Paraves Sereno, 1997
Definition- (Passer domesticus <- Oviraptor philoceratops)
(Holtz and Osmolska, 2004; modified from Sereno, 1998)
(Vultur gryphus <- Oviraptor philoceratops) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
= Troodontidae sensu Varricchio, 1997
Definition- (Troodon formosus, Saurornithoides mongoliensis, Borogovia
gracilicrus, Sinornithoides youngi <- Ornithomimus velox,
Oviraptor philoceratops)
= Paraves sensu Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Vultur gryphus <- Oviraptor philoceratops)
Comments- Paraves as a concept can apply to any topology including Oviraptor and birds, but explicit topologies were rare until the 1990s. Kurzanov (1981) described Avimimus
as closer to birds than known theropods, as also assumed by Chatterjee
(1991), so would count as a paravian. Paul (1984) placed
troodontids closer to birds than oviraptorosaurs, and in 1988 also
placed avimimids there, positioning Archaeopteryx and dromaeosaurids more basal in both works. Similarly, Thulborn (1984) placed Archaeopteryx, tyrannosaurids, troodontids, ornithomimids and Avimimus closer to birds than Oviraptor,
with dromaeosaurids more basal. Gauthier (1984, 1986) had a more
modern arrangement where deinonychosaurs (including dromaeosaurids and
troodontids) were closer to birds than caenagnathids (including Oviraptor)
and elmisaurids. Holtz (1992, 1994) on the other hand only placed
dromaeosaurids there, with troodontids closer to oviraptorids.
Makovicky (1995) had a similar topology but with dromaeosaurids and Ornitholestes
in what will be Paraves. Sereno (1997) first used Paraves as part
of a node-stem triplet sister to Oviraptorosauria, but without
explicitly stating its internal specifier. This was handled by
Sereno (1998) who defined Paraves as "all maniraptorans closer to
Neornithes than to Oviraptor."
Holtz and Osmolska (2004) later added species-level specifiers.
In the early 2000s with the discovery of bird-like basal troodontids
like Sinovenator, troodontids were consistantly paravians while Avimimus and Ornitholestes
were near universally excluded (25 and 29 steps to add in Hartman et
al.'s 2019 matrix; they fall out as a scansoriopterygid and
dromaeosaurid respectively). A heterodox exception is Maryanska
et al.'s (2002; also independently recovered by Lu et al., 2002)
hypothesis that oviraptorosaurs are closer to Aves than
deinonychosaurs, so that not even Archaeopteryx
is paravian, but this takes 12 more steps in Hartman et al.
(2019). More recently, taxa which can controversially be
paravians include alvarezsauroids (11 steps to add; which brings
therizinosaurs as well), Fukuivenator (11 steps to add), Yixianosaurus (7 steps to remove), Protarchaeopteryx (3 steps to add; increased to 5 steps with added oviraptorosaurs) and scansoriopterygids (9 steps to remove).
Despite
robust support for paravian monophyly, the interrelationship between
troodontids, dromaeosaurids and birds remains contentious with recent
studies alternatively favoring joining troodontids and dromaeosaurids
as Deinonychosauria (Hartman et al., 2019; Hu et al., 2018; Lef�vre et
al., 2017; Shen et al., 2017; Godefroit et al., 2013b; Senter et al.,
2012; Turner, Makovicky & Norell, 2012), placing troodontids closer
to Aves than dromaeosaurids (Gianechini et al., 2018; Cau et al., 2017;
Foth and Rauhut, 2017; Lee et al., 2014; Foth et al., 2014; Godefroit
et al., 2013a), or joining dromaeosaurids and avialans to form
Eumaniraptora to the exclusion of troodontids (Agnolin & Novas,
2013). As first demonstrated by Xu et al. (2011) and Agnolin and
Novas (2011), the consensus positions of archaeopterygids as birds and
unenlagiines as dromaeosaurids can easily change with small adjustments
to the TWiG matrix, making these clades active variables in paravian
topology as well. Indeed, Hartman et al. recovered the topology
used here as most parsimonious, but two alternatives were only one step
longer. In one, troodontids, unenlagiids and archaeopterygids are
successively closer to birds than dromaeosaurids. In another,
unenlagiids pair with dromaeosaurids, and troodontids pair with
archaeopterygids, all within Deinonychosauria. These should be
considered basically equally likely pending further analyses.
References- Kurzanov, 1981. On the unusual theropods from Upper Cretaceous
of Mongolia. In Resetov (ed.). Iskopaemye pozvonocnye Mongolii. Trudy, Sovmestnaa
Sovetsko-Mongolskaa paleontologiceskaa ekspedicia. 15, 39-50.
Gauthier, 1984. A cladistic analysis of the higher systematic
categories of the Diapsida. PhD thesis. University of California,
Berkeley. 564 pp.
Paul, 1984. The archosaurs: A phylogenetic study. Third Symposium on Mesozoic
Terrestrial Ecosystems, Short Papers. 175-180.
Thulborn, 1984. The avian relationships of Archaeopteryx, and the origin of birds. Zoological Journal of the Linnean Society. 82(1-2), 119-158.
Gauthier, 1986. Saurischian monophyly and the origin of birds. Memoirs of the Californian Academy of Sciences 8, 1-55.
Paul, 1988. Predatory Dinosaurs of the World. Simon & Schuster: New York
464 pp.
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.
Holtz, 1992. An unusual structure of the metatarsus of Theropoda (Archosauria:
Dinosauria: Saurischia) of the Cretaceous. PhD thesis. Yale University. 347
pp.
Holtz, 1994. The phylogenetic position of the Tyrannosauridae: Implications
for theropod systematics. Journal of Paleontology. 68(5), 1100-1117.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). Masters thesis, University of Copenhagen. 311 pp.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences. 25, 435-489.
Varricchio, 1997. Troodontidae. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. 749-754.
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.
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.
Holtz and Osmolska, 2004. Saurischia. In Weishampel, Dodson
and Osmolska (eds). The Dinosauria Second Edition. University of California
Press. 21-24.
Larson, 2009. Multivariate analyses of small theropod teeth and implications
for paleoecological turnovers through time. Journal of Vertebrate Paleontology.
29(3), 132A.
Agnolin and Novas, 2011. Unenlagiid theropods: Are they members of the
Dromaeosauridae (Theropoda, Maniraptora)? Anais da Academia Brasileira
de Ci�ncias. 83(1), 117-162.
Xu, You, Du and Han, 2011. An Archaeopteryx-like theropod from China and the origin of Avialae. Nature. 475, 465-470.
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.
Sorkin, 2012. Aerial ability in basal Deinonychosauria. Journal of Vertebrate
Paleontology. Program and Abstracts 2012, 176-177.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid
systematics and paravian phylogeny. Bulletin of the American Museum of
Natural History. 371, 1-206.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic
relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. Springer Netherlands. 96 pp.
Godefroit, Cau, Hu, Escuillie, Wu and Dyke, 2013a. A Jurassic avialan
dinosaur from China resolves the early phylogenetic history of birds.
Nature. 498, 359-362.
Godefroit, Demuynck, Dyke, Hu, Escuillie and Claeys, 2013b. Reduced
plumage and flight ability of a new Jurassic paravian theropod from
China. Nature Communications. 4(1), 1394.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Lefevre, Hu, Escuillie, Dyke and Godefroit, 2014. A new long-tailed
basal bird from the Lower Cretaceous of north-eastern China. Biological
Journal of the Linnean Society. 113, 790-804.
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.
Foth and Rauhut, 2017. Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs. BMC Evolutionary Biology. 17:236.
Shen, Lu, Liu, Kundrat, Brusatte and Gao, 2017. A new troodontid
dinosaur from the Lower Cretaceous Yixian Formation of Liaoning
Province, China. Acta Geologica Sinica. 91(3), 763-780.
Gianechini, Makovicky, Apestegu�a and Cerda, 2018. Postcranial skeletal anatomy of the holotype and referred specimens of Buitreraptor gonzalezorum Makovicky, Apestegu�a and Agnol�n 2005 (Theropoda, Dromaeosauridae), from the Late Cretaceous of Patagonia. PeerJ. 6:e4558.
Hu, Clarke, Eliason, Qiu, Li, Shawkey, Zhao, D'Alba, Jiang and Xu,
2018. A bony-crested Jurassic dinosaur with evidence of iridescent
plumage highlights complexity in early paravian evolution. Nature
Communications. 9, 217.
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
Avepectora Paul, 2002
Definition- (majority of the distal edge of strongly anteriorly facing
coracoids articulates with the anterior edge of a broad sternum at an angle
of approximately 45-90 degrees from the midline as in Dromaeosaurus albertensis)
(modified from Paul, 2002)
Comments- Paul (2002) named
this to cover a similar clade to his 1988 Protoavia, identical in
content to the consensus Maniraptoriformes. He cited Pelecanimimus'
sternal complex as supporting ornithomimosaurs' inclusion, but eventual
description of the postcrania suggests a more laterally oriented
coracoid-sternal articulation. That being said, Hartman
et al.'s 2019 topology resolves the strongly bent coracoid as being a
paravian character convergent in derived oviarptorids, derived
caenagnathids, Falcarius, Neimongosaurus, patagonykines and Archaeornithomimus.
Reference- Paul, 2002. Dinosaurs of the Air. The John Hopkins University Press, Baltimore and London. 460 pp.
Eumaniraptora Padian, Hutchinson and Holtz,
1997
Definition- (Deinonychus antirrhopus + Passer domesticus)
(Maryanska, Osmolska and Wolsan, 2002; modified from Padian, Hutchinson and Holtz, 1997)
Other definitions- (Passer domesticus <- Troodon formosus)
(modified from Agnolin and Novas, 2013)
(Dromaeosaurus albertensis + Passer domesticus) (Godefroit, Cau,
Hu, Escuillie, Wu and Dyke, 2013)
(Deinonychus antirrhopus + Vultur gryphus) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
(Passer domesticus <- Anchiornis huxleyi) (Lefevre, Cau, Cincotta, Hu, Chinsamy, Escuillie and Godefroit, 2017)
= Aves sensu Chiappe, 1992
Definition- (Archaeopteryx lithographica + Passer domesticus)
= Maniraptora sensu Holtz and Padian, 1995
Definition- (Dromaeosaurus albertensis + Passer domesticus)
= Avialae sensu Wagner and Gauthier, 1999
Definition- (Archaeopteryx lithographica + Vultur gryphus)
= Avemorpha Miller, 2004
= Palaeoaves Livezey and Zusi, 2007
= Ornithes Martyniuk, 2012
Definition- (Archaeopteryx lithographica + Passer domesticus)
= Avialae sensu Agnolin and Novas, 2013
Definition- (Archaeopteryx lithographica + Passer domesticus)
(modified)
= Eumaniraptora sensu
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Deinonychus antirrhopus + Vultur gryphus)
Comments- Holtz (1992) originally named Eumaniraptora in his unpublished thesis, but used
it for what would now be called Maniraptoriformes- a clade containing paravians,
oviraptorosaurs, ornithomimosaurs and also tyrannosaurids, but not Ornitholestes
or Compsognathus.
Padian et al. (1997) in an abstract named it as a new node "to unite
the stem taxa Deinonychosauria and Avialae", though without an explicit
definition. Their definitions of Deinonychosauria and Avialae
imply a definition of Deinonychus
+ Aves, however. It was first used outside an abstract by Padian
et al. (1999), and Maryanska et al. (2002) provided species-level
specifiers.
Miller (2004) proposed Avemorpha for the clade including dromaeosaurids and avialans (but not Archaeopteryx), but this is a junior synonym of Eumaniraptora.
Palaeoaves is used as a paraphyletic parvclass within Avialae by Livezey and
Zusi (2007) that includes Archaeopteryx, Rahonavis, Confuciusornis,
enantiornithines and Apsaravis, but not hesperornithines, Ichthyornis,
lithornithids or Aves sensu stricto.
Martyniuk (2012) erected Ornithes for "< Archaeopteryx lithographica & Passer domesticus",
which is not especially useful as it could include or exclude
unenlagiids, troodontids and dromaeosaurids in trees only a steps apart
in Hartman et al.'s analysis.
References- Chiappe, 1992. Enantiornithine (Aves) tarsometatarsi and the avian affinites
of the Cretaceous Avisauridae. Journal of Vertebrate Paleontology. 12(3), 344-350.
Holtz, 1992. An unusual structure of the metatarsus of Theropoda
(Archosauria: Dinosauria: Saurischia) of the Cretaceous. Unpublished PhD thesis.
Yale University. 347 pp.
Holtz and Padian, 1995. Definition and diagnosis of Theropoda and related taxa. Journal of Vertebrate Paleontology. 15(3), 35A.
Padian, Hutchinson and Holtz, 1997. Phylogenetic definitions and
nomenclature of the major taxonomic categories of the theropod
dinosaurs. Journal of Vertebrate Paleontology. 17(3), 68A.
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.
Wagner and Gauthier, 1999. 1,2,3 5 2,3,4: A solution to the
problem of the homology of the digits in the avian hand. Proceedings of the
National Academy of Sciences. 96, 5111-5116.
Maryanska, Osmolska and Wolsan, 2002. Avialan status for Oviraptorosauria. Acta
Palaeontologica Polonica. 47 (1), 97-116.
Miller, 2004. A new phylogeny of the Dromaeosauridae. 2004 Student Showcase Journal. 20, 123-158.
Livezey and Zusi, 2007. Higher-order phylogeny of modern birds (Theropoda, Aves:
Neornithes) based on comparative anatomy. II. Analysis and discussion. Zoological
Journal of the Linnean Society. 149 (1), 1-95.
Lamm, Ksepka, Stone and Clarke, 2008. Identifying differential size trends in
Mesozoic birds using new data and a novel method. Journal of Vertebrate Paleontology.
28(3), 103A.
Bell, Chiappe and O'Connor, 2009. Ecological diversity of Mesozoic birds: Morphometric
analysis with a phylogenetic perspective. Journal of Vertebrate Paleontology.
29(3), 61A.
Chinsamy-Turan, 2009. The bone microstructure of Mesozoic birds. Journal of
Vertebrate Paleontology. 29(3), 77A.
Nudds and Dyke, 2009. Estimating the flight capabilities of extinct Mesozoic
birds from their primary feather morphology. Journal of Vertebrate Paleontology.
29(3), 156A-157A.
O'Connor, 2009. A comprehensive phylogeny of Mesozoic birds. Journal of Vertebrate
Paleontology. 29(3), 157A.
Weishampel and Habib, 2009. Flight morphology and launch dynamics of basal birds,
and the potential for competition with pterosaurs. Journal of Vertebrate Paleontology.
29(3), 199A.
Wang, Dyke and Nudds, 2010. Inferring the flight styles of early birds and flight
evolution from primary feather length. Journal of Vertebrate Paleontology. Program
and Abstracts 2010, 183A.
Dececchi and Larsson, 2011. The origin of wings. Journal of Vertebrate Paleontology.
Program and Abstracts 2011, 97.
Wang, Dyke and Palmer, 2011. Scaling in size and stiffness of avian primary
feathers: Implications for the strength of Mesozoic bird feathers. Journal of
Vertebrate Paleontology. Program and Abstracts 2011, 211.
Dececchi, 2012. Patterns and processes at origin of birds: Macroevolutionary
tempo and mode. Journal of Vertebrate Paleontology. Program and Abstracts 2012,
85-86.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Vernon, New Jersey. Pan Aves. 189 pp.
Mitchell, Makovicky and Gao, 2012. Paleoecology of the Jehol birds inferred
from modern bird ecomorphology. Journal of Vertebrate Paleontology. Program
and Abstracts 2012, 143.
O'Connor, 2012. Dietary evolution in Mesozoic birds. Journal of Vertebrate Paleontology.
Program and Abstracts 2012, 151.
Agnolin and Novas, 2013. Avian ancestors: A review of the phylogenetic
relationships of the theropods Unenlagiidae, Microraptoria, Anchiornis and Scansoriopterygidae. Springer Netherlands. 96 pp.
Dececchi, Habib and Larsson, 2013. Testing wing assusted incline running (WAIR):
Investigating the terrestrial origin of the avian flight stroke. Journal of
Vertebrate Paleontology. Program and Abstracts 2013, 113.
Field and Lynner, 2013. Precise inference of avialan flight ability from shoulder
joint dimensions. Journal of Vertebrate Paleontology. Program and Abstracts
2013, 126.
Godefroit, Cau, Hu, Escuillie, Wu and Dyke, 2013. A Jurassic avialan
dinosaur from China resolves the early phylogenetic history of birds.
Nature. 498, 359-362.
Xu, Zhao and Han, 2013. Homeotic transformation in the evolution of the theropod
semilunate carpal. Journal of Vertebrate Paleontology. Program and Abstracts
2013, 241.
Naish, 2014. The fossil record of bird behaviour. Journal of Zoology. 292(4),
268-280.
O'Connor and Zhou, 2014. Earliest stages in the evolution of the modern avian
skeleton: Archaeopteryx and the Jehol avifauna compared. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 197.
Sanz, Serrano, Martin-Serra and Palmqvist, 2014. Revisiting size trends in early
stem birds. Journal of Vertebrate Paleontology. Program and Abstracts 2014,
221.
Serrano, Palmqvist, Martin-Serra and Sanz, 2014. Morphofunctional evolution
of the humerus in the avian lineage. Journal of Vertebrate Paleontology. Program
and Abstracts 2014, 228.
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.
Lefevre, Cau, Cincotta, Hu, Chinsamy, Escuillie and Godefroit, 2017. A
new Jurassic theropod from China documents a transitional step in the
macrostructure of feathers. The Science of Nature. 104:74.
Eumaniraptora incertae sedis
Cretaaviculus Bazhanov,
1969
C. sarysuensis Bazhanov, 1969
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Holotype- contour feather (17 mm)
Comments- This specimen is a small, asymmetric contour feather with a
length of 17 mm and a width of 4.8-5.5 mm. The barbs are at a 20 degree angle
to the shaft. The asymmetry suggests it is a paravian. Nessov (1992) considers
it indeterminate, which is true so far as feather characteristics cannot be
used to diagnose Mesozoic birds.
References- Bazhanov, 1969. [On the record of a bird remain living in
the Cretaceous in the USSR]. Tezisy Dokladov XV Sessii Vsesoyuznogo Paleontologicheskogo
Obshchestva. 5-6.
Shillin and Romanova, 1978. [Senonian floras of Kazakhstan]. Alma-Ata: Kairat
Publishing. 176 pp.
Nessov, 1992. Mesozoic and Paleogene birds of the USSR and their paleoenvironments.
in Campbell (ed). Papers in Avian Paleontology Honoring Pierce Brodkorb. 465-478.
Eumaniraptora indet. (Lambe, 1902)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith River Group, Alberta, Canada
Material- (CMN coll.) proximal caudal vertebra (Lambe, 1902)
Comments- The CMN vertebra was identified by Lambe (1902) as a posterior dorsal
tentatively referred to Struthiomimus (his Ornithomimus altus)
(plate XV figure 6-8), but is actually a caudal, and may be Troodon or
Saurornitholestes based on its low square articular face, amphicoelous centrum and lack of a pleurocoel.
References- Lambe, 1902. New genera and species from the Belly River Series
(Mid-Cretaceous). Geological Survey of Canada Contributions to Canadian Palaeontology.
3(2), 25-81.
unnamed eumaniraptoran (Dumont, Tafforeau, Bertin, Bhullar, Field, Schulp, Strilisky,
Thivichon-Prince, Viriot and Louchart, 2016)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
Material- (RTMP 94.31.32) tooth
Comments- Initially identified
as Aves indet. based on the small size, the completely serrated distal
edge suggests otherwise. Both edges are straight with a smooth
carina at the base of the mesial esdge.
Reference-
Dumont, Tafforeau, Bertin, Bhullar, Field, Schulp, Strilisky,
Thivichon-Prince, Viriot and Louchart, 2016. Synchrotron imaging of
dentition provides insights into the biology of Hesperornis and Ichthyornis, the "last" toothed birds. BMC Evolutionary Biology. 16:178.
unnamed possible eumaniraptoran (Gates, Zanno and Makovicky, 2015)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
Material- (FMNH PR2900) tooth (4.4x2.2x1 mm)
Comments- This is an anterior tooth which lacks serrations and a basal constriction
Reference- Gates, Zanno and Makovicky, 2015. Theropod teeth from the
upper Maastrichtian Hell Creek Formation "Sue" Quarry: New morphotypes
and faunal comparisons. Acta Palaeontologica Polonica. 60(1), 131-139.
Eumaniraptora indet. (Kessler and Jurcsak, 1984)
Late Berriasian-Early Valanginian, Early Cretaceous
Cornet bauxite, Bihor, Romania
Material- (MTCO 17558; = MTCO-P 207; paratype of Limnornis corneti)
proximal scapula
?(MTCO 17966) proximal radius
?(MTCO 17982) partial furcula
Comments- Kessler (1984) noted approximately sixty bird-like fragments
from the Cornet bauxite, six of which can "with certainty be attributed
to birds." These are the holotypes of Eurolimnornis corneti and
Palaeocursornis corneti in addition to other isolated fragments, but
most have been moved to Pterosauria or more generally Ornithodira and Archosauria
indet. by Dyke et al. (2011) and Agnolin and Varricchio (2012).
MTCO 17558 was originally (Kessler and Jurcsak, 1984) decribed as a partial
carpometacarpus and paratype of Limnornis corneti (later renamed Palaeocursornis
corneti), and later (Kessler and Jurcsak, 1986) a paratype of Eurolimnornis
corneti. Kurochkin (1995) used the carpometacarpus identification to argue
for avian affinities of Eurolimnornis due to supposed distal fusion and
tendinal sulcus, but Hope (2002) noted these are also present in Ichthyornis.
Hope also noted it couldn't be referred definitively to Eurolimnornis
or Palaeocursornis, and Dyke et al. reinterpreted it as a bird proximal
scapula (though they probably mistakenly wrote it is not identifiable as a bird).
MTCO 17966 and 17982 were stated to be birds by Dyke et al., but that identity
was doubted by Agnolin and Varricchio.
References- Kessler, 1984. Lower Cretaceous birds from Cornet, Roumania.
In Rief and Westphal (eds.). Third Symposium on Mesozoic Terrestrial Ecosystems,
Tubingen. 119-121.
Kessler and Jurcsak, 1984. Fossil bird remains in the bauxite from Cornet (Romania,
Bihor County). Travaux du Musee d'Histoire Naturelle, Grigore Antipa. 25, 393-401.
Kessler and Jurcsak, 1986. New contributions to the knowledge of the Lower Cretaceous
bird remains from Cornet (Romania). Travaux du Musee d'Histoire Naturelle, Grigore
Antipa. 28, 289-295.
Kurochkin, 1995. Synopsis of Mesozoic birds and early evolution of class Aves.
Archaeopteryx. 13, 47-66.
Hope, 2002. The Mesozoic radiation of Neornithes. In Chiappe and Witmer (eds.).
Mesozoic birds: Above the Heads of Dinosaurs. University of California
Press. 339-388.
Dyke, Benton, Posmosanu and Naish, 2011. Early Cretaceous (Berriasian) birds
and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology. 54(1),
79-95.
Agnolin and Varricchio, 2012 . Systematic reinterpretation of Piksi barbarulna
Varricchio, 2002 from the Two Medicine Formation (Upper Cretaceous) of western
USA (Montana) as a pterosaur rather than a bird. Geodiversitas. 34(4), 883-894.
unnamed Eumaniraptora (Pe�alver, Arillo, Delcl�s, Peris, Grimaldi, Anderson, Nascimbene and P�rez-de la Fuente, 2017)
Early Cenomanian, Late Cretaceous
Noije bum mines, Myanmar
Material-
(AMNH Bu JZC-F18) feather, feather fragments (Pe�alver, Arillo,
Delcl�s, Peris, Grimaldi, Anderson, Nascimbene and P�rez-de la Fuente,
2017)
(LV-0321) distal phalanx II-1, phalanx II-2 (4.7 mm), manual ungual II
(2 mm), manual claw sheath, remiges (Xing, McKellar and O'Connor, 2020)
Comments- Pe�alver et al.
(2017) describe an amber sample (AMNH Bu
JZC-F18) containing a feather, ixodid tick nymph and other
arthropods. The feather is asymmetrical with barbules, indicating
it belonged to a eumaniraptoran. St. Fleur's (2017) newspaper
article says "the tick was a nymph ... and that its host was most
likely some sort of fledgling dinosaur no bigger than a hummingbird,
which Dr. Grimaldi referred to as a "nanoraptor."" Yet nothing in
Pe�alver et al. suggests the age or size of the dinosaur, and the term
"nanoraptor" misleadingly suggests a deinonychosaur or oviraptorosaur
instead of an avialan. When the article later states "the host
was more likely a nonavian dinosaur and not a modern bird based on
molecular dating", its using 'avian' in the crown Aves sense without
explicitly stating so, correlating to Pe�alver et al.'s statement
"crown-group birds are excluded as possible hosts because their
inferred age is significantly younger than Burmese amber." Yet
there are several groups such as confuciusornithiforms and
enantiornithines, the latter known from multiple specimens in Myanmar
amber, that
are plausible sources of the feather and hosts of the ixodid ticks but
which would not be considered raptors in the colloquial sense.
It's also possible AMNH Bu JZC-F18 is from e.g. an archaeopterygid or
microraptorian, but nothing in the feather's structure or stratigraphy
has favored any particular option, and "nanoraptor" is here considered
a hypothetical animal akin to "proavis" and not a nomenclatural
suggestion for the taxon which grew the preserved feather.
Xing et al. (2020) described distal wing LV-0321 which has asymmetrical
remiges with barbules, so is eumaniraptoran. Given its size, this
is either an extremely young juvenile or an ornithothoracine, and the
plumage gives no indication of juvenile status. Tha size of the
manual ungual compared to the phalanx is less than deinonychosaurs
except Buitreraptor, but the phalanx is not incredibly elongate as it is in that genus. The ratio is similar to Fukuipteryx, Zhongornis and some ornithothoracines.
References- Pe�alver, Arillo, Delcl�s, Peris, Grimaldi, Anderson,
Nascimbene and P�rez-de la Fuente, 2017. Ticks parasitised feathered
dinosaurs as revealed by Cretaceous amber assemblages. Nature
Communications. 8: 1924.
St. Fleur, 2017. Ticks trapped in amber were likely sucking dinosaur blood. The New York Times. December 12.
Xing, McKellar and O'Connor, 2020. An unusually large bird wing in
mid-Cretaceous Burmese amber. Cretaceous Research. Journal Pre-proof
DOI: 10.1016/j.cretres.2020.104412
unnamed possible eumaniraptoran (Riff, Kellner, Mader and Russell, 2002)
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Material- (CMN 50852) incomplete dorsal vertebra (21 mm)
Comments- Riff et al. (2002, 2004) considered this most similar to Rahonavis,
though it differs in being more elongate, having a larger neural canal and lacking
pleurocoels or lateral fossae. Chiarenza and Cau (2016) stated the specimen
lacked unambiguous paravian or avialan characters, noting in particular the
large neural canal is present in many small theropods and crocodyliforms.
References- Riff, Kellner, Mader and Russell, 2002. On the occurence
of an avian vertebra in Cretaceous strata of Morocco, Africa. Anais da Academia
Brasileira de Ciencias. 74(2), 367-368.
Riff, Mader, Kellner and Russell, 2004. An avian vertebra from the continental
Cretaceous of Morocco, Africa. Arquivos do Museu Nacional. 62(2), 217-223.
Chiarenza and Cau, 2016. A large abelisaurid (Dinosauria, Theropoda) from Morocco
and comments on the Cenomanian theropods from North Africa. PeerJ. 4:e1754.
undescribed eumaniraptoran (Naish, Martill and Merrick, 2007)
Late Aptian, Early Cretaceous
Nova Olinda Member of the Crato Formation, Brazil
Material- (Senckenberg Museum coll.) carpals, three remiges (14, 81 mm)
Comments- Naish et al. (2007) discuss this as bird remains, but as e.g.
Microraptor has asymmetrical feathers too, it may be from another kind
of paravian.
Reference- Naish, Martill and Merrick, 2007. Birds of the Crato Formation.
In: Martill, Bechly and Loveridge (eds.). The Crato fossil beds of Brazil: Window
into an ancient world. Cambridge University Press. 525-533.
Deinonychosauria Colbert and Russell,
1969
Definition- (Deinonychus antirrhopus <- Passer domesticus)
(Holtz and Osmolska, 2004; Padian, 1997)
Other definitions- (Troodon formosus + Dromaeosaurus albertensis)
(Holtz and Osmolska, 2004; modified from Sereno, 1997)
(Troodon formosus + Velociraptor mongoliensis) (modified from
Sereno, 1998)
(Dromaeosaurus albertensis <- Passer domesticus) (Godefroit,
Demuynck, Dyke, Hu, Escuillie and Claeys, 2013)
(Troodon formosus + Velociraptor mongoliensis, - Passer domesticus)
(Hendrickx, Hartman and Mateus, 2015)
(Troodon formosus + Velociraptor mongoliensis, - Ornithomimus
edmontonicus, Passer domesticus) (Sereno, online 2005)
= Saurornithes Nicholson, 1878a/b
Definition- (Archaeopteryx lithographica <- Passer domesticus)
(Martyniuk, 2012)
= Dromaeosauria Bonaparte and Novas, 1985
= Archaeopteryx sensu Sereno, 1998
Definition- (Archaeopteryx lithographica <- Passer domesticus)
(modified)
= Archaeopterygidae sensu Sereno, online 2005
Defrinition- (Archaeopteryx lithographica <- Passer domesticus)
= Dromaeosauridae sensu Godefroit, Demuynck, Dyke, Hu, Escuillie and Claeys,
2013
Definition - (Dromaeosaurus albertensis <- Passer domesticus)
= Tetrapterygidae Chatterjee, 2015
Deinonychosauria defined- There have been two basic suggested definitions for Deinonychosauria,
one stem-based (Deinonychus <- Passer) by Padian (1997) and
the other node based (Troodon + dromaeosaurids) by Sereno (1997). I prefer Padian's because it is based on
the eponymous genus, and Colbert and Russell (1969) did not originally specify
the inclusion of troodontids. They only include dromaeosaurids in the taxon,
and only mention Dromaeosaurus, Deinonychus and Velociraptor
as members of that family. Furthermore, this gives a name to the
clade including archaeopterygids and unenlagiids in some most
parsimonious topologies and ensures Deinonychosauria does not self
destruct or include Aves.
Referral of isolated teeth-
With very limited exceptions, theropod teeth with constricted roots are
from Maniraptoriformes, and except for therizinosaurs basically all
serrated varieties are from deinonychosaurs. The only(?) known
exception is irregularily developed distal crenulations on Longipteryx
premaxillary teeth (Wang et al., 2015), with supposed avialan teeth
described by Sankey et al. (2002) from the Dinosaur Park Formation not
associated with cranial or skeletal material to verify their
identity. Thus serrated paravian teeth are here referred to
Deinonychosauria, though trees almost as short as the one used here
placed taxa with serrated teeth like troodontids as avialans.
Dromaeosauria- While the term
'dromaeosaur' is a common short form of dromaeosaurid, use of
Dromaeosauria itself is much rarer in the published literature.
The earliest example of the latter may be Bonaparte and Novas (1985)
who seem to treat Dromaeosauria (containing at least Dromaeosaurus)
as a theropod group on par with Carnosauria and Segnosauria.
Similarly, Dubois (2006) proposed Dromaeosauria alongside Aves at the
level of phalanx, between family and order, but his scheme of new
Linnaean levels was not widely adopted. Fossilworks (online 2007)
presents Chatterjee and Templin (2007) as the authors of Dromaeosauria,
who use it in a cladogram as a eumaniraptoran clade containing Microraptor and Pedopenna, sister to Aves (Avialae in official usage). Other examples are mistakes for Dromaeosauridae or Eudromaeosauria.
Tetrapterygidae- Chatterjee (2015) proposed the family Tetrapterygidae for a
clade including Microraptor, Xiaotingia, Anchiornis and
Aurornis. However, as noted by Martyniuk (online, 2015), according to
ICZN Article 11.7.11 a family-group name must be "formed from the stem
of an available generic name" which "must be a name then used as valid
in the new family-group taxon." As Chatterjee did not intend for Tetrapterygidae
to include a genus Tetrapteryx, it is invalid. Furthermore, Tetrapteryx
has already been used for a genus of gruid bird (Thunberg, 1818), corrently considered
a synonym of Anthropoides. Finally, if Microraptor is in a family, the
family must be named Microraptoridae because that family was already erected
by Longrich and Currie (2009).
References-
Thunberg, 1818. Tetrapteryx capensis, ett nytt Fogelslaegte.
Kongliga Svenska Vetenskaps Academiens nya Handlingar. 1818(2), 242-245.
Nicholson, 1878a. A Manual of Zoology for the Use of Students with a General
Introduction on the Principles of Zoology. Blackwood and Sons: Edinburgh and
London. 800 pp.
Nicholson, 1878b. Advanced Text-Book of Zoology for Junior Students. Blackwood
and Sons: Edinburgh and London. 405 pp.
Colbert and Russell, 1969. The small Cretaceous dinosaur Dromaeosaurus.
American Museum Novitates. 2380, 49 pp.
Bonaparte and Novas, 1985. Abelisaurus comahuensis,
n. g., n. sp., Carnosauria from the Late Cretaceous of Patagonia. Ameghiniana.
21, 259-265.
Padian, 1997. Avialae. In Currie and Padian (eds.). Encyclopedia of Dinosaurs. Elsevier Inc. 39-40.
Sereno, 1997. The origin and evolution of dinosaurs. Annual Review of Earth and Planetary Sciences. 25, 435-489.
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.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Godefroit, Demuynck, Dyke, Hu, Escuillie and Claeys, 2013b. Reduced
plumage and flight ability of a new Jurassic paravian theropod from
China. Nature Communications. 4(1), 1394.
Holtz and Osmolska, 2004. Saurischia. In Weishampel, Dodson
and Osmolska (eds.). The Dinosauria Second Edition. University of California
Press. 21-24.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php
[version 1.0, 2005 November 7]
Dubois, 2006. Proposed rules for the incorporation of nomina of
higher-ranked zoological taxa in the International Code of Zoological
Nomenclature. 2. The proposed rules and their rationale. Zoosystema.
28(1), 165-258.
Chatterjee and Templin, 2007. Biplane wing planform and flight performance of the feathered dinosaur Microraptor gui. Proceedings of the National Academy of Sciences, 104(5), 1576-1580.
Fossilworks, online 2007. http://fossilworks.org/?a=referenceInfo&reference_no=19832
Longrich and Currie, 2009. A microraptorine (Dinosauria - Dromaeosauridae) from
the Late Cretaceous of North America. Proceedings of the National Academy of
Sciences. 106(13), 5002-5007.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Vernon, New Jersey. Pan Aves. 189 pp.
Chatterjee, 2015. The Rise of Birds: 225 Million Years of Evolution. Second
Edition. Johns Hopkins University Press. 392 pp.
Hendrickx, Hartman and Mateus, 2015. An overview of non-avian
theropod discoveries and classification. PalArch's Journal of Vertebrate Palaeontology.
12(1), 1-73.
Chatterjee, 2015. The Rise of Birds: 225 Million Years of Evolution. Second
Edition. Johns Hopkins University Press. 392 pp.
Martynuik, online 2015. https://web.archive.org/web/20150927175357/http://dinogoss.blogspot.com/2015/05/the-crane-and-microraptor.html
Wang, Zhao, Shen, Liu, Gao, Cheng and Zhang, 2015. New material of Longipteryx
(Aves: Enantiornithes) from the Lower Cretaceous Yixian Formation of China with
the first recognized avian tooth crenulations. Zootaxa. 3941(4), 565-578.
undescribed Deinonychosauria (Ikejiri, Watkins and Gray, 2006)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Wyoming, US
Material- (WDC BS-641) tooth (9.1 x 9.2 x 4.2 mm)
(WDC BS-885) tooth (12.6 x 8.3 x 3.6 mm)
(WDC BS-889) tooth (12.2 x 7 x 3.2 mm)
Comments- Called
"deinonychosaurid(?)" in the measurements table, Ikejiri et al. (2006)
describe these are more recurved and transversely compressed than
Allosaurus, with relatively larger serrations, high DSDI and smooth
enamel. They furthermore state WDC BS-641 has mesial serrations
conficed to the apical fifth.
Reference- Ikejiri, Watkins and Gray, 2006. Stratigraphy, sedimentology,
and taphonomy of a sauropod quarry from the Upper Jurassic Morrison Formation
of Themopolis, central Wyoming. In Foster and Lucas (eds.). New Mexico Museum
of Natural History Bulletin. 36, 36-46.
unnamed deinonychosaur (Rodriguez de la Rosa and Cevallos-Ferriz, 1998)
Early Maastrichtian, Late Cretaceous
Ca�on del Tule Formation, Mexico
Material- (IGM-7711) pedal phalanx II-I (18.8 mm)
(IGM-7712) pedal phalanx II-2 (21.7 mm)
Comments- Although described as being from the Cerro del
Pueblo Formation, Aguillon Martinez (2010) found this and other
material from the El Pelillal locality belong to the later Ca�on del
Tule Formation.
These may belong to the same individual, which was tentatively
referred to Troodontidae by Rodriguez de la Rosa and Cevallos-Ferriz (1998)
based on the centrally placed collateral ligament pit, which is also found in
Saurornithoides and Troodon. However, the longer II-2
compared to II-1 is more similar to archaeopterygids and dromaeosaurids. It is from a different taxon
than IGM-7710 as it is not nearly as elongate, and different from IGM-7715 as
it has a centrally placed collateral ligament pit.
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, 751-764.
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.
unnamed deinonychosaur (Weigert 1995)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Material- (IPFUB GUI ARCH. 1) tooth
(IPFUB GUI ARCH. 3) tooth
(IPFUB GUI ARCH. 4) tooth
(IPFUB GUI ARCH. 5) tooth
(IPFUB GUI ARCH. 7) tooth
(IPFUB GUI ARCH. coll.) 98 teeth (1-2.6 mm)
Comments- Weigert (1995) assigned these teeth to cf. Archaeopteryx sp..
These teeth are constricted basally, with mesial and distal
carinae. Both lingual and labial surfaces are generally smooth, and the lingual
surface is concave apically. The labial surface sometimes has faint grooves,
which were believed to be from wear. Eighty-six teeth have both mesial and distal
serrations, while the other seventeen sometimes lack mesial serrations and have
carinae shifted lingually, so are probably premaxillary teeth. The teeth average
1.65 mm in crown height and .60 mm in FABL. Serrations are low and rounded or
slightly pointed, with a miniscule size of 24/mm.
The specimens differ from Archaeopteryx and most other archaeopterygids in
having serrations. Weigert (1995) states these may be present but hidden in
the former genus, since most Archaeopteryx teeth are only visible in
labial view, where the serrations cannot be seen in the Portuguese specimens.
Yet the London and Munich specimens of Archaeopteryx both expose serrationless
teeth in lingual view, showing Weigert is incorrect. He explained the absence
of serrations in the Munich specimen by claiming it was a distinct species (A.
bavarica) which may have differed from A. lithographica in this respect,
but this is special pleading, and the distinctness of A. bavarica is
very poorly supported (see discussion in Archaeopteryx entry). The diagonally
oriented carina on the apical half of the tooth is described as an archaeopterygiform
apomorphy, but this is true of any teeth which share the same stout, highly
recurved outline (e.g. Sinornithoides). The other cited archaeopterygiform
apomorphy is the lingually projected mesial carina, but this is found in other
taxa such as Sinornithosaurus' anterior teeth. The teeth do seem to be
maniraptoriform, based on the constricted crown base, and are not referrable
to several groups due to their serrations (Ornithomimosauria, Alvarezsauroidea,
Oviraptorosauria, Avialae), but also lack the enlarged serrations found in most
therizinosaurs and derived troodontids.
Reference- Weigert, 1995. Isolated teeth of cf. Archaeopteryx sp.
from the Upper Jurassic of the coalmine Guimarota (Portugal. Neues
Jahrbuch f�r Geologie und Pal�ontologie, Monatshefte. 9, 562-576.
unnamed deinonychosaur (Buffetaut, Marandat and Sige, 1986)
Late Cretaceous
Serviers, Gard, France
Material- (Universite des Sciences et Techniques de Languedoc SER 03) partial
tooth (~2.2 mm)
Comments- This tooth is relatively elongate and barely recurved. It has
14 distal serrations per mm, while the mesial serrations are indistinct. It
was assigned to Dromaeosauridae by Buffetaut et al. (1986), but is most similar
to supposed bird teeth from the Dinosaur Park Formation
in its small size, tiny serrations, lack of much recurvature, and constricted
base.
References- Buffetaut, Marandat and Sige, 1986. Decourvert de dents de
Deinonychosaures (Saurischia, Theropoda) dans le Creace superieur du Sud de
la France. Les Comptes rendus de l'Acad�mie des sciences. 303, Serie
II(15), 1393-1396.
unnamed deinonychosaur (Buffetaut, Marandat and Sige, 1986)
Late Campanian-Early Maastrichtian, Late Cretaceous
Champ-Garimond, Gard, France
Material- (Universite des Sciences et Techniques de Languedoc CHG 48) tooth
(1.9 mm)
Comments- This tooth is short and slightly recurved, with a constricted
base. There are 15 distal serrations per millimeter, while mesial serrations
are only present as fine outlines. Like SER 03, it was assigned to Dromaeosauridae
by Buffetaut et al. (1986), but is most similar to supposed bird teeth from
the Dinosaur Park Formation in its small size, tiny serrations
which are absent mesially, lack of much recurvature, and constricted base.
References- Buffetaut, Marandat and Sige, 1986. Decourvert de dents de
Deinonychosaures (Saurischia, Theropoda) dans le Creace superieur du Sud de
la France. Les Comptes rendus de l'Acad�mie des sciences. 303, Serie
II(15), 1393-1396.
undescribed deinonychosaur (Debeljak, Kosir and Otonicar, 1999)
Campanian-Maastrichtian, Late Cretaceous
Kozina, Slovenia
Material- (ACKK-D-8/088; 5th morphotype) tooth (~4.9x~3.2x? mm) (Debeljak, Kosir, Buffetaut and Otonicar, 2002)
Comments- This tooth is
slightly recurved with a constricted root, serrated both mesially (at
least apical half) and distally with rounded, small and subequal
denticles. While stated to be "more troodontid-like" and referred to
"? Troodontidae", the combination of characters is unlike that family
and the tooth is here referred to Deinonychosauria.
References-
Debeljak, Kosir and Otonicar, 1999. A preliminary note on dinosaurs and
non-dinosaurian reptiles from the Upper Cretaceous carbonate platform
succession at Kozina (SW Slovenia). Dissertationes / Academia
Scientiarum et Artium Slovenica. Classis IV, Historia naturalis. 40,
3-25.
Debeljak, Kosir, Buffetaut and Otonicar, 2002. The Late Cretaceous dinosaurs
and crocodiles of Kozina (SW Slovenia): A preliminary study. Memorie della Societa
Geologica Italiana. 57, 193-201.
unnamed Deinonychosauria (Csiki and Grigorescu, 1998)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Material- (FGGUB R.1318) tooth (12.5x8x5.3 mm)
(FGGUB R.1319) tooth (11x8x5 mm)
(FGGUB R.1320) tooth (5.3x2.9x? mm)
(MAFI v.12685a) tooth (11.2x5.8x4.1 mm)
(MAFI v.12685b) tooth (8x3.9x2.3 mm)
Comments- These
teeth are slightly recurved with constricted roots, poorly developed
blood pits and small rectanguilar serrations (~5/mm) which are present
and similarly sized (DSDI .91-1) on both carinae. While they were
described as troodontid-like, only Troodon and Hesperornithoides have
mesial serrations, and the Sinpetru teeth seem more similar to
Microraptor than to any of those (large and hooked serrations in
Troodon, highly recurved crowns with apically limited mesial serrations
and high DSDI in Hesperornithoides).
They are thus only referred to Deinonychosauria here, which have poorly
established identifications in Cretaceous Europe.
Reference- Csiki and Grigorescu, 1998. Small theropods from the Late Cretaceous of the
Hateg basin (western Romania) - An unexpected diversity at the top of the food
chain. Oryctos. 1, 87-104.
undescribed Deinonychosauria (Nessov, 1977)
Early Cenomanian, Late Cretaceous
Khodzhakul Formation, Uzbekistan
Material- teeth
Comments- From Sheikhdzheili II "teeth of Deinonychosauria
(identification by A. K. Rozhdestvensky from collections made by the
author)" and from Khodzhakulsai "teeth of ?Deinonychosauria** (Nessov,
1977)" ("Discoveries made by [Nessov]'s assistants or by [Nessov]
himself and identified by [Nessov] are marked with two asterisks").
References- Nessov, 1977. [Turtles and some other reptiles of the Cretaceous
of Karakalpakia]. Voprosy gyerpyetology. Chyetvyertaya Vsyesoyuznaya Gyerpyetologichyeskaya
konfyeryentsiya, Lyeningrad. Avtoryefyerat doklada. Lyeningrad, Nauka. 155-156.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University, St. Petersburg. 1-156.
unnamed deinonychosaur (Wang, Cau, Wang, Yu, Wu, Wang and Liu, 2023 online)
Early Aptian, Early Cretaceous
Pigeon Hill, Longjiang Formation, Inner Mongolia, China
Material- (LY 2022JZ3004)
fragmentary proximal caudal vertebrae, fragmentary sacrum/ilium,
proximal pubes, ischia (~50 mm), incomplete femora (~134 mm), proximal
tibia, proximal fibula, pedal phalanx (~15 mm), fragmentary pedal
phalanges, pedal ungual
Comments- This was discovered in 2022. Wang et al. (2023) note "the material is comparable in size to LY 2022JZ3001 [the Migmanychion
type], the lack of evidence supporting their direct association
prevents the referral of this material to the same individual."
The recovery of Migmanychion
here as a basal paravian using Hartman et al.'s maniraptoromorph matrix
makes this plausible. Wang et al. found that "the small size of
the ischium compared to the femur (ischium-to-femur ratio < 0.4) is
more consistent with some early-diverging paravian clades than with
oviraptorosaurs or other theropods (e.g., Anchiornithidae..." and
further that "The slight cranioventral orientation of the pubis is
plesiomorphic among maniraptorans and excludes this specimen from
late-diverging deinonychosaurs, therizinosaurids, parvicursorines or
avialans." Notably the large obturator process excludes this
specimen from Avialae in the current topology, making it likely to be a
deinonychosaur.
Reference- Wang, Cau, Wang, Yu,
Wu, Wang and Liu, 2023 online. A new theropod dinosaur from the Lower
Cretaceous Longjiang Formation of Inner Mongolia (China). Cretaceous
Research. Journal Pre-proof, 10565. DOI: 10.1016/j.cretres.2023.105605.
unnamed deinonychosaur (Han, Clark, Xu, Sullivan, Choiniere and Hone, 2011)
Late Oxfordian, Late Jurassic
Wucaiwan, Upper Shishugou Formation, Xinjiang, China
Material- (IVPP V15850) tooth (4.8x3.9x2.5 mm)
Comments- Discovered by the Sino-American expeditions between 2001 and 2010, this is called
Morphotype 7 by Han et al. (2011), who refer it to a troodontid. It
is placed in Deinonychosauria here based on the combination of
constricted base, recurvature and small serrations limited to part of
the distal carina.
Reference- Han, Clark, Xu,
Sullivan, Choiniere and Hone, 2011. Theropod teeth from the
Middle-Upper Jurassic Shishugou Formation of northwest Xinjiang, China.
Journal of Vertebrate Paleontology. 31(1), 111-126.
unnamed deinonychosaur (Dong, 1997)
Early Albian, Early Cretaceous
Upper Gray Beds of the Zhonggou Formation, Gansu, China
Material- (IVPP V11122-2) tooth (~.79x~.63x? mm)
Comments- This short recurved tooth with a basal constriction was reported
to have mesial and distal serrations, both of which are much smaller than in e.g. Sinornithoides. It was referred to Troodontidae, but is here placed more generically in Deinonychosauria.
Reference-
Dong, 1997. On small theropods from Mazongshan area, Gansu province,
China. In Dong (ed.). Sino-Japanese Silk Road Dinosaur Expedition.
China Ocean Press. 13-18.
unnamed possible deinonychosaur (Amiot, Buffetaut, Tong, Boudad and Kabiri,
2002)
Cenomanian, Late Cretaceous
Kem Kem beds, Morocco
Material- (M-ZA-017) tooth (14 mm) (Amiot, Buffetaut, Tong, Boudad and Kabiri,
2004)
Comments- This tooth is straight and short, with a BW/FABL of 0.6. The
lingual side is concave, and the labial side convex. Both carinae have serrations
which are apically inclined and lie on the midline. Mesial serrations (2/mm)
and distal serrations (2.1/mm) have shallow interdenticle pits.
Amiot et al. (2002) first referred this specimen to Troodontidae based
on the lens-shaped cross section, low DSDI, and large serrations which
are apically inclined. They later (2004) referred it to
Dromaeosauridae, as Currie said the base was too narrow and the
serrations resembled dromaeosaurids. The serrations are not as
large as troodontines, but many troodontids have small serrations
anyway. This may also be a noasaurid anterior tooth, based on
geography and resemblence to Vespersaurus.
References- Amiot, Buffetaut, Tong, Boudad and Kabiri, 2002. Laurasian
theropod dinosaur teeth from the Late Cretaceous of Morocco. Conference abstract,
Third Georges Cuvier Symposium, Montbeliard, France.
Amiot, Buffetaut, Tong, Boudad and Kabiri, 2004. Isolated theropod teeth from
the Cenomanian of Morocco and their palaeobiogeographical significance. Revue
de Paleobiologie, Geneve. 9, 143-149.
undescribed possible Deinonychosauria (Gallina, Apesteguia, Haluza and Canale,
2014)
Late Berriasian-Valanginian, Early Cretaceous
Bajada Colorada Formation, Neuquen, Argentina
Comments- Gallina et al. (2014) mentioned possible deinonychosaurs from
this formation.
Reference- Gallina, Apesteguia, Haluza and Canale, 2014. A diplodocid
sauropod survivor from the Early Cretaceous of South America. PLoS ONE. 9(5),
e97128.
undescribed Deinonychosauria (Franco-Rosas, 2001)
Campanian-Maastrichtian, Late Cretaceous
Parecis Group, Brazil
Material- teeth
Comments- These were initially noted as being from the Cambambe Formation
of the Bauru Group, but Bittencourt and Langer (2011) reassigned the locality
to the later Parecis Group.
References- Franco-Rosas, 2001. Dentes de teropodomorfos da Forma��o
Cambambe, Mato Grosso. Congresso Brasileiro de Paleontologia XVII. Boletim
de Resumos. 157.
Bittencourt and Langer, 2011. Mesozoic dinosaurs from Brazil and their biogeographic
implications. Anais da Academia Brasileira de Ci�ncias. 83(1), 23-60.
unnamed possible deinonychosaur (Delcourt and Grillo, 2014)
Campanian-Maastrichtian, Late Cretaceous
Vale do Rio do Peixe Formation, Brazil
Material- (DGM 930-R) (~3 m; ~40 kg) partial dorsal rib, several dorsal
rib fragments, partial proximal caudal vertebra (42 mm), incomplete mid caudal
centrum (~41 mm), ?ischial fragment, femoral fragment (~273 mm), partial pedal
ungual ?I/IV, fragments
Comments- Delcourt and Grillo (2014) referred this specimen to Maniraptora
(but not Oviraptorosauria or Alvarezsauridae), and tentatively to Deinonychosauria.
Reference- Delcourt and Grillo, 2014. On maniraptoran material (Dinosauria:
Theropoda) from Vale do Rio do Peixe Formation, Bauru Group, Brazil. Revista
Brasileira de Paleontologia. 17(3), 307-316.
unnamed Deinonychosauria (Franco-Rosas, 2002)
Turonian-Late Maastrichtian, Late Cretaceous
Adamantina, Marilia and/or Serra da Galga Formations of the Bauru Group, Brazil
Material- teeth
Description- These teeth have long serrations with varied slopes and
distal shapes, and slightly pronounced interdenticle slits. They are said to
be similar to troodontids and velociraptorines.
Reference- Franco-Rosas, 2002. Methodological parameters for the identification
and taxonomic classification of isolated theropodomorph teeth. Anais da Academia
Brasileira de Ciencias. 74(2), 367.
undescribed deinonychosaur (Bertini, Santucci and Arruda-Campos, 2001)
Late Maastrichtian, Late Cretaceous
Marilia Formation of the Bauru Group, Brazil
Material- (MPMA-73) tooth
Comments- This was listed as a "dente de teropodomorfo deinonycossauriano", but has not yet
been illustrated or described.
Reference- Bertini, Santucci and Arruda-Campos, 2001. Titanossauros (Sauropoda:
Saurischia) no Cretaceo Superior continental (Formaceo Marilia, Membro Echapora)
de Monte Alto, Estado de S�o Paulo, e correlacao com formas associadas
no Triangulo Mineiro. Geociencias, Sao Paulo. 20(1/2), 93-103.
Jinfengopteryginae Turner, Makovicky and Norell, 2012
Definition- (Jinfengopteryx elegans <- Sinovenator changii,
Troodon formosus, Passer domesticus) (Turner, Makovicky and Norell, 2012)
= "Jinfengopteryginae" Turner, 2008
Comments- Turner (2008; later published as Turner et al., 2012) created
this clade for Jinfengopteryx, IGM 100/1126 and Almas within Troodontidae.
Considering the ease at which Jinfengopteryx
moves in Hartman et al.'s (2019) maniraptoromorph analysis, this was
probably not the best genus to use as an internal specifier.
Currently the most parsimonious place for it in Troodontidae is sister
to Sinovenator, making the clade limited to Jinfengopteryx itself in that case or when it is a basal deinonychosaur.
References- Turner, 2008. Phylogenetic relationships of paravian Theropods.
PhD Thesis. Columbia University. 666 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
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
Jinfengopteryx Ji, Ji, Lu, You,
Chen, Liu and Liu, 2005
J. elegans Ji, Ji, Lu, You, Chen, Liu and Liu, 2005
Early Aptian, Early Cretaceous
Qiaotou Member of the Huajiying Formation, Hebei, China
Holotype- (CAGS-IG-04-0801) (548 mm) skull (~68 mm), sclerotic plates,
mandible, hyoids, twelve cervical vertebrae, cervical ribs, eleven dorsal vertebrae,
dorsal ribs, gastralia, twenty-four caudal vertebrae (273 mm), chevrons, scapulae
(47.09 mm), coracoid, furculae, humeri (49.22 mm), radii (42.44 mm), ulnae (43.31
mm), scapholunare, semilunate carpal, metacarpal I (8.77 mm), phalanx I-1 (18.83
mm), manual ungual I (17.75 mm), metacarpal II (21.37 mm), phalanx II-1 (15.26
mm), phalanx II-2 (21.07 mm), manual ungual II (17.32 mm), metacarpal III (~21
mm), phalanx III-1+III-2 (10 mm), phalanx III-3 (12.95 mm), manual ungual III
(~14 mm), partial ilium (~49 mm), pubes, partial ischium, femora (70.32 mm),
tibiae (100.5 mm), fibula (98.75 mm), proximal tarsus, metatarsus (~59 mm),
pedal digit II, pedal digit III, pedal digit IV, feathers, gastroliths/eggs/seeds
Comments- Jin et al. (2008) reassign Jinfengopteryx's horizon
to the Qiaotou Member of the Huajiying Formation, as opposed to the Qiaotau
Formation which was stated in Ji et al. (2005). Ji et al.'s labeled "pterygoid"
looks to be a parasphenoid rostrum, the "squamosal" part of the parietal.
Turner (2008) redescribed the skull in his thesis.
Though Ji et al. (2005) assign it to the Archaeopterygidae, I first noted a
resemblence to troodontids (Mortimer, DML 2005). Xu and Norell (2006) stated
Jinfengopteryx was a possible troodontid based on "general body
plan" and several undisclosed dental features. It has been more recently
placed in basal Troodontidae (Brusatte et al., 2014; Lee et al., 2014) or basal
Paraves (Foth et al., 2014). Hartman et al. (2019) found an unstable position in Troodontidae further from Troodon than Sinusonasus,
but one step could move it to basal Deinonychosauria, where later
alterations of the matrix placed it. Three steps move it to
Archaeopterygidae, while 4 steps move it to basal Paraves.
References- Ji, Ji, Lu, You, Chen, Liu and Liu, 2005. First avialan bird
from China (Jinfengopteryx elegans gen. et sp. nov.). Geological Bulletin
of China. 24(3), 197-205.
Mortimer, DML 2005. https://web.archive.org/web/20190918121926/http://dml.cmnh.org/2005Mar/msg00372.html
Xu and Norell, 2006. Non-avian dinosaur fossils from the Lower Cretaceous Jehol
Group of western Liaoning, China. Geological Journal. 41(3-4), 419-437.
Ji and Ji, 2007. Jinfengopteryx compared to Archaeopteryx, with
comments on the mosaic evolution of long-tailed avialan birds. Acta Geologica
Sinica (English Edition). 81(3), 337-343.
Jin, Zhang, Li, Zhang, Li and Zhou, 2008. On the horizon of Protopteryx
and the early vertebrate fossil assemblages of the Jehol Biota. Chinese Science
Bulletin. 53(18), 2820-2827.
Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis.
Columbia University. 666 pp.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
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 clade (Archaeopteryx lithographica + Dromaeosaurus albertensis)
Imperobator Ely and Case, 2019
I. antarcticus Ely and Case, 2019
Early Maastrichtian, Late Cretaceous
Upper Cape Lamb Member of the Snow Hill Island Formation, James Ross Island, Antarctica
Holotype-
(UCMP 276000) (~3-4 m) premaxillary, maxillary and/or dentary
fragments, more than two teeth, caudal vertebra, distal tibia (~60 mm
trans), (?)tibial fragment, distal fibulae fused to calcanea, medial
astragalus, distal tarsal III, metatarsal I (?), pedal ungual I (?),
partial metatarsals II, proximal phalanx II-1, partial pedal ungual II,
partial metatarsals III, proximal phalanx III-1, distal phalanx III-1,
fragmentary phalanx ?III-?, partial metatarsal IV, phalanx IV-1 (~61
mm), fragmentary phalanx ?IV-?, fragmentary pedal ungual IV, metatarsal
V (?), pedal elements
....(AMNH FARB 30894) fragmentary (?)cranial elements, partial tooth,
partial pedal ungual, several fragments (Lamanna, Case, Roberts,
Arbour, Ely, Salisbury, Clarke, Malinzak, West and O'Connor, 2019)
Comments- This was discovered
in December 2003 and first reported by Martin and Case (2005) who
stated it was a theropod where "the foot structure seems very primitive
for a carnivorous dinosaur existing at the end of the Mesozoic", which
"does not appear to be related to North American dinosaurs" and instead
"appears to be a primitive holdover of the original Gondwanan dinosaur
assemblage." Case et al. (2007) described the specimen, placing
it in Dromaeosauridae and stating it "more closely resembles Early
Cretaceous dromaeosaurs such as Deinonychus and Utahraptor rather than contemporaneous dromaeosaurids, Velociraptor and Dromaeosaurus or regional close (i.e. South American) species like Neuquenraptor ... or Buitreraptor"
and "may in fact be a latest Cretaceous remnant of the Early
Cretaceous, cosmopolitan, basal stock of dromaeosaurids." While
this echos Martin and Case's idea, Deinonychus and Utahraptor have universally been regarded as closer to Velociraptor and Dromaeosaurus
than unenlagiines. Their reasons for placing it basally are "the
juxtaposition of the ascending process of the astragalus and the
anterior distal tibia without a well-defined fossa on the tibia" which
is true for almost all pennaraptorans and "the incipient ginglymoidal
MtII/digit II joint" which is indeed unlike dromaeosaurids and
unenlagiines, as also noted by Turner et al. (2012) who thus assigned
it to Deinonychosauria incertae sedis. Ely and Case (2016)
recovered the taxon as the most basal deinonychosaur, sister to
troodontids plus dromaeosaurids, in an unpublished analysis. Ely
and Case (2019) later officially described and named the specimen,
including it in Gianechini et al.'s TWiG analysis where it fell out in
Paraves in a polytomy with eudromaeosaurian and anchiornithine
OTUs. While the authors seem to favor a basal position as in
their 2016 abstract, their analyses suggests an anchiornithine
troodontid or eudromaeosaurian dromaeosaurid placement are about
equally supported. Lamanna et al. (2019) state "Several of the
present authors are
currently undertaking a comprehensive reassessment of the morphology
and phylogenetic rel ationships of Imperobator
that will include a
description of all known material of this taxon." Entering data
from Ely and Case (2019) into the Hartman et al. maniraptoromorph
analysis indicates a deinonychosaurian placement more derived than Jinfengopteryx and outside Unenlagiinae and Troodontidae plus Dromaeosauridae.
The materials list has been inconsistent between publications, with
Case et al. describing "two teeth that were preserved in a fragment of
concretion", Ely and Case (2019) mentioning no cranial material, but
Lammana et al. (2019) reporting Case and Malinzak relocated "additional
material pertaining to UCMP 276000 at facilities of Eastern Washington
University and the South Dakota School of Mines and
Technology, respectively, including skull fragments (probably belonging
to at least the premaxilla, maxilla, and/or dentary), a caudal
vertebra, and additional teeth and pedal elements", similar to Lammana
et al. (2017)'s statement "undescribed craniodental fragments of this
theropod individual collected during [2003], initially thought missing,
were recently relocated in the collections of the South Dakota School
of Mines and Technology." Case et al. state "the left tibia is
the most complete of the two tibia fragments" but Ely and Case state
"only the distal left tibia is preserved." Ely and Case state
"potential material from digit I may be present (Fig. 7D). It is
distinguished by what may be a prominent flexor heel on the
proximoventral surface, morphologically similar to that of the
dromaeosaurid (avialan?) Balaur bondoc (Csiki et al. 2010)",
implying a pedal ungual I, but 7D seems to be the proximal half of a
non-ungual phalanx positioned as III-1 in Case et al.. In the
materials list, Ely and Case also state "material from metatarsal I,
and even metatarsal V may be preserved", but we get no description or
illustration of these. While the illustrated and described pedal
material is from the left pes, Ely and Case state "only a few fragments
from the right pes" are preserved, of which the only elements specified
in either paper are "distal ends of metatarsals II and III" by Case et
al.. Note in Case et al.'s photo of the left pes, distal
metatarsal IV and phalanx IV-I figured by Ely and Case are missing, but
three small fragments positioned as one part of digit III and two parts
of digit IV are shown by Case et al. but not mentioned by Ely and Case
except perhaps when they say "a small portion of the ungual [IV?] is
also present."
Lamanna et al. (2017) report "a partial tooth, possible craniodental
fragments, and part of a pedal ungual" were discovered at the type
locality in 2011 or 2016, belonging to the type individual.
Lamanna et al. (2019) confirms these were found in both years,
"including a tooth and several bone fragments (AMNH FARB 30894), a
partial pedal ungual, and fragmentary putative cranial remains."
References- Martin and Case, 2005. Fossil hunting in Antarctica. Geotimes. February 2005, 18-21.
Case, Martin and Reguero, 2007. A dromaeosaur from the Maastrichtian
of James Ross Island and the Late Cretaceous Antarctic dinosaur fauna. In Cooper
and Raymond (eds.). Online
Proceedings of the 10th ISAES X. USGS Open-File Report 2007-1047, Short Research
Paper 083, 4 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Ely and Case, 2016. A basal deinonychosaur from the Early
Maastrichtian, Antarctic peninsula and the biostratigraphy of the
latest Cretaceous dinosaur fauna of Antarctica. Journal of Vertebrate
Paleontology. Program and Abstracts 2016, 130.
Lamanna, O'Connor, Salisbury, Gorscak, Clarke, MacPhee, Roberts,
Malinzak, Ely and Case, 2017. New material of non-avian dinosaurs from
the Late Cretaceous of James Ross Island, Antarctica. Journal of
Vertebrate Paleontology. Program and Abstracts 2017, 147.
Ely and Case, 2019. Phylogeny of a new gigantic paravian (Theropoda; Coelurosauria;
Maniraptora) from the Upper Cretaceous of James Ross Island, Antarctica. Cretaceous Research. 101, 1-16.
Lamanna, Case, Roberts, Arbour, Ely, Salisbury, Clarke, Malinzak, West
and O'Connor, 2019. Late Cretaceous non-avian dinosaurs from the James
Ross Basin, Antarctica: Description of new material, updated synthesis,
biostratigraphy, and paleobiogeography. Advances in Polar Science.
30(3), 228-250.
Archaeopterygidae Huxley, 1871
Definition- (Archaeopteryx lithographica <- Unenlagia comahuensis, Dromaeosaurus albertensis, Troodon formosus, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019)
Other definitions- (Archaeopteryx lithographica <- Dromaeosaurus
albertensis, Passer domesticus) (Xu, You, Du and Han, 2011)
(Archaeopteryx lithographica <- Passer domesticus) (Sereno, online 2005)
= Sauriurae Haeckel, 1866
= Saururae Huxley, 1867
= "Archornithidae" Carus, 1875
= Saururi Vogt, 1879
= Ornithopappi Stejneger, 1885
= Archaeopteryges Furbringer, 1888
= Archaeopterygiformes Furbringer, 1888
= Archornithes Furbringer, 1888
= Saurura Steinmann and Doederlein, 1890
= Archornithiformes Shefeldt, 1903
= Archaeornithidae Petronievics, 1925
= Archaeopterygomorphi Hay, 1930
Ex-archaeopterygids- Several taxa have been referred to Archaeopterygidae
or Archaeopteryx itself in the past, but do not belong there. Janensch
(1914) first noted an Archaeopteryx-like supposed carpometacarpus (HMN
coll.) from the Tendaguru Formation of Tanzania, but this is based on a misreading
of Stremme (1916-1919), who was describing differences from Archaeopteryx.
This was named Stremmeia by Nopcsa (1930) and reidentified as a salientian
tibiofibulare, and while this identification has been doubted by late twentieth
century authors, the specimen is also highly dissimilar to maniraptoran elements.
Lambrecht (1933) noted a specimen labeled Archaeopteryx "vicensensis"
at the Museum zu Vicensa, but found it was a pterosaur after writing Kleinschmidt.
Jensen (1981) identified a proximal femur (BYU 2023) from the Morrison Formation
as Archaeopteryx, but this is placed as Ornithodesmiformes incertae
sedis here. Kessler and Jurcsak (1984) described an incomplete supposed humerus (MTCO 14422)
from the Early Cretaceous of Romania as Archaeopteryx sp., but this was
reidentified by Dyke et al. (2011) as a long bone of Archosauria indet.. Paul
(1988) referred all dromaeosaurids to Archaeopterygidae, which has not been
recovered in any phylogenetic analysis. "Proornis" was originally
called "the North Korean Archaeopteryx" (e.g. Anonymous, 1994),
but is actually a confuciusornithid (Gao et al., 2009). Weigert (1995) described
103 teeth from the Guimarota Formation of Portugal as cf. Archaeopteryx sp.,
but they may belong to a basal deinonychosaur instead. Protarchaeopteryx
was assigned to the family by Ji and Ji (1997) and Paul (2002), but is probably a basal
oviraptorosaur. Forster et al. (1998) found Rahonavis and Unenlagia
to clade with Archaeopteryx
in some most parsimonious trees, but these are now considered members
of their own clade Unenlagiidae which is closer to either
dromaeosaurids or birds. Rauhut (2002) referred Paronychodon to Archaeopterygidae, but this was based
on comparisons to the Guimarota teeth noted above and the genus is here assigned
to Troodontidae based on enamel microstructure. Ji et al. (2005) described Jinfengopteryx
as being more closely related to Archaeopteryx than to Aves, but is now
recognized as a basal troodontid or basal paravian. Xu et al. (2011) recently
used a version of Senter's TWiG analysis to refer Xiaotingia to Archaeopterygidae,
but the genus has also been placed as a basal troodontid and basal dromaeosaurid.
Sauriurae- Sauriurae was a group first used by Haekel (1866) for Archaeopteryx,
who placed modern birds in the Ornithurae instead. This taxonomy was followed
for over a century, with hesperornithines and ichthyornithines being added to
Ornithurae by later authors. Martin (1983) was the first author to place enantiornithines
in Sauriurae, which has been followed near universally by those who doubt the
dinosaur-bird relationship. Additional taxa have also been assigned to Sauriurae
including Confuciusornis (Hou et al., 1995), Sinosauropteryx (Ji
and Ji, 1996), Protarchaeopteryx (Ji and Ji, 1997), Yandangornis
(Cai and Zhao, 1999), Caudipteryx (Martin and Czerkas, 2000), Jeholornis
(Martin, 2004), Vorona (Kurochkin, 2006), and lately all deinonychosaurs
and oviraptorosaurs (Martin, 2004). The group has basically consisted of any
non-euornithine birds considered by the authors, with the notable exception
of Kurochkin (2006), who places Protoavis and confuciusornithids in Euornithes (his Ornithurae)
and views sauriurines as being theropods while euornithurines are not. While there
may be some evidence for placing confuciusornithids and enantiornithines in
a group exclusive of Aves, phylogenetic analyses are unanimous in rejecting
a clade of Archaeopteryx and enantiornithines which excludes Aves.
References- Haekel, 1866. Generelle Morphologie der Organismen: allgemeine
Grundz�ge der organischen Formen-Wissenschaft, mechanisch begr�ndet
durch die von Charles Darwin reformirte Descendenz-Theorie. Berlin: Reimer.
462 pp.
Huxley, 1867. On the classification of birds; and on the taxonomic value of
the modifications of certain of the cranial bones observable in that class.
Proceedings of the Zoological Society of London. 35, 415-472.
Carus, 1875. Handbuch der Zoologie. Erster Band: Wirbelthiere, Mollusken und
Molluskoiden. 894 pp.
Vogt, 1879. L'Archaeopteryx macroura, Un intermediaire entre les oiseaux
et les reptiles. La Revue Scientifique. 9(2), 241-248.
Stejneger, 1884. Classification of Birds. Science Record. 2(7), 154-155.
Furbringer, 1888. Untersuchungeb zur Morphologie und Systematik der Vogel. Amsterdam:
Holkema. 1751 pp.
Steinmann and Doederlein, 1890. Elemente der pal�ontologie bearbeitet.
Leipzig. 848 pp.
Shufeldt, 1903. On the classification of certain groups of birds. The American
Naturalist. 37, 33-64.
Janensch, 1914. Ubersicht uber die Wirbeltierfauna der Tendaguru-Schichten.
Archiv fur Biontologie. 3, 81-110.
Stremme, 1916-1919. Uber die durch Bandverknocherung hervorgerufene proximale
Verschmelzung zweier Mittelhand - oder Mittelfussknochen eines Reptils. Wissenschaftliche
Ergebnisse der Tendaguru-Expedition. Archiv fur Biontologie. 4, 143-144.
Petronievics, 1925. Uber die Berliner Archaeornis. Ann. Geol. Peninsule
Balkan. 8(1), 1-52.
Hay, 1930. Second bibliography and catalogue of the fossil Vertebrata of North
America. Publication 390 of the Carnegie Institute of Washington, Volume 2.
Nopcsa, 1930. Notes on Stegocephalia and Amphibia. Proceedings of the Zoological
Society of London. 1930, 979-995.
Lambrecht, 1933. Handbuch der Palaeornithologie. 1024 pp.
Brodkorb, 1963. Catalogue of fossil birds. Part 1 (Archaeopterygiformes through
Ardeiformes). Bull. Florida State Mus., Bioi. Sci.. 7, 179-293.
Jensen, 1981. Another look at Archaeopteryx as the worlds oldest bird.
Encyclia, The Journal of the Utah Academy of Sciences, Arts, and Letters. 58,
109-128.
Kessler and Jurcs�k, 1984. Fossil birds remains in the bauxite from Cornet
(Pa�durea Craiului Mountains, Romania). 75 years of the Laboratory of Paleontology,
University of Bucharest, Romania, Special Volume. 129-134.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster Co., New York.
464 pp.
Anonymous, 1994. Korean Pictorial. 1994(2).
Hou, Zhou, Gu and Zhang, 1995. Confuciusornis sanctus, a new Late Jurassic
sauriurine bird from China. Chinese Science Bulletin. 40(18), 1545-1551 [in
Chinese].
Weigert, 1995. Isolierte Zahne von cf. Archaeopteryx sp. aus dem Oberen
Jura der Kohlengrube Guimarota (Portugal). N. Jb., Geol. Palaont. Mh. 9, 562-576.
Ji and Ji, 1996. On discovery of the earliest bird fossil in China and the origin
of birds. Chinese Geology. 233, 30-33.
Ji and Ji, 1997. A Chinese archaeopterygian, Protarchaeopteryx gen. nov..
Geological Science and Technology. 238, 38-41.
Forster, Sampson, Chiappe and Krause, 1998. The theropod ancestry of birds:
New evidence from the Late Cretaceous of Madagascar. Science. 279, 1915-1919.
Cai and Zhao, 1999. A long tailed bird from the Late Cretaceous of Zhejiang.
Science in China (series D). 42(4), 434-441.
Martin and Czerkas, 2000. The fossil record of feather evolution in the Mesozoic.
American Zoologist. 40(4), 687-694.
Paul, 2002. Dinosaurs of the Air. The Johns Hopkins University Press, Baltimore.
460 pp.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca,
Spain. Cretaceous Research. 23, 255-263.
Martin, 2004. A basal archosaurian origin for birds. Acta Zoologica Sinica.
50(6), 978-990.
Ji, Ji, Lu, You, Chen, Liu and Liu, 2005. First avialan bird from China (Jinfengopteryx
elegans gen. et sp. nov.). Geological Bulletin of China 24(3): 197-205.
Sereno, online 2005. Stem Archosauria - TaxonSearch. http://www.taxonsearch.org/dev/file_home.php
[version 1.0, 2005 November 7]
Kurochkin, 2006. Parallel evolution of theropod dinosaurs and birds. Entomological
Review. 86(suppl. 1), S45-S58.
Gao, Li, Wei, Pak and Pak, 2009. Early Cretaceous birds and pterosaurs from
the Sinuiju series, and geographic extension of the Jehol biota into the Korean
peninsula. Journal of the Paleontological Society of Korea. 25(1), 57-61.
Dyke, Benton, Posmosanu and Naish, 2011. Early Cretaceous (Berriasian) birds
and pterosaurs from the Cornet bauxite mine, Romania. Palaeontology. 54(1),
79-95.
Xu, You, Du and Han, 2011. An Archaeopteryx-like theropod from China
and the origin of Avialae. Nature. 475, 465-470.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Vernon, New Jersey. Pan Aves. 189 pp.
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
Ostromia Foth and Rauhut, 2017
O. crassipes (Meyer, 1857) Foth and Rauhut, 2017
= Pterodactylus crassipes Meyer, 1857
= Rhamphorhynchus crassipes (Meyer, 1857) Meyer, 1857
= Archaeopteryx crassipes (Meyer, 1857) Ostrom, 1972a
= Scaphognathus crassipes (Meyer, 1857) Olshevsky, 1991
= Archaeornis crassipes (Meyer, 1857)
B�hler and Bock, 2002
Early Tithonian, Late Jurassic
Solnhofen Formation, Germany
Holotype-
(TM 6928; TM 6929; Haarlem specimen; Teyler specimen; fourth specimen of Archaeopteryx; holotype of
Pterodactylus crassipes) (350 day old juvenile) three or four posterior dorsal centra (6.5 mm), dorsal
rib fragments, gastralia, distal humerus, incomplete radii, incomplete ulnae, semilunate carpal, metacarpal I (10.5
mm), phalanx I-1 (23.1 mm), manual ungual I (9.5 mm), incomplete metacarpal
II, distal manual ungual II, incomplete metacarpal III (23.2 mm),
distal phalanx III-3, manual ungual III (8.3 mm), manual claw sheaths, distal pubis, distal femora (~54 mm), proximal
tibiae (~80 mm), proximal fibula, fibular fragment, phalanx I-1, partial pedal ungual I, distal
metatarsal II, phalanges II-1 (one distal), phalanges II-2, pedal unguals II, distal
metatarsal III (~48 mm), phalanges III-1, phalanx III-2, phalanx III-3, pedal
unguals III (7.8 mm), distal metatarsal IV, phalanx IV-2, phalanx IV-3, phalanx IV-4,
pedal ungual IV, two pedal phalanges, remiges
Diagnosis- (after Foth and
Rauhut, 2017) longitudinal furrows on both sides of all preserved
manual phalanges and at least metacarpal III (also in Anchiornis and Eosinopteryx); differs from Anchiornis and Eosinopteryx
in longer metacarpal I (45% of mcIII length vs. 41% and 42%), manual
ungual I shorter than metacarpal I, and longer metatarsus (~60% of
tibial length vs. 52% and 51%)
Other diagnoses- Meyer (1857)
initially identified the specimen as a pterosaur and distinguished it
from other known pterosaurs by the shorter metacarpals I-III, longer
manual digits I-III, longer metatarsus and smaller unguals.
Comments- Discovered in 1855, Meyer (1857) named TM 6928/6929 Pterodactylus crassipes, finding the manual proportions and ungual size most similar to Dimorphodon (then Pterodactylus macronyx). Due to incompleteness, Meyer wrote "it would not be impossible that Pterodactylus crassipes also represented a Rhamphorynchus, which will decide on the attachment of the head or tail. If the assumption is confirmed, the species would be called Rhamphorynchus crassipes."
As ICZN Article 11.5.1 states "A name proposed conditionally for a
taxon before 1961 is not to be excluded on that account alone", this is
an available name as well. Meyer (1859/1860) described it in more
detail and illustrated a slab. The only other author to consider the
specimen was Wellnhofer (1970), who believed it to be a
rhamphorhynchoid based on "the
short metacarpus, the long metatarsus, the shape of the prebubis and
the strikingly large claws on the hands and feet." Furthermore, he stated "the comparison of "Pt." crassipes with Scaphognathus crassirostris
reveals broad similarities both in size and morphology of individual
skeletal elements (prebubis, femur, claws). Although the existing
remains of "Pt." crassipes are not sufficient to make a specific association with Scaphognathus crassirostris, both specimens may certainly be considered congeneric" and on page 122 reidentified crassipes as Scaphognathus sp.. While the combination Scaphognathus crassipes
could be implied from Wellnhofer's statement, it was not used
explicitly until Olshevsky (1991) who listed it without attribution.
Ostrom
(1970) identified the specimen as Archaeopteryx cf. lithographica,
stating "the present remains are much too fragmentary for positive
species identification" before describing it in detail two years later
(1972b) as A. lithographica.
His in depth study concluded "With the exception of the apparent
difference in pubis position, the Teyler specimen does not differ in
any other significant way from the London and Solnhofen specimens of Archaeopteryx.
The limb elements are slightly more robust and apparently were slightly
longer than those of the Berlin specimen. In summary, I see no
morphologic evidence to exclude this specimen from the same taxon as
the other three." This was problematic because "the specific name crassipes is a senior synonym of lithographica", yet "the species name A. lithographica
is well known and firmly established in the zoologic literature. It
therefore seems undesirable to suppress that term in favor of the
senior subjective synonym." Thus Ostrom (1972a) petitioned the ICZN to
"(a) use its plenary powers to suppress the species name crassipes Meyer, 1857, as published in the binomen Pterodactylus crassipes, for purposes of the Law of Priority but not for those of the Law of Homonymy : (b) place the specific name crassipes
(as suppressed under the plenary powers in (a) above) on the Official
Index of Rejected and Invalid Specific Names in Zoology." Nye (1974)
and Eisenmann (1974) wrote Comments to the ICZN arguing that cases of
subjective synonymy such as this, the senior synonym should not be
completely suppressed in case it is later thought to be a distinct
taxon. Ostrom and the ICZN agreed, so that in 1977 Melville issued the
narrower Opinion 1070 that "the specific name lithographica von Meyer, 1861, as published in the binomen Archaeopteryx lithographica, is to be given precedence over the specific name crassipes von Meyer, 1857, as published in the binomen Pterodactylus crassipes by any zoologist who believes that the two specific names apply to the same taxon."
Most recently, Foth and Rauhut (2017) reinterpreted crassipes
as an anchiornithid based on longitudinally grooved manual elements and
pubic curvature, with proportional differences between it and Archaeopteryx, Anchiornis and Eosinopteryx supporting its validity as the new genus Ostromia. They and Hartman et al. (2019) included Ostromia
in analyses based on the TWiG and recovered it closest to
anchiornithines, Foth and Rauhut in a polytomy with included taxa, and
Hartman et al. in a polytomy with Serikornis and other archaeopterygids (including anchiornithines in their topology).
References- Meyer, 1857.
Beitr�ge zur n�heren Kenntniss fossiler Reptilien. Neues Jahrbuch f�r
Mineralogie, Geognosie, Geologie und Petrefakten-Kunde. 1857, 532-543.
Meyer, 1859/1860. Zur Falma der Vorwelt. Vierte Abteilung: Reptilien
aus dem lithographischen Schiefer des Jura in Deutschland und
Frankreich. Frankfurt. 144 pp.
Ostrom, 1970. Archaeopteryx: notice of a 'new' specimen. Science. 170,
537-538.
Wellnhofer, 1970. Die Pterodactyloidea (Pterosauria) der Oberjura-Plattenkalke
S�ddeutschlands. Abhandlung der Bayerischen Akademie der Wissenschaften,
Neue Folge. 141, 133 pp.
Ostrom, 1972a. Pterodactylus crassipes Meyer, 1857 (Aves): Proposed suppression
under the plenary powers. Z.N.(S.) 1977. Bulletin of Zoological Nomenclature. 29(1), 30-31.
Ostrom, 1972b. Description of the Archaeopteryx specimen in the Teyler
Museum, Haarlem. Proceedings Koninklijke Nederlandse Akademie Van Wetenschappen,
B. 75, 289-305.
Eisenmann, 1974. Comment on proposal to suppress Pterodactylus crassipes Meyer, 1857 and counter-proposal to recognize Archaeopteryx lithographica Meyer, 1861, and to fix its type-species. Z.N.(S.) 1977. Bulletin of Zoological Nomenclature. 31(3), 114-115.
Nye, 1974. Comment on: (B)--Proposed supression of Pterodactylus crassipes Meyer (Aves). Z.N.(S.) 1977. Bulletin of Zoological Nomenclature. 30(3/4), 140-141.
Melville, 1977. Opinion 1070. Conservation of Archaeopteryx lithographica von Meyer, 1861 (Aves).
Bulletin of Zoological Nomenclature. 33(3/4), 165-166.
Olshevsky, 1991. A Revision of the Parainfraclass Archosauria Cope,
1869, Excluding the Advanced Crocodylia. Mesozoic Meanderings. 2, 196
pp.
B�hler and Bock, 2002. Zur Archaeopteryx-Nomenklatur: Mi�verst�ndnisse
und L�sung. Journal f�r Ornithologie. 143, 269-286.
Wellnhofer, 2009. Archaeopteryx: The Icon of Evolution. Verlag Dr. Friedrich Pfeil. 208 pp.
Foth and Rauhut, 2017. Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs. BMC Evolutionary Biology. 17:236.
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
Serikornis Lefevre, Cau, Cincotta, Hu, Chinsamy, Escuillie and Godefroit, 2017
S. sungei Lefevre, Cau, Cincotta, Hu, Chinsamy, Escuillie and Godefroit, 2017
Oxfordian, Late Jurassic
Daxishan, Tiaojishan Formation, Liaoning, China
Holotype-
(PMOL-AB00200) (490 mm subadult) incomplete skull, incomplete
mandibles, hyoids, nine cervical vertebrae (anterior 8.1, posterior 9.9
mm), cervical ribs, six dorsal vertebrae (anterior 6.7, posterior 6.6
mm), dorsal
ribs, synsacrum, twenty-seven caudal vertebrae (proximal 7.7, mid 13,
distal 10.8 mm),
chevrons, scapula (37.8 mm), coracoids, furcula, humeri (one
incomplete; 60.7 mm), radii (one incomplete; 50.5 mm),
ulnae (one incomplete; 50.8 mm), semilunate carpal, metacarpals I (12 mm),
phalanges I-1 (one incomplete; 27.5 mm), manual ungual I (17 mm),
metacarpals II (one incomplete; 32 mm),
phalanges II-1 (20 mm), phalanges II-2 (one proximal; 23.5 mm), manual
unguals II (19 mm), metacarpals III (one incomplete; 30 mm),
phalanx III-1 (7.5 mm), phalanx III-2 (6.5 mm), phalanges III-3 (16
mm), manual unguals III (one proximal; 12.7 mm), ilium (33 mm), distal
pubes (63 mm), ischia (~16 mm), femora (67.4 mm), tibiae (tibiotarsus
95.2 mm), fibulae,
proximal
tarsals, metatarsals I (8.5 mm), phalanges I-1 (9 mm), pedal unguals I
(3 mm), metatarsals II (48.5 mm),
phalanges II-1 (9 mm), phalanges II-2 (11 mm), pedal unguals II (13
mm), metatarsals III (48.5 mm),
phalanges
III-1 (12 mm), phalanges III-2 (10 mm), phalanges III-3 (10 mm),
pedal ungual III (5.5 mm), metatarsals IV (53 mm), phalanges IV-1 (9
mm), phalanges IV-2 (8 mm), phalanges IV-3 (6 mm), phalanges
IV-4 (7 mm), pedal unguals IV (8 mm), metatarsals V (15.5 mm), body
feathers, remiges, metatarsal remiges
Diagnosis- (after Lefevre et
al., 2017) four anterior maxillary teeth twice as long as the others;
distal end of posteroventral coracoid process thicker than proximal
part to form a rounded bump; distal tip of ischium narrow and strongly
deflected dorsally to form a hook.
Other diagnoses- Lefevre et al.
(2017) listed "coracoid tuber well-developed and laterally projected
from the lateral margin of the coracoid and forming a subglenoid shelf
along the caudoventral margin of the bone" as diagnostic compared to Anchiornis, but this is due to perspective and so is true in Anchiornis
specimens with more posteriorly exposed coracoids (e.g. BMNHC
Ph 822). The "smooth ventral side of coracoid devoid of small
pits" is primitive compared to Anchiornis.
Comments- The holotype is one
of several paravian specimens acquired by the YFGP from a fossil dealer
prior to June 2012. It was labeled YFGP-T5201 in the poster of
Brougham (2013, online), which is probably a mistake (see entry for
YFGP-T5201).
Lefevre et al. (2017) assigned a subadult stage to the holotype based
on a mix of juvenile ("bone extremities are rough in appearance"; "an
uneven periosteal surface which indicates that periosteal growth was
still occurring at the time of death") and adult (completely fused
sacral vertebrae; completely fused caudal neurocentral sutures)
features.
Brougham (2013, online) recovered the specimen in a clade with Aurornis and Eosinopteryx using Turner's dromaeosaurid and Xu's Xiaotingia TWiG matrices and Cau's Aurornis matrix. Lefevre et al. (2017) used a version of Cau's matrix to recover this as a basal paravian sister to Eosinopteryx, in a clade with Aurornis and Pedopenna. Hartman et al. (2019) found it to be the most basal archaeopterygid using a composite TWiG analysis.
References- Brougham, 2013. Multi-matrix analysis of new Late Jurassic feathered
theropods from China supports troodontid-avialan clade. Symposium on
Vertebrate Palaeontology and Comparative Anatomy, Programme and
Abstracts. 49.
Brougham, 2013 online. https://www.researchgate.net/publication/280728942_SVPCA_Poster
Lefevre, Cau, Cincotta, Hu, Chinsamy, Escuillie and Godefroit, 2017. A
new Jurassic theropod from China documents a transitional step in the
macrostructure of feathers. The Science of Nature. 104:74.
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
Pedopenna Xu and Zhang,
2005
P. daohugouensis Xu and Zhang, 2005
Bathonian-Callovian, Middle Jurassic
Daohugou, Haifanggou Formation, Inner Mongolia, China
Holotype- (IVPP V12721) (<1 m) distal tibiotarsus, distal fibula,
metatarsal I, phalanx I-1 (8.6 mm), pedal ungual I (~7.6 mm), metatarsal II
(57.6 mm), phalanx II-1 (11.6 mm), phalanx II-2 (14.2 mm), pedal ungual II (~13
mm), metatarsal III (~57.8 mm), phalanx III-1 (~18 mm), phalanx III-2 (~11.3
mm), phalanx III-3 (~11.3 mm), pedal ungual III (10.1 mm), metatarsal IV (57.4
mm), phalanx IV-1 (11.3 mm), phalanx IV-2 (9.9 mm), phalanx IV-3 (6.2 mm), metatarsal
V (14.1 mm), feathers, scale impressions, pedal claw sheath
Diagnosis- (from Xu and Zhang, 2005) very slender pedal phalanx I-1 (length/mid-shaft-diameter
ratio about 7.2).
Comments- Recovered in 2002 from the Daohugou beds, which
have more recently been assigned to the Haifanggou Formation (e.g. Xu
et al., 2016).
Xu and Zhang (2005) recovered Pedopenna as a paravian
in a trichotomy with avialans and deinonychosaurs using a version of the TWiG
matrix. More recently, Brusatte et al. (2014) found it to be a scansoriopterygid,
while Foth et al. (2014) found it to be a basal avialan in a clade with Anchiornis
and Eosinopteryx. Hartman et al. (2019) recovered it as a scansoriopterygid, but only one step moved it to Archaeopterygidae.
References- Xu and Zhang, 2005. A new maniraptoran dinosaur from China
with long feathers on the metatarsus. Naturwissenschaften. 92, 173-177.
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.
Xu, Zhou, Sullivan, Wang and Ren, 2016. An updated review
of the Middle-Late Jurassic Yanliao Biota: Chronology, taphonomy, paleontology
and paleoecology. Acta Geologica Sinica. 90(6), 2229-2243.
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
Caihong Hu, Clarke, Eliason, Qiu, Li, Shawkey, Zhao, D'Alba, Jiang and Xu, 2018
C. juji Hu, Clarke, Eliason, Qiu, Li, Shawkey, Zhao, D'Alba, Jiang and Xu, 2018
Oxfordian, Late Jurassic
Gangou (= Zhuanshanzi), Tiaojishan Formation, Hebei, China
Holotype-
(PMOL-B00175) (400 mm, ~475 g, adult) skull (~67.6 mm), mandibles,
hyoids, (cervical series ~72 mm) about ten cervical vertebrae, cervical
ribs, (dorsal series ~87.8 mm) about twelve dorsal vertebrae, dorsal
ribs, gastralia, five sacral vertebrae (~31.5 mm), (caudal series ~178
mm) first to twenty-sixth
caudal vertebrae, chevrons, scapulae, coracoids, furcula, humeri (one
proximal; 42.1 mm), radii (one distal), ulnae (one distal; 47.2 mm),
metacarpals I (8.6, 9.3 mm), phalanges I-1 (18.2, 21.8 mm), manual
ungual I (7.5 mm), metacarpals II (23.2, 23.7 mm),
phalanges II-1 (11.6, 11.8 mm), phalanges II-2 (19.9, 20 mm), manual
ungual II (10.6 mm), metacarpals III (23.2, 23.5 mm),
phalanges III-1 (5.4, 5.9 mm), phalanges III-2 (5.7, 5.8 mm), phalanges
III-3 (13.3, 13 mm), manual unguals III (7.8, 7.8 mm), partial ilium
(31 mm), pubes (54.9 mm), ischium (20.5 mm), femora (70.9 mm), tibiae
(82.8, 81.6 mm), proximal
tarsals, metatarsal I (5.5 mm), phalanx I-1 (4.2 mm), pedal ungual I
(4.4 mm), metatarsals II (47.3, 47.6 mm),
phalanges II-1 (~8, 8.7 mm), phalanges II-2 (9.6 mm), pedal unguals II
(10.5 mm), metatarsal III (49 mm), phalanges
III-1 (12.2 mm), phalanges III-2 (8.4, 8 mm), phalanges III-3 (~8.9,
8.9 mm), pedal unguals III,
metatarsal IV (46.6 mm), phalanx IV-1 (8.2 mm), phalanx IV-2 (7.2 mm),
phalanx IV-3 (5.7 mm), phalanges
IV-4 (6.4, 6.2 mm), pedal unguals IV (8.3 mm), body feathers, remiges
(~97 mm), metatarsal remiges (~31 mm), retrices (112 mm)
Diagnosis- (after Hu et al.,
2018) accessory fenestra posteroventral to promaxillary fenestra;
lacrimal with prominent dorsolaterally oriented crests; robust dentary
with anterior tip dorsoventrally
deeper than its midsection; short ilium (<50% of the femoral length).
Comments- Collected by a farmer, the holotype was acquired by the PMOL in February 2014.
Hu et al. (2018) recovered Caihong as an anchiornithid closest to Xiaotingia
in Xu et al.'s 2011 TWiG analysis, and an anchiornithid in Brusatte's
TWiG analysis. Hartman et al. (2019) recovered it as a basal
archaeopterygid (including 'anchiornithines'), but found it moved to a
basal dromaeosaurid in two steps.
References- Hu, Clarke,
Eliason, Qiu, Li, Shawkey, Zhao, D'Alba, Jiang and Xu, 2018. A
bony-crested Jurassic dinosaur with evidence of iridescent plumage
highlights complexity in early paravian evolution. Nature
Communications. 9, 217.
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
Anchiornithinae Xu, Zhou, Sullivan, Wang and Ren, 2016
Definition- (Anchiornis huxleyi <- Archaeopteryx lithographica,
Troodon formosus, Epidexipteryx hui, Gallus gallus, Dromaeosaurus
albertensis, Unenlagia comahuensis) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019; modified after Xu, Zhou, Sullivan, Wang and Ren, 2016)
= Anchiornithidae Xu, Zhou, Sullivan, Wang and Ren, 2016 vide Foth and Rauhut, 2017
Definition- (Anchiornis huxleyi <- Passer domesticus, Archaeopteryx lithographica, Dromaeosaurus albertensis, Troodon formosus, Oviraptor philoceratops) (Foth and Rauhut, 2017)
Other definitions- (Anchiornis huxleyi <- Vultur
gryphus, Archaeopteryx lithographica, Dromaeosaurus albertensis,
Saurornithoides mongoliensis, Unenlagia comahuensis, Epidendrosaurus
ninchengensis) (Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017)
= Anchiornithidae sensu
Cau, Beyrand, Voeten, Fernandez, Tafforeau, Stein, Barsbold,
Tsogtbaatar, Currie and Godefroit, 2017
Definition- (Anchiornis huxleyi <- Vultur
gryphus, Archaeopteryx lithographica, Dromaeosaurus albertensis,
Saurornithoides mongoliensis, Unenlagia comahuensis, Epidendrosaurus
ninchengensis)
References- Xu, Zhou, Sullivan, Wang and Ren, 2016. An updated review
of the Middle-Late Jurassic Yanliao Biota: Chronology, taphonomy, paleontology
and paleoecology. Acta Geologica Sinica. 90(6), 2229-2243.
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.
Foth and Rauhut, 2017. Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs. BMC Evolutionary Biology. 17:236.
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
Anchiornis Xu, Zhao, Norell, Sullivan,
Hone, Erickson, Wang, Han and Guo, 2008
A. huxleyi Xu, Zhao, Norell, Sullivan, Hone, Erickson, Wang, Han
and Guo, 2008
Oxfordian, Late Jurassic
Tiaojishan Formation, Liaoning, China
Holotype- (IVPP V14378) (~340 mm; 110 g; subadult or young adult) posterior
cervical vertebrae (eighth cervical 5 mm), thirteen dorsal vertebrae (second
dorsal 4.3 mm, seventh dorsal 4.8 mm, eleventh dorsal 5.1 mm), nineteen dorsal
ribs, sacrum, seventeen caudal vertebrae (first caudal 2.8 mm, tenth caudal
7.4 mm), chevrons, scapulae (26.8 mm), coracoids, furcula, humeri (41.5 mm),
radius, ulna (37.1 mm), scapholunare, distal carpal III, metacarpal I, carpometacarpus, phalanx
I-1, manual ungual I, phalanx II-1, phalanx II-2, manual ungual II, phalanx
III-1, phalanx III-2, phalanx III-3, manual ungual III, incomplete ilium (~26.2
mm), partial ischium, femora (43.2 mm), tibiae (67.8 mm), metatarsal 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, pedal claw sheaths, feathers
Referred-
(41HIII 0404) (574 mm, adult) partial skull, mandibles (52.7 mm),
hyoids, basihyoid, six cervical vertebrae, cervical ribs,
anterior dorsal vertebrae, sixth to thirteenth dorsal vertebrae, dorsal
ribs, gastralia, (caudal series 290.6 mm) first caudal vertebra (6.1
mm), incomplete second caudal vertebra (6.5 mm), incomplete third
caudal vertebra (7.5 mm), partial fourth caudal vertebra, partial
fifth caudal vertebra, sixth caudal vertebra (9.7 mm), seventh caudal
vertebra (10.8 mm), eighth caudal vertebra (11.3 mm), ninth caudal
vertebra (11.4 mm), tenth caudal vertebra (11.4 mm), eleventh caudal
vertebra (11 mm), twelfth caudal vertebra, thirteenth caudal vertebra,
fourteeenth caudal vertebra, fifteenth caudal vertebra (11.6 mm),
sixteenth caudal vertebra (11 mm), seventeenth caudal vertebra,
eighteenth caudal vertebra (11.3 mm), nineteenth caudal vertebra (11
mm), twentieth caudal vertebra (10.3 mm), twenty-first caudal vertebra
(9.9 mm), twenty-second caudal vertebra (9.2 mm), twenty-third caudal
vertebra (8.2 mm), twenty-fourth caudal vertebra (7.6 mm), twenty-fifth
caudal vertebra (6.9 mm), twenty-sixth caudal vertebra, partial
twenty-seventh caudal vertebra, partial twenty-eighth caudal vertebra,
distal scapulae, ?coracoid fragment, furcula,
humeri (63.2 mm), radii (51 mm), ulnae (52.9, 52.9 mm), scapholunare,
semilunate carpal, metacarpal I (11.2 mm), phalanges I-1 (one partial;
24.4, 24.3 mm), manual ungual I (15.9 mm),
metacarpals II (30.9 mm), phalanges II-1 (one proximal; 19 mm), phalanx
II-2 (24.6 mm), manual unguals II (one incomplete; 16.4, 15.7 mm),
metacarpals III (28.7 mm), phalanges III-1 (7.6, 7.7 mm), phalanges
III-2 (one fragmentary; 6.8, 7.4 mm), phalanges III-3 (12.8, 13 mm),
manual unguals III (13.8 mm), manual claw sheaths, incomplete ilium
(36.3 mm), pubes (one partial; 53.7 mm), ischia (20.8 mm), femora
(65.6, 66.8 mm), tibiae (92.5 mm),
fibulae (90.6 mm), astragali, calcanea, metatarsal I (8.7 mm), phalanx
I-1 (6.6 mm), pedal unguals
I (6.2 mm), metatarsals II (49.2, 48.3 mm), phalanx II-1 (11.2 mm),
phalanges II-2 (11.3 mm), pedal unguals II,
metatarsals III (52.8, 49.5 mm), phalanges III-1 (13.4 mm), phalanges
III-2 (10.1, 10.8 mm), phalanx III-3 (9.4 mm), pedal ungual III (13
mm), metatarsals IV (52.2, 50.1 mm), phalanges IV-1 (9.8, 9.8 mm),
phalanges
IV-2 (one proximal; 5.6 mm), phalanx IV-3 (~13.2 mm), phalanx IV-4,
pedal ungual IV (12.1 mm), metatarsals V (13.1 mm), body feathers,
retrices (Guo, Xu and Jia, 2018)
(41HIII 0415) (375 mm, adult) skull (51.4 mm), mandibles, axis, third
to ninth cervical vertebrae,
third to thirteenth dorsal vertebrae, dorsal ribs, first to
twenty-fourth caudal vertebrae, scapulae (25.7 mm), coracoids, furcula,
humeri (43.8, 42.8 mm), radii (36.9, 34.6 mm), ulnae (37.3, 34.6 mm),
scapholunare, semilunate carpal, metacarpals I (8.3, 7.9 mm), phalanges
I-1
(21.6, 18 mm), manual unguals I (8.2, 8.6 mm),
metacarpals II (21.5 mm), phalanges II-1 (12.5 mm), phalanges II-2
(18.6, 19.3 mm), manual unguals II (9.5, 9.6 mm),
metacarpal III, phalanx III-1, phalanx III-2 (6.4 mm), phalanx III-3
(10.4 mm),
manual ungual III (7.7 mm), fragmentary ilium, ischia (13.6 mm), femora
(46.8 mm), tibiotarsi (71.5, 71.2 mm),
fibulae, metatarsals I (4.8, 3.9 mm), phalanges I-1 (5.5, 5.7 mm),
pedal unguals
I (2.7, 3.6 mm), metatarsals II (32.9, 36.2 mm), phalanges II-1 (7.7,
8.6 mm), phalanges II-2 (7.8, 7.8 mm), pedal unguals II (5.8 mm),
metatarsals III (35.7, 37.2 mm), phalanges III-1 (9.1 mm), phalanges
III-2 (7.6, 7.6 mm), phalanges III-3 (7.2, 6.9 mm), pedal unguals III
(6.4, 5.7 mm), metatarsals IV (32.8, 35.9 mm), phalanges IV-1 (5.8, 7.5
mm), phalanges
IV-2 (4.4, 5.5 mm), phalanges IV-3 (3.5, 4.9 mm), phalanges IV-4 (2.1,
5.4 mm), pedal ungual IV (4.1 mm), metatarsal V (Guo, Xu and Jia, 2018)
(BMNHC Ph 804) skull (43.5 mm), partial mandibles, hyoid, ten cervical
vertebrae, twelve dorsal vertebrae, dorsal ribs, gastralia, synsacrum,
thirty-one or thirty-two caudal vertebrae, scapulae (29.1, 29.1 mm),
coracoids, furcula, humeri (45.7, 44.5 mm), radii (39.1, 38.2 mm),
ulnae (39.8, 38.8 mm), scapholunare, semilunate carpals, metacarpals I, phalanx I-1, manual ungual
I, metacarpals II,
phalanx II-1 (11.4 mm), phalanx II-2 (18.1 mm), manual ungual II (10.1
mm), metacarpals III,
phalanx III-1 (6.9 mm), phalanx III-2 (6.6 mm), phalanx III-3 (9.5 mm),
manual ungual III (9 mm), ilium (25.1 mm), pubes (39.2 mm), ischia (~19
mm), femora (50.9 mm), tibiae (tibiotarsi 69.5, 69.1 mm), fibulae,
proximal
tarsals, metatarsal I, phalanges I-1 (5.3 mm), pedal unguals I (4.2
mm), metatarsals II (39.7 mm),
phalanges II-1 (8.7, 8.7 mm), phalanges II-2 (8.3, 7.6 mm), pedal
ungual II (9.1 mm), metatarsals III, phalanges
III-1 (9.8, 9.7 mm), phalanges III-2 (8, 7.8 mm), phalanges III-3 (7.1,
7 mm), pedal unguals III (7.9, 7.7 mm),
metatarsals IV (38.3, 38.3 mm), phalanges IV-1 (7.4, 7.4 mm), phalanges
IV-2 (5.2, 5.2 mm), phalanx IV-3 (5.1, 5.2 mm), phalanx
IV-4 (4.9, 5.1 mm), pedal unguals IV (6.1, 5 mm), metatarsal V, body
feathers, remiges, metatarsal remiges (Pei et al., 2017)
(BMNHC Ph 822) skull (61.2 mm), mandibles, ten cervical vertebrae,
cervical ribs, about twelve dorsal vertebrae, dorsal ribs, gastralia,
synsacrum, thirty to thirty-one caudal vertebrae, chevrons, scapula
(40.8 mm), coracoid, incomplete furcula, humeri (61.8, 64.9 mm), radii
(58.9 mm), ulnae (58.9, 60.8 mm), scapholunare, semilunate carpal, metacarpal I (12.6 mm), phalanges I-1 (29.6
mm), manual unguals I (18.6 mm), metacarpals II (36.2 mm),
phalanges II-1, phalanges II-2, manual unguals II (20.8 mm),
metacarpals III (32.7 mm),
phalanges III-1 (8.7 mm), phalanges III-2 (7.2 mm), phalanges III-3
(15.3 mm), manual unguals III (13.5 mm), ilium (34.8 mm), pubes (61.4
mm), ischium, femora (70.5 mm), tibiae (tibiotarsi 108.6, 108 mm),
fibulae, proximal
tarsals, metatarsal II (57.8 mm),
phalanges II-1 (11.2 mm), phalanges II-2 (12.2 mm), pedal unguals II
(13.3 mm), metatarsal III (58 mm), phalanges
III-1 (15.2 mm), phalanges III-2 (11.2 mm), phalanges III-3 (7.8 mm),
pedal unguals III,
metatarsal IV (56.1 mm), phalanges IV-1 (11.6 mm), phalanges IV-2 (8.9
mm), phalanges IV-3 (8.4 mm), phalanges
IV-4 (7.9 mm), pedal unguals IV (10.8 mm), pedal claw sheaths, body
feathers, remiges, metatarsal remiges (Pei et al., 2017)
(BMNHC Ph 823) skull (56 mm), mandibles, few posterior cervical
vertebrae, twelve dorsal vertebrae, synsacrum, at least thirty-one
caudal vertebrae, chevrons, scapulae (40.1 mm), coracoid, partial
furcula, humeri (65.2, 64.5 mm), partial radii, partial ulnae, distal
phalanx I-1, manual ungual I, phalanx II-1, phalanges II-2, manual
unguals II, distal metacarpal III,
phalanx III-1, phalanx III-2, phalanx III-3, manual unguals III, ilia
(40.9, 42.3 mm), ischia (22.1 mm), femora (68.7, 67.8 mm), tibiae
(tibiotarsi 95.2, 92.1 mm), fibula, proximal
tarsals, phalanges I-1 (7.5 mm), pedal unguals I, metatarsals II (49
mm),
phalanx II-1 (10.4 mm), phalanx II-2 (8.1 mm), pedal ungual II (13 mm),
metatarsals III (51.2 mm), phalanx
III-1 (12.8 mm), phalanx III-2 (7.8 mm), phalanx III-3 (11 mm), pedal
ungual III (13.2 mm),
metatarsals IV (48.5 mm), phalanx IV-2 (8.8 mm), phalanx IV-3 (7.5 mm),
phalanx
IV-4 (7.2 mm), pedal ungual IV (13.4 mm) (Pei et al., 2017)
(BMNHC Ph 828) skull (38.5 mm), incomplete mandibles, several dorsal vertebrae,
three gastralia, partial sacrum, scapulocoracoids (scapula ~23 mm), furcula,
humeri (40 mm), radii, ulnae (35.3 mm), scapholunares, semilunate carpals,
distal carpal III, metacarpals I (7.8 mm), phalanges I-1, manual unguals I, metacarpals II (20
mm), phalanges II-1, phalanges II-2, manual ungual II, metacarpals III (19.5
mm), phalanx III-1, phalanges III-2, phalanges III-3, manual ungual III, manual
claw sheaths, distal femur, tibiae (65.6 mm), astragalus, distal tarsals, metatarsals
II, phalanx II-1, phalanx II-2, pedal ungual II, phalanges III-1, phalanges
III-2, phalanges III-3, pedal ungual III, metatarsals IV, phalanges IV-1, phalanges
IV-2, phalanges IV-3, phalanges IV-4, pedal ungual IV, metatarsal V, pedal claw
sheaths, body feathers (15-20 mm), remiges (to 62 mm), metatarsal remiges (Li
et al., 2010)
(C.0502) (462 mm) skull (62.5 mm), mandibles (52.7 mm), hyoids,
(cervical series 62 mm) ten cervical vertebrae, cervical ribs, (dorsal
series ~69.6 mm) twelve dorsal vertebrae, dorsal
ribs, gastralia, (sacrum ~25.5 mm) five sacral vertebrae, (caudal
series 243.1 mm) thirty-three
caudal vertebrae (first 4.2, twelfth 10.2, twenty-second 8.9 mm),
chevrons, scapula (33.7 mm), coracoids, furcula, humeri (51.3, 51 mm),
radii (45, 46.2 mm), ulnae (45.2, 46.4 mm), scapholunares, semilunate
carpals, distal carpal III, metacarpals I (10.6, 11 mm), phalanges I-1
(21.3, 21.6 mm), manual
unguals I (14.2, 16.3 mm), metacarpals II (26.8, 26.9 mm),
phalanges II-1 (16.7, 17.6 mm), phalanges II-2 (20.6, 22.2 mm), manual
unguals II (15.1, 17.3 mm), metacarpals III (25.8, 26.1 mm),
phalanges III-1 (5, 6.6 mm), phalanges III-2 (6.2, 6.4 mm), phalanges
III-3 (12.9, 12.2 mm), manual unguals III (15.1, 13.4 mm),
manual claw sheaths, incomplete ilia, pubes (48.9, 49.1 mm), ischia,
femora (55.4, 56.4 mm), tibiae (tibiotarsi 81.8, 82 mm), proximal
tarsals, metatarsal I, phalanx I-1 (6.5 mm), pedal ungual I (5.2 mm),
metatarsals II (45.1, 44.9 mm),
phalanges II-1 (10.6, 10.7 mm), phalanges II-2 (10.4, 10.6 mm), pedal
unguals II (10.9, 10.3 mm), metatarsals III (48.2, 48.1 mm), phalanges
III-1 (11.8, 11.6 mm), phalanges III-2 (9, 9.5 mm), phalanges III-3
(8.6, 8.6 mm), pedal unguals III (11.4, 11.4 mm),
metatarsals IV (44.4, 44.4 mm), phalanges IV-1 (9.2, 9.4 mm), phalanges
IV-2 (6.9, 6.6 mm), phalanges IV-3 (6.2, 6.3 mm), phalanges
IV-4 (6.7, 6.8 mm), pedal unguals IV (8, 10 mm), metatarsals V (10.3,
13.4 mm), pedal claw sheaths, body feathers, remiges, retrices,
metatarsal
remiges (Jiang, 2011)
(LPM-B00169) skull (63.7 mm), mandibles (53.8 mm), eight cervical vertebrae
(series 66.8 mm), thirteen dorsal vertebrae (series 85.4 mm), sixteen dorsal
ribs, gastralia, sacrum (31 mm), nineteen caudal vertebrae (first caudal 5.2
mm, thirteenth caudal 14.2 mm, eighteenth caudal 13.5 mm), seventeen chevrons,
scapulae (one fragmentary; 45.2 mm) coracoid, furcula, humeri (69 mm), radii
(54 mm), ulnae (55.1 mm), scapholunares, pisiforms, semilunate carpals, metacarpals
I (12.4 mm), phalanges I-1 (26.2 mm), manual ungual I (15.6 mm), metacarpals
II (33.9 mm), phalanges II-1 (21 mm), phalanges II-2 (27 mm), manual unguals
II (20.2 mm), metacarpals III (30.5 mm), phalanges III-1 (7.2 mm), phalanges
III-2 (8.1 mm), phalanx III-3 (14.2 mm), manual ungual III (13.8 mm), manual
claw sheaths, ilia (one partial; 37.4 mm), pubes (61.4 mm), ischium (22.4 mm),
femora (66.2 mm), tibiae (106.4 mm), fibula, astragalus, distal tarsal IV, metatarsals
I (11.1 mm), pedal ungual I, metatarsals II (51.2 mm), phalanges II-1 (11.5
mm), phalanges II-2 (12.2 mm), pedal unguals II (14.9 mm), metatarsals III (55.2
mm), phalanges III-1 (12.9 mm), phalanges III-2 (11.1 mm), phalanges III-3 (10.5
mm), pedal unguals III (13.7 mm), metatarsals IV (51.9 mm), phalanges IV-1 (10.8
mm), phalanges IV-2 (8.8 mm), phalanges IV-3 (7 mm), phalanges IV-4 (7.6 mm),
pedal unguals IV (13.5 mm), metatarsals V (19.2 mm), pedal claw sheaths, body
feathers, remiges, metatarsal remiges (Hu et al., 2009)
(LPM coll.) (Hu et al., 2009- page S3)
(PKUP V1068) skull,
mandibles (one incomplete), ten cervical vertebrae, twelve dorsal
vertebrae, first sacral vertebra, fused second to fourth sacral
vertebrae, fifth sacral vertebra, twenty-two caudal vertebrae, scapulae
(~34,2 mm), coracoids, furcula, humeri (72.2 mm), radii (52, 52.8 mm),
ulnae (59, 58.5 mm), scapholunares, semilunate carpals, metacarpals I (12.4, 12.7 mm), phalanges
I-1 (25, 26.1 mm), manual unguals I (21.8 mm), metacarpals II (32.1, 35
mm),
phalanges II-1 (18.9, 18 mm), phalanges II-2 (26.8 mm), manual unguals
II (20.5 mm), metacarpals III (29.8 mm),
phalanges III-1 (7.2 mm), phalanges III-2 (6.9 mm), phalanges III-3
(14.7 mm), manual unguals III (14 mm), partial ilia (42.2, ~48.1 mm),
pubes (56.9, 55.2 mm), femora (88.5, 90.5 mm), tibiae (tibiotarsi 112,
117.7 mm), fibulae, proximal
tarsals, phalanx I-1, pedal ungual I, metatarsals II (50, 56.2 mm),
phalanges II-1 (13 mm), phalanges II-2 (13, 11.5 mm), pedal ungual II,
metatarsals III (one incomplete; 51.5, 56.4 mm), phalanges
III-1 (15, 15.1 mm), phalanx III-2 (10 mm), phalanx III-3 (11.3 mm),
pedal ungual III (16.8 mm),
metatarsals IV (one partial; 49.1, 55 mm), phalanges IV-1 (11, 11.5
mm), phalanges IV-2 (9.1 mm), phalanges IV-3, phalanges IV-4, pedal
ungual IV (10.2 mm) (Pei et al., 2017)
(STM 0-1) incomplete skeleton including femur (44 mm) and feathers (Zheng et
al., 2014)
(STM 0-2) incomplete skeleton including gastralia and femur (70 mm) (Zheng et
al., 2014)
(STM 0-3) incomplete skeleton including gastralia, femur (75 mm) and feathers
(Zheng et al., 2014)
(STM 0-4) incomplete skeleton including femur (65 mm) and feathers (Zheng et
al., 2014)
(STM 0-5) incomplete skeleton including femur (41 mm) and feathers (Zheng et
al., 2014)
(STM 0-6) material including gastralia (Zheng et al., 2014)
(STM 0-7) incomplete skeleton including premaxilla, mandible, caudal
vertebrae, humerus, radius, ulna, metacarpal II, manual phalanx II-2,
manual ungual II, metacarpal III, manual claw sheath, ischium, femur
(64 mm), tibia, fibula, metatarsal 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, phalanx IV-4, scales and feathers (Zheng et al., 2014)
(STM 0-8) (adult) incomplete skeleton including gastralia, femur (86 mm) and feathers
(Zheng et al., 2014)
(STM 0-9) incomplete skeleton including femur (65 mm) and feathers (Zheng et
al., 2014)
(STM 0-10) incomplete skeleton including gastralia and femur (47 mm) (Zheng
et al., 2014)
(STM 0-11) material including gastralia and feathers (Zheng et al., 2014)
(STM 0-12) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-13) incomplete skeleton including femur (73.5 mm) (Zheng et al., 2014)
(STM 0-14) incomplete skeleton including gastralia and femur (67 mm) (Zheng
et al., 2014)
(STM 0-15) material including gastralia (Zheng et al., 2014)
(STM 0-16) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-17) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-18) incomplete skeleton including gastralia, femur (74 mm) and feathers
(Zheng et al., 2014)
(STM 0-19) incomplete skeleton including gastralia, femur (68 mm) and feathers
(Zheng et al., 2014)
(STM 0-20) incomplete skeleton including gastralia and femur (52 mm) (Zheng
et al., 2014)
(STM 0-21) incomplete skeleton including femur (52 mm) and feathers (Zheng et
al., 2014)
(STM 0-22) incomplete skeleton including gastralia (Zheng et al., 2014)
(STM 0-23) incomplete skeleton including gastralia and femur (75 mm) (Zheng
et al., 2014)
(STM 0-24) material including gastralia and femur (50 mm) (Zheng et al., 2014)
(STM 0-25) material including gastralia and femur (62 mm) (Zheng et al., 2014)
(STM 0-26) incomplete skeleton including gastralia and femur (93 mm) (Zheng
et al., 2014)
(STM 0-27) incomplete skeleton including gastralia, femur (52 mm) and feathers
(Zheng et al., 2014)
(STM 0-28) incomplete skeleton including femur (75 mm) and feathers (Zheng et
al., 2014)
(STM 0-29) incomplete skeleton including gastralia, femur ( mm) and feathers
(Zheng et al., 2014)
(STM 0-30) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-31) incomplete skeleton including gastralia,scapula, coracoid, furcula, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-32) incomplete skeleton including gastralia, femur (40 mm) and feathers
(Zheng et al., 2014)
(STM 0-33) material including femur (50 mm) and feathers (Zheng et al., 2014)
(STM 0-34) incomplete skeleton including gastralia and femur (70 mm) (Zheng
et al., 2014)
(STM 0-35) incomplete skeleton including gastralia, femur (80 mm) and feathers
(Zheng et al., 2014)
(STM 0-36) incomplete skeleton including femur (71 mm) and feathers (Zheng et
al., 2014)
(STM 0-37) incomplete skeleton including gastralia, femur (67 mm) and feathers
(Zheng et al., 2014)
(STM 0-38) incomplete skeleton including femur (45 or 40.5 mm), feathers and gastric pellet (30 mm) (Zheng et
al., 2014)
(STM 0-39) incomplete skeleton including gastralia, femur (40 mm) and feathers
(Zheng et al., 2014)
(STM 0-40) incomplete skeleton including gastralia, femur (65 mm) and feathers
(Zheng et al., 2014)
(STM 0-41) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-42) incomplete skeleton including gastralia and femur (60 mm) (Zheng
et al., 2014)
(STM 0-43) material including femur (72 mm) (Zheng et al., 2014)
(STM 0-44) incomplete skeleton including gastralia and femur (60 mm) (Zheng
et al., 2014)
(STM 0-45) (Zheng et al., 2014)
(STM 0-46) material including gastralia and femur (60 mm) (Zheng et al., 2014)
(STM 0-47) incomplete skeleton including gastralia, femur (68 mm) and feathers
(Zheng et al., 2014)
(STM 0-48) incomplete skeleton including gastralia, femur (71 mm) and feathers
(Zheng et al., 2014)
(STM 0-49) material including gastralia and femur (67 mm) (Zheng et al., 2014)
(STM 0-50) incomplete skeleton including gastralia, femur (65 mm) and feathers
(Zheng et al., 2014)
(STM 0-51) material including gastralia and femur (~50 mm) (Zheng et al., 2014)
(STM 0-52) incomplete skeleton including dorsal vertebrae, dorsal ribs,
gastralia, coracoids, humerus, pubes, femur (57 mm) and feathers (Zheng
et al., 2014)
(STM 0-53) incomplete skeleton including gastralia, femur (56 mm) and feathers
(Zheng et al., 2014)
(STM 0-54) incomplete skeleton including femur (49.5 mm) and feathers (Zheng
et al., 2014)
(STM 0-55) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-56) incomplete skeleton including gastralia, femur (42 mm) and feathers
(Zheng et al., 2014)
(STM 0-57) incomplete skeleton including femur (44 mm) and feathers (Zheng et
al., 2014)
(STM 0-58) incomplete skeleton including femur (49 mm) and feathers (Zheng et
al., 2014)
(STM 0-59) incomplete skeleton including gastralia, femur (47 mm) and feathers
(Zheng et al., 2014)
(STM 0-60) material including dorsal vertebrae, dorsal ribs, gastralia,
sacral vertebrae, ilium, pubes, ischia, femur (45 mm) and feathers
(Zheng et al., 2014)
(STM 0-61) incomplete skeleton including femur (66 mm) (Zheng et al., 2014)
(STM 0-62) incomplete skeleton including femur (49 mm) and feathers (Zheng et
al., 2014)
(STM 0-63) material including gastralia, femur (~60 mm) and feathers (Zheng
et al., 2014)
(STM 0-64) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-66) specimen including femur (60 mm) and feathers (Zheng et al., 2014)
(STM 0-67) material including gastralia, femur (~43 mm) and feathers (Zheng
et al., 2014)
(STM 0-68) incomplete skeleton including gastralia and feathers (Zheng et al.,
2014)
(STM 0-69) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-70) material including femur (65 mm) and feathers (Zheng et al., 2014)
(STM 0-71) incomplete skeleton including gastralia, femur (64 mm) and feathers
(Zheng et al., 2014)
(STM 0-72) incomplete skeleton including gastralia and femur (70 mm) (Zheng
et al., 2014)
(STM 0-73) incomplete skeleton including gastralia and femur (62 mm) (Zheng
et al., 2014)
(STM 0-74) incomplete skeleton including gastralia and femur (50 mm) (Zheng
et al., 2014)
(STM 0-75) incomplete skeleton including gastralia and femur (50 mm) (Zheng
et al., 2014)
(STM 0-76) incomplete skeleton including gastralia, femur (55 mm) and feathers
(Zheng et al., 2014)
(STM 0-77) incomplete skeleton including gastralia and femur (49 mm) (Zheng
et al., 2014)
(STM 0-78) incomplete skeleton including gastralia and femur (51 mm) (Zheng
et al., 2014)
(STM 0-79) incomplete skeleton including gastralia, femur (49 mm) and feathers
(Zheng et al., 2014)
(STM 0-80) incomplete skeleton including gastralia, femur (~38 mm) and feathers
(Zheng et al., 2014)
(STM 0-81) material including gastralia and femur (~32 mm) (Zheng et al., 2014)
(STM 0-82) incomplete skeleton including gastralia, femur (47 mm) and feathers
(Zheng et al., 2014)
(STM 0-83) material including gastralia and femur (~39 mm) (Zheng et al., 2014)
(STM 0-84) material including gastralia and femur (52 mm) (Zheng et al., 2014)
(STM 0-85) incomplete skeleton including gastralia and femur (44 mm) (Zheng
et al., 2014)
(STM 0-86) incomplete skeleton including femur (60 mm) and feathers (Zheng et
al., 2014)
(STM 0-87) material including gastralia, femur (62 mm) and feathers (Zheng et
al., 2014)
(STM 0-88) material including gastralia, femur (~66 mm) (Zheng et al., 2014)
(STM 0-89) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-90) material including gastralia, femur (79 mm) and feathers (Zheng et
al., 2014)
(STM 0-91) material including gastralia and femur (70 mm) (Zheng et al., 2014)
(STM 0-92) incomplete skeleton including gastralia and femur (64 mm) (Zheng
et al., 2014)
(STM 0-93) incomplete skeleton including skull, cervical vertebrae,
dorsal vertebrae, dorsal ribs, gastralia, caudal vertebrae, scapula,
coracoid, furcula, humeri, radii, ulnae, manus, ilium, pubes, femora
(69 mm), tibiae, metatarsi and feathers (Zheng et al., 2014)
(STM 0-94) material including gastralia (Zheng et al., 2014)
(STM 0-95) material including gastralia and femur (60 mm) (Zheng et al., 2014)
(STM 0-96) incomplete skeleton including gastralia, femur (~48 mm) and feathers
(Zheng et al., 2014)
(STM 0-97) incomplete skeleton including gastralia and femur (58 mm) (Zheng
et al., 2014)
(STM 0-98) incomplete skeleton including feathers (Zheng et al., 2014)
(STM 0-99) incomplete skeleton including gastralia and femur (76 mm) (Zheng
et al., 2014)
(STM 0-100) incomplete skeleton including femur (74 mm) (Zheng et al., 2014)
(STM 0-101) material including femur (60 mm) (Zheng et al., 2014)
(STM 0-102) incomplete skeleton including gastralia and femur (66 mm) (Zheng
et al., 2014)
(STM 0-103) (Zheng et al., 2014)
(STM 0-104) incomplete skeleton including gastralia, femur (43 mm) and feathers
(Zheng et al., 2014)
(STM 0-105) incomplete skeleton including gastralia, femur (72 mm) and feathers
(Zheng et al., 2014)
(STM 0-106) incomplete skeleton including gastralia, femur (75 mm) and feathers
(Zheng et al., 2014)
(STM 0-107) material including feathers (Zheng et al., 2014)
(STM 0-108) material including gastralia, femur (62 mm) and feathers (Zheng
et al., 2014)
(STM 0-109) material including feathers (Zheng et al., 2014)
(STM 0-110) incomplete skeleton including gastralia, femur (57 mm) and feathers
(Zheng et al., 2014)
(STM 0-111) incomplete skeleton including gastralia, femur (65 mm) and feathers
(Zheng et al., 2014)
(STM 0-112) incomplete skeleton including gastralia, femur (~63 mm) and feathers
(Zheng et al., 2014)
(STM 0-113) material including gastralia and femur (48 mm) (Zheng et al., 2014)
(STM 0-114) material including dorsal ribs, caudal series, humerus,
radius, ulna, metacarpal II, metacarpal III, femur (50 mm), tibiae,
metatarsal I, phalanx I-1, pedal ungual I, metatarsi, phalanx II-1,
phalanx III-1, pes, scales, skin, propatagium and feathers (Zheng et
al., 2014)
(STM 0-115) material including gastralia and feathers (Zheng et al., 2014)
(STM 0-116) material including femur (41 mm) (Zheng et al., 2014)
(STM 0-117) material including feathers (Zheng et al., 2014)
(STM 0-118) incomplete skeleton including premaxilla, mandible,
gastralia, caudal vertebrae, humerus, radius, ulna, metacarpal II,
metacarpal III, pubis, ischium, femora (62 mm), tibiae, fibula,
metatarsal I, phalanx II-1, phalanx III-1, skin and feathers (Zheng et
al., 2014)
(STM 0-119) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-120) material including dorsal vertebrae, dorsal ribs, gastralia, humeri, femora (48 mm) and feathers (Zheng
et al., 2014)
(STM 0-121) incomplete skeleton including gastralia and femur (44 mm) (Zheng
et al., 2014)
(STM 0-122) material including gastralia and feathers (Zheng et al., 2014)
(STM 0-123) material including gastralia, femur (60 mm) and feathers (Zheng
et al., 2014)
(STM 0-124) material including feathers (Zheng et al., 2014)
(STM 0-125) material including caudal vertebrae, humerus, radius, ulna,
metacarpal II, metacarpal III, femur (59 mm), tibia, fibula, metatarsal
I, metatarsals, scales and feathers (Zheng et al., 2014)
(STM 0-126) incomplete skeleton including gastralia, femur (54 mm) and feathers
(Zheng et al., 2014)
(STM 0-127) material including humerus, radius, ulna, scapholunare,
semilunate carpal, metacarpal I, metacarpal II, metacarpal III,
ischium, femur (70 mm), tibia, fibula, metatarsal I, phalanx II-1,
phalanx III-1 skin and propatagium (Zheng et al., 2014)
(STM 0-128) material including gastralia and femur (~50 mm) (Zheng et al., 2014)
(STM 0-129) material including gastralia, femur (~46 mm) and feathers (Zheng
et al., 2014)
(STM 0-130) material including gastralia and femur (60 mm) (Zheng et al., 2014)
(STM 0-131) material including gastralia and feathers (Zheng et al., 2014)
(STM 0-132) incomplete skeleton including premaxilla, mandible,
gastralia, caudal vertebrae, radius, ulna, metacarpal I, phalanx I-1,
metacarpal II, metacarpal III, ischium, femur (40 mm), tibia, fibula,
metatarsal I, phalanx II-1, phalanx III-1, skin and feathers
(Zheng et al., 2014)
(STM 0-133) material including humerus, radius,
ulna, metacarpal II, metacarpal III, tibia, metatarsals, skin and
scales (Zheng et al., 2014)
(STM 0-134) incomplete skeleton including femur (40 mm) and feathers (Zheng
et al., 2014)
(STM 0-135) incomplete skeleton including gastralia and femur (60 mm) (Zheng
et al., 2014)
(STM 0-136) material including gastralia, femur (47 mm) and feathers (Zheng
et al., 2014)
(STM 0-137) incomplete skeleton including gastralia, femur (72 mm) and feathers
(Zheng et al., 2014)
(STM 0-138) incomplete skeleton including gastralia, femur (65 mm) and feathers
(Zheng et al., 2014)
(STM 0-139) incomplete skeleton including feathers (Zheng et al., 2014)
(STM 0-140) incomplete skeleton including gastralia, femur (43 mm) and feathers
(Zheng et al., 2014)
(STM 0-141) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-142) incomplete skeleton including gastralia, femur (41 mm) and feathers
(Zheng et al., 2014)
(STM 0-143) incomplete skeleton including gastralia, femur (68 mm) and feathers
(Zheng et al., 2014)
(STM 0-144) incomplete skeleton including premaxilla, mandible,
gastralia, caudal series, humerus, radius, ulna, carpals, metacarpal I,
phalanx I-1, manual ungual I, metacarpal II, phalanx II-1, phalanx
II-2, metacarpal III, phalanx III-1, phalanx III-2, phalanx III-3,
manual claw sheath, femur (~50 mm), tibia, fibula, metatarsal I,
phalanx II-1, phalanx III-1, skin, propatagium, body feathers and
remiges (Zheng et al., 2014)
(STM 0-145) incomplete skeleton including gastralia and femur (50 mm) (Zheng
et al., 2014)
(STM 0-146) material including gastralia (Zheng et al., 2014)
(STM 0-147) incomplete skeleton including premaxilla, mandible,
gastralia, radius, ulna, metacarpal II, metacarpal III, femur (~35 mm),
tibia, metatarsal I, metatarsus, phalanx II-1, phalanx III-1, pedal
phalanges, pedal ungual IV and skin (Zheng et al., 2014)
(STM 0-148) incomplete skeleton including gastralia and femur (40 mm) (Zheng
et al., 2014)
(STM 0-149) material including feathers (Zheng et al., 2014)
(STM 0-150) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-151) incomplete skeleton including femur (69 mm) and feathers (Zheng
et al., 2014)
(STM 0-152) incomplete skeleton including gastralia, femur (68 mm) and feathers
(Zheng et al., 2014)
(STM 0-153) material including gastralia, femur (44 mm) and feathers (Zheng
et al., 2014)
(STM 0-154) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-155) incomplete skeleton including gastralia, femur (73 mm) and feathers
(Zheng et al., 2014)
(STM 0-156) incomplete skeleton including gastralia and femur (50 mm) (Zheng
et al., 2014)
(STM 0-157) incomplete skeleton including gastralia, femur (54 mm) and feathers
(Zheng et al., 2014)
(STM 0-158) incomplete skeleton including gastralia, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-159) material including gastralia, femur (58 mm) and feathers (Zheng
et al., 2014)
(STM 0-160) incomplete skeleton including gastralia (Zheng et al., 2014)
(STM 0-161) incomplete skeleton including gastralia and femur (75 mm) (Zheng
et al., 2014)
(STM 0-162) incomplete skeleton including gastralia, femur (~53 mm) and feathers
(Zheng et al., 2014)
(STM 0-163) material including femur (48 mm) and feathers (Zheng et al., 2014)
(STM 0-164) incomplete skeleton including gastralia, femur (47 mm) and feathers
(Zheng et al., 2014)
(STM 0-165) incomplete skeleton including dorsal vertebrae, dorsal
ribs, gastralia, sacrum, scapulae, coracoid, furcula, humeri, radius,
ulna, manus, pubes, ischium, femora (63 mm) and feathers (Zheng et al.,
2014)
(STM 0-166) incomplete skeleton including gastralia, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-167) incomplete skeleton including gastralia and femur (45 mm) (Zheng
et al., 2014)
(STM 0-168) incomplete skeleton including gastralia and femur (56 mm) (Zheng
et al., 2014)
(STM 0-169) incomplete skeleton including gastralia, femur (72 mm) and feathers
(Zheng et al., 2014)
(STM 0-170) incomplete skeleton including gastralia, femur (64 mm) and feathers
(Zheng et al., 2014)
(STM 0-171) incomplete skeleton including gastralia, femur (53 mm) and feathers
(Zheng et al., 2014)
(STM 0-172) material including femur (70 mm) and feathers (Zheng et al., 2014)
(STM 0-173) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-174) incomplete skeleton including gastralia and femur (50 mm) (Zheng
et al., 2014)
(STM 0-175) incomplete skeleton including gastralia, femur (75 mm) and feathers
(Zheng et al., 2014)
(STM 0-176) incomplete skeleton including gastralia and femur (68 mm) (Zheng
et al., 2014)
(STM 0-177) incomplete skeleton including gastralia, femur (69 mm) and feathers
(Zheng et al., 2014)
(STM 0-178) incomplete skeleton including gastralia, femur (63 mm) and feathers
(Zheng et al., 2014)
(STM 0-179) incomplete skeleton including skull, mandibles, cervical
vertebrae, cervical ribs, femur (65 or 71 mm), feathers and gastric
pellet including at least three squamates (53 mm) (Zheng et al., 2014)
(STM 0-180) material including feathers (Zheng et al., 2014)
(STM 0-181) incomplete skeleton including gastralia, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-182) incomplete skeleton including gastralia, femur (43 mm) and feathers
(Zheng et al., 2014)
(STM 0-183) incomplete skeleton including gastralia and femur (50 mm) (Zheng
et al., 2014)
(STM 0-184) incomplete skeleton including gastralia (Zheng et al., 2014)
(STM 0-185) incomplete skeleton including femur (65 mm) (Zheng et al., 2014)
(STM 0-186) incomplete skeleton including gastralia, femur (46 mm) and feathers
(Zheng et al., 2014)
(STM 0-187) incomplete skeleton including gastralia, femur (42 mm) and feathers
(Zheng et al., 2014)
(STM 0-188) incomplete skeleton including gastralia, femur (75 mm) and feathers
(Zheng et al., 2014)
(STM 0-189) incomplete skeleton including gastralia and femur (44 mm) (Zheng
et al., 2014)
(STM 0-190) incomplete skeleton including gastralia and femur (54 mm) (Zheng
et al., 2014)
(STM 0-191) incomplete skeleton including gastralia and femur (68 mm) (Zheng
et al., 2014)
(STM 0-192) incomplete skeleton including gastralia and femur (75 mm) (Zheng
et al., 2014)
(STM 0-193) incomplete skeleton including gastralia, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-194) incomplete skeleton including gastralia, femur (46 mm) and feathers
(Zheng et al., 2014)
(STM 0-195) material including femur (43 mm) (Zheng et al., 2014)
(STM 0-196) material including gastralia and feathers (Zheng et al., 2014)
(STM 0-197) material including gastralia and feathers (Zheng et al., 2014)
(STM 0-198) incomplete skeleton including gastralia, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-199) material including gastralia and feathers (Zheng et al., 2014)
(STM 0-200) incomplete skeleton including gastralia (Zheng et al., 2014)
(STM 0-201) material including gastralia, femur (65 mm) and feathers (Zheng
et al., 2014)
(STM 0-202) incomplete skeleton including gastralia, femur (60 mm) and feathers
(Zheng et al., 2014)
(STM 0-203) incomplete skeleton including gastralia and femur (62 mm) (Zheng
et al., 2014)
(STM 0-204) incomplete skeleton including gastralia, femur (64 mm) and feathers
(Zheng et al., 2014)
(STM 0-205) material including femur (60 mm) (Zheng et al., 2014)
(STM 0-206) incomplete skeleton including gastralia, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-207) incomplete skeleton including gastralia, femur (51 mm) and feathers
(Zheng et al., 2014)
(STM 0-208) incomplete skeleton including femur (70 mm) and feathers (Zheng
et al., 2014)
(STM 0-209) incomplete skeleton including gastralia, femur (50 mm) and feathers
(Zheng et al., 2014)
(STM 0-210) incomplete skeleton including gastralia and feathers (Zheng et al.,
2014)
(STM 0-211) incomplete skeleton including femur (58 mm) and feathers (Zheng
et al., 2014)
(STM 0-212) incomplete skeleton including gastralia, femur (45 mm) and feathers
(Zheng et al., 2014)
(STM 0-213) incomplete skeleton including gastralia, femur (71 mm) and feathers
(Zheng et al., 2014)
(STM 0-214) incomplete skeleton including gastralia and feathers (Zheng et al.,
2014)
(STM 0-215) incomplete skeleton including femur (68 mm) and feathers (Zheng
et al., 2014)
(STM 0-216) incomplete skeleton including femur (67 mm) and feathers (Zheng
et al., 2014)
(STM 0-217) incomplete skeleton including gastralia, femur (70 mm) and feathers
(Zheng et al., 2014)
(STM 0-218) material including gastralia and femur (66 mm) (Zheng et al., 2014)
(STM 0-219) material including gastralia, femur (~46 mm) and feathers (Zheng
et al., 2014)
(STM 0-220) material including gastralia, femur (~50 mm) and feathers (Zheng
et al., 2014)
(STM 0-221) incomplete skeleton including femur (73 mm) and feathers (Zheng
et al., 2014)
(STM 0-222) material including femur (50 mm) (Zheng et al., 2014)
(STM 0-223) material including gastralia, femur (46 mm) and feathers (Zheng
et al., 2014)
(STM 0-224) skull, mandible, hyoid, cervical series, dorsal vertebrae,
dorsal ribs, gastralia, first to ~fifteenth caudal vertebrae, furcula,
humeri, radii, ulnae, metacarpals I, phalanges I-1, manual ungual I,
metacarpals II, phalanx II-1, phalanx II-2, manual ungual II,
metacarpals III, phalanax III-1, phalanx III-2, phalanx III-3, manual
ungual III, ilium, femora (80 or 70 mm), tibiae, metatarsi, pedal
phalanges and unguals, feathers and gastric pellet (39 mm) (Zheng et
al., 2014)
(STM 0-225) incomplete skeleton including gastralia and femur (60 mm) (Zheng
et al., 2014)
(STM 0-226) (Zheng et al., 2014)
(STM 0-227) gastric pellet including ptycholepid fish scales and bones (45 mm) (Zhang et al., 2018)
(STM 0-228) gastric pellet including ptycholepid fish scales and bones (35 mm) (Zhang et al., 2018)
(STM A0-4) skull, sclerotic plates, mandibles, cervical series, dorsal
series, partial dorsal ribs, sacrum, first to ~eighteenth caudal
vertebrae, scapulae, coracoids, humeri, radius, ulnae, metacarpals I,
phalanges I-1, manual unguals I,
metacarpals II, phalanges II-1, phalanges II-2, manual ungual II,
metacarpals III, phalanx III-1, phalanges III-2, phalanges III-3,
manual unguals III, manual claw sheaths, femora (~35 mm), tibia,
metatarsi, pedal phalanges, pedal unguals, gastric pellet (17 mm)
(Zheng et al., 2018)
(XHPM 1084) specimen including skull and manus (Foth and Rauhut, 2017)
(YFGP-T5199) (470 mm adult) skull (59.5 mm), mandibles, eight cervical vertebrae
(series 56.5 mm), cervical ribs, seven dorsal vertebrae (series 97.5 mm), dorsal
ribs, gastralia, caudal series (236.5 mm; proximal caudal 6.5, distal caudal
16 mm), chevron, scapulae (31 mm), coracoids, furcula, humeri (70 mm), radii,
ulnae (59.5 mm), scapholunares, semilunate carpal, distal carpals III, metacarpals I (14 mm),
phalanges I-1 (27 mm), manual unguals I (16.5 mm), metacarpals II (~38 mm),
phalanges II-1 (25 mm), phalanges II-2, manual ungual II, metacarpals III (~31
mm), phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ilia (35.5
mm), pubes (49.5 mm), ischium (16 mm), femora (69.5 mm), tibiotarsi (106 mm),
metatarsals I (12 mm), phalanges I-1, pedal unguals I, metatarsals II, phalanges
II-1, phalanges II-2, pedal unguals II, metatarsals III (61 mm), phalanges III-1,
phalanges III-2, phalanges III-3, pedal unguals III, metatarsals IV, phalanges
IV-1, phalanges IV-2, phalanges IV-3, phalanx IV-4, pedal ungual IV, metatarsal
V, body feathers, remiges, retrices (Lefevre et al., 2014; described by Lindgren et al., 2015)
Other diagnoses-
Xu et al. (2008) suggested a short ischium (less than one-fourth of the
femoral length) as being diagnostic, but the holotype's ischium is only
partially exposed and of published measured specimens only YFGP-T5199
has such a short ischium (23%). Other specimens (41HIII 0404 32%;
41HIII 0415 29%; BMNHC Ph 804 37%; BMNHC Ph 823 33%; LPM-B00169 34%) are
longer than Eosinopteryx (28%) and Aurornis (27%). They also listed 'ventral surface of coracoid sculptured
by numerous small pits' as diagnostic, but this seems present in Eosinopteryx as well (unexposed in Aurornis).
Hu et al. (2009) suggested an elongate tibiotarsus as diagnostic, but
the tibiofemoral ratios in e.g. 41HIII 0404 and BMNHC Ph 823 are the
same or less than Aurornis and/or Eosinopteryx.
Pei et al. (2017) listed several characters that were also stated to be
present in other 'anchiornithines'- straight nasal process of
premaxilla; short anterior ramus of maxilla; ventrally displaced
promaxillary fenestra; sheet-like posteroventral process of
dentary; deltopectoral crest no
longer than one-fourth of humeral shaft; straight ulna;
straight radius; fibula with
extremely expanded proximal end as anteroposteriorly broad as tibia.
Comments- The holotype
was discovered by a farmer (Hu et al., 2009) and given to the IVPP
before October 2008. Foth and Rauhut (2017) include high
resolution photos of the holotype manus.
Jiang (2011) describes complete skeleton C.0502 as a new species of Anchiornis
in a thesis. Though described as having thirteen dorsals and four
sacrals, the supposed last dorsal appears to be a first sacral.
The ilium is reconstructed with a very short preacetabular process, but
it is near certainly just broken.
While Xu et al. (2008) used a version of the Theropod Working
Group matrix with added characters from Xu's thesis to place Anchiornis
as an avialan more basal than Archaeopteryx, Hu et al. (2009) used Senter's
version of the TWiG matrix to place it as a troodontid more derived than Sinovenator
but less so than Mei and other taxa. Xu et al. (2011) found it to be
an archaeopterygid, with that group basal in Deinonychosauria. Hartman et al. (2019) found Anchiornis
to be an archaeopterygid, which resolved as basal deinonychosaurs, or
with one extra step sister to troodontids or basal avialans. Its
monophyly relative to Aurornis and Eosinopteryx has not been established.
References- Xu, Zhao, Norell, Sullivan, Hone, Erickson, Wang, Han and
Guo, 2008. A new feathered maniraptoran dinosaur fossil that fills a morphological
gap in avian origin. Chinese Science Bulletin. 54(3), 430-435.
Hu, Hou, Zhang and Xu, 2009. A pre-Archaeopteryx troodontid theropod
from China with long feathers on the metatarsus. Nature. 461, 640-643.
Li, Gao, Vinther, Shawkey, Clarke, D'Alba, Meng, Briggs and Prum, 2010. Plumage
color patterns of an extinct dinosaur. Science. 327 (5971), 1369-1372.
Jiang, 2011. A new discovery of Anchiornis from western of Liaoning, China. Masters thesis, Chengdu University of Technology. 101 pp.
Xu, You, Du and Han, 2011. An Archaeopteryx-like theropod from China
and the origin of Avialae. Nature. 475, 465-470.
Longrich, Vinther, Meng, Li and Russell, 2012. Primitive wing feather arrangement in Archaeopteryx lithographica and Anchiornis huxleyi. Current Biology. 22(23), 2262-2267.
Pei, Li, Meng, Norell and Gao, 2013. Excellently preserved new specimens of
Anchiornis and the implication of early evolution in Paraves. Journal
of Vertebrate Paleontology. Program and Abstracts 2013, 189.
Foth, 2014. Comment on the absence of ossified sternal elements in
basal paravian dinosaurs. Proceedings of the National Academy of
Sciences. 111(50), E5334.
Lefevre, Cau, Hu, Wu, Escuillie and Godefroit, 2014. New basal
Avialae from the Jurassic of China. Journal of Vertebrate Paleontology. Program
and Abstracts 2014, 167.
O'Connor, Wang, Zheng and Zhou, 2014. Reply to Foth: Preserved
cartilage is rare but not absent: Troodontid sternal plates are absent,
not rare. Proceedings of the National Academy of Sciences. 111(50),
E5335.
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.
Lindgren, Sj�vall, Carney, Cincotta, Uvdal, Hutcheson, Gustafsson, Lef�vre,
Escuilli�, Heimdal, Engdahl, Gren, Kear, Wakamatsu, Yans and Godefroit,
2015. Molecular composition and ultrastructure of Jurassic paravian feathers.
Scientific Reports. 5, 13520.
Pei, 2015. New paravian fossils from the Mesozoic of east Asia and
their bearing on the phylogeny of the Coelurosauria. PhD thesis,
Columbia University. 545 pp.
Foth and Rauhut, 2017. Re-evaluation of the Haarlem Archaeopteryx and the radiation of maniraptoran theropod dinosaurs. BMC Evolutionary Biology. 17:236.
Lefevre, Cau, Cincotta, Hu, Chinsamy, Escuillie and Godefroit, 2017. A
new Jurassic theropod from China documents a transitional step in the
macrostructure of feathers. The Science of Nature. 104:74.
Pei, Li, Meng, Norell and Gao, 2017. New specimens of Anchiornis huxleyi
(Theropoda: Paraves) from the Late Jurassic of northeastern China.
Bulletin of the American Museum of Natural History. 411, 66 pp.
Wang, Pittman, Zheng, Kaye, Falk, Hartman and Xu, 2017. Basal paravian
functional anatomy illuminated by high-detail body outline. Nature
Communications. 8:14576.
Guo, Xu and Jia, 2018. Morphological and phylogenetic study based on new materials of Anchiornis huxleyi (Dinosauria, Theropoda) from Jianchang, western Liaoning, China. Acta Geologica Sinica. 92(1), 1-15.
Rauhut, Foth and Tischlinger, 2018. The oldest Archaeopteryx (Theropoda: Avialiae): A new specimen from the Kimmeridgian/Tithonian boundary of Schamhaupten, Bavaria. PeerJ. 6:e4191.
Zheng, Wang, Sullivan, Zhang, Zhang, Wang, Li and Xu, 2018. Exceptional
dinosaur fossils reveal early origin of avian-style digestion.
Scientific Reports. 8:14217.
Aurornis Godefroit, Cau, Hu, Escuillie,
Wu and Dyke, 2013
A. xui Godefroit, Cau, Hu, Escuillie, Wu and Dyke, 2013
Oxfordian, Late Jurassic
Yaolugou, Tiaojishan Formation, Liaoning, China
Holotype- (YFGP-T5198) (510 mm adult) skull (57 mm), mandibles, seven
cervical vertebrae (series 62 mm), cervical ribs, eleven dorsal vertebrae (series
97 mm), dorsal ribs, synsacrum (30 mm), about thirty caudal vertebrae (series
265 mm), chevrons, scapulae (36.6 mm), coracoid, furcula, humeri (58 mm), radii,
ulnae (57 mm), scapholunare, pisiform, semilunate carpal, metacarpal I (12 mm), phalanx
I-1 (27 mm), manual ungual I, metacarpals II (34 mm), phalanx II-1 (14.5 mm),
incomplete phalanx II-2, metacarpals III (34.5 mm), phalanx III-1, phalanx III-2,
phalanx III-3, incomplete manual ungual III, ilia (34.8 mm), pubes (55 mm),
ischia (~18 mm), femora (66 mm), tibiotarsi (90.5 mm), partial metatarsal I, phalanx
I-1, pedal ungual I, metatarsi (44 mm), phalanges II-1 (7 mm), phalanx II-2
(9 mm), pedal ungual II, phalanges III-1 (12 mm), phalanx III-2 (6.5 mm), phalanx
III-3 (6 mm), pedal ungual III (7.5 mm), phalanges IV-1, phalanx IV-2, phalanx
IV-3 (5 mm), phalanx IV-4 (5 mm), pedal ungual IV (7 mm), body feathers
Diagnosis- (modified after Godefroit et al., 2013) manual phalanx I-1
distinctly more robust than radius; robust postacetabular process of ilium not
markedly deflected ventrally and with horizontal dorsal margin; distal end of
ischium dorsoventrally expanded and formed by hook-like ventral process delimiting
a prominent distal obturator notch and by a longer dorsal distal process.
Other diagnoses- Godefroit et al. also list "metatarsal I gracile
and elongate (about 30% of metatarsal III length)", but this is erroneous
(see below).
Pei et al. (2017) claimed "The posterior (= postacetabular) process of
the right ilium of YFGP-T5198 is more squared than in other Anchiornis,
but the left posterior (= postacetabular) process of YFGP-T5198 is
dorsoventrally shallow and has a posteroventrally sloping dorsal edge
as in other Anchiornis
specimens (e.g., IVPP V14378, LPMB00169, BMNHC Ph 804, BMNHC Ph 822, and
BMNHC Ph 823)", but the left ilium seems to be in dorsal view.
Comments- The identification of some elements by Godefroit et al. (2013)
is questionable. Instead of the surangular and angular being equal in depth
as reconstructed, there is a suture showing the angular was much shallower as
is usual for maniraptorans. The third manual digit is reconstructed as having
two elongate proximal phalanges and a partially preserved third phalanx that
would almost certainly make it longer than digit II. While the poor preservation
and photo quality make any identification uncertain, there is a lateral bend
halfway through and a short medial concavity that would support their phalanx
III-1 being both III-1 and III-2. Thus their III-2 would be III-3, and their
III-3 fragment would be most of ungual III, resembling the proportions of other
basal paravians. Finally, metatarsal I is shown as being a third as long as
III and untapered proximally, unlike any theropod. This apparent structure is
formed by parts of the main metatarsal shafts before their surface is broken
away more proximally. There does seem to be a small fragment of metatarsal I
conntected to phalanx I-1.
The
holotype is one of several paravian specimens acquired by the YFGP from
a fossil dealer prior to June 2012. Pei et al. (2017) considered this
probably a junior synonym of Anchiornis huxleyi.
The authors resolved Aurornis
as a basal avialan using a reduced version of Cau's matrix, similar to Lee et
al.'s (2014) later iteration of this matrix. Brusatte et al. (2014) found it
to be a basal troodontid, however. All of these trees had the genus closely
related to Eosinopteryx and Anchiornis, which are tentatively
placed as archaeopterygids here.
References- Godefroit, Cau, Hu, Escuillie, Wu and Dyke, 2013. A Jurassic
avialan dinosaur from China resolves the early phylogenetic history of birds.
Nature. 498, 359-362.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Pei, Li, Meng, Norell and Gao, 2017. New specimens of Anchiornis huxleyi
(Theropoda: Paraves) from the Late Jurassic of northeastern China.
Bulletin of the American Muiseum of Natural History. 411, 66 pp.
unnamed archaeopterygid (Lefevre, Cau, Hu, Wu, Escuillie and Godefroit, 2014)
Oxfordian, Late Jurassic
Tiaojishan Formation, Liaoning, China
Material-
(YFGP-T5200) (juvenile) skull, mandible, hyoids, cervical series,
dorsal vertebrae, dorsal ribs, gastralia, uncinate processes, sacrum,
first to ~nineteenth caudal vertebrae, scapulae, coracoids, furcula,
humeri, radii, ulnae, metacarpal I, phalanges I-1, manual unguals I,
metacarpals II, phalanges II-1, phalanges II-2, manual unguals II,
metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3,
manual unguals III, posterior ilia, pubes, femora (~46 mm), tibiae,
fibulae, proximal tarsals, metatarsals I, phalanges I-1, pedal unguals
I, metatarsals II, phalanx II-1, phalanx II-2, pedal ungual II,
metatarsals III, phalanges III-1, phalanx III-2, phalanges III-3 (one
distal), pedal unguals III, metatarsals IV, phalanges IV-1, phalanges
IV-2, phalanx IV-3, phalanx IV-4, pedal unguals IV, body feathers,
remiges, retrices, metatarsal remiges
Comments-
Brougham (2013) added "three new undescribed Tiaojishan theropods" to
three coelurosaur matrices, which his publically available poster
(Brougham, online 2013) shows are labeled as YFGP-T5199, YFGP-T5200 and
YFGP-T5201. However, YFGP-T5199 has since been described as an Anchiornis
specimen (Lindgren et al., 2015; also figured in Lefevre et al., 2017)
which is not the same as Brougham figured. Lefevre et al. (2014)
also lists these three specimen numbers, but states "rectrices are well
developed along the tail of YFGP-T5200", while Lefevre et al.'s (2015)
description of YFGP-T5200 corresponds to "YFGP-T5199" in Brougham's
poster. Thus Brougham's numbers are incorrectly associated with
specimens. Regardless, Brougham recovered all of these specimens
in a clade with Aurornis and Eosinopteryx, Lefevre et al. (2014) stated they were "basalmost Avialae" which is where Cau's matrix recovered Anchiornis-grade taxa at the time, and Lefevre et al. (2015) found YFGP-T5200 to be "a basalmost Avialae forming a basal clade with Aurornis." They are thus here assigned to Archaeopterygidae, where the Hartman et al. matrix recovered Aurornis and Eosinopteryx.
As described by Lefevre et al. (2015), YFGP-T5200 has symmetrical
remiges, metatarsal remiges and retrices, with no barbules and the
retrices extend down most of the tail as in Archaeopteryx.
References-
Brougham, 2013. Multi-matrix analysis of new Late Jurassic feathered
theropods from China supports troodontid-avialan clade. Symposium on
Vertebrate Palaeontology and Comparative Anatomy, Programme and
Abstracts. 49.
Brougham, 2013 online. https://www.researchgate.net/publication/280728942_SVPCA_Poster
Lefevre, Cau, Hu, Wu, Escuillie and Godefroit, 2014. New basal
Avialae from the Jurassic of China. Journal of Vertebrate Paleontology. Program
and Abstracts 2014, 167.
Lefevre, Cau, Hu, Wu, Escuillie and Godefroit, 2015. A Jurassic avialan
from China challenges the flight abilities in the earliest birds.
Abstracts of the 12th Symposium on Mesozoic Terrestrial
Ecosystems. [pp]
Lindgren, Sj�vall, Carney, Cincotta, Uvdal, Hutcheson, Gustafsson, Lef�vre,
Escuilli�, Heimdal, Engdahl, Gren, Kear, Wakamatsu, Yans and Godefroit,
2015. Molecular composition and ultrastructure of Jurassic paravian feathers.
Scientific Reports. 5, 13520.
Lef�vre, Cau, Cincotta, Hu, Chinsamy, Escuilli� and Godefroit, 2017. A
new Jurassic theropod from China documents a transitional step in the
macrostructure of feathers. Science of Nature. 104(9-10):74.
unnamed archaeopterygid (Lefevre, Cau, Hu, Wu, Escuillie and Godefroit, 2014)
Oxfordian, Late Jurassic
Tiaojishan Formation, Liaoning, China
Material- (YFGP-T5201) skull, mandibles, hyoid, cervical
series, dorsal series, dorsal
ribs, gastralia, uncinate processes, synsacrum, caudal series,
chevrons, scapulae, coracoids, furcula, humeri, radii (one proximal),
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,
fragmentary ilia, pubes, ischia, femora (~69 mm), tibiae, fibulae,
proximal
tarsals, 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, proximal phalanges III-3,
metatarsals IV, phalanges IV-1, phalanges IV-2, phalanx IV-3, phalanges
IV-4 (one proximal), pedal ungual IV
Comments- Brougham (2013) added "three new undescribed
Tiaojishan theropods" to
three coelurosaur matrices, which his publically available poster
(Brougham, online 2013) shows are labeled as YFGP-T5199, YFGP-T5200 and
YFGP-T5201. As discussed in the YFGP-T5200 comments, Brougham's
YFGP-T199 is actually YFGP-T5200, and the actual YFGP-T5199 Anchiornis specimen is not pictured. The other pictured specimens besides Aurornis (YFGP-T5198) and Eosinopteryx (YFGP-T5197) are the then-undescribed Serikornis (PMOL-AB00200; labeled YFGP-T5201) and an otherwise unpublished specimen (labeled YFGP-T5200). Thus unless Serikornis
originally had a YFGP number, YFGP-5201 is assumed to be the skeleton
in side view with no obvious remiges or retrices and an elongated
upcurved snout. Brougham recovered all of these specimens in a
clade with Aurornis and Eosinopteryx and Lefevre et al. (2014) stated they were "basalmost Avialae" which is where Cau's matrix recovered Anchiornis-grade taxa at the time. They are thus here assigned to Archaeopterygidae, where the Hartman et al. matrix recovered Aurornis and Eosinopteryx.
Lefevre et al. (2014) state "The skull of YFGP-T5201 displays several
characters previously thought to be synapomorphic for Troodontidae."
References-
Brougham, 2013. Multi-matrix analysis of new Late Jurassic feathered
theropods from China supports troodontid-avialan clade. Symposium on
Vertebrate Palaeontology and Comparative Anatomy, Programme and
Abstracts. 49.
Brougham, 2013 online. https://www.researchgate.net/publication/280728942_SVPCA_Poster
Lefevre, Cau, Hu, Wu, Escuillie and Godefroit, 2014. New basal
Avialae from the Jurassic of China. Journal of Vertebrate Paleontology. Program
and Abstracts 2014, 167.
Eosinopteryx Godefroit,
Demuynck, Dyke, Hu, Escuillie and Claeys, 2013
E. brevipenna Godefroit, Demuynck, Dyke, Hu, Escuillie and Claeys,
2013
Oxfordian, Late Jurassic
Yaolugou, Tiaojishan Formation, Liaoning, China
Holotype- (YFGP-T5197) (subadult) skull (43.2 mm), mandibles (40.2 mm),
seven cervical vertebrae (series 39.6 mm), cervical ribs, dorsal vertebrae (series,
57.9 mm), dorsal ribs, gastralia, partial synsacrum (25.1 mm), caudal vertebrae
1-3 (~5.1 mm), fourth caudal vertebra (5.7 mm), fifth caudal vertebra (5.6 mm),
sixth caudal vertebra (5.8 mm), seventh caudal vertebra (6.4 mm), eighth caudal
vertebra (6.4 mm), ninth caudal vertebra (6.4 mm), tenth caudal vertebra (7.6
mm), eleventh caudal vertebra (9.6 mm), twelfth caudal vertebra (9.6 mm), thirteenth
caudal vertebra (8.8 mm), fourteenth caudal vertebra (8 mm), fifteenth caudal
vertebra (7.6 mm), sixteenth caudal vertebra (8 mm), seventeenth caudal vertebra
(8 mm), eighteenth caudal vertebra (7.4 mm), nineteenth caudal vertebra (6.3
mm), twentieth caudal vertebra (9.2 mm), chevrons, scapulae (23.8 mm), coracoids,
partial furcula, humeri (37.9 mm), radii (39.5 mm), ulnae (42 mm), scapholunare,
semilunate carpal, metacarpals I, phalanges I-1 (one incomplete), manual unguals
I (one partial), metacarpals II, phalanges II-1, phalanges II-2, manual unguals
II, metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3, manual
ungual III, ilia (25 mm), incomplete pubis (35 mm), ischia (13.4 mm), femora
(48.5 mm), tibiae (69.5 mm), metatarsal I, phalanx I-1, metatarsals II, phalanges
II-1, phalanges II-2, pedal unguals II, metatarsals III (35.5 mm), phalanges
III-1 (8.8 mm), phalanges III-2 (7.1 mm), phalanx III-3 (6.5 mm), pedal ungual
III (6 mm), metatarsals IV, phalanges IV-1, phalanges IV-2, phalanges IV-3,
phalanx IV-4, pedal ungual IV, body feathers, remiges
Diagnosis- (after Godefroit et al., 2013)
short tail, composed of 20 caudal vertebrae, 2.7 times length of the femur.
Other diagnoses- Contra
Godefroit et al.'s (2013) character "short snout, about 82% the length
of the orbit", Pei et al. (2017) noted the snout "is broken and
incomplete, and thus cannot be used for comparison" and that "the
anteroposterior length of the preorbital portion of the skull is about
1.5-2 times that of the orbit, which is the same as in Anchiornis." Contra Godefroit et al.'s character "lacrimal with long posterior process participating in about
half the length of dorsal margin of orbit and a vestigial anterior process", Pei et al. say "The lacrimal of [Anchiornis
specimen] BMNHC Ph 804 has both long anterior and posterior processes,
but both processes are extremely slender and easy to break, so the
longer posterior process of the lacrimal is not a reliable diagnostic
feature of Eosinopteryx."
Contra Godefroit et al.'s character "chevrons reduced to small rod-like
elements below the 8th or 9th caudal", Pei et al. state "Reduced
chevrons below the proximal 8th or 9th caudal vertebrae are observed in
other Anchiornis specimens,
such as BMNHC Ph 804 and BMNHC Ph 822." Contra Godefroit et al.'s
character "ilium with proportionally long, low and distally tapering
postacetabular process (ratio 'length/height at midlength' = 5)", Pei
et al. state "a long and low posterior (= postacetabular) process of
the ilium is also present in Anchiornis
(e.g., IVPP V14378, BMNHC Ph 804, BMNHC Ph 822, and BMNHC Ph 823)."
Contra Godefroit et al.'s character "pedal unguals shorter than
corresponding penultimate phalanges", Pei et al. suggest "Anchiornis
specimens have variation in proportions of pedal phalanges, even within
the same specimen, such as in BMNHC Ph 804." Finally, contra Godefroit
et al.'s character "absence of rectrices (versus other paravians with
preserved plumage) and feathers on metatarsus and pes (versus other
troodontids with preserved plumage on the hindlimb)", Pei et al. state
"The absence of rectrices and feathers on the metatarsus is possibly a
preservational artifact, as many Anchiornis specimens only have feathers associated with few bones instead of the entire body."
Comments-
The holotype is one of several paravian specimens acquired by the YFGP
from a fossil dealer prior to June 2012. Plumage differences from e.g. Anchiornis are here seen as questionable, as other specimens
which should originally possess pennaceous feathers on wings/tail also lack
them, presumably due to preservation (Longipteryx holotype, NGMC 91,
Epidexipteryx). Godefroit et al. (2013) claimed neurocentral and tibiotarsal
fusion indicated a subdult or adult age, but Lefevre et al. (2014) found histology
showed it was immature.
The authors resolved Eosinopteryx as a basal troodontid using a version
of Senter's TWiG matrix. Brusatte et al. (2014) also found Eosinpteryx
to be a basal troodontid, while Foth et al. (2014) and Lee et al. (2014) both
recovered it as a basal avialan closer to birds than troodontids. All of these
trees had the genus closely related to Aurornis and Anchiornis,
which are tentatively placed as archaeopterygids here.
References- Godefroit, Demuynck, Dyke, Hu, Escuillie and Claeys, 2013.
Reduced plumage and flight ability of a new Jurassic paravian theropod from
China. Nature. 498, 359-362.
Brusatte, Lloyd, Wang and Norell, 2014. Gradual assembly of avian body plan
culminated in rapid rates of evolution across the dinosaur-bird transition.
Current Biology. 24(20), 2386-2392.
Foth, Tischlinger and Rauhut, 2014. New specimen of Archaeopteryx provides
insights into the evolution of pennaceous feathers. Nature. 511, 79-82.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Lefevre, Cau, Hu, Wu, Escuillie and Godefroit, 2014. New basal Avialae from
the Jurassic of China. Journal of Vertebrate Paleontology. Program and Abstracts
2014, 167.
Pei, Li, Meng, Norell and Gao, 2017. New specimens of Anchiornis huxleyi
(Theropoda: Paraves) from the Late Jurassic of northeastern China.
Bulletin of the American Muiseum of Natural History. 411, 66 pp.
unnamed archaeopterygid (Louchart and Pouech, 2017)
Early-Middle Berriasian, Early Cretaceous
Cherves-de-Cognac, France
Material- (CHVm03.514) tooth (1.25x.98x.43 mm)
Comments- Louchart and Pouech
(2017) described this as an archaeopterygid, but Rauhut et al. (2018)
believed it "differs from the typical teeth of Archaeopteryx
in the more compressed and less bulbous shape" and so referred it to
Avialae indet.. However, the compression (BW/FABL 43%) is similar
to e.g. the sixth maxillary tooth of the Daiting specimen (46%), so
Louchert and Pouech's identification is accepted here.
References- Louchart and
Pouech, 2017. A tooth of Archaeopterygidae (Aves) from the Lower
Cretaceous of France extends the spatial and temporal occurrence of the
earliest birds. Cretaceous Research. 73, 40-46.
Rauhut, Foth and Tischlinger, 2018. The oldest Archaeopteryx (Theropoda: Avialiae): A new specimen from the Kimmeridgian/Tithonian boundary of Schamhaupten, Bavaria. PeerJ. 6:e4191.
Archaeopteryx Meyer,
1861b
= Griphosaurus Wagner, 1862 (nomen rejectum)
= Griphornis Owen vide Woodward, 1862 (nomen rejectum)
= Archaeornis Petronievics vide Petroneivics and Woodward, 1917
= Jurapteryx Howgate, 1985
= Wellnhoferia Elzanowski, 2001
Other definitions- (Archaeopteryx lithographica <- Passer
domesticus) (modified from Sereno, 1998)
A. lithographica Meyer, 1861b
= Griphornis longicaudatus Owen vide Woodward, 1862 (nomen rejectum)
= Griphosaurus problematicus Wagner vide Woodward, 1862 (nomen rejectum)
= Archaeopteryx macrura Owen, 1863 (nomen rejectum)
= Archaeopteryx siemensii Dames, 1897
= Archaeornis siemensii (Dames, 1897) Petronievics vide Petroneivics
and Woodward, 1917
= Archaeopteryx oweni Petronievics, 1921
= Griphosaurus longicaudatus (Owen vide Woodward, 1862) Owen vide Brodkorb,
1963
= Archaeopteryx recurva Howgate, 1984
= Jurapteryx recurva (Howgate, 1984) Howgate, 1985
= Archaeopteryx bavarica Wellnhofer, 1993
= Wellnhoferia grandis Elzanowski, 2001
= Archaeopteryx albersdoerferi Kundr�t, Nudds, Kear, L� and Ahlberg, 2019
Early Tithonian, Late Jurassic
Upper Solnhofen Member of the Altmuhltal Formation, Germany
Neotype- (NHMUK 37001; London specimen; holotype of Griphosaurus;
holotype of Griphosaurus problematicus; holotype of Griphornis longicaudatus;
holotype of Archaeopteryx macrura; holotype of Archaeopteryx oweni)
(.46 m, 450 g, 460 day old subadult) premaxilla, maxillary fragment,
nasals, partial quadrate, braincase, cranial fragments, cervical
centrum (7 mm), cervical neural arch, ninth cervical vertebra, tenth
cervical vertebra, first dorsal vertebra (~7 mm), second dorsal
vertebra, third dorsal vertebra, fourth dorsal vertebra, fifth dorsal
vertebra, sixth dorsal vertebra (~7 mm), seventh dorsal vertebra (~7
mm), eighth dorsal vertebra, ninth dorsal vertebra, tenth dorsal
vertebra, eleventh dorsal vertebra (5.5 mm), twelfth dorsal vertebra
(5.5 mm), thirteenth dorsal vertebra (5.5 mm), dorsal ribs, gastralia,
synsacrum (~33 mm) fourth caudal vertebra, fifth caudal vertebra, sixth
caudal vertebra (6 mm), seventh caudal vertebra (6.5 mm), eighth caudal
vertebra (8 mm), ninth caudal vertebra (~10.5 mm), tenth caudal
vertebra (~10.5 mm), eleventh caudal vertebra (11.7 mm), twelfth caudal
vertebra (12.3 mm), thirteenth caudal vertebra (13 mm), fourteenth
caudal vertebra (12.3 mm), fifteenth caudal vertebra (12.5 mm),
sixteenth caudal vertebra (12.4 mm), seventeenth caudal vertebra (11.4
mm), eighteenth caudal vertebra (11.4 mm), nineteenth caudal vertebra
(10.5 mm), twentieth caudal vertebra (9.3 mm), twenty-first caudal
vertebra (8.5 mm), twenty-second caudal vertebra (7 mm), twenty-third
caudal vertebra (4.3 mm), scapulae (46 mm), coracoids, furcula, humeri
(75 mm), radii (65 mm), ulnae (67.5 mm), scapholunare, pisiform, semilunate
carpals, distal carpal III, partial metacarpal I, phalanx I-1, partial
manual ungual I, metacarpals II (34.4 mm), phalanx II-2, manual ungual
II, metacarpals III (one proximal), ilia (38 mm), pubes (51.5 mm),
ischium (~25.5 mm), femora (60.5 mm), tibiae (80.5 mm), fibulae,
astragalus, metatarsal I fragment, phalanx I-1 (8.8 mm), pedal ungual I
(~6.8 mm), metatarsal II (~40 mm), phalanx II-1 (11 mm), phalanx II-2
(11 mm), pedal ungual II (~11 mm), metatarsal III (44 mm), phalanx
III-1 (12.7 mm), phalanx III-2 (11 mm), phalanx III-3 (~9.5 mm), pedal
ungual III (~14 mm), phalanx IV-4 , pedal ungual IV (~11 mm), remiges
(130 mm), retrices (60-120 mm) (Meyer, 1861b)
Referred-
?(BMMS coll.; Ottmann and Steil specimen; chicken wing; ninth specimen) (420 day old juvenile) humerus (70.1
mm), radius (~59 mm), ulna (62 mm), semilunate carpal, metacarpal I (10 mm),
phalanx I-1 (22.5 mm), metacarpal II (31.3 mm), phalanx II-1 (~16.5 mm), phalanx
II-2 (21 mm), manual ungual II (~15 mm), metacarpal III (27.5 mm), phalanx III-1
(6.3 mm), phalanx III-2 (5 mm), phalanx III-3 (14.4 mm), manual ungual III (~10
mm), remiges (Wellnhofer and Roper, 2005)
?(BSP 1869 VIII 1; MB.Av.100; = HMN.Ab.100; holotype of Archaeopteryx lithographica) upper major primary covert
feather (70.3 mm) (Meyer, 1861a)
(BSP 1999 I 50; Munich specimen; Solnhofen-Aktien-Verein specimen; seventh specimen;
holotype of Archaeopteryx bavarica) (270 day old juvenile) partial skull
(~45 mm), mandibles (40 mm), atlas, axis, third cervical vertebra, fourth cervical
vertebra, fifth cervical vertebra, sixth cervical vertebra, seventh cervical
vertebra, eighth cervical vertebra, ninth cervical neural spine, tenth cervical
neural spine, cervical ribs, first dorsal neural spine, partial second dorsal
vertebra, third dorsal neural spine, seventh dorsal neural spine, eighth dorsal
vertebra, ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra,
twelfth dorsal vertebra, thirteenth dorsal vertebra, dorsal ribs, gastralia,
(sacrum 22 mm) first sacral centrum, second sacral centrum, fifth sacral centrum,
first caudal vertebra, second caudal vertebra, third caudal vertebra, fourth
caudal vertebra, fifth caudal vertebra, sixth caudal vertebra, seventh caudal
vertebra, eighth caudal vertebra, ninth caudal vertebra, tenth caudal vertebra,
eleventh caudal vertebra, twelfth caudal vertebra, thirteenth caudal vertebra,
fourteenth caudal vertebra, fifteenth caudal vertebra, sixteenth caudal vertebra,
seventeenth caudal vertebra, eighteenth caudal vertebra, nineteenth caudal vertebra,
twentieth caudal vertebra, twenty-first caudal vertebra, chevrons, scapulae
(34.5 mm), coracoids (15 mm), partial furcula, humeri (~55 mm), radii (53 mm),
ulnae (53 mm), pisiform, semilunate carpal, distal carpal III, metacarpal I (7
mm), phalanges I-1 (20 mm), manual unguals I (9.5 mm), metacarpals II (one distal;
25 mm), phalanges II-1 (12.5 mm), phalanges II-2 (18.5 mm), manual unguals II
(10 mm), metacarpals III (one distal; ~23 mm), phalanges III-1 (one partial;
4.9 mm), phalanx III-2 (4.2 mm), phalanges III-3 (12 mm), manual unguals III
(6.5 mm), manual claw sheaths, ilium (~27.5 mm), pubis (40 mm), ischia (~16
mm), incomplete femora (~46.5 mm), tibiae (~71.5 mm), fibula (69.5 mm), astragali,
calcanea, two distal tarsals, metatarsals I, phalanges I-1 (7.1 mm), pedal ungual
I (5.2 mm), metatarsals II (~36 mm), phalanges II-1 (~8 mm), phalanges II-2
(~8 mm), pedal unguals II (7, 7.4 mm), metatarsals III (40.5 mm), phalanges
III-1 (10.8 mm), phalanges III-2 (8.5 mm), phalanges III-3 (8.4 mm), pedal unguals
III (6.8, 7 mm), metatarsal IV (37 mm), phalanx IV-1 (8 mm), phalanx IV-2 (6
mm), phalanges IV-3 (~5.5 mm), phalanges IV-4 (7 mm), pedal unguals IV (6.2
mm), metatarsal V (~10 mm), pedal claw sheaths, remiges, retrices (Wellnhofer,
1993)
?(Opitsch coll.; Maxberg specimen; third specimen; lost) fourth
cervical centrum, fifth cervical centrum, sixth cervical centrum,
seventh cervical centrum, eighth cervical centrum, ninth cervical
centrum, tenth cervical centrum, first dorsal centrum, second dorsal
centrum, third dorsal centrum, fourth dorsal centrum, fifth dorsal
centrum, sixth dorsal centrum, seventh dorsal centrum, eighth dorsal
centrum, ninth dorsal centrum, tenth dorsal centrum, eleventh dorsal
centrum, twelfth dorsal centrum, thirteenth dorsal vertebra, dorsal rib
fragments, gastralia, first sacral vertebra, second sacral vertebra,
third sacral vertebra, fourth sacral vertebra, fifth sacral vertebra,
first caudal centrum, second caudal centrum, third caudal centrum,
partial scapulae, coracoid, incomplete furcula, humeri (one incomplete;
72 mm), radii (63 mm), ulnae (one incomplete; ~62 mm), metacarpals I
(~10 mm), distal phalanx I-1, manual unguals I (~12 mm), metacarpals II
(~33 mm), phalanges II-1 (19 mm), phalanges II-2 (~22 mm), manual
unguals II (~15 mm), metacarpals III (one proximal; 30 mm), phalanges
III-1, phalanges III-2, phalanx III-3 (~16 mm), manual unguals III,
manual claw sheath, partial ilium, pubes, ischium, femur (~58 mm),
tibiae (one partial; ~79.5 mm), incomplete fibula, distal tarsal III,
distal tarsal IV, metatarsal I, phalanx I-1, pedal ungual I, metatarsus
(II ~38 mm, III ~42 mm, IV ~39 mm), phalanx II-1 (~9.5 mm), phalanx
II-2 (~10 mm), pedal ungual II, phalanx III-1 (~11 mm), phalanx III-2
(~10.5 mm), phalanx III-3, pedal ungual III, phalanx IV-1, phalanx
IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV, remiges, hindlimb
feathers (Heller, 1959)
Early Tithonian, Late Jurassic
Lower Eichstatt Member of the Altmuhltal Formation, Germany
(MB.Av.101; = HMN MB. 1880/81; Berlin specimen; second specimen; holotype of Archaeopteryx
seimensii)
(.405 m, 340 g; 330 day old juvenile) skull (45 mm), sclerotic ring,
mandible (43.5 mm), hyoids, axis (5 mm), third cervical vertebra (6.5
mm), fourth cervical vertebra (9 mm), fifth cervical vertebra (9.5 mm),
sixth cervical vertebra (8 mm), seventh cervical vertebra (9 mm),
eighth cervical vertebra, ninth cervical vertebra (8 mm), tenth
cervical vertebra (7.7 mm), cervical ribs, first dorsal vertebra (7.3
mm), second dorsal vertebra (5.5 mm), third dorsal vertebra (5.9 mm),
fourth dorsal vertebra (5.7 mm), fifth dorsal vertebra (5.5 mm), sixth
dorsal vertebra (5.5 mm), seventh dorsal vertebra (6.1 mm), eighth
dorsal vertebra (6.1 mm), ninth dorsal vertebra (6.3 mm), tenth dorsal
vertebra (6.2 mm), eleventh dorsal vertebra (6.2 mm), twelfth dorsal
vertebra, thirteenth dorsal vertebra, dorsal ribs, gastralia, (sacrum
~31 mm) first sacral vertebra (6.5 mm), second sacral vertebra, first
caudal vertebra, second caudal vertebra (5.3 mm), third caudal vertebra
(~4 mm), fourth caudal vertebra (~4 mm), fifth caudal vertebra (6 mm),
sixth caudal vertebra (6.5 mm), seventh caudal vertebra (7.3 mm),
eighth caudal vertebra (9 mm), ninth caudal vertebra (10.5 mm), tenth
caudal vertebra (11.3 mm), eleventh caudal vertebra (11.3 mm), twelfth
caudal vertebra (~11.5 mm), thirteenth caudal vertebra (~11.5 mm),
fourteenth caudal vertebra (~10.5 mm), fifteenth caudal vertebra (10
mm), sixteenth caudal vertebra (10 mm), seventeenth caudal vertebra
(~10 mm), eighteenth caudal vertebra (~9.5 mm), nineteenth caudal
vertebra (8.5 mm), twentieth caudal vertebra (8 mm), twenty-first
caudal vertebra (7.2 mm), twenty-second caudal vertebra (5.5 mm),
scapulae (42 mm; one incomplete), coracoids (one fragmentary), furcular
fragments, humeri (63.5 mm), radii (54.4 mm), ulnae (55 mm),
scapholunares, pisiforms, semilunate carpals, distal carpal III,
metacarpals I (8.3 mm),
phalanges I-1 (20.5 mm), manual unguals I (~12.5 mm), metacarpals II
(28 mm), phalanges II-1 (15.5 mm), phalanges II-2 (19.4 mm), manual
unguals II (~16 mm), metacarpals III (24.8 mm), phalanges III-1 (6.4
mm), phalanges III-2 (4.2 mm), phalanges III-3 (12.3 mm), manual
unguals III (~9.5 mm), manual claw sheaths, ilia (~32 mm), pubes (48
mm), ischium (~20 mm), femora (52.2 mm), tibiae (71 mm), astragalus,
distal tarsal III, distal tarsal IV, metatarsal I, phalanx I-1 (5.2
mm), pedal ungual I (5.5 mm), partial metatarsal II (~35 mm), phalanx
II-1 (8.2 mm), phalanx II-2 (7 mm), pedal ungual II (12.5 mm),
incomplete metatarsals III (~37 mm), phalanges III-1 (9.6 mm),
phalanges III-2 (9 mm), phalanges III-3 (8.2 mm), pedal unguals III
(8.8 mm), incomplete metatarsal IV (~32.5 mm), phalanx IV-1 (7 mm),
phalanges IV-2 (6.4, 6.6 mm), phalanges IV-3 (4.9 mm), phalanges IV-4
(5.6, 5.8 mm), pedal unguals IV (6.8 mm), pedal claw sheaths,
propatagium, body feathers, remiges, retrices (Haberlein, 1877)
Early Tithonian, Late Jurassic
Upper Eichstatt Member of the Altmuhltal Formation, Germany
(BMMS 500; Solnhofen specimen; sixth specimen; holotype of
Wellnhoferia grandis) (1.1 kg, 530 day old subadult) partial skull (~65
mm), sclerotic plates, partial mandible (~61 mm), axis, third cervical vertebra
(~9 mm), partial fourth cervical vertebra, cervical ribs, partial eleventh dorsal
vertebra, twelfth dorsal vertebra (~8 mm), thirteenth dorsal vertebra (~8 mm),
fourteenth dorsal vertebra (8.8 mm), dorsal ribs, gastralia, (sacrum ~28 mm)
partial first sacral vertebra, second sacral neural spine, third sacral neural
arch, fourth sacral transverse process, fifth sacral neural arch, first caudal
vertebra (5.4 mm), second caudal vertebra (6.2 mm), third caudal vertebra (6.7
mm), fourth caudal vertebra (6.5 mm), fifth caudal vertebra (7.5 mm), sixth
caudal vertebra (9 mm), seventh caudal vertebra (9.7 mm), eighth caudal vertebra
(9.5 mm), ninth caudal vertebra (13 mm), tenth caudal vertebra (14.2 mm), eleventh
caudal vertebra (14.3 mm), twelfth caudal vertebra (15.5 mm), thirteenth caudal
vertebra (15 mm), fourteenth caudal vertebra (~11 mm), partial fifteenth caudal
vertebra (~11 mm), chevrons, incomplete scapulae (~51 mm), coracoids (24.5 mm),
partial furcula, incomplete humeri (83 mm), partial radii (~69 mm), partial
ulnae (~72 mm), partial metacarpals I, phalanges I-1 (28, 28.4 mm), manual unguals
I (17.5 mm), incomplete metacarpals II (~36.6 mm), phalanges II-1 (20.1 mm),
phalanges II-2 (27.5 mm), manual unguals II (17.8 mm), incomplete metacarpals
III, phalanx III-1 (8 mm), phalanges III-2 (6.5 mm), phalanges III-3 (17.8 mm),
manual unguals III (12 mm), manual claw sheaths, ilia (~38 mm), pubes (59.3
mm), partial ischium (~24.5 mm), femora (~67 mm), tibiae (92 mm), fibula (82.4
mm), astragali, calcanea, distal tarsals, metatarsals I (9.9 mm), phalanx I-1
(11 mm), pedal ungual I (~9.8 mm), metatarsus (II 45 mm, III ~47.5 mm, IV 45
mm), phalanges II-1 (12 mm), phalanges II-2 (12.5 mm), pedal unguals II (~14
mm), phalanges III-1 (one proximal; 13.7 mm), phalanx III-2 (11.8 mm), phalanx
III-3 (10.5 mm), pedal ungual III (~10 mm), phalanx IV-1 (10 mm), phalanx IV-2
(8.5 mm), phalanx IV-3 (9.5 mm), pedal ungual IV (~12.6 mm), metatarsal V (10.5
mm), pedal claw sheaths, remiges (Wellnhofer, 1988)
(JM SoS 2257; Eichstatt specimen; fifth specimen; holotype of Archaeopteryx
recurva) (.29 m, 180 g; 110 day old juvenile) skull (39 mm), sclerotic ring,
mandible (36.5 mm), atlas, axis (3.7 mm), third cervical vertebra (~4.5 mm),
fourth cervical vertebra (~6 mm), fifth cervical vertebra (~7 mm), sixth cervical
vertebra (~5.5 mm), seventh cervical vertebra (~5 mm), eighth cervical vertebra
(~5 mm), ninth cervical vertebra (~4.5 mm), tenth cervical vertebra (~3.5 mm),
cervical ribs, first dorsal vertebra (4 mm), second dorsal vertebra, partial
third dorsal vertebra, partial fifth dorsal vertebra (4 mm), sixth dorsal vertebra
(4 mm), seventh dorsal vertebra (~4 mm), eighth dorsal vertebra (4.5 mm), ninth
dorsal vertebra (~5 mm), tenth dorsal vertebra (~4.5 mm), eleventh dorsal vertebra,
twelfth dorsal vertebra (4 mm), thirteenth dorsal vertebra (4 mm), dorsal ribs,
gastralia, (sacrum ~16.5 mm), first sacral centrum (3.7 mm), second sacral centrum
(3.4 mm), third sacral centrum (3.1 mm), fourth sacral centrum (3.2 mm), fifth
sacral centrum, first caudal vertebra (3 mm), second caudal vertebra (3.3 mm),
third caudal vertebra (3.5 mm), fourth caudal vertebra (~4 mm), fifth caudal
vertebra (4.7 mm), sixth caudal vertebra (5.4 mm), seventh caudal vertebra (6.4
mm), eighth caudal vertebra (7.2 mm), ninth caudal vertebra (7.9 mm), tenth
caudal vertebra (8.4 mm), eleventh caudal vertebra (8.5 mm), twelfth caudal
vertebra (8.6 mm), thirteenth caudal vertebra (8.3 mm), fourteenth caudal vertebra
(8.3 mm), fifteenth caudal vertebra (8 mm), sixteenth caudal vertebra (7.7 mm),
seventeenth caudal vertebra (7.4 mm), eighteenth caudal vertebra (6.9 mm), nineteenth
caudal vertebra (6.6 mm), twentieth caudal vertebra (~5.5 mm), twenty-first
caudal vertebra (4.2 mm), twenty-second caudal vertebra (4 mm), twenty-third
caudal vertebra, seventeen chevrons, incomplete scapulae, partial coracoids,
humeri (41.5 mm), radii (35 mm), ulnae (36.5 mm), scapholunares, pisiform, semilunate
carpals, distal carpals III, metacarpals I (one proximal; 5.5 mm), phalanges
I-1 (one distal; 15.6 mm), manual unguals I (9.8 mm), metacarpals II (one incomplete;
17.8 mm), phalanges II-1 (10.2 mm), phalanges II-2 (15.1 mm), manual unguals
II (10.8 mm), metacarpals III (one incomplete; 16.5 mm), phalanx III-1 (4.2
mm), phalanx III-2 (3 mm), phalanx III-3 (9.8 mm), manual unguals III (6.5 mm),
manual claw sheaths, incomplete ilium (~20 mm), pubes (31.5 mm), ischia (14.5
mm), femora (37 mm), tibiae (~53 mm), fibula (50.5 mm), astragali, distal tarsal
III, distal tarsal IV, metatarsals I, phalanges I-1 (5.5 mm), pedal unguals
I (3.5 mm), partial metatarsals II (28.3 mm), phalanges II-1 (~7.1 mm), phalanges
II-2 (7 mm), pedal unguals II (5.8 mm), metatarsals III (30.2 mm), phalanges
III-1 (one proximal; 9 mm), phalanges III-2 (8 mm), phalanges III-3 (7 mm),
pedal unguals III (5.4, 4.8 mm), metatarsals IV (27.3 mm), phalanges IV-1 (6.1
mm), phalanges IV-2 (5 mm), phalanges IV-3 (4.6, 4.7 mm), phalanges IV-4 (4.9
mm), pedal unguals IV, metatarsals V (one partial; 6.5 mm), pedal claw sheaths, remiges, retrices (Wellnhofer, 1974)
(WDC-CSG-100; Thermopolis specimen; tenth specimen) (260 g, 1 year old juvenile)
skull (52.9 mm), sclerotic ring, mandibular fragment, five dentary teeth, partial
hyoid, six cervical vertebrae, partial seventh dorsal vertebra, partial eighth
dorsal vertebra, partial ninth dorsal vertebra, partial tenth dorsal vertebra,
partial eleventh dorsal vertebra, partial twelfth dorsal vertebra, partial thirteenth
dorsal vertebra, dorsal ribs, gastralia, first sacral centrum, second sacral
centrum, third sacral centrum, fourth sacral centrum, fifth sacral fragment,
first caudal vertebra, second caudal vertebra (3.8 mm), third caudal vertebra
(4.2 mm), fourth caudal vertebra (4.2 mm), fifth caudal vertebra (5.4 mm), sixth
caudal vertebra (6.5 mm), seventh caudal vertebra (7.9 mm), eighth caudal vertebra
(9.5 mm), ninth caudal vertebra, tenth caudal vertebra (~10 mm), eleventh caudal
vertebra (11.1 mm), twelfth caudal vertebra (10.9 mm), thirteenth caudal vertebra
(10.9 mm), fourteenth caudal vertebra (10.6 mm), fifteenth caudal vertebra (10.6
mm), sixteenth caudal vertebra (10.1 mm), seventeenth caudal vertebra (9.1 mm),
eighteenth caudal vertebra (9.1 mm), nineteenth caudal vertebra, partial twentieth
caudal vertebra, chevrons, scapulae (35 mm), coracoids, furcula, humeri (one
proximal; 56.9), radii, ulnae (50.9 mm), semilunate carpal, metacarpals I (6.6
mm), phalanges I-1 (19.5 mm), manual unguals I, metacarpals II (23.5 mm), phalanges
II-1 (12.8 mm), phalanges II-2 (18.6 mm), manual unguals II, metacarpals III
(22 mm), phalanges III-1 (4.8 mm), phalanges III-2 (4.2 mm), phalanges III-3
(12.9 mm), manual ungual III, manual claw sheaths, partial ilium, pubes, ischium,
femora (one incomplete; 50.3 mm), tibiae (one incomplete; 74.6 mm), fibula,
astragali, calcaneum, metatarsals I, phalanges I-1 (6.1 mm), pedal unguals I,
metatarsals II (35.1 mm), phalanges II-1 (10.6 mm), phalanges II-2 (one proximal;
8.8 mm), pedal ungual II, metatarsals III (39.6 mm), phalanges III-1 (one proximal;
10.8 mm), phalanx III-2 (9.6 mm), phalanx III-3 (7.8 mm), pedal ungual III,
metatarsals IV (36.3 mm), phalanges IV-1 (7.5 mm), phalanges IV-2 (one proximal;
6.6 mm), phalanx IV-3 (5.6 mm), phalanx IV-4 (5.6 mm), pedal ungual IV, pedal
claw sheaths, remiges, retrices (Mayr et al., 2005)
(Pohl coll; on long term loan to BSP; Altmuhl specimen; eleventh specimen) premaxillae, jugal,
dentaries, partial ?splenial, incomplete ?surangular, ?articular, axis, third
cervical vertebra, fourth cervical vertebra, fifth cervical vertebra, sixth
cervical vertebra, seventh cervical vertebra, eighth cervical vertebra, ninth
cervical vertebra, tenth cervical vertebra, cervical ribs, first dorsal vertebra,
second dorsal vertebra, third dorsal vertebra, fourth dorsal vertebra, fifth
dorsal vertebra, sixth dorsal vertebra, seventh dorsal vertebra, eighth dorsal
vertebra, ninth dorsal vertebra, tenth dorsal vertebra, eleventh dorsal vertebra,
twelfth dorsal vertebra, thirteenth dorsal vertebra, dorsal ribs, gastralia,
partial sacrum, first caudal vertebra, second caudal vertebra, third caudal
vertebra, fourth caudal vertebra, fifth caudal vertebra, sixth caudal vertebra,
eighth caudal vertebra, ninth caudal vertebra, tenth caudal vertebra, eleventh caudal vertebra,
twelfth caudal vertebra, thirteenth caudal vertebra, fourteenth caudal vertebra,
fifteenth caudal vertebra, sixteenth caudal vertebra, seventeenth caudal vertebra,
eighteenth caudal vertebra, nineteenth caudal vertebra, twentieth caudal vertebra,
twenty-first caudal vertebra, chevrons, scapula (44.4 mm), incomplete coracoid
(13.3 mm), partial furcula, incomplete humerus (65.6 mm), radius (~62 mm), ulna
(~62.5 mm), pisiform, semilunate carpal, partial metacarpal I (~9.9 mm), phalanx
I-1 (23.3 mm), manual ungual I, metacarpal II (~31.5 mm), phalanx II-1 (18.6
mm), phalanx II-2 (19.6 mm), manual ungual II, metacarpal III (~31.5 mm), phalanx
III-1 (6.4 mm), phalanx III-2 (5.3 mm), phalanx III-3 (~12.3 mm), manual ungual
III, manual claw sheaths, incomplete ilium (~37.1 mm), pubes (52.2 mm), ischium
(23.3 mm), femora (one distal; 55.3 mm), tibiae (76.3 mm), partial fibulae,
astragali, metatarsals I (~7.1 mm), phalanges I-1 (7.9, 7.4 mm), pedal unguals
I, metatarsals II (36.8, 38.4 mm), phalanges II-1 (9.9, 9.9 mm), phalanges II-2
(9.7, 9.5 mm), pedal unguals II, metatarsals III (40.3, 41.4 mm), phalanges
III-1 (11.6, 11.2 mm), phalanges III-2 (10 mm), phalanges III-3 (8.8 mm), pedal
unguals III, metatarsal IV (~37.9 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal unguals IV, pedal claw sheaths, body feathers, remiges,
retrices (Ravasz, online 2011; described by Foth et al., 2014)
Early Tithonian, Late Jurassic
M�rnsheim Formation, Germany
(SNSB BSPG VN-2010/1; Daiting specimen; The Phantom; eighth specimen; holotype of Archaeopteryx albersdoerferi)
(320 day old juvenile) incomplete skull, mandibles (one incomplete, one
partial), scapulae (35.5 mm), incomplete coracoids, furcula, humeri
(one incomplete; 55.5 mm), radius (46.7 mm), ulna (47.5 mm),
scapholunare, pisiform, incomplete carpometacarpus, phalanx I-1,
partial manual ungual
I, proximal femur, fibular fragment (Mauser, 1997)
Early Tithonian, Late Jurassic
Painten Formtion, Germany
(Dinosaurier Freiluftmuseum Altmuhltal coll.; DNWK 02924; Schamhaupten
specimen; twelfth specimen) skull (56 mm), mandibles (45.5 mm), mid
cervical vertebra, eighth cervical vertebra (8.1 mm), ninth cervical
vertebra (6.1 mm), tenth cervical vertebra (5.3 mm), first dorsal
vertebra, second dorsal vertebra, third dorsal vertebra, fourth dorsal
vertebra, fifth dorsal vertebra, sixth dorsal vertebra (6.2 mm),
seventh dorsal vertebra (6.2 mm), eighth dorsal vertebra (6 mm), ninth
dorsal vertebra (6 mm), tenth dorsal vertebra, eleventh dorsal
vertebra, twelfth dorsal vertebra, thirteenth dorsal vertebra, dorsal
ribs, gastralia, partial sacrum, proximal caudal vertebrae, fourth
caudal vertebra (~5.8 mm), fifth caudal vertebra (~6.1 mm), sixth
caudal vertebra (6.2 mm), seventh cuadal vertebra (7.5 mm), eighth
caudal vertebra (9 mm), ninth caudal vertebra (9.7 mm), tenth caudal
vertebra (9.9 mm), eleventh caudal vertebra (10.1 mm), twelfth caudal
vertebra (10.1 mm), thirteenth caudal vertebra (9.9 mm), fourteenth
caudal vertebra (9.9 mm), fifteenth caudal vertebra (9.7 mm), sixteenth
caudal vertebra (9.3 mm), seventeenth caudal vertebra (9 mm),
eighteenth caudal vertebra (8.7 mm), nineteenth caudal vertebra (6.8
mm), twentieth caudal vertebra (5.3 mm), twenty-first caudal vertebra
(3.2 mm), twenty-second caudal vertebra (2.4 mm), chevrons, scapulae
(43 mm), coracoid, incomplete furcula, humeri (61 mm), radii (54.4 mm),
ulnae (55 mm), semilunate carpal, metacarpals I (6.8 mm), phalanges I-1
(20.7 mm), manual unguals I (9.4 mm),
metacarpals II (28.2 mm),
phalanges II-1 (16 mm), phalanges II-2 (19.3 mm), metacarpals III (27.2
mm),
phalanges III-1 (7.1 mm), phalanges III-2 (4.6 mm), phalanges III-3 (12
mm), manual unguals III (7.5 mm), manual claw sheaths, incomplete
pubes, incomplete ischium, incomplete femora (~53 mm), tibiae (66 mm),
fibulae, astragalus, calcaneum, distal tarsal IV, metatarsals I (8.2
mm), phalanges I-1 (7 mm), pedal unguals I (6.3 mm), metatarsals II
(31.6 mm),
phalanges II-1 (8.4 mm), phalanges II-2 (8.3 mm), pedal unguals II (9.8
mm), metatarsals III (34 mm),
phalanges
III-1 (10.5 mm), phalanges III-2 (9.2 mm), phalanges III-3 (8 mm),
pedal unguals III (8.1 mm), metatarsals IV (33.1 mm), phalanges IV-1
(7.3 mm), phalanges IV-2 (6.5 mm), phalanges IV-3 (5.8 mm), phalanges
IV-4 (5.9 mm), pedal unguals IV (6.2 mm), pedal claw sheaths,
metatarsals V (Rauhut, Foth and Tischlinger, 2018)
Diagnosis- (after Elzanowski, 2002) 8-9 maxillary teeth (also in Dromaeosaurus);
premaxillary and anterior dentary teeth have the basal third to half of
the crown with straight to slightly convex mesial and distal margins
that slightly converge apically, before the apical part of the crown
forms a bulbous mesial and less marked distal expansion.
(after Rauhut et al., 2018) marked incision between the postorbital and quadratojugal processes of the jugal (also in Microraptor); depressed rim around the posterior margin of the trigeminal foramen (unknown in other archaeopterygids).
(after Kundrat et al., 2019) anterior-most extension of promaxillary
fenestra situated between fifth and sixth maxillary teeth; 10-12
dentary teeth.
Other diagnoses-
Elzanowski (2002) listed several characters of his
Archaeopterygidae that are plesiomorphic (four premaxillary teeth;
premaxilla projects anterior to mandible; 23 presacral vertebrae; five
sacral vertebrae; no hypocleidium on furcula). An absent external
mandibular fenestra is probably true in other archaeopterygids, a mid
dorsal ischial process is common in basal paravians.
Rauhut et al. (2018) note 'a longitudinal groove on the medial side of the suborbital process' of the jugal is also present in Anchiornis,
so this is an archaeopterygid character. Similarly, they state
'quadratojugal process of the jugal very slender, less than half the
height of the minimal height of the suborbital process' is present in Anchiornis, Xiaotingia and Microraptor. The presence of 'nine cervical and fourteen dorsal vertebrae' is left uncertain here given the poor preservation of Archaeopteryx' cervicodorsal vertebrae and the condition in related taxa.
London specimen and feather-
Meyer (1861a) first mentioned and described a feather (BSP 1869 VIII 1;
MB.Av.100) discovered in Spring or Summer 1861 (published August 16th), but did not name
it. Wellnhofer (2009) notes that
a discovery in 1860 is sometimes reported but that there is no evidence
it was that long prior to the September 15 publication of Meyer
1861a. Meyer later (1861b) referred to the feather and mentioned the
then undescribed London specimen noticed in Haberlein's collections the Summer of that year, stating that
"For the denomination of the animal I consider the term Archaeopteryx lithographica as appropriate", but exactly
which specimen (if not both) he considered "the animal" is ambiguous.
The title suggests the feather is the holotype, but this was written by an anonymous
editor. In his later (1862a) paper, he indicates the name Archaeopteryx lithographica
was meant to be tied to the feather, making it the holotype ("The fossil
feather presented by me may come from a similar animal, for which I have chosen
the denomination Archaeopteryx lithographica"). This was misunderstood
by many subsequent authors, including de Beer (1954), who have viewed the London
specimen as the holotype instead. Wagner (1962a) described the London specimen
as a new genus of pterosaur- Griphosaurus,
as Meyer's 1861b paper had not been published at the time of Wagner's
presentation (November 9, 1861). While Wagner's paper was dated 1861,
it wasn't published until January 20th, 1862 (Swinton, 1960), but an
English translation was published in April 1862 (Wagner, 1862b).
Woodward (1962) used the name Archaeopteryx lithographica, but
stated Owen would describe the London species under the name Griphornis longicaudatus.
A footnote indicates Owen had decided to retain the name Archaeopteryx
though, as indeed did happen in his 1863 paper. More confusingly, Woodward uses
the name "Griphosaurus problematicus Wagner, 1861" in the plate
label for the London specimen, though Wagner never used a species name in his
1862 ("1861") publication. Owen (1863a) made the London specimen the
holotype of his new species Archaeopteryx macrura (misspelled macrurus
in his 1863a publication, but corrected in 1863b), since he considered its feathers
different from the A. lithographica holotype. Several later authors followed
Owen's usage (Dames, 1884; Lambrecht, 1933). The new combination Griphosaurus
longicaudatus has been listed as deriving from Owen (1862), but Buhler and
Bock (2002) indicate this was a mistaken attribution by Brodkorb (1963). Petronievics
(1921) used the name Archaeopteryx oweni for the London specimen without
justification, so it is clearly a junior synonym. In 1960, Swinton petitioned
the ICZN to place Archaeopteryx lithographica on the Official List of
Generic/Specific Names in Zoology, and to add Griphosaurus, Griphornis
longicaudatus, Griphosaurus problematicus, Archaeopteryx macrurus
and Archaeopteryx oweni to the Official Index of Rejected and Invalid
Generic/Specific Names in Zoology, which was upheld by Riley and China in 1961
as Opinion #607. Eisenmann (1974) in a comment to the ICZN proposed ruling the London specimen is the type of Archaeopteryx lithographica, but the issue was not taken up by the commission at the time. Bock and Buhler (2007) petitioned the ICZN to
make the London specimen the neotype of Archaeopteryx lithographica,
since authors have interpreted Meyer's intent in different ways, and
nearly all references have based the taxon on the more diagnostic
skeleton as opposed to the feather. Kadolsky (2008) further noted
"the skeleton find (the later BMNH specimen no. 37001) is not
described, diagnosed or characterised by even a single word, nor is an
indication to such characterisation given" so that Archaeopteryx lithographica
the London specimen would be a nomen nudum if based on this
publication, so asked the ICZN "to use its plenary power to rule that
both the generic and specific names Archaeopteryx and lithographica have been made available by von Meyer, 1861b." The issue was decided by the ICZN in 2011 as Opinion 2283 in
favor of making the London specimen the neotype. The London specimen was most recently
described in detail by de Beer (1954).
The feather- The feather was redescribed
by Griffiths (1996), who noted it differed from the London and Berlin specimens
in being smaller, broader and more asymmetric. He proposed that both taphonomic
and taxonomic explanations were possible, and his analysis indicates the feather
cannot be definitively referred to Archaeopteryx lithographica. Benton and Gower (2002) reveal Walker had written a letter in 1985 also
wondering about the possibility the feather was taxonomically distinct from
the skeletons, though it was never published. The feather was reinterpreted as a retrix by Foth
and Rauhut (2013) and by Kaye et al. (2019) as a secondary much shorter than Archaeopteryx'.
Most recently, Carney et al. (2019; published in detail in 2020)
convincingly argued the feather is an upper major primary covert and
found it was identical in anatomy and size to that of the Altmuhl Archaeopteryx specimen. Unfortunately they dismissed an alternative taxonomic identification by proposing all Solnhofen paravians are Archaeopteryx, whereas this site supports the validity of 'anchiornithine' Ostromia and unenlagiine Alcmonavis. Yet given the differences between Archaeopteryx and Anchiornis feathers, and that unenlagiines are even more distant phylogenetically, a referral to Ostromia or the larger Alcmonavis is here considered less likely.
Berlin specimen-
The Berlin specimen was probably discovered in 1875 or 1874
(Tischlinger, 2005), though first announced in May 1877 (Haberlein,
1877) and variously regarded as conspecific with the London specimen
(Dames, 1884), or different due to size (Seeley, 1881). Later, Dames
(1897) made the holotype of a new species- Archaeopteryx siemensii due to dental and pelvic characters. Subsequently,
Petronievics (as a footnote in Petronievics and Woodward, 1917) separated it
further as a new genus- Archaeornis. This was due to his view the expanded
calcitic mass of the London specimen was not homologous to the pubic boot of
the Berlin specimen, that the latter lacked a pubic symphysis, and other unstated
pectoral and pelvic differences. Additional differences later provided by Petronievics
(1921, 1925) include teeth with a circular cross section, proximal carpals ossified,
metacarpal III with cylindrical section, unfused scapulocoracoid, narrow coracoid,
unfused tibiotarsus, pubis and ischium at a greater angle, obturator foramen
absent in pubis, straight ischium, and ischium poorly ossified distally. Petronievics
later (1927, 1950) went so far as to place Archaeopteryx as a pan-paleognath
and Archaeornis as a pan-neognath. Nopcsa (1923) stated most of these
characters were preservational (carpal number; pubic foot size; ischial shape)
or ontogenetic (length of pubic symphysis; ischial ossification; tibiotarsal
fusion), while others were misinterpretations (tooth cross section; scapulocoracoid
fusion; pubic angle; obturator foramen). He and de Beer (1954) synonymized it
with A. lithographica, which has been the consensus until the 2000s. In
2002, Elzanowski reinstated Archaeopteryx siemensii based on the smaller
size, lack of a cuppedicus fossa on the ilium, lack of a ventral hook on the
ilial preacetabular process, and pedal unguals without flexor tubercles. Senter
and Robins (2003) disagreed, finding flexor tubercles in all Archaeopteryx
specimens, and believing the ilial features to become more developed
with age. Mayr et al. (2007) confirmed the flexor tubercles are more
poorly developed and noted the metatarsus was more slender, but did not
comment on ilial morphology. Wang et al. (2017) confirmed the
presence of a propatagium.
Maxberg Specimen- The Maxberg specimen was discovered in 1956 and described
in 1959 by Heller. It was privately owned by Opitsch, though on display at the
Maxberg Museum from 1959-1982. Unfortunately, when Opitsch died in 1991, the
specimen could not be found and was seemingly stolen (Abbott, 1992). Mayr et
al. (2007) assigned it to A. lithographica (as opposed to A. siemensii)
due to its robust metatarsus.
Eichstatt specimen- The Eichstatt specimen was recognized in March 1951 and
originally identified as a juvenile Compsognathus. It was initially
described by Mayr (1973) as perhaps a new species of Archaeopteryx, and
later in detail by Wellnhofer (1974) as a juvenile A. lithographica.
Howgate (1984) separated it from the other specimens then known as a new species,
Archaeopteryx recurva. This was based on the small size, recurved teeth,
supposedly absent furcula, more vertical pubis, short pubic symphysis, elongate
tibia (1.42 times femoral length compared to 1.30-1.38), elongate metatarsus
(1.67 times femoral length compared to 1.43-1.47) and unfused metatarsus. He
later used these characters to separate it further as a new genus- Jurapteryx.
However, almost all later authors have believed it to be a juvenile specimen
of Archaeopteryx lithographica (Paul, 1988; Wellnhofer, 1992), with the
furcula missing due to taphonomy and the pubes of the London and Berlin specimens
rotated too far posteriorly. Houck et al. (1990) determined the limb proportions
of Archaeopteryx specimens were consistant with allometric variation
that would be expected due to ontogeny, which has been confirmed for subsequent
specimens by Senter and Robins (2003) and Bennett (2008). Christiansen's (2006)
allometric study concluded the data supported multiple species at least as well
as it did a single species, but his rationale was flawed (Bennett, 2008). Elzanowski
(2002) could not confirm separation of the Eichstatt specimen as a distinct
species, while Senter and Robins (2003) noted theropods such as Coelophysis
decrease curvature of teeth with ontogeny, suggesting the same explanation for
the Eichstatt specimen's teeth.
Solnhofen specimen- The Solnhofen specimen was recognized as Archaeopteryx in November 1987 in Muller's private
collection after being initially identified as a Compsognathus until
it was examined by Wellnhofer and described by him in 1988 as a new
specimen of Archaeopteryx lithographica.
Muller claimed it was probably recovered 10-15 years prior to 1987, but
a litigant onvolved in ownership of the fossil claimed it was actually
found in 1985. Longer descriptions of the material appeared later
in 1988 and in 1992. In 2001, Elzanowski separated the specimen as a
new taxon - Wellnhoferia grandis. This was based on the large size, short
tail as reconstructed (~16-17 vertebrae), unfused scapulocoracoid, shorter manual
ungual I compared to phalanx I-1 (33%), manual phalanges I-1 and II-2 deeper,
manual phalanges III-1 and III-2 sutured, more retroverted pubis (~128 degrees),
metatarsals II and IV subequal in length, pedal phalanx II-2 longer than II-1,
short pedal digit IV with only four phalanges, pedal ungual IV longest phalanx
in digit, and pedal ungual IV straight with flexor tubercle widely separated
from its base. In 2002, he added a proximally tapering metatarsal II to the
diagnosis. Senter and Robins (2003) agree it is distinct, and Mayr et al. (2007)
agree it is distinct from the Berlin and Munich specimens, but disputed differences
between it and the London specimen (synonymizing it with A. lithographica).
In particular, they note the proportions of manual digit I phalanges and number
of phalanges in pedal digit IV are not preserved in the London specimen, and
the tail length is uncertain in the Solnhofen specimen (it was estimated to
be short by Elzanowski due to the rapid decrease in central length distally).
The specimens are also both larger than other specimens and share additional
characters (premaxillary teeth with constricted crowns; similar limb proportions;
stout metatarsus; well developed flexor tubercles on pedal unguals). However,
the London specimen has a fused scapulocoracoid, pedal phalanx II-2 equal in
length to II-1, a more elongate pedal digit IV, and a curved pedal ungual IV.
Munich specimen- The next specimen was discovered on August 3 1992 and first called
the Solnhofen-Aktien-Verein specimen, then later the Munich specimen. It was
described in 1993 as a new species of Archaeopteryx, A. bavarica.
This was based on the small size, twelve dentary teeth, dentary interdental
plates, ossified sternum, elongate tibia (1.48 times femoral length), hindlimb
elongate compared to humerus (3.56 times). Elzanowski (2002) added further distinguishing
characters- anterior dentary tooth crowns compressed; third and fourth cervical
neural spines tall, equaling about a third of vertebral height; ulna elongate
compared to humerus (96%); ilium lacking ventral hook on preacetabular process;
no cuppedicus fossa on ilium; pedal unguals lack flexor tubercles. Senter and
Robins (2003) determined the tibial, hindlimb and ulnar lengths were all explainable
by allometry, believed the ilial features could be absent due to ontogeny, noted
tooth compression could be too (as in tyrannosaurids), and stated pedal ungual
flexor tubercles are actually present. Wellnhofer and Tischlinger (2004) determined
the supposed sternum of the Munich specimen is really a coracoid. Mayr et al.
(2007) noted their new specimen is intermediate between the Munich and Berlin
specimens in size, hindlimb and ulnar proportions, while its tibia is longer
than the Munich specimen. Considering the misidentified sternum, and that the
ilial and pedal ungual characters are supposedly shared with the Berlin specimen,
they synonymized A. bavarica with A. siemensii.
Daiting specimen- The eighth or
Daiting specimen was discovered in 1990 and initially identified as a
pterosaur, but not publicized until a cast was displayed in 1996. It
was originally noted by Mauser (1997) in a brief article and was
described by Tischlinger (2009) then in more detail by Kundrat et al.
(2019). Albersd�rfer bought it from a private collector in 2009
and placed it on long term loan to the BSPG as SNSB BSPG
VN-2010/1. It is the youngest of described specimens, the same
age as Alcmonavis. Kundrat et al. (2014) believed fused internasal
and interfrontal contacts and "numerous other cranial features" suggested
this could be a separate species from the Eichstatt and Thermopolis specimens, and described it as Archaeopteryx albersdoerferi
in 2019. Diagnostic characters listed by Kundrat et al. are-
angle between the nasal process and ventral margin of maxilla >50
degrees; antorbital fenestra and combined maxillary/promaxillary
fenestrae both higher than wide; anterior lacrimal process slightly
longer than posterior process; pneumatic recess present at anterior end
of jugal; incomplete postorbital bar; Y-shaped quadratojugal with
reduced dorsal process and expanded posterior process; quadrate with
incompletely bilobed head, massive lateral vertical rim and enlarged
medial condyle separated from lateral condyle by shallow depression;
maxillary and pterygoid palatine processes of equal length; heterodont
maxillary dentition comprising asymmetrical anterior and symmetrical
posterior crowns; teeth labiolingually compressed; coracoid with
lateral process as long as half of mid-coracoid width; coracoid with
tuber that does not project beyond dorsal margin; deltopectoral portion
of humeral shaft strongly angled (rather then curved) posteriorly;
distal carpal III fused with metacarpal II; semilunate carpal fused
with the metacarpals I and II. It was recovered as either an
archaeopterygid (Xu et al. 2011 TWiG parsimony analysis; Godefroit et
al. 2011 Cau analysis) or closer to Aves (Xu et al. 2011 TWiG Bayesian
analysis; Turner et al., 2012 TWiG analysis).
New specimens- The ninth specimen was initially announced by Roper (2004) after it was collected
in Spring of that year. Also known as the chicken wing or Ottmann and Steil specimen, it has immature bone grain. Wellnhofer
and Roper (2005) described it in detail, assigning it to A. lithographica
instead of A. bavarica (based on ulnar proportions).
The Thermopolis specimen was found in the estate of a fossil collector
in the 1970s, but not offered to the scientific community until
2001. It was briefly described in 2005 (Mayr, 2005; Mayr et al.,
2005) before being described in detail by Mayr et al. (2007). Though
initially only assigned to Archaeopteryx
sp. (Mayr et al., 2005), they later assigned it to A. siemensii,
in which they also include the Munich specimen. This was based on small size,
slender metatarsus, poorly developed pedal ungual flexor tubercles, as well
as characters differing from the Solnhofen specimen (long manual ungual I; metatarsals
II and IV not subequal; pedal digit IV with five phalanges; pedal ungual IV
shorter than phalanx IV-1). They further note that the ischium exhibits a different
morphology than the London specimen, which added evidence to separate siemensii
from lithographica.
An eleventh specimen or Altmuhl specimen was announced at the Munich Show - Mineralientage M�nchen
in 2011 (artdaily.cc, online 2011). The skeleton is essentially complete except for
most of the skull and one pectoral girdle and arm. Foth and Rauhut (2013) describe
the feathers in an abstract, and officially in Foth et al. (2014). Foth et al.
only refer the specimen to Archaeopteryx sp. noting the taxonomic controversy
surrounding the genus.
The twelfth specimen or Schamhaupten specimen was found in Summer of
2010 and described in depth by Rauhut et al. (2018). It is the
oldest of published specimens, from the earliest Tithonian Painten
Formation, and was referred to Archaeopteryx sp.. It's notable for having obvious postorbitals and prefrontals.
A thirteenth specimen, SNSB-BSPG 2017 I 133 or the Muhlheim specimen,
was found in 2017. It was described by Rauhut et al. (2019) as a
new taxon of avialan closer to Pygostylia, Alcmonavis poeschli.
Resolving the species problem-
As can be seen from the above discussions, recent papers have placed
the known specimens in one (Paul, 2002), two (Senter and Robins, 2004;
Mayr et al., 2007) or four (Elzanowski, 2002) species, often based on
similar characters. These will be examined below (minus the twelfth
specimen, described after text was written) to determine how many
species are based on good evidence.
Size is not evidence of multiple species, as no specimen is obviously adult,
and young theropods (including basal birds) were precocial in regard to their
ossification timing (e.g. Scipionyx, juvenile Yixian enantiornithines).
The latter disproves Howgate's (1984) suggestion that well ossified elements
indicate the Eichstatt specimen is an adult. More recently, Erickson et al.
(2009) have analyzed the histology of the Munich specimen and noted this and
other specimens (including the Solnhofen specimen) show fibrous surface texture
with long striae on long bones typical of young individuals.
Tooth recurvature is polymorphic in every specimen (Howgate, 1984). In the London
specimen, the third premaxillary tooth and maxillary teeth 1, 3 and 6 are straight,
but an isolated premaxillary tooth is recurved. In the Berlin specimen, the
first premaxillary tooth, and maxillary teeth 1 and 6 are straight, but premaxillary
teeth 2-4 and maxillary teeth 1 and 3-5 are recurved. In the Eichstatt specimen,
maxillary teeth 3 and 9, and dentary teeth 2, 4-6 and 10-11 are straight, but
all premaxillary teeth, maxillary teeth 1-2 and 4-6, and dentary tooth 7 are
recurved. In the Thermopolis specimen, premaxillary teeth 1-2 are straight,
but premaxillary teeth 3-4 and maxillary teeth 4-8 are recurved. In the Solnhofen
specimen, premaxillary teeth 1-2 and 4, maxillary teeth 4-7 and perhaps four
dentary teeth (4-7?) are straight, but premaxillary tooth 3 and maxillary teeth
2 and 3 are recurved. In the Daiting specimen, three maxillary teeth and two
dentary teeth are straight, while one dentary tooth is recurved. In the 11th
specimen, premaxillary teeth 1-2 and most dentary teeth are straight but premaxillary
teeth 3-4 and at least one dentary tooth are recurved. The narrow tips of the
Eichstatt specimen's teeth are directly due to the more acute angle of a recurved
crown, and some of its teeth (e.g. second premaxillary tooth) have apical wear
facets as in the London specimen. The London's and Munich's specimens teeth
appear flatter because they are exposed in lingual view, unlike the Berlin specimen.
The presence of one less dentary tooth in the Munich specimen compared to the
Eichstatt specimen is within the normal range of interspecific variation in
theropods (e.g. Currie, 2003). The presence of interdental plates in the Munich
specimen is probably a misinterpretation (Martin and Stewart, 1999), so it it
similar to the London and 11th specimens in this regard. So the only difference
is in the frequency of tooth recurvature, which is actually less in the Eichstatt
specimen than the Berlin and Thermopolis specimens, while the sample size of
the London specimen (four teeth) is too low for useful comparison. A large sample
of Saurornitholestes teeth shows generally decreasing curvature with
increasing size, which shows curvature could be ontogenetic as described by
Howgate (1984) for Varanus and Senter and Robins (2004) for Coelophysis.
There are a number of cranial differences between the Berlin and Eichstatt specimens
which may be due to ontogeny (the Berlin specimen has a more convex ventral
maxillary edge, differently sized and placed maxillary fenestrae, deeper dentary;
similar to ontogenetic changes in tyrannosaurids), though if the subnarial nasal
process being absent isn't preservational, it may be of taxonomic importance.
The convexity doesn't seem to be shared by the Solnhofen or Thermopolis specimens.
The Thermopolis specimen has a nasal subnarial process and maxillary fenestrae
that resemble the Eichstatt specimen's more, but the latter are still differently
shaped.
The neural spines of cervical vertebrae three and four are comparable in height
in the Eichstatt (3rd 28% of vertebral height; 4th 28%), Berlin (3rd 29%; 4th
28%) and Munich (3rd 29%; 4th 30%) specimens. While it's true the tail of the
Solnhofen specimen is broken through the fifteenth caudal, and thus the total
number of vertebrae is uncertain, Elzanowski (2001) based his lower estimate
on the sharp decrease in centrum length between the thirteenth and fourteenth
caudal (the latter is 73% of the former). This compares to 95% in the London
specimen, 92% in the Munich specimen, ~91% in the Berlin specimen, 100% in the
Eichstatt specimen, 97% in the Thermopolis specimen and ~97% in the 11th specimen.
Contrary to Elzanowski though, the fifteenth caudal is not necessarily as short,
since only the proximal half is preserved. Other Archaeopteryx specimens
can have isolated shorter vertebrae before the tail tip, for instance caudal
17 of the Munich specimen is 74% as long as caudal 16, even though caudals 18
and 19 are 88% and 105% as long as caudal 16. While Elzanowski argues the sudden
thinness of these last two vertebrae shows they were near the tail tip, Wellnhofer
(1992) notes the tail is twisted at caudals 12-13 to show the following vertebrae
in dorsal aspect. As in the Thermopolis specimen, distal caudal vertebrae are
taller than they are wide, explaining the thinness. Thus the tail of the Solnhofen
specimen was not necessarily shorter than the others'. One vertebral difference
in specimens that does seem real is the height of the dorsal neural spines,
which using the twelfth dorsal vertebra for comparison are 28% of vertebral
height in the Eichstatt specimen, 26% in the Solnhofen specimen, 38% in the
Munich specimen, ~29% in the Berlin specimen, ~29% in the Maxberg specimen and
~38% in the 11th specimen. Also, the Thermopolis specimen has its last dorsal
sutured to the sacrum, giving it six sacral vertebrae. This is certainly not
the case for the Eichstatt, Maxberg and London specimens, and doesn't seem to
be for the Munich specimen either. The Eichstatt specimen seems to lack caudal
neural spines, wheras they are present in the Munich, Solnhofen and 11th specimens.
The scapula and coracoid are apparently fused in the London specimen, tightly
connected in the Berlin and Maxberg specimens, and more loosely connected in
the Eichstatt, Solnhofen, Daiting and 11th specimens. They are also unfused
in the Munich and Thermopolis specimens. However, scapulocoracoid fusion is
variable within other taxa as well, including Struthiomimus, Dromiceiomimus
and Caudipteryx. Maniraptoran coracoids are complex bones, which makes
comparing their shapes difficult. For instance, the right coracoid of the Thermopolis
specimen and left coracoid of the London specimen (as illustrated by Ostrom,
1976) appear to be short, but that's probably because their proximal portion
is bent into the sediment. The proximal portion is illustrated in the London
specimen by Petronievics and Woodward (1917), and also makes the coracoid elongate
in the Munich specimen, while the bend can be observed in the lateral views
of the Solnhofen and Thermopolis left coracoids. The medial margin of the London
specimen's coracoid has two notches which are not seen in the Munich or Thermopolis
specimens. The absence of a furcula in the Eichstatt specimen is near certainly
taphonomic, while the supposed sternum in the Munich specimen is a coracoid.
Ulnohumeral ratios vary between ~86% (Maxberg), 87% (Berlin), ~87% (Solnhofen),
88% (Eichstatt and ninth), 89% (Thermopolis), 90% (London), ~92% (Daiting),
~95% (11th) and ~96% (Munich). While this a large amount of variation, is is
continuous and also known in Sapeornis. All specimens probably had four
carpals (scapholunare, pisiform, semilunate carpal, distal carpal III), though these
are easily lost. In particular, the London specimen is seemingly missing proximal
carpals, the Eichstatt specimen lacks the pisiform on the right hand (the bone
labeled as an ulnare is probably distal carpal III), the left manus of the Munich
specimen lacks the scapholunare, only the semilunates are visible in the Thermopolis
specimen, ninth and Daiting specimens, and the Maxberg specimen seems to lack
preserved carpals. Contrary to Elzanowski (2001), manual ungual I is not short
compared to phalanx I-1 in the Solnhofen specimen. However, manual phalanx I-1
of the Solnhofen specimen is 1.39 times as long as phalanx II-1, compared to
1.36 times in the ninth specimen, 1.60 times in the Munich specimen, 1.32 times
in the Berlin specimen, 1.53 times in the Eichstatt specimen, 1.52 times in
the Thermopolis specimen and 1.25 times in the 11th specimen. Similar variation
is known in other coelurosaurs however (Sapeornis, Dromiceiomimus,
Gorgosaurus). Elzanowski (2001) noted manual phalanx I-1 is more robust
in the Solnhofen and Haarlem specimens (minimum height 8.4% and 8.2% of phalanx
length respectively compared to the Berlin (6%), Eichstatt (5.84%) and Munich
(5.75%) specimens, and this is also true compared to the ninth (6.56%), Thermopolis
(7.4%) and 11th (7.0%) specimens. Yet the latter three also close the gap in
ratios, making them seem less important for dividing specimens. Similarly, the
Thermopolis specimen (~8.1%) fills the gap between the Solnhofen (8.9%) and
11th (8.7%) on one hand, and the Berlin (6.2%), Eichstatt (6.9%) and Berlin
(6.2%) specimens on the other when it comes to robusticity of manual phalanx
II-2. The cross section of metacarpal III was said to distinguish the London
and Berlin specimens, but this has not been studied in further specimens. Metacarpal
III is more bowed with an intermetacarpal gap in the Solnhofen, Munich, Thermopolis
and ninth specimens compared to the Berlin, Eichstatt, Daiting and 11th specimens,
but this may be due to its orientation. Manual phalanges III-1 and III-2 are
sutured in the Solnhofen specimen (as in Jinfengopteryx and the Didactylornis
type), but not in the Berlin, Maxberg, Eichstatt, Munich, ninth and Thermopolis
specimens. The ratio between phalanges III-2 and III-1 varies between 66% (Berlin),
~69% (Maxberg), 71% (Eichstatt), 79% (ninth), 81% (Solnhofen), 83% (11th), 86%
(Munich) and 88% (Thermopolis). While this is quite a range, it is also gradational
and seen in some other taxa (Struthiomimus, Dromiceiomimus).
The Eichstatt, Solnhofen, Berlin and possibly the 11th specimens have an unexpanded
preacetabular processes, the London and probably the Munich and Thermopolis
specimens have ventrally expanded processes. Senter and Robins (2003) suggested
this could be due to ontogeny, but juvenile tyrannosaurids, Juravenator,
Gallimimus, therizinosauroids, Microvenator, Bambiraptor
and Scansoriopteryx all have expanded preacetabular processes, though
the subadult Yangchuanosaurus shangyouensis holotype lacks one while
the larger Y. magnus holotype has one. The preacetabular process varies
in length between specimens, from 54% of pubic peduncle plus acetabulum length
in the Solnhofen specimen to 89% (Eichstatt), ~111% (Munich), 120% (London),
~133% (11th) and 160% (Berlin). Elzanowski (2002) claimed the Berlin and Munich
specimens lack an m. cuppedicus fossa on their ilia, unlike the London specimen,
but such a fossa is clearly visible in the Berlin specimen (Ostrom, 1976- figure
20) while the Munich specimen seems to not preserve a lateral surface (Wellnhofer,
1993- plate 9). The Eichstatt specimen's ilium is only visible in medial view,
while the Solnhofen specimen is too poorly preserved to tell. The London specimen
differs from the Berlin, Eichstatt, Munich and Solnhofen specimens in having
a dorsally concave ilium. The pubic angle of Archaeopteryx has been quite
controversial, but is articulated at 3 degrees posterior to vertical in the
Munich specimen. The pubis is disarticulated in the London, Berlin, Eichstatt,
Solnhofen, Thermopolis and 11th specimens, so its angle cannot be used to distinguish
species. The London and Thermopolis specimens have posterior pubic boots partially
replaced by calcite (originally from cartilage?), while the Eichstatt, Munich,
Berlin and 11th specimens seem to have more ossified pubic boots, but this could
be ontogenetic. The London specimen differs from the Haarlem, Berlin and Eichstatt
specimens (Ostrom, 1976) in having a transversely expanded pubic boot. The length
of the pubic apron varies between 36% (Berlin), 39% (Solnhofen), 40% (Eichstatt),
~46% (London) and ~47% (Thermopolis), but may be overestimated in the last two
due to their incompletely ossified distal ends. The obturator foramen identified
by Petronievics in the London specimen is a pneumatic fossa (Christiansen and
Bonde, 2000), and whether it is present in other specimens is uncertain due
to a lack of posterior pubic exposure. The ischia of specimens differ a great
deal, but this has not been recently and explicitly described. The London specimen
has an obturator foramen (unique among maniraptoriforms), while a fossa is present
in the Eichstatt and Munich specimen, and the Thermopolis and 11th specimens
seemingly lack either. The proximoventral edge of the ischium is convex in the
London specimen, unlike the Berlin, Eichstatt, Munich, Thermopolis and 11th
specimens. The obturator process projects ventrally in the Munich and Thermopolis
specimens, a bit less in the Eichstatt and 11th specimens, and not at all in
the London specimen. The distodorsal process (at midlength) varies from a rounded
bump in the London specimen and low angularity in the Munich specimen to a large
triangular spine in the Solnhofen, 11th and especially Eichstatt specimens,
to a large flange with a posterior notch in the Thermopolis specimen. Similarly,
the proximodorsal process varies from a low peak (Berlin, Eichstatt, Thermopolis,
11th), to a slender point (Solnhofen), to a massive block-like process (Munich,
London). The London specimen's further differs in having a notch proximally,
making it rectangular. Ischial variation has not been examined much in other
taxa, though Microraptor varies in the angle of the distal obturator
process edge, obturator process expansion, shaft depth and ventral convexity
proximal to the obturator process. Mirischia famously varies in the presence
of an obturator process and notch. Tyrannosaurus varies in obturator
process size and orientation, as well as proximodorsal process size, shaft curvature
and distal expansion. The differences observed in Archaeopteryx specimens
may be due to intraspecific variation as well.
Hindlimb ratios are explainable by allometry (Bennett, 2008), as are the more
robust metatarsi of large specimens (Maxberg, Solnhofen) compared to smaller
ones (Eichstatt, Munich, Thermopolis, 11th) (compare to juvenile tyrannosaurids
vs. adults). Tibiotarsal and tarsometatarsal fusion is controversial in Archaeopteryx.
It seems absent in the Eichstatt and Munich specimens. Tibiotarsal and metatarsal
fusion are absent in the Thermopolis specimen, though the astragalus and calcaneum
could be fused. In the Berlin and London specimens, there is no evidence for
metatarsal fusion and at least the ascending process of the astragalus remains
unfused to the tibia. Though Ostrom (1976) claimed the distal tarsal in the
Maxberg specimen was "at least partially fused" to the metatarsus,
he said the same of the Eichstatt specimen which seems to be untrue. Metatarsals
II-IV of the Maxberg specimen are indistinguishable proximally and fused as
determined by x-rays. The Solnhofen specimen lacks tibiotarsal and tarsometatarsal
fusion, but metatarsals II-IV seem to be partially fused proximally. Fusion
could be ontogenetic, of course, as the Solnhofen and Maxberg are among the
largest specimens. The metatarsal II/IV ratio varies between 97% (Thermopolis),
~97% (Munich, Maxberg, 11th), 100% (Solnhofen), 104% (Eichstatt), and ~108%
(Berlin). Tyrannosaurus has a similar variation (91-103%), and that of
Dromiceiomimus is slightly less (90-97%). While Elzanowski (2001) emphasized
the equal lengths in the Solnhofen specimen, it is intermediate between the
other specimens and thus not an outlier. The proximally tapering second metatarsal
in the right foot of the Solnhofen specimen is probably due to distortion as
it is not present in the left foot. The phalanx II-2/II-1 ratio varies between
83% (Thermopolis), 85% (Berlin), 96-98% (11th), ~99% (Eichstatt), ~100% (Munich),
100% (London), 104% (Solnhofen) and ~105% (Maxberg), so would seem to distinguish
the Berlin+Thermopolis specimens, as opposed to the Solnhofen specimen (contra
Elzanowski, 2001). Ratios in Velociraptor are more variable (103-129%),
showing this could be intraspecific variation. Pedal digit IV has four phalanges
in the Solnhofen specimen (as in the Didactylornis type), but five phalanges
in the Berlin, Eichstatt, Munich, Thermopolis and 11th specimens. This leads
to the fourth digit being shorter in the Solnhofen specimen. Though the London
specimen has an unknown number of phalanges in digit IV, it is long as in specimens
with five phalanges. Pedal ungual IV is longer than phalanx IV-1 in the Solnhofen
specimen, but not the Berlin, Eichstatt, Munich and Thermopolis specimens. Pedal
ungual IV of the Solnhofen specimen is also distinct from the London, Berlin,
Eichstatt, Munich, Thermopolis and 11th specimens in being straight ventrally.
Variation in pedal ungual curvature has been reported in Deinonychus
too (Ostrom, 1969). Pedal flexor tubercles are large in the London (I, II, III
and IV) and Solnhofen (I and III but not IV) specimens, but small (though not
absent) in the Berlin, Eichstatt, Munich, Thermopolis and 11th specimens. If
Zhongornis is a juvenile Confuciusornis, it would provide an example
of young specimens having smaller pedal flexor tubercles than adults.
When the characters above are entered into a matrix with Shenzhouraptor
and Anchiornis as outgroups to establish polarity, the London+Munich
specimens branch off first due to preacetabular length (also in the 11th), obturator
area of ischium at least depressed (also in Eichstatt) and large proximodorsal
ischial process. Other specimens are united with Shenzhouraptor by the
low dorsal neural spines (not in the 11th specimen) and large distodorsal ischial
process. The Berlin, Eichstatt, Thermopolis and 11th specimens are united by
non-enlarged pedal flexor tubercles (also in Munich), while the Maxburg, Solnhofen
and Shenzhouraptor are united by metatarsal fusion. The Solnhofen, ninth
and Shenzhouraptor are united by a short manual phalanx I-1 (also in
the Berlin specimen and some Anchiornis) and pedal ungual IV longer than
IV-1 (also in some Anchiornis). The fact most of these characters are
homoplasious shows that there is little evidence to subdivide the genus into
species containing more than one individual, despite individual Archaeopteryx
specimens (particularily the London and Solnhofen specimens) having several
unique features. The choices are similar to that entertained for Microraptor
on this site, either several species each represented by a single
specimen, or one species which is variable in morphology. The latter
more conservative approach is taken here, especially considering the
fact no specimens are adult. Additional evidence is provided by
the maniraptoromorph matrix of Hartman et al. 2019 where a topology of
((7(11(2,5)))(12((1,10)(6,8)))) is recovered, incompatable with any
suggested distribution of taxa.
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Ornithodesmiformes Martyniuk, 2012
Definition- (Ornithodesmus cluniculus, Dromaeosaurus albertensis,
Troodon formosus <- Archaeopteryx lithographica) (Martyniuk,
2012)
Comments- This clade was named
by Martyniuk (2012) presumedly based on the historical priority of
Ornithodesmidae, and actually fulfills a useful role in the present
topology assuming Ornithodesmus
is an unenlagiid as recovered by Hartman et al.. However, it self
destructs in the two alternate paravian topologies one step longer
where Troodon is closer to Archaeopteryx than Dromaeosaurus. This and the uncertain position of Ornithodesmus makes such a clade unwise to use.
Reference- Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other
Winged Dinosaurs. Pan Aves. 189 pp.
unnamed ornithodesmiform (Jensen, 1981)
Late Kimmeridgian, Late Jurassic
Brushy Basin Member of the Morrison Formation, Colorado, US
Material- (BYUVP 2023) femur
Comments- This was first referred to Archaeopteryx by
Jensen (1981), then to Theropoda indet. by Molnar (1985) and
Maniraptora indet. by Jensen and Padian (1989). Hartman et al.
(2019) included it in a phylogenetic analysis for the first time and
recovered it as a deinonychosaur in the Unenlagiidae+Dromaeosauridae
clade excluded from Halszkaraptorinae, Sinovenatorinae, Gobivenator+ other troodontids and Eudromaeosauria. It thus may belong to
"Paleopteryx" or Hesperornithoides, but can only be compared to the latter in that both lack fourth trochanters.
References- Jensen, 1981. [A new oldest bird?] Anima (Tokyo). 1981, 33-39.
Jensen, 1981. Another look at Archaeopteryx as the worlds oldest bird.
Encyclia, The Journal of the Utah Academy of Sciences, Arts, and Letters. 58,
109-128.
Molnar, 1985. Alternatives to Archaeopteryx; a survey of proposed early
or ancestral birds. In Hecht, Ostrom, Viohl and Wellnhofer (eds.). The Beginnings
of Birds. Freunde des Jura-Museums Eichstatt. 209-217.
Jensen and Padian, 1989. Small pterosaurs and dinosaurs from the
Uncomphagre fauna (Brushy Basin Member, Morrison Formation:
?Tithonian), Late Jurassic, western Colorado. Journal of Paleontology.
63(3), 364-373.
Padian, 1998. Pterosaurians and ?avians from the Morrison Formation (Upper Jurassic, western U.S.). Modern Geology. 23, 57-68.
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 ornithodesmiform (Rodriguez de la Rosa and Cevallos-Ferriz, 1998)
Early Maastrichtian, Late Cretaceous
Ca�on del Tule Formation, Mexico
Material- (IGM-7715) distal pedal phalanx II-2
Comments- Although described as being from the Cerro del
Pueblo Formation, Aguillon Martinez (2010) found this and other
material from the El Pelillal locality belong to the later Ca�on del
Tule Formation.
This was tentatively referred to Dromaeosauridae by Rodriguez
de la Rosa and Cevallos-Ferriz (1998) based on the dorsally placed collateral
ligament pit. However, this character is also present in basal troodontids (e.g.
IGM 100/44, Sinornithoides, Borogovia), Neuquenraptor and
Rahonavis. It is a different taxon than IGM-7710 and IGM-7712, both of
which have centrally placed collateral ligement pits.
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.
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.
Migmanychion Wang, Cau, Wang, Yu, Wu, Wang and Liu, 2023 online
M. laiyang Wang, Cau, Wang, Yu, Wu, Wang and Liu, 2023 online
Early Aptian, Early Cretaceous
Pigeon Hill, Longjiang Formation, Inner Mongolia, China
Holotype- (LY 2022JZ3001)
dorsal rib fragments, distal radius, distal ulna, distal carpal I,
metacarpal I (23.1 mm), phalanx I-1 (41.8 mm), manual ungual I (28.6
mm), metacarpal II (52.8 mm), phalanx II-1 (31.6 mm), phalanx II-2
(43.7 mm), manual ungual II (35.8 mm), metacarpal III (52.7 mm),
phalanx III-1 (15.1 mm), phalanx III-2 (15.0 mm), phalanx III-3 (24.9
mm), manual ungual III (18.5 mm)
Diagnosis- (after Wang et al.,
2023) differs from other taxa except Fukuivenator in- discoidal distal
carpal 1 capping exclusively metacarpal I (unknown in Fukuivenator);
sharp proximolateral flange on the ventral surface of metacarpal I;
manual phalanx II-1 with lateroventral ridge (also in some paravians);
manual ungual II dorsoventrally shallower but proximodistally longer
than manual ungual I; manual unguals I and III having a prominent
proximodorsal lip and the dorsal margin arched well above the level of
the articular facet when the latter is oriented vertically.
differs from Fukuivenator in- metacarpal I more than four times longer
than distally wide and narrower than metacarpal II; metacarpal III
being no more than 40% of metacarpal II width at mid-shaft (55% in
Fukuivenator); flexor tubercle manual ungual I distally displaced
relative to proximal articular facet; manual ungual II with dorsal
margin arched well above the level of the articular facet when the
latter is oriented vertically; manual ungual III half the size of
manual ungual II (in Fukuivenator ungual III is >80% the size of
ungual II).
Comments- This was discovered in 2022. Wang et al. (2023) used Cau's meagmatrix to recover Migmanychion sister to Fukuivenator, which is in turn sister to Pennaraptora. Two steps were needed to force it to be sister to Pennaraptora (which left Fukuivenator more basal), and three steps were needed to make it a therizinosaur (where it remains sister to Fukuivenator),
oviraptorosaur or within Eumaniraptora (as either an archaeopterygid or
dromaeosaurid). In Hartman et al.'s maniraptoromorph matrix, Migmanychion
resolves as a paravian either sister to other dromaeosaurids or
anywhere in Halszkaraptorinae. One step can make it an
archaeopterygid or unenlagiine, while two steps can make it a
troodontid, basalmost deinonychosaur, basalmost avialan, basalmost
oviraptorosaur, sister to Pennaraptora, sister to
Therizinosauria+Alvarezsauroidea, basalmost maniraptoran, basalmost
ornithomimosaur, sister to Maniraptoriformes or sister to
Ornitholestiinae (which includes Fukuivenator,
although making it sister to that genus takes three steps). Thus
its fragmentary preservation and unspecialized morphology make Migmanychion
plausibly fit many places between ornitholestiids and Euavialae,
although generally not within the known content of each subclade.
Notably the basal deinonychosaur position preferred here makes referral
of additional Pigeon Hill specimens LY 2022JZ3004 (pelvic area referred
to Paraves by Wang et al.) and LY 2022JZ3005 (subarctometatarsal
metatarsi with distally placed halluces referred to Coelurosauria by
Wang et al.) plausible.
Reference- Wang, Cau, Wang, Yu,
Wu, Wang and Liu, 2023 online. A new theropod dinosaur from the Lower
Cretaceous Longjiang Formation of Inner Mongolia (China). Cretaceous
Research. Journal Pre-proof, 10565. DOI: 10.1016/j.cretres.2023.105605.
Ornithodesmidae Hooley, 1913
Definition- (Ornithodesmus cluniculus <- Archaeopteryx lithographica,
Passer domesticus, Paronychodon lacustris, Pterodactylus antiquus)
(Martyniuk, 2012)
References- Hooley, 1913. On the skeleton of Ornithodesmus latidens;
an ornithosaur from the Wealden Shales of Athersfield (Isle of Wight). Quarterly
Journal of the Geological Society. 69, 372-421.
Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other Winged Dinosaurs.
Pan Aves. 189 pp.
Ornithodesmus Seeley, 1887
O. cluniculus Seeley, 1887
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (NHMUK R187) (sacrum- 96 mm) first sacral vertebra (17 mm),
second sacral vertebra (16 mm), third sacral vertebra (18 mm), fourth sacral
vertebra (15 mm), fifth sacral vertebra (16 mm), sixth sacral vertebra (13 mm)
Comments- The holotype was purchased from the Fox collection in 1884 (Lydekker, 1888). Seeley (1887) originally described Ornithodesmus as a bird,
but Hulke (in Anonymous, 1887) soon suggested it was pterosaurian. Seeley later
(1901) referred pterosaur skeleton NHMUK R176 to Ornithodesmus as a new species, O. latidens.
For over a century, the well known pterosaur Ornithodesmus latidens was
used as the standard example of the genus. This ended in 1993 when Howse and
Milner reidentified the Ornithodesmus cluniculus holotype as a theropod
(the pterosaur was later renamed Istiodactylus latidens by Howse et al.,
2001). Specifically they believed it to be a troodontid, based largely on comparison
to NHMUK R4463 (another supposed troodontid sacrum). Makovicky (1995) and Norell
and Makovicky (1997) identified NHMUK R4463 as Saurornitholestes, and
the latter reference noted Ornithodesmus resembles Dromaeosauridae in
possessing a well developed dorsal ridge formed by zygapophyses. Makovicky stated
the specimen was "probably neither a troodontid nor a dromaeosaurid"
while Naish et al. (2001) stated some characters argue against a dromaeosaurid
identity, such as transverse processes which are not dorsoventrally flattened.
That character is meaningless outside of a phylogenetic context however. As
an alternative, Naish et al. note resemblence to Coelophysis rhodesiensis
and Carnotaurus in the presence of six sacrals, neural spine lamina and
neural platform. Yet coelophysoids never have more than five sacrals, and dromaeosaurids
like Velociraptor have all three listed characters. In addition, no coelophysoids
or ceratosaurs have sacral pleurocoels or flattened ventral sacral surfaces
with a median groove, unlike Ornithodesmus
and dromaeosaurids. It was first included in the maniraptoromorph
analysis of Hartman et al. (2019) who recovered it in Unenlagiinae, but
after the addition of more data it can also fall out in
Halszkaraptorinae or as a basal dromaeosaurid with an equal number of
steps.
References- Seeley, 1887. On a sacrum, apparently indicating a new type
of bird, Ornithodesmus cluniculus, Seeley, from the Wealden of Brook.
Quarterly Journal of the Geological Society of London. 42, 206-211.
Anonymous, 1887. Discussion (on Ornithodesmus and Patricosaurus).
Quarterly Journal of the Geological Society of London. 43, 219-220.
Lydekker, 1888. Catalogue of the Fossil Reptilia and Amphibia
in the British Museum (Natural History), Cromwell Road, S.W., Part 1. Containing
the Orders Ornithosauria, Crocodilia, Dinosauria, Squamata, Rhynchocephalia,
and Proterosauria. British Museum of Natural History. 309 pp.
Seeley, 1901. Dragons of the Air. Methuen & Co..
239 pp.
Howse and Milner, 1993. Ornithodesmus - a maniraptoran theropod dinosaur
from the Lower Cretaceous of the Isle of Wight, England. Palaeontology. 36,
425-437.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). Masters Thesis, University of Copenhagen. 311 pp.
Norell and Makovicky, 1997. Important features of the dromaeosaur skeleton:
Information from a new specimen. American Museum Novitates. 3215, 28 pp.
Howse, Milner and Martill, 2001. Pterosaurs. In Martill and Naish (eds.). Dinosaurs
of the Isle of Wight. The Palaeontological Association. 324-335.
Naish, Hutt and Martill, 2001. Saurischia dinosaurs: Theropods. In Martill and
Naish (eds). Dinosaurs of the Isle of Wight. The Palaeontological Association. 242-309.
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
Pyroraptor Allain and Taquet,
2000
P. olympius Allain and Taquet, 2000
Late Campanian-Early Maastrictian, Late Cretaceous
La Boucharde, Bouches-du-Rhone, France
Holotype- (MNHN BO001) pedal ungual II (66 mm)
Paratypes- (MNHN BO002) pedal phalanx II-2 (23 mm)
(MNHN BO003) metatarsal II (118.7 mm)
(MNHN BO004) pedal ungual II
(MNHN BO005) ulna (112.8 mm)
(MNHN BO014) tooth
(MNHN BO015) tooth
Referred- (MNHN BO006-BO010) pedal phalanges including II-2 (~28 mm)
(Allain and Taquet, 2000)
(MNHN BO011) manual phalanx (Allain and Taquet, 2000)
(MNHN BO012) metatarsal I (Allain and Taquet, 2000)
(MNHN BO013) incomplete radius (Allain and Taquet, 2000)
(MNHN BO016) proximal caudal vertebra (24 mm) (Allain and Taquet, 2000)
(MNHN BO017) dorsal vertebra (Allain and Taquet, 2000)
(MNHN coll.) distal metatarsal III (Turner, Makovicky and Norell, 2012)
Late Campanian-Early Maastrichtian, Late Cretaceous
Montrebei, Tremp Formation, Spain
(DPM-MON-T1) tooth (6.3x3.3x2.4 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Late Campanian, Late Cretaceous
La�o, Sedano Formation, Spain
(MCNA 14623) dentary fragment, tooth (8x5.4x2.4 mm) (Torices, Currie, Canudo
and Pereda-Suberbiola, 2015)
(MCNA 14624) dentary fragment, tooth (6.2x4.3x1.9 mm) (Torices, Currie, Canudo
and Pereda-Suberbiola, 2015)
(MCNA 14625) tooth (3.9x3.1x1.4 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
(MCNA 14626) tooth (3.8x2.5x1.4 mm) (Torices, Currie, Canudo and Pereda-Suberbiola,
2015)
Diagnosis- (after Allain and Taquet, 2000) ventrally concave metatarsal
II.
Other diagnoses- Turner et al. (2012) noted the supposed deep m. brachialis
fossa on the ulna is due to crushing. The other characters listed in Allain
and Taquet's (2000) diagnosis are common in dromaeosaurids- teeth distally serrated
and with restricted mesial serrations; ulna subequal in lenth to metatarsus;
asymmetrically ginglymoid metatarsal II; strongly curved pedal ungual II.
Comments- The hypodigm was discovered from 1993-1998. At least two individuals are present in the type material,
based on a larger second pedal phalanx II-2 (Turner et al., 2012). Turner et
al. also noted a metatarsal III is present in the collection, and Allain and
Taquet misidentified metatarsal I as a distal metacarpal I. DPM-MON-T1 was first
described as Dromaeosauridae indet. 3 by Torices (2002). Torices et al. (2015)
identified several teeth and dentary fragments based on morphometric analysis,
plus spatiotemporal similarity.
Pyroraptor may be a junior synonym of the contemporary dromaeosaurid
Variraptor, but this cannot be established until the dorsal vertebra
MNHN BO017 or additional specimens are described. Alternatively, the paratypes
and referred specimens of Variraptor may belong to Pyroraptor
instead. They are not comparable presently.
Senter et al. (2004) included Pyroraptor in their phylogenetic analysis
of coelurosaurs and found it to be a dromaeosaurid excluded from Microraptoria
(based on the stout pedal phalanx II-2) and Dromaeosaurus+Utahraptor
(based on the high DSDI). Turner et al. found it to fall within Dromaeosauridae,
but outside Eudromaeosauria (including Bambiraptor
in their trees). Hartman et al. (2019) recovered it as an
unenlagiine, but with the addition of more data it can equally
parsimoniously be a microraptorian dromaeosaurid.
References- Allain and Taquet, 2000. A new genus of Dromaeosauridae (Dinosauria,
Theropoda) from the Upper Cretaceous of France. Journal of Vertebrate Paleontology.
20(2), 404-407.
Torices, 2002. Los dinosaurios ter�podos del Cret�cico Superior
de la Cuenca de Tremp (Pirineos Sur-Centrales, Lleida). Coloquios de Paleontolog�a.
53, 139-146.
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae.
Bulletin of Gunma Museum of Natural History. 8, 1-20.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 206 pp.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
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
Yaverlandia Galton, 1971
Y. bitholus Galton, 1971
Barremian, Early Cretaceous
Wessex Formation, England
Holotype- (MIWG 1530) frontals, orbitosphenoid
fragment
Diagnosis- (after Galton, 1971) frontals thickened with two domes; frontals with pitted surface.
(proposed) fused frontals.
Other diagnoses- Galton (1971)
also included frontal-orbit contact and the unrestricted supratemporal
fenestrae as diagnostic within Pachycephalosauria, but these are
typical of theropods.
Comments- This specimen was found in 1923 and originally considered to perhaps be referrable
to Vectisaurus (Watson, 1930; Swinton, 1936), though Watson also noted
resemblence to Stegoceras (his Troodon). Galton (1971) named the taxon Yaverlandia
bitholus, and assigned it to the Pachycephalosauridae, which was followed
by most authors until recently. When entered into cladistic analyses of Pachycephalosauria,
Yaverlandia was resolved at the base of the clade (Williamson and Carr,
2002) or as a basal pachycephalosaurid (Sereno, 2000). However, Hopson (1979)
and Giffin (1989) doubted a pachycephalosaurian identity, based on the structure
of the endocranium. Sullivan (2000, 2003) noted several characters incongruent
with a pachycephalosaurian identity- frontals with broad orbital contact; parietals
excluded from domes; supratemporal fenestrae contact frontals. Naish (2004)
determined Yaverlandia
is not a pachycephalosaur, though its identity remained elusive in his
abstract. Naish (2006) noted the taxon shares a few characters with
pachycephalosaurids but lacks pachycephalosaurian and marginocephalian
characters, suggesting either significant reversals or convergence. He
classified the genus as a maniraptoriform based on ventrally extending,
laterally concave ridges that form the lateral margins of the cerebral
concavity and ventrally concave dorsal orbital margins, and furthermore
as a troodontid. Yet most of the troodontid-like characters are also present in unenlagiines and Halszkaraptor, neither considered by Naish, so Yaverlandia is here assigned to Ornithodesmiformes instead.
References- Watson, 1930. Proceedings of the Isle of Wight Natural History
Society. 2, 60.
Swinton, 1936. The dinosaurs of the Isle of Wight. Proceedings of the Geologists'
Association. 47, 204-220.
Galton, 1971. A primitive dome-headed dinosaur (Ornithischia; Pachycephalosauridae)
from the Lower Cretaceous of England and the function of the dome of pachycephalosaurids.
Journal of Paleontology. 45(1), 40-47.
Hopson, 1979. Paleoneurology. In Gans, Northcutt and Ulinski (eds.). Biology
of the Reptilia (Neurology A). Academic Press. 9, 39-146.
Wall and Galton, 1979. Notes on pachycephalosaurid dinosaurs (Reptilia:
Ornithischia) from North America, with comments on their status as
ornithopods. Canadian Journal of Earth Sciences. 16(6), 1176-1186.
Giffin, 1989. Pachycephalosaur paleoneurology (Archosauria: Ornithischia). Journal
of Vertebrate Paleontology. 9(1), 67-77.
Sereno, 2000. The fossil record, systematics and evolution of pachycephalosaurs
and ceratopsians from Asia. In Benton, Shishkin, Unwin and Kurochkin (eds.).
The Age of Dinosaurs in Russia and Mongolia. Cambridge University Press. 480-516.
Sullivan, 2000. Prenocephale edmontonensis (Brown and Schlaikjer) new
comb. and P. brevis (Lambe) new comb. (Dinosauria: Ornithischia: Pachycephalosauria)
from the Upper Cretaceous of North America. New Mexico Museum of Natural History
and Science Bulletin. 17, 177-190.
Naish and Martill, 2001. Boneheaded and horned dinosaurs. In Martill and Naish
(eds.). Dinosaurs of the Isle of Wight. The Palaeontological Association, London.
133-146.
Williamson and Carr, 2002. A new genus of derived pachycephalosaurian from western
North America. Journal of Vertebrate Paleontology. 22(4), 779-801.
Sullivan, 2003. Revision of the dinosaur Stegoceras Lambe (Ornithischia,
Pachycephalosauridae). Journal of Vertebrate Paleontology. 23(1), 181-207.
Naish, 2004. So... what is Yaverlandia? SVPCA 2004 Abstracts. [pp]
Naish, 2006. The osteology and affinities of Eotyrannus lengi and Lower
Cretaceous theropod dinosaurs from England. PhD thesis, University of Portsmouth.
353 pp.
Sullivan, 2006. A taxonomic review of the Pachycephalosauridae (Dinosauria:
Ornithischia). New Mexico Museum of Natural History and Science Bulletin. 35,
347-365.
Naish, 2011. Theropod dinosaurs. In Batten (ed.). English Wealden Fossils. The
Palaeontological Association. 526-559.
Deinonychosauria sensu Sereno, 1997
Definition- (Troodon formosus + Dromaeosaurus albertensis) (modified)
= Deinonychosauria sensu Sereno, 1998
Definition- (Troodon formosus + Velociraptor mongoliensis) (modified)
undescribed deinonychosaur (Fortner, 2015)
Early Maastrichtian, Late Cretaceous
Aguja Formation, Texas, US
Material- partial postcranial skeleton
Comments- This was said to be "compatible with identification as
either Troodontidae or Dromaeosauridae." It may be Richardoestesia
or Paronychodon, if not a eudromaeosaur or troodontine.
Reference- Fortner, 2015. A small theropod dinosaur from the Aguja Formation
(Upper Cretaceous), Big Bend National Park, Texas. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 126.
Dromaeosauridae
Troodontidae Gilmore, 1924
Definition- (Troodon formosus <- Ornithomimus
velox, Mononykus olecranus, Therizinosaurus cheloniformes, Oviraptor
philoceratops, Archaeopteryx lithographica, Unenlagia comahuensis,
Velociraptor mongoliensis, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019)
Other definitions- (Troodon formosus, Saurornithoides mongoliensis,
Borogovia gracilicrus, Sinornithoides youngi <- Ornithomimus
velox, Oviraptor philoceratops) (Varricchio, 1997)
(Troodon formosus <- Velociraptor mongoliensis) (Makovicky
and Norell, 2004; modified from Sereno, 1998)
(Troodon formosus <- Ornithomimus velox, Mononykus
olecranus, Therizinosaurus cheloniformes, Oviraptor philoceratops, Velociraptor
mongoliensis, Passer domesticus) (modified from Senter, Barsbold, Britt and Burnham, 2004)
(Troodon formosus <- Ornithomimus edmontonicus, Velociraptor
mongoliensis, Passer domesticus) (Sereno, online 2005)
(Troodon formosus <- Deinonychus antirrhopus, Passer domesticus)
(Hu, Hou, Zhang and Xu, 2009)
(Troodon formosus <- Velociraptor mongoliensis, Passer domesticus)
(Turner, Makovicky and Norell, 2012)
(Troodon formosus <- Dromaeosaurus albertensis, Passer domesticus)
(Godefroit, Cau, Hu, Escuillie, Wu and Dyke, 2013)
= Saurornithoididae Barsbold, 1974
= Troodontidae sensu Sereno, 1998
Definition- (Troodon formosus <- Velociraptor mongoliensis)
(modified)
= Troodontidae sensu Sereno, online 2005
Definition- (Troodon formosus <- Ornithomimus edmontonicus,
Velociraptor mongoliensis, Passer domesticus)
= Troodontoidea Gilmore, 1924 vide Livezey and Zusi, 2007
= Troodontidae sensu Hu, Hou, Zhang and Xu, 2009
Definition- (Troodon formosus <- Deinonychus antirrhopus, Passer
domesticus)
= Troodontia Alifanov, 2012
= Troodontidae sensu Turner, Makovicky and Norell, 2012
Definition- (Troodon formosus <- Velociraptor mongoliensis,
Passer domesticus)
= Troodontidae sensu Godefroit, Cau, Hu, Escuillie, Wu and Dyke, 2013
Definition- (Troodon formosus <- Dromaeosaurus albertensis,
Passer domesticus)
Comments- The first troodontids discovered were assigned to the Megalosauridae (Saurornithoides in Osborn, 1924) and Coeluridae (Stenonychosaurus
in Sternberg 1932), both waste-basket families at the time.
Troodontidae was proposed for pachycephalosaurs by Gilmore (1924) who
believed Troodon to be a Stegoceras tooth, until Sternberg (1945) named Pachycephalosauridae and moved Troodon back to Theropoda. Barsbold's (1974) Saurornithoididae was used for a time until Currie (1987) synonymized Stenonychosaurus with Troodon, with Troodontidae being used ever since.
Two main alternatives have been proposed for the placement of
troodontids within Theropoda, with Ostrom (1969), Barsbold (1976) and
Gauthier (1984) combining them with dromaeosaurids in Deinonychosauria
based largely on the shared hyperextendable and enlarged pedal ungual
II, while Thulborn (1984), Currie (1985), Bakker (1986) and Holtz
(1992) placed them closer to ornithomimids in Bullatosauria based
largely on the shared inflated parasphenoid cultriform process.
While this was controversial throughout the 1990s, the description of
basal bird-like Sinovenator
in 2002 cemented maniraptoran and paravian troodontids, with Senter et
al. (2004) probably the last quantitative analysis to support
Bullatosauria (and only because Sinovenator as drawn away from other troodontids into Dromaeosauridae).
Troodontidae defined- Sereno's newest (online 2005) definition differs from his earlier
(1998) one which only had Velociraptor as an external specifier, and
Senter et al.'s (2004), which had Mononykus, Therizinosaurus and
Oviraptor as additional external specifiers. Keeping Oviraptor
seems useful, in the case Osmolska and Barsbold (1990), Russell and Dong (1994)
or Norell et al. (2001) are correct. I'm not aware of any topology placing Shuvuuia
or Therizinosaurus closer to Troodon
than the other taxa, but they both seem relatively plausible as
troodontid sister taxa. Hartman et al. (2019) found topologies pairing
troodontids with archaeopterygids were only a single step longer than
grouping troodontids with dromaeosaurids, so added Archaeopteryx as an external specifier so that Archaeopterygidae could not be a senior synonym of Troodontidae.
Troodontoidea- Livezey and Zusi (2007) in commentary for their large avian analysis list cite "Troodontoidea (Troodon and Saurornithoides)",
which while possibly a typo (as no other troodontid OTUs were used) is
also among many higher-level taxa stated as new that are used in this
work (e.g. Palaeoaves, Rahoinaviformes, Rahonavidae, Apsaraviformes,
Apsaravidae...). Bhullar et al. (2012) use Troodontoidea in their
cladogram without comment.
Microtroodontidae- Wiemann et
al. (2018) use the term microtroodontid without elaboration, first
stating "Egg colour pigments are preserved in eggshells from the
oviraptorid Heyuannia huangi,
Mongolian microtroodontids..." and indicating in Supplementary Table 2
that the two specimens considered microtroodontids are IGM 100/1323
(later made the holotype of Almas)
and MAE 14-20, which has never been mentioned outside of Wiemann's
eggshell dataset but is stated to also be from the Djadochkta
Formation. Shown as a clade exclusive of other troodontids
(a Two Medicine Troodon egg, two Djadokhta eggs and PFMM-0014003517 from the Chichengshan
Formation and also not mentioned outside Wiemann's datasets), this is
said to be "a composite topology (supertree) from previously published
phylogenies" 8, 31 and 32, but none of the three references include MAE
14-20. The term "microtroodontid" is later used in Norell et al.
(2020; supplementary info) and Jiang et al. (2023), but with no further
elaboration. According to ICZN Article 11.7.11 a
family-group name must be "formed from the stem of an available generic
name" which "must be a name then used as valid in the new family-group
taxon." As Wiemann et al. did not intend for a genus 'Microtroodon' to
be a microtroodontid, it is invalid. It also fails ICZN Article
13.1.1 ("be accompanied by a description or definition that states in
words characters that are purported to differentiate the taxon"), 16.1
("Every new name published after 1999, including new replacement names
(nomina nova), must be explicitly indicated as intentionally new") and
16.2 (" a new family-group name published after 1999 must be
accompanied by citation of the name of the type genus (i.e. the name
from which the family-group name is formed)").
Macrotroodontidae- Wiemann et
al. (2018) also use the term macrotroodontid without explanation,
stating "Both egg colour pigments were detected in eggshells from H. huangi,
the Mongolian microtroodontid (IGM 100/1323) and macrotroodontid (AMNH
FARB 6631)", which seems to be what they call "Large Troodontid" in
Supplementary Table 2 and includes Two Medicine Troodon egg YPM PU 23259, two Djadokhta eggs (AMNH 6631 and IGM 100/1003; perhaps Saurornithoides, Gobivenator and/or Byronosaurus) and PFMM-0014003517 from the Chichengshan
Formation. Unlike 'microtroodontid', the term was not used in
subsequent papers, but it is invalid due to the same ICZN Articles
(11.7.11, 13.1.1, 16.1 and 16.2).
No longer troodontids-
Note that in cases where specimens consist only of teeth, these are
only referred to Troodontidae here if they match those morphologies
found with known troodontid skulls (roots constricted and carinae
either unserrated, serrated only distally, or with large serrations
present on both carinae). Teeth with both carinae finely serrated
are known in Hesperornithoides,
but this early taxon can easily move to other positions within Paraves
(Dromaeosauridae, Archaeopterygidae, basal Avialae). Thus many
teeth with finely serrated mesial and distal carinae and constricted
bases are reassigned to Paraves here, often resembling some Microraptor
teeth most, but probably members of poorly known paravian clades in
most cases considering their temporal and geographic position.
That being said, some posterior teeth of Microraptor and Caihong
would also fall under the serrated troodontid morphospace, so some
teeth listed as troodontids below may end up being e.g. dromaeosaurids
or archaeopterygids.
Currie et al. (1990) said that troodontid teeth
reported from the Cedar Mountain Formation by Nelson and Crooks (1987) were
more likely velociraptorines because of the serrations are small (12/mm) and
elongate.
Supposed troodontid teeth from the Mussentuchit Member of the Cedar
Mountain Formation first noted by Parrish and Eaton (1991) and
described by Frederickson et al. (2018) are here identified as
microraptorian instead.
Bertini and
Franco-Rosas (2001) state "Troodontidae teeth were recognized" among
over 200 specimens from the Late Cretaceous Adamantina and Marilia
Formations of the Bauru Group in Brazil, probably originating with
Bertini's (1993) and
Franco's (1999) theses. In publications which describe supposedly
troodontid-like teeth from these formations, they are not
distinguishable from e.g. noasaurids which are more geographically
plausible. Thus they are referred to Averostra here.
Eaton (1999) reported troodontid teeth from the Iron Springs Formation
of Utah, but these were not present in the final report of Eaton et al.
(2014) and may have been based on dromaeosaurid tooth UMNH VP 24114 or
Theropod indet. tooth UMNH VP 24116.
Kirkland et al.
(1998) listed Troodontidae indet. from the basal Straight Cliffs
Formation and the Straight Cliffs Formation, while Eaton et al. (1999)
lists Troodontidae indet. from the Smoky Hollow Member (but not the
John Henry Member) of the Straight Cliffs Formation. However, Parrish
(1999) doesn't record any troodontid specimens from either member in
his study, suggesting they were misidentified.
Although Metcalf and Walker (1994) identified a tooth from the Bathonian Chipping
Norton Formation of England as a possible troodontid, it is more likely a dromaeosaurid.
Nessov (1995) cited a troodontid astragalocalcaneum from the Bostobe Formation
of Kazakhstan, but Averianov and Sues (2007) noted many other maniraptorans
have fused astragalocalcanea as well. Kordokova et al. (1996) cited troodontids
from the Zhirkindek and Bostobe Formations of Kazakhstan, but these were not
mentioned in their later (2001) paper. Averianov (2007) could not
confirm the presence of troodontids from either formation, but described a frontal as Troodontidae indet. in 2016..
Nessov (1995) identified CCMGE 484/12457 as a troodontid frontal, which he named
Saurornithoides isfarensis. Though they could not locate it, Averianov
and Sues (2007) reidentified the specimen as a hadrosaurid prefrontal.
Dong (1997) described tooth IVPP V11122-2 from the Zhonggou Formation
of China as Troodontidae, but it is here placed in Deinonychosauria.
Csiki and Grigorescu (1998) described five teeth from the Hateg
Formation of Romania (FGGUB R.1318, R.1319, R.1320 and MAFI v.12685a
and b) as "troodontid-like", more precisely "probably more closely
related to troodontids (?and "paronychodons") than to other small
theropods." However these more closely resemble the then-unknown Microraptor and are here assigned to Deinonychosauria.
Codrea et al. (2002) described teeth as troodontid-like from the
Sinpetru Beds of Romania (IRSNB coll.), but these are referred here to Averostra.
Debeljak et al. (2002) described a tooth from the Late Cretaceous of
Slovenia (ACKK-D-8/088) as ?troodontid, but it is here referred to
Deinonychosauria.
Codrea et al. (2012) described tooth UBB NgTh1 from the Rusca Mountain
Basin of Romania as being troodontid-like, but it is here placed as
?Averostra.
Ford (online 2015) placed tooth B of Huene (1934; CA coll.) in
Troodontidae without rationale, but it is a narrow D-shape unlike
members of that clade and is placed as Averostra indet. here
following Soto and Cambiaso (2006).
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Ford, online 2015. http://www.paleofile.com/Dinosaurs/Theropods/Troodonincertae.asp
Averianov, 2016 (online 2015). Frontal bones of non-avian theropod
dinosaurs from the Upper Cretaceous (Santonian-?Campanian) Bostobe
Formation of the northeastern Aral Sea region, Kazakhstan. Canadian
Journal of Earth Sciences. 53(2). 168-175.
Frederickson, Engel and Cifelli, 2018. Niche partitioning in pheropod
dinosaurs: Diet and habitat preference in predators from the uppermost
Cedar Mountain Formation (Utah, U.S.A.). Scientific Reports. 8:17872.
Wiemann, Yang and Norell, 2018. Dinosaur egg colour had a single evolutionary origin. Nature. 563, 555-558.
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
Norell, Wiemann, Fabbri, Yu, Marsicano, Moore-Nall, Varricchio, Pol and
Zelenitsky, 2020. The first dinosaur egg was soft. Nature. 583, 406-410.
Jiang, He, Elsler, Wang, Keating, Song, Kearns and Benton, 2023.
Extended embryo retention and viviparity in the first amniotes. Nature
Ecology & Evolution. DOI: 10.1038/s41559-023-02074-0
"Paronychodontidae"
Diagnosis- basoapical fluting on lateral teeth.
Comments- This family has yet to be officially named, but appears in
quotes in Ruiz-Omenaca et al. (1998), Pereda Suberbiola (1999) and Canudo and Ruiz-Omenaca (2003).
Reference- Ruiz-Ome�aca, Canudo, Cuenca-Besc�s and Amo,
1998. Theropod teeth from the Lower Cretaceous of Galve (Teruel, Spain). Third
European Workshop on Vertebrate Paleontology. 62-63.
Pereda Suberbiola, 1999. Las faunas Finicretacicas de dinosaurios Ibericos. Zubia. 17, 259-279.
Canudo and Ruiz-Omenaca, 2003. Los restos directos de dinosaurios terop�dos
(excluyendo Aves) en Espa�a. Ciencias de la Tierra. 26, 347-373.
unnamed 'paronychodontid' (Zinke and Rauhut, 1994)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Material- (IPFUB GUI D 1) dentary fragment, three teeth (Zinke and Rauhut,
1994)
(IPFUB GUI D 2) maxillary tooth (1.65 mm) (Zinke and Rauhut, 1994)
(IPFUB GUI D 3) tooth (Zinke and Rauhut, 1994)
(IPFUB GUI D 4-27) twenty-seven teeth (~1.67 mm) (Zinke, 1998)
Comments- IPFUB GUI D 1-3 teeth are recurved with one to two labial ridges
and one to four lingual ridges. They have distal serrations which vary in size
(6.5/mm in a tooth from GUI D 1; 14/mm in GUI D 2; 31/mm in GUI D 3), though
they increase in size basally. GUI D 2 may have a slight basal constriction,
and GUI D 3 is not flattened lingually. The mesial carina is unserrated (GUI
D 2, GUI D 3), though GUI D 1 has small pits apically that may be worn serrations.
Zinke notes all specimens have serrated distal carinae (6-13/mm) and some have
serrated mesial carinae (~18/mm). They were identified by Zinke and Rauhut as
cf. Paronychodon sp..
References-
Zinke and Rauhut, 1994. Small theropods (Dinosauria, Saurischia) from
the Upper Jurassic and Lower Cretaceous of the Iberian peninsula.
Berliner geowissenschaftliche Abhandlungen, E. 13, 163-177.
Zinke, 1998. Small theropod teeth from the Upper Jurassic coal mine of Guimarota
(Portugal). Palaontologische Zeischrift. 72(1/2), 179-189.
unnamed 'paronychodontid' (Ruiz-Ome�aca, Canudo, Cuenca-Besc�s
and Amo, 1998)
Late Hauterivian-Early Barremian, Early Cretaceous
Castellar Formation, Spain
Material- (MPZ 98-11) tooth (5.32 mm)
Comments- Though this tooth has two ridges on both lingual and labial
sides and no mesial serrations, it also has distal serrations (6/mm), so is
here excluded from Paronychodon.
References- Ruiz-Ome�aca, Canudo, Cuenca-Besc�s and Amo,
1998. Theropod teeth from the Lower Cretaceous of Galve (Teruel, Spain). Third
European Workshop on Vertebrate Paleontology. 62-63.
Ruiz-Omenaca, 2006. Restos directos de dinosaurios (Saurischia, Ornithischia)
en el Barremiense (Cretacico Inferior) de la Cordillera Iberica en Aragon (Teruel,
Espana). PhD Thesis. Universidad de Zaragoza. 584 pp.
unnamed 'paronychodontid' (Zinke and Rauhut, 1994)
Early Barremian, Early Cretaceous
Camarillas Formation, Spain
Material- (JHM POCA-H7) anterior tooth (3.3 mm)
(JHM POCA-H8) lateral tooth (2.08 mm)
Comments- These teeth have 2-3 labial ridges and 3-4 lingual ridges.
They lack mesial serrations, and have weak distal serrations (14-16/mm). Distal
serrations are rounded and directed slightly apically.
References- Zinke and Rauhut, 1994. Small theropods (Dinosauria, Saurischia)
from the Upper Jurassic and Lower Cretaceous of the Iberian Peninsula. Berliner
geowiss. Abh.. E 13, 163-177.
Ruiz-Omenaca, 2006. Restos directos de dinosaurios (Saurischia, Ornithischia)
en el Barremiense (Cretacico Inferior) de la Cordillera Iberica en Aragon (Teruel,
Espana). PhD Thesis. Universidad de Zaragoza. 584 pp.
unnamed 'paronychodontid' (Pol, Buscalioani, Carballeira, Frances, Martinez,
Marandat, Moratalla, Sanz, Sige and Villatte, 1992)
Maastrichtian, Late Cretaceous
Calizas de Lychnus Formation, Spain
Material- four teeth
Comments- These teeth were reported as cf. Paronychodon by Pol
et al. (1992), then referred to cf. Euronychodon sp. by Canudo and Ruiz-Omenaca
(2003), but they differ from both lacustris and portucalensis
in being serrated distally.
References- Pol, Buscalioani, Carballeira, Frances, Martinez, Marandat,
Moratalla, Sanz, Sige and Villatte, 1992. Reptiles and mammals from the Late
Cretaceous new locality Quintanilla del Coco (Burgos Province, Spain). Neues
Jahrbuch f�r Geologie und Pal�ontologie, Abhandlungen. 184(3), 279-314.
Canudo and Ruiz-Omenaca, 2003. Los restos directos de dinosaurios terop�dos
(excluyendo Aves) en Espa�a. Ciencias de la Tierra. 26, 347-373.
Paronychodon Cope, 1876
= Euronychodon Antunes and Sigogneau-Russell, 1991
= "Plesiosaurodon" Nessov vide Sues and Averianov, 2013
Diagnosis- serrations absent from teeth.
Comments- This genus is here restricted to 'paronychodontid' teeth without
serrations mesially or distally, following Currie et al. (1990). Estes (1964)
included serrated specimens with ridges and flattened lingual sides in the genus
as well, though these are here referred to Zapsalis abradens (= Saurornitholestes spp. and ?Dromaeosaurus
morphotype A of Sankey et al., 2002) and the unnamed troodontid of Sankey et
al. (2002). Many unstudied specimens catalogued or listed as Paronychodon
probably belong to these two taxa. For instance, one photographed tooth of AMNH
27122 has distal serrations in addition to ridges, so is probably a Zapsalis
specimen. Teeth matching the Paronychodon morphotype are known from the
Barremian-Maastrichtian of North America and Eurasia, indicating it was probably
a clade comparable to theropod families in scope, but which didn't show much
dental variation. Once cranial and/or postcranial remains are identified, additional
genera of 'paronychodontids' will probably need to be named.
Relationships- Paronychodon was first compared to plesiosaurs
by Cope (1876a), who soon (1876b) realized it was theropod based on comparison
to Zapsalis. Most authors retain the genus as Theropoda incertae sedis,
though some have tried to clarify its relationships further. Estes (1964) referred
it to Coeluridae, while Russell (1984) referred it to Dromaeosauridae, and Osmolska
and Barsbold (1990) to Troodontidae. These assignments were all done without
justification.
More recently, Zinke and Rauhut (1994) suggested a sister group relationship
to troodontids based on the large apically angled distal serrations of the Guimarota
'paronychodontid' teeth and the basal constriction in some teeth. These characters
are now known in basal dromaeosaurids too (e.g. Microraptor). Rauhut
and Zinke (1995) suggested assigning Una Formation Paronychodon teeth
to Pelecanimimus, but Rauhut later (2002) considered this unlikely after
communication with Perez-Moreno.
Rauhut suggested instead that at least the Una Paronychodon could be
archaeopterygids, based on the constricted base, lingually bent carinae (forming
mesial and distal grooves along the carinae lingually), and labiodistally twisted
tips. Yet the last two characters have yet to be reported from Late Cretaceous
Paronychodon, which differs from the Una specimens in several details
in any case.
Another hypothesis was given in an abstract by Sankey (2002), who purported
to show that Paronychodon is a morphotype of Richardoestesia? isosceles,
based on morphology and relative abundance. The details of this study have yet
to be published, though it does make sense stratigraphically, as both taxa first
appear in Late Jurassic Europe and spread to North America in the Albian, with
Late Cretaceous examples known from the Western North America, Central Asia
and Europe. It's also logical anatomically, as Richardoestesia? isosceles
would be expected to have some unserrated and possibly constricted teeth if
it were microraptorian. It should be noted Paronychodon has priority
over Richardoestesia, and lacustris and caperatus both
have priority over isosceles. Also, Euronychodon has priority
over Asiamericana, and portuculensis has priority over both asiatica
and asiaticus. So if this synonymy is proven, none of the names associated
with straight-toothed Richardoestesia will survive synonymization. Longrich
(2008) proposed such a synonymy based on the weak longitudinal ridges on some
Richardoestesia teeth, but Larson (2008) noted other contemporaneous
theropods sometimes have weak ridges too (tyrannosaurids, 'velociraptorines')
and that varied morphologies probably reflect positional variation in Paronychodon
teeth. Thus it is unlikely they existed in the same jaws as Richardoestesia
teeth.
Sues and Averianov (in prep.) will propose that Paronychodon are juvenile
deinonychosaurs, probably in part based on Zapsalis-like
specimens which mix 'paronychodontid' longitudinal ridges with
dromaeosaurid-like serrations and are dromaeosaurid premaxillary teeth
(Currie and Evans, 2019). It is true that Paronychodon teeth are smaller than Dromaeosaurus
teeth in the Dinosaur Park Formation, for instance, and that embryonic Troodon
has serrationless teeth. Yet apparently intermediate Zapsalis teeth are
not intermediate in size, as might be expected if they were subadults. This
has not been published besides a mention by Averianov (2007) though, so is difficult
to evaluate.
Hwang (2005, 2007) found that Paronychodon teeth are identical in enamel
microstructure to serrationless troodontids like Byronosaurus and IGM
100/1323, while Richardoestesia more closely matched dromaeosaurids.
Thus they are here placed as basal troodontids.
Pachycephalosaur fangs?- An odd possibility was suggested by Olshevsky
(DML, 1997), that some Paronychodon specimens, including the holotype,
may be anterior dentary fangs of "homalocephalid" pachycephalosaurs
(cf. Goyocephale) . However, the dentary fangs of Goyocephale
are serrated distally, polygonal in section, have a bulbous root, and seem to
only possess one lingual ridge. Premaxillary teeth of Goyocephale are
somewhat similar to type B teeth of Paronychodon, but lack ridges, are
much less labiolingually compressed, and have distal serrations apically. Stegoceras
teeth are even less similar, being serrated both mesially and distally with
no ridges. The supposed Middle Jurassic pachycephalosaur Ferganocephale
has vertical enamel ridges on the base of one side and lacks serrations, but
is otherwise highly distinct, being unrecurved, short and uncompressed labiolingually,
with a prominent cingulum. The ridges radiate from the base instead of the apex
and the entire tooth shape is distinctively ornithischian. Besides the anatomical
differences, stratigraphically Paronychodon and "homalocephalids"
are also mismatched, with the latter only known from the Campanian-Maastrichtian
of Mongolia and perhaps China. The utter lack of "homalocephalids"
in well sampled strata like the Dinosaur Park Formation is particularily telling.
Also notable is that each "homalocephalid" only had eight fang-like
teeth, but had around sixty-six leaf-shaped teeth. So we would expect more pachycephalosaur
teeth by a factor of 8:1 or so (even more in Maastrichtian formations where
pachycephalosaurines dominate, with no premaxillary teeth and greater numbers
of cheek teeth), but Baszio (1997) showed this is not the case. For instance,
he recorded 12 Paronychodon teeth from the Dinosaur Park Formation, and
only 16 pachycephalosaur teeth. Similarly, Baszio recorded 84 Paronychodon
teeth from the Milk River Formation, but only 16 pachycephalosaur teeth. Finally,
Zinke and Rauhut (1994) described 'paronychodontid' teeth within a theropod dentary
fragment, though these differ from Paronychodon in some details.
Junior synonyms?- Zapsalis was based on a tooth described by Cope
(1876) and synonymized with Paronychodon lacustris by Estes (1964).
However, Zapsalis falls outside the current concept of Paronychodon
in having serrations, and is more robust than teeth of that genus as well. Instead
it matches ?Dromaeosaurus morphotype A of Sankey et al. (2002) and was recently found by Currie and Evans (2019) to be identical to premaxillary teeth
of Saurornitholestes. Bambiraptor and Velociraptor also have lingual ridges, so that Zapsalis is here considered a morphotype of dromaeosaurid premaxillary teeth.
Tripriodon was based on teeth assigned to two species by Marsh (1889)-
the genotype T. coelatus, and T. caperatus. The former is a junior
synonym of the multituberculate Meniscoessus robustus (as first shown
by Osborn, 1891), while the second belongs to Paronychodon (as first
shown by Estes, 1964). Estes was incorrect in synonymizing Tripriodon
itself with Paronychodon however, as T. caperatus is only a referred
species. This also prevents Marsh's Tripriodontidae from being a theropod family.
Three teeth were originally referred to Paronychodon lacustris (Antunes
and Brion, 1988), but later described as the new genus Euronychodon by
Antunes and Sigogneau-Russell (1992). This was based on the supposed absence
of longitudinal depressions and a median ridge. Yet longitudinal depressions
appear to be present in the figure, while identical ridge patterns are seen
in some Paronychodon teeth (e.g. Milk River specimens). Euronychodon
is thus retained as a junior synonym of Paronychodon here (as in Sige
et al., 1997 and Rauhut, 2002). Numerous other teeth were later referred to
Euronychodon from the Barremian-Maastrichtian of Eurasia, but they are
here listed as Paronychodon or unnamed 'paronychodontids'.
Variation- Sankey et al. (2002) showed there are two morphotypes of Paronychodon
teeth. Type A teeth are elongate and straighter, with unconstricted bases. Type
B teeth are short and strongly recurved, with constricted bases. They interpreted
these are being due to positional variation instead of taxonomic varation, though
it should be noted that the holotypes of both Paronychodon lacustris
and Tripriodon caperatus are type A teeth. Thus if taxonomic variation
proves correct, these names should only be associated with type A teeth. Based
on comparison to other theropods, type B teeth may be more posterior in position.
Baszio (1997) noted that half the teeth are flat lingually, and the other half
are biconvex. He considered this due to positional variation, with biconvex
teeth being from the posterior portion of the jaw. The FABL, BW and FABL/BW
measurements of lingually flat teeth were not significantly different from those
of biconvex teeth. Again, the holotypes of both Paronychodon lacustris
and Tripriodon caperatus are flattened lingually, which should restrict
the names to lingually flat teeth if the variation is later shown to be taxonomic.
References- Cope, 1876a. Descriptions of some vertebrate remains from
the Fort Union Beds of Montana. Proceedings of the Academy of Natural Sciences
of Philadelphia. 28, 248-261.
Cope, 1876b. On some extinct reptiles and batrachia from the Judith River and
Fox Hills Beds of Montana. Proceedings of the Academy of Natural Sciences, Philadelphia.
28, 340-359.
Marsh, 1889. Discovery of Cretaceous mammalia. American Journal of Science,
3rd series. 38, 81-92.
Osborn, 1891. A review of the "Discovery of the Cretaceous Mammalia".
The American Naturalist. 25(295), 595-611.
Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern
Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Russell, 1984. A check list of the families and genera of North American dinosaurs.
Syllogeus. 53, 1-35.
Antunes and Brion, 1988. Le Cretace terminal de Beira Litoral, Portugal: remarques
stratigraphicques et ecologiques, etude complementaire de Rosasia soutoi
(Chelonii, Bothremydidate). Ciencias de Terra. 9, 153-200.
Osmolska and Barsbold, 1990. Troodontidae. In Weishampel, Dodson, and Osmolska,
eds.. The Dinosauria, Berkeley: University of California Press: 259-268.
Antunes and Sigogneau-Russell, 1992. La faune de petits dinosaures du Cretace
Terminal Portugais. Comun. Serv. Geol. Portugal. 78(1), 49-62.
Zinke and Rauhut, 1994. Small theropods (Dinosauria, Saurischia) from the Upper
Jurassic and Lower Cretaceous of the Iberian Peninsula. Berliner geowiss. Abh..
E 13, 163-177.
Rauhut and Zinke, 1995. A description of the Barremian dinosaur fauna from Una
with a comparison to that of Las Hoyas. In II International Symposium on Lithographic
Limestones, Lleida-Cuenca (Spain), 9th–16th July 1995, Extended Abstracts.
123-126.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Olshevsky, DML 1997. https://web.archive.org/web/20190623204714/http://dml.cmnh.org/1997Dec/msg00058.html
Sige, Buscalioni, Duffaud, Gayet, Orth, Rage and Sanz, 1997. Etat des donn�es
sur le gisement Cr�tac� sup�rieur continental de Champ-Garimond
(Gard, Sud de la France). Munchner Geowiss.. 34, 111-130.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca,
Spain. Cretaceous Research. 23, 255-263.
Sankey, 2002. Theropod dinosaur diversity in the latest Cretaceous (Maastrichtian)
of North America. Journal of Vertebrate Paleontology. 22(3), 103A.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Hwang, 2005. Phylogenetic patterns of enamel microstructure in dinosaur teeth.
Journal of Morphology. 266, 208-240.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern
Aral Sea region, Kazakhstan. Cretaceous Research.
Hwang, 2007. Phylogenetic patterns of enamel microstructure in dinosaur teeth.
PhD thesis. Columbia University. 274 pp.
Larson, 2008. Diversity and variation of theropod dinosaur teeth from the uppermost
Santonian Milk River Formation (Upper Cretaceous), Alberta: a quantitative method
supporting identification of the oldest dinosaur tooth assemblage in Canada.
Canadian Journal of Earth Science. 45, 1455-1468.
Longrich, 2008. Small theropod teeth from the Lance Formation of Wyoming, USA.
in Sankey and Baszio (eds). Vertebrate Microfossil Assemblages: Their Role in
Paleontology and Paleobiogeography. Indiana University Press, Bloomington, Ind..
pp. 135-158.
Sues and Averianov, 2013. Enigmatic teeth of small theropod dinosaurs from the
Upper Cretaceous (Cenomanian–Turonian) of Uzbekistan. Canadian Journal
of Earth Sciences. 50, 306-314.
Currie and Evans, 2019. Cranial anatomy of new specimens of Saurornitholestes langstoni (Dinosauria, Theropoda, Dromaeosauridae) from the Dinosaur Park Formation (Campanian) of Alberta. The Anatomical Record. DOI: 10.1002/ar.24241
P. lacustris Cope, 1876
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Holotype- (AMNH 3018) type A tooth (10 mm)
Referred - (AMNH 8522) type A tooth (13 mm) (Sahni, 1972)
(AMNH 8523) type B tooth (4.8 mm) (Sahni, 1972)
(AMNH 8524) nine teeth (Sahni, 1972)
(AMNH coll.) seven teeth (Sahni, 1972)
(MOR 017) tooth (MOR online)
Early Campanian, Late Cretaceous
Milk River Formation, Alberta, Canada
Material- (CMN coll.) teeth (Russell, 1935)
(UA MR-4: 46) tooth (Baszio, 1997)
(UA MR-47) tooth (Baszio, 1997)
(UA MR-48) type B tooth (Baszio, 1997)
(UA MR-49) type A tooth (Baszio, 1997)
(UA MR-50) type A tooth (Baszio, 1997)
(UA MR-51) type A tooth (Baszio, 1997)
(UA MR-52) tooth (Baszio, 1997)
(UA MR-53) type A tooth (Baszio, 1997)
(UA MR-54) type A tooth (Baszio, 1997)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith River Group, Alberta, Saskatchewan, Canada
Material- (RTMP 83.36.8) type A tooth (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 84.36.53) type A tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.57.62) type A tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.60.114) tooth (Ryan and Russell, 2001)
(RTMP 87.19.65) type B tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 87.112.11) type B tooth (4.3 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 88.211.66) type A tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 94.12.187) type A tooth (11.5 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.145.53) type B tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 95.177.50) type B tooth (~5.2 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 95.187.28) type B tooth (5.1 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
(RTMP 99.55.61) type A tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 99.55.230) type A tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 2000.19.2) type B tooth (5.2 mm) (Sankey, Brinkman, Guenther and Currie,
2002)
nine teeth (Baszio, 1997)
material (Tokaryk, 1988)
Late Campanian, Late Cretaceous
De-na-zin Member of the Kirtland Formation, New Mexico, US
Material- (SMP VP-1354) tooth (Sullivan and Lucas, 2006)
Late Maastrichtian, Late Cretaceous
Frenchman Formation, Saskatchewan, Canada
Material- four teeth (Baszio, 1997)
Diagnosis- (after Sahni, 1972) smaller than P. caperatus and with
less defined ridges.
Comments-
The holotype has a FABL of 4 mm and a BW of 2.4 mm. It is recurved and
flattened lingually with four lingual ridges and six labial ridges.
Serrations are absent. It is not from the Hell Creek Formation, contra
some sources. Though Sankey et al. (2002) claimed it is not
illustrated, Glut (1997) included a photo. Note the collection number
also includes other material such as the incomplete theropod tibia
photographed on the AMNH database website. Baszio (1997) stated
that Paronychodon teeth from the Judith River Group,
Milk River Formation and Frenchman Formations were identical. Sullivan and Lucas
(2006) in turn stated that SMP VP-1354 from the Kirtland Formation is identical
to Milk River teeth. All of these are smaller and with less defined ridges than
the Lance Formation species. They can be referred to the species P. lacustris,
while the Lance Formation material can be referred to P. caperatus. It's
probable other Campanian-Maastrichtian Paronychodon remains (listed as
P. sp. below) will be referrable to P. lacustris once they are
examined in more detail. Furthermore, this P. lacustris morphotype probably
belonged to several different species which differed in non-dental characters,
which explains its apparently wide distribution.
References- Cope, 1876. Descriptions of some vertebrate remains from
the Fort Union Beds of Montana. Proceedings of the Academy of Natural Sciences
of Philadelphia. 28, 248-261.
Russell, 1935. Fauna of the Upper Milk River beds, Southern Alberta. Transactions,
Royal Society of Canada. 3(29), 115-127.
Sahni, 1972. The vertebrate fauna of the Judith River Formation, Montana. Bulletin
of the AMNH. 147.
Tokaryk, 1988. Preliminary vertebrate faunal list of the Oldman Formation Saskatchewan.
Journal of Vertebrate Paleontology. 8(3), 28A.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Glut, 1997. Dinosaurs, the Encyclopedia. Mcfarland & Company, Inc..
1076 pp.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Sullivan and Lucas, 2006. The Kirtlandian land-vertebrate "age" -
faunal composition, temporal position and biostratigraphic correlation in the
nonmarine Upper Cretaceous of western North America. New Mexico Museum of Natural
History and Science Bulletin. 35, 7-29.
P. caperatus (Marsh, 1889)
Olshevsky, 1991
= Tripriodon caperatus Marsh, 1889
= Menisocoessus caperatus (Marsh, 1889)
= Dipriodon caperatus (Marsh, 1889) Currie, Rigby and Sloan, 1990
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Holotype- (YPM 10624; = YPM 11852) type A tooth
Referred- (AMNH 21550) tooth (AMNH online)
(AMNH 21551) tooth (AMNH online)
(AMNH 21811) twenty-four teeth (AMNH online)
(AMNH 21812) sixteen teeth and fragments (AMNH online)
(AMNH 21813) sixteen teeth (AMNH online)
(AMNH 21814) three teeth (AMNH online)
?(AMNH 21881) two teeth (AMNH online)
?(AMNH 21882) sixteen teeth (AMNH online)
?(AMNH 21883) tooth (AMNH online)
(AMNH 24930) eighteen teeth (AMNH online)
?(AMNH 24931) tooth (AMNH online)
?(AMNH 24932) four teeth (AMNH online)
(AMNH 27122 in part) sixteen teeth and fragments (AMNH online)
?(AMNH coll.) teeth (Estes, 1964)
(SDSM 12459) two teeth (Whitmore, 1988)
(SDSM 12460) tooth (Whitmore, 1988)
(SDSM 12461) tooth (Whitmore, 1988)
(SDSM 12707) tooth (Whitmore, 1988)
(SDSM 15101) tooth fragment (Whitmore, 1988)
(UA BTB: 134) tooth (Baszio, 1997)
(UA BTB: 136) tooth (Baszio, 1997)
(UA BTB: 137) tooth (Baszio, 1997)
(UA BTB: 138) type A tooth (Baszio, 1997)
(UA BTB: 146) tooth (Baszio, 1997)
?(UA BTB: 166) type B tooth (Baszio, 1997)
(UA BTB: coll.) tooth (Baszio, 1997)
(UCM 38288) (juvenile) tooth (4 mm) (Carpenter, 1982)
(UCM 38459) (juvenile) tooth (1.7 mm) (Carpenter, 1982)
(UCMP 53283) tooth (UCMP online)
(UCMP 73075) teeth (UCMP online)
(UCMP 73076) teeth (UCMP online)
(UCMP 85135) tooth (UCMP online)
(UCMP 124400) three teeth (UCMP online)
(UCMP 124401) four teeth (UCMP online)
(UCMP 186863) teeth (UCMP online)
(UCMP 186919) teeth (UCMP online)
(UCMP 187085-187134) fifty teeth (UCMP online)
(UW 14091) tooth (Breithaupt, 1982)
(UW 14092) tooth (Breithaupt, 1982)
?(YPM 10625) tooth (Marsh, 1889)
?(YPM 54481) (YPM online)
?(YPM 54490) (YPM online)
(YPM 54999) (YPM online)
(YPM 55021) (YPM online)
(YPM 55022) (YPM online)
(YPM 55498) (YPM online)
?(YPM 55501) (YPM online)
(YPM 55514) (YPM online)
?(YPM 55517) (YPM online)
?(YPM 55527) (YPM online)
(YPM 55528) (YPM online)
?(YPM 55533) (YPM online)
(YPM 55547) (YPM online)
(YPM 55548) (YPM online)
?(YPM 55585) (YPM online)
(YPM 55594) (YPM online)
(YPM 10630) partial type A tooth (Marsh, 1889)
(YPM coll.) tooth (Osborn, 1891)
tooth (Breithaupt, 2001)
Diagnosis- (after Sahni, 1972) larger than P. lacustris and with
better defined ridges.
Comments- Marsh (1889) originally described Tripriodon caperatus
based on several teeth and tooth fragments discovered that year, including supposed lower incisors.
The holotype is often listed as YPM 11853, but is 11852 in the YPM online catalog.
Although Marsh states the specimen came from Laramie beds, the Laramie Formation
has not yielded dinosaurs in Wyoming, and the YPM online catalog confirms it
was actually found in the Lance Formation. Marsh viewed T. caperatus
and the genotype T. coelatus as belonging to a new family Tripriodontidae
in his Allotheria, most closely related to Stereognathus (now recognized
as a tritylodontid). Tripriodon coelatus is based on a molar which is
now thought to belong to Meniscoessus robustus, a species of multituberculate.
This was first recognized by Osborn (1891), who believed T. caperatus
were lower incisors of Meniscoessus. Osborn also recognized supposed
lower incisors of Selenacodon brevis (YPM 10630) and Tripriodon coelatus
(YPM coll.) have the same morphology as T. caperatus. Selenacodon
brevis is now viewed as a junior synonym of the multituberculate Cimolemys
gracilis. Estes (1964) incorrectly believed T. caperatus to be the
genotype, and was the first of many authors to synonymize Tripriodon
with Paronychodon. He referred the T. caperatus holotype and YPM
10630 to Paronychodon lacustris. Currie et al. (1990) mistakenly listed
Dipriodon caperatus, but Dipriodon is another genus now viewed
as a synonym of Meniscoessus.
The holotype tooth seems nearly identical to the Paronychodon holotype
except for having a shorter crown. It clearly matches tooth type A of Sankey
et al. (2002). Baszio (1997) notes these Lance Formation teeth are larger than
Paronychodon lacustris from the Milk River, Dinosaur Park, Frenchman
and Horseshoe Canyon Formations, with more pronounced wrinkles. They are retained
as Paronychodon caperatus here, after Olshevsky (1991). UA BTB 166 is
unique in being biconvex with helical ridges.
References- Marsh, 1889. Discovery of Cretaceous Mammalia. American Journal
of Science, 3rd series. 38, 81-92.
Osborn, 1891. A review of the "Discovery of the Cretaceous Mammalia".
The American Naturalist. 25(295), 595-611.
Estes, 1964. Fossil vertebrates from the Late Cretaceous Lance Formation, eastern
Wyoming. University of California Publications in Geological Sciences. 49, 1-180.
Breithaupt. 1982. Paleontology and paleoecology of the Lance Formation, (Maastrichtian),
east flank of Rick Springs Uplift, Sweetwater County, Wyoming. Contributions
to Geology, University of Wyoming. 21(2), 123-151.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek
formations and a description of a new species of theropod. Contributions to
Geology, University of Wyoming. 20(2), 123-134.
Whitmore, 1988. The vertebrate paleontology of Late Cretaceous (Lancian) localities
in the Lance Formation, Northern Niobrara County, Wyoming. Unpublished Masters
Thesis. South Dakota School of Mines and Technology, Rapid City, South Dakota.
130pp.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.
Olshevsky, 1991. A Revison of the Parainfraclass Archosauria Cope, 1869, Excluding
the Advanced Crocodyila. Mesozoic Menanderings #2 (1st printing). iv + 196pp.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Breithaupt, 2001. Passport-in-time microvertebrate fossil project at the University
of Wyoming Geological Museum: Late Cretaceous Paleontological resources in the
public eye. in Santucci and McClelland (eds). Proceedings of the 6th fossil
resource conference. United States Department of the interior, National Park
Service, Geological Resource Division. 107-112.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Stokosa 2005. Enamel microstructure variation within the Theropoda. in Carpenter
(ed). The Carnivorous Dinosaurs. 163-178.
P. sp. (Peng, Russell and Brinkman, 2001)
Late Campanian, Late Cretaceous
Foremost Formation of the Judith River Group, Alberta, Canada
Material- (RTMP 96.62 coll.) three teeth
Reference- Peng, Russell and Brinkman, 2001. Vertebrate microsite assemblages
(exclusive of mammals) from the Foremost and Oldman Formations of the
Judith River Group (Campanian) of southeastern Alberta: An illustrated
guide. Provincial Museum of Alberta Natural History Occasional Paper. 25, 54 pp.
P. sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Oldman Formation of the Judith River Group, Alberta, Canada
Material- (RTMP 92.77.6) tooth
(RTMP coll.) teeth
Reference- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press,
Bloomington, Indiana. pp. 279-297.
P. sp. (Fanti and Miyashita, 2009)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
Material- (UALVP 48815) tooth (3.9x2.3x1.1 mm)
Reference- Fanti and Miyashita, 2009. A high latitude vertebrate fossil
assemblage from the Late Cretaceous of west-central Alberta, Canada: Evidence
for dinosaur nesting and vertebrate latitudinal gradient. Palaeogeography, Palaeoclimatology,
Palaeoecology. 275, 37-53.
P. sp. (Baszio, 1997)
Early Maastrichtian, Late Cretaceous
Horseshoe Canyon Formation, Alberta, Canada
Material- (RTMP 97.39.6) partial type A tooth (Ryan, Currie, Gardner, Vickaryous
and Lavigne, 1998)
(RTMP 1041) tooth (Baszio, 1997)
(RTMP 2009.139.4) tooth (Larson, Brinkman and Bell, 2010)
Comments- RTMP 97.39.6 is flat lingually and convex labially. It has
eight lingual ridges and ten labial ones. RTMP 2009.139.4 also has that convexity,
but also faint serrations.
References- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan, Currie, Gardner and Livigne, 1997. Baby hadrosaurid material associated
with an unusually high abundance of troodontid teeth from the Horseshoe Canyon
Formation (Early Maastrichtian), Alberta, Canada. Journal of Vertebrate Paleontology.
17(3), 72A.
Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998. Baby hadrosaurid material
associated with an unusually high abundance of Troodon teeth from the Horseshoe
Canyon Formation, Upper Cretaceous, Alberta, Canada. Gaia. 15, 123-133.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe
Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta,
with special reference to the Albertosaurus sarcophagus bonebed. Canadian
Journal of Earth Sciences. 47(9), 1159-1181.
P? sp. (Krumenacker and Scofield, 2015)
Late Albian-Cenomanian, Early-Late Cretaceous
Wayan Formation, Idaho, US
Material- (IMNH 2251/49862; Morph 5) tooth (?x~6.2x? mm)
References- Krumenacker and Scofield, 2015. A diverse theropod tooth
assemblage from the Mid-Cretaceous (Albian-Cenomanian) Wayan Formation of Idaho.
Journal of Vertebrate Paleontology. Program and Abstracts 2015, 158.
Krumenacker, Simon, Scofield and Varricchio, 2016. Theropod dinosaurs from the
Albian-Cenomanian Wayan Formation of eastern Idaho. Historical Biology. DOI:
10.1080/08912963.2015.1137913
P. sp. (Redman, Moore and Varricchio, 2015)
Late Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
Material- (MOR 221) tooth (MOR online)
(MOR 505) tooth (MOR online)
(MOR 516) teeth (MOR online)
teeth (Redman, Moore and Varricchio, 2015)
Reference- Redman, Moore and Varricchio, 2015. A new vertebrate microfossil
locality in the Upper Two Medicine Formation in the vicinity of Egg Mountain.
Journal of Vertebrate Paleontology. Program and Abstracts 2015, 201-202.
P. sp. (Estes, Berberian and Mesozoely, 1969)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, South Dakota, US
Material- (AMNH 21555) tooth (AMNH online)
(MCZ 3645) two teeth (Estes, Berberian and Mesozoely, 1969)
(UCMP 119922) tooth (UCMP online)
(UCMP 119923) tooth (UCMP online)
(UCMP 120076) tooth (UCMP online)
(UCMP 120192) tooth (UCMP online)
(UCMP 120254) tooth fragment (UCMP online)
(UCMP 123341) tooth (UCMP online)
(UCMP 124405) tooth (UCMP online)
(UCMP 124990) (juvenile) type B tooth (3.4 mm) (Carpenter, 1982)
(UCMP 124991) (juvenile) tooth (4.8 mm) (Carpenter, 1982)
(UCMP 124992) (juvenile) tooth (4.3 mm) (Carpenter, 1982)
(UCMP 128764) three teeth (UCMP online)
(UCMP 186867) teeth (UCMP online)
(UCMP 186876) teeth (UCMP online)
(UCMP 186879) teeth (UCMP online)
(UCMP 186884) teeth (UCMP online)
(UCMP 186898) teeth (UCMP online)
(UCMP 186910) teeth (UCMP online)
(UCMP 186923) teeth (UCMP online)
(UCMP 187080) tooth (UCMP online)
(UCMP 187081) tooth (UCMP online)
(UCMP 187082) tooth (UCMP online)
(UCMP 187135-187156) twenty-two teeth (UCMP online)
(YPM PU 20571) (Estes, Berberian and Mesozoely, 1969)
(YPM 56974) two teeth (YPM online)
teeth (Stenerson and O'Conner, 1994)
teeth (Triebold, 1997)
teeth (DePalma, 2010)
References- Estes, Berberian and Mesozoely, 1969. Lower vertebrates from
the Late Cretaceous Hell Creek Formation, McCone County, Montana. Breviora.
337, 1-33.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek
formations and a description of a new species of theropod. Contributions to
Geology, University of Wyoming. 20(2), 123-134.
Stenerson and O'Conner, 1994. The Late Cretaceous Hell Creek Formation of Northwestern
South Dakota and its Fauna. MAPS Digest. 17(4), 108-120.
Triebold, 1997. The Sandy Site: Small Dinosaurs from the Hell Creek Formation
of South Dakota. in Wolberg, Stump and Rosenberg (eds). Dinofest International,
Proceedings of a Symposium sponsered by Arizona State University. A Publication
of The Academy of Natural Sciences. 245-248.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous
bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis,
University of Kansas. 227 pp.
P. sp. indet.
Cretaceous
Montana, US
Material- (AMNH 2134) tooth (AMNH online)
P. sp. (Hoganson and Erickson, 2004)
Maastrichtian, Late Cretaceous
Fox Hills Formation, North Dakota, US
Reference- Hoganson and Erickson, 2004. Paleoecological implications
of the Fox Hills Formation (Maastrichtian) reptilian and amphibian fauna from
south-central North Dakota. Geological Society of America Rocky Mountain and
Cordilleran Sections Annual Meeting, Boise, Idaho, Abstracts with Programs.
36(4), 80.
P. sp. (Breithaupt, 1985)
Late Campanian, Late Cretaceous
Mesaverde Formation, Wyoming, US
Material- (AMNH 12881) tooth (Demar and Breithaupt, 2006)
(AMNH 12882) tooth (Demar and Breithaupt, 2006)
(UCMP 120848) tooth (UCMP online)
(UW 34819) tooth (Demar and Breithaupt, 2006)
References- Breithaupt, 1985. Nonmammalian vertebrates faunas from the
late Cretaceous of Wyoming. Thirty-Sixth Annual Field Conference-1985, Wyoming
Geological Association Guidebook. 159-175.
Demar and Breithaupt, 2006. The nonmammalian vertebrate microfossil assemblages
of the Mesaverde Formation (Upper Cretaceous, Campanian) of the Wind River and
Bighorn Basin, Wyoming. in Lucas and Sullivan (eds). Late Cretaceous Vertebrates
from the Western Interior. New Mexico Museum of Natural History & Science,
Bulletin. 35, 33-53.
P. sp. (Wroblewski, 1995)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
Material- teeth
References- Wroblewski, 1995. First report of changes in Lower Vertebrate
Faunas across the Cretaceous-Tertiary boundary, Western Hanna Basin, Wyoming.
Journal of Vertebrate Paleontology. 15(3), 61A.
Wroblewski, 1998. Changing paleoenvironments and paleofaunas across the K-T
boundary, Ferris Formation, Southcentral Wyoming. Tate Geological Museum, Casper
College, Casper Wyoming. Tate ’98. Life in the Cretaceous. 53-70.
P. sp. (Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten,
Eaton, Hasiotis and Lawton, 1997)
Late Cenomanian, Late Cretaceous
Dakota Formation, Utah, US
Material- teeth
Comments- These were listed as cf. Paronychodon sp. by Kirkland
et al. (1997).
Reference- Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton,
Hasiotis and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the
Central Colorado Plateau: a key to understanding 35 million years of tectonics,
sedimentology, evolution, and biogeography. Brigham Young University Geology
Studies. 42, 69-103.
P. sp. (Kirkland and Parrish, 1995)
Cenomanian-Early Turonian, Late Cretaceous
Mussentuchit Member of the Cedar Mountain Formation, Utah, US
Material- (CM 72650) tooth fragment (Fiorillo, 1999)
(NCSM 33277) incomplete type B tooth (~3.3x1.57x1.00 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
(NCSM 33298) type A tooth (2.11x1.16x0.77 mm) (Avrahami, Gates, Heckert, Makovicky and Zanno, 2018)
teeth (Kirkland et al., 1997)
partial type A tooth (Garrison et al., 2007)
Comments- Kirkland and Parrish (1995), Kirkland et al. (1997) and Cifelli
et al. (1999) list cf. Paronychodon sp. teeth. Fiorillo (1999) describes
a tooth fragment possessing a flattened side with longitudinal ridges which
he assigns to "Paronychodon", and believes is a dromaeosaurid
tooth based on its cross section. Garrison et al. (2007) describes a partial
tooth lacking serrations and a flattened side with three ridges, referring it
to cf. Paronychodon sp..
References- Kirkland and Parrish, 1995. Theropod teeth from the Lower
Cretaceous of Utah. Journal of Vertebrate Paleontology. 15(3), 39A.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis and Lawton,
1997. Lower to Middle Cretaceous dinosaur faunas of the Central Colorado Plateau:
a key to understanding 35 million years of tectonics, sedimentology, evolution,
and biogeography. Brigham Young University Geology Studies. 42, 69-103.
Cifelli, Nydam, Gardner, Weil, Eaton, Kirkland, Madsen, 1999. Medial Cretaceous
vertebrates from the Cedar Mountain Formation, Emery County, Utah: the Mussentuchit
Local Fauna. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 219-242.
Fiorillo, 1999. Non-mammalian microvertebrate remains from the Robison Eggshell
site, Cedar Mountain Formation (Lower Cretaceous), Emery County, Utah. in Gillette
(ed.). Vertebrate Paleontology in Utah. Utah Geological Survey, Miscellaneous
Publication. 99-1, 259-268.
Garrison, Brinkman, Nichols, Layer, Burge and Thayn, 2007. A multidisciplinary
study of the Lower Cretaceous Cedar Mountain Formation, Mussentuchit Wash, Utah:
a determination of the paleoenvironment and paleoecology of the Eolambia
caroljonesa dinosaur quarry. Cretaceous Research. 28, 461-494.
Avrahami, 2018. Paleobiodiversity of a new microvertebrate locality
from the Upper Cretaceous Mussentuchit Member, Cedar Mountain
Formation, Utah: Testing morphometric multivariate approaches for
quantifying shape variation in microvertebrate specimens. Masters
thesis, North Carolina State University. 181 pp.
Avrahami, Gates, Heckert, Makovicky and Zanno, 2018. A new
microvertebrate assemblage from the Mussentuchit Member, Cedar Mountain
Formation: Insights into the paleobiodiversity and paleobiogeography of
early Late Cretaceous ecosystems in western North America. PeerJ.
6:e5883.
P. sp. (Kirkland, Lucas and Estep, 1998)
Middle-Late Turonian, Late Cretaceous
Smoky Hollow Member of the Straight Cliffs Formation, Utah, US
Material- (MNA 995) tooth (Parrish, 1999)
(OMNH 24451) tooth (Parrish, 1999)
(OMNH 25415) tooth (Parrish, 1999)
Comments- These were listed as cf. Paronychodon by Kirkland et
al. (1998) and Parrish (1999).
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian)
of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
P. sp. (Kirkland, Lucas and Estep, 1998)
Coniacian-Santonian, Late Cretaceous
John Henry Member of the Straight Cliffs Formation, Utah, US
Comments- This is listed as cf. Paronychodon sp. by Kirkland et
al. (1998).
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the
Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous
Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin.
14, 79-89.
P. sp. (Kirkland, Lucas and Estep, 1998)
Early Campanian, Late Cretaceous
Wahweap Formation, Utah
Comments- This is listed as cf. Paronychodon sp. by Kirkland et
al. (1998).
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the
Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous
Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin.
14, 79-89.
P. sp. (Kirkland, Lucas and Estep, 1998)
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Material- (UCM 8304) tooth (Parrish, 1999)
(OMNH 24161) tooth (Parrish, 1999)
(OMNH 24164) tooth (Parrish, 1999)
Comments- These were listed as cf. Paronychodon by Kirkland et
al. (1998) and Parrish (1999).
Reference- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of the
Colorado Plateau. in Lucas, Kirkland and Estep (eds.). Lower and Middle Cretaceous
Terrestrial Ecosystems. New Mexico Museum of Natural History and Science Bulletin.
14, 79-89.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-. Judithian)
of southern Utah. in Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
P? sp. (Armstrong-Zeigler, 1978)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (MNA Pl. 1627) nine teeth (Armstrong-Ziegler, 1978)
(NMMNH P-27490) tooth (?x1.9x1.2 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30276) tooth (1.8x1.3x.5 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30329) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30332) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33479) tooth (~4.3x2x1.1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-38430) tooth (Williamson and Brusatte, 2014)
(NMMNH P-53360) tooth (Williamson and Brusatte, 2014)
Late Campanian, Late Cretaceous
Hunter Wash Member of Kirtland Formation, New Mexico, US
(NMMNH P-29132) tooth (Williamson and Brusatte, 2014)
(NMMNH P-30218) tooth (2.6x1.5x.8 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30233) tooth (?x2.8x1.5 mm) (Williamson and Brusatte, 2014)
(NMMNH P-30234) tooth (?x2.1x1.1 mm) (Williamson and Brusatte, 2014)
Comments- MNA Pl. 1627 were listed as Paronychodon lacustris by
Armstrong-Ziegler (1978), but later as Dromaeosauridae incertae sedis by Lucas
et al. (1987). Williamson and Brusatte (2014) note this differs from Dinosaur
Park Paronychodon in having "less pronounced apicobasal ridges and
in lacking ridges that anastomose from the apex to the base of the crown."
Some resemble Zapsalis except for lacking distal serrations, so may belong
to that taxon.
References- Armstrong-Zeigler, 1978. An aniliid snake and associated
vertebrates from the Campanian of New Mexico. Journal of Paleontology. 52(2),
480-483.
Lucas, Mateer, Hunt and O’Neill, 1987. Dinosaurs, the age of the Fruitland
and Kirtland Formations, and the Cretaceous-Tertiary boundary in the San Juan
Basin, New Mexico. in Fassett and Rigby (eds). The Cretaceous-Tertiary Boundary
in the San Juan and Raton Basins, New Mexico and Colorado. The Geological Society
of America Special Paper. 209, 35-50.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
P. sp. (Langston, Standhardt and Stevens, 1989)
Early Maastrichtian, Late Cretaceous
Aguja Formation, Texas, US
Material- (LSU 113:1310) type A tooth (~5.5 mm)
(LSU 113:1311) type A tooth
(LSU 113:5107) type A tooth (~3 mm)
(LSU 113:5993) type A tooth (~2.5 mm)
(LSU 113:5996) type A tooth
References- Langston, Standhardt and Stevens, 1989. Fossil vertebrate
collecting in the Big Bend - History and retrospective. in Vertebrate Paleontology,
Biostratigraphy and Depositional Environments, Latest Cretaceous and Tertiary,
Big Bend Area, Texas. Guidebook Field Trip Numbers 1 a, B, and 49th Annual Meeting
of the Society of Vertebrate Paleontology, Austin, Texas, 29 October - 1 November
1989. 11-21.
Sankey, Standhardt and Schiebout, 2005. Theropod teeth from the Upper Cretaceous
(Campanian-Maastrichtian), Big Bend National Park, Texas. in Carpenter (ed).
The Carnivorous Dinosaurs. 127-152.
P. portucalensis (Antunes
and Sigogneau-Russell, 1991) new comb.
= Euronychodon portucalensis Antunes and Sigogneau-Russell, 1991
Late Campanian-Early Maastrichtian, Late Cretaceous
unnamed unit, Taviero, Portugal
Holotype- (CEPUNL TV 20) type A tooth (1.8 mm)
Paratypes- (CEPUNL TV 18) type B tooth
(CEPUNL TV 19) tooth
Diagnosis- provisionally indeterminate within Paronychodon.
Comments- These teeth were originally referred to Paronychodon lacustris
(Antunes and Brion, 1988). They were later described as the new genus Euronychodon
by Antunes and Sigogneau-Russell (1992) based on the supposed absence of longitudinal
depressions and a median ridge. Yet longitudinal depressions appear to be present
in the figure, while identical ridge patterns are seen in some Paronychodon
teeth (e.g. Milk River specimens). Euronychodon is thus retained as a
junior synonym of Paronychodon here (as in Rauhut, 2002).
References- Antunes and Brion, 1988. Le Cretace terminal de Beira Litoral,
Portugal: remarques stratigraphicques et ecologiques, etude complementaire de
Rosasia soutoi (Chelonii, Bothremydidate). Ciencias de Terra. 9, 153-200.
Antunes and Sigogneau-Russell, 1992. La faune de petits dinosaures du Cretace
Terminal Portugais. Comun. Serv. Geol. Portugal. 78(1), 49-62.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca,
Spain. Cretaceous Research. 23, 255-263.
P. sp. nov. (Rauhut and Zinke, 1995)
Late Barremian, Early Cretaceous
Una (= Calizas de La Huergina) Formation, Spain
Material- (IPFUB Una Th 53, 55-61, 69) thirteen teeth (to 6 mm) (Rauhut
and Zinke, 1995)
(IPFUB Una Th 69) tooth (Rauhut, 2002)
Comments- These teeth are slightly recurved, with no mesial or distal
serrations. They have a flattened lingual side with a central ridge, while some
specimens also have a weak labial ridge. Some teeth have a slight basal constriction.
Rauhut and Zinke (1995) originally referred these specimens to cf. Euronychodon
sp., though they were later referred to cf. Paronychodon sp. by Rauhut
(2002).
References- Rauhut and Zinke, 1995. A description of the Barremian dinosaur
fauna from Una with a comparison to that of Las Hoyas. In II International Symposium
on Lithographic Limestones, Lleida-Cuenca (Spain), 9th–16th July 1995,
Extended Abstracts. 123-126.
Rauhut, 2002. Microrestos de dinosaurios del Cret�cico inferior de U�a,
Espa�a. Resumenes de las XVIII Jornadas Argentinas de Paleontolog�a
de Vertebrados. Bahia Blanca, Argentina. 36.
Rauhut, 2002. Dinosaur teeth from the Barremian of Una, Province of Cuenca,
Spain. Cretaceous Research. 23, 255-263.
P? sp. (Le Loeuff, 1992)
Late Campanian, Late Cretaceous
La�o, Sedano Formation, Spain
Material- (MCNA 14562) tooth (1.9x1.3x.9 mm)
(MCNA 14563) tooth (1.8x1.1x.7 mm)
Comments- These were listed as cf. Euronychodon sp.
by Suberbiola et al. (2000). Isasmendi et al. (2020) referred at
least two of the teeth called Coelurosauria indet. by Torices et al.
(2015) to cf. Paronychodon,
as they have longitudinal grooves. It is likely more La�o teeth
listed under unnamed Maniraptoriformes on this site belong here as well.
References- Le Loeuff, 1992. Les vert�br�s continentaux
du Cr�tac� sup�rieur d'Europe: Pal�o�cologie,
Biostratigraphie et Pal�obiog�ographie. M�moires des Sciences
de la Terre de l'Universit� Pierre et Marie Curie, Paris, (Th�se
d'Universit�, non publi�). 92-3, 273 pp.
Astibia, Murelaga, Pereda-Suberbiola, Elorza and Gomez-Alday, 1999. Taphonomy
and palaeoecology of the Upper Cretaceous continental vertebrate-bearing beds
of the La�o Quarry (Iberian Peninsula). Est. Mus. Cienc. Nat. de Alava.
14 (N�m. Espec. 1), 43-104.
Pereda-Suberbiola, Asibia, Murelaga, Elzorza and Gomez-Alday, 2000. Taphonomy
of the Late Cretaceous dinosaur-bearing beds of the La�o Quarry (Iberian Peninsula).
Palaeogeography, Palaeoclimatology, Palaeoecology. 157, 247-275.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod
dinosaurs from the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta
Palaeontologica Polonica. 60(3), 611-626.
Isasmendi, Torices, Canudo and Pereda-Suberbiola, 2020.
Paleobiodiversity of theropod dinosaurs from the Upper Cretaceous La�o
site, northern Iberian peninsula. The Society
of Vertebrate Paleontology 80th
Annual Meeting, Conference Program. 186-187.
P? sp. (Torices, Barroso-Barcenilla, Cambra-Moo, Perez and Serrano,
2011)
Late Campanian-Early Maastrichtian, Late Cretaceous
Villalba de la Sierra Formation, Spain
Material- teeth
Comments- Torices et al. (2011) mention cf. Paronychodon.
Reference- Torices, Barroso-Barcenilla, Cambra-Moo, Perez and Serrano,
2011. Vertebrate microfossil analysis in the palaeontological site of 'Lo Hueco'
(Upper Cretaceous, Cuenca, Spain). Journal of Vertebrate Paleontology. Program
and Abstracts 2011, 205.
P. sp. (Lopez-Martinez, Canudo, Ardevol, Pereda-Suberbiola, Orue-Etxebarria,
Cuenca-Bescos, Ruiz-Omenaca, Muerlaga and Feist, 2001)
Late Maastrichtian, Late Cretaceous
Blasi 2B, Tremp (=Arenisca) Formation, Spain
Material- (MPZ98/76) tooth (2.7x1.5x.6 mm)
(MPZ98/77) tooth (2.8x1.4x.6 mm)
(MPZ98/78) tooth (2.2x1.2x.8 mm)
Comments- These are strongly recurved and have three labial ridges. They
were described as unserrated. While MPZ 98-76 is listed as having 15.97 distal
serrations per mm by Lopez-Martinez et al. (2001), Torices et al. (2015) list
it as serrationless and provide a figure demonstrating this is so. They were
listed as cf. Euronychodon sp. by Lopez-Martinez et al., and cf. Paronychodon
sp. by Torices et al..
References- Canudo, Lopez Martinez and Ruiz Omenaca, 2001. Los dinosaurios
del Maastrichtiense Superior (Cretacico Superior) del pirineo de Huesca (Espana).
Actas de Las I Jornadas Internacionales sobre Paleontologia de Dinosaurios y
su entorno. 319-328.
Lopez-Martinez, Canudo, Ardevol, Pereda-Suberbiola, Orue-Etxebarria, Cuenca-Bescos,
Ruiz-Omenaca, Muerlaga and Feist, 2001. New dinosaur sites correlated with Upper
Maastrichtian pelagic deposits in the Spanish Pyrenees: Implications for the
dinosaur extinction pattern in Europe. Cretaceous Research. 22, 41-61.
Torices, Currie, Canudo and Pereda-Suberbiola, 2015. Theropod dinosaurs from
the Upper Cretaceous of the South Pyrenees Basin of Spain. Acta Palaeontologica
Polonica. 60(3), 611-626.
P. sp. (Garcia, Duffaud, Feist, Marandat, Tambareau, Villatte
and Sige, 2000)
Turonian-Maastrichtian, Late Cretaceous
La Nueve, Aix, France
Material- (LNE-D01) tooth
Reference- Garcia, Duffaud, Feist, Marandat, Tambareau, Villatte and
Sige, 2000. La neuve, gisement a plantes, invertebres et vertebres du Begudien
(Senonien superieur continental) du bassin d’Aix-en-Provence. Geodiversitas.
22(3), 326-348.
P? sp. (Sige, Buscalioni, Duffaud, Gayet, Orth, Rage and Sanz,
1997)
Campanian, Late Cretaceous
unnamed unit, Champ-Garimond, Gard, France
Material- teeth
Comments- These differ from most Paronychodon specimens in lacking
carinae. They were assigned to Paronychodon by Sige et al. (1997).
Reference- Sige, Buscalioni, Duffaud, Gayet, Orth, Rage and Sanz, 1997.
Etat des donn�es sur le gisement Cr�tac� sup�rieur
continental de Champ-Garimond (Gard, Sud de la France). Munchner Geowiss.. 34,
111-130.
P. sp. nov. (Csiki and Grigorescu, 1998)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Material- (FGGUB R.1431) type A tooth (5.1 mm) (Csiki and Grigorescu, 1998)
(IRSNB coll.) type A tooth (~2.8x~1.1x? mm) (Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke,
2002)
(IRSNB coll.) teeth (Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke, 2002)
Comments- FGGUB R.1431 differs from most Paronychodon
specimens in having only two lingual grooves and lacking labial
grooves. A tooth morphotype photographed by Codrea et al. (2002) is
elongate and strongly recurved, "lack serrations on both their mesial
and distal carinae, but, contrary to the Paronychodon morphotype, they are not very asymmetrical and their enamel is not ornamented." Csiki and Grigorescu refer FGGUB R.1431 to cf. Euronychodon,
while Codrea et al. refer to their teeth as Euronychodon morphotype.
References- Csiki and Grigorescu, 1998. Small theropods from the Late
Cretaceous of the Hateg Basin (Western Romania) - an unexpected diversity at
the top of the food chain. Oryctos. 1, 87-104.
Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke, 2002. Dinosaur
egg nests, mammals and other vertebrates from a new Maastrichtian site of the
Hateg Basin (Romania). Comptes Rendus Palevol. 1(3), 173-180.
P. sp. (Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van
Itterbeecke, 2002)
Late Maastrichtian, Late Cretaceous
Sinpetru Beds, Romania
Material- (IRSNB coll.) partial type A tooth (~4.5x~2.4x? mm)
(IRSNB coll.) teeth
Comments-
Codea et al. (2002) describe an assemblage collected in 2001. They
state "a number of teeth may be identified as theropods on the basis of
overall shape, but lack serrations entirely. Some of them closely
resemble the Paronychodon morphotype ... in being flat on one side and covered with coarse longitudinal ridges."
Reference- Codrea, Smith, Dica, Folie, Garcia, Godefroit and Van Itterbeecke,
2002. Dinosaur egg nests, mammals and other vertebrates from a new Maastrichtian
site of the Hateg Basin (Romania). Comptes Rendus Palevol. 1(3), 173-180.
P. asiaticus (Nessov, 1995)
Sues and Averianov, 2013
= Euronychodon asiaticus Nessov, 1995
= "Plesiosaurodon" sp. Nessov vide Sues and Averianov, 2013
Mid-Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Holotype- (CCMGE N 9/12454) type A tooth
Paratypes- (CCMGE coll.) six teeth
Referred- (ZIN PH 301/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1052/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1053/16) type A tooth (Sues and Averianov, 2013)
(ZIN PH 1054/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1055/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1056/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1057/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1058/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1059/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1060/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1061/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1062/16) premaxillary tooth (Sues and Averianov, 2013)
(ZIN PH 1063/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1064/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1065/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1066/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1067/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1068/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1069/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1070/16) premaxillary tooth (Sues and Averianov, 2013)
(ZIN PH 1072/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1147/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1149/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1150/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1151/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1152/16) type A tooth (Sues and Averianov, 2013)
(ZIN PH 1345/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1833/16) type A tooth (Sues and Averianov, 2013)
Middle Albian-Early Cenomanian, Early-Late Cretaceous
Khodzhakul Formation, Uzbekistan
(ZIN PH 1071/16) type B tooth (Sues and Averianov, 2013)
(ZIN PH 1220/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1221/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1228/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1229/16) tooth (Sues and Averianov, 2013)
(ZIN PH 1230/16) tooth (Sues and Averianov, 2013)
Diagnosis- (after Sues and Averianov, 2013) can differ from P. lacustris
in having a higher number of labial ridges (up to 17 versus up to 14).
Comments- These teeth were first described as?Plesiosauria (Nessov, 1985)
and Paronychodon cf. lacustris (Nessov, 1986) before being described
as a new species of Euronychodon by Nessov (1995). Nessov (1995) distinguished
it from Euronychodon portucalensis based on the presence of numerous
labial grooves, but the latter are seen in a paratype of E. portucalensis.
Sues and Averianov (2013) redescribed the teeth as Paronychodon asiaticus,
and noted some specimens were labelled "Plesiosaurodon" by Nessov.
Averianov (2007) first announced Paronychodon sp. from the Khodzhakul
Formation, which were also described by Sues and Averianov and referred to P.
asiaticus.
References- Nessov, 1985. [New mammals of the Cretaceous of the Kyzylkum].
Vyestnik Lyeningradskogo univyersityeta, Biologiya. 17, 8-18.
Nessov, 1986. [The first discovery of the Late Cretaceous bird Ichthyornis
in the Old World and some other bones of birds from the Cretaceous and Paleogene
of Middle Asia]. Trudy Zoologichyeskogo Instituta AN SSSR. 147, 31-38.
Nessov, 1995. Dinozavri severnoi Yevrazii: Novye dannye o sostave kompleksov,
ekologii i paleobiogeografii. Institute for Scientific Research on the Earth's
Crust, St. Petersburg State University, St. Petersburg. 156 pp.
Averianov, 2007. Theropod dinosaurs from Late Cretaceous deposits in the northeastern
Aral Sea region, Kazakhstan. Cretaceous Research. 28(3), 532-544.
Sues and Averianov, 2013. Enigmatic teeth of small theropod dinosaurs from the
Upper Cretaceous (Cenomanian-Turonian) of Uzbekistan. Canadian Journal of Earth
Sciences. 50, 306-314.
P? sp. (Averianov, Leshchinskiy, Skutschas, Fayngertz and Rezvyi,
2004)
Aptian-Albian, Early Cretaceous
Ilek (=Shestakovo) Formation, Russia
Material- teeth
Comments- Referred to cf. Paronychodon.
Reference- Averianov, Leshchinskiy, Skutschas, Fayngertz and Rezvyi,
2004. Dinosaurs from the Early Cretaceous Ilek Formations in West Siberia, Russia.
2nd EAVP Meeting. July 19-24, 2004. Brno, Czech Republic. Abstracts of papers
and posters with program, Excursion Guidebook. pg 6.
unnamed possible Troodontidae (Antunes and Sigogneau-Russell, 1992)
Maastrichtian, Late Cretaceous
Unnamed unit, Distrito do Coimbra, Portugal
Material- (AV 7) fragmentary tooth
(TV 43) fragmentary tooth
(TV 52) fragmentary tooth
Reference- Antunes and Sigogneau-Russell, 1992. La Faune de Petits Dinosaures
du Cretace Terminal Portugais. Comunica��es dos Serci�os geol�gicos de Portugal. 78(1), 49-62.
unnamed troodontid (Averianov, 2016)
Santonian, Late Cretaceous
Bostobe Formation, Kazakhstan
Material- (ZIN PH 45/49) partial frontal
Reference- Averianov, 2016 (online 2015). Frontal bones of non-avian theropod
dinosaurs from the Upper Cretaceous (Santonian-?Campanian) Bostobe
Formation of the northeastern Aral Sea region, Kazakhstan. Canadian
Journal of Earth Sciences. 53(2). 168-175.
undescribed troodontid (Suzuki and Watabe, 2000)
Cenomanian-Santonian, Late Cretaceous
Bayshin Tsav, Baynshire Formation, Mongolia
Material- (IGM coll.; 950728
BTs-Nar or 950728 BTs Theropod) partial skeleton including six dorsal
vertebrae, several dorsal ribs, gastralia, seven sacral or proximal
caudal vertebrae, manual
ungual I, phalanx II-2, manual ungual II, phalanx III-1, phalanx III-2,
manual claw sheath, manual phalanges, manual ungual, manual claw
sheath, pubis, incomplete femur, tibia, fibula, pedal phalanges, pedal
ungual, fragments
Comments- This specimen was
found on July 28 1995, listed as "A skeleton of small theropod
discovered near Bayshin Tsav IV" and "Small theropod skeleton" by
Suzuki and Watabe (2000). They photographed it as "manus of small
theropod at Bayshin Tsav". Tsogtbaatar (2004) states it was prepared in 2000 (as
950728 BTs-Nar) and identified as a troodontid, being stored at the IGM.
Reference- Suzuki and Watabe, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1995. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 45-57.
Tsogtbaatar, 2004. Fossil specimens prepared in Mongolian Paleontological Center 1993-2001. Hayashibara Museum of Natural
Sciences Research Bulletin. 2, 123-128.
undescribed possible troodontid (Norton, DML 2000)
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadochta Formation, Mongolia
Material- (IGM 97/155) specimen including manual phalanx III-3
Comments-
Norton (DML, 2000) noted this specimen was on display at the AMNH
Fighting Dinosaurs exhibit. Discovered in 1997 at Ukhaa Tolgod, it was
said to be "unpublished but noted as having "raptor like " traits".
Prieto-Marquez et al. (2011) state "troodontid specimen MAE 97-155" has
a manual phalanx III-3 identical to halszkaraptorine ISMD-VP09.
References- Norton, DML 2000. https://web.archive.org/web/20210603191138/http://dml.cmnh.org/2000Jun/msg00082.html
Prieto-Marquez, Bolortsetseg and Horner, 2011. A diminutive
deinonychosaur (Dinosauria: Theropoda) from the Early Cretaceous of
Oosh (Ovorkhangai, Mongolia). Alcheringa. 1-20.
unnamed possible troodontid (Dong, 1997)
Early Albian, Early Cretaceous
Upper Gray Beds of the Zhonggou Formation, Gansu, China
Material- (IVPP V11119) two teeth, two caudal vertebrae, partial tibia,
distal tarsal, incomplete metatarsal II, phalanx II-1 (17 mm), phalanx II-2
(13 mm), pedal ungual II (9 mm), distal metatarsal III, phalanx III-1, partial
metatarsal IV, phalanx IV-1
Comments- Dong (1997) referred this to Sinornithoides sp. nov.
based on undescribed distal tarsal similarities, subequal size, and the plesiomorphically
limited proximal extent of the distal articular surface on metatarsal III. However,
he notes the less robust metatarsal IV differs. This suggests the specimen was
outside the Sinornithoides+Troodon
clade and should not be referred to the former genus. Hartman et
al. (2019) recovered this as a coelurosaur outside several clades
including Troodontidae+Dromaeosauridae.
References-
Dong, 1997. On small theropods from Mazongshan area, Gansu province,
China. In Dong (ed.). Sino-Japanese Silk Road Dinosaur Expedition.
China Ocean Press. 13-18.
You, Morschhauser, Li and Dodson, 2018. Introducing the Mazongshan
dinosaur fauna. Journal of Vertebrate Paleontology. 38(supp. 1), 1-11.
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 troodontid (Kobayashi, Lu, Lee, Xu and Zhang, 2008)
Late Cretaceous
Qiupa Formation, Henan, China
Material- skeleton
Comments- Kobayashi et al. (2008) state "A new locality, yielded Luanchuanraptor,
in Tantou Basin in Luanchuan County exposes the Upper Cretaceous Qiupa
Formation and is rich in dinosaur eggs and bones, such as undescribed
skeletons of an ... troodontid." Note this is not the
subsequently described Xixiasaurus,
which is from the Majiacun Formation in Xixia County near Songgou village, not
the Qiupa Formation in Luanchuan County near Qiupa village.
Reference- 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.
Troodontinae sensu Hendrickx,
Mateus, Ara�jo and Choiniere, 2019
Definition- (Sinovenator changii + Troodon formosus)
Reference- 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.
Sinovenatorinae Shen, Lu, Liu, Kundrat,
Brusatte and Gao, 2017
Definition- (Sinovenator changii <- Jinfengopteryx elegans, Troodon formosus, Passer domesticus) (Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019)
Other definitions- (Sinovenator changii <- Troodon formosus,
Saurornithoides mongoliensis, Anchiornis huxleyi, Archaeopteryx
lithographica, Gallus gallus, Unenlagia comahuensis, Dromaeosaurus
albertensis) (Shen, Lu, Liu, Kundrat, Brusatte and Gao, 2017)
Comments- This clade was recovered by Shen et al. (2017) using Brusatte's
version of the TWiG analysis. In their topology it includes Sinovenator,
Mei, Daliansaurus and Sinusonasus. Hartman et al. (2019) redefined this clade to exclude Jinfengopteryx, previously given its own subfamily.
References- Shen, Lu, Liu, Kundrat, Brusatte and Gao, 2017. A new troodontid
dinosaur from the Lower Cretaceous Yixian Formation of Liaoning Province, China.
Acta Geologica Sinica. 91(3), 763-780.
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
Mei Xu and Norell, 2004
M. long Xu and Norell, 2004
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V12733; ?= IVPP V12744 of Xu et al. 2014) (530 mm; subadult)
incomplete skull (53 mm), sclerotic plates, incomplete mandible (56 mm), ten
cervical vertebrae (67 mm), dorsal vertebrae (92 mm), dorsal ribs, gastralia,
sacrum (30 mm), caudal vertebrae (257 mm), chevrons, scapulae (45 mm), coracoids
(11 mm), furcula, sternal rib fragments(?), humeri (42 mm), radii (39 mm), ulnae
(42 mm), semilunate carpal, (manus 67 mm) metacarpal I, phalanx I-1, manual
ungual I, metacarpal II, phalanx II-1, phalanx II-2, manual ungual II, phalanx
III-2, phalanx III-3, manual ungual III, ilia, pubes, ischium, femora (81 mm),
tibiae (106 mm), fibula, astragali, distal tarsals, metatarsal I, phalanx I-1,
pedal ungual I, metatarsal II (51 mm), phalanx II-1, phalanx II-2, pedal ungual
II, metatarsal III (58 mm), phalanx III-1, phalanx III-2, phalanx III-3, pedal
ungual III, metatarsal IV (55 mm), phalanx IV-1, phalanx IV-2, phalanx IV-3,
phalanx IV-4, pedal ungual IV, metatarsal V
Referred- (DNHM D2514) (~325 mm; ~420 g; adult) incomplete skull (~49
mm), mandible, fragmentary fourth cervical vertebra, fifth cervical vertebra,
sixth cervical vertebra, seventh cervical vertebra, eighth cervical vertebra,
ninth cervical vertebra, tenth cervical vertebra, incomplete eleventh cervical
vertebra, cervical ribs, first dorsal neural arch, second dorsal neural arch,
third dorsal neural arch, fourth dorsal neural arch, fifth dorsal neural arch,
incomplete eighth dorsal vertebra, incomplete ninth dorsal vertebra (5.8 mm),
incomplete tenth dorsal vertebra (6 mm), incomplete eleventh dorsal vertebra
(5.9 mm), incomplete twelfth dorsal vertebra (5.9 mm), incomplete thirteenth
dorsal vertebra, dorsal rib fragments, sacrum (22.2 mm), first caudal vertebra,
second caudal vertebra, third caudal vertebra, fourth caudal vertebra (4 mm),
fifth caudal vertebra (4 mm), sixth caudal vertebra (4.3 mm), seventh caudal
vertebra (4.9 mm), eighth caudal vertebra (5.5 mm), ninth caudal vertebra (8
mm), tenth caudal vertebra (10.3 mm), eleventh caudal vertebra (11.1 mm), twelfth
caudal vertebra (11 mm), thirteenth caudal vertebra (11.2 mm), fourteenth caudal
vertebra (10.9 mm), fifteenth caudal vertebra (10.9 mm), sixteenth caudal vertebra
(10.6 mm), seventeenth caudal vertebra (10.6 mm), eighteenth caudal vertebra,
eleven chevrons, scapulae (36, 40 mm), coracoid, humeri (36 mm), radii (one
incomplete), ulnae (one incomplete; ~34 mm), metacarpal I (4 mm), phalanx I-1
(14 mm), manual ungual I (7.3 mm), metacarpal II (13 mm), phalanx II-1 (7.8
mm), phalanx II-2 (13 mm), manual ungual II, metacarpal III (14.3 mm), phalanx
III-1 (8.1 mm), phalanx III-2 (8.1 mm), phalanx III-3 (10.7 mm), ilia (34.4
mm), proximal pubes, femora (~65 mm), tibiotarsi (~86.1 mm), fibulae, distal
metatarsal II (~42.5 mm), phalanx II-2, pedal ungual II, distal metatarsal III
(~49 mm), phalanx III-1 (12.8 mm), phalanx III-3, metatarsals IV (one distal;
45.7 mm), phalanx IV-1 (8.4 mm), phalanx IV-2 (6.4 mm), phalanx IV-3 (5.9 mm),
phalanx IV-4 (5.2 mm), metatarsal V (11 mm) (Gao et al., 2012)
Diagnosis- (after Xu and Norell, 2004) extremely large external nares
extending posteriorly over one half of the maxillary tooth row; closely packed
middle maxillary teeth; maxillary tooth row extending posteriorly to the level
of the preorbital bar; robust U-shaped furcula; most proximal end of the pubic
shaft is significantly compressed anteroposteriorly, and extends laterally just
ventral to the articulation with the ilium; lateral process on distal tarsal
IV.
(after Gao et al., 2012) posterior sacrum extremely wide with elongate fourth
and fifth transverse processes; ilium strongly sigmoid in dorsal view, with
a stronger lateral curve than in Velociraptor or Anchiornis.
Comments- The holotype was discovered prior to April 2004.
References- Xu and Norell, 2004. A new troodontid dinosaur from China
with avian-like sleeping posture. Nature. 431, 838-841.
Gao, Morschhauser, Varricchio, Liu and Zhao, 2012. A second soundly sleeping
dragon: New anatomical details of the Chinese troodontid Mei long with
implications for phylogeny and taphonomy. PLoS ONE. 7(9), e45203.
Xu, Han and Zhao, 2014. Homologies and homeotic transformation of the theropod
'semilunate' carpal. Nature Scientific Reports. 4, 6042.
Xiaotingia Xu, You, Du and Han,
2011
X. zhengi Xu, You, Du and Han, 2011
Oxfordian, Late Jurassic
Tiaojishan Formation?, Liaoning, China
Holotype- (STM 27-2) (820 g adult) skull (~61 mm to quadrate), mandible
(62 mm), (cervical series 80 mm) ten cervical vertebrae, cervical ribs, (dorsal
series 118 mm) thirteen dorsal vertebrae, dorsal ribs, gastralia, synsacrum,
three proximal caudal vertebrae, scapulae (55 mm), partial coracoid, furcula
(42 mm across), humeri (71 mm), radii (63 mm), ulnae (65 mm), semilunate carpal,
metacarpal I (10 mm), phalanges I-1 (21 mm), manual unguals I (14 mm), metacarpals
II (24 mm), phalanges II-1 (15 mm), phalanges II-2 (25 mm), manual unguals II
(14 mm), metacarpals III (24 mm), phalanges III-1 (8 mm), phalanges III-2 (4
mm), phalanges III-3 (15 mm), manual ungual III (11 mm), manual claw sheaths,
ilia (52 mm), partial pubis, incomplete ischium (~28 mm), incomplete femora
(~84 mm), incomplete tibia, incomplete fibula, metatarsal I (9 mm), phalanx
I-1 (6 mm), pedal ungual I (6 mm), distal metatarsal II, phalanx II-1 (12 mm),
phalanx II-2 (9 mm), pedal ungual II (13 mm), distal metatarsal III, phalanx
III-1 (17 mm), phalanx III-2 (13 mm), phalanx III-3, pedal ungual III, distal
metatarsal IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal
ungual IV, pedal claw sheaths, feathers
Diagnosis- (after Xu et al., 2011) maxillary posterior ramus has depth
at mid-length exceeding that of dentary; surangular has little lateral exposure
and forms a wide, flat dorsal surface over posterior part of mandible; extremely
large surangular foramen >6% of mandibular length; posterior end of mandible
blunt and dorsoventrally expanded; proximalmost caudal centra less than half
as long as posterior dorsal centra; metacarpal III more robust than metacarpals
I and II; manual phalanx II-2 longer than metacarpal II.
Comments- As the holotype was acquired from a dealer with no quarry information (prior to November 2010),
Xiaotingia may instead be from the Yixian Formation which outcrops in
the same area. Xu et al. (2011) used a version of Senter's TWiG matrix to place
Xiaotingia in Archaeopterygidae with Archaeopteryx and Anchiornis,
with the family in basal Deinonychosauria. More recently, Hartman et
al. (2019) used a far more extensive TWiG analysis to recover it as a
sinovenatorine troodontid. As they state, "troodontid characters
present in Xiaotingia
but not anchiornithines include distally positioned obturator process
(183:2), and characters shared with sinovenatorines include large
posterior surangular foramen (80:2), capital groove in humerus (458:1),
metacarpal III extending distally past metacarpal II (640:1), laterally
ridged ischium (182:2), and enlarged pedal ungual II (224:1). Forcing Xiaotingia
into Archaeopterygidae requires nine more steps, which strongly
suggests it is not a member considering we included all of the TWiG
data originally used to place it there. Alternative placements as a
non-anchiornithine avialan (Lee et al., 2014), a dromaeosaurid (Senter
et al., 2012), and a scansoriopterygid relative (Lef�vre et al., 2017)
are eight, five, and 26 more steps, respectively."
References- Xu, You, Du and Han, 2011. An Archaeopteryx-like theropod
from China and the origin of Avialae. Nature. 475, 465-470.
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.
Lee, Cau, Naish and Dyke, 2014. Sustained miniaturization and anatomical innovation
in the dinosaurian ancestors of birds. Science. 345(6196), 562-566.
Lefevre, Cau, Cincotta, Hu, Chinsamy, Escuillie and Godefroit, 2017. A
new Jurassic theropod from China documents a transitional step in the
macrostructure of feathers. The Science of Nature. 104:74.
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
Jianianhualong Xu, Currie, Pittman, Xing, Meng, Lu, Hu and Yu, 2017
J. tengi Xu, Currie, Pittman, Xing, Meng, Lu, Hu and Yu, 2017
Late Barremian-Early Aptian, Early Cretaceous
Yixian Formation, Liaoning, China
Holotype- (DLXH 1218) (~1.12 m; ~2.4 kg; adult) skull, sclerotic
plates, mandibles, hyoids, cervical series (~160 mm) mostly fused to
cervical ribs (except c3 and c6),
dorsal vertebrae (~170 mm), nine pairs of dorsal ribs, ~12 rows of
gastralia, sacrum,
first to twenty-third caudal vertebrae (~540 mm), chevrons, partial
scapulae, coracoid, furcula,
humeri, radii, ulnae, scapholunare, semilunate carpal, distal carpal III,
metacarpals I, phalanges I-1, manual unguals I,
metacarpals II, phalanges II-1, phalanges II-2, manual unguals II,
metacarpals III, phalanges III-1, phalanges III-2, phalanges III-3,
manual unguals III, manual claw sheaths, ilia (one fragmentary), pubes,
ischia, femora, distal tibiae,
distal fibula, fragmentary proximal tarsals, metatarsal I, phalanges
I-1, pedal unguals
I, distal tarsals III, distal tarsals IV, metatarsals II, phalanx II-1,
phalanx II-2, pedal ungual II,
metatarsals III, metatarsals IV, phalanx IV-1, pedal phalangeal
fragments, metatarsals V, body feathers (30-~75 mm),
remiges, retrices (~120 mm)
Diagnosis- (after Xu et al.,
2017) maxillary anterior ramus triangular and relatively tall;
maxillary ascending process extending posterodorsally at a high angle
(an angle of ~45 degrees to maxillary ventral margin); long manual
phalanx I-1 (slightly shorter than metacarpal II) with prominent
proximoventral heel, large groove along the medial surface of more than
proximal half; highly elongated manual phalanx II-2 (slightly longer
than metacarpal II); ilium with slightly concave dorsal margin in
lateral view; metatarsal IV without prominent ventral flange.
Comments- The holotype was discovered prior to Novemver 2016. Xu et al. recovered it closer to troodontines than Sinovenator
using Senter's version of the TWiG matrix, but Hartman et al. (2019)
incorporating more TWiG data found it to be a sinovenatorine
instead. Only a single step moves it closer to troodontines
however.
References- Xu, Currie,
Pittman, Xing, Meng, Lu, Hu and Yu, 2017. Mosaic evolution in an
asymmetrically feathered troodontid dinosaur with transitional
features. Nature Communications. 8:14972.
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 troodontid (Tsuihiji, Watabe, Barsbold, Suzuki and Tsogtbaatar, 2009)
Barremian-Albian?, Early Cretaceous
Khamaryn Ar, Mongolia
Material- (IGM 100/140; 080924 KhA OTGON) thirteen distal caudal vertebrae, five distal chevrons,
distal radius, distal ulna, scapholunare, semilunate carpal, metacarpal I (13.9 mm),
phalanx I-1 (30.7 mm), manual ungual I (20.4 mm), metacarpals II (one incomplete,
one distal), phalanges II-1 (18.8 mm), phalanges II-2 (30.1, 30.7 mm), manual
unguals II (22.9, 23.2 mm), metacarpals III (one incomplete, one distal), phalanges
III-1 (7.2 mm), phalanges III-2 (9.2 mm), phalanges III-3 (19.1 mm), manual
unguals III (18.5 mm), manual claw sheaths, phalanx I-1 (11.5 mm), pedal ungual
I (12 mm), partial metatarsal II, phalanx II-1 (20.4 mm), phalanx II-2 (14.9
mm), incomplete pedal ungual II, partial metastarsal III, phalanx III-1 (25.6
mm), phalanx III-2 (18 mm), phalanx III-3 (16.9 mm), incomplete pedal ungual
III, incomplete metatarsal IV, phalanx IV-1 (16.8 mm), phalanx IV-2 (15.1 mm),
phalanx IV-4 (13.7 mm), incomplete pedal ungual IV
Comments- IGM 100/140 was discovered on September 23-25
2008 (Tsubamoto et al., 2010), given the field number 080924 KhA OTGON
and photographed and mentioned by Tsubamoto et al. as "Dromaeosauridae
or Troodontidae" (fig. 9B). Tsuihiji et al. (2009) mention it in
an abstract, but it wasn't fully described until Tsuihiji et al.
(2015). This specimen was added to a version of Senter's TWiG
analysis by Tsuihiji et al. (2015) and found to be more derived than Byronosaurus. Hartman et al. (2019) included it in their more extensive TWiG analysis and recovered it as a sinovenatorine sister to Jianianhualong, but only a single step moves it closer to troodontines.
References- Tsuihiji, Watabe, Barsbold, Suzuki and
Tsogtbaatar, 2009. New material of a troodontid theropod (Dinosauria:
Saurischia) from the Lower Cretaceous of Mongolia. 4th International
Symposium of IGCP 507, Abstracts. 59.
Tsubamoto, Saneyoshi, Tsogtbaatar, Chinzorig, Khatanbaatar, Mainbayar
and Suzuki, 2010. Report of the HMNS-MPC Joint Paleontological
Expedition in 2008. Hayashibara Museum of Natural Sciences Research
Bulletin. 3, 29-39.
Tsuihiji, Barsbold, Watabe, Tsogtbaatar, Suzuki and Hattori,
2015. New material of a troodontid theropod (Dinosauria: Saurischia) from the
Lower Cretaceous of Mongolia. Historical Biology. 28(1-2), 128-138.
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
Sinovenator Xu, Norell, Wang,
Makovicky and Wu, 2002
S. changii Xu, Norell, Wang, Makovicky and Wu, 2002
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V12615) (adult) incomplete skull (~90 mm), dentary, supradentary,
surangulars, anterior cervical vertebra, mid cervical vertebra, mid cervical
vertebra, posterior cervical vertebra, third dorsal vertebra (9.8 mm), fourth
dorsal vertebra (9.8 mm), fifth dorsal vertebra (10 mm), sixth dorsal vertebra
(10.2 mm), seventh dorsal vertebra (10.9 mm), eighth dorsal vertebra (11.2 mm),
ninth dorsal vertebra (11.6 mm), tenth dorsal vertebra (11.7 mm), eleventh dorsal
vertebra (11.3 mm), twelfth dorsal vertebra, fragmentary thirteenth dorsal vertebra,
two partial dorsal ribs, sacrum, first caudal vertebra (7.2 mm), second caudal
vertebra (8 mm), third caudal vertebra (10 mm), fourth caudal vertebra (10 mm),
fifth caudal vertebra (10.2 mm), sixth caudal vertebra (~10.2 mm), incomplete
scapula (~66 mm), coracoid, distal humerus, partial radius, partial ulna, metacarpal,
manual ungual, ilium (~60 mm), pubes (97, 94 mm), ischium (40 mm), femur (117.5
mm), tibia (152.9 mm), incomplete fibulae, astragalus, metatarsal I (~19 mm),
metatarsal II (77 mm), phalanx II-2, pedal ungual II, metatarsal III (86.2 mm),
metatarsals IV (82.6, 83.4 mm), partial metatarsal V
Paratype- (IVPP V12583) (subadult) sacrum, first caudal vertebra (7 mm),
second caudal vertebra (8.2 mm), third caudal vertebra (8 mm), fourth caudal
vertebra (8 mm), fifth caudal vertebra (8 mm), sixth caudal vertebra (8 mm),
seventh caudal vertebra (8.6 mm), eighth caudal vertebra (10 mm), ninth caudal
vertebra (10.9 mm), tenth caudal vertebra (14.5 mm), eleventh caudal vertebra
(16.1 mm), twelfth caudal vertebra (19 mm), thirteenth caudal vertebra, fourteenth
caudal vertebra (18 mm), fifteenth caudal vertebra (18 mm), sixteenth caudal
vertebra (20 mm), seventeenth caudal vertebra (19 mm), eighteenth caudal vertebra
(18 mm), nineteenth caudal vertebra (17.9 mm), twentieth caudal vertebra (17.5
mm), twenty-first caudal vertebra (17.2 mm), twenty-second caudal vertebra (17.9
mm), twenty-third caudal vertebra (19 mm), twenty-fourth caudal vertebra (19
mm), twenty-fifth caudal vertebra (18.5 mm), twenty-sixth caudal vertebra (17.5
mm), proximal scapulae, fragmentary coracoids, humeri (71 mm), radius (~87 mm),
ulna (91 mm), semilunate carpal, distal carpal III, metacarpal I (12 mm), phalanx
I-1 (28 mm), metacarpal II (28.5 mm), two manual phalanges, pubis (87 mm), ischium
(42.5 mm), femur (105 mm), tibiotarsi (148, 146 mm), fibulae, metatarsal II
(79 mm), phalanges II-1 (15, 15 mm), phalanges II-2 (10, 10 mm), metatarsal
III (91.7 mm), phalanges III-1 (21.5, 22 mm), phalanx III-2 (14.6 mm), phalanx
III-3 (~13.5 mm), metatarsal IV (88 mm), phalanges IV-1 (15, 14.7 mm), phalanges
IV-2 (12, 12.8 mm), phalanx IV-3 (10.2 mm), phalanx IV-4 (10.2 mm), partial
metatarsal V
Referred- (IVPP V14009) (adult) specimen including semilunate carpal,
metacarpal I, metacarpal II and metacarpal III (Xu et al., 2014)
(IVPP V14322) mandible, cervical vertebrae, dorsal vertebrae, dorsal ribs, proximal
humerus, manual phalanges, manual unguals, pubis, ischium, femora, tibiae, partial
fibula, astragalus, calcaneum, metatarsal II, phalanx II-1, phalanx II-2, pedal
ungual II, metatarsal III, phalanx III-1, phalanx III-2, metatarsal IV, phalanx
IV-1, phalanx IV-2 (Manabe, 2005)
(IVPP coll.) femur (206 mm), tibia (147 mm), metatarsal II, phalanx II-1, phalanx
II-2, metatarsal III (92 mm), phalanx III-1, phalanx III-2, phalanx III-3 metatarsal
IV, phalanx IV-1, phalanx IV-2, phalanx IV-3, phalanx IV-4, pedal ungual IV
(White, 2009)
(PMOL-AD00102) (adult) posterior skull (~161 mm), stapes, posterior
mandibles, proatlases, atlantal intercentrum, atlantal neural arches,
atlantal ribs, axis, axial ribs, incomplete third cervical vertebra,
incomplete third cervical vertebra, incomplete third cervical vertebra,
sixth cervical prezygapophyses, partial cervical ribs (Yin et al., 2018)
? skull, cervical vertebrae, dorsal ribs, sacral vertebrae, caudal vertebrae,
scapula, humerus, radius, ulna, manus, ilia, femur, tibiae, fibula, pes (Fossil
Mall, online 2015)
Diagnosis- (after Xu et al., 2002) straight and vertical anterior margin
of antorbital fenestra (also in Sinusonasus);
surangular T-shaped in cross-section; prominent lateral cnemial crest continuous
with the fibular crest.
(after Yin et al., 2018) well-developed medial shelf on jugal; slender
bar in parasphenoid recess; lateral groove on pterygoid flange of
ectopterygoid; lateral surface of anterior cervical vertebrae bearing
two pneumatic foramina.
Other diagnoses- Xu et al.
(2002) proposed a frontal with a vertical lamina bordering the lacrimal
as diagnostic, but while present in the holotype this is missing in
PMOL-AD00102 (Yin et al., 2018).
Comments- Creisler (DML 2002) noted Sinovenator changii was named
after a woman, so suggested it be emended to S. changiae. Similarly, Xu (2002) states Li indicated it should be S. changae,
which is used in that thesis. However, the Fourth Edition of the
ICZN no longer requires emendations based on this reasoning (Article
31.1.3- "The original spelling of a name formed under Articles 31.1.1
and 31.1.2 is to be preserved [Art. 32.2] unless it is incorrect [Arts.
32.3, 32.4]"), with Article 32.3 saying "The correct original spelling
of a name is to be preserved unaltered, except where it is mandatory to
change the suffix or the gender ending under Article 34" where Article
34 involves matching genus and species genders, and Article 32.4 saying
"An original spelling is an "incorrect original spelling" if it must be
corrected as required in Article 32.5" and Article 32.5 saying "
Incorrect transliteration or latinization ... are not to be considered
inadvertent errors". If Sinovenator
"changiae" was published by Haubold (2003) while S.
"changae" was published by Long and Schouten (2008), but ICZN Article
32.2.3 states both are available names with their own authorships,
though objective junior synonyms of S. changii.
The holotype and paratype were discovered in Summer 2001 (Xu, 2002) and named and briefly described
by Xu et al. (2002). A more detailed description is in Xu's (2002) thesis, but
has not yet been published.
Senter (2007) mentioned an undescribed specimen whose photo was available on
www.dinosaur.net.cn (2004). This is IVPP V14322, and is illustrated in Manabe (2005).
White (2009) describes and illustrates the pes of an unregistered specimen at
the IVPP as Sinovenator sp.
An undescribed basically complete specimen is referred to Sinovenator
on the Fossil Mall website (2015), but may be at least partially faked due to its apparent
lumbar region and non-maniraptoran forelimb position.
Yin et al. (2018) described a posterior skull of a new specimen
(PMOL-AD00102) that differs from the smaller holotype in a few
characters- frontal without vertical lamina bordering the lacrimal;
presence of a septum between the basal tubera; presence of a
basisphenoid recess; deep sagittal crest; basipterygoid process
with a blunt distal end. They considered these possibly
ontogenetic or taphonomic.
Senter et al. (2004) corrected Xu et al.'s description of several characters,
finding the teeth to be unserrated, the antorbital fossa to lack a prominent
rim, the anterior and posterior dentary teeth to be subequal in size and density,
and the fourth metatarsal to be subequal in diameter to the second. Senter (2007)
later corrected his account of dentary tooth density, agreeing with Xu et al.
that the anterior teeth were more densely packed.
References- Creisler, DML 2002. https://web.archive.org/web/20210121093206/http://dml.cmnh.org/2002Feb/msg00579.html
Xu, 2002. Deinonychosaurian fossils from the Jehol Group of western Liaoning
and the coelurosaurian evolution. PhD Thesis. Chinese Academy of Sciences. 325
pp.
Xu, Norell, Wang, Makovicky and Wu, 2002. A basal troodontid
from the Early Cretaceous of China. Nature. 415, 780-784.
Haubold, 2003. Literaturbericht - Dinosauria 2002-2003. Zentralblatt
f�r Geologie und Pal�ontologie, Teil II. 2003(5/6), 467-524.
Dinosaur.net, online 2004. http://www.dinosaur.net.cn/_Kyohaku2004/show_fly.htm (not archived but but copied at http://projectos.cienciaviva.pt/pw011/jazidas/220[1].sinevenator.jpg)
Senter, Barsbold, Britt and Burnham, 2004. Systematics and evolution of Dromaeosauridae.
Bulletin of Gunma Natural History Museum. 8, 1-20.
Manabe, 2005. The Dinosaur Expo 2005: Evolution of Dinosaurs from their Origin to Birds. Asahi Shinbunsha. 149 pp.
Senter, 2007. A new look at the phylogeny of Coelurosauria. Journal of Systematic
Palaeontology. 5(4), 329-463.
Long and Schouten, 2008. Feathered Dinosaurs: The Origin of Birds. Oxford University Press. 193 pp.
White, 2009. The subarctometatarsus: Intermediate metatarsus architecture demonstrating
the evolution of the arctometatarsus and advanced agility in theropod dinosaurs.
Alcheringa. 33(1), 1-21.
Xu, Han and Zhao, 2014. Homologies and homeotic transformation of the theropod
'semilunate' carpal. Nature Scientific Reports. 4, 6042.
Fossil Mall, online 2015. http://www.fossilmall.com/Science/Sites/China/Sinovenator/Sinovenator.jpg
Ma and Rayfield, 2015. Reconstructing the cranial musculoskeletal anatomy of
two maniraptoran theropod dinosaurs and implications for avian evolution. Journal
of Vertebrate Paleontology. Program and Abstracts 2015, 170.
Yin, Pei and Zhou, 2018. Cranial morphology of Sinovenator changii (Theropoda: Troodontidae) on the new material from the Yixian Formation of western Liaoning, China. PeerJ. 6:e4977.
Liaoningvenator Shen, Zhao,
Gao, Lu and Kundrat, 2017
L. curriei Shen, Zhao, Gao, Lu and Kundrat, 2017
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (DNHM D3012) (4+ year old subadult) skull (97.6 mm), mandibles,
ten cervical vertebrae fused to cervical ribs (series ~138 mm; c3-8 13.6-16.1
mm [c5], c9 12.5, c10 12.1 mm), twelve dorsal vertebrae (series ~150 mm), several
partial to incomplete dorsal ribs, gastralia, sacrum, first to sixteenth caudal
vertebrae (series ~245 mm; c14 18.9 mm), chevron?, incomplete scapula (~59 mm),
coracoids, humerus (~65 mm), radii (one distal), ulnae (one distal), phalanx
I-1 (32.6 mm), manual ungual I, partial metacarpal II, phalanx II-1 (26.3 mm),
phalanges II-2 (one distal; ~34 mm), manual unguals II, partial metacarpal III,
partial phalanx III-1, phalanx III-2, phalanx III-3, manual ungual III, ilia
(71, 77 mm), partial pubis, distal ischia, femora (one partial; 111 mm), tibiotarsi
(162, ~152 mm), fibula (152 mm), distal tarsal III, distal tarsal IV, phalanx
I-1, pedal ungual I, metatarsals II (~82 mm), phalanx II-1, phalanx II-2, partial
pedal ungual II, metatarsals III (~93 mm), phalanx III-1, phalanx III-2, phalanx
III-3, metatarsal IV, phalanx IV-1, metatarsal V (~24 mm)
Referred- ?(CAGS-IG01-004) incomplete skeleton including skull (Hwang
et al., 2004)
Diagnosis- (after Shen et al., 2017) prominent slender triradiate postorbital;
transition point in caudal series starts from seventh caudal vertebra; deltopectoral
crest distinctly extended to distal half of humeral shaft; manual phalanx I-1
longer than metacarpal II, ratio about 1.49; no posterodistal process on ischium;
slender obturator process on ischium; metatarsus width distally distinctly decreases.
Comments- Shen et al. (2017) added Liaoningvenator to a version
of Senter's TWiG analysis and recovered it sister to Eosinopteryx as an
anchiornithine troodontid. Hartman et al. (2019) found it to be sister to IGM 100/1126, closer to troodontines than Anchiornis or sinovenatorines. Wwang et al. (2004) mentioned CAGS-IG01-004 as a specimen from the Lujiatun
as being similar to Almas, so this may be a second individual of Liaoningvenator.
It was stated to have small teeth with constricted bases and carinae lacking
serrations, an apparently absent postorbital (may be taphonomic), and no quadratojugal-squamosal
contact.
Based on figures, I believe when Shen et al. state "The manual phalanx
III-1 is the longest", they meant phalanx II-2.
References- Hwang, Norell, Ji and Gao, 2004. A new troodontid from the
lower Yixian Formation of China and its affinities to Mongolian troodontids.
Journal of Vertebrate Paleontology. 24(3), 26A.
Shen, Zhao, Gao, Lu and Kundrat, 2017. A new troodontid dinosaur (Liaoningvenator
curriei gen. et sp. nov.) from the Early Cretaceous Yixian Formation in
western Liaoning Province. Acta Geoscientica Sinica. 38(3), 359-371.
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 troodontid (Wang, Zhang, Tan, Jiangzuo, Zhang and Tan, 2021 online)
Late Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Material- (LH PV39) (adult) ~fourth cervical vertebra
(20.51 mm), ~fifth cervical vertebra (20.82 mm), ~sixth cervical
vertebra (19.37 mm), ~seventh cervical fragment, ~eighth cervical
vertebra (15.95 mm), ~ninth cervical vertebra (14.18 mm), synsacrum
(10.70, 9.80, 7.25, 7.53, 8.49 mm), four mid caudal vertebrae (23.12,
25.88, 24.81, 26.22 mm), two fragmentary chevrons, manual ungual I
(24.2 mm on curve), fragmentary manual ungual
Comments- This was discovered
in 2001 and first reported by Sereno (online 2001) with photos of two
cervicals and an ungual, misidentified as a dromaeosaur. Wang et
al. (2021) described it in detail, recovering it as the most basal
sinovenatorine using Pei's version of the TWiG analysis in a parsimony
analysis, and in a more derived position in a polytomy with Almas, IGM 100/1126, Talos
and troodontids closer to troodontines in a Bayesian analysis.
Adding it to Hartman et al.'s maniraptoromorph matrix results in a
position sister to Liaoningvenator,
then IGM 100/1126. Forcing it to be a sinovenatorine or slightly
closer to Troodontinae sister to the contemporaneous and also small Philovenator
both take a single additional step, suggesting its position within
Troodontidae is not strongly resolved and that it may belong to Philovenator.
References- Sereno, online 2001. https://web.archive.org/web/20020603194101/http://www.projectexploration.org/mongolia/u52701.htm
Wang, Zhang, Tan, Jiangzuo, Zhang and Tan, 2021 online. New troodontid
theropod specimen from Inner Mongolia, China clarifies phylogenetic
relationships of later-diverging small-bodied troodontids and paravian
body size evolution. Cladistics. Early View. 10.1111/cla.12467
undescribed troodontid (Dufeau, 2003)
Late Campanian, Late Cretaceous
Zos Wash, Djadokhta Formation, Mongolia
Material-
(IGM 100/1126; = IGM 100/1005, IGM 100/1128) (6+ year old adult)
incomplete skull (78.2 mm), sclerotic plates, mandibles (68.1, 66.3
mm), hyoids, synsacrum, first caudal vertebra, second caudal vertebra
(6.1 mm), third caudal vertebra (6.8 mm), fourth caudal vertebra (7.2
mm), fifth caudal vertebra (8.2 mm), sixth caudal vertebra (8.8 mm),
seventh caudal vertebra (8.9 mm), eighth caudal vertebra (9.5 mm),
ninth caudal vertebra (9.9 mm), tenth caudal vertebra (11 mm), eleventh
caudal vertebra (11.5 mm), twelfth caudal vertebra (12.6 mm), proximal
thirteenth caudal vertebra, chevrons, distal radius, distal ulna,
scapholunare, semilunate carpal, metacarpal I (11 mm), phalanx I-1 (~27
mm),
manual ungual I (12.8 mm), metacarpal II (~25 mm), phalanx II-1 (16.1
mm), phalanges II-2 (24.3, 26.6 mm), manual unguals II (~13, 13.2 mm),
metacarpal III (~24 mm), phalanx III-1 (6 mm), phalanx III-2 (5.8 mm),
phalanges III-3 (15.1, 17.3 mm), manual unguals III (8.5, 9.1 mm),
incomplete ilia, pubes (one distal; ~65 mm), ischia (~34 mm), femora
(87.5 mm), tibiotarsi (122.4 mm), fibulae, metatarsal I, phalanges I-1
(6.4, 6.5 mm), pedal unguals I (5, 4.9 mm), metatarsals II (one
incomplete; 71.8 mm), phalanges II-1 (12.5, ~12 mm), phalanges II-2
(6.4 mm), pedal unguals II (~14 mm), metatarsals III (one incomplete;
78.5 mm), phalanges III-1 (13, 12.5 mm), phalanx III-2 (9.8 mm),
phalanx III-3 (8.9 mm), pedal ungual III, metatarsals IV (one
incomplete; 76.2 mm), phalanges IV-1 (8.3, 8.8 mm), phalanges IV-2
(7.2, 7.6 mm), phalanx IV-3 (6.2 mm), phalanx IV-4 (~7 mm), pedal
ungual IV (8.5 mm), partial pedal phalanx III/IV-? (Dufeau, 2003)
(IGM 100/3500) (adult) maxilla, fused parietals, posterior braincase,
dentaries, proximal caudal vertebra, several fragmentary distal caudal
vertebrae, distal chevrons, incomplete femur, tibiotarsus, incomplete
metatarsal II, phalanx II-1, phalanx II-2, pedal ungual II, incomplete
metatarsal III, phalanx III-1, phalanx III-2, phalanx III-3, pedal
ungual III, incomplete metatarsal IV, phalanx IV-1, phalanx IV-2,
phalanx IV-3, phalanx IV-4, pedal ungual IV (Pei, 2015)
Comments- This specimens were collected in 1997. In the
Fighting Dinosaurs: New Discoveries from Mongolia exhibit at the AMNH in 2000,
it was displayed with troodontid postcrania. It is mislabeled on the AMNH website
that year as the Shuvuuia holotype, though it differs in many respects from that
taxon. A braincase analyzed by Franzosa (2004) as the Zos Canyon troodontid
and labeled IGM 100/1005 is the same specimen, as Dufeau (2003) includes a photo
of the skull labeled "IGM 100/1005 Undescribed troodontid". Hwang
et al. (2004) refer to a second Ukhaa Tolgod basal troodontid skull (in addition
to Almas),
which based on their description I infer to be this specimen. Erickson
et al. (2009) examined the femur of an undescribed troodontid they call
IGM 100/1129, which is a misprint for this same specimen (Pei,
2015). As of 2009, the skull was labeled IGM 100/1128 at the AMNH
(pers. obs. 6-21-2009), which is reflected by its number in Turner's
(2008) thesis. By the publications of Turner et al. 2011 and 2012
the entire specimen was called IGM 100/1126, as it is in its
unpublished description by Pei (2015). IGM 100/1126 was included
in Turner's TWiG analyses (2008; et al., 2011 and 2012), where it
emerged as a jinfengopterygine troodontid along with Almas. Pei used a version of Brusatte's TWiG analysis which recovered IGM 100/1126 and Almas closer to troodontines than Sinornithoides
or jinfengopterygines. Hartman et al. (2019) used another version of
the TWiG analysis and found an intermediate topology where Almas and IGM 100/1126 plus Liaoningvenator are closer to troodontines and Sinornithoides than sinovenatorines/Jinfengopteryx.
References- AMNH, 2000 online. http://paleo.amnh.org/gobi/gobi.swf [requires discontinued Shockwave Flash]
Dufeau, 2003. The cranial anatomy of the theropod dinosaur Shuvuuia deserti
(Coelurosauria: Alvarezsauridae), and its bearing upon coelurosaurian
phylogeny. Masters Thesis, The University of Texas at Austin. 275 pp.
Franzosa, 2004. Evolution of the brain in Theropoda (Dinosauria). PhD Thesis,
The University of Texas at Austin. 357 pp.
Hwang, Norell, Ji and Gao, 2004. A new troodontid from the lower Yixian Formation
of China and its affinities to Mongolian troodontids. Journal of Vertebrate
Paleontology. 24(3), 73A-74A.
Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis,
Columbia University. 666 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.
Turner, Pol and Norell, 2011. Anatomy of Mahakala omnogovae (Theropoda:
Dromaeosauridae), T�gr�giin Shiree, Mongolia. American Museum Novitates.
3722, 66 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 206 pp.
Pei, 2015. New paravian fossils from the Mesozoic of east Asia and
their bearing on the phylogeny of the Coelurosauria. PhD thesis,
Columbia University. 545 pp.
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
Almas Pei, Norell, Barta, Bever, Pittman and Xu, 2017
A. ukhaa Pei, Norell, Barta, Bever, Pittman and Xu, 2017
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadokhta Formation, Mongolia
Holotype-
(IGM 100/1323)
(subadult) incomplete skull (~82 mm), sclerotic plates, mandibles (one
anterior, one partial), gastralia, posterior three sacral vertebrae,
first caudal neural arch, second caudal vertebra, third caudal
vertebra, fourth caudal vertebra, fifth caudal vertebra, sixth caudal
vertebra, seventh caudal vertebra, eighth caudal vertebra, ninth caudal
vertebra, tenth caudal vertebra, eleventh caudal neural arch, five
chevrons, partial ilium (56.5 mm), pubes (one incomplete; 60.1 mm),
ischia (one partial; 38.5 mm), femora (one partial; 68.3 mm), tibiae
(tibiotarsi 94.5, 96.1 mm), fragmentary fibulae, astragalocalcaneum,
partial metatarsals II, partial metatarsal III, partial metatarsal IV,
metatarsal V, six eggshell fragments
Referred- (IGM 100/972) (juvenile) partial premaxillae, maxillae, nasals,
lacrimals, incomplete jugals, anterior frontal, palatal elements, incomplete
mandible (Norell, Clark, Demberelyin, Barsbold, Chiappe, Davidson, McKenna,
Perle and Novacek, 1994)
(IGM 100/974) (juvenile) incomplete skull, posterior mandible (Norell, Clark,
Demberelyin, Barsbold, Chiappe, Davidson, McKenna, Perle and Novacek, 1994)
?(IGM 100/1003) (adult) tooth
....(juvenile) skeleton including cervical vertebrae, dorsal ribs, incomplete
femora, tibiae, fibula, metatarsi, pedal phalanges, nineteen eggs, nest (Clark
et al., 2002)
Diagnosis- (after Pei et al., 2017) posteriorly curved pterygoid flange; absence
of lateral groove on anterior part of dentary; third chevron more than
three times as long as corresponding caudal vertebra; distinct
spike-like process extending from anterior edge of obturator process on
ischium.
Comments- The holotype of Almas was found in 1993, first published as similar to possible Liaoningvenator
specimen CAGS-IG01-004 in having teeth of the same number and
morphology, and a modified diapsid configuration (Hwang et al., 2004).
Hwang (2005; 2007) describes the tooth morphology, while Erickson et
al. (2009) examined femoral histology. Hwang (2007) also scored it in
an early TWiG analysis, finding it to emerge sister to Byronosaurus. Turner (2008; published as Turner
et al., 2012) also included it in a TWiG analysis, finding it to emerge with
Jinfengopteryx
and IGM 100/1126 as a jinfengopterygine troodontid. This was also found
in the related unpublished version shown in Turner et al. (2011). Pei
and Norell (2011) briefly described the holotype an an abstract, while
Pei (2015) later described Almas
in his thesis, which was published as Pei et al. (2017). Pei used a
version of Brusatte's TWiG analysis to recover it closer to
troodontines than Sinornithoides
or jinfengopterygines. The 2017 description doesn't include a
phylogenetic analysis but does describe characters placing Almas in
that derived position.
Baby Byronosaurus and nest? IGM 100/972 and 100/974 are two juvenile skulls found associated
with Citipati nest IGM 100/971 in 1993 (Norell et al., 1994). They were identified by Norell
et al. as Velociraptor, based on their long premaxillae (on IGM 100/974
at least). However, Norell and Makovicky (1999) stated that Norell et al. (in
press) will show they are actually troodontids. This later became Bever and
Norell (2008, 2009), who supported their assignment to Byronosaurus,
as did Grellet-Tinner (2005) for 100/972 at least. However, Pei and Norell (2011)
referred at least IGM 100/972 to the then-unnamed Almas, based on (taller?) snout shape and less maxillary teeth
compared to Byronosaurus, but these are juvenile characters expected
in any nestling. Adding to the possibilities, IGM 100/1126 from the Djadohkta has serrationless teeth, and embryonic
Troodon lack serrations, so young individuals of the contemporaneous
Saurornithoides might as well. Sues and Averianov (2007) mistakenly state IGM 100/974
was reidentified as an oviraptorid, and that the serrationless teeth were actually
maxillary palatal bumps, but they were confusing it with embryonic Citipati
specimen IGM 100/971.
Discovered in 1995, Norton (DML, 2000) first stated "troodontid nest
with 13 hatched eggs and one hatchling skeleton" IGM 100/1003 was on
display at the AMNH's Fighting Dinosaurs exhibit. Clark et al. (2002)
noted an undescribed Ukhaa Tolgod troodontid nest with juveniles and an
adult tooth as being the subject of an upcoming paper, cited as Norell
et al., in prep.. Only the eggs have so far been described, in
Grellet-Tinner's (2005) thesis.
Grellet-Tinner (2005) determined eggshell attached to IGM 100/972's
skull is the same as that from that eroded nest, and that 100/972 was
found in the same location as 100/1003 but a few meters downhill. As
neither 100/972 or 100/974 were in situ in the Citipati nest, it seems plausible they eroded out
from 100/1003. If true, and Pei and Norell are correct that 100/972 is not Byronosaurus,
this would eliminate referral of any nests or eggs to the latter genus.
The same thing? Pei and
Norell (2011) already suggested IGM 100/974 and Almas were the same
taxon, and Pei et al. (2017) state that among Djadokhta troodontids,
"the Ukhaa perinates IGM 100/972 and IGM 100/974 have a higher chance
of being closely related to Almas ukhaa." They correctly note 100/972
lacks a lateral groove on the anterior dentary, and Bever and Norell
(2009) also state it has a posteriorly curved lateral flange on its
pterygoid (both unpreserved in 100/974), both proposed synapomorphies
of Almas. Pei et al.'s objection to them being conspecific is "both
specimens were found in a nest of eggs with sizes that are too large to
be those of Almas ukhaa", but the perinates were found in a nest of
Citipati eggs and the size of the eggs in IGM 100/1003 has not been
reported (Grellet-Tinner, 2005).
References- Norell, Clark, Dashzeveg, Barsbold, Chiappe, Davidson, McKenna
and Novacek, 1994. A theropod dinosaur embryo, and the affinities of the Flaming
Cliffs dinosaur eggs. Science. 266, 779-782.
Norell and Makovicky, 1999. Important features of the dromaeosaurid skeleton
II: Information from newly collected specimens of Velociraptor mongoliensis.
American Museum Novitates. 3282, 1-45.
Norton, DML 2000. https://web.archive.org/web/20210603191138/http://dml.cmnh.org/2000Jun/msg00082.html
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria,
Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences.
38(12), 1753-1766.
Clark, Norell and Makovicky, 2002. Cladistic approaches to the relationships
of birds to other theropod dinosaurs. In Chiappe and Witmer (eds.). Mesozoic
Birds: Above the Heads of Dinosaurs. University of California Press. 31-64.
Norell and Hwang, 2004. A troodontid dinosaur from Ukhaa Tolgod (Late Cretaceous
Mongolia). American Museum Novitates. 3446, 9 pp.
Hwang, Norell, Ji and Gao, 2004. A new troodontid from the
lower Yixian Formation of China and its affinities to Mongolian troodontids.
Journal of Vertebrate Paleontology. 24(3), 73A-74A.
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.
Hwang, 2005. Phylogenetic patterns of enamel microstructure in dinosaur teeth.
Journal of Morphology. 266(2), 208-240.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
Hwang, 2007. Phylogenetic patterns of enamel microstructure in dinosaur teeth.
PhD Thesis, Columbia University. 274 pp.
Bever and Norell, 2008. Neonate troodontid skulls from the Upper Cretaceous
of Mongolia with observations on the cranial ontogeny of paravian theropods.
Journal of Vertebrate Paleontology. 28(3), 52A.
Turner, 2008. Phylogenetic relationships of paravian Theropods. PhD Thesis,
Columbia University. 666 pp.
Bever and Norell, 2009. The perinate skull of Byronosaurus (Troodontidae)
with observations on the cranial ontogeny of paravian theropods. American Museum
Novitates. 3657, 51 pp.
Erickson, Rauhut, Zhou, Turner, Inouye, Hu and Norell, 2009. Was dinosaurian
physiology inherited by birds? Reconciling slow growth in Archaeopteryx.
PLoS ONE. 4(10), e7390.
Pei and Norell, 2011. A new troodontid (Dinosauria: Theropoda) from the Late
Cretaceous Djadokhta Formation of Mongolia. Journal of Vertebrate Paleontology.
Program and Abstracts 2011, 172.
Turner, Pol and Norell, 2011. Anatomy of Mahakala omnogovae (Theropoda:
Dromaeosauridae), T�gr�giin Shiree, Mongolia. American Museum Novitates.
3722, 66 pp.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371,
1-206.
Pei, 2015. New paravian fossils from the Mesozoic of east Asia and
their bearing on the phylogeny of the Coelurosauria. PhD thesis,
Columbia University. 545 pp.
Pei, Norell, Barta, Bever, Pittman and Xu, 2017. Osteology of a new
Late Cretaceous troodontid specimen from Ukhaa Tolgod, �mn�govi Aimag,
Mongolia. American Museum Novitates. 3889, 47 pp.
Philovenator Xu, Zhao, Sullivan,
Tan, Sander and Ma, 2012
P. curriei Xu, Zhao, Sullivan, Tan, Sander and Ma, 2012
Late Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype- (IVPP V10597) (~.68 m; 1.1 kg; subadult) femur (86.5 mm), incomplete
tibiotarsus (~105 mm), incomplete fibula, metatarsal I (4 mm), proximal phalanx
I-1, tarsometatarsus (mtII 94 mm, mt III 107 mm, mtIV 105 mm), phalanx II-1
(10.5 mm), phalanx II-2 (8.4 mm), proximal pedal ungual II, phalanx III-1 (14.5
mm), phalanx III-2 (12 mm), proximal phalanx III-3, phalanx IV-1 (11 mm), phalanx
IV-2 (10.5 mm), phalanx IV-3 (9 mm), phalanx IV-4 (9 mm), pedal ungual IV (8
mm), metatarsal V (21.5 mm)
Diagnosis- (after Xu et al.,
2012) prominent process on medial side of femoral shaft slightly
proximal to the distal end; sheet-like cnemial crest that expands
significantly anteriorly; astragalocalcaneal condyles deep
anteroposteriorly and separated by deep and narrow groove; extremely
long and slender tarsometatarsus (tarsometatarsus/femur length ratio
1.25, tarsometatarsus length/width ratio 22.0); anteroposterior depth
much greater than transverse width at mid-shaft of tarsometatarsus;
prominent, elongate posterior flange that extends along most of
metatarsal IV; metatarsal IV flange about equal in depth to shaft.
Comments- The holotype was found in 1988, reported by Dong et al. (1989) as "a well preserved hindlimb of a juvenile Saurornoithoides", and originally described by Currie and Peng (1994) as a possible juvenile
Saurornithoides mongoliensis. Norell et al. (2009) believed this
specimen to be merely Troodontidae indet., as they stated hindlimb elements
were undiagnostic in Saurornithoides and Zanabazar. More recently,
Xu et al. (2012) have reexamined the specimen and found it to be a subadult of
a new taxon of troodontid named Philovenator,
recovered as a troodontine in a version of Senter's TWiG
analysis. A smaller analysis of troodontids and hindlimb
characters refined it as sister to Linhevenator in that clade.
References- Dong, Currie and Russell, 1989. The 1988 field program of the Dinosaur Project. Vertebrata Palasiatica. 27(3), 233-236.
Currie and Peng, 1994. A juvenile specimen of Saurornithoides
mongoliensis from the Upper Cretaceous of northern China. Canadian Journal
of Earth Sciences. 30(10), 2224-2230.
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria,
Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences.
38(12), 1753-1766.
Norell, Makovicky, Bever, Balanoff, Clark, Barsbold and Rowe, 2009. A review
of the Mongolian Cretaceous dinosaur Saurornithoides (Troodontidae: Theropoda).
American Museum Novitates. 3654, 63 pp.
Xu, Zhao, Sullivan, Tan, Sander and Ma, 2012. The taxonomy of the troodontid
IVPP V 10597 reconsidered. Vertebrata PalAsiatica. 50(2), 140-150.
Tamarro Sell�s, Vila, Brusatte, Currie and Galobart, 2021
T. insperatus Sell�s, Vila, Brusatte, Currie and Galobart, 2021
Late Maastrichtian, Late Cretaceous
Talarn Formation, Tremp Group, Spain
Holotype- (MCD-7073) (subadult) incomplete metatarsal II (~140 mm)
Diagnosis- (After Sell�s et
al., 2021) metatarsal II with marked plantar ridge; small foramen on
lateral surface of plantar ridge of metatarsal II;
subarctometatarsalian condition with the metatarsal III restricted to
the plantar margin on its proximal part.
Comments- This was discovered in September 2003. Sell�s et al. (2021) used Hartman et al.'s maniraptoromorph analysis to recover Tamarro as a jinfengopterygine troodontid in a trichotomy with IGM 100/140 and 100/1128, with Philovenator and Liaoningvenator successively further out.
Reference- Sell�s, Vila,
Brusatte, Currie and Galobart, 2021. A fast‑growing basal troodontid
(Dinosauria: Theropoda) from the latest Cretaceous of Europe.
Scientific Reports. 11:4855.
unnamed clade (Daliansaurus liaoningensis + Troodon formosus)
Comments- This clade contains
all troodontids with enlarged serrations, so isolated teeth with such
are listed here. Note some members have small or absent
serrations, however.
"Saurornithoides"
asiamericanus (Nessov, 1995) Olshevsky, 2000
= Pectinodon "asiamericanus" Nessov, 1985
= Saurornithoides "asiamericanus" (Nessov, 1985) Olshevsky,
1991
= Troodon asiamericanus Nessov, 1995
Early Cenomanian, Late Cretaceous
Khodzhakul Formation, Uzbekistan
Holotype- (CCMGE 49/12176) posterior dentary tooth (~3.5x~2.4x~1.1 mm)
Referred- (ZIN PH 1883/16) tooth (Averianov and Sues, 2016)
(ZIN PH 1884/16) tooth (Averianov and Sues, 2016)
(ZIN PH 1885/16) anterior dentary tooth (Averianov and Sues, 2007)
(ZIN PH 1886/16) anterior dentary tooth (Averianov and Sues, 2007)
(ZIN PH 1888/16) maxillary tooth (Averianov and Sues, 2007)
(ZIN PH 1889/16) tooth (Averianov and Sues, 2016)
(ZIN PH 1890/16) tooth (Averianov and Sues, 2016)
(ZIN PH 1891/16) tooth (Averianov and Sues, 2016)
?(ZIN PH 2355/16) distal caudal vertebra (Averianov and Sues, 2016)
?(ZIN PH 2356/16) distal caudal vertebra (Averianov and Sues, 2016)
?(ZIN PH 2357/16) distal caudal vertebra (Averianov and Sues, 2016)
?(ZIN PH 2358/16) distal caudal vertebra (Averianov and Sues, 2016)
Comments- The holotype tooth was originally distinguished from Troodon
formosus by its smaller size, more labiolingually compressed crowns, unserrated
mesial carina and smaller distal serrations (Nessov, 1995). However, these characters
are also seen in more basal troodontids such as Sinornithoides and Sinusonasus
(Averianov and Sues, 2004). The referred teeth are indistinguishable from the
holotype.
References- Nessov, 1985. New mammals from the Cretaceous of the Kyzylkum.
Vestnik Leningradskogo Universiteta. Seriya 7, 8-18.
Olshevsky, 1991. A revision of the parainfraclass Archosauria Cope, 1869, excluding
the advanced Crocodylia. Mesozoic Meanderings. 2, 196 pp.
Nessov, 1995. Dinosaurs of northern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust,
St. Petersburg State University. 156 pp.
Olshevsky, 2000. An annotated checklist of dinosaur species by continent. Mesozoic
Meanderings. 3, 157 pp.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
Averianov and Sues, 2016. Troodontidae (Dinosauria: Theropoda) from the Upper
Cretaceous of Uzbekistan. Cretaceous Research. 59, 98-110.
unnamed troodontid (Currie, Rigby and Sloan, 1990)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith River Group, Alberta, Canada
Material- (RTMP 79.8.635) tooth (3.3 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 85.30.1) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 2000.19.1) tooth (5.5 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 2000.20.1) tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 2000.21.11) tooth (5.2 mm) (Sankey, Brinkman, Guenther and Currie, 2002)
Late Maastrichtian, Late Cretaceous
Lance Formation, Montana, Wyoming, US
(AMNH 8114) tooth (Estes, 1964)
?(AMNH 27117) eleven teeth (Estes, 1964; AMNH online)
Diagnosis- (after Sankey et al., 2002) tooth crowns flattened or concave
on one side; enamel pitted on flat/concave side; two well-developed longitudinal
ridges present on flat surface, one ridge extending from the apex denticle to
near the base of the enamel and the second extending along the anterior carina;
mesial serrations always absent; distal serrations rounded.
Comments- This tooth morphology was considered a pathological varient
of Troodon formosus by Currie et al. (1990), who referred to it as "Paronychodon"
(Troodon). Sankey et al. (2002) considered it a valid taxon though, based
on its differing temporal and spatial distribution than Troodon formosus.
Specifically, this taxon is restricted to the upper Dinosaur Park Formation
within the Judith River Group, and has not been reported from the Prince Creek
Formation where T. formosus is common. Sankey et al. also referred teeth
described by Estes (1964) as cf. Saurornithoides sp. from the Lance Formation
to this species, based on figured specimen AMNH 8114. Troodon bakkeri
from that formation is not the same as this taxon, as it has pointed distal
serrations like Troodon formosus. It's highly possible additional Late
Cretaceous North American teeth currently identified as Troodon belong
to this taxon.
References- Estes, 1964. Fossil vertebrates from the Late Cretaceous
Lance Formation, eastern Wyoming. University of California Publications in Geological
Sciences. 49, 1-180.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. In Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press. 107-125.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
undescribed Troodontidae (Kirkland, Lucas and Estep, 1998)
Late Maastrichtian, Late Cretaceous
North Horn Formation, Utah, US
Material- (BYU coll.) nest, 13 eggs (~170x~76 mm) (Difley, Britt and Policelli, 2004)
(BYU coll.) nest, eggs (Difley, Britt and Policelli, 2004)
(OMNH coll.) teeth (Cifelli, Nydam, Eaton, Gardner, Kirkland, 1999)
Comments- Kirkland et al. (1998) and Cifelli et al. (1999) list Troodontidae
gen. et sp. indet. from the North Horn Formation. Difley et al. (2004) report two nests of Prismatoolithus eggs referrable to Troodontidae.
References- Kirkland, Lucas and Estep, 1998. Cretaceous dinosaurs of
the Colorado plateau. In Lucas, Kirkland and Estep (eds.). Lower and Middle
Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural History and
Science Bulletin. 14, 79-89.
Cifelli, Nydam, Eaton, Gardner, Kirkland, 1999. Vertebrate faunas of the North
Horn Formation (Upper Cretaceous-Lower Paleocene), Emery and Sanpete counties,
Utah. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological Survey,
Miscellaneous Publication. 99-1, 378-388.
Difley, Britt and Policelli, 2004. A troodontid nest in the North Horn Formation,
central Utah. Journal of Vertebrate Paleontology. 24(3), 115A.
unnamed Troodontidae (Estes and Sanchiz, 1982)
Late Hauterivian-Early Barremian, Early Cretaceous
Castellar Formation, Spain
Material- (MPT coll.) teeth (~1.2x~.9x? mm)
Comments- These were referred to Coeluridae by Estes and Sanchiz (1982) based on resemblence to cf. Saurornithoides
tooth AMNH 8114 of Estes (1964), now identified as a troodontine.
Estes and Sanchiz noted the Spanish specimens have smaller serrations
though, which closely matches dentary teeth of e.g. Daliansaurus and Sinornithoides. As in that taxon, mesial serrations are absent.
References- Estes, 1964. Fossil vertebrates from the Late Cretaceous
Lance Formation, eastern Wyoming. University of California Publications in Geological
Sciences. 49, 1-180.
Estes and Sanchiz, 1982. Early Cretaceous lower vertebrates
from Galve (Teruel), Spain. Journal of Vertebrate Palaeontology. 2(1), 21-39.
unnamed Troodontidae (Novikov, Lebedev and Alifanov, 1998)
Aptian-Albian, Early Cretaceous
Ilek (=Shestakovo) Formation, Russia
Material- (PM TGU 16/5-124) maxillary or posterior dentary tooth (~2.7x2.1x1 mm) (Averianov and Sues, 2007)
(PM TGU coll.) caudal vertebra (Averianov and Sues, 2007)
(PM TGU coll.) metacarpal I (Averianov and Sues, 2007)
(private coll.) ?dentary with teeth (~10 mm), ?tibiotarsus (Alifanov et
al., 1999)
Comments- Alifanov et al. (1999a, b) state "materials
included lower leg and jawbone with small teeth with a height of about
1 cm. The latter are characterized by a strong arcuate curvature of the
leading edge and the development of a rounded pre-root protrusion at
the posterior edge" (translated). Averianov and Sues (2007) report it
"was excavated at the Shestakovo 3 site by a commercial collector from
Novosibirsk and was unfortunately unavailable for study." They
describe an isolated tooth PM TGU 16/5-124 with 13 hooked
serrations (5.1/mm) only present distally, but say "the first
metacarpal and caudal from Shestakovo 1 will be described elsewhere."
References- Novikov, Lebedev and Alifanov, 1998. New Mesozoic vertebrate
fossil sites of Russia. Third European Workshop on Vertebrate Paleontology, p. 58.
Alifanov, Efimov, Novikov and Morales, 1999a. Novyy psittakozovrovyy
kompleks tetrapod iz nizhnemelovogo mestonakhozhdeniya Shestakovo
(yuzhnaya Sibir'). Doklady Akademii Nauk. 369(4), 491-493.
Alifanov, Efimov, Novikov and Morales, 1999b. A new psittacosaurian complex of
tetrapods from the Lower Cretaceous Shestakovo locality (southern Siberia).
Doklady Earth Sciences. 369A(9), 1228-1230.
Leshchinskiy, Voronkevich, Fayngertz, Maschenko, Lopatin and Averianov, 2001.
Early Cretaceous vertebrate locality Shestakovo, western Siberia, Russia: A
refugium for Jurassic relicts? Journal of Vertebrate Paleontology. 21(3), 73A.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
unnamed troodontid (Zhao, 2003)
Cenomanian, Late Cretaceous
Liantoutang Formation, Zhejiang, China
Material- (ZMNH M8711) two nests of five and fourteen eggs
Comments- These are referred to the ootaxon Prismatoolithus sp., suggesting they are more closely related to Troodon than Philovenator.
References- Zhao, 2003. The nesting behavior of troodontid dinosaurs.
Vertebrata PalAsiatica. 41, 157-168.
Varricchio, Jin and Jackson, 2015. Lay, brood, repeat: Nest reuse and site fidelity
in ecologic time for two Cretaceous troodontid dinosaurs. Journal of Vertebrate
Paleontology. 35(3), e932797. DOI: 10.1080/02724634.2014.932797
unnamed troodontid (Barsbold, Osmolska and Kurzanov, 1987)
Aptian-Albian, Early Cretaceous
Dzunbain Formation, Mongolia
Material- (IGM 100/44) quadrate, baincase fragment, dentary fragment,
posterior mandible, four teeth (2.5-3 mm), five partial cervical neural arches
(22-25 mm), semilunate carpal, metacarpal I (15 mm), phalanx I-1 (36 mm), manual
ungual I (18 mm), metacarpal II (40 mm), phalanx II-1 (26 mm), incomplete phalanx
II-2 (24 mm), manual ungual II, metacarpal III (~37 mm), manual ungual III,
distal femur, metatarsal I (13 mm), phalanx I-1 (13 mm), distal metatarsal II,
phalanx II-1 (25 mm), phalanx II-2 (17 mm), pedal ungual II (~20 mm), distal
metatarsal III (~108 mm), phalanx III-1 (29 mm), phalanx III-2 (17 mm), phalanx
III-3 (~12 mm), distal metatarsal IV, phalanx IV-1 (19 mm), phalanx IV-2 (~13
mm), pedal ungual IV fragment
Comments- Discovered in 1979,
this is notable as being the first described Early Cretaceous specimen
recognized as being troodontid, until the description of Sinornithoides
in 1994. The specimen has gained notoriety for being included in
all TWiG analyses since 2004 as the 'Early Cretaceous
troodontid'. While described by Barsbold et al. as Troodontidae
indet., its unique character combination in matrices indicates future
authors should compare IGM 100/44 in depth to the many Early Cretaceous
troodontids now known to determine its validity. Although
Barsbold et al. initially placed the specimen in the Barunbayaskaya
Svita, its Khamryn-Us locality has most recently been referred to the
Dzunbain Formation (e.g. description of the co-occuring Mongolostegus).
Reference- Barsbold, Osmolska and Kurzanov, 1987. On a new troodontid
(Dinosauria, Theropoda) from the Early Cretaceous of Mongolia. Acta Palaeontologica
Polonica. 32(1-2), 121-132.
Daliansaurus Shen, L�,
Liu, Kundr�t, Brusatte and Gao, 2017
D. liaoningensis Shen, L�, Liu, Kundr�t, Brusatte and Gao, 2017
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (DNHM D2885) (~1 m; 4 year old adult) partial skull (~138 mm), incomplete
mandibles, nine cervical vertebrae with fused cervical ribs, thirteen dorsal
vertebrae, fifteen dorsal ribs (seven partial), two uncinate processes, five
sacral vertebrae, first to twenty-eighth caudal vertebrae, seven chevrons, humeri
(82.9 mm), radii (65.2, 65.7 mm), ulnae (65, 64.7 mm), semilunate carpals, metacarpals
I (15.7, 14.1 mm), phalanges I-1 (28.7, 31.1 mm), manual unguals I (16.6, 25.4
mm), metacarpals II (30.5, 31.5 mm), phalanges II-1 (~20.5, 23.2 mm), phalanges
II-2 (27.5, 27.8 mm), manual unguals II (20.8, 19.9 mm), metacarpals III (~31,
32 mm), phalanx III-1 (7.5 mm), phalanx III-2 (8.7 mm), phalanges III-3 (19.2
mm), manual unguals III (19.7, 19.1 mm), ilia (82.3, 92.8 mm), distal ischium,
femur (130.8 mm), tibiotarsus (190.1 mm), fibula, metatarsal I, phalanx I-1
(11.3 mm), pedal ungual I (13.3 mm), metatarsal II (98.1 mm), phalanx II-1 (20.1
mm), phalanx II-2 (14.7 mm), incomplete pedal ungual II (22.7 mm), metatarsal
III, partial phalanx III-1, metatarsal IV (110.1 mm) , phalanx IV-1 (15.9 mm),
phalanx IV-2 (14.8 mm), phalanx IV-3 (13.3 mm), phalanx IV-4 (11.3 mm), pedal
ungual IV (20.8 mm)
Diagnosis- (after Shen et al., 2017) uncinate processes on dorsal ribs
(among troodontids); pedal ungual IV robust, deep, and approximately the same
size as pedal ungual II. Differs from Sinusonasus in that metatarsal
II terminates before trochlea of metatarsal IV begins; metatarsal IV lacks prominent
longitudinal flange.
Other diagnoses- Shen et al. (2017) state "Daliansaurus differs
from Sinovenator and Sinusonasus in that ... in that metacarpal
II is slightly shorter than metacarpal III (instead of longer)" but the
manus is unknown in Sinusonasus.
Comments- Discovered prior to June 2016, this was described as a sinovenatorine troodontid sister to
Sinusonasus by Shen et al. (2017) using Brusatte's version of the TWiG
dataset. Hartman et al. (2019) used a more extensive version of the TWiG analysis to recover it and Sinusonsus in a clade closer to troodontines than sinovenatorines. Forcing Daliansaurus into Sinovenatorinae takes 4 more steps.
References- Shen, L�, Liu, Kundr�t, Brusatte and Gao, 2017. A new troodontid
dinosaur from the Lower Cretaceous Yixian Formation of Liaoning Province, China.
Acta Geologica Sinica. 91(3), 763-780.
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
Shen, L�, Gao, Hoshino, Uesugi and Kundr�t, 2019. Forearm bone histology of the small theropod Daliansaurus liaoningensis (Paraves: Troodontidae) from the Yixian Formation, Liaoning, China. Historical Biology. 31(2), 253-261.
Tochisaurus Kurzanov and
Osmolska, 1991
T. nemegtensis Kurzanov and Osmolska, 1991
Early Maastrichtian, Late Cretaceous
Nemegt, Nemegt Formation, Mongolia
Holotype- (PIN 551-224) (~2.8 m) metatarsal II (233 mm), metatarsal III
(232 mm), metatarsal IV (242 mm)
Diagnosis- (after Kurzanov and Osmolska, 1991) long, slender metatarsus; strongly reduced metatarsal II.
Comments- This was found in 1948 and originally figured by Kurzanov (1987) as an unknown
theropod, then identified by Osmolska (1987) as a troodontid. Although Osmolska
speculated it might be a specimen of Borogovia, Kurzanov and Osmolska
(1991) noted the distal end of metatarsal II is far more reduced in Tochisaurus.
Similarly, Tochisaurus differs from Zanabazar in having metatarsal
II narrower proximally, and the proximal surface of the metatarsus inclined.
It could not be compared to the earlier Saurornithoides, however. Tochisaurus
was not included in published phylogenetic analyses until that of
Hartman et al. (2019), where it fell out sister to the earlier Daliansaurus. While this should be considered provisional given the incomplete holotype of Tochisaurus, it is notable that Daliansaurus has Kurzanov and Osmolska's two proposed Tochisaurus auatpomorphies but to a more extreme degree.
References- Kurzanov, 1987. Avimimidae and the problem of the origin
of birds. Trudy, Sovmestnaa Sovetsko-Mongolskaa paleontologiceskaa
ekspedicia. 31, 1-95.
Osmolska, 1987. Borogovia gracilicrus gen. et sp. n., a new troodontid
dinosaur from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica.
32, 133-150.
Kurzanov and Osm�lska, 1991. Tochisaurus nemegtensis gen. et sp.
n., a new troodontid (Dinosauria, Theropoda) from Mongolia. Acta Palaeontologia
Polonica. 36, 69-76.
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
Hesperornithoides Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019
H. miessleri Hartman, Mortimer, Wahl, Lomax, Lippincott and Lovelace, 2019
Late Kimmeridgian, Late Jurassic
Jimbo Quarry, Brushy Basin Member of the Morrison Formation, Wyoming, US
Holotype-
(WYDICE-DML-001; = WDC DML0001; Lori) (~.89 m; adult) incomplete skull
(~66 mm), incomplete mandibles, hyoids, partial axis, third cervical
vertebra (11.7 mm), fourth cervical vertebra (~14 mm), ?sixth cervical
vertebra (14.5 mm), eighth cervical vertebra (13.7 mm), cervical ribs,
first dorsal vertebra (11.7 mm), anterior dorsal rib, four mid caudal
vertebrae (two partial; 20.8, 22.3 mm), seven distal caudal vertebrae
(three fragmentary; 22.3, 23.6, 18.2, 17.5 mm), five chevrons, partial
scapula, partial coracoid, partial furcula, proximal humerus, distal
humerus, radius, ulna (46.4 mm), scapholunare, semilunate carpal, metacarpal
I (16.9 mm), manual ungual I (17.1 mm), metacarpal II, manual ungual II
(15.5 mm), metacarpal III, phalanx III-2 (7 mm), phalanx III-3 (18.2
mm), manual ungual III (14.4 mm), ilial fragment, incomplete femur,
tibiae (168 mm), fibulae, astragalus, calcaneum, distal tarsal,
metatarsal I, incomplete metatarsals II, pedal ungual II, incomplete
metatarsals III, phalanx III-1 (26.3 mm), phalanx III-2 (17.3 mm),
phalanx III-3, pedal ungual III (13.6 mm), incomplete metatarsals IV,
phalanx IV-1, phalanx IV-2 (13.2 mm), phalanx IV-3 (~8.5 mm), phalanx
IV-4 (6.9 mm), proximal pedal ungual IV, pedal claw sheath, metatarsal V
Diagnosis- (after Hartman et al., 2019) pneumatic jugal (also in Zanabazar
and some eudromaeosaurs among maniraptorans); short posterior lacrimal
process (<15% of ventral process length, measured from internal
corner; also present in Zanabazar, Archaeopteryx, and Epidexipteryx);
quadrate forms part of lateral margin of paraquadrate foramen; small
external mandibular fenestra (<12% of mandibular length; also in Zhenyuanlong and Dromaeosaurus
among non-avian paravians); humeral entepicondyle >15% of distal
humeral width (also in some avialans); manual ungual III subequal in
size to ungual II (also in Daliansaurus, IGM 100/44 and Mahakala); mediodistal corner of tibia exposed anteriorly (also in Archaeopteryx and Jeholornis).
Comments-
This specimen was discovered in Summer 2001 and announced in an
abstract (Lovelace, 2004). In Hartman et al.'s (2005) preliminary
analysis also reported via abstract and Wahl's (2006) thesis, it
emerged sister to Sinornithoides using the Mei
TWiG matrix. Hartman et al. (2019) officially described and named the
taxon, using a new extensive TWiG analysis to place it most
parsimoniously sister to Xixiasaurus plus Sinusonasus
slightly more stemward in Troodontidae. As they state "a
placement as the first branching dromaeosaurid is just two steps
longer, supported by the dorsally placed maxillary fenestra, mesial
dental serrations, and large lateral teeth. This may be more compatible
stratigraphically, but moving Hesperornithoides' clade to a more stemward position in Troodontidae outside Sinovenatorinae, the Liaoningvenator-like
taxa and derived troodontids is also only two steps longer. Similarly,
in trees two steps longer than the MPTs where troodontids are avialans,
Hesperornithoides can be the
first branching taxon closer to Aves than troodontids based on
homoplasic characters such as the short posterodorsal lacrimal process.
Two additional steps also place the taxon in contemporaneous
Archaeopterygidae, sister to Caihong
which shares [mesial dental serrations, large lateral teeth and
elongate but not hypertrophied distal caudal prezygapophyses]. Despite
the uncertainty of its position within Paraves, however, Hesperornithoides
is strongly supported as a member of the Deinonychosauria plus Avialae
clade, as even constraining it to the paravian stem requires 15
additional steps."
References- Lovelace, 2004. Taphonomy and paleoenvironment of a Late
Jurassic dinosaur locality in the Morrison Formation of east-central Wyoming.
Journal of Vertebrate Paleontology. 24(3), 152A.
Hartman, Lovelace and Wahl, 2005. Phylogenetic assessment of a maniraptoran
from the Morrison Formation. Journal of Vertebrate Paleontology. 25(3), 67A-68A.
Wahl, 2006. Osteology and phylogenetic relationships of a new small maniraptoran
from the Upper Jurassic of Wyoming. Masters Thesis, Fort Hays State University. 98 pp.
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
Xixiasaurus Lu, Xu, Liu,
Zhang, Jia and Ji, 2010
X. henanensis Lu, Xu, Liu, Zhang, Jia and Ji, 2010
Coniacian-Campanian, Late Cretaceous
Lower-Middle Majiacun Formation, Henan, China
Holotype- (41HIII-0201) anterior skull, anterior dentary, radial fragment,
ulnar fragment, phalanx I-1 (36 mm), manual ungual I, distal metacapal II, phalanx
II-1 (30 mm), proximal phalanx II-2, distal metacarpal III, distal manual ungual
Diagnosis- (after Lu et al., 2010) distinct opening on the lateral surface
of the base of the nasal process of the premaxilla; snout shows a more tapered
U-shape than in Byronosaurus; 22 maxillary teeth; mandibular symphyseal
region is slightly inflected medially.
Comments- Discovered prior to March 2009, Lu et al. (2010) proposed Xixiasaurus was most closely related to Byronosaurus and Urbacodon
based on the lack of dental serrations. However, Hartman et al.
(2019) found the distribution of serrations to be highly homoplasic
within troodontids, with serrationless teeth primitive for
maniraptoriforms but Xixiasaurus itself nested within a clade of serrated taxa (Sinusonasus, Hesperornithoides, Daliansaurus, IGM 100/44).
References- Lu, Xu, Liu, Zhang, Jia and Ji, 2010. A new troodontid (Theropoda:
Troodontidae) from the Late Cretaceous of central China, and the radiation of
Asian troodontids. Acta Palaeontologica Polonica. 55(3), 381-388.
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
Sinusonasus Xu and Wang, 2004
S. magnodens Xu and Wang, 2004
Early Aptian, Early Cretaceous
Lujiatun Beds of Yixian Formation, Liaoning, China
Holotype- (IVPP V 11527) (adult) incomplete skull (~109 mm), incomplete
mandible, several dorsal ribs, gastralia, sacrum, twenty-seven caudal vertebrae
(537 mm), chevrons, pubis (113 mm), ischia (61 mm), femur (141 mm), tibiae (186
mm), fibula (167 mm), metatarsal I (22.7, 19.9 mm), phalanx I-1 (10.2, 12.1
mm), pedal ungual I (15.7, 15.7 mm), metatarsal II (93.5 mm), phalanx II-1 (24.2,
21.2 mm) phalanx II-2 (17.6, 20.6 mm), pedal ungual II (31.1 mm), metatarsal
III (~108 mm), phalanx III-1 (28.5 mm), phalanx III-2 (19.1 mm), phalanx III-3
(19.4 mm), pedal ungual III (13.6 mm), metatarsal IV (106.6 mm), phalanx IV-1,
phalanx IV-2 (13.7 mm), phalanx IV-3 (14.7 mm), metatarsal V
Diagnosis- (after Xu and Wang, 2004) interantorbital canal absent; nasal
sinusoid in lateral view; middle maxillary teeth relatively large; distal chevrons
elongated to contact each other anteroposteriorly; femoral neck long.
Comments- The holotype was discovered prior to December 2003. This genus was consistantly mispelled Sinucerasaurus
by Xu and Norell (2006), though the latter was written over a year after the
publication of Sinusonasus. Xu and Norell (2004) proposed it was closer to troodontines than Sinovenator and Sinornithoides, but Hartman et al. (2019) recovered it intermediate between those two genera.
References- Xu and Wang, 2004. A new troodontid (Theropoda: Troodontidae)
from the Lower Cretaceous Yixian Formation of western Liaoning, China. Acta
Geologica Sinica. 78(1), 22-26.
Xu and Norell, 2006. Non-avian dinosaur fossils from the Lower Cretaceous Jehol
Group of western Liaoning, China. Geological Journal. 41(3-4), 419-437.
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
Sinornithoides Russell and
Dong, 1994
S. youngi Russell and Dong, 1994
Early Cretaceous
Ejinhoro Formation, Inner Mongolia, China
Holotype- (IVPP V9612) (1.1 m, juvenile) skull (109 mm), hyoid, atlantal neural arch,
fourth cervical centrum (18 mm), fifth cervical vertebra (18 mm), sixth cervical
prezygapophysis, ninth cervical vertebra (19.1 mm), ninth cervical rib, tenth
cervical vertebra (14.4 mm), tenth cervical rib, first dorsal vertebra (13 mm),
first dorsal rib, ten pairs of dorsal ribs, fifteen rows of gastralia, (sacrum-
72 mm) sixth sacral vertebra (12 mm), first caudal vertebra (~11.5 mm), first
chevron (16.7 mm), second caudal vertebra (~13.5 mm), second chevron (23.8 mm),
third caudal vertebra (14.5 mm), fourth caudal vertebra, fifth caudal vertebra,
sixth caudal vertebra, seventh caudal vertebra (11.6 mm), seventh chevron (9.2
mm), eighth caudal vertebra (14 mm), ninth caudal vertebra (17.5 mm), tenth
caudal vertebra (21.2 mm), eleventh caudal vertebra (21.5 mm), twelfth caudal
vertebra (23 mm), thirteenth caudal vertebra (24.5 mm), fourteenth caudal vertebra
(23.9 mm), fifteenth caudal vertebra (23 mm), sixteenth caudal vertebra (22.5
mm), seventeenth caudal vertebra (23.5 mm), eighteenth caudal vertebra (22.9
mm), nineteenth caudal vertebra (24 mm), twentieth caudal vertebra (23.8 mm),
twenty-first caudal vertebra (23 mm), twenty-second caudal vertebra (22 mm),
twenty-third caudal vertebra (18 mm), twenty-fourth caudal vertebra (22.4 mm),
twenty-fifth caudal vertebra (~20 mm), twenty-sixth caudal vertebra (~18 mm),
twenty-seventh caudal vertebra (~16.8 mm), chevrons, proximal scapula, coracoids
(~30 mm long, 27 mm deep), incomplete furcula, humeri (82.8 mm), radius (59.1
mm), ulna (65 mm), scapholunare, semilunate carpal, metacarpal I (11.3, 12.1 mm),
phalanx I-1 (29.3 mm), manual ungual I (19.7 mm), metacarpal II (37.1 mm), phalanx
II-1 (19 mm), phalanx II-2 (28.5, 28.1 mm), manual ungual II (22, 22 mm), metacarpal
III (35 mm), phalanx III-1 (4.2 mm), phalanx III-2 (8.2 mm), phalanx III-3 (20.3
mm), manual ungual III (15.5 mm), ventral ilia (73 mm), pubes (89 mm), ischia
(47.5 mm), femora (140 mm), tibiae (197.6 mm), fibulae, astragali (19.2 mm wide),
distal tarsal III, distal tarsal IV, metatarsal I (~12.3, 12.3 mm), phalanx
I-1 (11.4, 10 mm), pedal ungual I (10.7, 9.5 mm), metatarsal II (100.4, 100.6
mm), phalanx II-1 (20.2, 21.7 mm), phalanx II-2 (11.7 mm), pedal ungual II (19
mm), metatarsal III (111 mm), phalanx III-1 (27.6, 23 mm), phalanx III-2 (20,
19.2 mm), phalanx III-3 (16, 15.8 mm), pedal ungual III (14.6 mm), metatarsal
IV (110 mm), phalanx IV-1 (17.8, 18.2 mm), phalanx IV-2 (15.7, 13.9 mm), phalanx
IV-3 (13, 11.7 mm), phalanx IV-4 (12.5, 13 mm), pedal ungual IV (11.5 mm), metatarsal
V (41.8 mm)
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 was discovered on August 6
1988. Dong (1992) photographs and mentions this specimen as "Saurornithoides
new species (Russell and Dong in prep.)" before it was more fully
prepared. Russell and Dong (1994) initially proposed the taxon
was stemward of Troodon, Zanabazar, Saurornithoides and Borogovia, which has been recovered in all analyses since. Currie and Dong (2001) later described the skeleton in more detail.
Dong (1997) referred a fragmentary specimen (IVPP V11119) to Sinornithoides
sp. nov., but it seems to be more basal.
References- Dong, 1992. Dinosaurian Faunas of China. China Ocean Press. 188 pp.
Russell and Dong, 1994. A nearly complete skeleton of a new
troodont dinosaur from the Early Cretaceous of the Ordos Basin, Inner Mongolia,
People's Republic of China. Canadian Journal of Earth Sciences. 30(10), 2163-2173.
Dong, 1997. On small theropods from Mazongshan area, Gansu province,
China. In Dong (ed.). Sino-Japanese Silk Road Dinosaur Expedition.
China Ocean Press. 13-18.
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria,
Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences.
38, 1753-1766.
White, 2009. The subarctometatarsus: intermediate metatarsus architecture demonstrating
the evolution of the arctometatarsus and advanced agility in theropod dinosaurs.
Alcheringa. 33(1), 1-21.
Byronosaurus Norell, Makovicky
and Clark, 2000
= "Byranjaffia" Novacek, 2002
B. jaffei Norell, Makovicky and Clark, 2000
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadokhta Formation, Mongolia
Holotype- (IGM 100/983) partial skull, incomplete mandibles, posterior
axis, incomplete third cervical vertebra, anterior fourth cervical vertebra,
fourth cervical rib, two anterior dorsal centra, posterior dorsal vertebrae,
dorsal rib fragment, possible sacral centrum, four distal caudal fragments,
distal femur (~150 mm), proximal tibia, proximal fibula, limb bone fragments,
distal metatarsal II, phalanx II-2, distal phalanx III-2, proximal phalanx III-3
Paratype- (IGM 100/984) premaxillae, anterior maxillae, maxillary fragment,
lacrimals, posterior nasals, vomer fragment
Diagnosis-
(after Norell et al., 2000) teeth lacking serrations; interfenestral
bar not recessed from lateral plane of maxilla; interantorbital canal;
shallow groove along the ventrolateral margin the maxilla.
Comments- The holotype was discovered in 1993 and illustrated as an undescribed
troodontid in Novacek et al. (1994). Its unassociated remains were mixed with
those of ornithomimid specimen IGM 100/987. The paratype was found later on July 15
1996. Novacek (2002) used the name "Byranjaffia" in his book for Byronosaurus.
The former is either a misspelling or an unpublished name previously
intended for the genus. Both specimens were described briefly by
Norell et al. (2000) then later in detail by Makovicky et al. (2003).
Note the nests and juvenile remains referred to Byronosaurus by Bever and Norell (2008- IGM 100/972, 100/974) and Grellet-Tinner (2005- IGM 100/1003) are here placed in Philovenator.
References- Novacek, Norell, McKenna and Clark, 1994. Fossils of the Flaming Cliffs. Scientific
American. 271(6), 60-69.
Norell, Makovicky and Clark, 2000. A new troodontid theropod from Ukhaa Tolgod,
Mongolia. Journal of Vertebrate Paleontology. 20(1), 7-11.
Novacek, 2002. Time Traveler: In Search of Dinosaurs and Ancient Mammals from
Montana to Mongolia. Farrar, Strauss and Giroux. 368 pp.
Makovicky, Norell, Clark and Rowe, 2003a online. Byronosaurus jaffei, Digital Morphology.
http://digimorph.org/specimens/Byronosaurus_jaffei/rostrum/
Makovicky, Norell, Clark and Rowe, 2003b online. Byronosaurus jaffei, Digital Morphology.
http://digimorph.org/specimens/Byronosaurus_jaffei/braincase/
Makovicky, Norell, Clark and Rowe, 2003. Osteology and relationships of Byronosaurus
jaffei (Theropoda: Troodontidae). American Museum Novitates. 3402, 1-32.
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.
Bever and Norell, 2008. Neonate troodontid skulls from the Upper Cretaceous
of Mongolia with observations on the cranial ontogeny of paravian theropods.
Journal of Vertebrate Paleontology. 28(3), 52A.
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.
Troodontinae sensu van der Reest and Currie, 2017
Definition- (Gobivenator mongoliensis + Zanabazar junior)
Reference- van der Reest and Currie, 2017. Troodontids (Theropoda) from
the Dinosaur Park Formation, Alberta, with a description of a unique new taxon:
Implications for deinonychosaur diversity in North America. Canadian Journal
of Earth Sciences. 54, 919-935.
Gobivenator Tsuihiji, Barsbold,
Watabe, Tsogtbaatar, Chinzorig, Fugiyama and Suzuki, 2014
G. mongoliensis Tsuihiji, Barsbold, Watabe, Tsogtbaatar, Chinzorig,
Fugiyama and Suzuki, 2014
Campanian, Late Cretaceous
Dzamin Khond, Djadochta Formation, Mongolia
Holotype- (IGM 100/86; 070623 DK CHZ) incomplete skull, incomplete mandible, proatlas,
atlas, axis (14.6 mm), third to fourth cervical vertebrae, eighth to tenth cervical
vertebrae, cervical ribs, twelve dorsal vertebrae (d1 24 mm, d12 21.6 mm), dorsal
ribs, gastralia, synsacrum (205.4 mm), thirty-five caudal vertebrae (c1 15.8
mm, c34 19.4 mm), chevrons, scapula (107.9 mm), coracoids, humeri (108.9 mm),
proximal radius, proximal ulna, incomplete ilium, pubis (169.4 mm), ischium
(~92 mm), femur (~192 mm), proximal tibia, proximal fibula, astragalocalcaneum,
metatarsal I, phalanx I-1, pedal ungual I, distal tarsal III, distal tarsal
IV, metatarsal II (145.2 mm), phalanx II-1, phalanx II-2, metatarsal III (159.4
mm), metatarsal IV (163 mm), phalanx IV-1, metatarsal V
Campanian, Late Cretaceous
Tugrikin Shire, Djadochta Formation, Mongolia
Referred- ?(HMNS coll.)
incomplete skull, incomplete mandibles, partial skeleton including two
manual phalanges (Tsogtbaatar and Chinzorig, 2010)
Diagnosis-
(after Tsuihiji et al., 2014) pointed anterior end of fused parietal
that wedges into V-shaped notch between frontals; gracile anterior,
posterior and ventral rami of postorbital; fossa on surangular in front
of posterior surangular foramen; dorsoventrally elongated
proximal chevrons (up to 4.5 times as long as the height of the
preceding caudal vertebrae).
Comments- Gobivenator
was discovered on June 15 and/or 23 2007 (Saneyoshi et al., 2010),
mentioned as "jaw and associated skeletons of a Troodontidae" with
field number 070623 DK CHZ. Whether the partial dentary with
teeth shown in figure 8B as "Lower jaw of Troodontidae" belongs to the
holotype is unknown, as only the opposite mandible is shown in its
description. Tsuihiji et al. (2010) presented the specimen in an
abstract, recovering it as sister to Byronosaurus
in a preliminary phylogenetic analysis. Tsuihiji et al. (2014)
later described the taxon in more detail and named it, this time
recovering Gobivenator as sister to troodontines with Byronosaurus
very basal based on a version of Senter's TWiG matrix. Most
recently, Hartman et al.'s (2019) extensive TWiG matrix found it
similarly close to troodontines but with Byronosaurus just slightly more stemward.
A skull photographed by Tsogtbaatar and Chinzorig (2010) as "a troodontid from Tugrikin Shire" (Fig. 3) seems most similar to Gobivenator
(~88% of the holotype's size) in that it has the slender postorbital
and at least ventrally defined surangular fossa, while strongly
differing from Saurornithoides, Byronosaurus, Almas
and IGM 100/1126 in various features. Differences such as the
narrower interfenestral bar and undeveloped groove on the ventrolateral
maxilla, posteriorly angled dorsal jugal process and slender angular
could be individual variation or taxonomic. It was originally
discovered on August 1 1994 and was misidentified in the field as Velociraptor (field number 940801 TS-I WTB) (Watabe et al., 2000).
References-
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1994. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 30-44.
Saneyoshi, Watabe, Tsubamoto, Tsogtbaatar, Chinzorig and Suzuki, 2010.
Report of the HMNSMPC Joint Paleontological Expedition in 2007.
Hayashibara Museum of Natural Sciences Research Bulletin. 3, 19-28.
Tsogtbaatar and Chinzorig, 2010. Fossil specimens prepared in Mongolian
Paleontological Center: 2002–2008. Hayashibara Museum of Natural
Sciences Research Bulletin. 3, 155-166.
Tsuihiji, Watabe, Tsogtbaatar, Suzuki and Barsbold, 2010.
A new troodontid (Dinosauria: Theropoda) from the Late Cretaceous of the Gobi
Desert, Mongolia. Journal of Vertebrate Paleontology. Program and Abstracts
2010, 177A-178A.
Tsuihiji, Barsbold, Watabe, Tsogtbaatar, Chinzorig, Fugiyama and Suzuki, 2014.
An exquisitely preserved troodontid theropod with new information on the palatal
structure from the Upper Cretaceous of Mongolia. Naturwissenschaften. 101, 131-142.
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
Geminiraptor Senter, Kirkland,
Bird and Bartlett, 2010
G. suarezarum Senter, Kirkland, Bird and Bartlett, 2010
Barremian, Early Cretaceous
Yellow Cat Member of Cedar Mountain Formation, Utah, US
Holotype- (CEUM 73719) partial maxilla
Diagnosis- (after Senter et al., 2010) maxilla with extensive pneumatization
internal to antorbital fossa, inflating the bone so that it has a triangular
cross-section; large, anteroposteriorly elongate maxillary fenestra; promaxillary
fenestra visible in lateral view; anteroposteriorly narrow promaxillary strut
and interfenestral strut; small, square dental alveoli with bony septa between
them.
Comments- Discovered in 2004, the holotype is incorrectly listed as CEUM 7319 in the paper
(Carpenter, online 2010 Comment to Senter et al., 2010).
Senter et al. (2010) used a version of the TWiG matrix to place Geminiraptor
as a troodontid more derived than Sinovenator, but less than troodontines.
References- Carpenter, online 2010. https://journals.plos.org/plosone/article/comment?id=10.1371/annotation/1da07b25-d3db-45e3-a063-c57958012b28
Senter, Kirkland, Bird and Bartlett, 2010. A new troodontid
theropod dinosaur from the Lower Cretaceous of Utah. PLoS ONE. 5(12), e14329.
Senter, Kirkland, Deblieux and Madsen, 2010. Three new theropods from the Cedar
Mountain Formation (Lower Cretaceous) of Utah. Journal of Vertebrate Paleontology.
Program and Abstracts 2010, 162A.
Urbacodon Averianov and Sues,
2007
U. itemirensis Averianov and Sues, 2007
Cenomanian, Late Cretaceous
Dzharakuduk Formation, Uzbekistan
Holotype- (ZIN PH 944/16) dentary, six teeth
Diagnosis- (after Averianov and Sues, 2007) distinguished from Troodon, Saurornithoides, Sinornithoides, Sinovenator, Sinusonasus and IGM 100/44 by the absence of serrations on the teeth; from Byronosaurus
by the presence of fewer neurovascular foramina in the lateral groove
on the dentary and by more bulbous anterior dentary crowns; from Mei by much larger size.
Comments- The holotype was discovered on September 9 2004.
Reference- Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda)
from the Cenomanian of Uzbekistan, with a review of troodontid records from
the territories of the former Soviet Union. Journal of Vertebrate Paleontology.
27(1), 87-98.
U? sp. (Averianov and Sues, 2007)
Late Turonian, Late Cretaceous
Bissekty Formation, Uzbekistan
Material- (CCMGE 2/11822) tooth (Nessov, 1995)
(CCMGE 71/12455) premaxillary tooth (Nessov, 1993)
?(CCMGE 466/12457) partial braincase (Nessov, 1995)
?(CCMGE 475/12457) mid caudal vertebra (Nessov, 1995)
?(SPbGU VZ Din 7/2) incomplete posterior dorsal vertebra (23.4 mm) (Averianov
and Sues, 2016)
?(USNM 538124) distal caudal vertebra (29 mm) (Averianov and Sues, 2016)
?(ZIN PH 99/16) pedal phalanx II-2 (Averianov and Sues, 2016)
(ZIN PH 265/16) anterior dentary tooth (Averianov and Sues, 2007)
?(ZIN PH 893/16) incomplete posterior dorsal vertebra (26.6 mm) (Averianov and
Sues, 2016)
(ZIN PH 1899/16) posterior maxillary or dentary tooth (Averianov and Sues, 2007)
?(ZIN PH 2308/16) partial maxilla (Averianov and Sues, 2016)
?(ZIN PH 2339/16) incomplete anterior dorsal vertebra (27.7 mm) (Averianov and
Sues, 2016)?
?(ZIN PH 2340/16) incomplete distal caudal vertebra (38 mm) (Averianov and Sues,
2016)
?(ZIN PH 2341/16) distal metatarsal III (Averianov and Sues, 2016)
?(ZIN PH 2342/16) distal metatarsal III (Averianov and Sues, 2016)
?(ZIN PH 2343/16) manual phalanx I-1 (43 mm) (Averianov and Sues, 2016)
?(ZIN PO 4608) partial dentary (Nessov, 1992)
? cervical vertebrae, metacarpal I (Averianov and Sues, 2007)
Comments- Nessov (1992, 1997) identified ZIN PO 4608 as an ichthyornithine.
Nessov (1993, 1997) identified CCMGE 71/12455 as Deinonychosauria or Mammalia.
Nessov (1995) identified CCMGE 2/11822 as Theropoda indet., CCMGE 466/12457
as possibly dromaeosaurid, and CCMGE 475/12457 as possibly ornithomimid. Averianov
and Sues (2007) reidentified these and additional specimens as being troodontid,
assigned to Urbacodon sp. due to the resemblence of the dental and dentary
remains to the U. itemirensis holotype. The material was described in
depth by Averianov and Sues (2016), except for the braincase CCMGE 466/12457
which will be described in a future publication on troodontid endocrania. Using
a reweighted version of the TWiG analysis, the authors recovered Urbacodon
as sister to Gobivenator outside Troodontinae.
References- Nessov, 1981. Cretaceous salamanders and frogs of the Kyzylkum
Desert. Trudy Zoologicheskogo Instituta AN SSSR. 101, 57-88.
Nessov, 1992. Review of localities and remains of Mesozoic and Paleogene birds
of the USSR and the description of new finds. Russkii Ornitologicheskii Zhurnal.
1, 7-50.
Nessov, 1993. New Mesozoic mammals of Middle Asia and Kazakhstan and comments
about evolution of theriofaunas of the Cretaceous coastal plains of Asia. Trudy
Zoologicheskogo Instituta RAN. 249, 105-133.
Nessov, 1995. Dinosaurs of northern 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.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
Averianov and Sues, 2016. Troodontidae (Dinosauria: Theropoda) from the Upper
Cretaceous of Uzbekistan. Cretaceous Research. 59, 98-110.
Troodontinae Gilmore, 1924 vide Martyniuk,
2012
Definition- (Troodon formosus + Saurornithoides mongoliensis)
(Martyniuk, 2012)
Other definitions- (Gobivenator mongoliensis + Zanabazar junior)
(van der Reest and Currie, 2017)
(Sinovenator changii + Troodon formosus) (Hendrickx,
Mateus, Ara�jo and Choiniere, 2019)
= Saurornithoidinae Barsbold, 1974 vide Martyniuk, 2012
Comments-
Martyniuk (2012) used three troodontid subfamilies, Jinfengopteryginae,
Troodontinae and Saurornithoidinae, but did not define the
latter. If ICZN rules are followed so that a subfamily cannot
contain a subfamily, Saurornithoidinae cannot exist given Saurornithoides
is an internal specifier of Troodontinae. van der Reest and
Currie's (2017) more inclusive definition of Troodontinae is invalid as
it does not include Troodon,
the eponymous genus (Phylocode Article 11.7). Hendrickx et al.'s
(2019) definition includes all troodontids except anchiornithines if
they belong here instead of as archaeopterygids, and is unlikely to
catch on if Sinovenatorinae is used.
References- Martyniuk, 2012. A Field Guide to Mesozoic Birds and Other
Winged Dinosaurs. Pan Aves. 189 pp.
van der Reest and Currie, 2017. Troodontids (Theropoda) from the Dinosaur Park
Formation, Alberta, with a description of a unique new taxon: Implications for
deinonychosaur diversity in North America. Canadian Journal of Earth Sciences.
54, 919-935.
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.
"Saurornitholestes"
robustus Sullivan, 2006
Late Campanian, Late Cretaceous
De-na-zin Member of the Kirtland Formation, New Mexico, US
Holotype- (SMP VP-1955) frontal (62 mm) (Sullivan, 2006)
Referred- ?(NMMNH P-68396) tooth (4.8x5.5x2.9 mm) (Williamson and Brusatte,
2014)
Diagnosis- Provisionally indeterminate relative to Troodon formosus,
and incomparable to Talos.
Other diagnoses- Sullivan (2006) used the ratio of length (measured along
the midline) to thickness (posterior part of the frontal) being 6:1 to distinguish
robustus from S. langstoni, but this matches the condition in
Troodon formosus once wear in robustus is accounted for.
Comments- Discovered in summer 2005, Sullivan (2006) originally described the frontal as a new species
of the dromaeosaurid Saurornitholestes, distinguished from S. langstoni
by its greater thickness. Turner et al. (2012) stated the holotype lacks the
synapomorphies of Saurornitholestes, but listed no characters of that
genus. They instead placed it as Theropoda indet.. Evans et al. (2014) reexamined
the specimen and determined it was nearly identical to Troodon and indeterminate
among derived troodontids. The tooth was said to fall within the range of variation
of Dinosaur Park Troodon teeth by Williamson and Brusatte (2014).
References- Williamson, 2001. Dinosaurs from microvertebrate sites in
the Upper Cretaceous Fruitland and Kirtland Formations, San Juan Basin, New
Mexico. 2001 GSA abstracts.
Sullivan, 2006. Saurornitholestes robustus, n. sp. (Theropoda: Dromaeosauridae)
from the Upper Cretaceous Kirtland Formation (De-na-zin Member), San Juan Basin,
New Mexico. In Lucas and Sullivan (eds.). Late Cretaceous vertebrates from the
Western Interior. New Mexico Museum of Natural History and Science Bulletin.
35, 253-256.
Turner, Makovicky and Norell, 2012. A review of dromaeosaurid systematics and
paravian phylogeny. Bulletin of the American Museum of Natural History. 371, 206 pp.
Evans, Larson, Cullen and Sullivan, 2014. 'Saurornitholestes' robustus
is a troodontid (Dinosauria: Theropoda). Canadian Journal of Earth Sciences.
51(7), 730-734.
Larson, Cullen, Todd and Evans, 2014. Geometric morphometrics of small theropod
frontals from the Dinosaur Park Formation, Alberta. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 165.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
Koparion Chure, 1994
K. douglassi Chure, 1994
Early Tithonian, Late Jurassic
Quarry 94, Brushy Basin Member of the Morrison Formation, Utah, US
Holotype- (DINO 3353) maxillary tooth (2 mm)
Diagnosis- differs from Sinusonasus, Sinornithoides and
"Saurornithoides" asiamericanus in having mesial serrations
on lateral teeth; differs from Sinornithoides, "Saurornithoides"
asiamericanus, Saurornithoides and Troodon in having comparatively
smaller distal serrations; differs from "Saurornithoides" asiamericanus,
Saurornithoides and Troodon in having distal serrations which
are not hooked apically.
Comments-
Chure's (1995) reference to "teeth ... most similar to ... troodontids"
from Dinosaur National Monument in Utah probably refers to Koparion.
Although Chure no longer thinks Koparion is a troodontid
(pers. comm. to Harris, 1997; in Naish, DML 1997), there are no other suggested
phylogenetic placements which are supported by the evidence. Rauhut (2000) suggested
it might be a compsognathid, but Compsognathus differs in lacking mesial
serrations. Compsognathus-like teeth from Guimarota described by Zinke
(1998) do have mesial serrations, but Koparion differs in having blood
pits, and less serrations per mm on both carinae (8 and 7 on mesial and distal
vs. 10-17 and 10-15 in cf. Compsognathus) despite similar crown size.
These are more similar to derived troodontids. The only other theropods with
constricted tooth bases and serrated teeth are Richardoestesia-like taxa
and therizinosaurs. Richardoestesia and related taxa differ in having
highly elongate teeth with extremely numerous serrations with no blood pits.
Therizinosauroids have much larger serrations (~3/mm in Beipiaosaurus
and Alxasaurus), while Falcarius' serration size is comparable.
However, maxillary teeth of Falcarius have convex distal edges with serrations
smaller in comparison to tooth size. Smaller posterior dentary teeth are schematically
illustrated as being more recurved, so may be more comparable to Koparion.
Therizinosaurs also lack blood pits. Koparion differs from Sinusonasus
in having mesial serrations, though they are similar in having small distal
serrations without apically hooked tips. It differs from Sinornithoides
and "Saurornithoides" asiamericanus in having comparatively
smaller distal serrations (Currie and Dong, 2001), and having mesial serrations
on non-premaxillary teeth. It further differs from "Saurornithoides"
americanus in having distal serrations whose tips are not hooked.
References- Chure and Britt, 1993. New data on theropod dinosaurs from
the Late Jurassic Morrison Fm. (MF). Journal of Vertebrate Paleontology. 13(3),
30A.
Chure, 1994. Koparion douglassi, a new dinosaur from the Morrison Formation
(Upper Jurassic) of Dinosaur National Monument; The oldest troodontid (Theropoda:
Maniraptora). Brigham Young University Geological Studies. 40, 11-15.
Chure, 1995. The teeth of small theropods from the Morrison
Formation (Upper Jurassic: Kimmeridgian), UT. Journal of Vertebrate Paleontology.
15(3), 23A.
Naish, DML 1997. https://web.archive.org/web/20190623204709/http://dml.cmnh.org/1997Jan/msg00325.html
Zinke, 1998. Small theropod teeth from the Upper Jurassic coal mine of Guimarota
(Portugal). Palaontologische Zeitschrift. 72(1/2) 179-189.
Rauhut, 2000. The interrelationships and evolution of basal theropods (Dinosauria,
Saurischia). Ph.D. dissertation, University of Bristol, Bristol. 583 pp.
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria,
Theropoda) from the People’s Republic of China. Canadian Journal of Earth
Sciences. 38, 1753-1766.
Troodontinae indet. (Lambe, 1902)
Late Campanian-Maastrichtian, Late Cretaceous
Judith River or Horseshoe Canyon Formation, Alberta, Canada
(RTMP 81.16.321) anterior dentary tooth (6 mm) (Currie, 1987)
(RTMP 81.20.71) anterior premaxillary tooth (Currie, 1987)
(RTMP 82.20.47) posterior dentary tooth (Currie, 1987)
(RTMP 82.20.299) mid-dentary tooth (Currie, 1987)
(RTMP 84.168.5) maxillary tooth (Currie, 1987)
Late Campanian-Maastrichtian, Late Cretaceous
Judith River or Edmonton Group, Alberta, Canada
(CMN 1266) tooth (Lambe, 1902)
(CMN 1560) astragalus (Zanno et al., 2011)
(CMN 8841) tooth (Sternberg, 1945)
(CMN coll.) five teeth (Sternberg, 1945)
Late Cretaceous
US
(AMNH 21629) tooth (AMNH online)
(YPM 55009) (YPM online)
(YPM 55040) (YPM online)
Comments- These could be Troodon, Stenonychosaurus, Latenivenatrix, Talos or another taxon.
References- Lambe, 1902. New genera and species from the Belly River
Series (mid-Cretaceous). Contributions to Canadian Palaeontology, Geological
Survey of Canada. 3, 23-81.
Sternberg, 1945. Pachycephalosauridae proposed for domeheaded dinosaurs, Stegoceras
lambei n. sp., described. Journal of Paleontology. 19, 534-538.
Currie, 1987. Bird-like characteristics of the jaws and teeth of troodontid
theropods (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. 7, 72-81.
Zanno, Varricchio, O’Connor, Titus and Knell, 2011. A new troodontid theropod,
Talos sampsoni gen. et sp. nov., from the Upper Cretaceous western interior
basin of North America. PLoS ONE. 6(9), e24487.
undescribed Troodontidae (Eaton, Kirkland, Hutchison, Denton, O'Neill and Parrish, 1997)
Late Cenomanian, Late Cretaceous
Naturita Formation (= "Dakota Formation"), Utah, US
Material- (MNA or OMNH coll.) teeth
Comments- These are identified as cf. Troodon sp. indet. in Eaton
et al. (1997) and later references.
References-
Eaton, Kirkland, Hutchison, Denton, O'Neill and Parrish, 1997.
Nonmarine extinction across the Cenomanian-Turonian boundary,
southwestern Utah, with a comparison to the Cretaceous-Tertiary
extinction event. Geological Society of America Bulletin. 109(5),
560-567.
Kirkland, Britt, Burge, Carpenter, Cifelli, DeCourten, Eaton, Hasiotis
and Lawton, 1997. Lower to Middle Cretaceous dinosaur faunas of the
central Colorado plateau: A key to understanding 35 million years of
tectonics, sedimentology, evolution, and biogeography. Brigham Young
University Geology Studies. 42, 69-103.
unnamed possible Troodontinae (Schwimmer, Sanders, Erickson and Weems,
2015)
Middle Campanian, Late Cretaceous
Coachman Formation, South Carolina, US
Material- (ChM PV8689) tooth (3 mm)
(ChM PV9111) incomplete tooth
Comments- These were only assigned to Theropoda indet. by Schwimmer et
al. (2015), but the large serration size, particularly on the mesial carina,
suggests they belong to troodontines.
Reference- Schwimmer, Sanders, Erickson and Weems, 2015. A Late Cretaceous
dinosaur and reptile assemblage from South Carolina, USA. Transactions of the
American Philosophical Society. 105(2), 157 pp.
unnamed possible troodontine (Rodriguez de la Rosa, 1996)
Early Maastrichtian, Late Cretaceous
Ca�on del Tule Formation, Mexico
Material- (IGM-7710) pedal phalanx II-2 (22.5 mm)
Diagnosis- shaft more elongate than other troodontids; proximoventral
heel bifurcated.
Comments- Although described as being from the Cerro del
Pueblo Formation, Aguillon Martinez (2010) found this and other
material from the El Pelillal locality belong to the later Ca�on del
Tule Formation.
Rodriguez de la Rosa and Cevallos-Ferriz (1998) refer this
to the Troodontidae because the collateral ligament pit is centrally placed.
This is only seen in Saurornithoides and Troodon though, not more
basal forms (IGM 100/44, Sinornithoides, Borogovia). Thus IGM-7710
may be more closely related to the former two genera. Evans et al. (2014) believe
this may belong to a turtle instead.
References- Hernandez, Aguillon, Delgado and Gomez, 1995. The Mexican
Dinosaur National Monument. Journal of Vertebrate Paleontology. 15(3), 34A.
Rodriguez de la Rosa, 1996. Vertebrate remains from a Late Cretaceous locality
(Campanian, Cerro del Pueblo Formation), Coahuila, Mexico. Journal of Vertebrate
Paleontology. 16(3), 60A.
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.
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.
Evans, Larson, Cullen and Sullivan, 2014. 'Saurornitholestes' robustus
is a troodontid (Dinosauria: Theropoda). Canadian Journal of Earth Sciences.
51(7), 730-734.
unnamed possible troodontine (Zinke, 1998)
Early Kimmeridgian, Late Jurassic
Alcobaca Formation, Portugal
Material- (IPFUB GUI D 93-97, 100, 102, 104, 111, 114-116, 156, 190) 14
teeth (~1.02 mm; FABL ~.92 mm)
Comments- These teeth can be identified as troodontid based on their
constricted roots, enlarged serrations and hooked distal serrations. The latter
are particularily derived characters among troodontids, as is the presence of
serrations on most mesial teeth.
Reference- Zinke, 1998, Small theropod teeth from the Upper Jurassic
coal mine of Guimarota (Portugal). Palaontologische Zeischrift. 72(1/2), 179-189.
unnamed Troodontinae (Averianov and Sues, 2007)
Early Santonian, Late Cretaceous
Yalovach Formation, Tajikistan
Material- (ZIN PH 1/66) anterior dentary tooth
(ZIN PH 2/66) dentary fragment
(ZIN PH 3/66) anterior dentary tooth
(ZIN PH 4/66) anterior maxillary tooth
(ZIN PH 5/66) posterior maxillary tooth
(ZIN PH 7/66) posterior dentary tooth
(ZIN PH 8/66) premaxillary tooth
?(ZIN PH 13/60) tooth
(ZIN PH coll.) at least four teeth
Comments- ZIN PH 13/60 may belong to another taxon because it is unserrated.
The dentary fragment ZIN PH 2/66 lacks tooth crowns, so may be referrable to
the taxon with serrated teeth or the same one as ZIN PH 13/60.
Reference- Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda)
from the Cenomanian of Uzbekistan, with a review of troodontid records from
the territories of the former Soviet Union. Journal of Vertebrate Paleontology.
27(1), 87-98.
unnamed troodontine (Nessov and Golovneva, 1990)
Middle Maastrichtian, Late Cretaceous
Kakanaut Formation, Russia
Material- (ZIN PH 1/28) partial tooth
Diagnosis- (after Averianov and Sues, 2007) no distinct pits between
the bases of the distal serrations; larger serrations (1.64/mm) than T. formosus.
Comments- This was referred to cf. Troodon sp. by Nessov and Goloneva
(1990) and Nessov (1992), and Troodon cf. formosus by Averianov and Sues
(2007) because Troodon is known from the Maastrichtian of Alaska, but
the mesial carina is missing so it is unknown if serrations existed mesially.
References-
Nessov and Golovneva, 1990. [History of the flora, vertebrates and
climate in the late Senonian of the north-eastern Koriak uplands]. In
Krasilov (ed.). [Continental Cretaceous of the USSR]. Dal’nevostochnoe
Otdelenie AN SSSR. 191-212.
Nesov, 1992. Maastrichtian dinosaurs of NE Asia and climate changes caused by
vertical oceanic circulation. International Conference on Arctic Margins, Abstracts.
43.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
undescribed Troodontidae (Bolotsky and Moiseenko, 1988)
Late Maastrichtian, Late Cretaceous
Udurchukan Formation of the Tsagayan Group, Russia
Material- teeth
Comments- These teeth have been noted as Troodontidae indet. (Bolotsky
and Moiseenko, 1988; Nessov and Golovneva, 1990; Nessov, 1995), Troodon cf.
formosus (Moiseenko et al., 1997) and Troodon sp. (Alifanov and Bolotsky,
2002), but have yet to be described.
References- Bolotsky and Moiseenko, 1988. [On Dinosaurs of the Amur river region]. Amur KNII DVO AN SSSR. 38 pp.
Nessov and Golovneva, 1990. [History of the flora, vertebrates and
climate in the late Senonian of the north-eastern Koriak uplands]. In
Krasilov (ed.). [Continental Cretaceous of the USSR]. Dal’nevostochnoe
Otdelenie AN SSSR. 191-212.
Nessov, 1995. Dinosaurs of nothern Eurasia: New data about assemblages, ecology,
and paleobiogeography. Institute for Scientific Research on the Earth's Crust. 156 pp.
Moiseenko, Sorokin and Bolotsky, 1997. [Fossil reptiles of the Amur river area.]
Amurskii Nauchnyi Tsentr DVO RAN. 54 pp.
Alifanov and Bolotsky, 2002. New data about the assemblages of the Upper Cretaceous
carnivorous dinosaurs (Theropoda) from the Amur region. In Kirillova (ed.).
Fourth International Symposium of IGCP 434. 25-26.
Averianov and Sues, 2007. A new troodontid (Dinosauria: Theropoda) from the
Cenomanian of Uzbekistan, with a review of troodontid records from the territories
of the former Soviet Union. Journal of Vertebrate Paleontology. 27(1), 87-98.
undescribed Troodontidae (Currie and Eberth,
1993)
Middle-Late Campanian, Late Cretaceous
Erenhot, Iren Dabasu Formation, Inner Mongolia, China
Material- (AMNH 6570 in part; paratype of Ornithomimus asiaticus) (juvenile or subadult) axis, third cervical vertebra, fifth cervical vertebra (Makovicky,
1995)
?(AMNH 6576 in part; paratype of Ornithomimus asiaticus) pedal ungual I (~17 mm) (pers. obs.)
(AMNH 21751) distal metatarsals III (Currie and Eberth,
1993)
(AMNH 21772) metatarsal II (~183 mm) (Currie and Eberth, 1993)
(AMNH 25570) three vertebrae (AMNH online)
(AMNH 30261) proximal metatarsal (AMNH online)
(AMNH 30262) proximal tibial fragment (AMNH online)
(AMNH 30263) proximal tibial fragment (AMNH online)
(AMNH 30264) tibial fragment, fibular fragment (AMNH online)
(AMNH 30265) proximal tibia (AMNH online) 142
(AMNH 30266) proximal fibula (AMNH online) 142
(AMNH 30267) proximal tibia (AMNH online)
(AMNH 30268) proximal fibula (AMNH online)
(AMNH 30269) proximal fibula (AMNH online) 142
(AMNH 30270) proximal fibula (AMNH online)
(AMNH 30271) partial astragalus (AMNH online)
(AMNH 30272) partial astragalus (AMNH online)
(AMNH 30273) partial astragalus (AMNH online)
(AMNH 30274) partial astragalus (AMNH online)
(AMNH 30275) distal humerus (AMNH online)
(AMNH 30276) distal humerus (AMNH online)
(AMNH 30277) distal humerus (AMNH online)
(AMNH 30278) proximal humerus (AMNH online)
(AMNH 30279) proximal humerus (AMNH online)
(AMNH 30280) proximal ulna (AMNH online)
(AMNH 30281) distal radius (AMNH online)
(AMNH 30282) distal radius (AMNH online)
(AMNH 30283) proximal scapula (AMNH online)
(AMNH 30284) proximal scapula (AMNH online)
(AMNH 30285) scapular blade (AMNH online)
(AMNH 30286) pedal ungual I (~19 mm) (AMNH online)
(AMNH 30287) proximal manual ungual (AMNH online)
(AMNH 30288) two posterior cervical or proximal caudal vertebrae (AMNH online)
(AMNH 30289) distal metatarsal IV (AMNH online)
(AMNH 30290) distal metatarsal IV (AMNH online)
(AMNH 30291) distal metatarsal IV (AMNH online)
(AMNH 30292) distal metatarsal III (AMNH online)
(AMNH 30293) distal metatarsal III (AMNH online)
(AMNH 30294) distal metatarsal II (AMNH online)
(AMNH 30295) distal metatarsal II (AMNH online)
(AMNH 30296) distal metatarsal II (AMNH online)
(AMNH 30297) distal metatarsal II (AMNH online)
(AMNH 30300) partial ilium (AMNH online) 142
(AMNH 30301) proximal ?pubis (AMNH online) 142
(AMNH 30302) ?ilial fragment (AMNH online)
(AMNH 30303) partial synsacrum (AMNH online) 142
(AMNH 30304) proximal ?pubis (AMNH online)
?(AMNH 30305) last sacral vertebra (~25 mm) (AMNH online)
(AMNH 30306) partial synsacrum (AMNH online)
(AMNH 30307) synsacral fragment(AMNH online)
(AMNH 30308) partial posterior cervical vertebra (AMNH online)
(AMNH 30309) partial posterior cervical vertebra (AMNH online)
(AMNH 30310) partial anterior dorsal vertebra (AMNH online)
(AMNH 30311) partial anterior dorsal vertebra (AMNH online)
(AMNH 30312) partial anterior dorsal centrum (AMNH online)
(AMNH 30313) incomplete anterior dorsal centrum (AMNH online)
(AMNH 30314) partial anterior dorsal centrum (AMNH online)
(AMNH 30315) incomplete anterior dorsal centrum (AMNH online)
(AMNH 30316) partial anterior dorsal vertebra (AMNH online)
(AMNH 30317) incomplete anterior dorsal centrum (~24 mm) (AMNH online)
(AMNH 30318) anterior dorsal centrum (~26 mm) (AMNH online)
(AMNH 30320) anterior dorsal centrum (~26 mm) (AMNH online) 142
(AMNH 30321) partial anterior dorsal vertebra (~26 mm) (AMNH online)
?(AMNH 30322) anterior dorsal centrum (~32 mm) (AMNH online)
(AMNH 30323) incomplete posterior dorsal centrum (~22 mm) (AMNH online)
(AMNH 30324) incomplete dorsal centrum (AMNH online)
(AMNH 30325) posterior dorsal centrum (~26 mm) (AMNH online)
(AMNH 30326) posterior dorsal centrum (~27 mm) (AMNH online)
(AMNH 30327) posterior dorsal centrum (~27 mm) (AMNH online)
(AMNH 30328) incomplete posterior dorsal vertebra (~27 mm) (AMNH online)
(AMNH 30329) incomplete posterior dorsal vertebra (~29 mm) (AMNH online)
(AMNH 30330) proximal caudal centrum (~23 mm) (AMNH online)
(AMNH 30336) ?central fragment (AMNH online)
(AMNH 30337) incomplete distal caudal vertebra (~35 mm) (AMNH online)
(AMNH 30338) mid caudal vertebra (~32 mm) (AMNH online)
(AMNH 30339) incomplete mid caudal vertebra (~31 mm) (AMNH online)
(AMNH 30340) incomplete mid caudal vertebra (AMNH online)
(AMNH 30341) partial distal caudal vertebra (AMNH online)
(AMNH 30342) mid caudal vertebra (~29 mm) (AMNH online)
(AMNH 30343) mid caudal vertebra (~30 mm) (AMNH online)
(AMNH 30344) distal caudal vertebra (~32 mm) (AMNH online)
(AMNH 30345) incomplete mid caudal vertebra (~25 mm) (AMNH online)
(AMNH 30346) partial distal caudal vertebra (AMNH online)
(AMNH 30347) partial distal caudal vertebra (AMNH online)
(AMNH 30348) distal caudal vertebra (~31 mm) (AMNH online)
(AMNH 30349) fragmentary distal caudal vertebra (AMNH online)
(AMNH 30350) distal caudal vertebra (~33 mm) (AMNH online)
(AMNH 30351) incomplete distal caudal vertebra (~31 mm) (AMNH online)
(AMNH 30352) partial distal caudal vertebra (AMNH online)
(AMNH 30353) incomplete distal caudal vertebra (AMNH online)
(AMNH 30354) distal caudal vertebra (~23 mm) (AMNH online)
?(AMNH 30355) mid caudal vertebra (~32 mm) (AMNH online)
(AMNH 30356) proximal caudal centrum (~28 mm) (AMNH online)
(AMNH 30357) incomplete proximal caudal vertebra (~25 mm) (AMNH online)
(AMNH 30358) proximal caudal vertebra (~24 mm) (AMNH online)
(AMNH 30359) incomplete proximal caudal centrum (AMNH online)
(IVPP 230790-16; = IVPP 230090-16 of Currie and Eberth, 1993) metatarsal III (Currie and Eberth, 1993)
Comments-
Currie and Eberth (1993) stated "Troodontid bones are rare, but include
distinctive third metatarsals (AMNH 21751, 21772, IVPP 230090-16), in
which the distal articulation extends onto the posterior surfrace of
the bone in a broad tongue." However, Currie and Dong (2001)
corrected the identification of the second specimen, stating "AMNH
21772 is the proximal end of a second metatarsal. It is identified as a
troodontid on the basis of its contact surface for the fourth
metatarsal, its size, and especially its lateromedial
compression." The AMNH online catalogue photo indicates most of
the element is preserved and the locality info is "8 mi. E. of station"
indicating it was found in localities 140-149 in 1923 or 1928.
Currie and Dong describe AMNH 21751 as "two distal ends of third
metatarsals [that] are about the same size and represent left and right
elements. Although they may represent the same individual, the two
fossils are different colours, which suggests they may not have been
found together." They indicate these were "Collected in the 1920s
by the third Central Asiatic Expedition from exposures of the Iren
Dabasu Formation (?Santonian) near Erenhot." Currie and Dong list
IVPP 230790-16 (presumably the correct field number for Currie and
Eberth's 'IVPP 230090-16') as a metatarsal "Collected in 1990 from
exposures of the Iren Dabasu Formation (?Santonian) near Erenhot",
which would make it found during the second Sino-Canadian
expedition. They state "the tongue-like extensions of the third
metatarsals from Iren Dabasu are flat like those of Troodon ... , Borogovia ... , and Tochisaurus" but unlike the grooved surface of Sinornithoides or the distally restricted surface of Philovenator. This has since been identified in Bissekty Urbacodon sp. ZIN PH 2342/16, and it should be noted the extension of Tochisaurus is much shorter, while Mei, IGM 100/44, 100/140 and 100/1126 have a condition like Sinornithoides. Thus as hypothesized by Dong and Currie, at least AMNH 21751 and IVPP 230790-16 are closer to Troodon than Sinornithoides. Currie and Eberth stated "These bones are provisionally referred to Saurornithoides" (at the time a concept including Zanabazar)
without rationale, but Currie and Dong instead classified them as "an
unknown species of troodontid", stating they "cannot be identified
further without additional material."
Makovicky (1995) stated "A probable troodontid axis (AMNH 6570),
articulated with a third cervical vertebra, is present in the
collections of the American Museum of Natural History. This
identification is based on the morphology of the associated third
cervical and a probable fifth cervical, possibly from the same
individual, which strongly resembles those of Troodon.
The axis is from an immature individual as seen from absence of both
the odontoid and axial intercentrum." This specimen number
includes over two hundred paratype Archaeornithomimus
elements from the Kaisen Quarry AMNH locality 140, and the cervicals
described were not recognized in the material catalogued under it in
July 2009 (pers. obs.). However, a small ungual was noticed in
AMNH 6576 (which includes almost a hundred paratype Archaeornithomimus
elements from the Johnson Quarry AMNH locality 141) that most closely
resembles a troodontid pedal ungual I in the slight curvature,
proximally placed flexor tubercle and posterodorsal extent being less
than its posteroventral extent.
The AMNH online catalogue lists AMNH 25570 as "Troodon ?",
consisting of "3 vertebrae." A large number of elements (AMNH
30261-30297, 30300-30318, 30320-30330, 30336-30359) are labeled are
labeled "Troodontid" on the AMNH online catalogue, each from the same
location ("8 mi. E. of station") and from AMNH Quarry 142 specifically
when visible in the photo (AMNH 30265, 30266, 32069, 30300, 30301,
30303, 30320). Given the similar preservation and number of
elements preserved, it is possible these represent two individuals, and
that several other specimens only identified to the level of Saurischia
in the online catalogue (AMNH 30245, 30247-30260, 30298-30299, 30360)
that are also from "8 mi. E. of station" may belong to them as
well. Note AMNH 30267 is incorrectly identified as a proximal
fibula, while 30288 is called "Proximal end of metatarsal IV" but seems
to be two vertebrae instead, AMNH 30301 is called an "Ilium fragment."
but may be a proximal pubis (posterior edge downward in photo), AMNH
30302 is labeled as "Acetabulum fragment." and indeed may be the
ischial peduncle and postacetabular base of a left ilium, AMNH 30304 is
labeled "Prox. end of ischium" but more closely resembles a proximal
troodontid pubis in the diverging peduncles and shallowly concave
acetabular edge, AMNH 30305 is a last sacral vertebra with a convex
posterior central face and 30322 is an anterior dorsal with convex
anterior central face so both may be alvarezsaurid instead.
Scoring the material as photographed in the online catalogue (with AMNH
30305 and 30322 excluded, and 30301 and 30304 interpreted as pubes)
into Hartman et al.'s maniraptoromorph matrix does result in it being
troodontid, but note examination of the specimens themselves would
provide far more data for each element and that it's currently only an
assumption that they belong to the same taxon.
References- Currie and Eberth, 1993. Palaeontology, sedimentology and
palaeoecology of the Iren Dabasu Formation (Upper Cretaceous), Inner Mongolia,
People's Republic of China. Cretaceous Research. 14, 127-144.
Makovicky, 1995. Phylogenetic aspects of the vertebral morphology of Coelurosauria
(Dinosauria: Theropoda). Masters thesis, University of Copenhagen. 311 pp.
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria,
Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences.
38(12), 1753-1766.
Averianov and Sues, 2012. Correlation of Late Cretaceous continental vertebrate
assemblages in middle and central Asia. Journal of Stratigraphy. 36(2), 462-485.
unnamed possible troodontid (Mathur and Srivastava, 1987)
Maastrichtian, Late Cretaceous
Lameta Formation, India
Material- (GSI 19996) tooth (7x4x2 mm)
Comments- This was described by Mathur and Srivastava (1987) as (?) Megalosaurus
type E. It was listed under Troodontidae by Ford (online 2015). The
tooth is strongly recurved and short with subequal (10 per 5 mm), blunt
serrations on the mesial and distal carinae, absent apically. It
differs from Kallamedu Formation tooth DUGF/52 in being narrower, with
a more concave distal edge and serrations absent apically.
References- Mathur and Srivastava, 1987. Dinosaur teeth from Lameta Group
(Upper Cretaceous) of Kheda District, Gujarat. Journal of the Geological Society
of India. 29, 554-566.
Ford, online 2015. http://www.paleofile.com/Dinosaurs/Theropods/Troodonincertae.asp
unnamed troodontid (Goswami, Prasad, Benson, Verma and Flynn, 2012)
Late Maastrichtian, Late Cretaceous
Kallamedu Formation, India
Material- (DUGF/52) tooth (3.5 x 2.8 mm)
References- Goswami, Prasad, Benson, Verma and Flynn, 2012. New vertebrates
from the Late Cretaceous Kallamedu Formation, Cauvery basin, south India, including
a troodontid dinosaur, a gondwanatherian mammal, and a Simosuchus-like
notosuchian crocodyliform. Journal of Vertebrate Paleontology. Program and Abstracts
2012, 102.
Goswami, Prasad, Verma, Flynn and Benson, 2013. A troodontid dinosaur from the
latest Cretaceous of India. Nature Communications. 4, 1703.
Borogovia Osmolska, 1987
B. gracilicrus Osmolska, 1987
Early Maastrichtian, Late Cretaceous
Altan Uul IV, Nemegt Formation, Mongolia
Holotype- (ZPAL MgD-I/174) incomplete tibiotarsi (27 mm wide distally),
proximal fibula, distal metatarsals II, phalanges II-1 (32 mm), phalanges II-2 (13
mm), pedal unguals II (31 mm), distal metatarsal III, distal phalanges III-1, phalanx
III-2 (23 mm), phalanges III-3 (22 mm), pedal ungual III (15 mm), distal metatarsals
IV, phalanges IV-1 (18 mm), phalanges IV-2 (17 mm), phalanges IV-3 (15 mm), phalanges
IV-4 (16 mm), pedal unguals IV (21.5 mm; one distal)
Diagnosis- (after Osmolska,
1987) very slender and long tibiotarsus; second toe with very short
phalanx II-2 and straight ungual; third toe much thinner and weaker
than the second and fourth toes.
Comments- This specimen was discovered in 1971 and first mentioned by Osmolska (1982) as "personal observation on Saurornithoides sp. in the ZPAL collection" and "ZPAL Saurornithoides
sp. specimen from the Nemegt Formation of the Gobi Desert (personal observation)." It may be a junior synonym of Zanabazar as posited by Osmolska
(1987), as the known material of the two doesn't overlap.
References- Osmolska, 1982. Hulsanpes perlei n.g.n.sp. (Deinonychosauria,
Saurisichia, Dinosauria) from the Upper Cretaceous Barun Goyot Formation of
Mongolia. Ne�es Jahrbuch fur Geologie und Palaontologie Monatschefte. 1982(7),
440-448.
Osmolska, 1987. Borogovia gracilicrus gen. et sp. n., a new troodontid
dinosaur from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica.
32, 133-150.
Saurornithoides Osborn, 1924b
= "Ornithoides" Osborn, 1924a
S. mongoliensis Osborn,
1924b
= "Ornithoides oshiensis" Osborn, 1924a
= Troodon mongoliensis (Osborn, 1924b) Paul, 1988
Late Campanian, Late Cretaceous
Bayn Dzak, Djadokhta Formation, Mongolia
Holotype- (AMNH 6516) (1.56 m; 13 kg) incomplete skull (192 mm), three
sclerotic plates, incomplete mandibles, ninth dorsal vertebra, tenth dorsal
vertebra (23 mm), eleventh dorsal vertebra (22 mm), incomplete twelfth dorsal
vertebra (23 mm), (sacrum 147 mm) incomplete first sacral vertebra (23 mm),
partial second sacral centrum (24 mm), partial third sacral centrum (25 mm),
partial fourth sacral centrum (25 mm), partial fifth sacral vertebra (25 mm),
incomplete sixth sacral vertebra (25 mm), incomplete first caudal vertebra (22
mm), incomplete second caudal vertebra (23 mm), partial third caudal vertebra,
partial fourth caudal vertebra, incomplete first chevron, partial second chevron,
partial third chevron, ilial fragment, incomplete pubes, ischia (114 mm), proximal
femur (~198 mm), tibial mold (~243 mm), metatarsal I fragments, phalanx I-1
(16 mm), incomplete pedal ungual I, partial metatarsal II, phalanx II-1 (27.5
mm), phalanx II-2 (14 mm), pedal ungual II (29.3 mm), distal metatarsal III
(~139 mm), phalanx III-1 (~33 mm), phalanx III-2 (24 mm), proximal phalanx III-3,
partial metatarsal IV, phalanx IV-1 (20.5 mm), proximal phalanx IV-2
Late Campanian, Late Cretaceous
Khermeen Tsav,
Baron Goyot Formation, Mongolia
Referred- ?(IGM coll.; 970719 KmT AMRM-3) partial skeleton (Watabe and Suzuki, 2000)
Late Campanian, Late Cretaceous
Ukhaa Tolgod, Djadokhta Formation, Mongolia
?(IGM 100/1083) (1.85 m; 21 kg) maxillary fragment, incomplete
quadrate, incomplete anterior cervical vertebra, partial anterior dorsal vertebra,
sacral fragment, three distal caudal vertebrae, distal tarsal IV, pedal ungual
II (34.7 mm), distal metatarsal III, proximal metatarsal IV, four pedal phalanges
(Norell and Hwang, 2004)
Late Campanian, Late Cretaceous
Wulansuhai Formation (= Bayan Mandahu Formation), Inner Mongolia, China
?(IVPP V10599) partial sacrum, caudal vertebrae, chevrons, pelvis (Currie and
Dong, 2001)
Diagnosis- More derived than Sinornithoides and Sinusonasus
due to the absent promaxillary fenestra, large serrations on teeth, and six
sacral vertebrae. Less derived than Zanabazar and Troodon due
to the dorsal tympanic recess.
Other diagnoses- Osborn (1924b) was the first to publish a diagnosis,
but as the specimen was the first troodontid known from more than teeth, the
generic diagnosis is now far too generalized. The presence of five pairs of
external cranial fenestrae is incorrect, as Osborn did not take the supratemporal
fenestrae into account, but six pairs are primitive for tetanurines anyway.
The external mandibular fenestra is primitive for archosaurs, while the fairly
homodont maxillary series is primitive for an even more inclusive group. The
presence of nineteen premaxillary and maxillary teeth is incorrect, as there
are actually twenty-three. The "less raptorial" teeth are also found
in other maniraptoriforms except dromaeosaurids. Teeth which have only distal
serrations are also now known in several other troodontids (e.g. Sinornithoides,
Sinusonasus, Zanabazar). Maxillary teeth which lack replacement
gaps are also present in Mei, Sinovenator and basal avialans,
so may be plesiomorphic. Of his listed "specific characters", the
posteriorly increasing size of maxillary teeth, closely spaced dentary teeth,
and teeth with a sub-acute tip are common in troodontids. The premaxillary teeth
do not decrease in size posteriorly though, as the first tooth is small, the
second unpreserved, and the third and fourth subequal (Norell et al., 2009).
The illusion of decreasing size is due to the length of the roots which are
exposed. Flattened, recurved teeth are primitive for archosaurs
Norell et al. (2009) listed several characters in their diagnosis, as they differ
from Zanabazar. While Saurornithoides is smaller than Zanabazar,
Norell et al. note all other named troodontids except perhaps Troodon
are as well. Norell et al. also distinguish it from Zanabazar by its
low tooth count (19 maxillary teeth and ~31-33 dentary teeth), but Sinornithoides
(18 maxillary teeth), Sinusonasus (~19 maxillary teeth) and Urbacodon
(32 dentary teeth) are similar. A jugal with a straight ventral edge below the
front of the orbit is also present in Mei and seemingly Sinornithoides,
and is somewhat uncertain in Saurornithoides due to breakage in any case.
The dorsal tympanic recess is plesiomorphic, being present in Byronosaurus
and Sinovenator as well. Norell et al. note their last listed character
(maxillary teeth with only slight posterior increase in height) is also present
in Mei, Sinovenator and basal avialans, so may be plesiomorphic.
Comments- The holotype was discovered on July 9 1923 and initially announced in a magazine article (Osborn, 1924a) as Ornithoides oshiensis,
"a dinosaur birdlike in its skull form", "a name given in allusion to
the fact that the reptile was found in the basin Oshih, with numerous
teeth." A partial sacrum, three mid caudal vertebrae and two
partial ilia were originally included in the holotype, but were
reidentified as protoceratopsian by Norell et al. (2009) and placed in
the new number AMNH 30613. Currie and Peng (1994) described
hindlimb IVPP V10597 as a possible juvenile Saurornithoides mongoliensis,
but this was made the holotype of Philovenator curriei by Xu et al. in 2012. Watabe and Suzuki (2000) mention "a Saurornithoides
partial skeleton" found on July 18 1997 at Khermeen Tsav, given field
number 970719 KmT AMRM-3. It has not been described yet.
Norell and Hwang (2004) described a fragmentary specimen found in 1993
they provisionally referred to Saurornithoides mongoliensis,
as it was identical except for being larger. Norell et al. noted definitive
referral was not possible, though they did state the tooth replacement was more
similar to Saurornithoides than to Byronosaurus. IVPP V10599 was mentioned by Currie and
Dong as being referrable to Saurornithoides mongoliensis
and having a sixth sacral vertebra derived from the caudal series, but
details including a rationale for the referral are lacking. It
was discovered in 1988 as part of the Dinosaur Project, and may be
referrable to the troodontines Papiliovenator or Linhevenator later described from the same formation.
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.
Russell, 1969. A new specimen of Stenonychosaurus from the Oldman Foramtion
of Alberta. Canadian Journal of Earth Sciences. 6, 595-612.
Barsbold, 1974. Saurornithoididae, a new family of small theropod dinosaurs
from Central Asia and North America. Palaeontologia Polonica. 30, 5-22.
Paul, 1988. Predatory Dinosaurs of the World. Simon and Schuster Co., New York.
464 pp.
Osmolska and Barsbold, 1990. Troodontidae. In Weishampel, Dodson, and Osmolska
(eds). The Dinosauria, Berkeley: University of California Press. 259-268.
Currie and Peng, 1994. A juvenile specimen of Saurornithoides mongoliensis
from the Upper Cretaceous of northern China. Canadian Journal of Earth Sciences.
30(10), 2224-2230.
Watabe and Suzuki, 2000. Report on the Japan - Mongolia Joint
Paleontological Expedition to the Gobi desert, 1997. Hayashibara Museum
of Natural Sciences Research Bulletin. 1, 69-82.
Currie and Dong, 2001. New information on Cretaceous troodontids (Dinosauria,
Theropoda) from the People's Republic of China. Canadian Journal of Earth Sciences.
38(12), 1753-1766.
Norell and Hwang, 2004. A troodontid dinosaur from Ukhaa Tolgod (Late Cretaceous
Mongolia). American Museum Novitates. 3446, 9 pp.
Norell, Makovicky, Bever, Balanoff, Clark, Barsbold and Rowe, 2009 online. Saurornithoides mongoliensis, Digital Morphology. http://digimorph.org/specimens/Saurornithoides_mongoliensis/
Norell, Makovicky, Bever, Balanoff, Clark, Barsbold and Rowe, 2009. A review
of the Mongolian Cretaceous dinosaur Saurornithoides (Troodontidae: Theropoda).
American Museum Novitates. 3654, 63 pp.
Zanabazar Norell, Makovicky,
Bever, Balanoff, Clark, Barsbold and Rowe, 2009
= "Mongolodon" Franzosa, 2004
Z. junior (Barsbold, 1974) Norell, Makovicky, Bever, Balanoff,
Clark, Barsbold and Rowe, 2009
= Saurornithoides junior Barsbold, 1974
= "Mongolodon" junior (Barsbold, 1974) Franzosa, 2004
Early Maastrichtian, Late Cretaceous
Bugin Tsav, Nemegt Formation, Mongolia
Holotype- (IGM 100/1) (2.28 m, 27 kg, adult) skull (280.6 mm), anterior
mandibles, (sacrum- 200 mm) first sacral vertebra (~34 mm), second sacral vertebra
(~31 mm), third sacral vertebra (31 mm), fourth sacral vertebra (31 mm), fifth
sacral vertebra (36 mm), sixth sacral vertebra (36.2 mm), proximal caudal vertebra
(~31.5 mm), proximal caudal vertebra (33 mm), proximal caudal vertebra (32 mm),
proximal caudal vertebra (32 mm), proximal caudal vertebra (33 mm), proximal
caudal vertebra (32.8 mm), mid caudal vertebra (31.5 mm), mid caudal vertebra
(34 mm), mid caudal vertebra (37 mm), distal caudal vertebra (38 mm), distal
caudal vertebra (38 mm), distal caudal vertebra (39 mm), distal caudal vertebra
(39.5 mm), distal caudal vertebra (39.5 mm), two proximal chevrons, mid chevron,
five distal chevrons, distal tibia (68 mm wide), astragalocalcaneum, distal
tarsal III, distal tarsal IV, proximal metatarsal II, proximal metatarsal III,
proximal metatarsal IV
Late Cretaceous
Mongolia
Referred- ?(IGM 100/2) postcrania (Barsbold, 1983)
Diagnosis- (after Norell et al., 2009) exoccipital forms large part of
posttemporal fenestra (unknown in other troodontids except Troodon); deep paroccipital
processes, with proximodorsal edge above foramen magnum.
Other diagnoses- Barsbold (1974) originally only listed "large Saurornithoides with 20 maxillary and 35 dentary teeth" in
his diagnosis. The large size is also seen in Troodon. The maxillary
tooth count (19-20) overlaps Saurornithoides (19) and Sinusonasus
(~19), while the dentary tooth count (35) is the same as Troodon.
Norell et al. (2009) also listed several other characters in their diagnosis. The absent
dorsal tympanic recess is shared with Troodon. They note the foramen
magnum is more transversely compressed than Troodon, but that Byronosaurus
and Sinovenator are similar, making it plesiomorphic. Similarly, the
lack of a separate canal for the ophthalmic branch of the trigeminal nerve is
also seen in Byronosaurus and most dromaeosaurids, meaning it is plesiomorphic
as well.
Comments- The holotype was discovered in 1964 and initially described by Barsbold (1974)as a new species of Saurornithoides, from the earlier Djadochta Formation. Contrary to Barsbold, Norell et al. note the teeth lack
mesial serrations, the caudal vertebrae are probably not a continuous series,
and the distal fibula is unpreserved. However, they also fail to note three
proximal caudal vertebrae in their materials list and description. Franzosa
(2004) assigned this species to a new genus "Mongolodon" without comment
in his unpublished thesis. However, Norell et al. (2009) later redescribed the
species and made it the type of their new genus Zanabazar. This was largely
done because of the absence of characters supporting a sister relationship with
Saurornithoides mongoliensis, though Norell et al. do not explicitly
support a closer relationship of either species to Troodon either. Borogovia
may be a junior synonym (Osmolska, 1987), as its holotype cannot be compared
to IGM 100/1. Tochisaurus is not a junior synonym however (Kurzanov and
Osmolska, 1991), though this was proposed as possible by Osmolska.
Barsbold (1983) states "besides the holotype, remains of the
postcranial skeleton of another specimen, no. 100/2" can be referred to
the taxon, but it has not been mentioned in other references.
Norell et al. (2009) even state "the holotype remains the only known
specimen of this species" with Barsbold as a coauthor.
References- Barsbold, 1974. Saurornithoididae, a new family of small
theropod dinosaurs from Central Asia and North America. Palaeontologia Polonica.
30, 5-22.
Barsbold, 1983. Carnivorous dinosaurs from the Cretaceous of Mongolia. Transactions
of the Joint Soviet-Mongolian Paleontological Expedition. 19, 5-119.
Osmolska, 1987. Borogovia gracilicrus gen. et sp. n., a new troodontid
dinosaur from the Late Cretaceous of Mongolia. Acta Palaeontologica Polonica.
32, 133-150.
Osmolska and Barsbold, 1990. Troodontidae. In Weishampel, Dodson, and
Osmolska (eds.). The Dinosauria. University of California Press.
259-268.
Kurzanov and Osm�lska, 1991. Tochisaurus nemegtensis gen. et sp.
n., a new troodontid (Dinosauria, Theropoda) from Mongolia. Acta Palaeontologia
Polonica. 36, 69-76.
Franzosa, 2004. Evolution of the Brain in Theropoda (Dinosauria). PhD Thesis.
The University of Texas at Austin. 357 pp.
Norell, Makovicky, Bever, Balanoff, Clark, Barsbold and Rowe, 2009 online. Zanabazar junior, Digital Morphology. http://digimorph.org/specimens/Zanabazar_junior/
Norell, Makovicky, Bever, Balanoff, Clark, Barsbold and Rowe, 2009. A review
of the Mongolian Cretaceous dinosaur Saurornithoides (Troodontidae: Theropoda).
American Museum Novitates. 3654, 63 pp.
Papiliovenator Pei, Qin, Wen, Zhao, Wang, Liu, Guo, Liu, Ye, Wang, Yin, Dai and Xu, 2021
P. neimengguensis Pei, Qin, Wen, Zhao, Wang, Liu, Guo, Liu, Ye, Wang, Yin, Dai and Xu, 2021
Late Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype- (BNMNH-PV030) (subadult) incomplete skull (~120
mm), incomplete mandible, hyoids, atlantal neural arch, axis, partial
third cervical vertebra, partial fourth cervical vertebra, two
fragmentary mid cervical vertebrae, partial eighth cervical vertebra,
fragmentary ninth cervical vertebra, partial tenth cervical vertebra,
incomplete first dorsal vertebra, second dorsal vertebra, incomplete
third dorsal vertebra, partial fourth dorsal vertebra, few fragmentary
dorsal ribs, partial scapulae, partial coracoid, incomplete
humeri (~90 mm), incomplete radius, partial ulna, phalanx I-1, manual
ungual I, phalanx II-1, fragmentary pelvis, incomplete femur, partial
tibia,
incomplete fibula, partial metatarsal II, incomplete phalanx II-1,
partial phalanx II-2, pedal unguals II, incomplete metatarsal III,
incomplete metatarsal IV
Diagnosis- (after Pei et al.,
2021) lateral groove of dentary not posteriorly expanded; deep
surangular fossa hosting the surangular foramen anteroventral to
glenoid fossa; ventral ridge of surangular fossa mainly on
surangular; anterolaterally broadened and butterfly-shaped neural
arches of the first and second dorsal vertebrae in dorsal view.
Comments- This was discovered
in 2018. Pei et al. (2021) used a version of the TWiG analysis to
recover it as a troodontid sister to Byronosaurus, Gobivenator, Xixiasaurus and troodontines. In the Hartman et al. maniraptoromorph matrix it is the sister of Zanabazar, with a position similar to Pei et al.'s less parsimonious by two steps. Forcing it to be sister to Linhevenator or Philovenator from the same formation requires four more steps each.
Reference- Pei, Qin, Wen, Zhao,
Wang, Liu, Guo, Liu, Ye, Wang, Yin, Dai and Xu, 2021 online. A new
troodontid from the Upper Cretaceous Gobi basin of Inner Mongolia,
China. Cretaceous Research. Journal Pre-proof. DOI:
10.1016/j.cretres.2021.105052
Troodon Leidy, 1856
?= Polyodontosaurus Gilmore, 1932
?= Stenonychosaurus Sternberg, 1932
?= Pectinodon Carpenter, 1982
?= Latenivenatrix van der Reest and Currie, 2017
T. formosus Leidy, 1856
?= Laelaps cristatus Cope, 1876
?= Dryptosaurus cristatus (Cope, 1876) Hay, 1902
?= Deinodon cristatus (Cope, 1876) Osborn, 1902 non (Marsh, 1892) Hay,
1902
?= Dromaeosaurus cristatus (Cope, 1876) Matthew and Brown, 1922
?= Polyodontosaurus grandis Gilmore, 1932
?= Stenonychosaurus inequalis Sternberg, 1932
?= Saurornithoides inequalis (Sternberg, 1932) Carpenter, 1982
= Stegoceras formosum (Leidy, 1856) Olshevsky, 1991
?= Troodon cristatus (Cope, 1876) Olshevsky, 1995
?= Troodon inequalis (Sternberg, 1932) Snively and Russell, 2002
?= Latenivenatrix mcmasterae van der Reest and Currie, 2017
Late Campanian, Late Cretaceous
Judith River Formation, Montana, US
Holotype- (ANSP 9259) posterior premaxillary tooth
Referred- (AMNH 3954; syntypes of Laelaps cristatus) two maxillary
teeth (11 mm)
(AMNH 8519) tooth (Sahni, 1972)
(AMNH 8520) tooth (3.4 mm) (Sahni, 1972)
(AMNH 21760) tooth (AMNH online)
(ANSP 15937) dentary tooth (Fiorillo and Currie, 1994)
(ANSP 15947) maxillary tooth (Fiorillo and Currie, 1994)
(ANSP 15950) maxillary tooth (Fiorillo and Currie, 1994)
(ANSP 15964) dentary tooth (Fiorillo and Currie, 1994)
(ANSP 17642) premaxillary tooth (Fiorillo and Currie, 1994)
(ANSP 17780) premaxillary tooth (Fiorillo and Currie, 1994)
(ANSP 17795) premaxillary tooth (Fiorillo and Currie, 1994)
(ANSP 18005) maxillary tooth (Fiorillo and Currie, 1994)
(MOR 170) vertebra (MOR online)
(MOR 320) tooth (MOR online)
(MOR 993) (embryo) tooth, centrum, limb elements, partial egg (Varricchio and
Jackson, 2004)
Late Campanian, Late Cretaceous
Oldman Formation of the Judith River Group, Alberta, Canada
(RTMP 89.77.5) tooth (Ryan and Russell, 2001)
(RTMP 94.157.1) eggshells (Zelenitsky, Modesto and Currie, 2002)
(RTMP 94.157.2) eggshells (Zelenitsky, Modesto and Currie, 2002)
(RTMP 94.157.4) eggshells (Zelenitsky, Modesto and Currie, 2002)
(RTMP 94.157.5) eggshells (Zelenitsky, Modesto and Currie, 2002)
(RTMP 138.319) tooth (Ryan, 2003)
(RTMP coll.) tooth (Chiba, Ryan, Braman, Eberth, Scott, Brown, Kobayashi and
Evans, 2015)
teeth (Ryan and Russell, 2001)
(embryos and adults) material (Zelenitsky, Modesto and Currie, 2002)
Late Campanian, Late Cretaceous
Dinosaur Park Formation of the Judith River Group, Alberta, Canada
(AMNH 6174; Latenivenatrix morph) frontals (61.5 mm), parietals, laterosphenoid (Russell, 1969)
(AMNH 21598) tooth (AMNH online)
(AMNH 21714) tooth (AMNH online)
(CMN 199) distal tibia (58.6 mm wide), astragalus (Russell, 1969)
(CMN 1267) premaxillary tooth (Lambe, 1902)
(CMN 1650) distal manual phalanx, pedal ungual I (52 mm), phalanx II-1 (47.6
mm), phalanx II-2 (24.8, 24.6 mm), pedal ungual II (~70 mm), pedal ungual IV
(~45 mm) (Russell, 1969)
(CMN 2506) pedal phalanx II-1 (Russell, 1969)
(CMN 8539; holotype of Stenonychosaurus inequalis) six distal caudal
vertebrae, metacarpal I (36.5 mm), distal phalanx I-1, distal metacarpal II,
partial phalanx II-1, distal manual phalanx, distal tibia, astragalus, metatarsal
I, phalanx I-1 (~29 mm), pedal ungual I (~45 mm), metatarsal II (202.4 mm),
phalanx II-1 (~50.5 mm), phalanx II-2 (29 mm), pedal ungual II, metatarsal III
(253.6 mm), phalanx III-1 (~66 mm), phalanx III-2 (41.2 mm), phalanx III-3 (39.3
mm), pedal ungual III (~55 mm), metatarsal IV (244.5 mm), phalanx IV-1 (~34.2
mm), phalanx IV-2 (~28.3 mm), phalanx IV-3 (~23.6 mm), phalanx IV-4 (~24.5 mm),
pedal ungual IV (~47 mm) (Sternberg, 1932)
(CMN 8540; holotype of Polyodontosaurus grandis; Latenivenatrix morph) dentary (117.5 mm) (Gilmore,
1932)
(CMN 12340; holotype of Latenivenatrix mcmasterae)
postorbital, frontals (60.8 mm), parietals, basioccipital,
basisphenoid, dorsal centrum fragment, four dorsal ribs, seven
gastralia, three distal caudal vertebra fragments, three chevrons,
proximal radius, ulnae (one distal) (131 mm), semilunate carpal,
incomplete phalanx II-1, manual ungual, incomplete femur, astragali,
metatarsal I, phalanx I-1 (26.3 mm), pedal ungual I (46.5 mm),
metatarsal II fragments, phalanx II-1 (46.7 mm), pedal ungual II (65,
66 mm), metatarsal III fragments, phalanx III-1, phalanx III-3 (32.8
mm), pedal ungual III, metatarsal IV fragments, phalanx IV-1 (33.5 mm),
phalanx IV-2 (27.3 mm), pedal ungual IV (~43 mm) (Russell, 1969)
(CMN 12355) frontal (Sues, 1978)
(CMN 12392) anterior maxilla, fragments (Russell, 1969)
(CMN 12425) astragalus (58.4 mm wide) (Russell, 1969)
(CMN 12433) ulna (138 mm) (Russell, 1969)
(CMN 12434) pedal phalanx II-1 (Russell, 1969)
(CMN coll.) distal metacarpal I (?), astragalus, calcaneum(?), distal metatarsal
I (?), pedal ungual II (Lambe, 1902)
(ROM 1445) partial dentary, second dentary tooth, thirteenth dentary tooth,
twentieth dentary tooth (Russell, 1948)
(RTMP 65.23.32) maxillary tooth (10 mm) (Currie, Rigby and Sloan, 1990)
(RTMP 67.14.39; = PMAA P67.14.39; Latenivenatrix morph) partial dentary (Sues, 1977)
(RTMP 79.8.1; Latenivenatrix morph) frontals (58.4 mm), parietals, laterosphenoid (Currie, 1985)
(RTMP 79.8.635) (juvenile) posterior dentary tooth (4 mm) (Currie, 1987a)
(RTMP 79.8.1171) tooth (Baszio, 1997)
(RTMP 80.16.1473) parietals (Currie, 1985)
(RTMP 80.16.1478; mistyped RTMP 80.16.1748 in van der Reest and Currie, 2017; Latenivenatrix morph) incomplete frontals (~140 mm), mesethmoid (Currie, 1985)
(TMP 1981.016.0231) incomplete frontal (Currie, 1992)
(RTMP 81.22.66) (Currie, 1985)
(RTMP 81.37.15) tenth cervical vertebra (Makovicky, 1995)
(RTMP 82.16.124) frontals, parietals (Currie, 1985)
(RTMP 82.16.138) partial dentary (Currie, 1987a)
(RTMP 82.16.282) premaxillary tooth (Currie, 1987a)
(RTMP 82.19.23; Latenivenatrix morph) lacrimal, postorbitals, squamosals, frontals, parietals, braincase
(Currie, 1985)
(RTMP 82.19.151; mistyped RTMP 86.19.151 in van der Reest and Currie, 2017; Stenonychosaurus strat) partial dentary (Currie, 1987a)
(RTMP 82.20.259) premaxillary tooth (Currie, 1987a)
(RTMP 83.12.11; Latenivenatrix morph) incomplete dentary (Currie, Rigby and Sloan, 1990)
(RTMP 83.36.214) maxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 83.36.215) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 83.45.7) maxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 83.45.8) tooth (Currie, Rigby and Sloan, 1990)
(RTMP 84.65.1) distal metatarsal II, incomplete metatarsal III, distal metatarsal
IV (Wilson and Currie, 1985)
(RTMP 85.6.3) premaxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 85.6.186) maxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 86.36.4; Latenivenatrix morph?) frontals, parietals (Currie, 1987b)
(RTMP 86.36.457; Stenonychosaurus strat) incomplete braincase (Currie and Zhao, 1993)
(RTMP 86.49.10; Stenonychosaurus morph) frontal (62.4 mm) (Currie, 1987b)
(RTMP 86.54.66) premaxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 86.78.40; Stenonychosaurus morph) frontal (Evans et al., 2017)
(RTMP 86.177.8) maxillary tooth (Sankey, Brinkman, Guenther and Currie, 2002)
(RTMP 88.50.58) tooth (Baszio, 1997)
(RTMP 88.50.88; Stenonychosaurus morph) partial frontals, partial parietals (van der Reest and Currie, 2017)
(RTMP 88.96.2) maxillary tooth (Currie, Rigby and Sloan, 1990)
(RTMP 89.36.268) tooth (Ryan and Russell, 2001)
(RTMP 89.76.50) tooth (Baszio, 1997)
(RTMP 89.77.5) tooth (Baszio, 1997)
(RTMP 89.89.4) tooth (Baszio, 1997)
(RTMP 89.116.63) tooth (Baszio, 1997)
(RTMP 90.34.1) tooth (Baszio, 1997)
(RTMP 91.36.690; Stenonychosaurus morph?) frontal (van der Reest and Currie, 2017)
(RTMP 92.36.575; Latenivenatrix
morph) incomplete dentary (~160 mm), partial tibia, metatarsal II,
distal metatarsal III (258.8 mm), metatarsal IV (266.2 mm), metatarsal
V (74.1 mm) (Rauhut, 2003)
(RTMP 92.36.1212) (adult) posterior cervical vertebra (Makovicky, 1995)
(RTMP 93.36.86; Latenivenatrix morph) frontal (62.5 mm) (Evans et al., 2017)
(RTMP 94.12.438) third dorsal vertebra (Makovicky, 1995)
(RTMP 97.133.8; Latenivenatrix morph) metatarsal III (van der Reest and Currie, 2017)
(RTMP 98.68.90; Stenonychosaurus morph) metatarsal III (van der Reest and Currie, 2017)
(RTMP 98.93.1; Latenivenatrix morph) frontal (55.1 mm) (Evans et al., 2017)
(UALVP 5282; Stenonychosaurus morph) incomplete frontal (Russell, 1969)
(UALVP 5284) distal metatarsal III (Russell, 1969)
(UALVP 52611; Stenonychosaurus morph) frontals (60.5 mm), parietals (Evans et al., 2017)
(UALVP 55285; Latenivenatrix morph?) frontal (van der Reest and Currie, 2017)
(UALVP 55804; Latenivenatrix strat) sacrum (207.4 mm), ilia (one partial; ~308 mm), partial pubes (van der Reest and Currie, 2017)
(YPM PU 23414; Latenivenatrix morph) parietals (Currie, 1985)
teeth (Brinkman, 1990)
Campanian, Late Cretaceous
Two Medicine Formation, Montana, US
(MOR 246) nest (Horner and Weishampel, 1988)
....(MOR 246-1) (embryo) teeth, two cervical centra (4.4 mm), cervical neural
arch, sacral centrum (5.1 mm), scapula (14.7 mm), fragmentary coracoid, humerus
(20.3 mm), partial tibiae, partial fibula, egg
....(MOR 246-2) (embryo) humerus, egg
....(MOR 246-3) egg
....(MOR 246-5) (embryo) bone, egg
....(MOR 246-7) (embryo) bone, egg
....(MOR 246-8) (embryo) femur, egg
....(MOR 246-9) (embryo) bone, egg
....(MOR 246-10) (embryo) bone, egg
....(MOR 246-11) (embryo) (skull ~50 mm) maxilla (~18 mm), quadrate (12.1 mm),
basioccipital, teeth (1 mm), vertebrae, ilia, pubes (~23 mm), ischial fragments,
femur (34.9 mm), tibiae (46 mm), partial fibula, metatarsal II, metatarsal III,
metatarsal IV (33.5 mm)
....(MOR 246-12) egg
....(MOR 246-13) (embryo) bone, egg
....(MOR 246-14) egg
....(MOR 246-15) egg
....(MOR 246-16) (embryo) bone, egg
....(MOR 246-17) (embryo) bone, egg
....(MOR 246-18 (embryo) bone, egg
(MOR 247) egg (Varricchio, Horner and Jackson, 2002)
(MOR 299) eggs (Varricchio, Horner and Jackson, 2002)
(MOR 323) tooth (MOR online)
(MOR 363-7-4-83-1) nest, twenty-two eggs (Varricchio, Jackson, Borkowski and
Horner, 1997)
(MOR 393) nest, twenty-two eggs (Varricchio, Horner and Jackson, 2002)
(MOR 430) (several month old juvenile) partial skeleton including premaxilla,
caudal vertebrae, femur (126 mm), tibia (173 mm), metatarsal IV (105 mm) (Currie,
Rigby and Sloan, 1990)
(MOR 493) metatarsals, pedal phalanges (MOR online)
(MOR 510) tooth (MOR online)
(MOR 511) teeth (MOR online)
(MOR 512) teeth (MOR online)
(MOR 513) teeth (MOR online)
(MOR 553) anterior dorsal vertebra, distal caudal vertebrae (Makovicky, 1995)
(MOR 553-1) metatarsal II, metatarsal III, metatarsal IV (Varricchio, 1993)
(MOR 553-2) metatarsal (Varricchio, 1997)
(MOR 553-6.21.9) fourth cervical vertebra (Makovicky, 1995)
(MOR 553-7.7.91.19) tibia (Zanno et al., 2011)
(MOR 553-7.16.0.61) femur (Varricchio, 1993)
(MOR 553-7.21.92.46) (adult) ninth or tenth cervical vertebra (Makovicky, 1995)
(MOR 553-7.24.8.64) tibia (425 mm) (Varricchio, 1993)
(MOR 553-7.28.91.236) tenth cervical vertebra (Makovicky, 1995)
(MOR 553-7.20.91.120) anterior dorsal vertebra (Makovicky, 1995)
(MOR 553-7.23.91.36) first caudal vertebra (Makovicky, 1995)
(MOR 553-8.3.92.141) (adult) eighth cervical vertebra (Makovicky, 1995)
(MOR 553-8.8.92.186) first dorsal vertebra (Makovicky, 1995)
(MOR 553-8.10.92.203) (adult) twelfth dorsal vertebra (Makovicky, 1995)
(MOR 553-8.11.92.204) third cervical vertebra (Makovicky, 1995)
(MOR 553-8.12.92.222) (juvenile) eighth or ninth cervical vertebra (Makovicky,
1995)
(MOR 553-8.19.92.212) fourth cervical vertebra (Makovicky, 1995)
(MOR 553-8.20.92.305) (adult) sixth or seventh dorsal vertebra (Makovicky, 1995)
(MOR 553D) partial sacrum (Makovicky, 1995)
(MOR 553L-7.25.89.313) pedal ungual II (Wolff, Varricchio and Hanna, 2015)
(MOR 553L-7.27.8.87) ischium (Hutchinson, 2001)
(MOR 553S) material including more precise specimens below and the following
which may be those specimens- four humeri (129.4, 143.4, 146.7, 178.7 mm), pubis,
ischium, five femora (218.0, 233.0, 238.0, 273.0, 302.0 mm), six tibiae (283.0, 317.0, 323.0,
361.0, 382.0, 431.0 mm) and three metatarsals III (114.3, 124.2, 130.7 mm) (Carrano,
1998)
(MOR 553S-7.1.9.9) metatarsal IV (Zanno et al., 2011)
(MOR 553S-7.8.91.28) metatarsal II (Zanno et al., 2011)
(MOR 553S-7.16.0.61) (adult) femur (Zanno et al., 2011)
(MOR 553S-7.18.92.5) metatarsal II (Zanno et al., 2011)
(MOR 553S-7.20.91.120) dorsal vertebra (Wolff, Varricchio and Hanna, 2015)
(MOR 553S-7.21.92.47) maxilla (MOR online)
(MOR 553S-7.28.8.102) metatarsal IV (Zanno et al., 2011)
(MOR 553S-7.28.91.236) cervical vertebra (Wolff, Varricchio and Hanna, 2015)
(MOR 553S-7.28.91.239) (adult) femur (Zanno et al., 2011)
(MOR 553S-7.29.92.113) metatarsal III (Zanno et al., 2011)
(MOR 553S-8.2.91.303) humerus (MOR online)
(MOR 553S-8.2.92.131) braincase (MOR online)
(MOR 553S-8.3.9.387) pubis (Zanno et al., 2011)
(MOR 553S-8.3.92.141) cervical vertebra (Wolff, Varricchio and Hanna, 2015)
(MOR 553S-8.12.92.219) ulna (Zanno et al., 2011)
(MOR 553S-8.13.92.237) metacarpal II (Wolff, Varricchio and Hanna, 2015)
(MOR 553S-8.17.92.260) metatarsal IV (Wolff, Varricchio and Hanna, 2015)
(MOR 553S-8.17.92.265) fibula (Zanno et al., 2011)
(MOR 553S-8.20.92.305) mid dorsal vertebra (Zanno et al., 2011)
(MOR 553S-8.20.92.311) astragalus (Zanno et al., 2011)
(MOR 553S-8.28.92.270) mid dorsal vertebra (Zanno et al., 2011)
(MOR 553S-8.69.406) metatarsal III (Zanno et al., 2011)
(MOR 553S-11.1.01.1) astragalus (Zanno et al., 2011)
(MOR 553S-11.1.01.8; = MOR 553S 1.11.01.8 of Wolff, Varricchio and Hanna, 2015?)
metatarsal IV (Zanno et al., 2011)
(MOR 553S-92.260) metatarsal IV (Zanno et al., 2011)
(MOR 558) braincase, three cervical vertebrae, elements (MOR online)
(MOR 563) (year old juvenile) skeleton including tibia, metatarsal III (Varricchio,
1993)
?(MOR 564) (embryo) maxilla (MOR online)
?(MOR 584) partial skeleton (MOR online)
(MOR 615) eggs (MOR online)
(MOR 646) jaw fragment (MOR online)
(MOR 675) eggs (Varricchio, Horner and Jackson, 2002)
(MOR 676) two nests including eleven and eight eggs (Varricchio, Horner and
Jackson, 2002)
(MOR 702) two eggs (Varricchio and Jackson, 2004)
(MOR 721) specimen including femur (156.9 mm) and tibia (224 mm) (Carrano, 1998)
(MOR 748) (12 year old adult) nine proximal caudal vertebrae, eight distal caudal
vertebrae, partial pelvis, femur (320 mm), tibia (362 mm), fibula, astragalus,
metatarsal I, metatarsal II, metatarsal III (206.5 mm), metatarsal IV (227 mm),
four pedal phalanges, nest, at least ten eggs (Varricchio, Jackson, Borkowski
and Horner, 1997)
(MOR 750) eggs (Varricchio, Horner and Jackson, 2002)
(MOR 796) five vertebrae, partial ribs (MOR online)
(MOR 963) nest, twenty-four eggs (Varricchio, Jackson, Borkowski and Horner,
1997)
(MOR 964) eggs (MOR online)
(MOR 1006) partial egg (MOR online)
(MOR 1115) twenty-four eggs (MOR online)
(MOR 1138) eggs (MOR online)
(MOR 1139) eggs (Varricchio, Horner and Jackson, 2002)
(MOR 1170) tibia (MOR online)
(MOR coll.) caudal series (Wolff, Varricchio and Hanna, 2015)
(YPM PU 22445) (YPM online)
(YPM PU 23246-23248) (YPM online)
(YPM PU 22544) (Hirsch and Quinn, 1990)
(YPM PU 22594) (Hirsch and Quinn, 1990)
(YPM PU 23259) (YPM online)
(YPM PU 23408) (YPM online)
(YPM PU 23409) (YPM online)
teeth (Redman, Moore and Varricchio, 2015)
Diagnosis- (after Currie, 1987a; compared to Saurornithoides and
Zanabazar) round anterior border of antiorbital fenestra; sculpturing less
extensive on nasal process of maxilla; postorbital region of skull longer; no
sulcus between parasphenoid capsule and rectangular platform between basipterygoid
processes; mesethmoid located further anteriorly; very strong, caudoventrally
shifted basal tubera; otic recess extends further posteroventrally; dentary
symphysis more extensive; mesial serrations extend to tip of carina on maxillary
teeth.
(after van der Reest and Currie, 2017; for Latenivenatrix, but unknown in Stenonychosaurus)
17� retroverted pubis; anteriorly curving pubic shaft; large muscle
scar on lateral surface of pubic shaft slightly proximal to pubic boot.
Other diagnoses- Leidy's (1856) original description of Troodon
included both generalized theropod characters ("compressed, curved,
conical crown with trenchant edges ... outer side is more convex than
the inner") and those now known to be present in other troodontines
("trenchant edges are coarsely denticulated; the denticulations
themselves being compressed conical, with trenchant edges, and are bent
in such a manner that their apices are directed towards the summit of
the crown").
Ornithischian or theropod?- Troodon was originally discovered in October 1855 and identified
as a lizard by Leidy (1856) based on the holotype tooth, but reidentified as
a theropod by Cope (1877). Nopcsa (1901) and Hay (1902) placed it in Megalosauridae.
Brown (1908) placed it in Ankylosauridae, while Gilmore (1924) believed it was
a pachycephalosaurid and a senior synonym of Stegoceras. Thus pachycephalosaurs
were classified under Gilmore's new family Troodontidae until Sternberg (1945)
placed Troodon back in Theropoda and named Pachycephalosauridae. Russell
(1948) described a partial dentary (ROM 1445) of Troodon, assigning it
to Troodontidae within the Theropoda. A further confusion with ornithischians
occured when Galton (1983) suggested "Laosaurus" minimus should
be assigned to Troodon and that the latter is a carnivorous ornithopod
(first suggested by Baird, 1981). Galton cited pers. comm. with Horner alluding
to undescribed Two Medicine Formation material. The latter turned out to be
adults of the ornithopod Orodromeus (Horner and Weishampel, 1988) associated
with Troodon teeth, eggs and embryos (Horner and Weishampel, 1996). The
idea of Troodon as a carnivorous ornithopod was never established in
the technical literature, but was widespread in the popular literature in the
1980's. Instead, it was almost always assigned to Theropoda after 1945.
Laelaps cristatus- Cope (1876) described two teeth as a new species
of Laelaps (in which he placed all Judithian theropods). Once Marsh provided
the replacement name Dryptosaurus for the preoccupied Laelaps,
Hay (1902) moved cristatus to that genus. Osborn (1902) meanwhile referred
it to Deinodon, though this is not the same taxon as Hay's (1902) Deinodon
cristatus, which was a new combination of Aublysodon cristatus, a
tyrannosaurid premaxillary tooth. Matthew and Brown (1922) questionably referred
the species to their new genus Dromaeosaurus, probably based on size,
for it was the smallest of their 'deinodontids'. Olshevsky (1995) most recently
referred it to Troodon, though most workers have ignored it. This tooth
can indeed be identified as Troodon by its large serrations (2/mm distally,
and slightly smaller mesially), presence of mesial serrations and absence of
longitudinal ridges. The size (FABL 6 mm) and BW/FABL (.50) are also comparable
to Troodon maxillary teeth (Currie et al., 1990), as is the height/FABL
(1.83). The serrations are smaller than in premaxillary teeth, which is why
Cope kept it separate from Troodon (at the time only known from the type
premaxillary tooth). The tooth is larger than dentary teeth, posterior examples
of which also differ in lacking mesial serrations. As it is from the same formation
as T. formosus and lacks distinguishing characters, it is referred to
that species here.
Polyodontosaurus grandis- Sternberg collected a dentary (CMN 8540)
from the same formation as Leidy's specimen, which Gilmore (1932) described
as a new genus of lizard- Polyodontosaurus. Sternberg later (1951) assigned
this genus to Troodontidae, noting it may be congeneric with Troodon.
The latter synonymization was formalized by Romer in 1966, who assigned both
to the Coeluridae. On the other hand, Russell (1969) synonymized Polyodontosaurus
with Stenonychosaurus inequalis in the Troodontidae.
Stenonychosaurus and Saurornithoides- Sternberg (1932)
named Stenonychosaurus inequalis as a coelurid, based on a fragmentary
skeleton (CMN 8539). Sternberg later (1951) suggested Stenonychosaurus
and Troodon might be synonymous, but no comparable elements were known
which could prove this. Russell (1969) described several specimens found in
1968 as Stenonychosaurus inequalis, assigning it to the Troodontidae
along with Saurornithoides and Troodon. He did not synonymize
Troodon with Stenonychosaurus or Saurornithoides due to
the fragmentary condition of the former. Ostrom (1969) assigned both Saurornithoides
and Stenonychosaurus to the Dromaeosauridae based on their raptorial
second pedal digit, though perhaps deserving subfamilial status. Barsbold (1974)
noted differences between the holotype of Troodon and teeth of Saurornithoides,
while feeling the cranial and postcranial resemblences between Stenonychosaurus
and Saurornithoides justified a close relationship. He assigned the latter
two genera to his new family Saurornithoididae, while retaining Troodon
in Troodontidae. This was followed until Currie (1987a). Sues (1977) described
two new dentaries as saurornithoidids, though he was uncertain if they belonged
to Stenonychosaurus or Saurornitholestes (which was assigned to
Dromaeosauridae once he described it the next year). Stenonychosaurus
was generally used for the Dinosaur Park troodontid through the 1980's (e.g.
Currie, 1985), though Carpenter (1982) synonymized it with Saurornithoides,
as Saurornithoides inequalis. This was done without justification however,
referencing a Carpenter and Paul in prep. publication which never emerged. Paul
did later (1988) synonymize Saurornithoides with Troodon (including
Stenonychosaurus), but this is a subjective decision which has not been
accepted by later authors.
Estes (1964) originally referred several teeth from the Lance Formation of Wyoming
to Saurornithoides sp.. Carpenter (1982) named Pectinodon bakkeri
from the same formation, believing some of Estes' specimens to be referrable
to that species, some to his new combination Saurornithoides inequalis,
and others to an unnamed third taxon. He assigned Pectinodon to the Saurornithoididae.
Currie's breakthrough- Currie (1987a) cleared up the American troodontid
situation, determining the differences between known dentaries were ontogenetic,
and dental differences between Troodon, ROM 1445, Saurornithoides
and the Lance Formation specimens were largely due to variation within the tooth
row. Troodon's holotype is a premaxillary tooth, the teeth of Saurornithoides
which Barsbold compared were maxillary (as were Carpenter's Saurornithoides
inequalis examples), Pectinodon was based on posterior dentary teeth,
while ROM 1445 contains a few dentary teeth. Currie described intermediates
between all of these morphologies, although he noted the maxillary mesial serrations
are more extensive in Troodon than Saurornithoides, extending
to the tip of the tooth. He synonymized Stenonychosaurus inequalis and
Polyodontosaurus grandis with Troodon formosus, and provisionally
did the same for Pectinodon bakkeri. This has remained the consensus,
along with the assignment of Saurornithoides, Troodon and related
taxa to the Troodontidae.
Overlumped?- Recently the concept of all Campanian-Maastrichtian North
American troodontids being one taxon has been questioned. Currie et al. (1990)
noted that though Horseshoe Canyon Formation troodontid teeth are essentially
identical to those from the Judith River Group/Formation, teeth from the Frenchman,
Hell Creek, Lance, Prince Creek and Scollard Formations are different and may
prove to be separate species. Baszio (1997) elaborated, describing differences
between teeth from the Judith River, Horseshoe Canyon and Scollard Formations.
Olshevsky (1991) created the new combination Troodon bakkeri for the
Lance Formation material, while several papers have used the combination Troodon
inequalis for the Dinosaur Park Formation taxon (Snively, 2002; Osmolska,
2004; Currie, 2005). Currie (2005) stated that it is more conservative to retain
inequalis for Dinosaur Park specimens, but keeping taxa separate based
on a political boundary (Montana vs. Alberta) has no biological value.
Morhardt
et al. (2013) and Evans et al. (2017) both describe cranial differences between
Dinosaur Park and Horseshoe Canyon specimens, Evans et al. naming frontals from
the latter formation Albertavenator curriei. Sankey et al. (2002) have
split Dinosaur Park Formation troodontids into Troodon formosus and cf.
Troodontidae indet., with the latter being a rare form only present in
some formations and distinguished by various dental characters. If they are
correct, most of the non-dental specimens (including the Stenonychosaurus
holotype and CMN 12340) could not be definitively assigned to either taxon,
and many poorly or undescribed Troodon teeth could end up not being referrable
to that genus. Conversely, Evans et al. (2017) analyzed teeth morphometrically
and recovered the Troodon holotype very close to an apparent premaxillary
tooth from the Horseshoe Canyon Formation, which may indicate premaxillary teeth
of Albertavenator cannot be distinguished from Troodon and thus
the latter is indeterminate. They note the frontals can be distinguished, and
that CMN 12340 allows connecting frontal morphology with the supposedly diagnostic
pedal anatomy of Stenonychosaurus inequalis. This could support a valid
Stenonychosaurus and Albertavenator. However, there is only stratigraphic
evidence to connect any teeth with Albertavenator, so it's also possible
e.g. that particular Hoseshoe Canyon tooth is Troodon, and that the three
widely separated premaxillary teeth in the lower left of their figure 6j are
Albertavenator.
Most recently, van der Reest and Currie (2017) have used frontal and
metatarsal III differences correlated with stratigraphy to divide
Dinosaur Park troodontids into smaller, earlier Stenonychosaurus inequalis
(diagnosis- L-shaped frontal with a flat shallowly anteroposteriorly
rippled nasofrontal contact; convex anterior surface of metatarsal III)
and larger, later Latenivenatrix mcmasterae
(diagnosis- triangular frontal with a single deep groove in the
frontonasal contact surface; concave anterior surface of metatarsal
III) with CMN 12340 as the holotype. Further noted differences
are anteriorly expanded parietal sagittal crest and anteriorly limited
medial recurvature of dentary in Stenonychosaurus. In addition to their specimen identifications, RTMP 86.49.10 has a Stenonychosaurus
morphology (Currie, 1987b), while RTMP 67.14.39 (Sues, 1977), RTMP
83.12.11 (Evans et al., 2017), RTMP 93.36.86 (Evans et al., 2017) and
YPM PU 23414 (Currie, 1985) have a Latenivenatrix morphology. Notably, this does not allow any teeth including the Troodon holotype to be referred to either taxon, nor any described material from the Judith River Formation of Montana where T. formosus' holotype is from. Also, the holotype dentary of Polyodontosaurus grandis was reported to be placed stratigraphically high in the Latenivenatrix
range (Gilmore, 1932) although van der Reest and Currie bring up
potential sources for error. Its extensive and strong medial
curvature also matches Latenivenatrix more than Stenonychosaurus, but this is also present in e.g. ?Albertavenator and Zanabazar so is not an autapomorphy, and the Stenonychosaurus dentary is only referred to that taxon based on stratigraphy. So there is evidence Polyodontosaurus is a senior synonym of Latenivenatrix,
but this is uncertain. A final factor is the identity of the Two
Medicine Formation troodontid, which has yet to be described in
detail. van der Reest and Currie (2017) report that it and Talos share the anteriorly convex distal metatarsal III with Stenonychosaurus, unlike Latenivenatrix, Linhevenator and Gobivenator. The skull roof and dentary are not yet described. If this taxon is described as Stenonychosaurus or a new taxon, it's likely Troodon will be broken down on this site into several genera with the incomparable Dinosaur Park specimens and current T? sp. being made into unnamed/undescribed/indet. Troodontinae.
Troodon eggs and embryos- Horner (1982) first reported ten nests
in the Two Medicine Formation which he ascribed to ornithopods, "closely
allied to the Hypsilophodontidae". Horner and Weishampel later (1988) described
a new genus of ornithopod from the site, Orodromeus makelai. They believed
abundant nest with eggs and embryos belonged to Orodromeus, though this
was disproven by Horner and Weishampel (1996) due to the small size of adult
Orodromeus and the reidentification of an embryo as Troodon. Zelenitsky
and Hillis (1996) named eggs from the Oldman Formation of Alberta Prismatoolithus
levis (Prismatoolithidae), while Zelenitsky (2000) identified the Two Medicine
Formation Troodon eggs as belonging to this oospecies as well. Most of
the publications dealing with Troodon since that time have concerned
these nests, eggs and embryos.
One confusing aspect is that Continuoolithus canadensis eggs were provisionally
assigned to Troodon by Horner (1984), based on an embryo. This was followed
by Hirsch and Quinn (1990) and Horner (1994), until 1996. Horner (1997) found
the embryo to be indeterminate, while Varricchio and Jackson (2004) found the
eggshell to be theropod. The precise identity of Continuoolithus is still
unknown.
Canadian therizinosaur frontals?- Sues (1978) identified frontal CMN 12349 as Dromaeosaurus
and frontal CMN 12355 as Theropoda indet., although note his plates 7
and 8 are switched so that they are given each others' captions. Currie
(1987b) accepted CMN 12349 as Dromaeosaurus, but with regard to CMN 12355 stated that
"comparison with Mongolian specimens suggests that it may represent Erlicosaurus (Currie, in preparation)." Currie (1992) again stated CMN 12355 "may represent the segnosaurid Erlicosaurus"
(although note that as in Sues' paper it is mislabeled 12349 in his
Figure 2) and also referred TMP 1981.016.0231 to Segnosauridae. Currie
(2005) listed CMN 12349 as a tentative "therizinosauroid similar to Erlikosaurus", but figured CMN 12355 so probably meant that specimen. Indeed, it seems CMN 12349 was correctly identified as Dromaeosaurus
in the first place as it is similar to the holotype and has only been
referred to Therizinosauroidea accidentally due to Sues' original plate
caption mistake. Larson et al. (2014) performed a morphometric
analysis of Dinosaur Park coelurosaur frontals, and found that CMN
12355 grouped with Troodon, so is troodontid instead. Yet without including a therizinosaur such as Erlikosaurus
itself, the study only had so much explanatory power. More recently,
Cullen et al. (2020) expanded the analysis to include both Erlikosaurus
and
the Bissekty therizinosaurid, still recovering CMN 12355 within the
Dinosaur Park troodontid range and far from therizinosaurids. Both it
and the less complete TMP 1981.016.0231 are here referred to Troodon formosus sensu lato.
Reinterpreted records- The supposed Struthiomimus manual ungual in plate XV
figure 10-11 of Lambe (1902) is not ornithomimid and may be a Troodon
pedal ungual II instead. Found with this and apparently similar unguals and
manual phalanges were an astragalus, calcaneum, pedal phalanges and two elements
identified by Lambe as distal ends of metacarpal I and metatarsal I. A metatarsal
I would exclude ornithomimids from consideration, but a calcaneum would exclude
troodontids. It's probable some material was incorrectly associated or misidentified.
Though Britt (1993) described the posterior cervical vertebrae RTMP 81.37.15 and 92.36.1212
as ornithomimids, Makovicky (1995) indicated they were actually Troodon.
Although Currie et al. (1990) reported Troodon
from the Milk River Formation, Baszio (1997) notes the ROM specimens noted were
collected in the Milk Creek area, but probably in the Judith River Group. A
large sample of theropod teeth from the Milk River Formation did not include
Troodon.
Rowe et al. (1992) and Sankey (1997, 1998) identified Troodon in the
Aguja Formation of Texas based on teeth (TMM 43057-323 and LSUMG 140:6117 respectively),
but these were later identified as pachycephalosaurian by Sankey (2001). This
would mean Troodon is unknown from the Aguja Formation, except that Montellano
et al. (2009) later reported cf. Troodon teeth in an abstract. Whether
these are truly troodontid awaits proper description.
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131, 123-168.
Hutchinson, 2001. The evolution of femoral osteology and soft tissues on the
line to extant birds (Neornithes). Zoological Journal of the Linnean Society.
131, 169-197.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
Sankey, 2001. Late Campanian southern dinosaurs, Aguja Formation, Big Bend,
Texas. Journal of Vertebrate Paleontology. 75(1), 208-215.
Varricchio, 2001. "Beautiful wounding tooth": ontogeny and osteology
in the theropod Troodon formosus. Journal of Vertebrate Paleontology.
21(3), 110A.
Varricchio, Horner and Jackson, 2002. Embryos and eggs for the Cretaceous theropod
Troodon formosus. Journal of Vertebrate Paleontology. 22, 564-576.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Snively and Russell, 2002. Kinematic model of tyrannosaurid (Dinosauria: Theropoda)
arctometatarsus function. 255(2), 215-227.
Zelenitsky, Modesto and Currie, 2002. Bird-like characteristics of troodontid
theropod eggshell. Cretaceous Research. 23, 297-305.
Ryan, 2003. Taxonomy, systematics and evolution of centrosaurine ceratopsids
of the Campanian Western Interior basin of North America. Unpublished PhD thesis.
Department of Biological Sciences, Calgary, Alberta. 578pp.
Osmolska, 2004. Evidence on relation of brain to endocranial cavity in oviraptorid
dinosaurs. Aacta Paleontologica Polonica. 49(2), 321-324.
Varricchio and Jackson, 2004. Two eggs sunny-side up: reproductive physiology
in the dinosaur Troodon formosus. in Currie, Koppelhus, Shugar and Wright
(eds.). Feathered Dragons: Studies on the Transition from Dinosaurs to Birds.
Indiana University Press, Bloomington. pp 215-233.
Varricchio and Jackson, 2004. A phylogenetic assessment of prismatic dinosaur
eggs from the Cretaceous Two Medicine Formation of Montana. Journal of Vertebrate
Paleontology. 24(4), 931-937.
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.
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.
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.
Grellet-Tinner, Chiappe, Norell and Bottjer, 2006. Dinosaur eggs and nesting
behaviors: A paleobiological investigation. Palaegeography, Palaeoclimatology,
Palaeoecology. 232, 294-321.
Jackson, Horner and Varricchio, 2009. A study of a Troodon egg containing
embryonic remains using epifluorescence micriscopy and other techniques. Journal
of Vertebrate Paleontology. 29(3), 121A.
Montellano, Monroy, Hernandez-Rivera and Torres, 2009. Late Cretaceous microvertebrate
fauna from the Northern state of Coahuila, Mexico. Journal of Vertebrate Paleontology.
29(3), 151A.
Jackson, Jackson, Varricchio and Zelenitsky, 2010. Uncovering theropod eggs:
Water vapor conductance and nesting strategy of Troodon. Journal of Vertebrate
Paleontology. Program and Abstracts 2010, 110A.
Porfiri, Calvo and dos Santos, 2011. A new small deinonychosaur (Dinosauria:
Theropoda) from the Late Cretaceous of Patagonia, Argentina. Anais da Academia
Brasileira de Ci�ncias. 83(1), 109-116.
Zanno, Varricchio, O’Connor, Titus and Knell, 2011. A new troodontid theropod,
Talos sampsoni gen. et sp. nov., from the Upper Cretaceous western interior
basin of North America. PLoS ONE. 6(9), e24487.
Morhardt, Ridgely, Varricchio and Witmer, 2013. New studies of braincase anatomy,
brain size, and brain structure in the Late Cretaceous theropod Troodon formosus
(Dinosauria: Saurischia) based on CT scanning and 3D visualization. Journal
of Vertebrate Paleontology. Program and Abstracts 2013, 180.
Varricchio, 2013. Wounding tooth grows up: Ontogeny in the Cretaceous theropod
Troodon formosus. Journal of Vertebrate Paleontology. Program and Abstracts
2013, 230.
Larson, Cullen, Todd and Evans, 2014. Geometric morphometrics of small theropod
frontals from the Dinosaur Park Formation, Alberta. Journal of Vertebrate Paleontology.
Program and Abstracts 2014, 165.
Varricchio, Jackson and Jin, 2014. Lay-brood-repeat: Nesting site fidelity in
ecologic time for two Cretaceous troodontid dinosaurs. Journal of Vertebrate
Paleontology. Program and Abstracts 2014, 246.
Chiba, Ryan, Braman, Eberth, Scott, Brown, Kobayashi and Evans, 2015. Taphonomy
of a monodominant Centrosaurus apertus (Dinosauria: Ceratopsia) bonebed
from the upper Oldman Formation of southeastern Alberta. Palaios. 30, 655-667.
Germano and Varricchio, 2015. Taphonomic description of three recently discovered
Troodon clutches from Egg Mountain. Journal of Vertebrate Paleontology.
Program and Abstracts 2015, 131.
Redman, Moore and Varricchio, 2015. A new vertebrate microfossil locality in
the Upper Two Medicine Formation in the vicinity of Egg Mountain. Journal of
Vertebrate Paleontology. Program and Abstracts 2015, 201-202.
Templeman, Moore, Atudorei and Varricchio, 2015. Stable isotope evidence for
dinosaur ecology from Campanian eggshell at the Egg Mountain locality, western
Montana, USA. Journal of Vertebrate Paleontology. Program and Abstracts 2015,
224.
Varricchio, Jin and Jackson, 2015. Lay, brood, repeat: Nest reuse and site fidelity
in ecologic time for two Cretaceous troodontid dinosaurs. Journal of Vertebrate
Paleontology. 35(3), e932797. DOI: 10.1080/02724634.2014.932797
Wolff, Varricchio and Hanna, 2015. Initial work on the cursorial pathology of
Troodon formosus. Journal of Vertebrate Paleontology. Program and Abstracts
2015, 240.
Evans, Cullen, Larson and Rego, 2017. A new species of troodontid theropod (Dinosauria:
Maniraptora) from the Horseshoe Canyon Formation (Maastrichtian) of Alberta,
Canada. Canadian Journal of Earth Sciences. 54, 813-826.
van der Reest and Currie, 2017. Troodontids (Theropoda) from the Dinosaur Park
Formation, Alberta, with a description of a unique new taxon: Implications for
deinonychosaur diversity in North America. Canadian Journal of Earth Sciences.
54, 919-935.
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.
T? bakkeri (Carpenter, 1982) Olshevsky,
1991
= Pectinodon bakkeri Carpenter, 1982
Late Maastrichtian, Late Cretaceous
Lance Formation, Wyoming, US
Holotype- (UCM 38445; holotype of Pectinodon bakkeri) tooth (6.2x3.7x?
mm)
Paratypes- (UCM 38446) (juvenile) tooth (1.8x2x? mm)
(UCMP 73098) (juvenile) tooth (2.8x1.8x? mm)
(UCMP 125239; was part of UCMP 73098) (juvenile) tooth (3.2x2.5x? mm)
Referred- (SDSM 12456) four dentary teeth (Whitmore, 1988)
(SDSM 12458) maxillary tooth (Whitmore, 1988)
(SDSM 15098) three dentary teeth (Whitmore, 1988)
(SDSM 15099) maxillary tooth (Whitmore, 1988)
(UA 156) tooth (Baszio, 1997)
(UA 157) tooth (Baszio, 1997)
(UCM 41666) (juvenile) anterior dentary (Carpenter, 1982)
(UCM 43218) (juvenile) basioccipital (Carpenter, 1982)
(UCMP 84990) tooth (UCMP online)
(UCMP 125240; = UCMP 124402) tooth (Carpenter, 1982)
(UCMP 125241; = UCMP 124394) tooth (Carpenter, 1982)
(UCMP 125242; = UCMP 124394) tooth (Carpenter, 1982)
(UCMP 125243; = UCMP 124394) tooth (Carpenter, 1982)
(UCMP 125244; = UCMP 124394) tooth (Carpenter, 1982)
(UCMP 125245; = UCMP 124394) tooth (Carpenter, 1982)
(UCMP 125246; = UCMP 124394) tooth (Carpenter, 1982)
(UCMP 125247; = UCMP 124394) tooth (Carpenter, 1982)
(UCMP 186864) teeth (UCMP online)
(UCMP 186886) teeth (UCMP online)
(UCMP 186916) tooth (UCMP online)
(UCMP 186979) tooth (UCMP online)
(UCMP 187050-187056) seven teeth (UCMP online)
(UCMP 187058-187079) twenty-two teeth (UCMP online)
(UCMP 187180) tooth (UCMP online)
(UCMP 187181) tooth (UCMP online)
?(UCMP 189686) ungual (UCMP online)
(UCMP 214059) tooth (UCMP online)
(YPM 54491) (YPM online)
(YPM 55041) (YPM online)
(YPM 55521) (YPM online)
(YPM 55536) (YPM online)
(YPM 55545) (YPM online)
(YPM 55546) (YPM online)
(YPM 55583) (YPM online)
(YPM 55584) (YPM online)
(YPM 55591) (YPM online)
(YPM 55592) (YPM online)
(YPM 55627) (YPM online)
teeth (Derstler, 1995)
(juvenile) elements, eggshells (Derstler, 1995)
teeth (Spencer, Turner and Chadwick, 2001)
Late Maastrichtian, Late Cretaceous
Lance Formation, Montana, US
(CM 30748) partial humerus, ulna, femora, tibia (McIntosh,
1981)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, US
(UCMP 364730; = UCMP 556335) tooth (UCMP online)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, South Dakota, US
teeth (DePalma, 2010)
Other diagnoses- (after
Carpenter, 1982) crowns of teeth strongly compressed labiolingually and
recurved; mesial margin without serrations or sharp, translucent
carina; mesial edge usually rounded, but may have low, blunt, opaque
carina; distal margin with large serrations having translucent edges;
serrations largest in middle, and may be subequal to crown's tip in
size; distal serrations perpendicular to vertical axis of tooth; crown
tip directed distally, almost parallel to crown base; crown tip does
not function as a piercing tip, but as first serration; first
definitive serration occurs immediately below crown tip and differs
from other deinonychosaurs in that it is not significantly smaller than
crown tip; near tooth's base, two small serrations are crowded
together; FABL of tooth almost equal to height.
Comments- Estes (1964) referred teeth to Saurornithoides sp.,
but these seem to belong to a different species of troodontid (cf. Troodontidae
indet. of Sankey et al., 2002). Carpenter (1982) described several teeth
as Pectinodon bakkeri, believing them to be distinct from Judith River
troodontids, though Currie showed this was due to their position in the jaw
(posterior dentary). Virtually no Lance Formation troodontid material has been
described except this material, which Currie (1987) felt was indistinguishable
from Judith River Troodon, though Currie et al. (1990) stated they were
somewhat different. Derstler (1995) considered his juvenile material and eggshells
to belong to a new taxon of troodontid. Olshevsky (1991) created the new combination
Troodon bakkeri for Lance Formation troodontid teeth, which is used here
because the basal tubera are large and the dentary symphysis seems more robust
than Zanabazar, though not as much as in T. formosus (due to ontogeny?). All Lance Formation troodontid material is listed here.
References- Estes, 1964. Fossil vertebrates from the Late Cretaceous
Lance Formation, eastern Wyoming. University of California Publications in Geological
Sciences. 49, 1-180.
McIntosh, 1981. Annotated catalogue of the dinosaurs (Reptilia, Archosauria)
in the Collections of Carnegie Museum of Natural History. Bulletin of the Carnegie
Museum of Natural History. 18, 1-67.
Carpenter, 1982. Baby dinosaurs from the Late Cretaceous Lance and Hell Creek
formations and a description of a new species of theropod. Contributions to
Geology, University of Wyoming. 20(2), 123-134.
Currie, 1987. Bird-like characteristics of the jaws and teeth of troodontid
theropods (Dinosauria, Saurischia). Journal of Vertebrate Paleontology. 7, 72-81.
Whitmore, 1988. The vertebrate paleontology of Late Cretaceous (Lancian) localities
in the Lance Formation, Northern Niobrara County, Wyoming. Masters
Thesis. South Dakota School of Mines and Technology.
130 pp.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.
Olshevsky, 1991. A Revison of the Parainfraclass Archosauria Cope, 1869, Excluding
the Advanced Crocodyila. Mesozoic Menanderings #2 (1st printing). iv + 196pp.
Derstler, 1995. The Dragons’ Grave - an Edmontosaurus bonebed containing
theropod egg shells and juveniles, Lance Formation, (Uppermost Cretaceous),
Niobrara County, Wyoming. Journal of Vertebrate Paleontology. 15(3), 26A.
Baszio, 1997. Investigations on Canadian dinosaurs: Systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Spencer, Turner and Chadwick, 2001. A remarkable vertebrate assemblage from
the Lance Formation, Niobrara County, Wyoming. GSA Annual Meeting and Exposition
Abstracts. 33(6), A-197.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
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.
T? sp. (Currie, Rigby and Sloan, 1990)
Campanian, Late Cretaceous
Prince Creek Formation, Alaska, US
Material- (AK83-V-095) maxillary or dentary tooth (Fiorillo and Gangloff,
2000)
(AK138-V-128) (subadult) partial braincase (Fiorillo et al., 2009)
(AK233-V-054) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK282-V-001) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK282-V-010) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK282-V-052) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK282-V-056) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK283-V-017) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK283-V-115) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK284-V-024) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK285-V-008) premaxillary tooth (Fiorillo and Gangloff, 2000)
(AK285-V-013) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK285-V-037a) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK299-V-134) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK300-V-017) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK300-V-021) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK300-V-042) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK300-V-055) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK300-V-060) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK300-V-129) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK335-V-012FT) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK335-V-076) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK382-V-015) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK382-V-105) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK383-V-018) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK383-V-137) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK383-V-140) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK383-V-176) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK383-V-183) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK385-V-001) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK385-V-002) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK387-V-000FT) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK387-V-017) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK388-V-002) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK388-V-082) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK392-V-007) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK459-V-011) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK498-V-001) tooth (Fiorillo, 2008)
(AK490-V-004) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK490-V-008FL) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK490-V-086FT) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK491-V-143) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK491-V-168) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK497-V-002) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK498-V-001) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK498-V-002) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK498-V-003) maxillary or dentary tooth (Fiorillo and Gangloff, 2000)
(AK coll.) thirty-three teeth (Fiorillo, 2008)
(DMNH 22158) (adult) partial braincase (Fiorillo et al., 2009)
(UCMP 140624; lost) three teeth (UCMP online)
skull fragments (Gangloff, 1998)
Comments- The two braincases were described as T. formosus, apparently
identical to that species except for some pneumatic features, which are known
to exhibit marked individual and symmetry variation. These teeth have not been
described in detail, though Currie et al. (1990) noted they were somewhat different
than Judith River Troodon teeth, and Sankey et al. (2002) said that no
examples of their cf. Troodontidae indet. were present in this formation. Fiorillo
(2008) found the teeth to be larger on average than T. formosus (9.78
mm in length vs. 4.96 mm), but indentical in length vs. FABL. The Alaskan teeth
range from 5.4-14.3 mm in length and 4.3-9 mm in FABL.
References- Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith
River Formation of southern Alberta, Canada. in Carpenter and Currie (eds.).
Dinosaur Systematics: Perspectives and Approaches. Cambridge University Press,
New York. pp. 107-125.
Nelms, 1992. Paleoecological implications of a North Slope Dinosaurian assemblage.
International Conference on Arctic Margins, Abstracts. 42.
Gangloff, 1998. Arctic Dinosaurs with Emphasis on the Cretaceous Record of Alaska
and the Eurasian-North American Connection. Kirkland, Lucas and Estep (eds).
Lower and Middle Cretaceous Terrestrial Ecosystems. New Mexico Museum of Natural
History and Science, Bulletin. 14, 211-220.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation
(Cretaceous) of Northern Alaska, with speculations on Arctic dinosaur paleoecology.
Journal of Vertebrate Paleontology. 20(3), 41A.
Fiorillo and Gangloff, 2000. Theropod teeth from the Prince Creek Formation
(Cretaceous) of Northern Alaska, with speculations on Arctic dinosaur paleoecology.
Journal of Vertebrate Paleontology. 20(4), 675-682.
Sankey, Brinkman, Guenther and Currie, 2002. Small theropod and bird teeth from
the Late Cretaceous (Late Campanian) Judith River Group, Alberta. Journal of
Paleontology. 76(4), 751-763.
Fiorillo, 2008. On the occurence of exceptionally large teeth of Troodon
(Dinosauria: Saurischia) from the Late Cretaceous of Northern Alaska. Palaios.
23, 322-328.
Fiorillo, Tykoski, Currie, McCarthy and Flaig, 2009. Description of two partial
Troodon braincases from the Prince Creek Formation (Upper Cretaceous),
North Slope Alaska. Journal of Vertebrate Paleontology. 29(1), 178-187.
T? sp. (Ryan and Russell, 2001)
Late Campanian, Late Cretaceous
Wapiti Formation, Alberta, Canada
(RTMP 89.55.1008) metatarsal (Ryan and Russell, 2001)
(RTMP 2004.23.3) posterior premaxillary or anterior maxillary tooth (Fanti and
Miyashita, 2009)
(UALVP 48749) tooth (5.5x4x2.2 mm) (Torices et al., 2014)
(UALVP 48750) posterior dentary tooth (3.7x3.7x2 mm) (Fanti and Miyashita, 2009)
(UALVP 48753) anterior maxillary tooth (10.8x7.5x3.5 mm) (Fanti and Miyashita,
2009)
(UALVP 48755) premaxillary tooth (4.8x3.1x3.1 mm) (Fanti and Miyashita, 2009)
(UALVP 48757) tooth (8.9x5.9x3.6 mm) (Torices et al., 2014)
(UALVP 48764) tooth (10.8x6.5x3.8 mm) (Torices et al., 2014)
(UALVP 48765) tooth (8.5x5.8x3 mm) (Torices et al., 2014)
(UALVP 48776) tooth (10.5x7.1x3.7 mm) (Torices et al., 2014)
(UALVP 48777) tooth (9.4x7.5x3.6 mm) (Torices et al., 2014)
(UALVP 48779) tooth (11x6.5x3.8 mm) (Torices et al., 2014)
(UALVP 48780) tooth (10.2x6.7x4.8 mm) (Torices et al., 2014)
(UALVP 48785) tooth (8.4x6.3x3.3 mm) (Torices et al., 2014)
(UALVP 52599) tooth (Fanti, Currie and Burns, 2015)
(UALVP 53514) tooth (Fanti, Currie and Burns, 2015)
eighteen teeth (Fanti and Miyashita, 2009)
Comments- Torices et al. (2014) could not statistically distinguish teeth
from the Wapiti, Dinosaur Park or Horseshoe Canyon Formations.
References- Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive
of Aves). in Tanke and Carpenter (eds.). Mesozoic Vertebrate Life: New Research
Inspired by the Paleontology of Philip J. Currie. Indiana University Press,
Bloomington, Indiana. pp. 279-297.
Fanti and Miyashita, 2009. A high latitude vertebrate fossil assemblage from
the Late Cretaceous of west-central Alberta, Canada: Evidence for dinosaur nesting
and vertebrate latitudinal gradient. Palaeogeography, Palaeoclimatology, Palaeoecology.
275, 37-53.
Torices, Funston, Kraichy and Currie, 2014. The first appearance of Troodon
in the Upper Cretaceous site of Danek Bonebed, and a reevaluation of troodontid
quantitative tooth morphotypes. Canadian Journal of Earth Sciences. 51(11),
1039-1044.
Fanti, Currie and Burns, 2015. Taphonomy, age, and paleoecological implication
of a new Pachyrhinosaurus (Dinosauria: Ceratopsidae) bonebed from the
Upper Cretaceous (Campanian) Wapiti Formation of Alberta, Canada. Canadian Journal
of Earth Sciences. 52(4), 250-260.
T? sp. (Langston, 1975)
Maastrichtian, Late Cretaceous
St. Mary River Formation, Alberta, Canada
(CMN 10649) tooth (Langston, 1975)
(CMN 10674) partial tooth (Langston, 1975)
References- Langston, 1975. The ceratopsian dinosaurs and associated
lower vertebrates from the St. Mary River Formation (Maestrichtian) at Scabby
Butte, Southern Alberta. Canadian Journal of Earth Science. 12, 1576-1608.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
T? sp. nov. (Baszio, 1997)
Late Maastrichtian, Late Cretaceous
Scollard Formation, Alberta, Canada
(RTMP 94.106.1) tooth (Ryan and Russell, 2001)
(UA 109) tooth (Baszio, 1997)
teeth (Ryan and Russell, 2001)
Comments- Baszio (1997) reported UA 109 was similar to Judith River T.
formosus in size, but differed in having larger mesial serrations (up to
three times wider than distal serrations).
References- Baszio, 1997. Investigations on Canadian dinosaurs: systematic
palaeontology of isolated dinosaur teeth from the Latest Cretaceous of south
Alberta, Canada. Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan and Russell, 2001. The dinosaurs of Alberta (exclusive of Aves). in Tanke
and Carpenter (eds.). Mesozoic Vertebrate Life: New Research Inspired by the
Paleontology of Philip J. Currie. Indiana University Press, Bloomington, Indiana.
pp. 279-297.
T? sp. (Estes, Berberian and Mesozoely, 1969)
Late Maastrichtian, Late Cretaceous
Hell Creek Formation, Montana, South Dakota, US
(FMNH PR2901) tooth (5.9x2.8x1.3 mm) Gates, Zanno and Makovicky, 2015)
(MCZ 3694) tooth (Estes, Berberian and Mesozoely, 1969)
(UCMP 186856) tooth (UCMP online)
(UCMP 186868) teeth (UCMP online)
(UCMP 186873) tooth (UCMP online)
(UCMP 186885) teeth (UCMP online)
(UCMP 186904) tooth (UCMP online)
(UCMP 187057) tooth (UCMP online)
(UCMP 187168-187174) seven teeth (UCMP online)
(UCMP 187178) tooth (UCMP online)
(UCMP 187184) tooth (UCMP online)
(UCMP 187186) tooth (UCMP online)
(UCMP 214060-214062) three teeth (UCMP online)
teeth (Stenerson and O'Conner, 1994)
material (Triebold and Russell, 1995; Triebold, 1997)
material (Jacobson and Sroka, 1998)
four teeth (Larson, Nellermoe and Gould, 2003)
teeth and elements (DePalma, 2010)
Comments- Estes et al. (1969) referred MCZ 3694 to Coeluridae, while
Stenerson and O'Conner (1994) called their teeth Saurornithoides sp..
None have been described yet, though the UCMP specimens were identified as Troodon
by Sankey. Gates et al. (2015) noted FMNH PR2901 resembles Pectinodon
except for its prominent mesial keel.
References- Estes, Berberian and Mesozoely, 1969. Lower vertebrates from
the Late Cretaceous Hell Creek Formation, McCone County, Montana. Breviora.
337, 1-33.
Stenerson and O'Conner, 1994. The Late Cretaceous Hell Creek Formation of Northwestern
South Dakota and its Fauna. MAPS Digest. 17(4), 108-120.
Triebold and Russell, 1995. A new small dinosaur locality in the Hell Creek
Formation: Journal of Vertebrate Paleontology. 15(3), 57A.
Triebold, 1997. The Sandy Site: Small Dinosaurs from the Hell Creek Formation
of South Dakota. in Wolberg, Stump and Rosenberg (eds). Dinofest International,
Proceedings of a Symposium sponsered by Arizona State University. A Publication
of The Academy of Natural Sciences. 245-248.
Jacobson and Sroka, 1998. Preliminary assement of a Hell Creek Dinosaurian Fauna
from sites in Corson County, South Dakota. Journal of Vertebrate Paleontology.
18(3), 53A.
Larson, Nellermoe and Gould, 2003. A study of theropod teeth from a low-species-density
hadrosaur bone bed in the Lower Hell Creek Formation in Carson, Co., S.D.. Journal
of Vertebrate Paleontology. 23(3), 70A-71A.
DePalma, 2010. Geology, taphonomy, and paleoecology of a unique Upper Cretaceous
bonebed near the Cretaceous-Tertiary boundary in South Dakota. Masters thesis,
University of Kansas. 227 pp.
Gates, Zanno and Makovicky, 2015. Theropod teeth from the upper Maastrichtian
Hell Creek Formation "Sue" Quarry: New morphotypes and faunal comparisons.
Acta Palaeontologica Polonica. 60(1), 131-139.
T? sp. (Stokosa, 2005)
Maastrichtian, Late Cretaceous
Fox Hills Formation, South Dakota, US
Material- (KUVP 67265) (two individuals) (~1.3 m) incomplete femur (~160 mm), incomplete tibia (Brewer, 2016)
(~.85 m) incomplete femur (~100 mm), incomplete tibia (Brewer, 2016)
(SDSM 14518) dentary tooth (Stokosa, 2005)
Comments- SDSM 14518 was identified by Stoksad (2005) as Troodon cf.
formosus. Its macroscopic structure has not been described.
KUVP 67265 was found in 1980 and "tentatively identified as Troodon formosus in the absence of skull material" by Brewer (2016) in an SVP abstract.
References- Stokosa 2005. Enamel microstructure variation within the Theropoda. In Carpenter (ed.). The Carnivorous Dinosaurs. 163-178.
Brewer, 2016. New specimens of Troodon formosus hind limbs from the Fox Hills Formation of South Dakota. Journal of Vertebrate Paleontology. Program and Abstracts, 104.
T? sp. (Wroblewski, 1998)
Late Maastrichtian, Late Cretaceous
Ferris Formation, Wyoming, US
Reference- Wroblewski, 1998. Changing paleoenvironments and paleofaunas
across the K-T boundary, Ferris Formation, Southcentral Wyoming. Tate Geological
Museum, Casper College, Casper Wyoming. Tate ’98. Life in the Cretaceous.
53-70.
T? sp. (Hall, 1991)
Late Campanian, Late Cretaceous
Fossil Forest Member of the Fruitland Formation, New Mexico, US
Material- (KUVP 96932) tooth
References- Hall, 1991. Lower vertebrate paleontology of the upper Fruitland
Formation, Fossil Forest area, New Mexico, and implications for Late Cretaceous
terrestrial biostratigraphy. Masters Thesis. University of Kansas. 126 pp.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
T? sp. nov. (Wel and Williamson, 2000; described by Williamson
and Brusatte, 2014)
Late Maastrichtian, Late Cretaceous
Naashoibito Member of Ojo Alamo Formation, New Mexico, US
Material- (NMMNH P-22566; = UNM FKK-014) tooth (6.8x5.3x2.6 mm) (Williamson
and Brusatte, 2014)
(NMMNH P-32746) tooth (Williamson and Brusatte, 2014)
(NMMNH P-32772) tooth (8.4x6x3.3 mm) (Williamson and Brusatte, 2014)
(NMMNH P-32784) tooth (Williamson and Brusatte, 2014)
(NMMNH P-33520) tooth (?x?x3.1 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33521) tooth (3.9x3.3x1.9 mm) (Williamson and Brusatte, 2014)
(NMMNH P-33901) tooth (Williamson and Brusatte, 2014)
(SMP VP-3341) tooth (Jasinski, Sullivan and Lucas, 2011)
Comments- Williamson and Brusatte (2014) found these to be statistically
different from Dinosaur Park specimens.
References- Weil and Williamson, 2000. Diverse Maastrichtian terrestrial
vertebrate fauna of the Naashoibito Member, Kirtland Formation (San Juan Basin,
New Mexico) confirms “Lancian” faunal heterogeneity in western North
America. Geological Society of America Abstracts with Programs. 32, A-498.
Williamson, 2001. Dinosaurs from microvertebrate sites in the Upper Cretaceous
Fruitland and Kirtland Formations, San Juan Basin, New Mexico. 2001 GSA abstracts.
Jasinski, Sullivan and Lucas, 2011. Taxonomic composition of the Alamo Wash
local fauna from the Upper Cretaceous Ojo Alamo Formation (Naashoibito Member),
San Juan Basin, New Mexico. In Sullivan, Lucas and Spielmann (eds.). Fossil
Record 3. New Mexico Museum of Natural History and Science Bulletin. 53, 216-271.
Williamson and Brusatte, 2014. Small theropod teeth from the Late Cretaceous
of the San Juan Basin, Northwestern New Mexico and their implications for understanding
Latest Cretaceous dinosaur evolution. PLoS ONE. 9(4), e93190.
T? sp. (Langston, Standhardt and Stevens, 1989)
Late Maastrichtian, Late Cretaceous
Lower Javelina Formation, Texas, US
Reference- Langston, Standhardt and Stevens, 1989. Fossil vertebrate
collecting in the Big Bend - History and retrospective. in Vertebrate Paleontology,
Biostratigraphy and Depositional Environments, Latest Cretaceous and Tertiary,
Big Bend Area, Texas. Guidebook Field Trip Numbers 1 a, B, and 49th Annual Meeting
of the Society of Vertebrate Paleontology, Austin, Texas, 29 October - 1 November
1989. 11-21.
T? sp. (Montellano, Monroy, Hernandez-Rivera and Torres, 2009)
Late Campanian, Late Cretaceous
Aguja Formation, Mexico
Material- teeth
Reference- Montellano, Monroy, Hernandez-Rivera and Torres, 2009. Late
Cretaceous microvertebrate fauna from the Northern state of Coahuila, Mexico.
Journal of Vertebrate Paleontology. 29(3), 151A.
T? sp. (Hernandez-Rivera, 1997)
Late Campanian, Late Cretaceous
El Gallo Formation, Mexico
Material- (LACM 42637/HJG 689) tooth (Hilton, 2003)
(42675/HJG 696) tooth (Hilton, 2003)
Comments- These were identified as Saurornithoides by Hilton (2003).
References- Hernandez-Rivera, 1997. Mexican Dinosaurs. in Currie and
Padian (eds). Encyclopedia of Dinosaurs. Academic Press. 433-437.
Hilton, 2003. Dinosaurs and other Mesozoic reptiles of California. University
of California Press. 318pp.
Albertavenator Evans, Cullen,
Larson and Rego, 2017
A. curriei Evans, Cullen, Larson and Rego, 2017
Early Maastrichtian, Late Cretaceous
Horsethief Member of the Horseshoe Canyon Formation, Alberta, Canada
Holotype- (RTMP 93.105.1) incomplete frontal (46.8 mm)
Paratype- (RTMP 96.5.8) incomplete frontal
Referred- ?(RTMP 74.1.1) tooth (6.3x4.6x2.5 mm) (Larson and Currie, 2013)
?(RTMP 74.1.2) tooth (8.4x6.2x3.4 mm) (Larson and Currie, 2013)
?(RTMP 83.12.11) anterior dentary, four teeth (Currie, 1987)
?(RTMP 83.45.5) tooth (7.5x5.7x3.2 mm) (Larson and Currie, 2013)
?(RTMP 83.45.6) tooth (7.4x5.9x3.5 mm) (Larson and Currie, 2013)
?(RTMP 85.12.2) tooth (4.1x3.5x1.7 mm) (Larson and Currie, 2013)
?(RTMP 85.30.32) tooth (5.4x3.1x1.4 mm) (Larson and Currie, 2013)
?(RTMP 85.31.19) tooth (6.6x5.3x2.8 mm) (Larson and Currie, 2013)
?(RTMP 87.12.14) tooth (7.8x5.3x2.8 mm) (Larson and Currie, 2013)
?(RTMP 87.12.19) tooth (7.4x6.1x3.3 mm) (Larson and Currie, 2013)
?(RTMP 88.96.2) tooth (8.4x6.3x3.3 mm) (Larson and Currie, 2013)
?(RTMP 88.96.7) tooth (8.9x6.3x3.1 mm) (Larson and Currie, 2013)
?(RTMP 89.125.14) tooth (4.5x2.4x2 mm) (Larson and Currie, 2013)
?(RTMP 90.82.24) tooth (7.8x6x3.3 mm) (Larson and Currie, 2013)
?(RTMP 90.82.25) tooth (4.9x3.4x1.6 mm) (Larson and Currie, 2013)
?(RTMP 90.82.26) tooth (7.3x5.5x3.1 mm) (Larson and Currie, 2013)
?(RTMP 93.12.13) tooth (7x5.53.3 mm) (Larson and Currie, 2013)
?(RTMP 94.9.4) tooth (7.4x5.7x2.8 mm), tooth (6.1x5.5x2.8 mm) (Larson and Currie,
2013)
?(RTMP 95.2.23) tooth (6.2x5.5x2.5 mm) (Larson and Currie, 2013)
?(RTMP 96.5.8) (Eberth et al., 2013)
?(RTMP 96.5.29) (Eberth et al., 2013)
?(RTMP 96.5.30) (Eberth et al., 2013)
?(RTMP 96.29.29) posterior maxillary tooth (Ryan, Currie, Gardner, Vickaryous
and Lavigne, 1998)
?(RTMP 97.7.3) tooth (7x5.5x3.2 mm) (Larson and Currie, 2013)
?(RTMP 97.39.3) posterior dentary tooth (Ryan, Currie, Gardner, Vickaryous and
Lavigne, 1998)
?(RTMP 97.39.5) dentary tooth (Ryan, Currie, Gardner, Vickaryous and Lavigne,
1998)
?(RTMP 97.39.7) posterior premaxillary or anterior maxillary tooth (Ryan, Currie,
Gardner, Vickaryous and Lavigne, 1998)
?(RTMP 97.77.4) tooth (7x5.6x3.4 mm) (Larson and Currie, 2013)
?(RTMP 98.63.43) tooth (10.7x6.1x3.6 mm) (Larson et al., 2010)
?(RTMP 98.68.156) tooth (8.5x6.4x3.2 mm), tooth (7.6x5.8x3 mm), tooth (8.1x5.6x3.2
mm) (Larson and Currie, 2013)
?(RTMP 99.50.114) tooth (6.8x5.6x3.7 mm) (Larson et al., 2010)
?(RTMP 99.50.115) tooth (10.7x5.8x3.7 mm) (Larson et al., 2010)
?(RTMP 2000.45.10) tooth (Larson et al., 2010)
?(RTMP 2000.45.24) tooth (4.4x3.6x2.2 mm) (Larson et al., 2010)
?(RTMP 2000.45.41) tooth (Larson et al., 2010)
?(RTMP 2000.45.90) tooth (5.2x3.5x2 mm) (Larson et al., 2010)
?(RTMP 2000.45.91) tooth (10x5.4x3.4 mm) (Larson et al., 2010)
?(RTMP 2001.45.80) tooth (12.1x7x4.1 mm) (Larson et al., 2010)
?(RTMP 2001.45.81) tooth (6.6x4.8x2.8 mm) (Larson et al., 2010)
?(RTMP 2002.45.48) tooth (7.7x5.6x? mm) (Larson et al., 2010)
?(RTMP 2003.45.57) tooth (Larson et al., 2010)
?(RTMP 2003.45.58) tooth (7.1x5x2.7 mm) (Larson et al., 2010)
?(RTMP 2003.34.1) tooth (Larson et al., 2010)
?(RTMP 2009.122.1) tooth (9.4x6.3x3.7 mm) (Larson and Currie, 2013)
?(RTMP coll.) six teeth (Baszio, 1997)
?(RTMP coll.) nine premaxillary teeth, thirty-nine maxillary teeth, thirteen
dentary teeth (Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998)
?(RTMP or UALVP coll.) metatarsal (Eberth and Currie, 2010)
?(UALVP 48639) tooth (9.5x?x2.8 mm), tooth (9.3x6x3.3 mm), tooth (9.5x6.2x3.7
mm) (Torices et al., 2014)
?(UALVP 53921) tooth (10x6.2x3.7 mm) (Torices et al., 2014)
?(UALVP 55456) tooth (7.4x5.7x3.3 mm) (Torices et al., 2014)
?(UALVP 55489) tooth (10x6.2x3.6 mm) (Torices et al., 2014)
? partial braincase (Morhardt et al., 2013)
Diagnosis- (after Evans et al., 2017) primary supraciliary foramen truncated
anteriorly by lacrimal contact; superficial (ectocranial) surface of frontal
proportionally shorter than other troodontids, length to width ratio less than
1.3; frontoparietal contact in which an enlarged lappet of the frontal extends
medially to extensively overlap the lateral region of the interfrontal process
of the parietal.
Comments- Currie (1987) described a dentary with teeth that can be referred
to Troodon because of its extensive mandibular symphysis. Although Currie
et al. (1990) state Horseshoe Canyon Formation teeth are essentially identical
to Judith River teeth, Baszio (1997) notes they are are generally smaller than
Dinosaur Park (Judith River) T. formosus, but identical in morphology.
Ryan et al. (1998) state they could not distinguish Horseshoe Canyon teeth from
Judith River teeth, but referred to the former as Troodon sp. due to
the stratigraphic difference. Similarly, Torices et al. (2014) could not distinguish
Horseshoe Canyon and Dinosaur Park teeth statistically. However, Larson and
Currie (2013) found "troodontid teeth from the Horseshoe Canyon Formation
are distinct from the Dinosaur Park Formation teeth." Morhardt et al. (2013)
examined Dinosaur Park and Two Medicine braincases and found "two distinct
morphologies associated with the occipital sinus, in one case being 'peaked'
(dorsally extended, mediolaterally compressed) and the other case being 'rounded'
(dorsal surface gently curves, shows no dorsal extension, and is not mediolaterally
compressed)." The peaked morphology was said to be more similar to Zanabazar.
Most recently, Evans et al. (2017) found Dinosaur Park and Judith River teeth
largely overlapped the morphospace of Horseshoe Canyon teeth, though the spaces
that ignore the 5% of outliers have a more limited degree of overlap. They also
agreed with Currie that Dinosaur Park and Horseshoe Canyon dentaries cannot
be distinguished, but described two Horseshoe Canyon frontals as the types of
their new taxon Albertavenator curriei based on several difference from
Dinosaur Park frontals. Discovered in 1993, the holotype was listed along with
the paratype as Troodon formosus elements by Eberth et al. (2013). The
lack of any associated specimens means that only the frontals can be confidently
assigned to Albertavenator, as it is possible multiple troodontid taxa
coexisted in the formation.
References- Currie, 1987. Bird-like characteristics of the jaws and teeth
of troodontid theropods (Dinosauria, Saurischia). Journal of Vertebrate Paleontology.
7, 72-81.
Currie, Rigby and Sloan, 1990. Theropod teeth from the Judith River Formation
of southern Alberta, Canada. in Carpenter and Currie (eds.). Dinosaur Systematics:
Perspectives and Approaches. Cambridge University Press, New York. pp. 107-125.
Baszio, 1997. Investigations on Canadian dinosaurs: systematic palaeontology
of isolated dinosaur teeth from the Latest Cretaceous of south Alberta, Canada.
Courier Forschungsinstitut Senckenberg. 196, 33-77.
Ryan, Currie, Gardner and Livigne, 1997. Baby hadrosaurid material associated
with an unusually high abundance of troodontid teeth from the Horseshoe Canyon
Formation (Early Maastrichtian), Alberta, Canada. Journal of Vertebrate Paleontology.
17(3), 72A.
Ryan, Currie, Gardner, Vickaryous and Lavigne, 1998. Baby hadrosaurid material
associated with an unusually high abundance of Troodon teeth from the
Horseshoe Canyon Formation, Upper Cretaceous, Alberta, Canada. Gaia. 15, 123-133.
Evans, Lam, Maddin and Conacher, 2003. Taphonomy of the Prehistoric Park Quarry,
Horseshoe Canyon Formation, Drumheller, Alberta. Alberta Palaeontological Society,
Seventh Annual Symposium, "Fossils in Motion" Abstracts. 25-28.
Eberth and Currie, 2010. Stratigraphy, sedimentology, and taphonomy of the Albertosaurus
bonebed (upper Horseshoe Canyon Formation; Maastrichtian), southern Alberta,
Canada. Canadian Journal of Earth Sciences. 47(9), 1119-1143.
Larson, Brinkman and Bell, 2010. Faunal assemblages from the upper Horseshoe
Canyon Formation, an early Maastrichtian cool-climate assemblage from Alberta,
with special reference to the Albertosaurus sarcophagus bonebed. Canadian
Journal of Earth Sciences. 47(9), 1159-1181.
Eberth, Evans, Brinkman, Therrien, Tanke, Russell and Sues, 2013. Dinosaur biostratigraphy
of the Edmonton Group (Upper Cretaceous), Alberta, Canada: Evidence for climate
influence. Canadian Journal
of Earth Sciences. 50(7), 701-726.
Larson and Currie, 2013. Multivariate analyses of small theropod dinosaur teeth
and implications for paleoecological turnover through time. PloS ONE. 8(1),
e54329.
Morhardt, Ridgely, Varricchio and Witmer, 2013. New studies of braincase anatomy,
brain size, and brain structure in the Late Cretaceous theropod Troodon formosus
(Dinosauria: Saurischia) based on CT scanning and 3D visualization. Journal
of Vertebrate Paleontology. Program and Abstracts 2013, 180.
Torices, Funston, Kraichy and Currie, 2014. The first appearance of Troodon
in the Upper Cretaceous site of Danek Bonebed, and a reevaluation of troodontid
quantitative tooth morphotypes. Canadian Journal of Earth Sciences. 51(11),
1039-1044.
Evans, Cullen, Larson and Rego, 2017. A new species of troodontid theropod (Dinosauria:
Maniraptora) from the Horseshoe Canyon Formation (Maastrichtian) of Alberta,
Canada. Canadian Journal of Earth Sciences. 54, 813-826.
Linhevenator Xu, Tan, Sullivan,
Han and Xiao, 2011
L. tani Xu, Tan, Sullivan, Han and Xiao, 2011
Campanian, Late Cretaceous
Wulansuhai Formation, Inner Mongolia, China
Holotype- (LH V0021) (~23 kg adult) incomplete skull (~220 mm), premaxillary
tooth, partial surangular, lateral tooth, six incomplete anterior or mid dorsal
vertebrae (~29 mm), incomplete scapula (~170 mm), incomplete humerus (~95 mm),
incomplete ischium, femur (~240 mm), metatarsal I (~25 mm), incomplete phalanx
I-1 (~23 mm), incomplete pedal ungual I (~31 mm), incomplete metatarsal II (~120
mm), incomplete phalanx II-1 (28 mm), incomplete phalanx II-2 (14 mm without
heel), incomplete pedal ungual II (~50 mm), metatarsal III (150 mm), phalanx
III-1 (~40 mm), incomplete phalanx III-2 (~29 mm), incomplete phalanx III-3
(~25 mm), incomplete pedal ungual III (~31 mm), incomplete metatarsal IV (~145
mm), phalanx IV-1 (~35 mm), phalanx IV-2 (~18 mm), incomplete phalanx IV-3 (~16
mm), incomplete phalanx IV-4 (~17 mm), incomplete pedal ungual IV (~25 mm)
Diagnosis- (after Xu et al., 2011) jugal with lateral flange; surangular
crest anteroventrally oriented; presence of medial expansion near distal end
of femur; wide longitudinal groove present along distal third of dorsal surface
of metatarsal III.
Comments- Discovered in 2009. This was included in a version of Senter's
TWiG coelurosaur analysis and found to be a derived troodontid in a polytomy with
Troodon and Saurornithoides+Zanabazar.
Reference- Xu, Tan, Sullivan, Han and Xiao, 2011. A short-armed troodontid
dinosaur from the Upper Cretaceous of Inner Mongolia and its implications for
troodontid evolution. PLoS ONE. 6(9), e22916.
Talos Zanno, Varricchio, O’Connor,
Titus and Knell, 2011
T. sampsoni Zanno, Varricchio, O’Connor, Titus and Knell,
2011
Late Campanian, Late Cretaceous
Kaiparowitz Formation, Utah, US
Holotype- (UMNH VP 19479) (4 year old subadult) mid dorsal vertebra (22.86
mm), partial mid dorsal centrum, partial posterior dorsal centrum, dorsal fragments,
first sacral centrum (19.98 mm), caudal fragment, proximal chevron fragments,
radial fragment, ulna (93.4 mm), partial ilium, ilial fragments, incomplete
pubes, incomplete ischia, partial femur, partial tibia, partial fibula, astragalus
(one fragmentary), calcanea (one fragmentary), metatarsal I, phalanx I-1 (18.7
mm), pedal unguals I (22.89 mm), metatarsals II (158.86 mm), phalanx II-1 (32.29
mm), phalanges II-2 (22.29 mm), pedal ungual II, partial metatarsals III, phalanges
III-1 (35.58 mm), phalanges III-2 (25.96 mm), phalanges III-3 (25.68 mm), pedal
ungual III, metatarsals IV (175.88 mm), phalanges IV-1, phalanges IV-2, phalanx
IV-3 (17.12 mm), phalanges IV-4 (18.02 mm), pedal unguals IV (18.45 mm)
Referred- ?(ALF coll.) teeth (Zanno et al., 2011)
?(OMNH 21958) tooth (Parrish, 1999)
?(RAM coll.) teeth (Zanno et al., 2013)
?(UCM 83253) tooth (Parrish, 1999)
?(UCM 8659; in part) tooth (Parrish, 1999)
?(UCMP 149171) partial squamosals, parietals, basioccipital, proximal tibia,
fragmentary metatarsals, pedal phalanges, pedal unguals (Zanno et al., 2009;
described by Zanno et al., 2011)
?(UMNH VP 11806) dentary tooth (Zanno et al., 2011)
?(UMNH VP 12507) maxillary tooth (Zanno et al., 2011)
?(UMNH VP 16303) frontal (65.2 mm) (Zanno, 2007; described by Zanno et al., 2011)
?(UMNH VP coll.) distal caudal vertebra (Zanno et al., 2009; described by Zanno
et al., 2011)
?(UMNH VP coll.) teeth (Zanno et al., 2011)
Diagnosis- (after Zanno et al., 2011) acetabular margin of ischium strongly
concave dorsally; notch between lateral condyle and ascending process of astragalus
in lateral view, poorly developed; intercondylar bridge of astragalus hyperconstricted;
proximal groove separating astragalar body and ascending process craniocaudally
wide and v-shaped; cranioproximal groove on astragalar condyles absent; astragalar
condyles subequal in cranial extent; shaft of metatarsal II markedly compressed
(midshaft length-to-transverse width ratio 38.8); metatarsal III distally ginglymoid;
pronounced, rounded tab on dorsolateral aspect of metatarsal III proximal to
distal condyle; distal corner of lateral collateral ligament pit on metatarsal
IV with small protuberance, creating rounded depression between extensor aspects
of distal condyles.
Comments- Discovered in 2008 and initially announced by Zanno et al.
(2009). The holotype was included in a version of Norell et al.'s 2000 troodontid
analysis and found to be at least as derived as Byronosaurus.
Parrish and Eaton (1991) list troodontid remains, which Hutchison et al. (1997) lists as Troodon sp.. The partial skeleton UCMP 149171 was discovered in 1994. Zanno et al. (2009)
mistakenly cite it as UCMP 143270, which is a Parasaurolophus specimen.
Zanno et al. (2011) briefly describe the remains and note they may be referrable
to the contemporaneous Talos. Zanno (2007) first mentioned UMNH VP 16303
as a taxon distinct from Troodon, though it was later relegated to Troodontidae
indet.. Evans et al. (2017) find it to be outside the range of variation of Dinosaur
Park frontals (Latenivenatrix/Stenonychosaurus) and distant from Albertavenator
from the Hoseshoe Canyon Formation as well. The tooth UMNH VP 12507
has large distal serrations (12) and no mesial serrations.
References- Parrish and Eaton, 1991. Diversity and evolution of dinosaurs
in the Cretaceous of the Kaipirowits plateau, Utah. Journal of Vertebrate Paleontology.
11(3), 50A.
Hutchison, Eaton, Holroyd and Goodwin, 1997. Larger vertebrates of the
Kaiparowits Formation (Campanian) in the Grand Staircase-Escalante
National Monument and adjacent areas. In Hill (ed.). Learning from the
Land: Grand Staircase-Escalante National Monument Science Symposium
Proceedings. U.S. Department of the Interior, Bureau of Land
Management. 391-398.
Parrish, 1999. Dinosaur teeth from the Upper Cretaceous (Turonian-Judithian)
of southern Utah. In Gillette (ed.). Vertebrate Paleontology in Utah. Utah Geological
Survey, Miscellaneous Publication. 99-1, 319-321.
Zanno, 2007. A new troodontid theropod from the Late Campanian Kaiparowits Formation,
southern Utah. Journal of Vertebrate Paleontology. 27(3), 170A-171A.
Zanno, Varricchio, Titus, Wilkins and Knell, 2009. A new troodontid (Theropoda:
Paraves) specimen from the Upper Campanian Kaiparowitz Formation, Southern Utah:
Estimating the taxonomic diversity of North American Troodontidae. Journal of
Vertebrate Paleontology. 29(3), 205A.
Zanno, Varricchio, O’Connor, Titus and Knell, 2011. A new troodontid theropod,
Talos sampsoni gen. et sp. nov., from the Upper Cretaceous western interior
basin of North America. PLoS ONE. 6(9), e24487.
Zanno, Loewen, Farke, Kim, Claessens and McGarrity, 2013. Late
Cretaceous theropod dinosaurs of southern Utah. In Titus and Loewen
(eds.). Advances in Late Cretaceous Western Interior Basin Paleontology
and Geology. Indiana Press. 504-525.
Evans, Cullen, Larson and Rego, 2017. A new species of troodontid theropod (Dinosauria:
Maniraptora) from the Horseshoe Canyon Formation (Maastrichtian) of Alberta,
Canada. Canadian Journal of Earth Sciences. 54, 813-826.