenotes and Elmisaurus); less curved posterior coracoid margin (also in Anzu; unknown in other caenagnathids except Chirostenotes); coracoid tubercle placed less than halfway down coracoid (unknown in other caenagnathids except Chirostenotes); metacarpal
I less than half as long as metacarpal II (unknown in other caenagnathids except Hagryphus and Elmisaurus); manual phalanx I-1 longer than II-1 or II-2 (unknown in other caenagnathids except Hagryphus, Anzu, Chirostenotes and Elmisaurus);
manual phalanx II-1 longer than II-2 (unknown in other caenagnathids except Hagryphus, Chirostenotes and Elmisaurus).
Other diagnoses- Differences between palatines in this and Epichirostenotes
(longer anteroventral process, anterodorsal process less upturned)
noted by Funston and Currie (2016) may be due to incompletely preserved
edges in both. They stated "the dentary-surangular suture does not
extend posteriorly to the coronoid process", supposedly unlike other
caenagnathids, but this is true in Chirostenotes and Anzu
as well. Funston and Currie also said that "the laminae connecting the
neural spine to the transverse processes of C6 and C7 are not as well
developed in Apatoraptor as in Epichirostenotes,
especially in their posterior extent", but it is uncertain which
laminae these would be since oviraptorosaurs lack cervical
spinodiapophyseal laminae. Funston and Currie list "ventral flange of
angular underlying posteroventral dentary process", but this seems true
in other caenagnathids (Anzu, Chirostenotes TMP 20010.012.0012) with its seeming absence in the one preserved side of Caenagnathus being due to a dorsally displaced angular. They also list "medial fossa anterior
to articular region of mandible" which is present in Caenagnathus, and unknown in other caenagnathids (unreported but possible to check in Chirostenotes and Anzu). The lack of rib fusion in the third cervical unlike Epichirostenotes (and Anzu) could be ontogenetic based on size.
Comments- Funston (2014) comments on TMP 1993.051.0001, a specimen discovered
in 1993 and first believed to be ornithomimid. It is listed in Funston et al.
(2015) as "unnamed taxon", is said to be from the Horseshoe Canyon
Formation by Eberth et al. (2013) and tentatively assigned to Epichirostenotes
there. Currie et al. (2016) incorrectly state it's from the Dinosaur Park Formation
however, when commenting on its scapulocoracoid and humerus. The specimen was
officially named and described by Funston and Currie (2016), recovered as closer
to Elmisaurus rarus than to Citipes, Caenagnathasiaor
any of the larger caenagnathids. Funston's (2019) description in
his thesis includes CT scans not available in the published
version. Simon and Evans (2021) via histological study found the
specimen is a young subadult and proposed it may be a young Epichirostenotes,
which is based on a larger specimen from the same formation. This
would make the proposed cervical differences either ontogenetic or
individual variation.
References- Eberth, Evans, Brinkman, Therrien, Tanke and Russell, 2013.
Dinosaur biostratigraphy of the Edmonton Group (Upper Cretaceous), Alberta,
Canada: Evidence for climate influence. Canadian Journal of Earth Sciences.
50(7), 701-726.
Funston, 2014. Understanding Alberta’s caenagnathids: Insights on anatomy
and taxonomy from a new, articulated specimen. R.E. Peter Biology Conference.
Abstract 10.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied
caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of
Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osm�lska, 2016 (online 2015). New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica.
61(1), 143-157.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Simon and Evans, 2021. Osteohistology supports immature ontogenetic status of North American oviraptorosaurs Apatoraptor pennatus and Chirostenotes pergracilis. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 236-237.
Chirostenotes Gilmore, 1924
= "Steneodactylus" Gilmore vide Holtz, DML 1998
C. pergracilis Gilmore,
1924
= Caenagnathus sternbergi Cracraft, 1971
= "Steneodactylus pergracilis" Gilmore vide Holtz, DML 1998
= Chirostenotes sternbergi (Cracraft, 1971) Snively, Currie, Brinkman,
Ryan, Braman, Gardner, Lam, Spivak and Neuman, 2001
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (CMN 2367) distal metacarpal I, phalanges I-1 (one incomplete; 63
mm), manual unguals I (44 mm), partial metacarpal II, phalanges
II-1 (one incomplete; 65 mm), phalanges II-2 (one incomplete; 72 mm), manual
ungual II (62 mm), distal phalanges III-3 (44 mm; one lost), manual unguals III (36 mm; one lost)
Referred- (CMN 2690; holotype of Caenagnathus sternbergi) posterior
mandible (Cracraft, 1971)
?(TMP 1979.014.0499) manual ungual II (83 mm) (Currie and Russell, 1988)
(TMP 1979.020.0001) (~67 kg, 2 year old juvenile) (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 (58.1 mm
tall), distal metacarpal I, phalanx I-1 (65.4 mm), manual ungual I (48
mm on curve), phalanx II-1 (71.9 mm), phalanx II-2 (75.8 mm), manual
ungual II (69 mm; 83 mm on curve), phalanx III-1 (30.3 mm; lost),
phalanx III-3 (39.3 mm), ilium (255 mm), ischium (138 mm), femur (304
mm), tibia (367 mm), metatarsal I (42.4 mm), phalanx I-1 (39.8 mm),
distal tarsal III fragments, distal tarsal IV, metatarsal II (181 mm),
metatarsal III (207 mm), phalanx III-1 (54.1 mm), metatarsal IV (186
mm), metatarsal V (33 mm) (Currie and Russell, 1988)
(TMP 1985.043.0070) (1 year old juvenile) anterior dentaries (symph 19
mm) (Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020)
(TMP 1990.056.0006) dentaries (symph 46.5 mm) (Currie, Godfrey and Nessov, 1993)
?(TMP 1991.144.0001) incomplete dentaries (symph 27.2 mm) (Currie, Godfrey and Nessov, 1993)
?(TMP 1992.036.0390) (juvenile?) incomplete dentaries (symph 29.6 mm) (Currie, Godfrey and Nessov, 1993)
(TMP 1992.036.1237) (7 year old subadult) anterior dentaries (symph
38.5 mm) (Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020)
?(TMP 1992.040.0044) (juvenile?) anterior dentaries (symph 32 mm) (Currie, Godfrey and Nessov, 1993)
(TMP 1993.036.0181) distal tarsal III, distal tarsal IV, metatarsal II (221 mm), metatarsal IV (221 mm) (Funston and
Currie, 2014b; described by Funston et al., 2016)
(TMP 1996.012.0142) partial dentaries (symph 25.3 mm) (Funston, 2020)
(TMP 2001.012.0012) incomplete mandibles (188 mm; symph 32.8 mm) (Currie, 2005; described by
Funston and Currie, 2014a)
(TMP 2002.012.0103) incomplete ilium (Funston and Currie, 2021)
(UALVP 59400) (9 year old adult) incomplete mandibles, seventh cervical
vertebra (~49 mm), eighth cervical vertebra (~57 mm), fragmentary ninth
cervical vertebra, cervical vertebra, fifth caudal vertebra, sixth
caudal vertebra, seventh caudal vertebra, eighth caudal vertebra (27.1
mm), ninth caudal vertebra (25.2 mm), tenth caudal vertebra (22.3 mm),
eleventh caudal vertebra (21.4 mm), twelfth caudal vertebra (19.5 mm),
thirteenth caudal vertebra (19.4 mm), fourteenth caudal vertebra (16.1
mm), fifteenth caudal vertebra (20.4 mm), sixteenth caudal vertebra
(20.9 mm), seventeenth caudal vertebra (16.9 mm), nine chevrons,
proximal pubic fragment, fragmentary ischia, femoral and/or tibial
fregments, distal tibia, partial astragalocalcanea, distal tarsal IV,
feathers (Funston and Currie, 2021)
Middle-Late Campanian, Late Cretaceous
Belly River Group, Alberta, Canada
?(TMP 1979.008.0622) (juvenile?) anterior dentaries (symph 21 mm) (Currie, Godfrey and Nessov, 1993)
Late Campanian, Late Cretaceous
Upper Two Medicine Formation, Montana, US
?(MOR 1107) articular (symph ~28.5 mm scaled from TMP 2001.012.0012) (Varricchio, 2001)
Diagnosis- (after Cracraft, 1971) smaller than coeval Caenagnathus collinsi; articular ridge higher (unknown in other caenagnathids except Anzu, Caenagnathus and Apatoraptor); articular medial process not as elongated anteroposteriorly (unknown in other caenagnathids except Anzu, Caenagnathus and Apatoraptor).
(after Currie, 1989) metatarsal IV much shorter than III (<93%) (also in Caenagnathus?; unknown in other caenagnathids except Elmisaurus and Citipes).
(after Varricchio, 2001) pedal phalanx III-1 >30% of metatarsal II in length (also in Caenagnathus?; unknown in other caenagnathids except Elmisaurus and Citipes).
(after Longrich et al., 2013) short dentary symphysis (median length
<100% of transverse width at posterior edge), causing moderately
divergent lateral dentary edges (~30-35 degrees) (also in Citipes, Elmisaurus and Kuszholia; unknown in other caenagnathids except Anzu and Caenagnathus); tip of beak with four lingual ridges (also in Elmisaurus and some Kuszholia; unknown in other caenagnathids except Caenagnathus;
absent in TMP 1990.056.0006); dorsal edge of external mandibular
fenestra everted laterally (unknown in other caenagnathids except Anzu, Caenagnathus and Apatoraptor); manual ungual I slender and weakly arched, flexor tubercles distally positioned (unknown in other caenagnathids except Hagryphus, Anzu, Caenagnathus, Apatoraptor and Elmisaurus).
(after Funston, 2020) occlusal tip of dentary upturned at approximately
45 degrees (taphonomically absent in TMP 1990.056.0006?; also in Elmisaurus and Kuszholia; unknown in other caenagnathids except Caenagnathus);
posterior chevrons anteroposteriorly elongate at proximal end, as long
or longer anteroposteriorly than corresponding caudal vertebrae
(unknown in other caenagnathids except Anzu);
tall ilium (height above acetabulum subequal to length between ventral
ischial peduncle tip and anterior edge of pubic peduncle) (also in Citipes; unknown in other caenagnathids except Caenagnathus); distal tarsals and proximal metatarsals not coossified at maturity (also in Caenagnathus and metatarsals of Anzu; unknown in other caenagnathids except Elmisaurus and Citipes); metatarsal V strongly procurving (unknown in other caenagnathids except Anzu, Ceanagnathus?, Elmisaurus and Citipes).
Other diagnoses- (after Gilmore, 1924) differs from Struthiomimus
in shorter manual digits I and III, shorter metacarpal I, deeply
ginglymoid metacarpals I and II, less medially divergent digit I,
longer phalanx II-1, and more curved unguals. The shorter
metacarpal I, and deeply ginglymoid metacarpals I and II also diagnosed
to compared to Ornithomimus. The supposedly different deeply ginglymoid metacarpal III and short phalanges of digit III in Dromaeosaurus are due to mistaking its metatarsal II and pedal phalanges. It differs from Ornitholestes (including Tanycolagreus manus AMNH 587) in the longer metacarpal and phalanx I-1, and more divergent I-1.
Cracraft (1971) included the following character in his diagnosis compared to Caenagnathus collinsi's
holotype, but this does not seem true of e.g. TMP 2001.012.0012-
"portion of ramus immediately anterior to articular region much less
robust and apparently relatively thinner dorsoventrally."
Almost all of Currie and Russell's (1988) characters are now known to
be typical of caenagnathids, and often caenagnathoids in general- six
sacral vertebrae with pleurocoels; digit III of manus longer than
digit I, but with slender phalanges, having a diameter of less than
half that of phalanges in other digits; well-developed posterodorsal
lip on manual unguals; dolichoiliac, propubic pelvis; preacetabular
portion of iliac blade longer than postacetabular; metatarsal III is
proximally pinched between metatarsals II and IV, but only the proximal
tip is excluded from the anterior surface of the metatarsus. Their
final character, tarsometatarsus may have fused in old individuals, is
currently considered false.
Currie (1989) proposed that in Citipes elegans the "preserved portion of metatarsal III is triangular in section, in contrast with Chirostenotes pergracilis, where it is diamond shaped", but these were being measured at different places along the shaft (see comments under Leptorhynchos). He also proposed a short metatarsal II differed from Elmisaurus rarus and elegans, but elegans specimens TMP 1982.016.0006 and TMP 2000.012.0008 have short metatarsals II as well (89% of mtIII) (as do Kol and Macrophalangia, the latter of which Currie viewed as synonymous with Chirostenotes).
Following the identification of dentaries used here, Longrich et al.'s 'Leptorhynchus elegans' examples are actually Chirostenotes pergracilis.
As such, the anteriorly projecting beak tip in some specimens is
individual or taphonomic variation, as is the varience of posterior
divergence between dentaries (~30-35 degrees), and the symphyseal
length/width ratio (49-87% vs. 58% in Citipes and 128% in Caenagnathus), although the divergence and symphyseal width are still different from Caenagnathus.
Contrary to Longrich et al., the dentary is not excluded from the
dorsal margin of the external mandibular fenestra by the surangular
(TMP 2001.012.0012, 1990.056.0006), instead being similar to Caenagnathus with a small dorsal border. The beak tip can also be as elongate as Caenagnathus
(e.g. UALVP 59400, TMP 1990.056.0006). Finally, the manual
phalanges are not extremely long and slender compared to the one known
specimen of Caenagnathus (I-1 UALVP 59921).
Varricchio (2001) listed chorda tympani foramen/slot absent as diagnostic of Caenagnathus sternbergi compared to C. collinsi,
which Currie et al. (1993) had restoted CMN 8776 with. However,
the feature is just an irregular transverse crack on each side at the
posterior base of the glenoid (photo courtesy of McFeeters) instead of
the anteroposterior groove on the dorsal glenoid surface which Currie
et al. illustrated in Anzu specimen BHM 2033.
Funston (2020) included dentary fusion as a character, but this is true
of all caenagnathids. A deep lateral dentary fossa is only
present in TMP 2001.012.0012. Funston also listed cervical
vertebrae with low neural spines and small epipophyses as diagnostic,
but this is true for all oviraptorosaurs. Distal caudal vertebrae with
anteriorly directed transverse processes are also present in Nomingia and Citipati, with the condition in Anzu (the only other caenagnathid to preserve the area) unreported.
Comments- Chirostenotes pergracilis was named in 1924 based on two manus collected in 1914, and thought by Gilmore to be a a coelurid intermediate between Ornitholestes and ornithomimids.
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 holotype 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 found in 1928, then 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) described Elmisaurus which preserved a manus and pes of similar morphology. An
alternative hypothesis, the synonymy of Chirostenotes with Dromaeosaurus,
was suggested by Ostrom (1969; 1990), but is clearly incorrect given Dromaeosaurus' preserved metacarpal I and the recently described partial skeleton UALVP 59400. Macrophalangia is here provisionally referred to Caenagnathus collinsi, as in Longrich et al. (2013).
Currie and Russell (1988) first suggested Chirostenotes was synonymous with the oviraptorosaurian mandible Caenagnathus, which was strengthened by Sues' (1994)
Horseshoe Canyon specimen with Chirostenotes-like postcrania and an edentulous maxilla and shown to be true at the family level by Anzu (Lamanna et al., 2011) which preserved skeletons with Caenagnathus-like mandibles. As noted above, Longrich
et al. (2013) separate Caenagnathus collinsi from Chirostenotes pergracilis
based on differences between the type mandible of the former and Caenagnathus sternbergi,
which they synonymized with pergracilis (as partial skeleton UALVP 59400 later confirmed).
Cracraft (1971) named Caenagnathus sternbergi, known from a posterior
mandible that differs from C. collinsi in a few 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. Currie (2005) illustrated
a new mandible (TMP 2001.012.0012) which confirms Currie et al. were correct to refer the dentaries
to C. sternbergi. This was described
in detail by Funston and Currie (2014a). Longrich et al. referred TMP 1990.056.0006 to Chirostenotes, but 1979.008.0622, 1991.144.0001 and 1992.036.0390 to their Leptorhynchos elegans. Funston et al. (2019) reported histological work on dentaries which
showed small cf. Leptorhynchos elegans UALVP 55639 was older than larger Chirostenotes TMP
1985.043.0070 and 1992.036.1237. They further referred TMP 1992.040.0044 to Chirostenotes, but Funston (2020) later placed this as cf. elegans
instead. As UALVP 55639 with a 22.1 mm symphyseal length is
completely remodeled and thus quite old, I have tentatively reassigned
all larger dentaries (including TMP 1991.144.0001, 1992.036.0390 and
1992.040.0044) to Chirostenotes
and smaller TMP 1979.008.0622 to Caenagnathidae indet. pending
histological examination of each specimen. This is supported by
Funston et al. reporting that fibrous external bone texture is present
in histologically juvenile TMP 1985.043.0070 and also 1992.036.0390 and
1992.040.0044. Notably, the sternbergi holotype matches pergracilis TMP 2001.012.0012 in size, whereas it would be much smaller if it was a synonym of elegans.
TMP 1979.020.0001 was discovered in 1979 and described by Currie and Russell in 1988
as Chirostenotes pergracilis. They distinguished between two kinds of metatarsus
(robust CMN 8538; gracile ROM 781 and TMP 1979.020.0001), tentatively believing them
to be sexual morphs. Ironically, Currie and Russell had mentioned TMP 1982.039.0004
as an American Elmisaurus specimen although they did not yet refer the elegans
holotype to that genus. Currie (1989) expanded on the separation, referring some specimens
to small and fused Elmisaurus elegans (ROM 781, 37163 and TMP 1982.039.0004) and others to large and unfused
Chirostenotes pergracilis (CMN 8538 and 9570, and TMP 1979.020.0001). Currie and Russell had proposed Caenagnathus sternbergi belonged to the
elegans
morph, but this was eventually disproven by Funston and Currie (2021)
who described UALVP 59400, a partial skeleton combining a sternbergi-type mandible with a histologically adult Chirostenotes-type tarsometatarsus. Currie and Russell had referred TMP 1979.020.0001 to their elegans
morph 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 only be a separate species, which he called Chirostenotes
elegans. Currie and Russell
described several characters consistant with TMP 1979.020.0001 being pergracilis instead of elegans-
unfused tarsometatarsus; metatarsals II and IV much shorter than III
(87 and 90% respectively); proximal end of metatarsal III
diamond-shaped. 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 TMP 1979.020.0001 compared
to CMN 8538. Sullivan et al. (2011) questioned the referral of TMP
1979.020.0001 to Chirostenotes pergracilis,
based on "some minor morphological differences, including a broader curvature
of unguals I and II, a deeper ungual (measured from the dorsal and plantar extremity
of the articular facet as per Senter, 2007), and a shorter phalanx III-3."
Yet such differences are common within theropod species known from multiple
specimens. Thus TMP 1979.020.0001 is assigned to Chirostenotes pergracilis
here, as it has been by other more recent authors. A potential
complication was described by Simon and Evans (2021) who used histology
to determine TMP 1979.020.0001 was a juvenile with two LAGS, despite
being slightly larger than the histologically adult UALVP 59400.
They suggested the former may be a juvenile Caenagnathus instead, but if this is true it could also be the case for the Chirostenotes
holotype which has never been examined histologically, and could thus
raise significant taxonomic issues. Currie and Russell referred
manual ungual II TMP 1979.014.0499 to Chirostenotes, but it has not been described or figured. The size is identical to TMP 1979.020.0001 so it is tentatively retained in Chirostenotes here, although this would also make sense for a young Caenagnathus. Notably, differences between manual ungual II in Caenagnathus and Chirostenotes have since been discovered, so it should be possible to assign TMP 1979.014.0499 eventually.
Funston (2020) redescribed the holotype and TMP 1979.020.0001.
Note that for the latter specimen, Funston lists manual ungual I as III
in Table 2 and recognized distal tarsals Currie and Russel had not
commented on. A phalanx III-3 and ungual III had been lost from
the holotype, and manual phalanx III-1 from TMP 1979.020.0001.
Funston described anterior dentaries TMP 1996.012.0142 as
Caenagnathidae indet., but gievn their large size and distinction from Caenagnathus (short symphysis, upturned dentary tip), they are referred to Chirostenotes here. The partial skeleton UALVP 59400 was found on June 23 2016 and described as Chirostenotes pergracilis
by Funston and Currie (2021). Note their Figure 3 has the
articulated cervical vertebrae identified as 4, 5 and 6 but the text
and Table 1 place them as 7,8 and 9. It is notable for combining
mandibular and pedal material in the Dinosaur Park Formation, showing
the sternbergi mandible type is associated with the tarsal morphology of Macrophalangia.
Varricchio (2001) described MOR 1107, an articular from the Two Medicine Formation, as Caenagnathus sternbergi. The specimen does have sternbergi's articular characters as diagnosed by Cracraft, so is here referred to Chirostenotes pergracilis which it is contemporaneous with. While Citipes does not preserve articular material, comparison with complete Chirostenotes mandible TMP 2001.012.0012 shows MOR 1107 is comparable in size and much larger than Citipes.
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.
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, 165 pp.
Cracraft, 1971. Caenagnathiformes: Cretaceous birds convergent in jaw mechanism
to dicynodont reptiles. Journal of Paleontology. 45(5), 805-809.
Osmolska, 1981. Coossified tarsometatarsi in theropod dinosaurs
and their bearing on the problem of bird origins. Palaeontologia Polonica. 42,
79-95.
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(7), 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda)
from North America. Canadian Journal of Earth Sciences. 26(6), 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. 107-125.
Ostrom, 1990. Dromaeosauridae. In Weishampel, Dodson and Osmolska (eds.). The
Dinosauria. University of California Press. 269-279.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
Sues, 1994. New evidence concerning the phylogenetic position of Chirostenotes
(Dinosauria: Theropoda). Journal of Vertebrate Paleontology. 14(3), 48A.
Sues, 1997. On Chirostenotes, a Late Cretaceous oviraptorosaur (Dinosauria:
Theropoda) from western North America. Journal of Vertebrate Paleontology. 17(4),
698-716.
Holtz, DML 1998. https://web.archive.org/web/20210603185802/http://dml.cmnh.org/1998Aug/msg00743.html
Snively, Currie, Brinkman, Ryan, Braman, Gardner, Lam, Spivak and Neuman, 2001.
Alberta's dinosaurs and other fossil vertebrates: Judith River and Edmonton
groups (Campanian-Maastrichtian). In Hill (ed.). Mesozoic and Cenozoic Paleontology
in the western plains and Rocky Mountains. Guidebook for the Field Trips of
the Society of Vertebrate Paleontology 61st Annual Meeting. Museum of the Rockies
Occasional Paper No. 3. 47-75.
Varricchio, 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from
Montana. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University
Press. 42-57.
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.
Lamanna, Sues, Schachner and Lyson, 2011. A new caenagnathid oviraptorosaur
(Theropoda: Maniraptora) from the Upper Cretaceous (Maastrichtian) Hell Creek
Formation of the western United States. Journal of Vertebrate Paleontology.
Program and Abstracts 2011, 140.
Sullivan, Jasinski and van Tomme, 2011. A new caenagnathid Ojoraptorsaurus
boerei, n. gen., n. sp. (Dinosauria, Oviraptorosauria), from the Upper Cretaceous
Ojo Alamo Formation (Naashoibito Member), San Juan Basin, New Mexico. New Mexico
Museum of Natural History and Science Bulletin. 53, 418-428.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian
Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin
of the Peabody Museum of Natural History. 54(1), 23-49.
Funston and Currie, 2014a. A previously undescribed caenagnathid mandible from
the late Campanian of Alberta, and insights into the diet of Chirostenotes
pergracilis (Dinosauria: Oviraptorosauria). Canadian Journal of Earth Sciences.
51(2), 156-165.
Funston and Currie, 2014b. New Elmisaurus (Dinosauria: Oviraptorosauria)
material from Mongolia and Alberta, Canada, and its bearing on North American
caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts,
2014. 134.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian)
of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution.
Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020 (online
2019). Histology of caenagnathid (Theropoda, Oviraptorosauria)
dentaries and implications for development, ontogenetic edentulism, and
taxonomy. The Anatomical Record. 303(4), 918-934.
Funston and Currie, 2021 (online 2020). New material of Chirostenotes pergracilis
(Theropoda, Oviraptorosauria) from the Campanian Dinosaur Park
Formation of Alberta, Canada. Historical Biology. 33(9), 1671-1685.
Simon and Evans, 2021. Osteohistology supports immature ontogenetic status of North American oviraptorosaurs Apatoraptor pennatus and Chirostenotes pergracilis. The Society of
Vertebrate Paleontology Virtual Meeting Conference Program, 81st Annual
Meeting. 236-237.
Elmisaurinae Osmolska, 1981 vide Currie, 2000
Definition- (Elmisaurus rarus <- Caenagnathus collinsi)
(Hendrickx, Hartman and Mateus, 2015)
References- Osmolska, 1981. Coossified tarsometatarsi in theropod dinosaurs
and their bearing on the problem of bird origins. Palaeontologia Polonica. 42,
79-95.
Currie, 2000. Theropods from the Cretaceous of Mongolia. The Age of Dinosaurs in Russia and Mongolia. 434-455.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian
theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology.
12(1), 1-73.
Elmisaurus Osmolska, 1981
Other diagnoses- Osmolska (1981) listed proximolateral
process on metatarsal IV as diagnostic at genus level, but it is also present in Chirostenotes and Citipes and has not been shown to be absent in any caenagnathid.
Currie (1989) stated metatarsal II is subequal in length to III (>93%) in both Elmisaurus and Citipes (his E. elegans),
but this is not true in some Citipes specimens (II is 89% in TMP 1982.016.0006 and TMP 2000.012.0008) and Avimimus shows this as well (93%). Similarly, IV being close to III in length is true of all caenagnathids where known (Elmisaurus, Citipes, Kol, Avimimus) except Chirostenotes.
Metatarsal II curving anterodistally is present in all
caenagnathids. While Currie stated the proximal end of metatarsal
III is triangular in Elmisaurus, unlike Chirostenotes, the latter
does have a triangular section at some points. The only differences in cross
sectional shape between caenagnathid taxa in the literature is due to proximodistal
position along the metatarsal (see detailed discussion under Leptorhynchos
comments).
Currie et al. (2016) listed the "large, compound proximal protuberance
on the posterior surfaces of metatarsals II-IV" as being diagnostic for elmisaurines including Elmisaurus and Citipes, but
this is present in Caenagnathus (based on mtII) and Chirostenotes as well. They also proposed better development of m. tibialis cranialis tubercles on the dorsal surface
of metatarsals II-IV differed from Chirostenotes, but while these are present in the proximal middle section of II-IV Elmisaurus' and Citipes' holotypes, variation exists in other specimens. Elmisaurus IGM 102/6 has them on III and IV but not II, Elmisaurus IGM 102/7 lacks them on II and IV at least, and Citipes
TMP 2000.012.0008 lacks them on III and IV at least. Confusingly,
Currie et al. also refer to scars on the distal ends of metarsals as
being these insertions (e.g. Figure 9A1), which is the only place
Funston et al. (2016) describe them as being. Such distal scars
must be very faint however, since they were not illustrated in either
holotype and are not obvious in most photos either. Due to this,
they are not used in diagnoses here.
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(6), 1319-1324.
Currie, Funston and Osm�lska, 2016 (online 2015). New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica.
61(1), 143-157.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
E. rarus Osmolska, 1981
= Chirostenotes rarus (Osmolska, 1981) Paul, 1988
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Holotype- (ZPAL MgD-I/172) (1.68 m) tarsometatarsus (163 mm; II 147, III 157, IV 147 mm)
Paratypes- (ZPAL MgD-I/20) proximal tarsometatarsus (mtIII ~173 mm)
(ZPAL MgD-I/98) rib fragments, gastralial fragments, two partial mid sacral
centra (37 mm), proximal scapula, metacarpal I (45 mm), phalanx I-1 (65 mm),
manual ungual I (44 mm), incomplete metacarpal II (~63 mm), phalanx II-1 (66
mm), phalanx II-2 (66 mm), phalanx III-1 (30 mm), phalanx III-2 (30 mm), phalanx
III-3 (43 mm), proximal manual ungual, proximal pubes, partial ischia, partial
femora (~308 mm), partial tibia (~340 mm), phalanx I-1 (26 mm), pedal ungual
I (~23 mm), distal metatarsal II, phalanx II-1 (45 mm), partial phalanx II-2
(34 mm), fragmentary pedal ungual II (~31 mm), distal metatarsal III (~151 mm),
phalanx III-1 (46 mm), phalanx III-2 (32 mm), phalanx III-3 (30 mm), distal
metatarsal IV, fragmentary phalanx IV-1, phalanx IV-2 (22 mm), phalanx IV-3
(19 mm), phalanx IV-4 (18 mm), fragmentary pedal ungual IV (~26 mm), fragments
Referred- (IGM 102/6; field number PJC 2000.1) tarsometatarsus (194 mm;
II 172.4, III 185, IV 175.7 mm), metatarsal V (70.3 mm) (Currie, 2001; described by Currie, Funston and
Osm�lska, 2016)
(IGM 102/7; field number PJC 2000.2) (~23 kg adult) frontal (50.5 mm), ~third
to fifth cervical neural arch, partial posterior cervical vertebra (26.4 mm),
incomplete anterior dorsal vertebra (27.7 mm), three proximal dorsal ribs, dorsal
rib fragments, gastralial fragments, vertebral fragment, distal phalanx II-2,
incomplete manual ungual II (40 mm), phalanx III-1 (28 mm), phalanx III-3 (40
mm), proximal femur (~246 mm), tibiae (320, 327 mm), pedal phalanx I-1 (23.1 mm),
incomplete pedal ungual I (30 mm), distal tarsal III, (tarsometatarsus ~176
mm) metatarsal II (161.8 mm), partial metatarsal III (~172 mm), metatarsal IV
(162.2 mm) (Currie, 2001; described by Currie, Funston and Osm�lska,
2016)
(IGM 102/8; field number PJC 2000.3) metatarsal IV (164 mm) (Currie, 2001; described
by Currie, Funston and Osm�lska, 2016)
(IGM 102/9; field number PJC 2001.8) proximal tarsometatarsus (Currie, Funston and Osm�lska, 2016)
(IGM 102/10; field number PJC 2002.4) vertebra, tibia (Currie, Funston and Osm�lska,
2016)
?(?IGM coll.) femur (Currie, 2001)
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
?(IGM 102/107) anterior dentaries (symph 20.63 mm) (Tsuihiji, Watabe, Tsogtbaatar and Barsbold,
2016)
?(IGM coll.; field number NatGeo.2018.040) partial ?postorbital,
dentary fragment, partial angular, ?anterior cervical vertebral
fragment, partial scapula, coracoid, pubes (one incomplete, one
fragmentary), partial astragalus, proximal metatarsal III, incomplete
metatarsal IV, metatarsal fragments (Funston, 2019)
? Early Maastrichtian, Late Cretaceous
? Nemegt Formation, Mongolia
? ventral skull (www.paleofile.com)
Diagnosis- (after Osmolska, 1981) metacarpal I ~66% [69%] the length
of phalanx I-1 (76% in Hagryphus; 50% in Apatoraptor; but unknown in other caenagnathids); tarsometatarsal fusion (also in Citipes; unknown in other caenagnathids except Anzu, Caenagnathus and Chirostenotes); posterior surface of metatarsus deeply concave (also in Anzu and Citipes; unknown in other caenagnathids except Caenagnathus and Chirostenotes); length/width ratio of
metatarsus 0.18 (also in Citipes; unknown in other caenagnathids except Anzu, Caenagnathus? and Chirostenotes).
(after Currie et al., 2016) posterior medial and lateral
ridges of metatarsal III midshaft close to trochlear ridges (also in Citipes; unknown in other caenagnathids except Anzu and Chirostenotes).
Other diagnoses- Osmolska (1981) listed several supposedly diagnostic
characters- digit III markedly thinner than
digits I and II both of which are equally thick (untrue as digit I is more gracile
than II, and conditions are the same in other caenagnathids); ventroposterior
portions of manual phalanges distinctly thickened at proximal surfaces (also
in other caenagnathids); metatarsal III visible anteriorly for ~90% of its length (also in
the Citipes holotype, but not in Elmisaurus IGM 102/7).
Currie et al. (2016) stated at least manual unguals I and II of Elmisaurus (III is unknown) were more
robust and curved than Chirostenotes, with lower proximodorsal lips,
but they are comparable to ungual I of Caenagnathus, Hagryphus and Apatoraptor (II unknown except for Apatoraptor).
(suggested) ventral edge of ischium convex distal to obturator process.
Comments- Discovered in 1970, listed as "small theropod gen. et sp. nov." by Osmolska (1980) and described in 1981, Elmisaurus rarus
was the first caenagnathid discovered with both manual and pedal remains. This
allowed the synonymy between Chirostenotes and Macrophalangia
to be demonstrated. Elmisaurus has been viewed as a relative of Chirostenotes
(first in Elmisauridae and later in Caenagnathidae) from its discovery until
the 2000s, when this was questioned by Maryanska et al. (2002). They noted it
differs from Chirostenotes in having a vascular foramen between metatarsals
III and IV, an m. tibialis cranialis tubercle on the dorsal surfaces of metatarsals
II-IV, a deeply concave posterior side, and a proximolateral process on metatarsal
IV. Yet these are all apomorphies that tell us nothing about Elmisaurus'
relationships unless we find non-caenagnathid taxa that share them. Maryanska
et al. stated pygostylians have the first two characters, and Avimimus
the last, but Elmisaurus otherwise resembles oviraptorosaurs and is quite
dissimilar to any paravian. The relationship with Avimimus is possible,
as found in my modified version of the Hartman et al. matrix, but that taxon is
a caenagnathid in that tree anyway. It is especially confusing that Maryanska
et al. continue to refer Citipes to Caenagnathidae, as it shares the
characters they describe for Elmisaurus. Citipes is
similar enough to Chirostenotes pergracilis to be synonymized by some
authors (e.g. Currie and Russell, 1988), and Elmisaurus has been synonymized
with Chirostenotes by others (e.g. Paul, 1988). It is completely unwarranted
to widely separate the two genera, and Elmisaurus has more recently been
reestablished as a caenagnathid (e.g. Longrich et al., 2013; Currie et al.,
2016).
Currie (2001, 2002) first mentioned new specimens of Elmisaurus rarus,
most of which were later noted in an abstract (Funston and Currie, 2014). Currie
et al. (2016) described these specimens, as well as additional material of ZPAL
MgD-I/98. They identified the manual ungual of the latter as "almost certainly"
from digit I, which was the possible identification of Osmolska as well despite
her figure 2 placing it on digit II. While Funston and Currie state the frontal
"suggests the presence of a high crest", which is also assumed by
Currie et al., no evidence is presented to support this and the frontal
shows no dorsal projection. Only one specimen mentioned by Currie (2001/2002)
was not described in Currie et al.- a femur referred to cf. Elmisaurus rarus.
Sniovely (2000) uses a metatarsus of "Elmisaurus sp. (TMP:PJC}" in his
thesis, which based on the outline in his Figure 2.11 is the holotype
or a cast of it. Tsuihiji et al. (2016) described a partial
mandible that is here referred to E. rarus based on stratigraphy, but it cannot be distinguished from Chirostenotes based on morphology. Ford (2015 online)
reported a partial skull photographed in "a Japanese guide book".
References- Osmolska, 1980. The Late Cretaceous vertebrate assemblages of the Gobi
Desert, Mongolia. Memoires de la Societe Geologique de France. 139,
145-150.
Osmolska, 1981. Coossified tarsometatarsi in theropod dinosaurs
and their bearing on the problem of bird origins. Palaeontologia Polonica. 42,
79-95.
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(7), 972-986.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster. 464 pp.
Snively, 2000. Functional morphology of the tyrannosaund arctometatarsus. Masters Thesis, University of Calgary. 273 pp.
Currie, 2001. Nomadic Expeditions, Inc. report on fieldwork in Mongolia, September
2000. Alberta Palaeontological Society, Fifth Annual Symposium. 12-16.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. Alberta Palaeontological
Society, Sixth Annual Symposium.
8-12.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Longrich, Barnes, Clark and Millar, 2013. Caenagnathidae from the Upper Campanian
Aguja Formation of West Texas, and a revision of the Caenagnathinae. Bulletin
of the Peabody Museum of Natural History. 54(1), 23-49.
Funston and Currie, 2014. New Elmisaurus (Dinosauria: Oviraptorosauria)
material from Mongolia and Alberta, Canada, and its bearing on North American
caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts,
2014. 134.
Ford, 2015 online. http://www.paleofile.com/Dinosaurs/Theropods/Elmisaurussp.asp
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied
caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of
Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osm�lska, 2016 (online 2015). New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica.
61(1), 143-157.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
Tsuihiji, Watabe, Tsogtbaatar and Barsbold, 2016. Dentaries of a caenagnathid
(Dinosauria: Theropoda) from the Nemegt Formation of the Gobi Desert in Mongolia.
Cretaceous Research. 63, 148-153.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
E? sp. (Funston, Currie and Burns, 2016)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- (RSM P2600.1) ?scapula, proximal tibia, distal tibia, proximal
fibula, astragalocalcaneum, distal metatarsal II, distal metatarsal III (~15.8 mm trans)
Comments- RSM P2600.1 was found in 1989 and described as Leptorhynchos sp. (a genus in which they included what is now Citipes) by Funston et al. (2016). They noted "posterior surface has two cruciate ridges" as in Elmisaurus and Citipes (also now known in Anzu), but claimed it differed from elegans in that the latter has distal metatarsal III deeper than wide. However all Citipes elegans
have a surface wider than deep as in RSM P2600.1. The strongly
flaring distal tibia (176% of shaft width) is most similar to Elmisaurus, while the age matches Elmisaurus and Anzu. It is tentatively referred to Elmisaurus sp. here.
Reference- Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
E? sp. (Varricchio, 2001)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
Material- (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)
Comments- Varricchio (2001) referred a pes (MOR 752) to Elmisaurus elegans (now Citipes). This was based on lacking the long phalanges of Chirostenotes, being more slender than Elmisaurus rarus,
and possessing "a small but distinct anterolateral process just
proximal to the distal articulation" of metatarsal II. The latter two
characters have proven to not separate the taxa once more specimens of
each were recovered (see Other diagnoses under Citipes). However the dorsoventral expansion of distal condyles on pedal phalanges and short phalanx III-2 are similar to Elmisaurus rarus (ZPAL MgD-I/98) instead of Citipes elegans
(TMP 2000.012.0008), and indeed the shapes of phalanges II-1, III-1 and
III-2 are nearly identical. Funston et al. (2016) provisionally retained it in elegans, although they noted that elegans specimen TMP 2000.012.0008 lacks a phalanx III-3 that is longer than III-2 as seen in MOR 752. This is also unlike Elmisaurus, but intriguely present in Macrophalangia (= Caenagnathus?). Considering everything and the close age, MOR
752 is here provisionally referred to Elmisaurus sp..
References-
Varricchio, 2001. Late Cretaceous oviraptorosaur (Theropoda) dinosaurs from
Montana. In Tanke and Carpenter (eds.). Mesozoic Vertebrate Life. Indiana University
Press. 42-57.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
Citipes Funston, 2020
C. elegans (Parks, 1933) Funston, 2020
= Ornithomimus elegans Parks, 1933
= Macrophalangia elegans (Parks, 1933) Koster, Currie, Eberth, Brinkman,
Johnston and Braman, 1987
= Chirostenotes elegans (Parks, 1933) Currie and Russell, 1988
= Elmisaurus elegans (Parks, 1933) Currie,
1989
= "Leptorhynchos" elegans (Parks, 1933) Longrich, Barnes, Clark
and Millar, 2013a
= Leptorhynchos elegans (Parks, 1933) Longrich, Barnes, Clark and Millar,
2013b
Late Campanian, Late Cretaceous
Dinosaur Park Formation, Alberta, Canada
Holotype- (ROM 781) partial distal tarsal III, distal tarsal IV, metatarsal
II (155 mm), partial metatarsal III (161 mm), metatarsal IV (157 mm)
Referred- (ROM 37163) distal metatarsal II (Currie, 1989)
?(TMP 1981.023.0034) partial ilium (Rhodes, Funston and Currie, 2020)
....(TMP 1981.023.0035) partial ilium (Rhodes, Funston and Currie, 2020)
....(TMP 1981.023.0039) last sacral vertebra (Funston, 2020)
(TMP 1982.016.0006) tarsometatarsus (mtII 152.4, mtIII 172.2, mtIV 160.5, mtV 44.3 mm)
(Funston et al., 2016)
(TMP 1982.039.0004) proximal tarsometatarsus (Currie, 1989)
?(TMP 1984.163.0036) distal metatarsal III (Funston et al., 2016)
?(TMP 1986.036.0186) distal metatarsal III (Funston et al., 2016)
(TMP 1988.036.0104) distal metatarsal II (Funston et al., 2016)
?(TMP 1992.036.0674) partial ilium (Funston, 2020)
?(TMP 1993.036.0630) distal metatarsal III (Funston et al., 2016)
?(TMP 1994.012.0880) tibia (279.66 mm) (Funston
et al., 2016)
(TMP 1996.012.0141) incomplete tarsometatarsus (mtII ~130 mm, mtIV ~135 mm) (Currie, 2005; described by Funston
et al., 2016)
(TMP 2000.012.0008) (tarsometatarsus ~185 mm) distal tarsal III fused to metatarsal II (164.8 mm), phalanx II-1, pedal
ungual II, distal tarsal IV fused to metatarsal III (185.3 mm), phalanx III-1 (~44 mm), phalanx
III-2, phalanx III-3, pedal ungual III, phalanx IV-?2 (22.3 mm) (Funston et al., 2016)
?(TMP 2005.049.0190) metatarsal III (Funston
et al., 2016)
(UALVP 55585) (subadult) partial metatarsal III (Funston et al., 2016)
(UALVP 55639) (adult) anterior dentaries (symph 22.1 mm) (Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020)
(UALVP 59606) (6 year old adult) distal tarsal IV, metatarsal IV (146 mm) (Funston, 2020)
Late Campanian, Late Cretaceous
Aguja Formation, Texas, US
?(TMM 43057-36; paratype of Leptorhynchos gaddisi) distal tarsal IV fused to proximal metatarsal IV (~22.6 mm trans) (Longrich, Barnes, Clark and Millar, 2013a)
Diagnosis- (after Funston, 2020) distal tarsals III and IV coossified with each other and proximal metatarsus at maturity (also in Elmisaurus; unknown in other caenagnathids except Anzu, Caenagnathus and Chirostenotes); metatarsal III with prominent cruciate ridges on posterior surface (also in Anzu and Elmisaurus; unknown in other caenagnathids except Chirostenotes).
(proposed) distal tibia flared transversely <150% of shaft width (unknown in other caenagnathids except Anzu, Chirostenotes and Elmisaurus); distal condyles of pedal phalanges (II-1, III-1 and III-2 at least) less dorsoventrally expanded than Elmisaurus (unknown in other caenagnathids except Chirostenotes);
pedal phalanx III-2 elongate (posterior height <50% of length
between condyles and cotyles) (unknown in other caenagnathids except Caenagnathus? and Elmisaurus).
Other diagnoses-
Currie (1989) proposed "close to the distal articular surfaces, small
processes of metatarsals II and IV overlap metatarsal III" is
diagnostic of elegans, but
they are not present in TMP 2000.012.0008. Currie used the
character "posteromedial corner of the tarsometatarsus is more deeply
emarginated than that of Elmisaurus rarus" to distinguish elegans based on the rarus holotype versus TMP 1982.039.0004, but the deep emargination is also seen in Elmisaurus rarus specimen IGM 102/6 and not in elegans specimens ROM 781, TMP 1982.016.0006 or TMP 1996.012.0141. Currie stated metatarsal II was more slender in Citipes (maximum shaft diameter 9% of metatarsal length in holotype) than Elmisaurus (12% in holotype), but Elmisaurus rarus
specimen IGM 102/6 has a ratio of 8%. He also listed
"longitudinal, ridge-like posterolateral margin of metatarsal IV is not
as powerfully developed proximally as that of Elmisaurus rarus", but while this appears true in his drawing of the elegans
type, the ridge seems as well developed in all preserved areas although
far proximally it is restored in plaster (Funston et al., 2016: Fig.
3C3).
Longrich et al. (2013a) proposed several differences in beak shape from Chirostenotes based on the assumption TMP 1992.036.0390 belonged to elegans- tip of beak strongly upturned, with anterior occlusal margin projecting
vertically; anterior margin of symphysis straight; chin squarish in lateral
view. Here 1992.036.0390 is referred to Chirostenotes (see below), so that these are individual variation and/or taphonomic.
Funston et al. (2016) claimed that "on metatarsal III of Elmisaurus rarus,
there is rugosity on the distal base of the lateral cruciate ridge, and
a small longitudinal ridge that bisects the longitudinal sulcus between
the cruciate ridges proximally" and that "both of these features are
absent in Leptorhynchos elegans." Yet these are not visible in Elmisaurus
specimens IGM 102/6 or 102/7 (Fig. 10 of Currie et al., 2016) or
available images of the holotype, so are not used here. Funston
et al. also proposed the posterior extension of the proximal
tarsometatarsus is smaller than Elmisaurus, but this is not true in TMP 2000.012.0008. They also claim in elegans
"the distal third of the shaft of metatarsal II curves medially", but
this is only true in TMP 2000.012.0008 and not the holotype.
Similarly they say "metatarsal IV is straight along its length", but
this is untrue in the later described UALVP 59606. Finally, they
claim the "distal condyles of metatarsal III are deeper
anteroposteriorly than wide mediolaterally" in elegans,
but show the opposite in their Table 1 for TMP 1982.016.0006 and
2000.012.0008, which is the case in the holotype as well (Currie, 1989:
Fig. 2r).
Comments- The holotype was discovered in 1926 and 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 TMP 1982.039.0004), 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) (without justification), who called the species Chirostenotes
elegans. Maryanska et al. (2002) and Osmolska et al. (2004) also assigned
elegans to Chirostenotes instead of Elmisaurus, though
they never state their rationale.
When it came to cranial material, Currie and Russell (1988) first proposed the synonymy of Caenagnathus collinsi
with Chirostenotes pergracilis, and Caenagnathus sternbergi with Citipes (then Chirostenotes) elegans. The new partial skeleton UALVP 59400 instead shows sternbergi mandibles went with pergracilis tarsometatarsi. Longrich et al. (2013a) placed elegans in their new genus Leptorhynchos,
based on the Aguja Formation dentaries L. gaddisi. While the gaddisi and
elegans material is similar in size and tarsometatarsal fusion, their reasons for separating these from
Elmisaurus are flawed (see Leptorhynchos comments), and there was no reason to refer elegans to Leptorhynchos over Elmisaurus
or vice versa. Funston and Currie (2014b) agreed and called it Elmisaurus elegans, but Currie et
al. (2016) and Funston et al. (2020) referred elegans to Leptorhynchos without justification. Funston (2020) solved the issue by giving elegans its own genus Citipes,
which functions regardless of phylogeny. Notably, recently
recovered specimens show proposed differences between tarsometatarsi of
Citipes and Elmisaurus
vary individually within these taxa (see Other diagnoses above).
However, the tibiae and pedal phalanges are different showing along
with stratigraphy that they are not synonymous.
Longrich et al. (2013a) referred dentaries TMP 1979.008.0622, 1991.144.0001 and 1992.036.0390 to their Leptorhynchos elegans,
and Funston et al. (2020) referred the third specimen as well.
This was based on putative differences in beak shape (see Other
diagnoses above) of 1992.036.0390, but it is similar in size to RTMP
1990.056.0006 which these authors considered to be Chirostenotes, while Citipes metatarsi are much smaller than those of Chirostenotes.
Furthermore, the large TMP 2001.012.0012 has an intermediate beak
shape, and 1990.056.0006 is taphonomically deformed to be assymetrical
which may have straightened its beak tip. Adding further
evidence, Funston et al. (2020) examined dentaries histologically and
found small
(symphyseal length 22.1 mm) UALVP 55639 to be a mature adult.
Thus Funston (2020) concluded the TMP dentaries "cannot be confidently
referred to Citipes elegans
without osteohistological analysis showing that they are adults" and
assigned them all to Caenagnathidae indet.. I've provisionally
referred them all to Chirostenotes
here, as all but one are larger than the confirmed mature UALVP 55639,
two have fibrous external bone texture and two have poorly developed m.
genioglossus attachments, both characters Funston et al. observed in
histologically juvenile Chirostenotes TMP 1985.043.0070. This leaves UALVP 55639 the only recognized Citipes cranial element which is unfortunate as it is quite fragmentary. Funston and Currie (2014b) announced
several new specimens in an SVP abstract, later describing them in
Funston et al. (2016). One of these, partial tarsometatarsus TMP
1993.036.0181, was later reidentified by Funston and Currie (2021) as Chirostenotes pergracilis based on its large size and lack of fusion. Funston (2020) refers a few new specimens to Citipes-
ilia and associated sacral TMP 1981.023.0034, 0035 and 0039, ilium TMP
1992.036.0674 and metatarsal IV UALVP 59606, describing them and
figuring the metatarsal. The ilia and sacral are figured in
Funston's (2019) thesis.
Note TMP 1992.036.0674 is incorrectly listed on the TMP online catalogue as Dromaeosauridae.
Here, the Leptorhynchos gaddisi
holotype is considered indeterminate and possibly juvenile (see
entry). However, among the referred material from the same
formation (Longrich et al., 2013a) the distal tarsal IV and metatarsal
IV (TMM 43057-36) are like Citipes and Elmisaurus in fusion and size. It's provisionally referred to the contemporeneous Citipes elegans 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(7), 972-986.
Currie, 1989. The first records of Elmisaurus (Saurischia, Theropoda)
from North America. Canadian Journal of Earth Sciences. 26(6), 1319-1324.
Currie, Godfrey and Nessov, 1993 (published 1994). New caenagnathid (Dinosauria: Theropoda) specimens
from the Upper Cretaceous of North America and Asia. Canadian Journal of Earth
Sciences. 30(10), 2255-2272.
Currie, 2005. Theropods, including birds. In Currie and Koppelhus (eds). Dinosaur
Provincial Park, a spectacular ecosystem revealed, part two, flora and fauna
from the park. Indiana University Press. 367-397.
Funston, Currie and Murray, 2013. Examining the diet of a toothless dinosaur:
Evidence supporting a herbivorous diet in Caenagnathus (Dinosauria: Oviraptorosauria).
Journal of Vertebrate Paleontology. Program and Abstracts 2013, 131.
Longrich, Barnes, Clark and Millar, 2013a. Caenagnathidae from the Upper Campanian
Aguja Formation of west Texas, and a revision of the Caenagnathinae. Bulletin
of the Peabody Museum of Natural History. 54(1), 23-49.
Longrich, Barnes, Clark and Millar, 2013b. Correction to "Caenagnathidae
from the Upper Campanian Aguja Formation of west Texas, and a revision of the
Caenagnathinae". Bulletin of the Peabody Museum of Natural History. 54(2),
263-264.
Funston and Currie, 2014a. A previously undescribed caenagnathid mandible from
the late Campanian of Alberta, and insights into the diet of Chirostenotes
pergracilis (Dinosauria: Oviraptorosauria). Canadian Journal of Earth Sciences.
51(2), 156-165.
Funston and Currie, 2014b. New Elmisaurus (Dinosauria: Oviraptorosauria)
material from Mongolia and Alberta, Canada, and its bearing on North American
caenagnathid taxonomy. Journal of Vertebrate Paleontology, Program and Abstracts,
2014. 134.
Currie, Funston and Osm�lska, 2016 (online 2015). New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica.
61(1), 143-157.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian)
of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution.
Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Funston, Wilkinson, Simon, Leblanc, Wosik and Currie, 2020 (online
2019). Histology of caenagnathid (Theropoda, Oviraptorosauria)
dentaries and implications for development, ontogenetic edentulism, and
taxonomy. The Anatomical Record. 303(4), 918-934.
Rhodes, Funston and Currie, 2020. New material reveals the pelvic
morphology of Caenagnathidae (Theropoda, Oviraptorosauria). Cretaceous
Research. 114, 104521.
Avimimidae Kurzanov, 1981
Definition- (Avimimus portentosus <- Oviraptor philoceratops,
Elmisaurus rarus, Caenagnathus collinsi) (Martyniuk, 2012)
Diagnosis- hyperarctometatarsus (unknown in Shixinggia).
References-
Kurzanov, 1981. On the unusual theropods from Upper Cretaceous of
Mongolia. Fossil Reptiles of Mongolia. Sovmestnaya
Sovetsko-Mongol'skaya Paleontologicheskaya Ekspeditsiya, Trudy. 24,
39-50.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Vernon, New Jersey. Pan Aves. 189 pp.
Shixinggia Lu and Zhang, 2005
= "Shixinggia" Lu, 2004
S. oblita Lu and Zhang, 2005
= "Shixinggia oblita" Lu, 2004
Late 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 collected in 1995 and mentioned as an
oviraptorid in Lu et al.'s (2003) abstract. It 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's
oviraptorosaur matrix, and sister taxon to Heyuannia within Oviraptoridae
in a modified version of the TWiG matrix. Funston et al. (2016) and Hartman et al. (2019) recover it as sister to the Khaan+Heyuannia clade. Adding taxa to the latter analysis moves Shixinggia to Avimimidae.
References- Lu, Zhang and Li, 2003. A new oviraptorid dinosaur from the Late Cretaceous
of Shixing, Nanxiong Basin of Guangdong Province, Southern China. Journal of Vertebrate Paleontology. 23(3),
73A.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
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.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Oviraptoridae Barsbold, 1976a
Definition- (Oviraptor philoceratops <- Caenagnathus collinsi)
(Maryanska, Osmolska and Wolsan, 2002; modified from Sereno, 1998)
Other definitions- (Oviraptor philoceratops <- Chirostenotes
pergracilis) (Sereno, online 2005; modified from Padian, Hutchinson and Holtz, 1999)
= Ingeniidae Barsbold, 1981
= Oviraptoridae sensu
Padian, Hutchinson and Holtz, 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, 1976a. O novum pozdnemelovom semeystve melkikh teropod
(Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3),
685-688.
Barsbold, 1976b. A new Late Cretaceous family of small
theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk
SSSR. 226, 221-223.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological
Expedition Transactions. 15, 28-39.
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.
Sereno, 1998. A rationale for phylogenetic definitions, with application to
the higher-level taxonomy of Dinosauria. Neues Jahrbuch f�r Geologie und
Pal�ontologie Abhandlungen. 210(1), 41-83.
Padian, Hutchinson and Holtz, 1999. Phylogenetic definitions
and nomenclature of the major taxonomic categories of the carnivorous Dinosauria
(Theropoda). Journal of Vertebrate Paleontology. 19(1), 69-80.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php
[version 1.0, 2005 November 7]
Alifanov and Averianov, 2006. On the finding of ornithomimid dinosaurs (Saurischia,
Ornithomimosauria) in the Upper Cretaceous beds of Tajikistan. Paleontological
Journal. 40(1), 103-108.
Smith and Molnar, 2006. Jaw musculature and function in oviraptorosaurs. Journal
of Vertebrate Paleontology. 26(3), 126A.
Shanyangosaurus
Xue, Zhang and Bi, 1996
S. niupanggouensis Xue, Zhang and Bi, 1996
Middle-Late Maastrichtian, Late Cretaceous
Shanyang Formation, Shaanxi, China
Holotype- (NWUV 1111) (~1.7 m) uncinate processes(?), partial synsacrum
(centra 32 mm), proximal scapula, humeri (116 mm), femur (258 mm), tibia (327
mm), metatarsals IV (137 mm; one proximal), partial phalanx, pedal ungual
Comments- Xue et al. (1996) report "ribs with horizontal
hooks", but no ribs are mentioned in the material list, nor are any
shown in the plates. If these are uncinate processes, they would
support a pennaraptoran identity.
While Shanyangosaurus was originally identified only to the
level of Theropoda, it is most similar to oviraptorids (Mortimer, DML
2000; followed by Holtz et al., 2004) in the low acromion, absent
fourth trochanter, cnemial crest shape, unfused metatarsus whose fourth
metatarsal is wider than deep, and an elevated femoral head. Hartman et
al. (2019) were the first authors to include it in a phylogenetic
analysis, finding it emerged in Caudipteridae. Yet as it moves to
Oviraptoridae in one step, the cnemial crest shape was not analyzed,
and the age is more congruent with oviraptorids it is tentatively
placed there on this site. If it is constrained as an oviraptorid in
an updated version of the analysis, it falls outside the "Huanansaurus" plus Oviraptor clade.
References- Xue, Zhang, Bi, Yue and Chen, 1996. The development and environmental
changes of the intermontane basins in the eastern part of Qinling Mountains.
Geological Publishing House, Beijing. ISBN 7-116-02125-6. 179 pp.
Mortimer, DML 2000. https://web.archive.org/web/20210506115234/http://dml.cmnh.org/2000Sep/msg00125.html
Holtz, Molnar and Currie, 2004. Basal Tetanurae. In Weishampel, Dodson and Osmolska
(eds.). The Dinosauria Second Edition. University of California Press. 71-110.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
undescribed possible Oviraptoridae (Kirkland, Hernandez Rivera, Aguillon Mart�nez,
Delgado de Jesus, Gomez Nunez and Vallejo, 2000)
Campanian, Late Cretaceous
Cerro del Pueblo Formation, Mexico
Comments- Kirkland et al. (2000) stated "theropods are represented
by isolated bones (Fig. 13C), claws, and teeth indicating the presence
of tyrannosaurids, dromaeosaurids, ornithomimids, and
oviraptorids." Figure 13C is a hadrosaur femur, so the authors
apparently meant figure 12C consisting of theropod phalanges.
Some of these appear to be elements from the El Pelillal locality (e.g.
IGM-7710 and 7712) described by Rodriguez-De La Rosa and
Cevallos-Ferriz (1998), but Aguillon Martinez (2010) noted this
locality is actually from the younger Ca�on del Tule Formation.
Although Aguillon Martinez stated "small vertebrates from the Cerro del
Pueblo Formation remain undocumented", Rodriguez-De La Rosa and
Cevallos-Ferriz do indicate "Theropoda Indet. family" remains are known
from several areas of the Cerro del Pueblo Formation- Presa San
Antonio, Agua de la Mula, La Rosa and Rincon Colorado area. That
being said, no oviraptorosaur material was noted by Aguillon Martinez
or Rivera-Sylva and Carpenter (2014) and so may have been misidentified.
References-
Rodriguez-De La Rosa and Cevallos-Ferriz, 1998. Vertebrates of the El
Pelillal locality (Campanian, Cerro del Pueblo Formation), southeastern
Coahuila, Mexico. Journal of Vertebrate Paleontology. 18(4), 751-764.
Kirkland, Hernandez-Rivera, Aguillon-Mart�nez, de Jesus,
Gomez-Nunez and Vallejo, 2000. The Late Cretaceous Difunta Group of the Parras
Basin, Coahuila, Mexico and its vertebrate fauna. Universidad Aut�noma
del Estado de Hidalgo, Avances en Investigaci�n. 3, 133-172.
Aguillon Martinez, 2010. Fossil vertebrates from the Cerro
del Pueblo Formation, Coahuila, Mexico, and the distribution of Late Campanian
(Cretaceous) terrestrial vertebrate faunas. MS thesis, Dedman College Southern
Methodist University. 135 pp.
Rivera-Sylva and Carpenter, 2014. Mexican saurischian dinosaurs. In
Rivera-Sylva, Carpenter and Frey (eds.). Dinosaurs and Other Reptiles
from the Mesozoic of Mexico. Indiana University Press. 143-155.
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.
undescribed Oviraptoridae (Fanti et al., 2012)
Late Cretaceous(?)
Mongolia or China(?)
Material- (IGM coll.) specimen including skull (147.3 mm), mandible (120
mm), metacarpal I (24 mm), phalanx I-1 (30.1 mm), manual ungual I (31.7 mm),
phalanx II-2 (17.8 mm), manual ungual II (81.8 mm), femur (238 mm), tibia (275
mm), phalanx III-3 (17 mm), phalanx IV-2 (13.3 mm), phalanx IV-3 (10 mm), phalanx
IV-4 (9.3 mm), pedal ungual IV (30 mm) (Fanti et al., 2012)
(IGM coll.) specimen including skull (162 mm), humerus (113 mm), radius (90
mm), ulna (~98 mm), metacarpal I (22.4 mm), phalanx I-1 (29.2 mm), manual ungual
I (35.2 mm), metacarpal II (40.6 mm), phalanx II-1 (19.6 mm), phalanx II-2 (15
mm), manual ungual II (18 mm), femur (220 mm), tibia (265 mm), phalanx III-3
(15.8 mm), phalanx IV-2 (13.6 mm), phalanx IV-3 (9.9 mm), phalanx IV-4 (8.9
mm), pedal ungual IV (27.3 mm) (Fanti et al., 2012)
skull, mandible, five anterior cervical vertebrae (www.dino-pantheon.com, online 2007)
Comments- Fanti et al. (2012) listed measurements for two "oviraptorid
incertae sedis" specimens at the IGM. Analyzing their proportions may enable
identification. Numerous unpublished oviraptorid specimens are known and have
been photographed online. Most are referred to Oviraptor, Conchoraptor
or "Ingenia", but this is generally based on the pre-1980's
convention of calling all oviraptorids Oviraptor or the pre-2000 convention
of calling all crestless oviraptorids Conchoraptor or "Ingenia".
The more complete skeletons, especially when mounted, often contain faked portions.
References- www.dino-pantheon.com, online 2007. http://www.dino-pantheon.com/museumreport/photo/oviraptorhead.jpg
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia
from the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS
ONE. 7(2), e31330.
undescribed oviraptorid (U.S. Attorney's Office Southern District of New York, online 2014)
Late Cretaceous(?)
Mongolia
Material- (CMMD coll.) skeleton
Comments- U.S. Attorney's
Office Southern District of New York (online 2014) report on a legal
case where smuggled dinosaurs were being returned to the CMMD in
Mongolia, including "two freestanding Oviraptors." Only one has
been photographed online to my knowledge, so the identity and amount of
real fossil in the second mount remains unknown.
Reference- U.S. Attorney's Office Southern District of New York, online 2014. Manhattan U.S. Attorney Announces Return To Mongolia Of Fossils Of Over 18 Dinosaur Skeletons. July 10.
undescribed oviraptorid (Jensen, 2008)
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav, Barun Goyot Formation, Mongolia
Material-
?(CMMD coll.) skull, mandibles, eleven cervical vertebrae, cervical
ribs, thirteen dorsal vertebrae, dorsal ribs, sacrum, caudal series,
chevrons, scapulae, coracoids, furcula, humeri, radii, ulnae,
semilunate carpal, metacarpals I, phalanges I-1, manual unguals I,
metacarpals II, phalanges II-1, phalanges II-2, manual ungual II,
metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3,
manual unguals III, ilium, pubes, ischia, femora, tibiae, fibulae,
astragali, metatarsals II, phalanges II-1, phalanges II-2, pedal
unguals II, metatarsals III, phalanges III-1, phalanges III-2,
phalanges III-3, pedal unguals III, metatarsals IV, phalanges IV-1,
phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal unguals IV
(private coll.; PMO X678 cast; UALVP 49394 cast) skull (112 mm),
mandibles (99.8 mm), hyoids, eight partial cervical vertebrae, dorsal rib fragments
(Knutsen, 2008)
(private coll.; UALVL 49393 cast) skull (~126 mm), sclerotic ring, mandibles (139 mm), hyoids, phalanx
I-1 (57.5 mm), manual ungual I (47 mm), incomplete metacarpal II, phalanx II-1
(56 mm), phalanx II-2 (55 mm), manual ungual II (47 mm), incomplete metacarpal
III, phalanx III-1 (38 mm), phalanx III-2, phalanx III-3, manual ungual III
(Fanti et al., 2012)
Comments- This taxon is known from three specimens with similar high, pointed
crests. Casts are common, with Prehistoric Planet Store (online, 2019) listing one as being from Khermeen
Tsav. This specimen was analyzed by Jensen (2008), who called it Oviraptor
sp..
The second specimen includes a manus preserved next to the skull and is
listed as Oviraptor philoceratops
on the Witmer Lab website (Witmer, 2012 online). As noted by Rey
(DML, 2003), Gee and Rey (2003) based their fictional oviraptorosaur
"Ronaldoraptor" on this skull, but as the taxon is not obviously
nonfictional in the book it is not considered actual nomenclature
here. Fanti et al. (2012) used UALVP casts (Oviraptor in their online catalogue) to include
measurements of both specimens in their dataset as "oviraptorid
incertae sedis".
U.S. Attorney's Office Southern District of New York (online 2014)
report on a legal case where smuggled dinosaurs were being returned to
the CMMD in Mongolia, including "two freestanding Oviraptors."
One of these is photographed by Todd (2017 online) and has a cranial
crest similar in shape to the above specimens. While at least
some of the mounted skeleton must be real for it to be illegal to own
(and indeed the e.g. proximal scapula, some dorsal ribs and ilium seem
to be actual fossils), the texture of the forelimbs, feet otherwise
complete but both lacking digit I and metatarsal V, and low
cervicodorsal ratio suggest at least some is a cast. This
includes the possibility the skull is cast, which would eliminate the
only reason it is listed under this entry here, and it should be noted
the manus strongly differs from the UALVL 49393 specimen as well, with
shorter phalanges and a larger ungual I. The manus may be cast as
well of course. Making things even more ambiguous, the specimen
is listed as being from the Djadokhta Formation on its CMMD plaque, but
the associated point on the map is far to the west in Gurvantes, close
to Nemegt and Baynshiren loacilities but far from Djadokhta localities.
References- Gee and Rey, 2003. A Field Guide to Dinosaurs: The Essential Handbook for Travelers in the Mesozoic. Barron's Educational Series. 144 pp.
Rey, DML 2003. https://web.archive.org/web/20210603185800/http://dml.cmnh.org/2003Mar/msg00210.html
Jensen, 2008. Beak morphology in oviraptorids, based on extant
birds and turtles. Masters Thesis. University of Oslo. 48 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Witmer, 2012 online. https://people.ohio.edu/witmerl/collections/Theropods/oviraptor.htm
U.S. Attorney's Office Southern District of New York, online 2014. Manhattan U.S. Attorney Announces Return To Mongolia Of Fossils Of Over 18 Dinosaur Skeletons. July 10.
Todd, 2017 online. https://www.flickr.com/photos/101561334@N08/35724849645/in/album-72157683092773953/
Prehistoric Planet Store online, 2019. https://web.archive.org/web/20210120030224/http://www.prehistoricstore.com/item.php?item=1839
Banji Xu and Han, 2010
B. long Xu and Han, 2010
Early Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Holotype- (IVPP V16896) (juvenile) skull (65 mm), mandibles (one partial,
one incomplete)
Diagnosis- (after Xu and Han, 2010) premaxillonasal crest with stepped
posterior end; premaxillonasal crest with two longitudinal grooves and numerous
oblique striations on lateral surface; elongate external naris placed posteriorly;
deep fossa on dorsal surface of palatal process of pterygoid; several longitudinal
grooves on the posterodorsal dentary; several tubercles on the dorsal surangular
shelf.
Comments- Banji was discovered by an amateur collector prior to September 2009. Xu and Han (2010) initially recovered Banji
as a basal oviraptorid based on a Maryanska et al. analysis, while a
more modern version (Funston and Currie, 2016) moves it closer to heyuannines than Citipati or Rinchenia, sister to Wulatelong. Hartman et al. (2019) recover it in the more basal position however.
Reference- Xu and Han, 2010. A new oviraptorid dinosaur (Theropoda: Oviraptorosauria)
from the Upper Cretaceous of China. Vertebrata PalAsiatica. 48(1), 11-18.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
"Huanansaurus" Lu, Pu, Kobayashi,
Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015
"H. ganzhouensis" Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee,
Kundrat and Shen, 2015
Early Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Material- (HGM 41HIII-0443) incomplete skull (206.1 mm), mandible (182.8
mm), atlas, axis, third cervical vertebra, fourth cervical vertebra, fifth cervical
vertebra, incomplete sixth cervical vertebra, partial seventh cervical vertebra,
gastralia, incomplete humerus (195 mm), fragmentary radius (190 mm), fragmentary
ulna, fragmentary carpals, fragmentary metacarpal I (40 mm), phalanx I-1 (80
mm), incomplete manual unguals I (70 mm), incomplete metacarpal II (85 mm),
phalanges II-1 (one partial; 55 mm), phalanges II-2 (65 mm), manual unguals
II (65 mm), incomplete metacarpal III (84 mm), phalanx III-1 (32 mm), phalanx
III-2 (30 mm), phalanges III-3 (44 mm), manual unguals III (52 mm), distal femur,
proximal tibia, incomplete metatarsal II, phalanx II-1 (45 mm), phalanx II-2
(35 mm), pedal ungual II (55 mm), incomplete metatarsal III, phalanx III-1 (45
mm), phalanx III-2 (39 mm), phalanx III-3 (35 mm), pedal ungual III (50 mm),
incomplete metatarsal IV, phalanx IV-1 (33 mm), phalanx IV-2 (30 mm), phalanx
IV-3 (21 mm), phalanx IV-4 (20 mm), pedal ungual IV (50 mm)
Diagnosis- (after Lu et al., 2015) posterodorsal process of the premaxillae
contact lacrimals (also in Citipati and Conchoraptor); distinct
opening near posteroventral corner of distal end of posterodorsal premaxillary
process; circular supratemporal fenestra much smaller than laterotemporal fenestra;
nuchal crest pronounced; mandibular condyles of quadrate posterior to occipital
condyle; pneumatized dentaries; anterodorsal tip of dentary projecting anterodorsally
at an angle of 45 degrees or less relative to the ventral margin of symphysis;
length of dentary symphysis between 20% and 25% of mandible length; dentary
portion of dorsal external mandibular fenestra margin strongly concave; posteroventral
dentary process twisted so that lateral surface faces somewhat ventrally; angular
contributes extensively to border of external mandibular fenestra; metacarpal
I long and slender, diameter 20% of length; proximodorsal lip on all manual
unguals prominent.
Comments-
Lu et al. (2015) state "Huanansaurus" "was unearthed from the current
construction site of the Ganzhou Railway Station." It was
described on July 2 2015 as a new taxon
of oviraptorid. However, this paper has no mention of ZooBank
and as of February 6 2020
"Huanansaurus" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Huanansaurus ganzhouensis" Lu
et al., 2015 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
Lu et al. (2014) announced this specimen in an abstract as
a sister taxon to Citipati. It was fully described and named by Lu et
al. (2015), and emerged in a trichotomy with Citipati osmolskae
and IGM 100/42 in their analysis derived from Maryanska et al.'s
oviraptorosaur matrix. Cau et al. (2017) also recovered it sister
to C. osmolskae (but not IGM 100/42) while Hartman et al. (2019) find it more basal.
References- Lu, Kobayashi, Pu, Chang, Zhang, Shang and Liu, 2014. A new
oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous
of southern China and its paleogeographical implications. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 164.
Lu, Pu, Kobayashi, Xu, Chang, Shang, Liu, Lee, Kundrat and Shen, 2015. A new
oviraptorid dinosaur (Dinosauria: Oviraptorosauria) from the Late Cretaceous
of southern China and its paleobiogeographical implications. Scientific Reports.
5, 11490.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399. DOI: 10.1038/nature24679
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
unnamed oviraptorid "NXMV" (Lu, 2004)
Late Maastrichtian, Late Cretaceous
Pingling Formation, Guangdong, China
Material- (E-1) partial skull, mandible (150 mm)
....(K2-1) two posterior cervical vertebrae (30 mm), nine dorsal vertebrae (20
mm), dorsal rib fragment
....(K2-12) incomplete pes
Comments- These specimens probably belong to the same individual, called
NXMV by Lu (2004). Wang et al. (2013) believes it may belong to Ganzhousaurus, but Hartman et al. (2019) recover it as sister to "Corythoraptor" in Oviraptoridae, while Ganzhousaurus
is a non-caenagnathoid. Adding additional taxa leaves it in the
same area of the tree, but in an uncertain position relative to "Corythoraptor", Nankangia, "Lingyuanosaurus" and Yulong.
References-
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Lu, 2005. Oviraptorid Dinosaurs from Southern China. Geological Publishing House,
Beijing. 200 pp.
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda)
from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
"Corythoraptor" Lu, Li, Kundrat, Lee, Sun, Kobayashi, Shen, Teng and Liu, 2017
"C. jacobsi" Lu, Li, Kundrat, Lee, Sun, Kobayashi, Shen, Teng and Liu, 2017
Early Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Material- (JPM-2015-001) (>7 year old adult) incomplete
skull (~174.4 mm), incomplete mandibles (~143.2 mm), hyoids, atlas,
axis (52 mm without odontoid), third cervical vertebra (55.3 mm),
fourth cervical vertebra (67.8 mm), fifth cervical vertebra (66.4 mm),
sixth cervical vertebra (71.7 mm), seventh cervical vertebra (62.6 mm),
eighth cervical vertebra (63.7 mm), ninth cervical vertebra (64.5 mm),
tenth cervical vertebra (65.7 mm), eleventh cervical vertebra (71.4
mm), twelfth cervical vertebra (53.2 mm) (dorsal series ~400 mm) first
through sixth dorsal vertebrae, several dorsal ribs, gastralia, sacrum
(~210 mm), first through fifth caudal vertebrae, dorsal third chevron,
distal scapula, sternal plate, humerus (212.9 mm), radii (205 mm),
ulnae (210 mm), carpals, metacarpals I (39.4 mm), phalanges I-1 (69.5
mm), manual unguals I (~60.5 mm), metacarpals II (96.5 mm), phalanges
II-1 (64.5 mm), phalanges II-2 (63.5 mm), manual ungual II (60.9 mm),
metacarpals III (93.8 mm), phalanges III-1 (36.3 mm), phalanx III-2
(31.1 mm), phalanges III-3 (50.9 mm), manual unguals III (55.6 mm),
partial ilia (300 mm), pubes (360 mm), ischia (220 mm), femora (330.5
mm), tibiae (one incomplete; 392 mm), proximal fibuila, partial
astragalus, distal tarsal III, distal tarsal IV, metatarsal I, phalanx
I-1 (37.2 mm), pedal ungual I (36.9 mm), metatarsals II (one
incomplete; 148.3 mm), phalanges II-1 (49.2 mm), phalanges II-2 (30.4
mm), pedal unguals II (52.1 mm), metatarsals III (157 mm), phalanges
III-1 (48 mm), phalanges III-2 (30.5 mm), phalanges III-3 (30.5 mm),
pedal unguals III (54.2 mm), metatarsals IV (156 mm), phalanges IV-1
(33.2 mm), phalanges IV-2 (24.4 mm), phalanges IV-3 (22.9 mm),
phalanges IV-4 (28.9 mm), pedal unguals IV (49.6 mm), metatarsal V (45
mm)
Diagnosis- (after Lu et al.,
2017) ratio of length of tomial margin of premaxilla to premaxilla
height (ventral to the external naris) 1.0-1.4; inclination of
anteroventral margin of premaxilla relative to horizontally positioned
jugal ventral margin posterodorsal; dorsal process of premaxilla bears
two processes- a short
posterodorsally extending process, forming the anterodorsal margin of
the external naris, and a long process, forming most of the
anterodorsal process of the premaxilla; distinct cassowary-like helmet
on skull; external naris much longer than tall; long axis of external
naris parallel to dorsal margin of
antorbital fenestra; antorbital fossa bordered anteriorly by maxilla;
infratemporal fenestra dorsoventrally elongate, narrow
anteroposteriorly; straight anterodorsal margin of dentary in lateral
view; deep fossa, sometimes with associated pneumatopore, on lateral
surface of dentary; no pleurocoels on second through fourth cervical
vertebrae; length of neck twice as long as dorsal vertebral
column, and slightly longer than forelimb length; less pronounced
deltopectoral crest, forming an arc rather than being quadrangular;
manus / humerus + radius ratio between 0.50 and 0.65; manual ungual III
less curved than other unguals; anterior trochanter completely fused
with greater trochanter; distal ends of metatarsal II straight and
metatarsal IV laterally deflected.
Comments- Discovered in 2013 (Geggel, 2017), this was later described by Lu et al. on July 27 2017 as a new taxon
of oviraptorid. However, this paper has no mention of ZooBank
and as of February 6 2020
"Corythoraptor" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Corythoraptor jacobsi" Lu
et al., 2017 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically..
Lu et al. (2017) recovered this as sister to "Huanansaurus", similar to where Hartman et al. (2019) found it.
References- Geggel, 2017. Newfound dino looks like the creepy love child of a turkey and an ostrich. Live Science. https://www.livescience.com/59958-newfound-dinosaur-has-cassowary-like-crest.html
Lu, Li, Kundrat, Lee, Sun, Kobayashi, Shen, Teng and Liu, 2017.
High diversity of the Ganzhou oviraptorid fauna increased by a new
"cassowary-like" crested species. Scientific Reports. 7: 6393. DOI:
10.1038/s41598-017-05016-6
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Nankangia Lu, Yi, Zhong and
Wei, 2013
N. jiangxiensis Lu, Yi, Zhong and Wei, 2013
Early Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Holotype- (GMNH F10003) anterior mandibles, five mid dorsal vertebrae
(38, 37, 38, 39, 40 mm), partial penultimate sacral vertebra, last sacral vertebra
(43 mm), three proximal caudal vertebrae (38, 44, 42 mm), proximal caudal neural
arch, six mid caudal centra (41, 39, 38, 41, 38, 37 mm), incomplete mid caudal
vertebra, two mid chevrons, scapulocoracoids (scapula 260 mm), furcular fragments,
incomplete humerus (240 mm), ilia (one incomplete; 330 mm), pubes (one incomplete;
380 mm), ischia (205 mm), femora (350, 380 mm), tibia (400 mm), fibular fragment,
astragalus, calcaneum
Diagnosis- (after Lu et al., 2013) rostral end of mandibular symphyseal
region not downturned (also in caenagnathids, Incisivosaurus, Luoyanggia
and Ganzhousaurus); two infradiapophyseal fossae on ventral surface near
base of transverse process of dorsal vertebrae; pneumatic fossae on sacral vertebrae
slit-like; neural spines of proximal caudal vertebrae wider transversely than
anteroposteriorly, forming a large posterior fossa with rugose central area;
large fossa on anterior surface (infraprezygapophyseal fossa) and another (infradiapophyseal
fossa) on ventral surface of base of transverse process of the proximal caudals;
femur longer than ilium (also in Yulong and Khaan); ratio of height
to length of ilium 0.36; femoral neck extending dorsomedially at about an angle
of 90 degrees to the shaft; femur and tibia approximately the same length.
Comments- The holotype was donated to the GMNH by a farmer in 2010. Lu
et al. (2013) added Nankangia to a version of the Maryanska et al. analysis
and found it to be the basalmost oviraptorid except Gigantoraptor, but incorrectly placed Oviraptoridae at a less inclusive node. A more recent version (Funston and Currie, 2016) finds Nankangia as the most basal oviraptorid, having moved Gigantoraptor
to Caenagnathidae. Cau et al. (2017) recovers it in
Caenagnathidae while Hartman et al. (2019) find it to be a basal
oviraptorid.
References- Lu, Yi, Zhong and Wei, 2013. A new oviraptorosaur (Dinosauria:
Oviraptorosauria) from the Late Cretaceous of southern China and its paleoecological
implications. PLoS ONE. 8(11), e80557.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017. Synchrotron scanning reveals
amphibious ecomorphology in a new clade of bird-like dinosaurs. Nature.
552, 395-399. DOI: 10.1038/nature24679
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
"Lingyuanosaurus" Yao, Liao, Sullivan and Xu, 2019
"L. sihedangensis" Yao, Liao, Sullivan and Xu, 2019
Early Albian, Early Cretaceous
Sihedang, Jiufotang Formation, Liaoning, China
Material- (IVPP V23589) (~12 kg, juvenile) cervical centrum
(~30 mm), posterior dorsal neural arch, several dorsal ribs, mid sacral
centrum (~25 mm), proximal caudal vertebra (~15 mm), three partial
vertebrae, proximal humerus, distal humerus, manual ungual II, manual
ungual III, ilium (143 mm), proximal (?)ischium, femur (200 mm),
incomplete tibia, partial astragalus, fragments
Diagnosis- (after Yao et al.,
2019) posterior dorsal vertebrae with prominent paradiapophyseal lamina
separating anterior and posterior infradiapophyseal fossae; ventral
part of hyposphene expanding transversely to form intumescence which
extends as far as postzygapophysis posteriorly and beyond
postzygapophysis laterally; dorsal centroprezygapophyseal lamina
extending anteriorly considerably beyond level of prezygapophysis and
fusing with opposite lamina along midline; ilium with most of dorsal
margin strongly convex but posterior part of dorsal margin distinctly
concave; subtriangular cuppedicus fossa immediately dorsal to pubic
peduncle on medial surface of ilium.
Comments- "Lingyuanosaurus" was
discovered before July 2018 and described on March 22 2019 in
Scientific Reports. However, this paper has no mention of ZooBank
and as of February 5 2020
"Lingyuanosaurus" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Lingyuanosaurus sihedangensis"
Yao et al., 2019 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
Yao et al. (2019) analyzed "Lingyuanosaurus" in a version of Zanno's therizinosaur matrix and resolved it between Beipiaosauirus and Alxasaurus
as a therizinosauroid. However, when Mortimer added the taxon to
Hartman et al.'s (2019) analysis, it emerged as an oviraptorid.
Of the supposedly therizinosaurian characters listed by Yao et al.,
Zanno's "dorsal vertebrae with a complex laminar structure" refers to
mid-dorsals ("anterior dorsals" in Zanno's nomenclature) which are not
preserved in "Lingyuanosaurus"
and nor was that character (their 274) scored for the taxon.
Oviraptorosaurs can have both "laterally flattened manual unguals with
dorsally positioned collateral grooves" (e.g. Chirostenotes)
and "a highly modified ilium with a deep preacetabular process, a
reduced postacetabular process, a preacetabular process whose ventral
margin is dorsally displaced relative to the acetabulum" (e.g. Rinchenia).
The slender pubic peduncle is therizinosaur-like, however. It
only takes one more step to move the taxon to Therizinosauria, and four
more to move to Ornithomimosauria. Notably, Zanno's
oviraptorosaur sample was limited due to Balanoff working on the group
at the time.
References-
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Mortimer, 2019. https://theropoddatabase.blogspot.com/2019/03/what-is-lingyuanosaurus.html
Yao, Liao, Sullivan and Xu, 2019. A new transitional therizinosaurian
theropod from the Early Cretaceous Jehol Biota of China. Scientific
Reports. 9:5026. DOI: 10.1038/s41598-019-41560-z
Yulong Lu, Currie, Xu, Zhang, Pu and
Jia, 2013
Y. mini Lu, Currie, Xu, Zhang, Pu and Jia, 2013
Late Cretaceous
Qiupa Formation, Henan, China
Holotype- (HGM 41HIII-0107) (<1 year old young juvenile) skull (48.16
mm), mandibles (51.97 mm), atlas, axis, five cervical vertebrae, posterior dorsal
vertebrae, dorsal ribs, about twenty proximal and mid caudal vertebrae, scapula
(42 mm), clavicles, humeri (44.6 mm), radii (44.5 mm), ulnae (44.7 mm), distal
phalanx I-1, manual ungual I, metacarpal II (22.8 mm), phalanx II-1, phalanx
II-2, manual ungual II, metacarpal III, phalanx III-2, phalanx III-3, manual
ungual III, incomplete ilium (55 mm), partial femur (72 mm), tibiae (88.8 mm),
incomplete fibulae, astragali, metatarsal I, partial phalanx I-1, metatarsals
II, metatarsals III (43.3 mm), metatarsals IV, pedal phalanges including III-3
(11.7 mm), pedal unguals
Paratypes- (HGM 41HIII-0301) (embryo) skeleton including radius (22.6
mm), ulna (21 mm), ilium (39.3 mm), femur (46 mm), 26 eggs (19 not purchased),
nest
(HGM 41HIII-0108) (young juvenile) posterior skull, posterior mandibles
(HGM 41HIII-0109) (young juvenile) skull, mandibles (48.54 mm), partial postcranial
skeleton including scapula (50.26 mm), humerus (35.9 mm)
(HGM 41HIII-0110) (young juvenile) incomplete skull, incomplete mandibles, few
cervical centra
(HGM 41HIII-0111) (young juvenile) ilium
Diagnosis- (after Lu et al., 2013) posterodorsal corner of antorbital
fenestra and anteroventral corner of external naris at the same level (also
in Citipati); distinct opening in premaxilla anteroventral to the external
naris; antorbital fossa partly bordered by premaxilla anterodorsally; subnarial
process of premaxilla does not contact anterior process of lacrimal; parietal
almost as long as frontal; in dorsal view, posterior margin forms a straight
line between postzygapophyses in fourth and fifth cervical vertebrae; femur
longer than ilium.
Comments- Kobayashi et al. (2008) first mentioned Qiupa oviraptorids, and the Yulong material was found before 9-20-2012.
While the holotype and HGM 41HIII-0108 were found associated, the others were
found in separate quarries up to 4 km apart. The diagnostic characters may merely
be characteristic of juvenile oviraptorids, and Yulong's recovered poisition
in Lu et al.'s (2013) Maryanksa et al.-based analysis (between Gigantoraptor
and more derived oviraptorids) may be basal due to ontogenetically variable
characters, as juvenile theropods commonly end up more basally than adults.
References- Kobayashi, Lu, Lee, Xu and Zhang, 2008. A new basal ornithomimid
(Dinosauria: Theropoda) from the Late Cretaceous in Henan province of China.
Journal of Vertebrate Paleontology. 28(3), 101A.
Lu, Currie, Xu, Zhang, Pu and Jia, 2013. Chicken-sized oviraptorid dinosaurs
from central China and their ontogenetic implications. Naturwissenschaften.
100(2), 165-175.
Oviraptorinae Barsbold, 1976a 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) retained 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. It's apparent the original
concept of Oviraptorinae is probably not monophyletic with respect to "Ingeniinae",
and using Osmolska et al.'s definition leads to situations where useful similarity
to their or Barsbold's concept of the taxon is lost. A more useful definition
would be (Oviraptor philoceratops <- Heyuannia yanshini), or with the recent replacement of "Ingeniinae" with Heyuanninae (Oviraptor philoceratops <- Heyuannia huangi).
References- Barsbold, 1976a. O novum pozdnemelovom semeystve melkikh teropod
(Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3),
685-688.
Barsbold, 1976b. A new Late Cretaceous family of small
theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk
SSSR. 226, 221-223.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological
Expedition Transactions. 15, 28-39.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. In Weishampel, Dodson
and Osmolska (eds.). The Dinosauria. University of California Press. 249-258.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Osm�lska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel,
Dodson and Osm�lska, (eds.). The Dinosauria, Second Edition. University
of California Press. 165-183.
Citipatiinae Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
Comments- Funston et al. (2020) used a version of Maryanska et al.'s oviraptorosaur analysis to recover "Nankangia jiangxiensis, Oviraptor philoceratops, and Yulong mini
as successive outgroups to two subfamilies of oviraptorids." "One
of the two groups is comprised of forms with elongate manus with digits
roughly equal in robustness and an elongate third digit" which "is
hereby designated Citipatiinae because Citipati
is the oldest valid genus named in the family [sic]." While not
explicitly listed as fam. nov. as suggested by ICZN Recommendation 16A,
the authors do later refer to "this new citipatiine clade", which is
considered to cover ICZN Article 16.1 ("Every new name published after
1999, including new replacement names (nomina nova), must be explicitly
indicated as intentionally new"). While no phylogenetic
definition is given, the wording and symbols in their Figure 6 suggests
"Citipati osmolskae <- Heyuannia huangi" was the intention.
Reference- Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new
two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.
Royal Society Open Science. 7: 201184.
"Anomalipes" Yu, Wang, Chen, Sullivan, Wang, Wang and Xu, 2018
"A. zhaoi" Yu, Wang, Chen, Sullivan, Wang, Wang and Xu, 2018
Campanian, Late Cretaceous
Xingezhuang or Hongtuya Formation, Wangshi Group, Shandong, China
Material- (ZCDM V0020)
incomplete femur (~300 mm), incomplete tibia (~360 mm), partial fibula,
pedal ungual II (51.4 mm), metatarsal III (167 mm), phalanx IV-1 (37.4
mm)
Diagnosis- (after Yu et al.,
2018) femoral head anteroposteriorly narrow and somewhat deflected
posteriorly; accessory trochanter low; lateral ridge present on femur;
weak fourth trochanter present; metatarsal III with triangular proximal
articular surface; prominent anterior flange near proximal end of
metatarsal III; metatarsal III medial condyle much narrower than
lateral condyle; longitudinal groove on distal articular surface of
metatarsal III; pedal ungual II with lateral collateral groove deeper
and more dorsally located than medial groove.
Comments- This was described by Yu et al. on March 22 2018 as a new taxon
of caenagnathid. However, this paper has no mention of ZooBank
and as of February 6 2020
"Anomalipes" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Anomalipes zhaoi" Yu
et al., 2018 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
Yu et al. (2018) used a version of Maryanska et al.'s oviraptorosaur analyusis to place this sister to Gigantoraptor in basal Caenagnathidae. In Hartman et al. (2019) it is an oviraptorid sister to Citipati.
References- Yu, Wang, Chen,
Sullivan, Wang, Wang and Xu, 2018. A new caenagnathid dinosaur from the
Upper Cretaceous Wangshi Group of Shandong, China, with comments on
size variation among oviraptorosaurs. Scientific Reports. 8:5030. DOI:
10.1038/s41598-018-23252-2
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
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
Ukhaa Tolgod, Djadochta Formation, Mongolia
Holotype- (IGM 100/978) (~105 kg) complete skeleton including skull (160
mm), sclerotic ossicles, mandibles (151 mm), hyoids, twelve cervical vertebrae,
cervical ribs, seven dorsal vertebrae, two uncinate processes, thirty caudal vertebrae (first caudal
38.2 mm), pygostyle, chevrons, scapulocoracoid, furcula, sternal plates, humerus
(230 mm), radius (189 mm), ulna (200 mm), femur (345 mm), tibia (397 mm), astragalus,
metatarsal I (37 mm), tarsometatarsus (II 176 mm; III 192 mm; IV 188 mm)
Paratypes- (IGM 100/971) (embryo) partial skull (~40 mm), mandibles,
cervical vertebra, three fragmentary dorsal vertebrae, several dorsal rib fragments,
three sacral centra, few distal caudal vertebrae, scapulae, coracoids, furcula,
humeri (20 mm), proximal radius, proximal ulna, partial ilia, femur (36 mm),
tibia (50 mm), fibula, astragali, partial metatarsal II, partial metatarsal
III, partial metatarsal IV, pedal phalanges, nest, eggshell fragments (Norell
et al., 1994)
?(IGM 100/979; Big Mama) distal cervical rib, anterior dorsal vertebra, distal
dorsal ribs 1-6, four uncinate processes, gastralia, incomplete furcula, sternal
plates, four sternal ribs, partial scapulae, coracoids, humeri (215 mm; one
proximal), radii (198 mm; one distal), ulnae (214 mm; one distal), proximal
carpal, semilunate carpal, metacarpal I (~45 mm), phalanx I-1 (88.4 mm), manual
ungual I (91.1 mm), metacarpal II (89.5 mm), phalanx II-1 (65.5 mm), phalanx
II-2 (61.5 mm), manual ungual II (76 mm), metacarpal III (89.4 mm), phalanx
III-1 (33.8 mm), phalanx III-2 (~31 mm), phalanx III-3 (~45 mm), manual ungual
III, distal pubes, distal ischia, distal femur, incomplete tibia, fibulae (~415
mm), astragalus, calcaneum, metatarsal I, phalanx I-1, pedal ungual I, metatarsal
II (~173 mm), phalanx II-1, phalanx II-2, pedal ungual II, metatarsal III (~190
mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal
IV (~177 mm), phalanx IV-1, phalanx IV-2 (24.9 mm), phalanx IV-3 (20.1 mm),
phalanx IV-4 (19 mm), pedal ungual IV (48.4 mm), nest, fifteen eggs (Clark et
al., 1999)
Referred-
?(IGM 100/1004; Big Auntie) (13 year old adult) third to thirteenth
cervical vertebrae fused with cervical ribs, ten dorsal vertebrae, ten
dorsal ribs, four uncinate processes, at least four sacral vertebrae,
incomplete scapulacoracoids (scapula 292.8 mm), furcula, sternal
fragment, three sternal ribs, humeri (230.1 mm), radii (212.4 mm),
ulnae (211.5 mm), scapholunare, semilunate carpal, phalanx I-1 (92.9 mm),
manual ungual I (84.1 mm), metacarpal II (103.5 mm), phalanx II-1 (68.3
mm), phalanx II-2 (75.6 mm), manual ungual II, metacarpal III, phalanx
III-1 (44.5 mm), phalanx III-2 (48.2 mm), phalanx III-3 (53.7 mm),
manual ungual III (57.8 mm), partial ilium (~253 mm), proximal pubis,
partial ischia, femora (one partial; ~402.4 mm), tibiae (one
incomplete), fibulae (one incomplete), astragali, metatarsal I (42.4
mm), phalanx I-1 (31.9 mm), pedal ungual I (35.7 mm), distal metatarsal
II, phalanx II-1 (53.2 mm), phalanx II-2 (37 mm), distal metatarsal
III, distal metatarsal IV, phalanx IV-1 (38.6 mm), phalanx IV-2 (35.7
mm), phalanx IV-3 (31.6 mm), phalanx IV-4 (29 mm), pedal ungual IV
(49.7 mm), twelve eggs (~181x~66.8 mm) (Webster, 1996)
(IGM 100/1009) (juvenile) skull (Norton, DML 2000)
? Late Campanian, Late Cretaceous
? Ukhaa Tolgod, Djadochta Formation, Mongolia
?(IGM 100/1125) two eggs (Grellet-Tinner, 2005)
Diagnosis- (after Clark et al., 2002) anterodorsally sloping occiput
and quadrate; parietal much longer along the midline than the frontal and reaching
nearly to the level of the anterior end of the orbit; ascending process of jugal
perpendicular to the horizontal ramus (rather than extending posterodorsally);
external naris nearly circular; nasal process of premaxilla vertical rather
than sloping posterodorsally; cervical vertebrae elongate (approximately twice
as long as they are wide).
Comments- The first published specimen of this species was an embryo
discovered in 1993 in a nest of eggs (Norell et al., 1994), later described
in detail by Norell et al. (2001). Originally assigned to Oviraptoridae indet.,
it was tentatively referred to the then unnamed Citipati osmolskae by
Norell et al. (2001) and formally referred to that taxon by Clark et al. (2001).
Two partial juvenile paravian skulls were found with the specimen, originally
identified as Velociraptor (Norell et al., 1994) and later troodontids
(Norell and Makovicky, 1999).
Norell et al. (1995) announced a brooding oviraptorid on a nest found in 1993,
which was initially referred to Oviraptor based on manual resemblences.
It was mentioned in the popular press as "Big Mama". The specimen
was later described in detail (Clark et al., 1999) and said to be an oviraptorid
most closely related to Oviraptor, before being officially referred to
Citipati osmolskae by Clark et al. (2001).
A second brooding Citipati specimen found in 1995 is known as "Big
Auntie" (Clark pers. comm. to Auditore). It was mentioned by Webster (1996)
and Clark et al. (1999). A photograph of this specimen is present in Codd (2004),
incorrectly identified as IGM 100/42. Similarly, it is photographed and incorrectly
identified as Oviraptor philoceratops
holotype AMNH 6517 by Codd et al. (2007). Erickson et al. (2009)
examined it histologically, Grellet-Tinner (2005) described the eggs in
detail in his thesis, and the specimen was fully described by Norell et
al. (2018). The eggs are most similar to Elongatoolithus frustrabilis.
The holotype of the genus, a virtually complete specimen found in 1994, was initially identified
as Oviraptor philoceratops
(Webster, 1996) until it was preliminarily described and officially
named by Clark et al. (2001). The skull and mandibles were later
described in detail by Clark et al. (2002), though the postcrania
remains largely undescribed and unillustrated, with only Nesbitt et al.
(2009) and Balanoff and Norell (2012) showing the furcula and
describing some details. More recently, Persons et al. (2014) describe
the tail and pygostyle, while Norell et al. (2018) illustrate the third
to fifth cervical vertebrae and two uncinate processes. Balanoff
et al. (2018) described the endocranium based on Balanoff's (2011)
thesis.
Norton (DML, 2000) noted IGM 100/1009 was discovered in 1993
at Ukhaa Tolgod and was on display at the AMNH Fighting Dinosaurs exhibit. Bhullar et al. (2012) listed it as a Citipati embryo.
References- Gibbons, 1994. Dino embryo recasts parents' image. Science. 266, 731.
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.
Monastersky, 1996. Nesting dinosaur discovered in Mongolia. Science News. 149, 7.
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. Journal of Vertebrate Paleontology. 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 online. Citipati osmolskae, Digital Morphology. http://digimorph.org/specimens/Citipati_osmolskae/
Clark, Norell and Rowe, 2002. Cranial anatomy of Citipati osmolskae (Theropoda,
Oviraptorosauria), and a reinterpretation of the holotype of Oviraptor philoceratops.
American Museum Novitates. 3364, 1-24.
Codd, 2004. The uncinate processes in birds and their implications for the breathing
mechanics of maniraptoran dinosaurs. Dissertation zur Erlangung des Doktotgrades
der Mathematisch-Naturwissenschaftlichen Fakultat der Rheinischen Friedrich-Wilhelms-Universitat
Bonn. 108 pp.
Grellet-Tinner, 2005. A phylogenetic analysis of oological characters: A case
study of saurischian dinosaur relationships and avian evolution. PhD thesis,
University of Southern California. 221 pp.
Grellet-Tinner, Chiappe, Norell and Bottjer, 2006. Dinosaur eggs and nesting
behaviors: A paleobiological investigation. Palaegeography, Palaeoclimatology,
Palaeoecology. 232, 294-321.
Codd, Manning, Norell and Perry, 2007. Avian-like breathing mechanics in maniraptoran
dinosaurs. Proceedings of the Royal Society B. 275(1631), 157-161.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian
physiology inherited by birds? Reconciling slow growth in Archaeopteryx.
PLoS ONE. 4(10), e7390.
Nesbitt, Turner, Spaulding, Conrad and Norell, 2009. The theropod furcula. Journal
of Morphology. 270, 856-879.
Norton, DML 2000. https://web.archive.org/web/20210603191138/http://dml.cmnh.org/2000Jun/msg00082.html
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution.
PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria:
Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Bhullar, Marugan-Lobon, Racimo, Bever, Rowe, Norell and Abzhanov, 2012.
Birds have paedomorphic dinosaur skulls. Nature. 487, 223-226.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Wang, Sun, Sullivan and Xu, 2013. A new oviraptorid (Dinosauria: Theropoda)
from the Upper Cretaceous of southern China. Zootaxa. 3640(2), 242-257.
Persons, Currie and Norell, 2014 (online 2013). Oviraptorosaur tail forms and functions. Acta
Palaeontologica Polonica. 59(3), 553-567.
Pittman and Mallison, 2014. Tail function in oviraptorosaur dinosaurs: Insights
from a 3D tail model of Citipati osmolskae (Theropoda: Oviraptorosauria).
Journal of Vertebrate Paleontology. Program and Abstracts 2014, 205-106.
Moyer, Zheng, Norell and Schweitzer, 2015. Microscopic and immunohistochemical
analyses of the claw of the nesting dinosaur, Citipati osmolskae. Journal
of Vertebrate Paleontology. Program and Abstracts 2015, 186.
Balanoff, Norell, Hogan and Bever, 2018. The endocranial cavity of
oviraptorosaur dinosaurs and the increasingly complex, deep history of
the avian brain. Brain, Behavior and Evolution. 91, 125-135.
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati osmolskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.
C? sp. nov. (Barsbold, 1981)
Late Campanian, Late Cretaceous
Dzamin Khond, Djadochta Formation, Mongolia
Material- (IGM 100/42; Zamyn Khondt oviraptorid) incomplete skull (180mm),
mandibles (153, ~156 mm), fiftteen cervical vertebrae, cervical ribs, seven
dorsal vertebrae, three dorsal centra, eighteen dorsal ribs, sacrum, thirty
caudal vertebrae (first caudal 32 mm), twenty-three chevrons, scapulacoracoids
(scap ~238 mm), furcula, sternal plates (~79, ~84 mm), humeri (205 mm), radius
(180 mm), ulna (188 mm), semilunate carpal, metacarpal I (47 mm), phalanx I-1
(95 mm), manual ungual I (73 mm), metacarpal II (104 mm), phalanx II-1 (58 mm),
phalanx II-2 (73 mm), manual ungual II (64 mm), metacarpal III (98 mm), phalanx
III-1 (42 mm), phalanx III-2 (40 mm), phalanx III-3 (56 mm), manual ungual III
(54 mm), ilia (290 mm), pubes (320 mm), ischia, femora (305 mm), tibiae (380
mm), fibula, astragalus, calcaneum, metatarsal I, metatarsal II, phalanx II-1,
phalanx II-2, pedal ungual II, metatarsal III (168 mm), phalanx III-1 (39 mm),
phalanx III-2, phalanx III-3 (31 mm), pedal ungual III, metatarsal IV, phalanx
IV-1, phalanx IV-2 (24 mm), phalanx IV-3 (30.5 mm), phalanx IV-4 (26 mm), pedal
ungual IV (45 mm), metatarsal V
Diagnosis- (after Clark et al., 2001) anterior edge of the premaxilla
is concave in lateral view; vomers unfused; cervical vertebrae are not elongate.
Comments- This specimen was originally identified as Oviraptor philoceratops
(Barsbold, 1981), an identification which persisted unquestioned through the
80's and 90's (e.g. Barsbold, 1983, 1986; Barsbold et al., 1990; Maryanska et
al., 2002). The completeness relative to the O. philoceratops holotype
resulted in IGM 100/42 becoming the standard example of the species. In 2001,
Clark et al. realized the specimen was more similar to the recently named Citipati
osmolskae, and may represent another species of that genus. Clark et al.
(2002) reiterated this, while Osmolska et al. (2004) found it to be the sister
taxon of Citipati osmolskae in their cladistic analysis. However, Lu
(2004) found that IGM 100/42 was the sister taxon to Oviraptor or Conchoraptor
(depending on taxa included) using Maryanska et al.'s (2002) characters. This
and the resemblence of the more recently described Nemegtomaia (which
was more closely related to Citipati than to IGM 100/42 or Oviraptor
in Lu's analysis) suggests we use caution when assigning IGM 100/42 to a particular
genus.
The specimen has never been described in detail, though many elements have been
illustrated in varied sources. Barsbold (1981) illustrated the skull, mandible,
furcula, sternal plates and manual phalanges. He later (1983) illustrated the
skull in ventral view, mandible in medial view and scapulocoracoid. Barsbold
et al. (1990) illustrated the skull in dorsal view, humerus, complete manus,
and metatarsus. Jensen (2008) analyzed the skull for his thesis and presented
detailed photographs. The almost complete skeleton is mounted, but publically
available photos are too small to be useful. The data matrices of Norell et
al. (2001), Maryanska et al. (2002) and Osmolska et al. (2004) are the most
useful published sources of information besides the figures. Fanti et al. (2012)
and Currie et al. (2016) provide measurements.
References- Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian
Paleontological Expedition Transactions. 15, 28-39.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy,
Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120.
Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva
(ed.). Herpetologische Untersuchungen in Der Mongolischen
Volksrepublik. Academia Nauk SSSR. 210-223.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. In Weishampel, Dodson
and Osmolska (eds.). The Dinosauria. University of California Press. 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. 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, 24 pp.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47(1), 97-116.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Osm�lska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel,
Dodson and Osm�lska (eds.). The Dinosauria, Second Edition. University
of California Press. 165-183.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles.
Masters Thesis. University of Oslo. 48 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Currie, Funston and Osm�lska, 2016 (online 2015). New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica.
61(1), 143-157.
cf. Citipati (Kurzanov, 1987)
Middle-Late Campanian, Late Cretaceous
Iren Dabasu Formation, Inner Mongolia, China
Material- ?(AMNH 6570 in part; paratype of Ornithomimus asiaticus) partial manual ungual ?I (~39 mm) (pers. obs.)
(PIN 2549-100) femur (~160 mm) (Kurzanov, 1987)
Comments- Kurzanov (1987) briefly described and figured "a
left avimimid femur from the Upper Cretaceous Iren-Nor locality in
China (specimen PIN, no. 2549-100)" (translated), which Currie and
Eberth (1993) indicate was found by the Sino-Soviet expedition, which means it was collected between June 14 and July
17 at their localities K (= AMNH locality 141?), L or P. Kurzanov
referred it to Avimimidae based on the "accessory condyle" (which is
just the lateral condyle being separated from the ectocondylar tuber by
a fibular groove as in most theropods), the broad intercondylar flexor
groove and similarities of the trochanteric crest. Indeed, he stated "the
only not very significant difference is expressed in the fusion of the
large and small trochanters, while in Avimimus they are separated by a
narrow gap." On the other hand, Osmolska (1996) stated "There is a
great resemblance between the femur in [Bagaraatan] ostromi and the femur PIN
2549-100" in that "Both femora have similarly shaped proximal and
distal ends, ... well pronounced articular heads and femoral necks, the
poorly delimited lesser trochanters, which are as high as the greater,
and in the presence of the protuberances on the lateral surface."
Additionally, "The distal ends of femora are also similarly shaped in
both compared forms" with "distinctive tibiofibular crests ('condylus
lateralis' in Kurzanov 1987)." While PIN 2549-100 is similar in shape
to both Iren Dabasu Avimimus and Bagaraatan, the latter both have
accessory trochanters (usually misidentified as a large, distally
placed anterior trochanter in Avimimus) which are absent in PIN
2549-100, Avimimus differs from PIN 2549-100 and Bagaraatan in lacking
a distal ectocondylar notch defining the tuber, Bagaraatan differs from
PIN 2549-100 and Avimimus in having a narrow flexor groove, and PIN
2549-100 differs from at least Avimimus in having a fourth trochanter
reduced to "a
slight roughness, located almost under the head of the femur on its
medial side" (Kurzanov, 1987) (unknown in Bagaraatan). Currie and
Eberth (1993) believed PIN 2549-100 "is probably from a troodontid" and
"provisionally referred to Saurornithoides", but Averianov and Sues
(2012) concluded it "is probably troodontid but cannot be definitely
referred to Saurornithoides" and "should be listed as Troodontidae
indet." However, scoring this in Hartman et al.'s maniraptoromorph analysis results in identical scorings to Citipati osmolskae, with one more step needced to move it sister to Avimimus and two more to move it to Troodontidae (as sister to Linhevenator). Given the stratigraphic and geographic proximity, it is provisionally assigned to cf. Citipati here pending description of the holotype's femur (IGM 100/979 and 1004 have crushed and poorly exposed femora).
The hypodigm of Archaeornithomimus asiaticus includes two collections
of largely undescribed and unassociated specimens, AMNH 6570 from Third
Asiatic Field site 140 and AMNH 6576 from site 141, discovered between April 22 and May 25 1923. Based on personal
examination (July 2009), multiple elements in these collections do not
belong to Archaeornithomimus, among which is an ungual in a box of
phalanges and calcanea under AMNH 6570. It is moderately curved with a
large, proximally placed flexor tubercle and resembles both
Sinornithoides' pedal ungual I and Citipati's
manual ungual I in its
preserved portion. As it is twice the length of Iren Dabasu's
troodontid pedal unguals I but scales well to PIN 2549-100, the latter
identification is provisionally preferred here.
References- Kurzanov, 1987. Avimimidae and the problem of the origin
of birds [in Russian]. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa
ekspedicia. 31, 1-95.
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.
Osmolska, 1996. An unusual theropod dinosaur from the Late Cretaceous Nemegt
Formation of Mongolia. Acta Palaeontologica Polonica. 41, 1-38.
Averianov and Sues, 2012. Correlation of Late Cretaceous continental vertebrate
assemblages in middle and central Asia. Journal of Stratigraphy. 36(2), 462-485.
unnamed citipatiine (Dong and Currie, 1995)
Late Campanian, Late Cretaceous
Wulansuhai Formation (= Bayan Mandahu 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
Comments- IVPP V9608 was referred to Oviraptor philoceratops by
Dong and Currie (1996), but Longrich et al. (2010) referred
it to Oviraptorinae indet. and Hartman et al. (2019) recovered it as an oviraptorine in the Conchoraptor+Citipati clade excluded from Khaan+Conchoraptor. After additional taxa were added to the Hartman et al. matrix, it resolved sister to Citipti and "Anomalipes".
Reference- 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(4), 631-636.
Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from
the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5),
945-960.
Rinchenia Barsbold, 1997
R. mongoliensis (Barsbold, 1986) Barsbold, 1997
= Oviraptor mongoliensis Barsbold, 1986
= Citipati mongoliensis (Barsbold, 1986) Paul, 2010
Early Maastrichtian, Late Cretaceous
Altan Uul, Nemegt Formation, Mongolia
Holotype-
(IGM 100/32A) skull (156 mm), mandibles (150 mm), atlantal
intercentrum, atlantal neuropophyses, axial intercentrum, axis,
third-eleventh cervical
vertebrae, first-eleventh dorsal vertebrae, gastralia, synsacrum
(217 mm), twenty-seven caudal vertebrae, fourteen chevrons, scapulae
(~203 mm), coracoid, furcula, partial sternal plate, four sternal ribs,
humerus (163 mm),
incomplete radius (~103.3 mm), incomplete ulna (~116 mm), phalanx I-1
(~72 mm), proximal manual ungual I, three metacarpal or phalangeal
fragments, ilium (225 mm), proximal pubis, proximal ischium, femur (287
mm), incomplete tibia (~370 mm), proximal
fibula, metatarsal I (~22 mm), phalanx I-1 (~30 mm), pedal ungual I
(~22 mm), distal phalanx II-1, pedal ungual II (~28 mm), proximal and
distal metatarsal III, proximal phalanx III-1, phalanx III-2 (~36 mm),
phalanx III-3 (23.8 mm), pedal ungual III (~32 mm), proximal and distal
metatarsal IV, partial phalanx IV-1, phalanx IV-2 (17 mm), phalanx IV-3
(12 mm), phalanx IV-4 (11.9 mm), pedal ungual IV (33 mm)
Diagnosis- (after Funston et
al., 2018) tall, domed cranial crest composed primarily of nasals;
frontal taller than anteroposteriorly long; postorbital with vertical
frontal process parallel to jugal process; interfingering
jugal-quadratojugal contact; ventral process of dentary extending
posterior to external mandibular fenestra (also in Citipati); angular not contributing to retroarticular process (also in Citipati);
rounded hypapophyses on anterior dorsals; six sacral vertebrae;
plate-like distal chevrons; unfused scapulocoracoid with straight
posteroventral process of coracoid; deltopectoral crest of humerus
ventrally tapering; ilium anterodorsally expanded; preacetabular
process of ilium ventrally hooked and pointed; tall but
anteroposteriorly restricted brevis fossa; low trochanteric crest of
femur; popliteal fossa of femur overhung by medial distal condyle.
Other diagnoses- Barsbold (1986) originally distinguished mongoliensis from Oviraptor philoceratops (including IGM 100/42) based on its smaller size, more robust build, and dome-shaped ridge on the skull roof.
Barsbold (1997) later distinguished Rinchenia from Oviraptor
(still including IGM 100/42) based on its "higher dome-like crest that
incorporates the parietals", "more lightly built" postcranium and lower
cervical vertebrae.
Comments- The holotype was discovered in 1984. Barsbold (1986) originally described this specimen as a new
species of Oviraptor, but later (1997) assigned it to a new genus, Rinchenia.
While Olshevsky (DML, 1997) declared it a nomen nudum because "the
redescription of the species and formal assignment as type species of
the new genus is in press", these are not valid reasons according to
the ICZN. In particular, it satisfies Article 11, Article 13.1.1
by being "accompanied by a description or definition that states in
words characters that are purported to differentiate the taxon" ("about
the same size as Oviraptor,
but the skull has a higher dome-like crest that incorporates the
parietals in addition to the premaxillae, nasals, and frontals. The
postcranium is more lightly built, and the cervical vertebrae are
rather low in comparison with those of Oviraptor";
contra Funston et al., 2018), and Article 13.3 by being "accompanied by
the fixation of a type species in the original publication" via Article
68.3 ("When an author establishes a new nominal genus-group taxon for a
single taxonomic species and denotes that species by an available name,
the nominal species so named is the type species").
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. (2000). Funston et al. (2018) photographs most
elements except the manus and some vertebrae. Snively (2000)
studies and illustrates a metatarsus as Rinchenia
mongoliensis, but the specimen number (IGM 100/42) shows this is actually
Citipati sp. nov.. Funston et al. also provides a brief description and states "a complete anatomical description is forthcoming."
A detailed description is present in his 2019 thesis. Fanti et al. (2012) provide measurements.
References- Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva
(ed.). Herpetologische Untersuchungen in Der Mongolischen
Volksrepublik. Academia Nauk SSSR. 210-223.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. In Weishampel, Dodson
and Osmolska (eds.). The Dinosauria. University of California Press. 249-258.
Barsbold, 1997. Oviraptorosauria. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. 505-509.
Olshevsky, DML 1997. https://web.archive.org/web/20210605031503/http://dml.cmnh.org/1997Sep/msg00863.html
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. Masters Thesis, University of Calgary. 273 pp.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy,
diversity and ecology in the Nemegt Basin. Palaeogeography,
Palaeoclimatology, Palaeoecology. 494, 101-120.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Wulatelong Xu, Tan, Wang, Sullivan,
Hone, Han, Ma, Tan and Xiao, 2013
W. gobiensis Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao,
2013
Late Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype- (IVPP V18409) (29 kg; adult) incomplete skull, partial mandible,
eleven partial dorsal vertebrae, four partial dorsal ribs, first sacral centrum,
sixteen partial caudal vertebrae, incomplete scapulocoracoids, sternal plates
(one partial, one incomplete), partial humerus, metacarpal I (~35 mm), incomplete
phalanx I-1 (78 mm), manual ungual I, metacarpal II (~76 mm), phalanx II-1 (35
mm), phalanx II-2 (33 mm), partial manual ungual II, phalanx III-1 (51 mm),
partial phalanx III-2, phalanx III-3 fragment, manual ungual III fragment, incomplete
ilium (~225 mm), incomplete pubis (~250 mm), partial ischium, incomplete femur
(255 mm), incomplete tibia (325 mm), incomplete fibula, proximal tarsus, distal
tarsal IV, incomplete metatarsal II, phalanx II-2 (27 mm), pedal ungual II (45
mm), incomplete metatarsal III (143 mm), phalanx III-1 (43 mm), phalanx III-2
(30 mm), phalanx III-3 (24 mm), pedal ungual III, metatarsal IV (139 mm), phalanx
IV-1 (30 mm), phalanx IV-2 (23 mm), phalanx IV-3 (20 mm), phalanx IV-4 (16 mm),
pedal ungual IV (35 mm), metatarsal V
Diagnosis- (after Xu et al., 2013) ventral extremity of external naris
located below midheight of premaxilla; strap-like jugal process of maxilla extends
well beyond preorbital bar posteriorly and overlaps jugal; anterodorsal process
of surangular basally constricted in lateral view.
Comments- The holotype was discovered in 2009. This entry switches the
identification of manual digits II and III (III and IV of the authors) based
on figure 2A, as there is no room for a third non-ungual phalanx in the uppermost
digit. The length of the complete phalanx in the other digit and fragmentary
nature of more distal phalanges makes this uncertain, however. Xu et al. (2013)
proposed Wulatelong was a basal oviraptorid, but it clades with Oviraptor when additional taxa are added to the Hartman et al. analaysis.
Reference- Xu, Tan, Wang, Sullivan, Hone, Han, Ma, Tan and Xiao, 2013.
A new oviraptorid from the Upper Cretaceous of Nei Mongol, China, and its stratigraphic
implications. Vertebrata PalAsiatica. 51(2), 85-101.
"Tongtianlong" Lu, Chen, Brusatte,
Zhu and Shen, 2016
"T. limosus" Lu, Chen, Brusatte, Zhu and Shen, 2016
Early Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Material- (DYM-2013-8) skull (~189 mm), mandible (119 mm), (cervical
series 410 mm) atlas, axis (17 mm), third cervical vertebra ( mm), fourth cervical
vertebra ( mm), fifth cervical vertebra ( mm), sixth cervical vertebra ( mm),
seventh cervical vertebra ( mm), eighth cervical vertebra ( mm), ninth cervical
vertebra ( mm), tenth cervical vertebra ( mm), eleventh cervical vertebra (
mm), (dorsal series 350 mm) several partial dorsal vertebrae, dorsal ribs, sacrum,
nineteen caudal vertebrae, chevrons, incomplete scapulae (185 mm), coracoids
(72 mm), furcula, partial sternum, humeri (131 mm), incomplete radius (102 mm),
fragmentary ilium, partial pubes, partial ischia, partial femur (250 mm), incomplete
tibia (320 mm), incomplete fibula, astragalus, calcaneum, partial tarsometatarsus
(II 125, III 135, IV 110 mm), phalanx II-1, phalanx II-2, pedal ungual II, phalanx
III-1, phalanx III-2, phalanx III-3, pedal ungual III, metatarsal V (38 mm)
Diagnosis- (after Lu et al., 2016) dome-like skull roof with highest
point located above posterodorsal corner of orbit; anterior margin of premaxilla
highly convex in lateral view; distinct process at middle of anterior parietal
margin; lacrimal shaft anteroposteriorly long in lateral view, with flat lateral
surface; foramen magnum smaller than occipital condyle (also in Incisivosaurus
and Anzu); absence of dentary symphyseal ventral process; absence of
posterolateral sternal process.
Comments- "Tongtianlong" was first announced by Lu et al. (2015) was later described by Lu et al. on November 10 2016 as a new taxon
of oviraptorid. However, this paper has no mention of ZooBank
and as of February 6 2020
"Tongtianlong" lacks an entry on the ZooBank website. Thus
according to ICZN Article 8.5.3 (an electronic work must "be
registered in the Official Register of Zoological Nomenclature
(ZooBank) (see Article 78.2.4) and contain evidence in the work itself
that such registration has occurred"), "Tongtianlong limosus" Lu
et al., 2016 is a nomen nudum that will only be technically valid
pending action on behalf of the authors or ICZN as its journal is not
published physically.
Lu et al. (2016) recovered this as a heyuannine
sister to Wulatelong+Banji in their analysis using a modified
version of Maryanska et al.'s oviraptorosaur matrix. In Hartman et al.'s matrix this clades with Oviraptor and Wulatelong when more taxa are added.
References-
Lu, Chen, Kobayshi and Lee, 2015. A new oviraptorid dinosaur (Dinosauria: Oviraptorosauria)
from the Late Cretaceous of southern China. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 168.
Lu, Chen, Brusatte, Zhu and Shen, 2016. A Late Cretaceous
diversification of Asian oviraptorid dinosaurs: Evidence from a new species
preserved in an unusual posture. Scientific Reports. 6, 35780.
Oviraptor Osborn, 1924b
= "Fenestrosaurus" Osborn, 1924a
O. philoceratops Osborn, 1924b
= "Fenestrosaurus philoceratops" Osborn, 1924a
Late Campanian, Late Cretaceous
Bayn Dzak, Djadochta Formation, Mongolia
Holotype- (AMNH 6517) incomplete skull (~179 mm), mandibles (195 mm), cervical
vertebrae 4-12, dorsal vertebrae 1-7, several dorsal ribs, four uncinate processes,
scapula (231 mm), partial coracoid, furcula, sternal ribs, humerus (185 mm),
radius (163 mm), ulna (147 mm), phalanx I-1 (79 mm), carpometacarpus (107 mm;
mcII 95 mm), phalanx II-1 (54 mm), phalanx II-2 (68 mm), manual ungual II (~47
mm), metacarpal III (99 mm), phalanx III-1 (39 mm), phalanx III-2 (37 mm), phalanx
III-3 (28 mm), manual ungual III, partial ilium
Referred- ....(AMNH 6508) ~15 eggs, nest (Osborn, 1924b)
....(AMNH 33092) (embryo or nestling) tibia (58.7 mm), metatarsal III,
metatarsal IV (Norell, Balanoff, Barta and Erickson, 2018)
Late Cretaceous?
Mongolia?
?(IGM 100/791) (Balanoff and Norell, 2012)
Other diagnoses- (after Osborn,
1924b) eight craniofacial and mandibular fenestrations [all
plesiomorphic- external naris, antorbital fenestra, promaxillary
fenestra, orbit, laterotemporal fenestra; supratemporal fenestra;
external mandibular fenestra- except "md.f.1" which is probably an
eroded surangular fossa]; cranium exceeding facial region in length
[also in other oviraptorosaurs]; edentulous [also in other
caenagnathoids]; skull extremely abbreviate, orbit and fenestrations
exceptionally large [vague but at least applicable to other
oviraptorids]; large interclavicle [actually the furcula, also in other
oviraptorids]; manus tridactyl [also in other maniraptoromorphs];
metacarpals abbreviated [unclear]; digits irregularly elongated as in
the Ornithomimidae [plesiomorphic and not like ornithomimids]; elements
of digits not compressed laterally [plesiomorphic, stated different
from Tanycolagreus and Chirostenotes].
Comments- The first oviraptorid discovered (on July 3, 1923), it was initially annouced as Fenestrosaurus philoceratops in an article by Osborn (1924a). Osborn describes it as "a small birdlike dinosaur,
remains of which were
found resting on top of
one of the nests of dinosaur eggs; hence the specific name philoceratops, signifying "ceratops lover", "egg-eating or ovivorous. This dinosaur was without teeth." Oviraptor was originally
referred to the Ornithomimidae by Osborn (1924b).
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. Norell et al. (2018) reported and illustrated an
embryo or nestling hindlimb associated with the specimen (AMNH 33092)
to "be described in detail in another paper."
The holotype was only briefly illustrated and described by Osborn
(1924b), but a detailed redescription has not yet appeared. Smith's
(1992) attempt made many errors, which were corrected by Clark et al.
(2002) in their description of the skull. Makovicky (1995) added
information about the preserved vertebrae, though the appendicular
remains remain largely ignored. Carrano (1998) listed the femoral
length as 282.0 mm, dspite the hindlimb being unpreserved. Norell
et al. (2018) included
photos of the cervical and pectoral areas.
Osborn (1924b) initially assigned Oviraptor to Ornithomimidae, and even
after the naming of Oviraptoridae and discovery of more genera, was not placed
in a cladistic context until Barsbold et al. (1990). The latter authors positioned
it outside Heyuanninae. This is found
in nearly all analyses since, despite the massive increase in taxon number.
The remaining unknown is whether Oviraptor is basal to Citipati+Heyuannia,
or closer to some taxa with short dentaries.
Not Oviraptor- Barsbold (1976) erected Oviraptoridae
for the holotype 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). 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 almost all oviraptorids
being Oviraptor in modern phylogenies. Dong and Currie (1996) referred
a fragmentary skeleton (IVPP V9608) to Oviraptor philoceratops, but this
may be Citipati instead and was referred to Oviraptorinae indet. by Longrich
et al. (2010). Morell (1997) labeled IGM 100/1002 Oviraptor before it
was made a paratype of Khaan by Clark et al. (2001). Jensen (2008) calls
the tall-crested cast PMO X678 Oviraptor sp., which may be the same taxon
as a skull and manus on the Witmer Lab website (Witmer, 2012 online) labeled Oviraptor philoceratops,
but how taxon this related to Oviraptor requires further study. Balanoff and Norell (2012) list IGM 100/791 as a specimen of Oviraptor philoceratops, but it is mentioned nowhere else except a similar list in Balanoff's (2011) thesis. Numerous
other oviraptorid specimens, have been called Oviraptor, but only the
holotype can be properly referred to the genus.
References- Osborn, 1924a. The discovery of an unknown continent. Natural
History. 24(2), 133-149.
Osborn, 1924b. Three new Theropoda, Protoceratops zone, central Mongolia.
American Museum Novitates. 144, 1-12.
Brown and Schlaikjer, 1940. The structure and relationships of Protoceratops.
Annals of the New York academy of Sciences. 40(3), 133-266.
Barsbold, 1976a. O novum pozdnemelovom semeystve melkikh teropod
(Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3),
685-688.
Barsbold, 1976b. A new Late Cretaceous family of small
theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk
SSSR. 226, 221-223.
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. The predatory dinosaurs - Oviraptors. In Vorobyeva
(ed.). Herpetologische Untersuchungen in Der Mongolischen
Volksrepublik. Academia Nauk SSSR. 210-223.
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. 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 f�r 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 (eds.). Encyclopedia of Dinosaurs. 505-509.
Morell, 1997. The Origin of Birds: the Dinosaur Debate. Audubon. March-April,
36-45.
Carrano, 1998. The evolution of dinosaur locomotion: Functional morphology,
biomechanics, and modern analogs. PhD thesis, The University of Chicago. 424
pp.
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.
Osm�lska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel,
Dodson and Osm�lska, (eds). The Dinosauria, Second Edition. University
of California Press (Berkeley). pp. 165-183.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles.
Masters Thesis. University of Oslo. 48 pp.
Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria: Theropoda) from
the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology. 53(5),
945-960.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution.
PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria:
Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Witmer, 2012 online. https://people.ohio.edu/witmerl/collections/Theropods/oviraptor.htm
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati osmolskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.
Heyuanninae Yun, 2019
Definition- (Heyuannia huangi + Conchoraptor gracilis) (Yun, 2019)
= "Ingeniinae" Barsbold, 1981
Definition- ("Ingenia" yanshini + Conchoraptor
gracilis) (Osmolska, Currie and Barsbold, 2004)
Comments- Barsbold (1981) created Ingeniinae to separate his new genus Ingenia from Oviraptor (to which he referred specimens now distinguished as Conchoraptor
and Citipati sp.).
However, as Taylor (DML 2004) noted, the genus is preoccupied by a
tripyloidid nematode (Gerlach, 1957) so any family level taxon based on
it must refer to nematodes. A search of the nematode literature
suggests that the internal struicture of Tripyloididae reamins
unresolved and that no nematode subfamily Ingeniinae has been
proposed. Once Barsbold 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. Osmolska et al. (2004) officially defined the taxon, as "Conchoraptor gracilis, Ingenia yanshini, their most recent common ancestor, and all descendants." More recently, in Funston and Currie (2016) Machairosaurus, Jiangxisaurus, Nemegtomiaia and Heyuannia would also fall under this definition. Adding taxa to Hartman et al.'s matrix would also include Khaan and Gobiraptor.
Yun
(2019) noted that the subfamily needed to be renamed, so suggested
Heyuanninae as a replacement with the definiotion "the least inclusive
clade containing Heyuannia huangi and Conchoraptor gracilis." As Heyuannia is almost always sister to "Ingenia"in
oviraptorosaur topologies, the definitions usually cover the same
taxa. Although the correct stem would be Heyuanniinae, ICZN
Article 29.5 states "If a spelling of a family-group name was
not formed in accordance with Article 29.3 but is in prevailing usage,
that spelling is to be maintained, whether or not it is the original
spelling and whether or not its derivation from the name of the type
genus is in accordance with the grammatical procedures in Articles
29.3.1 and 29.3.2." More recently Funston et al. (2020) used
Heyuanninae as a stem-based group opposing their new subfamily
Citipatiinae although they did not explicitly define it as such.
If such a definition is proposed in the future, Oviraptor
needs to be an additional external specifier because it cannot be a
heyuannine according to the ICZN (it must be an oviraptorine).
References- Gerlach, 1957. Die Nematodenfauna des Sandstrandes an der K�ste von
Mittelbrasilien (Brasilianische Meerse-Nematoden IV). Mitteilungen aus
dem Zoologischen Museum in Berlin. 33(2), 411-459.
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. 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. PhD thesis, Southern Methodist University. 249 pp.
Osm�lska, Currie and Barsbold, 2004. Oviraptorosauria. In Weishampel,
Dodson and Osm�lska, (eds.). The Dinosauria, Second Edition. University
of California Press. 165-183.
Taylor, DML 2004. https://web.archive.org/web/20200928000123/http://dml.cmnh.org/2004Sep/msg00022.html
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Yun, 2019. Heyuanninae clade nov., a replacement name for the
oviraptorid subfamily "Ingeniinae" Barsbold, 1981. Zootaxa. 4671(2),
295-296.
Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new
two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.
Royal Society Open Science. 7: 201184.
undescribed heyuannine (Maryanska et al., 2002)
Early Maastrichtian, Late Cretaceous
Barun Goyot or Nemegt Formation, Mongolia
Material- (ZPAL MgD-I/106) material including partial skull, mandible
and
postcranium including cervical vertebrae, dorsal vertebrae, caudal
vertebrae, chevrons, partial scapulacoracoid, sternal plate, manual ungual
I, manual phalanges III-?, ilium, pubis, ischium, partial femur, metatarsal
II, metatarsal III and metatarsal IV
Comments- This crestless specimen was listed as Conchoraptor gracilis
by Maryanska et al. (2002) and Fanti et al. (2012), the latter of which
provide a few measurements. Balanoff (2011) scored it in her
thesis and found it resolved as a heyuannine between Heyuannia huangi and H. yanshini. The specimen number indicates it is either from the Barun Goyot
or Nemegt Formation.
References-
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution.
PhD thesis. Columbia University. 522 pp.
undescribed heyuannine (U.S. Attorney's Office Southern District of New York, online 2014)
Late Campanian(?), Late Cretaceous
Khulsan or Red Beds of Khermeen Tsav, Barun Goyot Formation, Mongolia
Material- (CMMD coll.) (upper left) (juvenile) skull,
mandibles, hyoid, several cervical vertebrae, twelve dorsal vertebrae,
several dorsal ribs, gastralia, first sacral vertebra, posterior sacral
or proximal caudal centrum, scapula, furcula(?), humerus, radius, ulna,
partial ilia, pubes, ischia, femora, tibiae, fibula, astragali,
metatarsal I, phalanx I-1, pedal ungual I, metatarsals II, phalanges
II-1, phalanges II-2, pedal unguals II, metatarsal III, phalanx III-1,
phalanx III-2, phalanx III-3, pedal ungual III, metatarsals IV,
phalanges IV-1, phalanges IV-2, phalanx IV-3, phalanx IV-4, pedal
ungual IV, metatarsal V
....(upper right) (juvenile) fragmentary skull, mandibles, six posterior cervical
vertebrae, few cervical ribs, five anterior dorsal vertebrae, posterior
dorsal vertebral fragments, partial dorsal ribs, two posterior sacral
centra, humerus, ulna, partial ilium, pubes, ischial fragment, femora,
tibiae, fibulae, astragalus, calcaneum, metatarsals I, pedal ungual I,
distal tarsal IV, metatarsals II, phalanges II-1, phalanges II-2, pedal
unguals II, metatarsals III, phalanges III-1, phalanges III-2,
phalanges III-3, pedal unguals III, metatarsals IV, phalanges IV-1,
phalanges IV-2, phalanx IV-3, phalanges IV-4, pedal ungual IV
....(cast UALVP 54983-1; lower left) (juvenile) skull, sclerotic plates, mandibles, hyoid, several
cervical vertebrae, nine dorsal vertebrae, several dorsal ribs, two
uncinate processes, gastralia, sacrum, ~twenty-six caudal vertebrae (c1 23, c3 19, c4 20, c5 18, c6 18 mm),
nine chevrons, scapula, coracoids, partial sternum, humerus, radius,
ulna, ilia, pubes, ischia, femur, tibia, fibula, astragalus, calcaneum,
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
....(cast UALVP 54983-3; lower right) (juvenile) posterior dorsal centrum, five posterior dorsal ribs,
gastralia, three sacral centra, first to eighth caudal vertebrae (c4 24, c5 24, c6 24 mm), first
to eighth chevrons, ilia, pubes (one distal), ischial fragments, femora
(one partial), tibiae (one proximal, one distal), fibulae (one
proximal, one distal), astragalus, metatarsals II (one partial),
partial metatarsals III, partial metatarsals IV
....(center right) (juvenile) fragmented skull, mandibles, several presacral
vertebrae, few dorsal ribs, uncinate process, gastralia, scapulae,
partial coracoid, humeri, radii, ulnae, carpals, semilunate carpal,
metacarpal I, manual ungual I, metacarpal II, phalanx II-?, metacarpal
III, phalanx III-2, phalanx III-3, manual unguals III
....
(cast UALVP 54983-2; upper middle) (juvenile) eight proximal caudal
vertebrae (c3 20, c4 19, c5 18, c6 18 mm), distal caudal fragments,
eight chevrons
Comments- This oviraptorid
bonebed was privately owned until a legal case (U.S. Attorney's Office
Southern District of New York,
online 2014) where smuggled dinosaurs were being returned to the CMMD
in Mongolia, including "a rock matrix containing at least four
Oviraptors." Casts are available at e.g. Gaston Design (online
2017) as Conchoraptor from the REd Beds of Khermeen Tsav. One is catalogued as UALVP 54983 (Funston, 2019) where they are listed as being Conchoraptor
from Khulsan in the online catalogue. Funston et al. (2015) lists
caudal and femoral measurements for two individuals, while Persons et
al. (2015) lists caudal measurements for three individuals (as 54983-1,
54983-2 and 54983-3), in both references as Conchoraptor gracilis.
Unfortunately, the size of each speciem is so similar that it's unsure
which skeletons had first caudal and femoral measurements of 23.9 and
213 mm and 22.9 and 212 mm respectively in Funston et al.'s (2015)
table.
References- U.S. Attorney's Office Southern District of New York, online 2014. Manhattan U.S. Attorney Announces Return To Mongolia Of Fossils Of Over 18 Dinosaur Skeletons. July 10.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied
caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of
Alberta, Canada. Cretaceous Research. 54, 179-187.
Persons, Funston, Currie and Norell, 2015. A possible instance of
sexual dimorphism in the tails of two oviraptorosaur dinosaurs.
Scientific Reports. 5:9472.
Gaston Design, 2017 online. https://www.gastondesign.com/product/818/
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Conchoraptor Barsbold, 1986
C. gracilis Barsbold, 1986
= Citipati gracilis (Barsbold, 1986) Paul, 2010
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav of Barun Goyot Formation, Mongolia
Holotype- (IGM 100/20) skull (~108 mm), posterior mandible
Paratype?* (see Comments)- (IGM 100/38) metacarpal I (24.6 mm), phalanx I-1 (32.9 mm),
manual ungual I (~26 mm), metacarpal II (42.8 mm), distal phalanx II-1, phalanx
II-2 (26.3 mm), manual ungual II (~23 mm), metacarpal III (41.4 mm), phalanx
III-1 (17.6 mm), phalanx III-2 (~19 mm), phalanx III-3 (~25 mm), manual ungual
III (~22 mm)
Referred- (IGM 100/21) mandible (93 mm) (Barsbold, 1976)
(IGM 100/36) material including skull, furcula, ilium (176 mm), pubis (145 mm),
femur (182 mm), tibia (215 mm), metatarsal III (102.2 mm), phalanx III-1 (31.1
mm), phalanx III-3 (14.6 mm), phalanx IV-2 (13.5 mm), phalanx IV-3 (9.6 mm),
phalanx IV-4 (7.3 mm), pedal ungual IV (18.9 mm) (Barsbold 1981)
(IGM 100/39) material including metacarpal I (~23 mm), phalanx I-1 (41 mm),
manual ungual I (~27 mm), phalanx II-2 (~31 mm), manual ungual II (~31 mm),
metacarpal III (44 mm), phalanx III-3 (~24 mm), manual ungual III (20 mm) (Maryanska
et al., 2002)
(IGM 100/46) material including (?)metatarsal II, (?)metatarsal III, pedal phalanx
III-3 (18.2 mm), (?)metatarsal IV, phalanx IV-2 (15.9 mm), phalanx IV-3 (11.5
mm), phalanx IV-4 (11.2 mm), pedal ungual IV (24.9 mm), (?)two pedal unguals,
(?)metatarsal V (Maryanska et al., 2002)
(IGM 100/47) materal including ilium (173 mm), femur (195 mm) and tibia (258
mm) (Maryanska et al., 2002)
(IGM 100/1275) partial skeleton including sacrum, twenty-seven caudal vertebrae
(first caudal 24.5 mm), pygostyle, femur (212 mm), tibia (250 mm), metatarsal
III (107.5) and phalanx III-1 (35.3 mm) (Lu et al., 2013)
(IGM 102/2) four mid caudals, nine mid chevrons (Fanti et al., 2012)
(IGM 102/3) material including partial skull, posterior dorsal fragments,
dorsal ribs, uncinate processes, gastralia, last three sacral
vertebrae, first to tenth caudal vertebrae, first to
ninth chevrons, partial scapula, sterna, humerus, radius (99 mm), ulna (100.1 mm), metacarpals I (24 mm),
phalanges I-1 (31 mm), manual unguals I (42 mm), metacarpal II (~44 mm), phalanx II-1 (18.3 mm), phalanx
II-2 (15.2 mm), manual ungual II (22 mm), metacarpal III, manual ungual III, posterior
ilia (226 mm), pubes, ischium, femora (240 mm), tibiae (277 mm), fibula, astragali, calcanea,
distal tarsals III, distal tarsals IV, metatarsal I, phalanges I-1,
pedal unguals I, metatarsals II, phalanges II-1, phalanges II-2, pedal
unguals II, metatarsals III (121 mm), phalanx III-1 (31 mm), phalanx III-2, phalanges
III-3 (17.2 mm), pedal unguals III, metatarsals IV, phalanges IV-1, phalanx IV-2 (14.4 mm),
phalanx IV-3 (10.9 mm), phalanges IV-4 (8.7 mm), pedal unguals IV (33 mm), metatarsals V (Fanti et al., 2012)
(IGM MAE 97-212 Block 1B) material including ilium (74.5 mm) and femur (81.2
mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 2B) material including ilium (75 mm), femur (80 mm), metatarsal
III (52.2 mm) and phalanx III-1 (14.1 mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 7A) material including ilium (77.7 mm) and femur (82.2
mm) (Lu et al., 2013)
(IGM MAE 97-212 Block 7B) material including femur (87.3 mm) and tibia (110
mm) (Lu et al., 2013)
(ZPAL MgD-I/95) incomplete skull (98 mm), mandibles (86 mm), metatarsus (141
mm) (Osmolska, 1976)
(ZPAL MgD-I/101) skull, postcranial fragments (ZPAL online 2006)
?(private coll.) skull, mandible, several cervical vertebrae, few
dorsal fragments, dorsal ribs, sacral fragments, over ten caudal
vertebrae, at least eight chevrons, scapulae, partial furcula,
coracoid, humerus, radius, ulna, metacarpal I, phalanx I-1, manual
ungual I, metacarpal II, phalanx II-1, metacarpal III, phalanx III-1,
distal digits II and III, ilia, pubes, ischia, femora, tibiae, fibulae
(one proximal), tarsus, metatarsal II, phalanges II-1, phalanges II-2,
pedal ungualsII, metatarsal III, phalanges III-1, phalanges III-2,
phalanges III-3, pedal ungual III, metatarsal IV, phalanges IV-1,
phalanges IV-2, phalanx IV-3, phalanx IV-4, pedal unguals IV,
metatarsal (Witmer, 2012 online 1)
?(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 (Gaston Designs, 2019 online 2)
Late Campanian, Late Cretaceous
Khulsan, Baron Goyot Formation, Mongolia
(UALVP 54984; cast?) material including first-sixth caudal vertebrae
(c2 23, c3 23, c4 23, c5 25, c6 23 mm) and four chevrons (Persons et
al., 2015)
(UALVP 54986; cast?) material including first-sixth caudal vertebrae
(c1 19, c2 18, c3 18, c4 18, c5 17, c6 17 mm) and four chevrons
(Persons et al., 2015)
(UALVP 54987; cast?) material including first-sixth caudal vertebrae
(c2 20, c3 23, c4 21, c5 24, c6 23 mm) and three chevrons (Persons et
al., 2015)
(UALVP 54988; cast?) material including first-sixth caudal vertebrae
(c1 26, c3 22, c4 21, c5 20, c6 19 mm) and three chevrons (Persons et
al., 2015)
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadochta Formation, Mongolia
(IGM 100/1203) material including posterior axis, third cervical
vertebra, dorsal ribs, uncinate processes, scapula, ilium (158 mm),
ischium, femur (154 mm), tibia (184 mm), metatarsal III (83.5 mm) and
phalanx III-1 (24 mm) (Balanoff, 2011)
(IGM 100/3006) material including partial skull, proximal caudal vertebrae,
chevrons, scapula, coracoid, femur (152.7 mm), metatarsal III (87.5 mm) and
phalanx III-1 (24.8 mm) (Balanoff, 2011; braincase described by Balanoff
et al., 2014)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(IGM uncatalogued) material including incomplete skull, mandibles (Funston, online 2018)
Late Campanian(?) or Early Maastrichtian, Late Cretaceous
Barun Goyot or Nemegt Formation, Mongolia
(ZPAL MgD-I/99) material including scapula and ilium (270 mm) (Fanti et al.,
2012)
(ZPAL MgD-I/100) material including partial scapula, humerus (108 mm), phalanx
I-1 (18 mm), manual ungual I (19 mm), phalanx II-1 (21 mm), phalanx II-2 (22
mm), manual ungual II (24 mm), ilium (210 mm), partial femur (Maryanska et al.,
2002)
Late Cretaceous
Mongolia
(FPDM-V6232) skull, skeleton (Goto, Ichishima and Ji, 2005)
(FPDM-V6234) skull, skeleton (Goto, Ichishima and Ji, 2005)
(IGM 97/212) specimen including femur (250 mm) (Erickson et al., 2009)
(IGM 100/19) (Balanoff and Norell, 2012)
(IGM 100/42; note not the same as the Citipati sp. complete skeleton)
specimen including tibia (225 mm), phalanx III-3 (17.4 mm), phalanx IV-2 (18.2
mm), phalanx IV-3 (13.1 mm), phalanx IV-4 (11.9 mm), phalanx IV-5 (25.5 mm)
(Fanti et al., 2012)
(IGM 100/97) material including femur (185 mm) and tibia (258 mm) (Fanti et
al., 2012)
(IGM 102/1) material including partial tibia, phalanx IV-2 (15.2 mm) (Fanti
et al., 2012)
(IGM 110/7) material including skull (94 mm), mandible (87.2 mm) (Fanti et
al., 2012)
(IGM 110/10) material including mandible (~55 mm) (Fanti et al., 2012)
(IGM 110/11) material including partial skull (Fanti et al., 2012)
(IGM 110/12) material including skull (~101 mm), partial dentary (Fanti et al.,
2012)
(IGM 110/18; actually two individuals, as listed below) material including distal
caudal vertebrae, pubis and ischium (Lu, 2004)
----(IGM 110/18a) material including humerus (91.1 mm), radius (87.3 mm), ulna
(88.2 mm) (Fanti et al., 2012)
----(IGM 110/18b) material including radius (~78.9 mm), ulna (75.1 mm) (Fanti
et al., 2012)
(IGM 110/19) material including thirty-two caudal vertebrae, ilium (~115 mm)
(Barsbold et al., 2000)
(IGM 110/20) material including partial skull (Fanti et al., 2012)
(IGM 110/21) material including mandible (68.2 mm), humerus (61.3 mm), radius
(~54.6 mm), ulna (55 mm), metacarpal I (22.8 mm), partial metacarpal II, tibia
(164 mm) (Fanti et al., 2012)
(IGM 110/22) material including partial skull (Fanti et al., 2012)
(IGM 110/25) material including scapula (Fanti et al., 2012)
(IGM 110/26) material including pedal phalanx III-3 (10.6 mm), phalanx IV-2
(13.9 mm), pedal ungual IV (17.9 mm) (Fanti et al., 2012)
(IGM 110/30) material including metacarpal I (26 mm), phalanx I-1 (38 mm), manual
ungual I (~30 mm), phalanx II-1 (~27.5 mm), phalanx II-2 (33 mm), manual ungual
II (25 mm) (Fanti et al., 2012)
(PIN coll.) skull (92 mm), mandibles (68 mm) (Maryanska et al., 2002)
(PMO X677; cast) skull, mandibles (Jensen, 2008)
(UALVP 49391; cast) skull (118 mm), mandible (89 mm) (Fanti et al., 2012)
(UALVP 49392; cast) skull (99 mm), mandible (86.6 mm) (Fanti et al., 2012)
(cast) skull, mandibles (Witmer, 2012 online 2)
(cast) skull, mandibles (Witmer, 2012 online 3)
Diagnosis- (after Funston et
al., 2018) maxilla with large accessory antorbital fenestra; nasals
with three dorsal fenestrae per side; postorbital extending nearly to
the bottom of the orbit.
Other diagnoses- (after
Barsbold, 1986) unreduced second and third manual digits
(plesiomorphic); narrow and straight manual unguals II and III
(plesiomorphic).
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) noted cranial characterics of Khermeen Tsav oviraptorids,
probably including IGM 100/20 as well as Heyuannia yanshini 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 a skull and manus as that specimen
number. Funston et al. (2018) also list 100/20 as including the
manus, but Fanti et al.'s (2012) measurement table would suggest the
manus (with incomplete unguals I and II and partial phalanx II-1) is
IGM 100/38. Whether 100/36 was separated from 100/20 after 1986,
Fanti et al. incorrectly listed 100/20 measurements as 100/36 or some
other explanation is correct is unknown. Barsbold et al. (1990)
illustrated the skull in dorsal view and also a metatarsus and two
pedal unguals, which may belong to IGM 100/46 since it is the only
original specimen given pedal measurements by Fanti et al.. The skull
was photographed incorrectly mounted on Heyuannia yanshini postcrania in Psihoyos (1994), as later specifically noted by Funston et al. (2018).
This same mount (with a mandible that is neither Conchoraptor's nor yanshini's holotype) was displayed at the Nakasato
Dinosaur Center as Ingenia and is now at the IGM. Fanti et al. listed IGM 100/36 among yanshini
specimens and incorrectly listed IGM 100/80-D as the holotype skull of Conchoraptor,
though Funston et al. (2018) state that holotype skull is labeled IGM
100/80-1 on the IGM mount. Funston et al. (2018) photograph and
describe the holotype skull and mandible, further described in Funston's (2019) thesis.
Osmolska (1976) described a skull and mandibles (ZPAL MgD-I/95) as Oviraptor
sp.. She also noted an additional skull and postcranial fragments from the
same formation, and a fragmentary skull from the Nemegt Formation. She believed
they and the IGM specimens noted by Barsbold (1976) belonged to a new species.
The Nemegt skull (probably ZPAL MgD-I/96) is crested and thus not Conchoraptor,
but the other specimens may be. Possible specimen numbers for the second Khermeen
Tsav skull are MgD-I/101 (listed on the ZPAL website as oviraptorid), and MgD-I/100
or MgD-I/106 (listed as Conchoraptor by Maryanska et al., 2002; but not
given cranial measurements by Fanti et al., making them less likely candidates).
ZPAL MgD-I/95 was illustrated as Oviraptor yanshini by Paul (1988), while
Elzanowski (1999) described its palate in detail and called it Oviraptor
sp..
Specimen numbers indicate all ZPAL specimens are from the Barun Goyot
or Nemegt Formations. Maryanska and Osmolska (1997) stated it was
probably Conchoraptor
or yanshini, 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 Funston et al. (2018) 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.
Fanti et al. (2012) lists MPC-D 102/03 as a specimen of Conchoraptor and provides measurements, although they say "a pair of Ingenia
collected at Khermeen Tsav [MPC-D 102/02, MPC-D 102/03] by an earlier
"Dinosaurs of the Gobi" expedition." Funston et al. (2016) cites
the specimen as "Conchoraptor gracilis
(MPC-D 102/3; M. A. Norell pers. comm.)" containing multiple
individuals, and his thesis confirms they were found at Khermeen Tsav
in 2002. Thus the "Ingenia"
pair IGM 110/2 and 110/3 in Fanti et al.'s list are probably different,
and indeed 102/3 is about twice the size of 110/3, and Funston's (2019)
figures confirm 102/3 matches that number in Fanti et al.'s measurement
table. Funston reported online (2018) that he was describing an
incomplete
skull (IGM uncatalogued) found in 2018 and
postcranial skeletion (which is IGM 102/3- Funston, 2019) for his 2019
thesis (7-5-2019 Facebook). The
first manual digit seems far more robust than the holotype/paratype,
but further evaluation pends on publication of the thesis.
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 yanshini
by Glut (1997) and Witmer (online, 2012 4), as Witmer notes it is from the PIN.
It was also photographed in Currie (2001). Osmolska et al. (2004) illustrate
a skull as Conchoraptor that appears to be neither the holotype, ZPAL
MgD-I/95 or the PIN specimen, and may therefore be one of the other specimens
mentioned above. Jensen (2008) analyzed a cast for his thesis (PMO X677) which
he referred to Conchoraptor gracilis. Erickson et al. (2009) examined
the histology of IGM 97/212, which they referred to Conchoraptor gracilis.
Balanoff and Norell (2012) mention many postcranial characters, largely
from IGM 100/1203 and 100/3006, noting an in prep. paper.
They also list IGM 100/19 as a specimen of Conchoraptor gracilis,
but it is mentioned nowhere else except a similar list in Balanoff's
(2011) thesis. This thesis states both are from Ukhaa Tolgod in
the Djadochta Formation, although Funston et al. (2018) stated 100/3006
is from Khulsan in the Barun Goyot Formation. Cervicals and the
ribcage of IGM 100/1203 are photographed in Norell et
al. (2018). Fanti et al. and Lu et al. (2013) listed numerous
specimens as belonging to this species and provided measurements. Among
these is IGM 100/1275, which Balanoff and Norell (2012) mention
numerous details of and Persons et al. (2013) describe the tail of.
The latter authors note this specimen is one of many partial skeletons
found together and catalogued as IGM MAE 97-212. Other specimens such
as IGM 100/1203 and 3006 may also belong here, and Balanoff et al.
(2014) have described the braincase of IGM 100/3006 based on Balanoff's
(2011) thesis.
It is especially common to find crestless privately held oviraptorid specimens
being referred to Conchoraptor.
One such incomplete skeleton ("Conchoraptor Juvenile in 3-D matrix
block" on Gaston Design) 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 (cast is
offered by Gaston Design as "Conchoraptor Adult in matrix 44"" - Gaston
Design, 2019 online 2) is more difficult to identify as it doesn't
preserve arms, though the constricted third metatarsal is unlike Heyuannia yanshini. Witmer Labs has a cast offered by Gaston
Designs as "Conchoraptor Juvenile skeleton in 2-D matrix Block".
Witmer (2012 online labs) has two skulls referred to Conchoraptor. Persons et al. (2015) list four Conchoraptor
UALVP specimens' proximal caudal measurements, which are likely casts
such as these. The UAPVP online catalogue assigns each to
Khulsan. A slab of six juvenile skeletons at the CMMD (cast UALVP
54983) has been referred to Conchoraptor in the literature (e.g. Funston et al., 2015), but is here placed as a separate heyuannine.
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. For instance, Funston et al.
(2018) provided the diagnosis listed above and listed ZPAL MgD-I-95 as
referrable, but the promaxillary fenestra size has yet to be shown in
the holotype, the number of nasal fossae might be seen in the poor
quality photo of Osmolska (1976) and the postorbital extent cannot be
measured in that specimen. This is particularily worrisome given
the discovery of additional uncrested oviraptorids such as Khaan and Heyuannia, and the possibility young specimens of
crested oviraptorids may lack crests (e.g. Yulong, Banji). Although some information
can be found in Barsbold (1976), Barsbold et al. (1990), Lu (2004), Osmolska
et al. (2004), Funston et al (2018), the specimens examined are not specified (a recent exception
is Balanoff and Norell, 2012). Similarly, the matrices of Norell et al. (2001),
Maryanska et al. (2002), Lu (2004), Fanti et al. (2012) and Lu et al. (2013)
are extremely informative and indicate basically every element is represented
in at least one specimen, but obviously don't indicate which specimens were
used to code each character. Maryanska et al. do list several specimens they
used to code their Conchoraptor OTU, but ironically the holotype was
not among them. The measurement tables in Fanti et al. and Currie et al. (2016)
will prove useful to refer specimens based on ratios once they are examined.
References- Barsbold, 1976a. O novum pozdnemelovom semeystve melkikh teropod
(Oviraptoridae fem. n.) Mongolii. Doklady Akademii Nauk SSSR. 226(3),
685-688.
Barsbold, 1976b. A new Late Cretaceous family of small
theropods (Oviraptoridae n. fam.) in Mongolia. Doklady Akademia Nauk
SSSR. 226, 221-223.
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.
Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva
(ed.). Herpetologische Untersuchungen in Der Mongolischen
Volksrepublik. Academia Nauk SSSR. 210-223.
Paul, 1988. The Predatory Dinosaurs of the World. Simon and Schuster. 464 pp.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. In Weishampel, Dodson
and Osmolska (eds.). The Dinosauria. University of California Press. 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. 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. 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. PhD thesis, Southern Methodist University. 249 pp.
Goto, Ichishima and Ji, 2005. The Flying Dinosaurs. Catalog of an
exhibition held at Fukui Kenritsu Kyōryū Hakubutsukan, July
15-November 3, 2005; also to be held at Asahikawa Kagakukan and at
Hamamatsu Kagakukan in 2006. Fukui Kenritsu Kyōryū Hakubutsukan. 118 pp.
ZPAL, online 2006. http://www.paleo.pan.pl/collect.htm#Mon-reptilia
Kundrat, 2007. Avian-like attributes of a virtual brain model of the oviraptorid
theropod Conchoraptor gracilis. Naturwissenschaften. 94, 499-504.
Kundrat and Janacek, 2007. Cranial pneumatization and auditory perceptions of
the oviraptorid dinosaur Conchoraptor gracilis (Theropoda, Maniraptora)
from the Late Cretaceous of Mongolia. Naturwissenschaften. 94(9), 769-778.
Jensen, 2008. Beak morphology in oviraptorids, based on extant birds and turtles.
Masters Thesis. University of Oslo. 48 pp.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian
physiology inherited by birds? Reconciling slow growth in Archaeopteryx.
PLoS ONE. 4(10), e7390.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution.
PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria:
Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Witmer, 2012 online 1. https://people.ohio.edu/witmerl/collections/Theropods/conchoraptor3.htm
Witmer, 2012 online 2. https://people.ohio.edu/witmerl/collections/Theropods/conchoraptor2.htm
Witmer, 2012 online 3. https://people.ohio.edu/witmerl/collections/Theropods/conchoraptor1.htm
Witmer, 2012 online 4. https://people.ohio.edu/witmerl/collections/Theropods/Ingenia.htm
Balanoff, Bever and Norell, 2014. Reconsidering the avian nature of the oviraptorosaur
brain (Dinosauria: Theropoda). PLoS ONE. 9(12), e113559.
Persons, Currie and Norell, 2014 (online 2013). Oviraptorosaur tail forms and functions. Acta
Palaeontologica Polonica. 59(3), 553-567.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied
caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of
Alberta, Canada. Cretaceous Research. 54, 179-187.
Persons, Funston, Currie and Norell, 2015. A possible instance of
sexual dimorphism in the tails of two oviraptorosaur dinosaurs.
Scientific Reports. 5:9472.
Currie, Funston and Osm�lska, 2016 (online 2015). New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica.
61(1), 143-157.
Funston, Currie, Eberth, Ryan, Chinzorig, Badamgarav and Longrich,
2016. The first oviraptorosaur (Dinosauria: Theropoda) bonebed:
Evidence of gregarious behaviour in a maniraptoran theropod. Scientific
Reports. 6:35782. DOI: 10.1038/srep35782
Funston, 2018 online. https://gregfunston.com/2018/10/12/introducing/
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy,
diversity and ecology in the Nemegt Basin. Palaeogeography,
Palaeoclimatology, Palaeoecology. 494, 101-120.
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati osmolskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Gaston Design, 2019 online 1. https://www.gastondesign.com/product/conchoraptor-juvenile-skeleton-matrix-block/
Gaston Design, 2019 online 2. https://www.gastondesign.com/product/conchoraptor-adult-matrix-2-d/
Balanoff and Norell, in prep.
Jiangxisaurus Wei, Pu,
Xu, Liu and Lu, 2013
J. ganzhouensis Wei, Pu, Xu, Liu and Lu, 2013
Early Maastrichtian, Late Cretaceous
Nanxiong Group, Jiangxi, China
Holotype- (HGM41HIII0421) (subadult) skull (150 mm), mandibles (130 mm),
hyoid (60 mm), partial atlas, axis (31 mm), third cervical vertebra (~32 mm),
fourth cervical vertebra (33 mm), fifth cervical vertebra (33 mm), sixth cervical
vertebra (32 mm), seventh cervical vertebra (28 mm), eighth cervical vertebra
(25 mm), three dorsal vertebrae (25, 26, 28 mm), nine partial dorsal ribs, first
caudal vertebra (25 mm), second caudal vertebra (25 mm), third caudal vertebra
(25 mm), fourth caudal vertebra (25 mm), fifth caudal vertebra (25 mm), sixth
caudal vertebra (22 mm), seventh caudal vertebra (20 mm), eighth caudal vertebra
(20 mm), ninth caudal vertebra (16 mm), three chevrons, scapula, coracoids,
incomplete furcula, partial sternal plates, four sternal ribs, humerus (136
mm), radius (96 mm), ulna (95 mm), scapholunare, semilunate carpal, metacarpal I
(25 mm), phalanx I-1 (30 mm), manual ungual I, metacarpal II (45 mm), phalanx
II-?, metacarpal III (35 mm), phalanx III-?, ilial fragment, pubic fragment,
ischial fragment
Diagnosis- (after Wei et al., 2013) elongated mandible (height 20% of
length); weakly downturned mandibular symphysis; surangular with elongate and
concave lateral surface; radiohumeral radio ~70%.
Comments- Wei et al. (2013) described this taxon as an oviraptorid of
uncertain phylogenetic placement, but Funston and Currie (2016) recovered it as a heyuannine closest to Nemegtomaia and Heyuannia. Hartman et al. (2019) also recovered it as a heyuannine but closest to Conchoraptor. It was discovered prior to November 2012.
References- Wei, Pu, Xu, Liu and Lu, 2013. A new oviraptorid dinosaur
(Theropoda: Oviraptorosauria) from the Late Cretaceous of Jiangxi Province,
Southern China. Acta Geologica Sinica (English Edition). 87(4), 899-904.
Wei, 2014. A new oviraptorid (Dinosauria: Theropoda) from the Upper
Cretaceous of Nankang city, Jiangxi Province. Masters thesis,
[university]. [pp]
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019. A new
paravian dinosaur from the Late Jurassic of North America supports a
late acquisition of avian flight. PeerJ. 7:e7247. DOI: 10.7717/peerj.7247
Oksoko Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
O. avarsan Funston, Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020
Early Maastrichtian, Late Cretaceous
Nemegt Formation, Mongolia
Holotype- (IGM
102/110a) (~45 kg, >1 year old juvenile) skull (150 mm), scleral
plates, mandible (120 mm),
hyoid (55.91 mm), axis, third-fourth cervical vertebrae, posterior
dorsal vertebrae, partial dorsal ribs, gastralia, synsacrum,
first-seventh
caudal vertebrae, proximal chevrons, sternal plates, sternal ribs,
radius (92 mm), ulna (92 mm),
scapholunare, semilunate carpals, two sesamoids, metacarpals I (23.95,
23.5 mm), phalanges I-1 (32.22, 30.1 mm), manual unguals I (31.2, 31.7
mm),
metacarpals II (40.09, 37.35 mm), phalanges II-2 (20.82, 20.96 mm),
manual unguals II (18.7, 17.7 mm), metacarpals III (28.25, 23.73 mm),
phalanx III-1 (9.07 mm), pubes, ischia, femora (235 mm),
tibiae (275 mm), fibulae, astragali (46 mm trans), calcaneum,
metatarsals I (26 mm), phalanges I-1, pedal unguals I, metatarsals II
(112 mm), phalanges II-1,
phalanges II-2, pedal unguals II, metatarsals III (127.5 mm), phalanges
III-1,
phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV
(119 mm),
phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal
unguals IV, metatarsal V (38 mm)
Paratypes- (IGM 102/11-A) (~33 kg, ~1 year old juvenile)
incomplete skull, posterior mandible, atlantal neurapophyses,
incomplete axis, third cervical vertebra, fourth cervical vertebra,
seventh to
tenth dorsal vertebrae, sacrum (146.9 mm), (caudal series ~435 mm)
first (20.6 mm) to twenty-seventh caudal
vertebrae, ten chevrons, ilia (216 mm), pubes (226 mm), incomplete
ischia (166 mm), femora (one
proximal; 210 mm), tibiae (one distal; 259 mm), fibulae (one distal),
astragali (40.2 mm trans), calcaneum, metatarsals I (20.3 mm),
phalanges I-1, pedal unguals I, distal
tarsals III, distal tarsals IV, metatarsals II (102 mm), phalanges
II-1,
phalanges II-2, pedal unguals II, metatarsals III (117 mm), phalanges
III-1,
phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV
(112 mm),
phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal
unguals IV
(IGM 102/11-B) (>33 kg) postorbital, quadratojugal, quadrate
(IGM 102/110b) (~45 kg, ~1 year old juvenile) skull (162 mm), scleral
plates, mandible, axis, third-fourth cervical vertebrae, dorsal ribs,
sacrum, first-fourth caudal
vertebrae, two chevrons, humerus (113 mm), radius (90 mm), ulna (94
mm), metacarpal I (24 mm), phalanx
I-1 (31.26 mm), manual ungual I (35.2 mm), metacarpal II (39.71 mm),
phalanx II-1 (20.6 mm), phalanx II-2 (15 mm), manual
ungual II (18 mm), phalanx III-1 (9.23 mm), pubes, ischia (169 mm),
femur (224 mm), tibiae (270 mm), fibula, astragalus (42.6 mm trans),
metatarsals I (30.2 mm), phalanges I-1, pedal unguals I, metatarsals II
(104 mm), phalanges II-1,
phalanges II-2, pedal unguals II, metatarsals III (118 mm), phalanges
III-1,
phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV
(110 mm),
phalanges IV-1, phalanges IV-2, phalanges IV-3, phalanges IV-4, pedal
unguals IV, metatarsal V (45 mm)
(IGM 102/110c) (~45 kg) three sacral vertebrae, first-seventh caudal
vertebrae, proximal chevrons, partial ilium, distal tibia,
astragalocalcaneum
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
(IGM 100/33; paratype of "Ingenia" yanshini)
(subadult) (atlas-axis 29.4 mm) atlas, axis, third cervical vertebra
(22.7 mm), fourth cervical vertebra (24.7 mm), fifth cervical vertebra
(26.6 mm), sixth cervical vertebra (27.1 mm), (dorsal series 258.9 mm)
first-tenth dorsal vertebrae, synsacrum (162.2 mm), (caudal series ~511
mm) twenty-three caudal vertebrae, chevrons, (scapulocoracoid 205 mm)
scapulae, coracoids, sternal plates (~60 mm), furcula, humeri (129.3,
128.7 mm), radii (94.7 mm), ulnae (99 mm), metacarpal I (23.8 mm),
phalanges I-1 (27.7 mm), manual unguals I (41.7 mm), metacarpal II
(41.7 mm), phalanges II-1 (17.5 mm), phalanges II-2 (14.5 mm), manual
ungual II (20 mm), metacarpal III (32 mm), pubes (210 mm), ischia (177
mm), femora (233 mm), tibiae (267 mm), fibula, astragalus (48 mm
trans), metatarsals I (29.2 mm), phalanges I-1, pedal unguals I,
metatarsals II (111.5 mm), phalanges II-1, phalanges II-2, pedal
unguals II, metatarsals III (124.5 mm), phalanges III-1 (34.3 mm),
phalanges III-2, phalanges III-3 (17.4 mm), pedal unguals III,
metatarsals IV (114.5 mm), phalanges IV-1, phalanges IV-2 (13.9 mm),
phalanges IV-3 (9.8 mm), phalanges IV-4 (9.2 mm), pedal unguals IV
(30.2 mm), metatarsals V (44 mm)
Early Maastrichtian, Late Cretaceous
Gurilin Tsav, Nemegt Formation, Mongolia
(IGM
102/12; 980808 GT SZK Oviraptr) (~74 kg, >5 year old adult) fourth
cervical vertebra (30.9 mm), fifth cervical vertebra (30.4 mm), sixth
cervical vertebra (30.5 mm), seventh cervical vertebra (28.9 mm),
eighth cervical vertebra (29.8 mm), ninth cervical vertebra (28.1 mm),
tenth cervical vertebra (27.8 mm), eleventh cervical neural arch,
(dorsal series 293.95 mm) first-tenth dorsal
vertebrae, four proximal dorsal ribs, (caudal series ~527 mm) fourth to
twenty-seventh caudal
vertebrae, eight chevrons, pygostyle, manual ungual I (62 mm),
incomplete ilium,
incomplete ischum, femur (280 mm), tibia (315 mm), fibulae (one
partial), astragalocalcaneum, distal
tarsals III, distal tarsal IV, metatarsal II (130 mm), metatarsals III
(146.4 mm), phalanx
III-1, metatarsal IV (134.2 mm)
Diagnosis- (after Funston et
al., 2020) apically thickened, dome-shaped cranial crest composed
equally of nasals and frontals; nasal recesses housed in a depression;
postorbital with dorsally directed frontal process; cervical vertebrae
with large epipophyses; functionally didactyl manus; accessory ridge of
brevis fossa of ilium; anteriorly curving pubis; large proximodorsal
process of distal tarsal IV.
Comments- Funston et al. (2017)
mention "three articulated juvenile skeletons,
representing a new species of oviraptorid theropod" confiscated from
poachers in December 14 2006. They state the taxon has "a domed
cranial
crest, a functionally didactyl hand, and a short tail." A brief
writeup of the presentation by Watson (2017) photographs the specimens,
noting they were found at Bugin Tsav and estimated at 45 kg.
Funston et al. (2018)
calls the taxon represented by all of these specimens the Guriliin Tsav
oviraptorid and stated it "will be
described elsewhere (Funston et al., submitted)", which was the
Funston et al. in review manuscript cited by Funston (2020) and
published as Funston et al. (2020). The latter paper states that
the precise locality of discovery within the Nemegt Formation is
unknown. The specimens are identified as IGM 102/110a, 110b and
110c in order of completeness.
Discovered in 1974, Barsbold (1981) first mentioned IGM 100/33 as a paratype of "Ingenia" yanshini,
then illustrated its furcula and sternal plates in 1983. Barsbold et
al. (2000) illustrated the nineteenth caudal vertebra and pelvis. The
latter appears more similar to the figure in Barsbold et al. (1990)
than to the yanshini
holotype, perhaps indicating the 1990 figure was based on IGM 100/33.
Funston et al. (2018) noted differences from the yanshini holotype such
as only six sacral vertebrae (instead of eight), non-hatchet-shaped
caudal transverse processes, unfused sterna, a more reduced metacarpal
III and no fibulotarsal contact. They also note that Barsbold (1983)
expanded the range of yanshini
to include Bugin Tsav (Nemegt) in
addition to Khermeen Tsav (Barun Goyot), and that "it is possible that
MPC-D 100/33 comes from Bugiin Tsav, but there is no detailed locality
data for the specimen." By Funston et al. (2020) it was stated to
be from Bugin Tsav without reservation. Funston et al. (2018)
stated "the sacral count, enlarged first manual
digit, reduced third metacarpal, and lack of fibulocalcaneal contact"
are shared between the then unnamed Oksoko and IGM 100/33, and the specimen was referred to that specimen when it was described in Funston et al. (2020).
IGM 102/12 (originally HMNS-IGM field number 980807 GT
SZK Oviraptr) is from Gurilin Tsav and was
said along with 980807 GT Coy Oviraptr to "belong to a single
species, which is similar to the partial skeleton of oviraptorosaurians
found at Bugin Tsav in 1994 by our party" (Suzuki and Watabe,
2000). However, 980807 GT Coy Oviraptr has a shallower postacetabular
process with greater invagination of the ischial peduncle, so is a
different taxon. IGM 102/12 was found in August 7 1998, initially photographed it in situ as
"Oviraptorosaurian skeleton
in Gurilin Tsav" by Suzuki and Watabe, then was later
photographed as figure 13 of Tsogtbaatar and Chinzorig (2010) once
prepared. In Funston et al. (2015) IGM 102/12 is listed as cf. Ingenia yanshini and given caudal and femoral measurements. Funston and Currie (2016) mentions IGM 102/12 as Ingenia
and state it has "downturned, rounded preacetabular blades." The
skeleton is photographed on Funston's blog (Funston, 2018 online). Funston et al. (2017)
mention it as "an unpoached, partial adult skeleton from Guriliin Tsav", and it was officially referred to their new genus Oksoko by Funston et al. (2020). The latter state "manual ungual I-2 was recovered in 2018 when the site was revisited."
IGM 102/11 was listed the same way as 102/11 by Funston et al. (2015)
and Funston and Currie (2016). Funston et al. (2016) figure its
distal metatarsal III as "Ingenia yanshiniBarsbold,
1981 (MPC-D 102/011), provenance unknown: confiscated
specimen." It was labeled "an unnamed oviraptorid
from the Nemegt Formation of Mongolia" by Funston and Currie (2018),
who figured its distal tibiotarsus. The skeleton is photographed
on Funston's blog (Funston, 2018 online), who assigned it to the same
species as IGM
102/12. This shows it is "the second poached skeleton [of] an
even younger individual" confiscated in December 14 2006 mentioned in
Funston et
al.'s (2017) abstract on Oksoko and was referred to the genus by
Funston et al. (2020). They note that "associated with it are the
postorbital, quadrate and quadratojugal of a slightly larger
individual" which they call IGM 102/11-B.
Funston
(2019) describes all of these specimens in detail in his thesis, which
also includes more figures than the published description. Funston et
al. (2020) added Oksoko to the Maryanska et al.-based oviraptorosaur analysis to find it a heyuannine sister to Jiangxisaurus, then Banji.
References-
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological
Expedition Transactions. 15, 28-39.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy,
Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120.
Barsbold, Maryanska and Osmolska, 1990. Oviraptorosauria. In Weishampel, Dodson
and Osmolska (eds.). The Dinosauria. University of California Press. 249-258.
Barsbold, Currie, Myhrvold, Osmolska, Tsogtbaatar and Watabe, 2000. A pygostyle
from a non-avian theropod. Nature. 403, 155-156.
Suzuki and Watabe, 2000. Report on the Japan-Mongolia joint paleontological
expedition to the Gobi desert, 1998. Hayashibara Museum of Natural
Sciences Research Bulletin. 1, 83-98.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian
Paleontological Center: 2002-2008. Hayashibara Museum of Natural
Sciences Research Bulletin. 3, 155-166.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied
caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of
Alberta, Canada. Cretaceous Research. 54, 179-187.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Funston, Currie and Burns, 2016 (online 2015). New elmisaurine specimens from North America
and their relationship to the Mongolian Elmisaurus rarus. Acta Palaeontologica
Polonica. 61(1), 159-173.
Funston, Currie and Tsogtbaatar, 2017. A new oviraptorid (Dinosauria:
Theropoda) provides a rare glimpse into social behaviour in dinosaurs.
Journal of Vertebrate Paleontology. Program and Abstracts 2017, 116.
Watson, 2017. Roosting dinosaurs are a fossil first. Nature. 548, 510.
Funston, 2018 online. https://gregfunston.com/2018/09/24/mongolia-monday-travel-log-5/
Funston and Currie, 2018. A small caenagnathid tibia from the Horseshoe
Canyon Formation (Maastrichtian): Implications for growth and lifestyle
in oviraptorosaurs. Cretaceous Research. 92, 220-230.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy,
diversity and ecology in the Nemegt basin. Palaeogeography,
Palaeoclimatology, Palaeoecology. 494, 101-120.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Funston, 2020. Caenagnathids of the Dinosaur Park Formation (Campanian)
of Alberta, Canada: Anatomy, osteohistology, taxonomy, and evolution.
Vertebrate Anatomy Morphology Palaeontology. 8, 105-153.
Funston,
Chinzorig, Tsogtbaatar, Kobayashi, Sullivan and Currie, 2020. A new
two-fingered dinosaur sheds light on the radiation of Oviraptorosauria.
Royal Society Open Science. 7: 201184.
Machairasaurus Longrich,
Currie and Dong, 2010
M. leptonychus Longrich, Currie and Dong, 2010
Late Campanian, Late Cretaceous
Wulansuhai Formation (= Bayan Mandahu Formation), Inner Mongolia, China
Holotype- (IVPP V15979) distal radius, distal ulna, scapholunare, semilunate
carpal, metacarpal I (24 mm), phalanx I-1 (30 mm), manual ungual I (29 mm),
metacarpal II (38 mm), phalanx II-1 (21 mm), phalanx II-2 (22 mm), manual ungual
II (24 mm), metacarpal III (34 mm), phalanx III-1 (12 mm), phalanx III-2, phalanx
III-3, manual ungual III, manual fragments, several pedal phalangeal fragments,
pedal ungual I, pedal ungual IV
Referred- (IVPP V15980) dorsal ribs, caudal vertebrae, chevrons, metacarpal
I (32 mm), phalanx I-1 (79 mm), manual ungual I (50 mm), phalanx II-1 (60 mm),
phalanx II-2 (66 mm), phalanx III-1 (38 mm), partial tibia, fragmentary metatarsal
II, phalanx III-3 (28 mm), fragmentary metatarsal IV, phalanx IV-2 (27 mm) (Longrich,
Currie and Dong, 2010)
Diagnosis- (after Longrich et al., 2010) manual unguals I–III elongate
and bladelike in lateral view (length of claw approximately 400 per cent the
height of the proximal articular surface).
Comments- Discovered in 1988 and 1990, the type specimens of Machairasaurus
leptonychus were described by Longrich et al. in 2010 as a new taxon. They
used a version of Osmolska et al.'s oviraptorosaur matrix which placed Machairasaurus
in Heyuanninae, more derived than Khaan and Conchoraptor
but more basal than Heyuannia.
This holds for future iterations of that matrix as well, such as
Funston and Currie (2016), but after adding taxa to Hartman et
al.'s matrix the genus falls out closer to Conchoraptor. Fanti et
al. (1012) provide measurements.
References- Longrich, Currie and Dong, 2010. A new oviraptorid (Dinosauria:
Theropoda) from the Upper Cretaceous of Bayan Mandahu, Inner Mongolia. Palaeontology.
53(5), 945-960.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Funston and Currie, 2016. A new caenagnathid (Dinosauria: Oviraptorosauria)
from the Horseshoe Canyon Formation of Alberta, Canada, and a reevaluation of
the relationships of Caenagnathidae. Journal of Vertebrate Paleontology. 36(4),
e1160910.
Khaan Clark, Norell and Barsbold,
2001
K. mckennai Clark, Norell and Barsbold, 2001
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadochta Formation, Mongolia
Holotype-
(IGM 100/1127) (female?) skull (~127 mm), mandible (108.4 mm), hyoids, twelve
cervical vertebrae, cervical ribs, several dorsal vertebrae, dorsal
ribs, uncinate process, gastralia, sacrum, twenty-seven caudal
vertebrae (first caudal 18.3 mm), chevrons, scapulacoracoids (154.1 mm;
scap 124.7 mm), furcula, sternal plates (58.8 mm), humeri (110.5, 108.6
mm), radii (~102 mm), ulnae (96.5 mm), scapholunare, semilunate carpal,
metacarpal I, phalanx I-1, manual ungual I, metacarpal II (47.3 mm),
phalanx II-1, phalanx II-2, manual ungual II, metacarpal III (44.4 mm),
phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ilia
(182 mm), pubis, ischium, femur (185 mm), tibia (212 mm), fibula (201
mm), metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal
ungual III, metatarsal IV (~95 mm), phalanx IV-1, phalanx IV-2, phalanx
IV-3, phalanx IV-4, pedal ungual IV
Paratypes- (IGM 100/973) skull (118.8 mm), mandibles (104 mm), hyoids,
atlas, axis, ilia (193.5, 187.5 mm), pubis (156.8 mm), ischium (133.1 mm), femora
(188.8, 199 mm), tibiae (221.4, 224 mm), fibula, astragali, calcanea, distal
tarsal III, metatarsals I (26.7, ~23 mm), phalanges I-1, pedal ungual I, metatarsals
II (82.8, 89.9 mm), phalanges II-1, phalanges II-2, pedal unguals II, metatarsals
III (97.7, 98.8 mm), phalanges III-1, phalanges III-2, phalanges III-3, pedal
unguals III, metatarsals IV (87.3, 93.9 mm), phalanges IV-1, phalanges IV-2,
phalanges IV-3, phalanges IV-4, pedal unguals IV, metatarsal V (35.8 mm)
(IGM 100/1002) (male?) incomplete skull (113.8 mm), mandible (101.6 mm), hyoids, twelve
cervical vertebrae, cervical ribs, ten dorsal vertebrae, dorsal ribs, gastralia,
sacrum, seven caudal vertebrae, four chevrons, scapulocoracoids (167 mm; scap
138.6 mm), furcula, sternal plates (55.1 mm), sternal ribs, humerus (117.5 mm),
radii (95.3 mm), ulnae (~101.6 mm), scapholunare, semilunate carpal, metacarpal I
(24.7 mm), phalanx I-1 (38 mm), manual ungual I (44.5 mm), metacarpal II (50.4
mm), phalanx II-1 (26 mm), phalanx II-2 (28.3 mm), manual ungual II (37 mm),
metacarpal III (47.6 mm), phalanx III-1 (16.2 mm), phalanx III-2, phalanx III-3,
manual ungual III, ilium (186 mm), pubis (~167 mm), ischium (~124 mm), femur
(196 mm), tibia (232 mm), fibula (214 mm), astragalus, distal tarsal IV, metatarsal
I (23.3 mm), phalanx I-1, pedal ungual I, metatarsals II (91.2 mm), phalanges
II-1, phalanges II-2, pedal unguals II, metatarsals III (105.6 mm), phalanges
III-1, phalanges III-2, phalanges III-3 (15.3 mm), pedal unguals III, metatarsals
IV (98.3 mm), phalanges IV-1, phalanges IV-2 (14 mm), phalanges IV-3 (11.5 mm),
phalanges IV-4 (10.7 mm), pedal unguals IV (31 mm), metatarsal V (35.9 mm)
Referred- (IGM 100/3616) multiple individuals (Norell et al., 2018)
Late Cretaceous?
Mongolia?
?(IGM 100/1253) partial skull (Balanoff, 2011)
Diagnosis- (after Clark et al.,
2001) metacarpal III is not expanded proximally and does not contact
the distal carpals. Differs from Conchoraptor in that the long axis of
the external naris is more horizontally oriented and the nasals are
fused.
Comments- IGM 100/973 was discovered in 1993 and first photographed in
Novacek et al. (1994). It was photographed in Dashzeveg et al. (1995) as cf.
Ingenia, and labeled Ingenia yanshini in Webster (1996). IGM 100/1127
and 100/1002 were discovered in 1995, and the latter was photographed in Morell
(1997) and labeled Oviraptor. Clark et al. (2001) made these their new
taxon Khaan mckennai.
IGM 100/1002 and 100/1127 were found in close proximity to each other,
and have been called "Romeo and Juliet." Balanoff and Norell
(2012)
recently described the taxon in depth and Balanoff et al. (2018)
described the endocranium, both based on Balanoff's (2011)
thesis. Person et al. (2015) noted the proximal chevrons of IGM
100/1002 were more elongate and distally expanded unlike 100/1127,
which they considered plausibly sexual dimorphism with 100/1127 being
the female.
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). Balanoff (2011) included the partial skull IGM
100/1253 in her oviraptorosaur analysis and recovered it as either
sister to Khaan or to Citipati plus Gigantoraptor, but considered it most likely a Khaan specimen.
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 and Perle, 1995. Extraordinary 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.
Dyke and Norell, 2005. Caudipteryx as a non-avialan theropod rather than
a flightless bird. Acta Palaeontologica Polonica. 50(1), 101-116.
Balanoff and Norell, 2009. Adult morphology and variation within the oviraptorid
Khaan mckennai (Theropoda: Oviraptorosauria). Journal of Vertebrate Paleontology.
29(3), 57A.
Balanoff, 2011. Oviraptorosauria: Morphology, phylogeny, and endocranial evolution.
PhD thesis. Columbia University. 522 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria:
Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Persons, Funston, Currie and Norell, 2015. A possible instance of
sexual dimorphism in the tails of two oviraptorosaur dinosaurs.
Scientific Reports. 5:9472.
Balanoff, Norell, Hogan and Bever, 2018. The endocranial cavity of
oviraptorosaur dinosaurs and the increasingly complex, deep history of
the avian brain. Brain, Behavior and Evolution. 91, 125-135.
Norell, Balanoff, Barta and Erickson, 2018. A second specimen of Citipati osmolskae associated with a nest of eggs from Ukhaa Tolgod, Omnogov Aimag, Mongolia. American Museum Novitates. 3899, 44 pp.
Heyuannia Lu, 2002
= "Ingenia" Barsbold, 1981
preoccupied Gerlach, 1957
= Ajancingenia Easter, 2013a
Comments- The genus Ingenia is preoccupied by the tripyloidid nematode Ingenia mirabilis, as noted
by Taylor (DML, 2004). Barsbold is aware of the situation, but Easter (2013a)
did not consult him, proposing his own replacement name Ajancingenia in a publication which
largely consisted of copied and reworded information from this website used
without my permission. While the initial objection to citing The Theropod Database
was editorial, Easter did not object to this, lied about consulting with Barsbold
in an online forum and took credit for my work in that same forum. Based on
my objection, an erratum was published (Easter, 2013b), and while his name remains
technically valid according to the ICZN, I do not support its use due to Easter's
unprofessional and unethical actions. Funston et al. (2018) proposed a solution where Ajancingenia was synonymized with Heyuannia, its sister taxon in most analyses. Thus "Ingenia" yanshini becomes Heyuannia yanshini, which is accepted here.
References- Gerlach, 1957. Die Nematodenfauna des Sandstrandes an der K�ste von
Mittelbrasilien (Brasilianische Meerse-Nematoden IV). Mitteilungen aus
dem Zoologischen Museum in Berlin. 33(2), 411-459.
Barsbold, 1981. Toothless dinosaurs of Mongolia. Joint Soviet-Mongolian Paleontological
Expedition Transactions. 15, 28-39.
Lu, 2002. A new oviraptorosaurid (Theropoda: Oviraptorosauria) from the Late
Cretaceous of southern China. Journal of Vertebrate Paleontology. 22(4), 871-875.
Taylor, DML 2004. https://web.archive.org/web/20200928000123/http://dml.cmnh.org/2004Sep/msg00022.html
Easter, 2013a. A new name for the oviraptorid dinosaur "Ingenia"
yanshini (Barsbold, 1981; preoccupied by Gerlach, 1957). Zootaxa. 3737(2),
184-190.
Easter, 2013b. Erratum. Zootaxa. 3750(1), 100.
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy,
diversity and ecology in the Nemegt Basin. Palaeogeography,
Palaeoclimatology, Palaeoecology. 494, 101-120.
H. huangi Lu, 2002
= Citipati huangi (Lu, 2002) Paul, 2010
= Conchoraptor huangi (Lu, 2002) Paul, 2016
Maastrichtian, Late Cretaceous
Zhutian (=Dalangshan) Formation, Guangdong, China
Holotype- (HYMV1-1) partial skull (~150 mm), mandible (~150 mm), hyoid (50
mm), thirteen cervical vertebrae (520 mm; fourth 30 mm; sixth ~45 mm), twelve
dorsal vertebrae (320 mm), dorsal ribs (30-160 mm), uncnate processes, first
sacral vertebra (27 mm), second sacral vertebra (30 mm), third sacral vertebra
(30 mm), fourth sacral vertebra (~30 mm), fifth sacral vertebra (~30 mm), sixth
sacral vertebra (30 mm), seventh sacral vertebra (25 mm), eighth sacral vertebra
(25 mm), first caudal vertebra (26 mm), second caudal vertebra (26 mm), third
caudal vertebra (26 mm), fourth caudal vertebra (26 mm), fifth caudal vertebra
(26 mm), sixth caudal vertebra (25 mm), seventh caudal vertebra (25 mm), first
chevron (60 mm), second chevron (100 mm), third chevron (100 mm), fourth chevron
(90 mm), fifth chevron, sixth chevron, proximal scapula, coracoid fragment,
furcula, partial ilia (260 mm), pubes (260 mm), ischia (195 mm), femora (255
mm), tibiae (320 mm), proximal fibula, astragalus, calcaneum, metatarsal I (30
mm), phalanx I-1 (19 mm), pedal ungual I (30 mm), metatarsal II (110 mm), phalanx
II-1 (35 mm), phalanx II-2 (20 mm), pedal ungual II (45 mm), metatarsal III
(135 mm), phalanx III-1 (37 mm), phalanx III-2 (27 mm), metatarsal IV (120 mm),
phalanx IV-1 (25 mm), phalanx IV-2 (20 mm), phalanx IV-3 (10 mm), phalanx IV-4
(10 mm)
Paratypes- (HYMV1-2) two dorsal ribs, gastralia, scapula (~175 mm), incomplete
coracoid, furcula, partial sternum, sternal ribs, humerus (130 mm), radius (110
mm), ulna (127 mm), scapholunare, semilunate carpal, metacarpal I (32 mm), phalanx
I-1, proximal metacarpal II, partial phalanx II-1, phalanx II-2, proximal metacarpal
III
(HYMV1-3) phalanx I-1, manual ungual I, distal metacarpal II, phalanx II-1,
phalanx II-2, manual ungual II, distal metacarpal III, phalanx III-1
(HYMV1-4) partial pubis, distal femur, proximal tibia, proximal fibula
(HYMV1-5) incomplete manus
Referred- (HYMV1-6) partial skull, mandibular fragment, posterior cervical
vertebrae, anterior dorsal vertebrae, dorsal ribs, uncinate processes, gastralia,
last ten caudal vertebrae, partial scapulae (~165 mm), coracoids (50 mm; one
partial), incomplete furcula, partial sternum, three partial sternal ribs, humeri
(one proximal), partial radius, partial ulna, proximal femur, tibia (Lu, 2004)
(HYMV1-7) last three caudal vertebrae, distal metacarpal I, phalanx I-1, manual
ungual I, metacarpal II fragment (Lu, 2004)
(HYMV2-1) humerus (105 mm), incomplete radius, ulna (103 mm), scapholunare,
semilunate carpal, pisiform, metacarpal I (29 mm), phalanx I-1 (32 mm), manual ungual
I (35 mm), metacarpal II (57 mm), phalanx II-1 (17 mm), phalanx II-2 (16 mm),
manual ungual II (21 mm), metacarpal III (52 mm) (Lu, 2004)
(HYMV2-2) partial humerus, partial radius, partial ulna (Lu, 2004)
(HYMV2-3) femur, tibia, fibula, astragalus (Lu, 2004)
(HYMV2-4) pectoral girdle (Lu, 2004)
(HYMV2-5) pelvis, partial hindlimb including metatarsal V fragment (Lu, 2004)
(HYMV2-6) ten mid caudal vertebrae (Lu, 2004)
(HYMV2-7) partial pelvis (Lu, 2004)
(HYMV2-8) radius, ulna, metacarpal I (Lu, 2004)
? bones and eggs (Qiu and Huang, 2001)
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav of Barun Goyot Formation, 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 (Gaston Design, 2019 online)
Diagnosis-
(after Lu, 2002) quadratojugal articular surface of quadrate
groove-like; quadrate diverticulum enters bone anterolaterally;
increased cervical vertebral count; pneumatic foramina present on
cervical neural arches and ribs; decreased dorsal vertebral count;
increased sacral vertebral count; proximal end of metacarpal I wraps
around metacarpal II in ventral view; pubis as long as ilium.
(after Lu, 2004) angle
of the fused scapula and coracoid approximately 145�; ratio of coracoid
length to the scapular length approximately 0.35; pubis as long as ischium; ratio of femur length to tibia length
0.8.
Comments- First discovered in the summer of 1999, Lu (2002) briefly described Heyuannia, then described
it in more depth including additional specimens in his 2004 thesis (published as a book
in 2005). The forelimb and pectoral girdle were described further by Lu et al.
(2005), though they incorrectly illustrate HYMV1-6 as 1-4. 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 cast and widely distributed as Conchoraptor ("Conchoraptor Juvenile in 3-D matrix block" on Gaston Design),
but its manual proportions more nearly resemble Heyuannia. This it is
provisionally referred to Heyuannia huangi here, though it is supposedly from
the Red Beds of Khermeen Tsav like Conchoraptor and Heyuannia yanshini.
Cheng et al. (2008) referred two Macroolithus yaotunensis eggs with embryos
(Chimei Museum 41 and NMNS-0015726-F02-embryo-01) from the Nanxiong Group
of Jiangxi to Heyuannia huangi "or an oviraptorosaurian of similar kind"
based on subarctometatarsaly. Yet this is true in most other oviraptorids as
well except for Gigantoraptor, "Tongtianlong" and Heyuannia yanshini. Wiemann et al. (2015a, b) used Cheng
et al.'s paper to justify assigning all M. yaotunensis eggs to Heyuannia,
including specimens from the Hugang Formation of Henan, the Yuanpu or Pingling
Formation of Guangdong and the Nanxiong Group of Jiangxi.
References- Qiu and Huang, 2001. Dinosaur fossils from the Heyuan Basin
in Guangdong Province, China. In Deng and Wang (eds.). Proceedings of the Eighth
Annual Meeting of the Chinese Society of Vertebrate Paleontology. 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. PhD thesis, Southern Methodist University. 249 pp.
Lu, 2005. Oviraptorid dinosaurs from Southern China. Geological Publishing House,
Beijing. 200 pages + 8 plates.
Lu, Huang and Qiu, 2005. The pectoral girdle and the forelimb of Heyuannia
(Dinosauria: Oviraptorosauria). In Carpenter (ed.). The Carnivorous Dinosaurs.
Indiana University Press. 256-273.
Cheng, Ji, Wu and Shan, 2008. Oviraptorosaurian eggs (Dinosauria) with embryonic
skeletons discovered for the first time in China. Acta Geologica Sinica. 82(6),
1089-1094.
Paul, 2010. The Princeton Field Guide to Dinosaurs. Princeton University Press.
320 pp.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, M�ller and Sander,
2015a. The blue-green eggs of dinosaurs: How fossil metabolites provide insights
into the evolution of bird reproduction. PeerJ PrePrints. DOI: 10.7287/peerj.preprints.1080v1
Wiemann, Yang and Sander, 2015b. The colorful eggs of dinosaurs: How fossil
metabolites reveal nesting behavior. Journal of Vertebrate Paleontology. Program
and Abstracts 2015, 237.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd Edition. Princeton University Press.
360 pp.
Wiemann, Yang, Sander, Schneider, Engeser, Kath-Schorr, M�ller and
Sander, 2017. Dinosaur origin of egg color: Oviraptors laid blue-green
eggs. PeerJ. 5:e3706.
Gaston Design, 2019 online. https://www.gastondesign.com/product/817/
H. yanshini (Barsbold, 1981)
Funston, Mendonca, Currie and Barsbold, 2018
= "Ingenia" yanshini Barsbold, 1981
= Oviraptor yanshini (Barsbold, 1981) Paul, 1988
= Ajancingenia yanshini (Barsbold, 1981) Easter, 2013
= Conchoraptor yanshini (Barsbold, 1981) Paul, 2016
Late Campanian(?), Late Cretaceous
Red Beds of Khermeen Tsav of Barun Goyot Formation, Mongolia
Holotype-
(IGM 100/30) parietal (lost), braincase (lost), posterior mandible,
atlas, axis, third cervical vertebra, fourth cervical vertebra with
fused ribs, fifth cervical vertebra, sixth cervical vertebra, seventh
cervical vertebra, eighth cervical vertebra, ninth cervical vertebra,
fourteen dorsal ribs, sacrum, thirty caudal vertebrae (first caudal
25.3 mm), twelve chevrons, scapulae (~145 mm), coracoids, furcula,
sternum (~72 mm), humeri (141 mm), ulnae (116 mm), carpometacarpus (mcI
31.6, mcII 50.3, mcIII 46.5 mm), manual ungual I (49.2 mm), phalanx II-1
(21.3 mm), phalanx II-2 (19.8 mm), manual ungual II (25.8 mm), phalanx
III-1 (12.8 mm), phalanx III-2 (11 mm), tibiae (290 mm), fibulae,
astragalocalcanea, distal tarsal III, metatarsal I, metatarsal II,
metatarsal III (125 mm), metatarsal IV
Paratypes-
(IGM 100/31) frontal fragment (lost), parietals (lost), mandible (105
mm), postcrania including eight sacral vertebrae (?- see comments),
radii (102.6 mm), metacarpal I (30.3 mm), phalanx I-1 (33.5 mm),
manual ungual I (46.9 mm), metacarpal II (46.2 mm), phalanx II-1 (18.7
mm), phalanx II-2 (16.2 mm), manual ungual II (25.7 mm), metacarpal III
(44 mm), phalanx III-1 (7.2 mm), phalanx III-2 (8 mm), phalanx III-3 (3
mm), ilia (228 mm), pubes (244 mm), ischia, femora (241 mm), tibia,
fibula, metatarsal I, phalanx I-1, pedal ungual I, metatarsal II,
phalanges II-1, phalanges II-2, pedal unguals II, metatarsal III (119.7
mm), phalanges III-1, phalanges III-2, phalanges III-3 (18.7 mm), pedal
unguals III, metatarsal IV, phalanges IV-1, phalanges IV-2 (14.5 mm),
phalanges IV-3 (11.5 mm), phalanges IV-4 (10.6 mm), pedal unguals IV
(33.3 mm)
(IGM 100/32) mandible (~120 mm), postcrania including cervical vertebrae, twenty-seven
caudal vertebrae, partial scapula, humerus (140 mm), radius (110.5 mm), ulna
(115.2 mm), semilunate carpal, phalanx I-1 (34.9 mm), manual ungual I (48.9
mm), metacarpal II (50.5 mm), phalanx II-1 (21.9 mm), phalanx II-2 (18.9 mm),
manual ungual II (25.7 mm), metacarpal III (41.6 mm), phalanx III-1 (12.6 mm),
ilium (242 mm), pubis (220 mm), femur (254 mm), tibia (294 mm), metatarsus (132
mm), phalanx III-1 (33.1 mm), phalanx III-3 (18.9 mm), phalanx IV-2 (16 mm),
phalanx IV-3 (1.6 mm), phalanx IV-4 (11.2 mm), pedal ungual IV (32 mm)
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Referred-
?(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)
Late Cretaceous(?)
Mongolia(?)
(BHI coll.) skull, mandible (Fanti et al., 2012)
(FDPM-V6240) skull, skeleton (Goto, Ichishima and Ji, 2005)
(IGM 100/34) material including scapula, partial humerus (69 mm), ilium (119
mm), femur (135 mm), tibia (162 mm), metatarsus (75.8 mm), phalanx III-1 (20.6
mm), phalanx III-3 (11.8 mm), phalanx IV-2 (10.8 mm), phalanx IV-3 (6.4 mm),
phalanx IV-4 (6.2 mm), pedal ungual IV (18 mm) (Snively, 2000)
(IGM 100/35) material including femur (135 mm), partial tibia and phalanx III-1
(20.4 mm) (Maryanska et al., 2002)
?(IGM 100/80-1; = IGM 100/20?; see comments) material including skull (115.6 mm), eleven cervical vertebrae
(Lu, 2004)
(IGM 110/02) (Fanti et al., 2012)
(IGM 110/03) material including humerus (73.5 mm), radius (54.7 mm), ulna (55.3
mm), metacarpal I (15.4 mm), phalanx I-1 (16.8 mm), manual ungual I (19 mm),
metacarpal II (23.8 mm), phalanx II-1 (9.5 mm) (Fanti et al., 2012)
(IGM PJC2002.17) skull, mandible (Fanti et al., 2012)
?(IGM coll.) material including skull (147.3 mm), mandible (120 mm), femur (238
mm), tibia (275 mm), metatarsal III (127.5 mm) and phalanx III-1 (34.5 mm) (Lu
et al., 2013)
?(IGM coll.) material including skull (162 mm), humerus (113 mm), radius (90
mm), ulna (98 mm), metacarpal II (40.6 mm), femur (220 mm), tibia (265 mm),
metatarsal III (118 mm) and phalanx III-1 (34.3 mm) (Lu et al., 2013)
Diagnosis- (after Funston et al., 2018) low ridge on medial side of surangular; eight sacral vertebrae (also in Nemegtomaia); chevrons with distal bulb; ulna with large lateral distal process; manual ungual I longer than metacarpal I.
Other diagnoses- (after Barsbold, 1981) highly reduced, slightly curved and compressed unguals of the second and third fingers.
Comments- 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) and is probably IGM 100/31 based on the scale bar and measurement in Fanti et al. (2012).
"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 (IGM 100/31-33). Barsbold (1983) didn't expand on the
description, but did illustrate the holotype's braincase, furcula and
sternum. Barsbold (1986) later illustrated the humerus, femur and
metatarsus of the holotype. Barsbold et al. (1990) added some further
information and illustrated the humerus in different views, radius,
ulna, complete manus, pelvis and complete hindlimb missing only
metatarsal V, pedal phalanx II-2 and digit I. These may all be from the
holotype as well, but the specimen numbers were not listed. Psihoyos
(1994) includes a photograph later revealed to be a composite of the
holotype, IGM 100/31 and 100/32 (Funston et al., 2018), which is
largely complete except for the dorsal series, though the skull of the Conchoraptor holotype is mounted on
it. This mount was displayed at the Nakasato Dinosaur Center with a
mandible that is neither 100/30, 100/31 or 100/20 (previously online
photos). Barsbold et al. (2000) illustrated the twenty-seventh
caudal vertebra of IGM 100/32. Lu (2004) noted many anatomical details,
including noting some of the elements preserved in paratype specimens.
Osmolska et al. (2004) illustrated a scapulocoracoid that is different
from the holotype 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 (Nemegt instead of Barun Goyot).
Funston et al. (2018) is the first published source to identify the IGM
mount as a composite of IGM 100/30, 100/31 and 100/32 and provides the
sources of all elements. They state the braincase of 100/30 and
skull roof of 100/31 are lost. Funston et al. also say "There are
eight sacral vertebrae (labeled MPCD 100/30), which appear to be fused
with the ilia (labeled MPC-D 100/ 31). It is possible that these
elements were artificially connected during the creation of the mount,
or that the labels are incorrect." This brings Lu's (2004)
statement IGM 100/31 has eight sacral vertebrae into question, as it
may have been based on this mounted specimen. Fanti et al. (2012)
and Currie et al. (2016) provide measurements of numerous elements.
Snively (2000) studied and illustrated the metatarsi of IGM 100/32 and 100/34 as Ingenia yanshini based on photos and measurements from Currie (PJC 1998 II).
Maryanska et al. (2002) lists the latter specimen and IGM 100/35 as I. yanshini, while Lu (2004)
notes cervical and cranial characters of IGM 100/80-1. The latter is notable
as previously the only yanshini cranial material thought to
exist was the braincase of the holotype (and possibly different skull roof of IGM 100/31), whereas Lu's statements imply much
of the skull is present in IGM 100/80-1. Confusingly, Fanti et al. list IGM
100/80-D as a skull and the holotype of Conchoraptor, and Funston et al, (2018) state the mounted Conchoraptor holotype skull at the IGM is labeled 100/80-1. So perhaps Lu was mistaken in assigning the skull to yanshini, but he also stated IGM 100/80-1 has eleven cervical vertebrae when the IGM mount only has nine cervicals from the yanshini holotype. 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 Nomingia, Gobiraptor, Rinchenia or Nemegtomaia
which are present in that formation. As with Conchoraptor,
no evidence has ever been published defending the placement of paratypes or
referred material in yanshini
and several specimens have been removed (see below). The
measurements and other referred specimens are all based on measurement
tables in Fanti et al. (2012), except two IGM specimens Lu et al.
(2013) call c.f. Ingenia Confiscated 1 and 2. PJC.2001.2 was referred to cf. Ingenia sp.
by Currie (2002).
Not "Ingenia"- "Ingenia" yanshini
paratype IGM 100/33 was illustrated by Barsbold (1983) and Barsbold et
al. (2000), but was removed by Funston et al. (2018) due to having six
sacrals, unfused sterna and no fibulotarsal contact among other
characters. Paul (1988) illustrated ZPAL MgD-I/95 as Oviraptor yanshini, a combination no other workers follow as it would
result in placing most oviraptorids in Oviraptor given recent phylogenies.
ZPAL MgD-I/95 is currently assigned to Conchoraptor. 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 (Witmer, online 2012).
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 more probably Heyuannia huangi
instead of H. yanshini. IGM 110/11 and 110/12 were listed by Funston et al. (2015) as c.f. Ingenia yanshini, but have been reidentified as the unnamed Guriliin Tsav oviraptorid (Funston, 2018 online).
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.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Trudy,
Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 19, 1-120.
Barsbold, 1986. The predatory dinosaurs - Oviraptors. In Vorobyeva
(ed.). Herpetologische Untersuchungen in Der Mongolischen
Volksrepublik. Academia Nauk SSSR. 210-223.
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. 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. 1076 pp.
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.
Masters Thesis, Institute of Vertebrate Paleontology and Paleoanthropology. 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. Masters Thesis, University of Calgary. 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. 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. 59-63.
Currie, 2002. Report on fieldwork in Mongolia, September 2001. Alberta Palaeontological
Society, sixth annual symposium. 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. 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. 33.
Lu, 2004. Oviraptorid dinosaurs from southern China. PhD thesis, Southern Methodist University. 249 pp.
Osmolska, 2004. Evidence on relation of brain to endocranial cavity in oviraptorid
dinosaurs. Acta Palaeontologica Polonica. 49(2), 321-324.
Weishampel, Barrett, Coria, Le Loeuff, Xu, Zhao, Sahni, Gomani and Noto, 2004.
Dinosaur Distribution. In Weishampel, Dodson and Osmolska (eds.). The Dinosauria Second
Edition. University of California Press. 861 pp.
Goto,
Ichishima and Ji, 2005. The Flying Dinosaurs. Catalog of an exhibition
held at Fukui Kenritsu Kyōryū Hakubutsukan, July 15-November 3, 2005;
also to be held at Asahikawa Kagakukan and at Hamamatsu Kagakukan in
2006. Fukui Kenritsu Kyōryū Hakubutsukan. 118 pp.
Balanoff and Norell, 2012. Osteology of Khaan mckennai (Oviraptorosauria:
Theropoda). Bulletin of the American Museum of Natural History. 372, 1-77.
Fanti, Currie and Badamgarav, 2012. New specimens of Nemegtomaia from
the Baruungoyot and Nemegt Formations (Late Cretaceous) of Mongolia. PLoS ONE.
7(2), e31330.
Witmer, 2012 online. https://people.ohio.edu/witmerl/collections/Theropods/Ingenia.htm
Easter, 2013. A new name for the oviraptorid dinosaur "Ingenia"
yanshini (Barsbold, 1981; preoccupied by Gerlach, 1957). Zootaxa. 3737(2),
184-190.
Lu, Currie, Xu, Zhang, Pu and Jia, 2013. Chicken-sized oviraptorid dinosaurs
from central China and their ontogenetic implications. Naturwissenschaften.
100(2), 165-175.
Funston, Persons, Bradley and Currie, 2015. New material of the large-bodied
caenagnathid Caenagnathus collinsi from the Dinosaur Park Formation of
Alberta, Canada. Cretaceous Research. 54, 179-187.
Currie, Funston and Osm�lska, 2016 (online 2015). New specimens of the crested theropod
dinosaur Elmisaurus rarus from Mongolia. Acta Palaeontologica Polonica.
61(1), 143-157.
Paul, 2016. The Princeton Field Guide to Dinosaurs 2nd Edition. Princeton University Press.
360 pp.
Funston, 2018 online. https://gregfunston.com/2018/09/24/mongolia-monday-travel-log-5/
Funston, Mendonca, Currie and Barsbold, 2018 (online 2017). Oviraptorosaur anatomy,
diversity and ecology in the Nemegt Basin. Palaeogeography,
Palaeoclimatology, Palaeoecology. 494, 101-120.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Gobiraptor Lee, Lee, Chinsamy, Lu, Barsbold and Tsogtbaatar, 2019
= "Gobiraptor" Lee, 2018
G. minutus Lee, Lee, Chinsamy, Lu, Barsbold and Tsogtbaatar, 2019
= "Gobiraptor minutus" Lee, 2018
Early Maastrichtian, Late Cretaceous
Altan Uul III, Nemegt Formation, Mongolia
Holotype- (IGM 102/111) (juvenile) partial skull, incomplete
mandibles, last sacral vertebra (25.5 mm), first caudal vertebra (23.1
mm), second caudal vertebra (24.3 mm), seven proximal caudal vertebrae
(22.8, 24.2, 23.6, 23.6, 23.2, 21.4 mm), proximal chevron fragments,
proximal scapula, coracoid fragment(?), proximal humerus, ilia (one
partial, one incomplete), incomplete pubis, partial ischium, femora
(195.7 mm), distal tarsal III, distal tarsal IV, metatarsal I (28.8
mm), phalanx I-1 (23.1 mm), pedal ungual I, metatarsal II (108.2
mm), metatarsal III (123 mm), phalanx III-1 (36.3 mm), phalanx III-2
(29 mm), proximal phalanx III-3, metatarsal IV (112.4 mm), phalanx IV-1
(20.5 mm), phalanx IV-2 (18.6 mm), phalanx IV-3 (16.2 mm), phalanx IV-4
(12.2 mm), pedal ungual IV, metatarsal V (4.2 mm)
Diagnosis- (after Lee et al.,
2019) flat articular surface for quadratojugal on quadrate;
anteroposteriorly elongate dentary anterior to external mandibular
fenestra; extremely thickened anterodorsal end of mandibular
symphysis with posterior expansion of its dorsal surface; rudimentary
lingual triturating shelf on dentary bearing small occlusal foramina;
weakly developed lingual ridge on each lingual shelf; anterior end of
coronoid wedging into ventral surface of dorsal dentary process.
Comments- Discovered in 2008,
Lee first named and described this in his 2018 thesis before officially
describing and naming it in 2019. In both papers, it emerged as a
heyuannine sister to Jiangxisaurus, Banji and "Tongtianlong". When added to the Hartman et al. matrix it emerges sister to Heyuannia yanshini, so may end up falling under the genus Heyuannia if future studies support this.
References- Lee, 2018. Two new
maniraptorans (Dinosauria: Theropoda) from the Nemegt Formation (early
Maastrichtian) of Mongolia. Masters Thesis, Seoul National University
School of Earth and Environmental Sciences. 280 pp.
Funston, 2019. Anatomy, systematics, and evolution of Oviraptorosauria
(Dinosauria, Theropoda). PhD thesis, University of Alberta. 774 pp.
Lee, Lee, Chinsamy, Lu, Barsbold and Tsogtbaatar, 2019. A new baby
oviraptorid dinosaur (Dinosauria: Theropoda) from the Upper Cretaceous
Nemegt Formation of Mongolia. PLoS ONE. 14(2), e0210867.